1

Great Clarendon Street, Oxford ox2 6dp

Oxford University Press is a department of the University of Oxford.
It furthers the University’s objective of excellence in research, scholarship,
and education by publishing worldwide in
Oxford New York
Auckland Cape Town Dar es Salaam Hong Kong Karachi
Kuala Lumpur Madrid Melbourne Mexico City Nairobi
New Delhi Shanghai Taipei Toronto
With offices in
Argentina Austria Brazil Chile Czech Republic France Greece
Guatemala Hungary Italy Japan Poland Portugal Singapore
South Korea Switzerland Thailand Turkey Ukraine Vietnam
Oxford is a registered trade mark of Oxford University Press
in the UK and in certain other countries
Published in the United States
by Oxford University Press Inc., New York
© Daniel Zohary, Maria Hopf, and Ehud Weiss 2012
The moral rights of the authors have been asserted
Database right Oxford University Press (maker)
First edition published 1988
Second edition published 1993
Third edition published 2000
Fourth edition published 2012
All rights reserved. No part of this publication may be reproduced,
stored in a retrieval system, or transmitted, in any form or by any means,
without the prior permission in writing of Oxford University Press,
or as expressly permitted by law, or under terms agreed with the appropriate
reprographics rights organization. Enquiries concerning reproduction
outside the scope of the above should be sent to the Rights Department,
Oxford University Press, at the address above
You must not circulate this book in any other binding or cover
and you must impose the same condition on any acquirer
British Library Cataloguing in Publication Data
Data available
Library of Congress Cataloging in Publication Data
Data available
Typeset by SPI Publisher Services, Pondicherry, India
Printed and bound by
CPI Group (UK), Ltd, Croydon, CR0 4YY

ISBN 978–0–19–954906–1

1 3 5 7 9 10 8 6 4 2
Dedicated to the memory of Dr Maria Hopf
This page intentionally left blank
 Preface to the fourth edition
Sadly, Dr Maria Hopf passed away since the publi-
cation of the third edition. As a token of our enor-
mous appreciation of her warm, kind personality,
her work in the compilation and updating of this
book, and her other scientific achievements, we
dedicate this edition to her memory.
A considerable amount of new data and research
avenues came to light over the past decade, trigger-
ing the preparation of this new edition. Many new
archaeological sites, some from untreated regions,
have been excavated and analyzed archaeobotani-
cally, and new research tools developed. These
tools, mostly in the field of molecular analysis of
living plants (and animals), drove many enthusias-
tic research groups to explore questions relating to
the domestication process and the relationship
between crops and their wild ancestors. Such new
representative data are included.
Main changes in the fourth edition
The field of radiocarbon dating improved dramati-
cally in the last generation, and dating labs are col-
laborating more extensively with each other and
research. Although not all sites mentioned in this
edition were dated to the same methodical degree,
we now have a much more accurate baseline for
comparison. Therefore, this edition makes use of
calibrated dates, when they are available. As a
result, our ability to compare sites has improved,
and greater accuracy gained in our understanding
of the movement of south-west Asian assemblage
crops beyond their ‘core area’.
In an attempt to improve our knowledge about
the transfer of the Neolithic package to Europe,
Asia, and Africa, we consulted our colleagues for
the most representative sites in each country. As a
result, Chapter 10 ‘Plant remains in representative
archaeological sites’, has been significantly updated.
Thirty-eight sites were deleted and sixty-four were
added (36% new sites out of a total of 179 sites).
North Africa has been added to the geographical
coverage of this edition, but the Indian subconti-
nent was excluded. Many research projects flour-
ished in the Indian subcontinent and eastern Asia in
the last decade, and we believe that these regions
deserve specific treatment.
We have made the text easier to read, with
renewed artwork, figures, tables, and maps.
Nineteen colour plates and eight new black and
white figures have been added, as well as a new
map, (Map 2) which summarizes the spread of the
south-west Asian Neolithic crop assemblage in
Europe, west Asia, and North Africa.
The concluding chapter now appears at the start of
the book, and is called, ‘The current state of the art.’
We also decided to adhere to geographers’ current
recommendation to use the term ‘south-west Asia’
instead of ‘Middle East’ and ‘Near East,’ which are
both ambiguous and Eurocentric geographic terms.
Acknowledgements
We wish to express our gratitude to the many col-
leagues who helped us in the preparation of this
edition. We are particularly indebted to those who
advised us on the current state of research in their
regions; this was essential in our ability to present
a much revised edition. Among them are: Felix
Bittmann, Ksenija Borojevic, Laurent Bouby, Otto
Brinkkemper, Ramon Buxó, Ahmed Fahmy,
Andrew Fairbairn, Ferenc Gyulai, Maria Hajnalova,
Roman Hovsepyan, Stefanie Jacomet, Glynis
Jones, Sabine Karg, Mordechai Kislev, Marianne
viii PREFACE TO THE FOURTH EDITION
Kohler-Schneider, Angela Kreuz, Terttu Lempiäinen,
Elena Marinova, Felicia Monah, Dominique de
Moulins, Aldona Mueller-Bieniek, Mark Nesbitt,
Simone Riehl, David Earle Robinson, Mauro
Rottoli, Anaya Sarpaki, Margareta Tengberg, João
Tereso, Tania Valamoti, Marijke van der Veen,
Karin Viklund, George Willcox. Mordechai Kislev,
Moshe Feldman, and particularly Mark Nesbitt,
offered valuable criticism in reading an initial
revised draft.
We are grateful to Ori Fragman, Mordechai
Kislev, Linda Learn (Class Act Fabrics), Oxford
University Press, for providing new pictures for
this edition, to Reuven Soffer from Soffer
Cartography inc., Jerusalem, for producing all
maps, and to the Photography and Graphic Design
Departments, Weizmann Institute of Science, for
the new and renewed pictures and tables. Also, to
Dorian Fuller for his permission to photograph
Aegilops tauschii from his reference collection, and
to Sue Colledge for sharing with us her Access
database of archaeobotanical finds across south-
west Asia and Europe.
Special thanks are due to Elisabetta Boaretto for
her assistance with updating radiocarbon dating,
and to Aaron Rottenberg for his assistance with
updating the molecular analysis. We also want to
thank Anat Hartmann-Shenkman, Yael Mahler-
Slasky, Leon Cherniaev and Ravit Ferera for their
assistance in preparing this edition.
D.Z. Jerusalem
E.W. Ramat-Gan & Rehovot
2011
 Preface to the third edition
In the last seven years, considerable progress has
been made in our understanding of the origin and
spread of cultivated plants in west and central Asia,
in the Mediterranean basin, and in the temperate
parts of Europe. Today the wild ancestors of the
crops that initiated and sustained food production
in this part of the world are already well-identified.
Moreover, the archaeobotanical evidence assembled
on the origins and spread of these cultivated plants
is today much more extensive and convincing than
the data available only a few years ago.
In this third edition, an attempt is being made to
integrate the new evidence and to update the book.
Significantly, the additional information does not
contradict the main conclusions presented in the
first (1988) and second edition (1993). Moreover, it
adds considerably to the clarity of the portrayed
picture.
To keep the book more or less to its original size,
we decided to focus on domestication, and to omit
the chapter on fruit collected from the wild that
appeared in earlier editions. The knowledge about
collection from the wild has increased greatly over
the last few years; we believe the subject deserves to
be treated separately.
We wish to express our gratitude to the many col-
leagues who helped us in the preparation of this
edition. We are particularly indebted to Corrie
Bakels, Dorian Q. Fuller, David R. Harris, Gordon
C. Hillman, Stefanie Jacomet, Mordechai E. Kislev.
Karl-Heinz Knötrzer, Helmut Kroll, Desanka Kučan,
Naomi F. Miller, Mark Nesbit, Jurgen Schultze-
Motel, Krystyna Wasyliltowa. George H. Willcox
and Willem van Zeist.
D.Z. Jerusalem
M.H. Mainz
2000
Contents

1 Current state of the art 1

Beginnings of domestication 1
Neolithic south-west Asian crop assemblage 1
Wild progenitors 3
The spread of south-west Asian crops 4
Availability of archaeological evidence 4
Early domestication outside the ‘core area’ 5
Beginning and spread of horticulture 5
Vegetables 6
Weeds and crops 7
Migrants from other agricultural regions 7

2 Sources of evidence for the origin and spread of domesticated plants 9

Archaeological evidence 9
Evidence from the living plants 13
Radiocarbon dating and dendrochronology 17

3 Cereals 20
Wheats: Triticum 23
Einkorn wheat: Triticum monococcum 34
Emmer and durum-type wheats: Triticum turgidum 39
Bread wheat: Triticum aestivum 47
Timopheev’s wheat: Triticum timopheevii 51
Barley: Hordeum vulgare 51
Rye: Secale cereale 59
Common oat: Avena sativa 66
Broomcorn millet: Panicum miliaceum 69
Foxtail millet: Setaria italica 71
Latecomers: sorghum and rice 72

4 Pulses 75
Lentil: Lens culinaris 77
Pea: Pisum sativum 82
Chickpea: Cicer arietinum 87
Faba bean: Vicia faba 89
Bitter vetch: Vicia ervilia 92

xiii
xiv CONTENTS

Common vetch: Vicia sativa 95

Grass pea: Lathyrus sativus 95
Spanish vechling: Lathyrus clymenum 97
Fenugreek: Trigonella foenum-graecum 97
Lupins: Lupinus 98

5 Oil- and fibre-producing crops 100

Flax: Linum usitatissimum 101
Hemp: Cannabis sativa 106
Old World cottons: Gossypium arboreum and G. herbaceum 107
Poppy: Papaver somniferum 109
Gold of pleasure: Camelina sativa 111
Other cruciferous oil crops 112
Sesame: Sesamum indicum 112

6 Fruit trees and nuts 114

Olive: Olea europaea 116
Grapevine: Vitis vinifera 121
Fig: Ficus carica 126
Sycamore fig: Ficus sycomorus 130
Date palm: Phoenix dactylifera 131
Pomegranate: Punica granatum 134
Apple: Malus domestica 135
Pear: Pyrus communis 138
Plum: Prunus domestica 140
Cherries Prunus avium and P. cerasus 143
Latecomers: apricot, peach, and quince 144
Carob: Ceratonia siliqua 145
Citrus fruits 146
Almond: Amygdalus communis 147
Walnut: Juglans regia 149
Chestnut: Castanea sativa 150
Hazelnut: Corylus avellana 151
Pistachio: Pistacia vera 151

7 Vegetables and tubers 153

Watermelon: Citrullus lanatus 153
Melon Cucumis melo 154
Leek: Allium porrum 155
Garlic: Allium sativum 156
Onion: Allium cepa 157
Lettuce: Lactuca sativa 157
Chufa or rush nut: Cyperus esculentus 158
Cabbage: Brassica oleracea 158
Turnip: Brassica rapa 159
Beet: Beta vulgaris 159
Carrot: Daucus carota 160
Celery: Apium graveolens 160
 CONTENTS xv

Parsnip: Pastinaca sativa 161

Asparagus: Asparagus officinalis 161

8 Condiments 163
Coriander: Coriandrum sativum 163
Cumin and dill: Cuminum cyminum and Anethum graveolens 164
Black cumin: Nigella sativa 164
Saffron: Crocus sativus 165

9 Dye crops 166

Woad: lsatis tinctoria 166
Dyer’s rocket: Reseda luteola 167
Madder: Rubia tinctorum 167
True indigo: Indigofera tinctoria 168
Safflower: Carthamus tinctorius 168

10 Plant remains in representative archaeological sites 169

Iran 169
Iraq 170
Turkey 171
Syria 172
Israel and Jordan 173
Egypt 174
Libya 176
Morocco 176
Caucasia and Transcaucasia 176
Central Asia 176
Cyprus 177
Greece 177
Crete 179
Former Yugoslavia 179
Bulgaria 179
Rumania 181
Moldavia and Ukraine 181
Hungary 182
Austria 183
Italy 184
Poland 185
Czech Republic and Slovakia 185
Switzerland 186
Germany 188
The Netherlands 189
Belgium 189
Denmark 189
Sweden 190
Norway 190
Finland 190
Britain and Ireland 190
xvi CONTENTS

France 191
Spain 192
Portugal 193

Appendix A: Site orientation maps 194

Appendix B: Chronological chart for the main geographical regions mentioned
in the book 197
Appendix C: Information on archaeological sites which appear on Map 2 200
References 201
Index 237
 OUP CORRECTED PROOF – FINAL, 02/08/2012, SPi
CHAP TER 1
Current state of the art
The aim of this book is to review available informa-
tion on the origin and spread of domesticated plants
in south-west Asia, Europe, and the Mediterranean
Basin. Two sources of evidence exist: firstly, infor-
mation obtained by the analysis of plant remains
retrieved from archaeological excavations, where
early archaeological contexts—namely Epipalaeo-
lithic/Mesolithic, Neolithic, and Bronze Age cul-
tures—are the main source; and secondly, data
provided by living plants, particularly by the wild
progenitors of domesticated plants. This chapter
presents the conclusions of the book as determined
from the combined information provided by these
two sources (relevant data and references will be
presented in the following chapters).
Beginnings of domestication
The first definite signs of domesticated plants in the
Old World appear in a string of Early Pre-Pottery
Neolithic B (PPNB) farming villages that developed
in south-west Asia (Map 1) by ca. 10,500–10,100
calibrated years before present (cal BP). Spikelet
forks of emmer and einkorn wheat with telltale,
rough disarticulation scars (pp. 24, 30–31) provide
the most convincing evidence that these cereals
were already domesticated by this time, and in this
area. The contemporary appearance of relatively
plump kernels further supports this notion, but
cannot be regarded as a fully reliable indication of
the early stage of domestication. These remains and
further evidence of pre-domestication cultivation
suggest that the actual beginning of wheat cultiva-
tion in this area should have been even earlier. No
convincing pre-PPNB domesticated plants have yet
been found.
There is a scholarly debate as to whether agricul-
ture originated in several places across a wide area,
including the Levant and northern Fertile Crescent
(e.g. Weiss et al. 2006; Willcox et al. 2008), or whether
it evolved in only one part of the Fertile Crescent,
such as south-east Turkey (e.g. Lev-Yadun et al.
2000). Although current archaeobotanical data sup-
port the first view, this critical question requires
more archaeobotanical and radiocarbon dating evi-
dence to support any definitive finding.
Neolithic south-west Asian crop
assemblage
The crops of early Neolithic agriculture in south-
west Asia are fairly well recognized. The most
numerous vegetable remains in early farming vil-
lages come from three cereals: emmer wheat
(Triticum turgidum subsp. dicoccum), einkorn wheat
(T. monococcum subsp. monococcum), and barley
(Hordeum vulgare). Diagnostic morphological traits
(non-brittle ears, broad kernels) traceable in the
archaeological finds indicate that by 10,500–10,100
cal BP, these domesticated annual grasses were
intentionally sown and harvested in a string of Pre-
Pottery Neolithic B sites in south-west Asia. Emmer
wheat and barley seem to have been the more com-
mon crops. Einkorn wheat is somewhat less
apparent.
Several grain legumes appear as constant com-
panions of the cereals (see Map 2—Plate 6). The most
frequent pulses in the early Neolithic south-west
Asian contexts are lentil (Lens culinaris) and pea
(Pisum sativum). Two more local legume crops are
bitter vetch (Vicia ervilia) and chickpea (Cicer arieti-
num). In contrast to the cereals, archaeological
1
 OUP CORRECTED PROOF – FINAL, 02/08/2012, SPi
2 DOMESTICATION OF PLANTS IN THE OLD WORLD
Asikli Höyük
10,500-10,000 BP
10,000-9,500 BP
9,500-9,000 BP
einkorn wheat Kissonerga-Mylouthkia +
Shillourokambos
emmer wheat
barley Yiftah'el
chickpea Jericho
flax
lentil
pea
bitter vetch
0 100 200 miles
0 200 400km
Map 1 Archaeological sites in which the earliest south-west Asian domesticated grain crops were reliably identified.
remains of pulses usually lack morphological fea-
tures by which initial stages of domestication can be
recognized. Clear indications of lentil domestication
appear at about 10,100–9,700 cal BP; and of pea,
chickpea, and bitter vetch, at about 9,900–9,500
cal BP. Probably all four legumes were cultivated
somewhat earlier, either together with wheats and
barley or soon after the domestication of those cere-
als. Finally, flax (Linum usitatissimum) belongs to the
south-west Asian group of founder crops. It is
impossible to decide whether the material obtained
from Early Neolithic layers represents collected wild
flax or the remains of domesticated forms. Yet, as in
the case of the legumes both direct and circumstan-
tial evidence indicates that by 9,900–9,500 cal BP,
flax was already domesticated in south-west Asia.
Evidence for early domestication of additional
plants in south-west Asia is much less convincing.
Grass pea (Lathyrus sativus) might have been such a
crop, yet the bulk of its early remains comes from
Çayönü
Cafer Höyük
Tell Abu Jarmo
Hureyra
Tell Aswad
‘Ain Ghazal
Ali Kosh
eighth and seventh millennia BP sites in Greece and
Bulgaria. Signs that rye (Secale cereale) was a south-
west Asian Neolithic crop are much rarer. The ori-
gin and early spread of the faba bean (Vicia faba) is
even less clear.
The plant remains from south-west Asian Pre-
Pottery Neolithic B (PPNB) sites reveal another fea-
ture: as a rule, not a single crop but rather a
combination of cereals, pulses, and flax appears in
these early farming villages. Moreover, the assem-
blage seems to be similar throughout the Fertile
Crescent (see Map 2—Plate 6). In other words, a
common package of grain crops characterizes the
development of agriculture in this ‘core area’.
At almost the same time, signs of herding appear,
implying that sheep and goats had also been
brought under human control. Shortly after, cattle
and pig domestication took place (Zeder 2011).
Thus, an effective south-west Asian Neolithic
food-production ‘package’ was formed, comprising
 OUP CORRECTED PROOF – FINAL, 02/08/2012, SPi

CURRENT STATE OF THE ART 3

60 Legened
61
einkorn wheat
emmer wheat
58 barley
flax
71 62
lentil
pea
2,500-2,000 BP
63 3,000-2,500 BP
3,500-3,000 BP
72 59 4,000-3,500 BP
4,500-4,000 BP
5,000-4,500 BP
5,500-5,000 BP
70 66 6,000-5,500 BP
69 6,500-6,000 BP
65 40 7,000-6,500 BP
56 53
68 7,500-7,000 BP
67 57 38 8,000-7,500 BP
39 21 8,500-8,000 BP
52,55 9,000-8,500 BP
73 35 37 9,500-9,000 BP
41
54 36 10,000-9,500 BP
51 10,500-10,000 BP
48 49 47
33 22
50 34 1
44 23
81 32 4
82
64 74
75 25
78 6
31 24
77 42 29
28 5
43
27 7
79 30 26
45
76 18

80 19 8
3
46
17 9 2

20 14
10
13
11

12

16

Scale 1:16,000,000 15
0 125 250 375 500 miles

0 250 500 km
83

Map 2 The spread of the south-west Asian Neolithic crop assemblage in Europe, west Asia, and north Africa. For details on the numbered sites,
see Appendix C (p. 200). These are the earliest sites in which domesticated grain crops were found, in each country. (See Plate 6.)

vegetative crops as well as domestic animals. Basin are already well identified. The distribution
Indeed, the remains uncovered in south-west Asian areas and the main ecological preferences of most of
PPNB sites indicate a major shift in food practices. them are also well known. Comparison of this evi-
While in Epi-Palaeolithic contexts, gathering and dence with the archaeological findings reveals that
hunting of a wide spectrum of wild species is appar- with practically all early crops, the first signs of
ent, the PPNB farmers already appear to focus on domestication appear in the same general areas
domesticates as their principal source of food. A where the wild ancestral stocks abound today.
large proportion of the remains retrieved from these The geographic distribution of the wild progeni-
early farming sites belong to the crops mentioned tors of Neolithic grain crops is significant. Apart
above and domestic animals. There is also a sharp from flax and barley, the wild ancestors of the
quantitative and qualitative drop in the wild-species founder crops have a rather limited distribution.
intake. An important confirmation of this ‘package’ Wild emmer wheat and wild chickpea are endemic
concept occurred recently with the discovery of just to the Fertile Crescent. Assuming that their distri-
such an ensemble of plants and animals in Early bution did not change drastically during the last ten
PPNB Cyprus, although some of them were not yet millennia, the domestication of these crops could
strictly domesticated. only have taken place in this restricted area. Because
domesticated emmer wheat appears to be the most
important Neolithic crop throughout south-west
Wild progenitors
Asia, Europe, and the Mediterranean Basin, the
The wild ancestors of most of the food plants of confinement of its wild progenitor to the Fertile
south-west Asia, Europe, and the Mediterranean Crescent delimits the place of origin of this
 OUP CORRECTED PROOF – FINAL, 02/08/2012, SPi
4 DOMESTICATION OF PLANTS IN THE OLD WORLD
domesticated cereal. It also marks the rather
restricted geographic area where Old World
Neolithic agriculture could have originated. Wild
forms of einkorn wheat, lentil, pea, and bitter vetch
have a somewhat wider distribution, but all, includ-
ing barley, are centered in the Fertile Crescent; that
is, the region in which the earliest farming villages
have been discovered.
The spread of south-west Asian crops
A most remarkable feature of south-west Asian
Neolithic agriculture is its rapid expansion soon
after establishment in the nuclear area (see Map
2—Plate 6). The quality and quantity of available
archaeobotanical evidence varies considerably from
region to region. Comprehensive information is
available for most parts of Europe, but there is much
sparser and frequently incomplete documentation
from Caucasia, Eastern Europe, and central Asia. In
Africa, critical data on plant remains are available
only for Egypt (but a few current projects might add
vital data for north Africa). In spite of the uneven
documentation, the following main features of the
diffusion of agriculture seem apparent.
The spread of agriculture from its south-west
Asian core to Europe and central Asia involves the
species contained in the Neolithic crop assemblage.
Map 2 (Plate 6) summarizes the information about
the six most important south-west Asian crops:
emmer wheat (including its free-threshing deriva-
tives), einkorn wheat, barley, lentil, pea, and flax.
From the data presented in this map and in Chapter
10, it is evident that crops domesticated in the
south-west Asian core area were the initiators of
food production in Europe, central Asia, and the
Mediterranean Basin (including the Nile Valley).
The earliest farming cultures in these vast regions
always contain wheat and barley, with one, two, or
more of the other south-west Asian founder crops
frequently present as well.
Establishment of the south-west Asian crop
assemblage in the Fertile Crescent and its spread
both west (to Europe) and east (to central Asia and
to the Indian subcontinent) was rapid (see Map
2—Plate 6). From the first farming communities in
the ‘Levantine Corridor’ at ca. 10,500–10,200 cal BP,
it was found to cover the whole Fertile Crescent by
9,500–9,000 cal BP. By ca. 9,000–8,500 cal BP, agricul-
ture had already appeared in Crete and Greece. By
the end of the ninth millennium BP, these crops
were grown in Obre in Bosnia-Hercegovina and in
Jeitun in Turkmenia. Soon after, agriculture appears
as far west as Balma Margineda in Andorra, Spain,
and Sacarovca in Moldavia—and as far south as
Grotta dell’Uzzo in Sicily. By the second half of the
eighth millennium BP, the Linearbandkeramik
farming culture was already firmly established in
loess soil regions throughout central Europe,
extending to Poland in the east, to northern France,
and Germany in the west. At the same time, early
Neolithic farming villages appeared in south Spain,
the Nile Valley, and in Chokh in Caucasia.
Substantial information on the age and spread of
early farming cultures is available for Europe,
where radiocarbon dating of sites exhibiting evi-
dence of early farming enabled the reconstruction
of the diffusion of agriculture. The evidence from
Caucasia, central Asia, and eastern Europe is much
more fragmentary. Yet the finds retrieved from sites
including Jeitun (p. 176) demonstrate that the diffu-
sion of the south-west Asian crops towards central
Asia happened relatively early, although it took
longer to reach Transcaucasia and the Nile Valley.
All over these vast areas, the start of food produc-
tion involved the same south-west Asian crops.
Availability of archaeological evidence
Any attempt to reconstruct the origins and diffu-
sion of agriculture in Eurasia and Africa must
address the uneven archaeological record. As
already mentioned, plant remains of Europe,
south-west Asia, and the Mediterranean Basin pro-
vide us with a reasonable overview of the begin-
nings and development of agriculture in these major
areas. In contrast, the archaeobotanical evidence
from central and eastern parts of Asia and from east-
ern Europe is much less complete. It is very poor in
Africa north of the Sahara. Consequently, while the
early stages of food production in south-west Asia
are relatively well documented, most founder crops
are adequately identified, and the expansion to
Europe and west Asia are convincingly elucidated,
there are far fewer solid facts on crop domestication
and the development of farming in east Asia (Smith
 OUP CORRECTED PROOF – FINAL, 02/08/2012, SPi
1998). However in the last few years, archaeobo-
tanical findings in these agricultural domains have
improved considerably. The history of crop domes-
tication in the African Savanna belt is still largely
uncharted and we still know very little about the
evolution of the unique crop assemblage of this
region (Harlan 1992a).
The time and place of origin of the majority of the
east and south Asian crops, and of practically all the
sub-Saharan African crops, are yet not fully estab-
lished. In numerous cases, the wild progenitors
have not yet been satisfactorily identified or they
are only very superficially known. However, critical
archaeobotanical information has been assembled
on at least two principal crops; rice (Oryza sativa)
and foxtail millet (Setaria italica). Their essential role
in the independent rise of farming in China is now
well documented.
At present, our picture of crop-plant evolution in
Eurasia and Africa is unbalanced. While there is
relatively reliable information on its development
in the classical Old World, we are largely unin-
formed of events south and east of this area. We also
know relatively little about the early interactions
between west Asia and the major agricultural prov-
inces in east and south Asia, and in Africa south of
the Sahara.
Early domestication outside
the ‘core area’
Signs of additional domesticants start appearing
soon after the introduction of south-west Asia
agriculture to Europe, central Asia, and the
Mediterranean Basin. Addition of some of these
crops obviously took place outside south-west
Asia, but they developed within the already estab-
lished agriculture of the south-west Asian crop
assemblage. The poppy, Papaver somniferum, pro-
vides a well-documented example of such domes-
tication. Both the area of distribution of the wild
poppy and the archaeological finds (p. 109–111)
indicate that P. somniferum was brought into
domestication in west Europe. It was added to the
south-west Asian grain-crop assemblage after the
latter’s establishment in western Europe. Chufa,
Cyperus esculentus, is another example of an early
local addition, this time in the Nile Valley (p. 158).
CURRENT STATE OF THE ART 5
Its dry tubers were found in large quantities in
Egypt from pre-dynastic times on. The early
appearance of broomcorn millet, Panicum mil-
iaceum, in the Caspian basin and the Czech
Republic (pp. 69–70) might indicate another local
addition. However, since the archaeological evi-
dence from central and east Asia is still inadequate,
it is impossible to decide whether the Caspian P.
miliaceum was added to the expanding south-west
Asian crop assemblage after it reached central
Asia, or whether this cereal represents an east
Asiatic domestication independent of the south-
west Asian diffusion.
Beginning and spread of horticulture
Olive, grapevine, fig, and date palm seem to have
been the first principal fruit crops domesticated in
the Old World. Definite signs of olive and date-palm
domestication appear in Chalcolithic Levant about
6,800–6,300 cal BP. Indications of date-palm domes-
tication are also available from contemporary lower
Mesopotamia. We still do not know the extent of
Chalcolithic horticulture. Except for the Israel-
Jordan area, the archaeobotanical information from
seventh–sixth millennia BP sites in the Levant is
still insufficient. The picture changes drastically in
the Early Bronze Age (first half of the fifth millen-
nium BP). From this time on, olives, grapes, and figs
emerge as important additions to grain agriculture,
initially in the Levant and soon after, in Greece.
These crops were subsequently planted throughout
the Mediterranean Basin. The extensive Bronze Age
cultivation of olives and grapes is indicated by the
appearance of numerous presses and remains of
storage facilities for olive oil and wine. At the same
time, dates were domesticated on the southern
fringes and the warm river basins of the south-west
Asia, and they abound in the Nile Valley during the
New Kingdom.
Apple, pear, plum, and cherry seem to have been
added much later to Old World horticulture, as
definite signs of their domestication appear only
in the first millennium BC. Their culture is almost
entirely based on grafting, so they could have been
domesticated extensively only after the introduc-
tion of this sophisticated method of vegetative
propagation.
 OUP CORRECTED PROOF – FINAL, 02/08/2012, SPi
6 DOMESTICATION OF PLANTS IN THE OLD WORLD
Remains of fruit trees rarely show diagnostic ana-
tomical traits enabling archaeobotanists to distin-
guish between fruits collected from the wild or
those harvested from domesticated orchards. To a
large extent, recognizing domestication in fruit
crops is based on circumstantial evidence, such as
the finding of fruit remains in areas in which the
wild forms do not occur or on the quantitative anal-
ysis of artefacts associated with fruit products (e.g.
oil, wine). It is difficult, therefore, to determine the
initial stage of fruit crop domestication: in other
words, it might well be that olive, grape, fig, or date
cultivation did not originate in the Chalcolithic
(sixth millennium BP), but was already active in the
late Neolithic (seventh millennium BP).
Despite these uncertainties, the following have
been confirmed: (a) the earliest definite signs of fruit
tree domestication appear in the south-west Asia;
(b) horticulture developed only after the firm estab-
lishment of grain agriculture; (c) as with grain crops,
several local wild fruits were taken into domestica-
tion at about the same time; (d) domestication of
fruit crops relied heavily on the invention of vegeta-
tive propagation; (e) planting of perennial fruit trees
is a long-term investment, promoting a fully settled
way of life; (f) soon after its successful establish-
ment, horticulture spread from its original ‘core
area’ into new territories in the Mediterranean Basin
and south-west Asia; and (g) after the introduction
of grafting (pp. 114–115), the domestication of a
whole group of ‘second-wave’ fruit crops became
possible.
Available archaeobotanical evidence of the
beginning of fruit-crop domestication can also be
supported by information on the wild relatives.
Wild olive, grapevine, fig, and date are widely
distributed over the Mediterranean and south-
west Asia. They have a wide geographic distribu-
tion, so this by itself does not provide critical
values for a precise delimitation of the place of
origin of these fruit crops. Yet it is reassuring to
know that forms from which domesticated clones
could have been derived thrive in wild niches in
the east Mediterranean basin. Therefore, evidence
from the living plants complements the archaeo-
logical finds. Most probably olive, grapevine,
date, fig, as well as pomegranate and almond,
were first brought into domestication in the same
general area where, several millennia earlier, grain
agriculture was successfully established in the
Old World. Thus, during the sixth millennium BP,
eastern Mediterranean Basin human societies
belonging to the Chalcolithic and Bronze Age cul-
tures, were introduced to the use of copper and
bronze, and they also mastered horticulture.
Vegetables
This is the least-known group of domesticated food
plants of the Old World. Vegetable material consists
almost entirely of perishable soft tissues, which
stand a meagre chance of charring and surviving as
archaeological remnants (p. 153). Consequently,
only few vegetable remains have been detected in
excavations. The exceptions here are Egyptian and
Judean Desert caves. In Egypt, especially arid coun-
try vegetables placed in pyramids and graves com-
monly survived by desiccation, and show that
garlic, leek, onion, lettuce, melon, watermelon, and
chufa were cultivated in the Nile Valley in the sec-
ond and the first millennia BC. As amply described
by Keimer (1924, 1984), vegetable gardens consti-
tuted an important element of food production in
Egyptian dynastic times.
Beyond Egypt there are almost no early archaeo-
botanical finds of vegetable crops. However, early
literary sources show that by the start of the second
millennium BC, vegetable gardens flourished not
only in the Nile Valley but also in Mesopotamia.
Furthermore, in both areas the crops grown were
more or less the same. The only major exception
was chufa which was restricted, almost entirely, to
Egypt.
In summary, available evidence makes it clear
that by the Bronze Age vegetable crops were part of
food production both in Lower Mesopotamia and
in Egypt. It is very likely that this geographic pat-
tern is not accidental. In both regions, we are faced
with the dense human settlement of very arid envi-
ronments. Survival in these zones depends on utili-
zation of limited areas of irrigated or flooded land
which is bordered by large, barren deserts. Areas
with no vegetation have little to offer in the way of
supplementary resources of green wild plants. This
shortage invites human initiative. The early devel-
opment of vegetable gardens might have been
 OUP CORRECTED PROOF – FINAL, 02/08/2012, SPi
caused by such needs. It must be taken into consid-
eration that this picture is partly skewed by the lack
of evidence in other regions.
Weeds and crops
Several Old World grain plants, oil producers, and
vegetables seem to be ‘secondary crops’; that is,
they first evolved as weeds and were only later
established as crops (p. 16). Oat, Avena sativa, rye,
Secale cereale subsp. cereale, and gold of pleasure,
Camelina sativa, are well-documented examples of
this mode of evolution under domestication.
Turnip, lettuce, carrot, beet, leek, and several other
vegetables are also very likely to have entered
domestication through the same ‘back door’. The
incorporation of secondary crops into Old World
food production seems to have happened rather
late, since definite signs of their domestication
appear in Europe and west Asia only in the second
and first millennia BC.
Migrants from other agricultural regions
With few exceptions, the classical ‘Old World’
(south-west Asia, the Mediterranean Basin, and
temperate Europe) received crops from other agri-
cultural regions rather late in its agricultural his-
tory. Foreign crops that arrived in this area (in
pre-Columbian times) fall into the following geo-
graphical groups (Zohary 1998):
(a) Temperate climate crops from central
and/or east Asia
Broomcorn millet (Panicum miliaceum) and foxtail
millet (Setaria italica) seem to represent the earliest
arrivals. The origin of P. miliaceum is not fully
understood, but it was probably taken into domes-
tication in central Asia–north China (p. 69–71). It
already appears in Caucasia and in central Europe
in sites around the first half of the eighth millen-
nium BP. S. italica, now recognized as a founder
crop of north China agriculture (p. 71), appeared
in central Europe in the first half of fourth millen-
nium BP, some four thousand years later. For
millet, as well, the available information suggests
arrival from the east (p. 69). However, the pos-
CURRENT STATE OF THE ART 7
sibility of independent domestication of foxtail
millet in the west has not been ruled out yet. Hemp
(Cannabis sativa) reached Anatolia and Europe
much later. Its remains appear (pp. 106–107) from
the eighth century BC onwards. Apricot (Armeniaca
vulgaris) and peach (Persica vulgaris) could have
been taken into domestication either in central
Asia or in China (pp. 154–155); the domesticated
pistachio (Pistacia vera) must have originated in
central Asia (pp. 151–152). The peach seems to
have reached the Mediterranean Basin by the mid-
dle of the first millennium BC. Apricot and pista-
chio arrived only in Roman times.
(b) Warm-weather crops from south and/or
east Asia
A group of more tropical crops (sensitive to freezing
temperatures) that originated in south and/or east
Asia, seem to have migrated into the south-west
Asia and the Mediterranean Basin from the Indian
subcontinent. Many of these cultigens were already
grown in India and Pakistan in the second millen-
nium BC. Sesame (Sesamum indicum) is apparently
the earliest of these migrants (pp. 112–113).
Undisputed remains of this Indian oil crop already
appear in south-west Asia in Iron Age (ca. 900–600
BC) contexts. The citron (Citrus medica) was grown
in the east Mediterranean basin (p. 146) by the end
of the fourth century BC. Asian rice (Oryza sativa)
seems to have arrived (pp. 73–74) in Hellenistic or
early Roman times. The cucumber (Cucumis sativus)
might also have been introduced at the same time
(p. 155). Finally, Old World cottons (Gossypium
arboreum and/or G. herbaceum) could have already
spread from the Indian subcontinent into the south-
west Asia (pp. 107–109) during Roman rule.
However, a fully developed cotton industry
appeared in this area only in Early Islamic times.
An impressive introduction of Indian and south-
east Asian crops was undertaken by the Arabs soon
after their conquests (Watson 1983; Zohary 1998).
The Early Islamic diffusion (eigth–eleventh centu-
ries AD) includes lemon (Citrus limon), lime
(C. aurantiifolia), bitter orange (C. aurantium), pum-
melo (C. maxima), and indigo (Indigofera tinctoria)—
all of which are discussed in this book. It also
involves sugar cane (Saccharum officinarum) and
 OUP CORRECTED PROOF – FINAL, 02/08/2012, SPi
8 DOMESTICATION OF PLANTS IN THE OLD WORLD
sugar extraction technology, banana and plantain
(Musa cultivars), aubergine (Solanum melongena),
and taro (Colocasia esculenta (L.) Schott), although
these crops are not surveyed here.
(c) Warm-weather crops from Africa south
of the Sahara
Although there are several good reasons to assume
(Harlan 1992) that indigenous agriculture was
already well developed in sub-Saharan Africa by
1,000 BC (or even earlier), surprisingly few of the
native African cultigens spread north into the
Mediterranean Basin. This is even more puzzling
since several African grain crops, namely sorghum
(Sorghum bicolor), pearl millet (Pennisetum glaucum),
and cow pea (Vigna unguiculata), seem to have
reached the Indian subcontinent already by the sec-
ond millennium BC (Possehl 1998; Fuller 2000;
Manning et al. 2011). In contrast, only few arrivals
are recorded north of the Sahara. Domesticated sor-
ghum, was grown in Egyptian Nubia from ca 100
AD onwards (pp. 72–73), yet there are no signs of its
spread further north. Advanced durra-type sor-
ghum cultivars appear in south-west Asia only in
Early Islamic times, and as Harlan and Stemler
(1978) argue, they might have arrived not from
Africa but from India. In addition, cowpea (Vigna
unguiculata) is known to have come from Egypt in
Hellenistic and Roman times (Germer 1985, p. 88).
 C H A PTER 2
Sources of evidence for the origin and
spread of domesticated plants
The study of the origin and spread of domesticated
plants is an interdisciplinary venture based on evi-
dence from numerous sources. Several disciplines,
such as archaeology, botany, genetics, chemistry,
anthropology, agronomy, and linguistics are
involved (for review, see Harlan and de Wet 1973),
yet the different sources of evidence vary consider-
ably in reliability and relative weight. The modern
synthesis leans heavily on two principal sources:
(i) information obtained through examination of
plant remains retrieved from archaeological
excavations; and
(ii) evidence gathered from living plants, particu-
larly the main traits that evolved under domes-
tication (‘domestication syndrome’) and the
genetic affinities between the crops and their
wild relatives.
Since 1950s major discoveries have radically
changed our view on the origin of domesticated
plants. They have transformed a realm of plenti-
ful speculation and few solid facts into a well-
documented field.
In the light of this, the contributions of several clas-
sical tools had to be fundamentaly reconsidered. Some
sources of evidence, such as N.I. Vavilov’s monumen-
tal studies of variation and distribution of crops
(Vavilov 1949–50; 1987), had to be considerably revised
and re-evaluated. Others, such as linguistic compari-
sons (widely used by de Candolle 1886), appear to
have retained some relevance, even today. However,
with the flood of archaeological documentation,
genetic information, and new techniques from molec-
ular science, those discoveries now carry less weight.
Readers who wish to acquaint themselves with
some of the more recent research should consult
Pearsall (2000) for archaeobotany, and Hancock
(2004) for plant evolution, genetics, and crop sci-
ences. All serve as a good sources for initial
orientation.
The following sections review the main sources of
evidence on which the modern assessment of crop-
plant evolution is based. A list is given in Table 1.
Archaeological evidence
The primary contribution of archaeology to the
understanding of crop-plant evolution is by the
recovery of plant remains in archaeological excava-
tions, and by identifying to what crop species they
belong. The accumulated evidence contributes to
answering the following questions:
(i) When and where do we find the earliest signs
of domesticated crops?
(ii) How and when did the crops spread to attain
their present distributions?
(iii) What were the early cultigens like?
(iv) What were the main changes in the crops once
they were introduced into cultivation?
(v) Where and when did these changes take
place?
Obviously the key to all answers is the availability of
‘fossil evidence’; that is, sufficient amounts of dated,
culturally defined plant remains amenable to analysis.
The following sections survey the main conditions
under which plant material survives in archaeological
contexts. These are also listed in Table 2.
9
10 DOMESTICATION OF PLANTS IN THE OLD WORLD

Table 1 Sources of evidence on the origin and spread of domesticated plants

I. Archaeological evidence
1. Archaeobotany: Identification of plant remains retrieved from archaeological excavations in connection with cultural associations and
14
C-dating. Determination of the earliest signs of domestication in these plants and their subsequent spread. Changes in crops in time and
space. Crop assemblages in various cultures.
2. Additional evidence
(a) Artefacts. Evaluation of: (i) dated tools associated with cultivation, harvesting, and processing of crops; (ii) cultivation artefacts such as
irrigation canals, terraces, lynchets, plough marks, and cultivation boundaries.
(b) Art. Early drawings, paintings, and reliefs of domesticated plants.
(c) Palynology. Appearance of pollen grains of crops and weeds in dated cores or site contexts.
(d) Weeds associated with agriculture.
(e) Examination of ancient DNA extracted from plant remains.
(f) Chemical analysis. Identification of crops by specific organic residues retained in charred seeds, ancient vessels, charcoal, etc.
(g) Starch analysis. Identification of plant remains and usage of tools by the remains of starch granules.

II. Evidence from the living plants

1. Search for the wild progenitors. Identification of the nearest wild relatives of the domesticated crops by use of:
(a) comparative morphology and comparative anatomy (classical taxonomy).
(b) determination of genetic affinities by cytogenetic analysis.
(c) determination of genetic affinities by DNA and protein resemblances.
2. Distribution and ecology of the wild progenitors
(a) Geographic distribution of the wild relatives (including weedy forms).
(b) Characterization of the habitats and the main adaptations of the wild relatives.
3. Evolution under domestication
Main trends of morphological, physiological and chemical changes. The range and the structuring of genetic variation in the crops and in their wild
progenitors. Development of crop complexes (wild forms, weedy races, and cultigens). Methods of planting, maintenance, and usage.
4. Additional evidence
(a) Genetic systems: characterization of the main systems operating under domestication, especially reproductive systems (including vegetative
propagation).
(b) Genetic interconnections between cultivars and wild relatives.
(c) Intentional and unconscious selections.

III. Other pertinent sources

1. Historical information. Representation of the plants in art, documentation in inscriptions, tablets, manuscripts, and books.
2. Linguistic comparisons. Names of crops in various languages.
3. Circumstantial evidence: Geological, climatic, hydrological, limnological, dendrochronological, anthropological, and zoological indications on the
initiation and spread of agriculture.

material decays rapidly. Carbonized plant remains in

Charred remains
Charred (carbonized) remains are the commonest
source of plant material in archaeological excava-
tions that are available for analysis. Carbonization
occurs on exposure to high temperatures, in most
cases due to fires. Such heating (under a limited sup-
ply of oxygen) converts the plant’s organic com-
pounds into charcoal. Since bacteria, fungi, or other
decomposing organisms do not affect charcoal, car-
bonized plant remains survive in most environments.
This includes wet places where ordinary organic
archaeological contexts are therefore not products of
geological carbonization (true fossils). They repre-
sent only ‘subfossil’ elements charred by fire.
When slowly and mildly charred, wood, seeds,
nuts, and sometimes even fleshy fruits, parenchy-
matous storage tissues, or ears of cereals, can still
retain most of their morphological and anatomical
features. The morphology and the microscopic ana-
tomical structures are frequently preserved in
astonishing clarity. This allows a reliable identifica-
tion of the plant remains.
 SOURCES OF EVIDENCE FOR THE ORIGIN AND SPREAD OF DOMESTICATED PLANTS 11

Table 2 Preservation of plant remains in archaeological ing appear also after the charring of certain seeds.
excavations Moreover, some organs do not generally survive
I. Charred remains
charring (e.g. Boardman and Jones, 1989, Märkle and
Rösch, 2008, Kislev and Rosenzweig, 1991). Thus, the
1. Charred during handling:
seed coats in leguminous plants or the glumes and
(a) near a hearth/oven
(b) in a drying kiln
pales in cereals are only recovered on special occa-
(c) in a storage pit/silo (when cleaned) sions because they disintegrate into powder in most
(d) in pottery fired in a pottery kiln cases. The intensity of the deformation depends,
2. Charred by conflagration: among other things, on the amount of humidity
(a) stored material present in the seed (the drier the grains, the less they
(b) material embedded in daub, unfired bricks, and floors are deformed), the spread of the heating, and the
(c) thatching material temperatures reached.
(d) scattered or dumped material Substantial information on the effects of heating
II. Plant impressions
on the seed of various plants has been gained exper-
imentally by simulation of charring in laboratory
1. In pottery
ovens. Grains of various cereals and seeds of sev-
2. In bricks and daub
eral pulses and flax have been the main elements
III. Parched remains tested. A determination of the amount of shrinkage
1. In arid regions: in the seed of various crops also provides a better
(a) in caves idea of the actual life-size dimensions of charred
(b) in tombs and pyramids seed discovered in excavations (e.g. Märkle and
(c) in clay Rösch, 2008, Braadbaart and van Bergen, 2005).
2. In temperate regions: Such experiments found that the degree of shrink-
(a) in sealed containers age or expansion of seeds vary according to the
(b) in offerings embedded in walls burning circumstances—temperature, time, degree
IV. Waterlogged remains of sealing from oxygen.
1. In lakes Charred plant material is recovered from the
2. In bogs excavated sediment either by direct collection or by
3. In wells separation techniques. There are lucky discoveries
4. In sites covered by rising seawater level of hoards of burnt grains stored in containers or
silos, which sometimes contain almost pure grains.
V. Biomineralization (Phytoliths)
In order to recover scattered remains embedded in
1. Opal
site deposits, the excavator frequently resorts to
2. Calcium oxalate
separation by flotation. Water flotation is the sim-
VI. Metal-oxide preservation plest and cheapest technique, and usually separates
1. Near silver the scattered charred remains present in the depos-
2. Near copper or bronze its effectively. This frequently includes relatively
3. Near iron large amounts of cereal chaff and wood charcoal as
VII. Petrified remains
well as other types of plant material that rarely
appear in silos. The introduction of flotation in the
1. Siliceous mineralization
late 1960s, and especially flotation machines, revo-
2. Calcareous mineralization
lutionized archaeobotany by allowing excavators to
search for seeds rather than rely on caches, and also
At fairly high temperatures (between 200 and 400 improved the efficiency of such separation.
°C), carbonization causes characteristic deforma-
tions. In cereals, the most obvious changes are shrink-
Impressions on pottery, daub, and bricks
age in the length of the kernel together with a relative
increase or ‘puffing’ in its circumference. Size reduc- Imprints of grains and other plant parts on pottery
tions and specific patterns of swelling and/or crack- contribute to documentation of crop plants in
12 DOMESTICATION OF PLANTS IN THE OLD WORLD
archaeological sites. Such imprints are found par-
ticularly on handmade vessels. Pottery is one of the
main diagnostic objects in archaeology, and imprints
on pottery therefore have an obvious advantage,
since once detected, they can be culturally classified
and dated. However, imprints are frequently
pressed into gritty, rough pottery (the common type
of ceramics in early periods). On such a background
the print is rather blurred, and unequivocal inter-
pretation of such findings is often difficult.
Daub and bricks provide another source of plant
impressions. Straw, chaff, and similar dry plant
material is often added to the wet clay to act as a
tempering element. Plant parts can also become
embedded in the clay by chance, and even if the
organic matter does not survive well, the impres-
sions remain intact in the dried or fired clay. They
can serve as negative moulds for casting and repro-
ducing the former inclusions.
Desiccated plant remains
Preservation by desiccation, which blocks the proc-
esses of bacterial and fungal decomposition, occurs
only under extreme dryness, so this source of evi-
dence is confined to very arid areas. Such desiccated
remains can be of particular importance because of
their perfect preservation.
Outstandingly rich remains of dried plants have
been discovered in Egypt. There, grains, fruits, veg-
etables, corms, and other parts of plants placed in
pyramids and tombs give an excellent account of
plant cultivation in the Nile valley during pre-
dynastic and dynastic times. During the last decade
several later sites, from the Iron Age onward, were
discovered along the Red Sea. The finds include soft
parts of vegetables, leaves, and flowers, which
hardly ever survive under other conditions. Several
discoveries of desiccated material were also made
in caves in the Dead Sea basin.
Waterlogged preservation
In Europe, valuable information has been obtained
by examining plant material sunk in peat bogs or
buried in the mud at the bottom of lakes, seas, or
wells. Anaerobic conditions in these environments
(and the presence of humic acids in bogs) act as
effective preservatives, and plant remains in such
places frequently retain their most delicate features.
Excellent examples of waterlogged preservation
have been found in lake-shore dwellings, sub-
merged coastal areas, bottoms of old wells; as well
as in the stomach contents of several human corpses
retrieved from bogs in Denmark, Holland, and
Germany. In some cases, the starch content of
waterlogged-seed remains had vanished.
Preservation by oxides of metals
Bronze, silver, and iron occasionally act as effective
preservatives for plant material buried close to them.
In humid situations they produce metal oxides,
which impregnate the plant remains. Because cop-
per-, silver-, and iron-oxides are highly toxic to bac-
teria and fungi, they block decomposition.
Mineralization
This type of preservation is brought about by filling
of cell cavities by inorganic substances or by replace-
ment of the content of cell walls by minerals. The
most common is mineralization by calcium carbon-
ate (CaCO3), silica, or phosphate.
Seed coats and fruit shells of several plants
undergo natural mineralization. For example,
stones of hackberry (Celtis) contain large quantities
of CaCO3 and the nutlets of several Boraginaceae
accumulate silica. They sometimes survive in
archaeological deposits without further means of
outside preservation.
Phytoliths
Plants deposit the mineral opal (SiO2.nH2O)
between or within their cell walls, which creates
minute silica bodies. As the silica is an inorganic
material, it does not decompose in the soil/sedi-
ment and therefore, it is one of the most dominant
botanical finds in archaeological excavations
worldwide.
Digested or partly digested remains
Preserved human feces (coprolites) constitute an
only partly exploited source of evidence. Since
 SOURCES OF EVIDENCE FOR THE ORIGIN AND SPREAD OF DOMESTICATED PLANTS
humans cannot digest cellulose, woody plant frag-
ments, and shelled seed, they frequently retain their
features after passing through the alimentary tract.
Therefore when feces are charred, desiccated, or
waterlogged, they often contain numerous identifi-
able plant fragments, which indicate the contents of
the food in the tested human culture. Coprolite
examination has already contributed significantly
to American environmental archaeology. In the Old
World this source of evidence has not yet been
exploited extensively, although some results are
already available (e.g. Hillman 1986; Dickson et al.
2000).
Chemical tests
Tar compounds present in charred plant remains
and organic residues precipitated in ancient vessels
can be identified by gas liquid chromatography,
infrared spectroscopy, and other tests used by
organic chemists. Such detection is possible even
when these substances survived in minute traces.
Significantly, some of these chemical compounds
are specific to a single crop species or a single plant
product. They can be used as diagnostic traits for
crop identification (e.g. Evershed 2008).
Evidence from the living plants
Several principal contributions to the understand-
ing of the crop-plant evolution are made by the
study of the living plants.
A major contribution is the identification of the
wild progenitors from which the various domesti-
cated plants could have been derived. Once the
wild ancestry of the crop has been determined the
following examinations can be carried out:
(i) comparison of the cultivated varieties (‘culti-
vars’) with their wild counterparts in order to
determine the main morphological, physiologi-
cal, chemical, and genetic changes that took
place under domestication;
(ii) assessment of the range and the structure of
genetic variation (chromosomal, protein, and
DNA polymorphisms) present in the wild pro-
genitor and those found in the domestic
derivatives;
13
(iii) assessment of changes in adaptation. Answers
can be sought to the following questions: Which
adaptations, that are vital under wild situa-
tions, have broken down under domestication?
What are the new ‘syndromes’ that have
evolved under domestication? Which selective
forces are responsible for these changes?
(iv) delimitation of the distribution areas of the
wild progenitors. This often provides informa-
tion on the place of origin of the crops.
A second major contribution comes from examina-
tion of the crops and the ways they are handled,
particularly under traditional (‘primitive’) systems
of agriculture. Such studies include:
(i) patterns of variation in each crop and their
geographies;
(ii) methods of cultivation and uses;
(iii) genetic systems operating in the various crops
and the breeding traditions used; and
(iv) examination of the interconnections between
the domesticated varieties and their wild rela-
tives. The role of weedy races accompanying
the crops is significant for this topic.
The following sections survey the main tools used
for identification of wild progenitors. They also deal
with some of the complications and problems
involving the use of the wild relatives for elucida-
tion of crop-plant evolution.
Discovery of wild progenitors
A principal goal in the study of the living plants is
the identification of the wild progenitors of the
crops; that is, the wild stocks from which the domes-
ticated plants could have evolved. Plant domestica-
tion is a relatively recent evolutionary event.
Therefore, one can expect that most wild ancestors
are still alive and include forms similar to those that
existed in pre-agricultural times. Indeed, the wild
progenitors of the majority of the world’s main food
plants have already been identified. Many of them
became known only during the last thirty-five
years.
Several complementary tests are available for the
identification of the wild progenitors of crops. They
14 DOMESTICATION OF PLANTS IN THE OLD WORLD
all seek to determine which of the wild species, usu-
ally grouped together with the crop in the same bio-
logical genus, is most closely related to the
domesticated plant.
(i) The classical taxonomic approach recognizes the
wild progenitor by its close morphological resem-
blance to the crop. It sorts out the wild progenitor
from among all the other wild taxa grouped in the
crop’s genus by its closest morphological and ana-
tomical affinities to the cultigen. This is the oldest
method. In some cases morphological comparison
provides sound clues for the determination of
ancestry. However, in numerous cases the wild
background is taxonomically complex. Moreover,
many crops exhibit a bewildering morphological
variation, very different from the patterns present
in wild plants, and this can be confusing in relation-
ship analysis. Critical evaluation in such cases
necessitates genetic verification; which can be
obtained through cytogenetic analysis and by com-
parative molecular (DNA and/or protein) tests.
(ii) Cytogenetic analysis aims at elucidating the
chromosomal affinities between the domesticated
plant and the wild species. It also tests whether or
not these wild taxa are separated from the crop (and
isolated from one another) by hybrid sterility or
other reproductive isolation barriers. Since evolu-
tionary domestication is a recent development, the
crop and its wild progenitor should retain a consid-
erable amount of homology in their chromosomes.
In contrast, other species grouped in the genus were
probably formed long before the beginning of agri-
culture. As a result, they could have diverged con-
siderably in their chromosomal constitution.
The principal tool of cytogenetics is a program of
crosses between species followed by examination of
inter-specific hybrids. Chromosome pairing in mei-
osis indicates the degree of chromosomal homology
between the two parents. In most crops (particu-
larly in grain crops), the cultivars show full homol-
ogy and complete inter-fertility with only one of the
wild species in the tested genus. Such a wild (con-
generic) type is recognized as the ancestor (wild
progenitor) of the crop. Together they comprise the
‘primary’ gene pool of the crop. In contrast, other
members of the genus are frequently chromosoma-
lly distinct and are separated from the crop by
strong reproduction isolation barriers such as cross-
incompatibility, hybrid inviability, or hybrid steril-
ity. Such species are often called ‘alien species’ and
their chromosomes ‘alien chromosomes’. They com-
prise the ‘secondary’ and ‘tertiary’ gene pools of the
crops (Harlan and de Wet 1971, p. 107; Harlan
1992a).
To summarize, fully fertile hybrids showing nor-
mal chromosome pairing in meiosis point to close
genetic relationship between the tested parents and
implicate the wild plant in the ancestry of the crop.
Lack of chromosome homology and the presence of
strong reproductive isolation barriers indicate long-
established genetic divergence and rule out the tested
wild plant from being a progenitor of the crop.
Chromosome analysis of domesticated plants has
frequently also to deal with complications due to
polyploidy; i.e. the formation of new subspecies (or
even new species) by doubling of chromosome
numbers. Evolution by polyploidy is common in the
plant kingdom. Many wild plants (including pro-
genitors of domesticated plants) are not standard
diploids but polyploid entities. One class comprises
auto-polyploids which increased their chromosome
number from the standard of two dosages (diploid
condition) to three sets (triploids), four sets (tetra-
ploids), or even higher levels. Such increases are not
uncommon among vegetatively propagated crops
(corm and tuber plants, ornamentals, and some fruit
trees). A second class includes allo-polyploids; i.e.
types formed by inter-specific hybridization fol-
lowed by chromosome doubling. This combines the
genetic contents of two (or even more) donor species
in a new hybrid species. Bread wheat is a product
of such fusion under domestication (pp. 47–48).
Cultivated tobacco and the New World cottons had
a similar mode of origin. In such crops, a special
cytogenetic test known as ‘genome analysis’ helps to
elucidate the polyploid origin and to identify the
parental stocks, which donated their chromosomes
to the new polyploid entities.
(iii) Advances in molecular biology provide critical
tools for assessing the range of variation, and finding
the structuring of genetic variation (genetic polymor-
phism) in crops and their wild relatives. This infor-
mation can be used for determining the genetic
 SOURCES OF EVIDENCE FOR THE ORIGIN AND SPREAD OF DOMESTICATED PLANTS
affinities and phylogenetic relationships between the
cultivars and their wild relatives. As major break-
throughs in this field occurred since the publication
of the third edition of this book some ten years ago, it
receives extensive treatment in the present edition.
Since the 1960s, critical results have been obtained
by testing protein variation in crops and their wild
relatives, particularly enzyme variants (isozyme
and allozyme polymorphism) and/or variability in
storage proteins deposited in seeds (Soltis and Soltis
1989). Proteins are the primary products of the
genes, and therefore their variability reflects differ-
ences in the hereditary material. Gel electrophoresis
separation makes it possible to discern variation
and differences in numerous proteins. Lately, detect-
ing protein variability has become an outdated
technique although it is time- and cost-efficient. Its
relatively low output makes this technique less
attractive than DNA-based marker systems.
Since the late seventies, large-scale analysis of
DNA variation became possible in a wide range of
techniques. The first major breakthrough for varia-
bility analysis was the development of restriction
fragment length polymorphism (RFLP) technology.
Restriction enzymes cleave the DNA strands at spe-
cific sites (‘restriction’ sites’) into identifiable frag-
ments. Individuals with identical site arrangements
yield identical DNA fragments, while those that
carry mutations in these sites produce different
fragment patterns. Because restriction sites along
the DNA strands of genes are numerous, polymor-
phism in these sites is enormous both within and
between populations of crops and their wild pro-
genitors (e.g. Havey and Muehlbauer 1989).
Another major breakthrough was made by devel-
opment of the polymerase chain reaction technology
(PCR), by which quantities of DNA fragments, large
enough for variation analysis, are amplified from
minute samples of target DNA and their specific
primers. Indeed, PCR is used in almost all DNA
polymorphism analyses. In the mid-nineties, com-
bining PCR and RFLP resulted in a most popular
technique, namely AFLP (amplified fragment length
polymorphism, Mueller and Wolfenbarger 1999).
Although it is still relatively expensive, this method
is highly reliable and informative, and indeed many
of the recent elucidations on crop origins such as
15
wheats, barley, and chickpea were available through
this procedure (Heun et al. 1997b; Badr et al. 2000;
Özkan et al. 2002; Nguyen et al. 2004; Duc et al. 2010).
Another technique that uses arbitrary primers for
the PCR and needs minute amounts of genetic mate-
rial is random amplified polymorphic DNA (RAPD).
It is a straightforward method, but is relatively less
reliable and informative. However, it reveals genetic
diversity easily, as found, for example, in flax and in
lentils (Sharma et al. 1996; Fu et al. 2002).
Another highly informative technique known as
microsatellites, involves short DNA repeats that
show high variation in repeat-number between
individuals (Varshney et al., 2005). Also known as
SSR (simple sequence repeat), this method produces
highly unique patterns that in fact are the basis for
individual DNA fingerprinting in many different
organisms including crops (Molina-Cano et al. 2005)
and humans.
Direct DNA sequencing of the genetic text at tar-
get sites may also reveal variability between indi-
viduals. In general, it retrieves phylogenetic
information from diversity in specific loci in the
DNA, as manifested (for example) in the sad2 locus
in flax (Allaby et al. 2005), or the btr1/2 loci in barley
(Komatsuda et al. 2007).
Remarkable advances in sequencing technology
have recently enabled comparisons of large DNA
sequences that accordingly embed variations or
point mutations in the form of single nucleotide
polymorphisms (SNPs, Rafalski 2002). Such point
mutations are abundant throughout the genome.
They reflect DNA variability and were used recently
to study diversity between and within crops and
their ancestors as done for example in barley
(Kanazin et al. 2002).
Today, the bewildering array of DNA markers
and the ability of automatic sequencing of genes of
interest in hundreds of individuals can be carried
out in a very short time. This revolutionized our
ability to trace and assess genetic and evolutionary
relationships and construct reliable phylogenetic
trees for crops and their progenitors.
Distribution of the wild progenitors
The wild relatives can frequently provide critical
information about where domestication occurred.
16 DOMESTICATION OF PLANTS IN THE OLD WORLD
In many crops, the progenitors occupy limited geo-
graphic territories—much smaller distribution areas
than those of their domesticates. Because domesti-
cation is a recent development, it is safe to assume
that the distributions of the wild forms (weeds
excluded) have not undergone drastic changes since
the beginning of cultivation. Delimitation of the
wild relative’s distribution thus marks the territory
in which the crop could have been taken into culti-
vation. The narrower the distribution area, the more
accurate the placement.
Fortunately, the distribution of the wild progeni-
tor of emmer wheat—a principal Old World ‘founder
crop’—is confined to the Fertile Crescent (Map 4, p.
42). It is thus possible to plot, fairly accurately, the
area where Neolithic agriculture could have started.
The archaeological records have fully corroborated
this supposition. The delimitation of the place of ori-
gin of the chickpea is even more precise. Its wild
ancestor is endemic to south-east Turkey (Map 10, p.
88). However, not all wild progenitors have such a
limited distribution. Some (e.g. the wild relatives of
the foxtail millet, oat, flax, and numerous fruit trees)
are distributed over extensive territories. The use of
their distributions for the determination of places of
origin is much less accurate.
Weeds and domestication
Some crops seem to have entered cultivation not
directly but by first evolving weedy forms. The
establishment of tilled fields (as well as other dis-
turbed habitats) gave an opportunity to numerous
unwanted plants to invade the newly made habitats
and to evolve as weeds. Weed evolution went hand
in hand with crop cultivation and from the very
start the control of these invaders seems to have
been a major problem in agriculture. Noxious weeds
are plants that have successfully adapted them-
selves to the ecology of the tilled ground. They are
independent only because they retain their wild
mode of seed dispersal, and germinate and develop
in spite of the efforts of the cultivator to eradicate
them. But if any such weeds turns out to produce a
valuable commodity, it can eventually change its
relationship with humans. The cultivator may fol-
low the rule ‘if you can’t beat them, join them’, and
start to utilize the weed by intentionally planting its
seed, harvesting its fruits, and selecting the better
yielders. Several Old World crops are such ‘second-
ary crops’, i.e. plants that entered domestication
through the back door of weed evolution (Vavilov
1949–50, 1987). They were added to the crop assem-
blage only after the establishment of the principal
seed crops. Well-documented cases are those of the
oat, Avena sativa (pp. 66–69), and of the gold of
pleasure, Camelina sativa (p. 111). Several other
plants seem to have followed a similar evolution
under domestication.
Classification and botanical names
Orientation in crop plant evolution is frequently
complicated by inconsistencies in species delimita-
tion and by proliferation of botanical names. As
already noted, cultivated plants are, as a rule, very
variable. Furthermore, evolution under domestica-
tion commonly involves drastic modifications in
organs and traits that stay fairly uniform in wild
plants. Traditional taxonomic treatments of crops
suffered from over-splitting, since they were based
almost entirely on morphological comparisons.
Frequently, inter-fertile crop varieties were ranked as
separate species and called by different botanical
names because they looked so different. For example,
classical cereal taxonomists recognized twelve to fif-
teen species of cultivated wheats (see Table 3, p. 29).
Barley and common oat were each split into two or
more species (Table 5, p. 57). Similar splitting and
species ranking characterized numerous other crops.
With the accumulation of cytogenetic informa-
tion, it has become increasingly clear that the tradi-
tional classification of many crops is inadequate
and even misleading. Frequently two, three, or even
half a dozen ‘species’ were found to be inter-fertile,
chromosomally homologous, and genetically inter-
connected. Moreover, in many cases the conspicu-
ous morphological distinctions turned out to be
governed by single mutations (Table 7, p. 61).
Ranking such types as independent species is unjus-
tified. They represent only varieties within species
and deserve only intra-specific ranking. In wheats,
modern taxonomic revision has reduced the species
number to five (Table 3, p. 29). All cultivated barleys
are grouped in a single species (Table 5, p. 57), as are
all common oats.
The discovery of the wild progenitors necessi-
tated another nomenclature change. Because the
 SOURCES OF EVIDENCE FOR THE ORIGIN AND SPREAD OF DOMESTICATED PLANTS
wild plants and their cultivated derivatives are
genetically interconnected, they cannot be regarded
as fully diverged species. According to internation-
ally agreed taxonomic rules, once a wild ancestor is
satisfactorily recognized, the crop and its wild rela-
tive cease to be treated as two separate species.
Instead, they should be lumped in a single collec-
tive species, frequently also including related weed
types. In other words, the wild and crop types are
considered as subspecies or varieties of a single bio-
logical species and botanically named accordingly.
However, habits die hard. Old names and tradi-
tional classifications are still widely used by many
researchers. Wild progenitors, in particular, are
commonly referred to as independent species. To
avoid confusion, botanical orientation in crops
should begin with the following questions:
(i) What are the main cultivated, weedy, and wild
elements in the crop complex?
(ii) What botanical names are used by different
people for these intra-specific taxa of the crop
complex?
(iii) What are the other fully divergent (‘alien’) spe-
cies placed in the same genus?
Radiocarbon dating and
dendrochronology
Radiocarbon (14C) dating was developed by W.F.
Libby at the University of Chicago soon after the
Second World War and created a real breakthrough
in archaeology (Libby et al. 1949). Previously, one
could date archaeological remains only by relative
chronology based on stratigraphy and cultural
associations. The introduction of radiocarbon-
dating methods brought about absolute dates and
made possible age comparisons between cultures in
the various parts of the world.
Until the 1980s, radiocarbon dating demanded
relatively large samples of charred material. Most
tests were made on carbon sources (such as wood
charcoal or hoards of grain) obtained from secured
archaeological contexts. By then, radiocarbon accel-
erator mass spectrometry (AMS) technique had
been developed (Nelson et al. 1977).
This technique, based on counting the atoms
instead of the decay product, made it possible to
reduce the size of the sample to a few milligrams.
17
Consequently, it became possible to date accurately
individual charred grains or to reduce the amount
of material removed from rare or very important
objects. The application of the AMS test is some-
times critical (Harris 1986). Beside dating, radiocar-
bon serves to detect—and eliminate—errors in
chronologically associated samples to the wrong
strata or archaeological level. These errors are due
to intrusion; i.e. the occasional displacement of
plant remains from one layer or context to another
as a result of boring by animals or other interfer-
ences (Boaretto 2007, 2009). Better precision and
accuracy of archaeological dating brought about by
several improvements in this field:
(i) the improved techniques of chemical pre-treat-
ment for the elimination of contaminations
from the sample material;
(ii) the ability to reduce the size of the sample to
few milligrams, extend the range of possible
material to be dated, the care taken to date
archaeological finds from secure, sealed,
contexts;
(iii) the building of a calibration curve up to the
range of radiocarbon dating’ 50,000 year
(Reimer et al. 2009).
The principle of radiocarbon dating is based on
the measurement of the radioactive 14C isotope con-
centration in sample material, in comparison to the
stable carbon isotope 12C. The production of 14C, due
to the interaction of cosmic rays with molecules,
takes place in the atmosphere as neutron absorption
on 14N atoms.
These 14C forms CO2 molecules, are then intro-
duced into different natural reservoirs (e.g. hydro-
sphere, biosphere) by different physical and chemical
processes. In these reservoirs the relative 14C and 12C
concentration is in equilibrium as long as the
exchange with the atmosphere stays open. When
this exchange ceases (e.g. when the organism dies),
the 14C concentration starts to decrease due to decay
of the radioactive 14C. Therefore, the measured con-
centration of 14C in material examined depends upon
the time elapsed from when the exchange with the
atmosphere stopped. The radiocarbon age is calcu-
lated from the measured 14C concentration using the
decay law. The smaller the proportion of 14C in the
tested organic remains, the older the sample.
18 DOMESTICATION OF PLANTS IN THE OLD WORLD

The concentration of 14C in the atmosphere was the limit of radiocarbon dating which is 50,000 years
not constant in the past (Suess 1970), and some fluc- (Reimer et al. 2009). A relevant part of the current
tuations seem to have occurred in the concentration calibration curve is shown in Fig. 1. One can see that
of this isotope in the atmosphere. This means that except for the last three thousand years, the radio-
age estimates based on conventional radiocarbon carbon age represents somewhat reduced estimates.
timescale are in need of some calibration. More pre- Moreover, the differences increase in time. Thus for
cise dating was made possible by establishing the radiocarbon dates 3,000–4,000 years before present
sequences of annual rings in wood remains of trees (BP), calibration adds 200–400 years. For older
(oaks, bristlecone pine), and currently also in corals; radiocarbon dates (6,000–9,000 BP), the addition is
and radiocarbon dating of the rings in these already 700–1,000 years. For still older radiocarbon
sequences. Recently, these sequences have been dates the calibration differences are even greater.
updated and extended to 50,000 years BP (Reimer et The result of the calibration process is usually
al. 2004). By plotting radiocarbon ages against tree- presented in the form of probability distribution of
ring ages, calibration curves have been constructed the radiocarbon age (black curve) as it is shown in
correlating radiocarbon dates with dendro- or cal- Fig. 2. The radiocarbon determination, with one
endar times (Fig. 1). Thus by means of dendro- standard deviation, is given in the center of the
chronology, the radiocarbon timescale can be upper side of the plot. The distribution of the radi-
calibrated against annual tree-ring chronology to ocarbon determination (red curve on the left-hand
calendar dates (Stuiver et al. 1986; Stuiver and Y axis) is projected on the calibration curve (an
Reimer 1993; Baillie 1995). Joined by other dating uneven double blue line on the X axis, in calibrated
methods (U-Th ages from corals), calibration reaches years BP), to produce the radiocarbon age (solid

Atmospheric data from Reimer et al (2004); Oxcal v3.10 Bronk Ramsey (2005); cub r:5 sd:12 prob usp[chron]

25000BP
Radiocarbon determination

20000BP

15000BP

10000BP

5000BP

0BP
25000CaIBP 20000CaIBP 15000CaIBP 10000CaIBP 5000CaIBP
Calibrated date

Fig. 1 The radiocarbon calibration curve for the past 25,000 years based on trees and corral annual ring sequences. Adapted from OxCal 3.10
software Bronk-Ramsey 2005 (Bronk-Ramsey 1995; Bronk-Ramsey 2001). The straight thin line represents the ideal 1:1 correspondence between
radiocarbon age and calendar age assuming constancy of 14C concentration in the atmosphere. If 14C years were the equivalent to calendar years,
all of the data would fall on the diagonal straight line.
 SOURCES OF EVIDENCE FOR THE ORIGIN AND SPREAD OF DOMESTICATED PLANTS 19

Atmospheric data from Reimer et al (2004); Oxcal v3.10 Bronk Ramsey (2005); cub r:5 sd:12 prob usp[chron]
Copyright © 2012. OUP Oxford. All rights reserved. May not be reproduced in any form without permission from the publisher, except fair uses permitted under U.S. or

9200±45BP
9500BP
68.2% probability
9400BP 10420BP (68.2%) 10260BP
95.4% probability
10500BP (95.4%) 10240BP
Radiocarbon determination

9300BP

9200BP

9100BP

9000BP

8900BP

10800CalBP 10600CalBP 10400CalBP 10200CalBP 10000CalBP

Calibrated date

Fig. 2 Presentation of radiocarbon results—probability distribution of a sample that was measured 9200±45 14C year BP (=Before Present);
see text for explanations of this result. The calibration is performed using the software OxCal 3.10 Bronk-Ramsey 2005 (Bronk-Ramsey 1995;
Bronk-Ramsey 2001).

black distribution). The probability of dates is given
in the top-right corner in two different levels of
confidence, 68.2% probability (±1σ) and 95.4%
probability (±2σ). For each probability, the range of
dates is given, from left to right, with the percent-
age in the middle representing how much this
range covers. When there is more than one range of
probabilities for each 1 or 2σ probabilities, each
interval appears separately. A straight line is placed
underneath the black curve to make it easy to deter-
mine the dates as can be seen on the X axis. In case
of more than one range of probabilities for each 1 or
2σ probabilities, such straight line is placed under
each curve.
It is important to note that radiocarbon dates are
given as a probability range, which means the actual
dates could be between two possible dates, rather
applicable copyright law.
ranges. Disturbing such area occur around 10,000
uncal BP—a period of great importance for the
understanding of the beginnings of agriculture.
These areas in the calibration curve are still widely
misunderstood aspects of radiocarbon dating.
As a rule, dates mentioned in this book are cali-
brated radiocarbon dates before present (cal BP).
Therefore, the use of ‘BP’ and ‘cal BP’ denote cali-
brated 14C dates (years before present, i.e. before
1950). BC (years Before Common Era/Before Christ)
denotes uncalibrated dates, either from Bronze Age
and later radiocarbon dates, when calibration has
little impact, when dates drawn from traditional
methods of stratigraphic dating, or when calibrated
dates were unavailable.
Also, progress of evidences in this book is pre-
sented on a timetable moving from older to newer

than the median dates between them. In addition,
as Figs 1 and 2 show, calibration curve is far from
being 1:1 straight line, and several wiggles and flat
areas can be seen. As a result, due to the fact they
fall into a flat area in the calibration curve, several
different radiocarbon dates can produce similar
dates, from earliest to recent finds. Therefore, when
we separate millennia to first and second halves (i.e.
‘the second half of the eleventh millennium cal. BP’
or ‘the first half of the ninth millennium cal. BP’),
the first half will be older and the second one,the
younger.

EBSCO Publishing : eBook Collection (EBSCOhost) - printed on 1/31/2022 10:31 AM via UNIVERSITAET TUEBINGEN
AN: 440843 ; Daniel Zohary, Maria Hopf (deceased), Ehud Weiss.; Domestication of Plants in the Old World : The
Origin and Spread of Domesticated Plants in Southwest Asia, Europe, and the Mediterranean Basin
Account: s9512272
 C H A PTER 3
Cereals
Cereals (annual grasses grown for their grains) have
been the principal crops of most Old World’s civili-
zations. Their grains constitute the main source of
calories for mankind. Different parts of the world
depended on cultivation of different cereals as sta-
ple foods. Food production in the Mediterranean
basin, Europe, the non-tropical parts of Asia, and
(to some extent) the highlands of Ethiopia, was
based primarily on wheat and barley. South and
south-east Asia had rice, broomcorn, and foxtail
millets. America grew maize. Africa south of the
Sahara raised sorghum, pearl millet, and several
other native grasses.
Cereals thrive on open ground (the cultivated
field) and usually complete their life cycle in less
than a year (Plate 1). The nutritive value of their
grains is generally high, and the seed can be stored
for relatively long periods. In most cereals the grains
are packed with starch, and in some, such as in
wheats or oats, they also contain an appreciable
amount of protein. Compared with grain crops
belonging to other plant families (e.g. legumes),
yields in cereals are relatively high. However, they
depend heavily on soil fertility and the availability
of fixed nitrogen.
Wheats and barley are the principal ‘founder
crops’ that started food production in south-west
Asia and Europe. The first definite signs of wheat
and barley domestication appeared in the Fertile
Crescent in the second half of the eleventh millen-
nium cal BP. Later, grains of these cereals consti-
tuted the bulk of plant remains retrieved from
south-west Asian Neolithic, Chalcolithic, and
Bronze Age contexts.
Wheats and barley were the main domesticates
that led the explosive expansion of the Neolithic
20
agriculture from its ‘core area’ to the vast territories
of west Asia, Europe, and north Africa.
Several other grasses entered agriculture in west
Asia and Europe, but apparently, only after the firm
establishment of the ‘first wave’ in south-west Asia.
Such an early domesticate is broomcorn millet, fol-
lowed by common oat.
Two major ‘alien’ cereal crops, namely rice (native
to south-east Asia) and sorghum (native to Africa
south of the Sahara) arrived as well, sometime in
Hellenistic or Roman times. Apparently, their estab-
lishment in Mediterranean agriculture was much
later.
Wheats, barley, and several other prominent cere-
als, differ from the majority of flowering plants in
their system of pollination. While the great majority
of the world’s plant species are allogamous (cross-
pollinated), most domesticated seed crops are pre-
dominantly self-pollinated, or ‘selfing’ (Zohary
1999). Significantly, selfing is also a characteristic
feature of their wild progenitors. In wheat, barley,
and most other grasses, it is brought about by pre-
cocious shedding of the pollen inside the florets;
that is, before they open.
Was it a mere chance that the first grain plants
that were successfully domesticated in the Old
World were selfers? Several facts suggest that pre-
dominantly self-pollinated plants were better suited
to domestication than cross-pollinated candidates.
One major advantage of self- over cross-pollination
in incipient domesticants is the fact that selfing
automatically brings about reproductive isolation
between the crop and its wild progenitor. This isola-
tion barrier enables the farmer to grow a desirable
cultivar in the same area in which its wild relatives
abound, without endangering the cultivar by

genetic swamping. If indeed domestication of
south-west Asian cereals started in areas where the
wild progenitors were common, genetic separation
between tame and wild could have been a major
asset. Under cross-pollination the initial small
patches of cultivation would have been exposed to
large quantities of pollen grains from its wild rela-
tives. Safeguarding the identity of the domesticated
varieties under a cross-pollination system would
have become difficult or even impossible (Harlan
1995b).
A second advantage of self-pollination lies in the
genetic structure maintained within the crop. Selfing
results in splitting the crop’s gene pool into inbred
homozygous lines. Variation is thus structured in
the form of numerous true breeding cultivars. Since
these cultivars are automatically ‘fixed’ by the pol-
lination system, they can easily be maintained by
the farmer, even when planted side by side. In con-
trast, the preservation of varietal identity in cross-
pollinated plants is much more problematic. It
requires repeated selections towards the desired
norms, constant care to avoid the mixing of types,
and the prevention of pollination from undesirable
plants. It is therefore not surprising that early suc-
cesses in plant domestication involve selfers. In fact,
the ‘first wave’ domesticants in south-west Asia,
namely emmer wheat, einkorn wheat, barley, pea,
lentil, chickpea, bitter vetch, and flax, are all self-
pollinators. Cross-pollinated crops entered agricul-
ture later and comprise only a small minority among
the traditional grain crops. Rye and faba bean are
the only important cross-pollinated grain crops
indigenous to south-west Asia.
It is noteworthy that wheats, barley, and all other
Fertile Crescent ‘founder crops’, are not obligatory
selfers but predominantly self-pollinated plants in
which rare events of cross-pollination occur. Such
pollination systems are admirably suited for rapid
grain-crop evolution, because, being annual, they
produce new types recurrently. Occasional cross-
pollination provides the crop with genetic flexibil-
ity and serves to combine and reshuffle genes from
different sources. The numerous cycles of inbreed-
ing that follow the rare events of crossing lead to the
fixation of many new recombinant lines. The grower
can easily pick up the more attractive ones among
them.
CEREALS 21
Cereal cultivation comprises several essential
stages: sowing seeds in tilled fields; weeding; reap-
ing the mature spikes or panicles; threshing; win-
nowing; and storage.
The introduction of these practices automatically
initiated drastic changes in grasses taken into culti-
vation, setting them apart from their wild progeni-
tors (Harlan et al. 1973; de Wet 1975). Most
conspicuous is the selection for types in which the
mature dispersal units are retained on the mother
plant, and in which the wild-type adaptation for
seed dissemination has broken down (Zohary 1969).
In cereals, this implies a shift from shattering ears
or panicles (in wild forms), to non-shattering
types (in domesticated forms). In the wild forms,
survival depends on seed dispersal and the various
wild cereals have evolved elaborate seed dissemi-
nation devices. In wild wheats (pp. 34, 40) and wild
barley (p. 53), the seed-dispersal unit (diaspore)
comprises a single internode of the ear. Disar-
ticulation of the spike’s rachis is thus an essential
element of the wild-type seed dispersal. Plants in
wild populations are constantly selected for quick
shattering of their mature ears. In contrast, the
introduction of planting and harvesting brings
about automatic selection in exactly the opposite
direction. Under the new system, a sizeable propor-
tion of the seed produced by plants with brittle ears
will shatter and would not be included in the har-
vest. In contrast, practically all grains produced by
non-shattering mutants ‘wait on the stalks’ to be
reaped by the grower. Therefore, under cultivation,
non-shattering individuals have a much better
chance to contribute their seeds to the subsequent
generation. Consequently, non-brittle (or less brit-
tle) mutants (which were disadvantaged under
wild conditions) become highly successful under
the new human-controlled system. Thus, when
wild cereals are brought into cultivation, one should
expect selection for non-brittle forms whether or
not the cultivator is aware of this trait. Furthermore,
the incorporation of non-brittle mutants makes the
crop fully dependent on humans, as non-shattering
plants lose their seed-dispersal ability, and can no
longer survive under wild conditions.
Thus, a symbiotic relationship has been estab-
lished between humans and their crops (Rindos
1984).
22 DOMESTICATION OF PLANTS IN THE OLD WORLD
The appearance of non-shattering mutants in
cereal crops was probably a swift process. Genetic
considerations indicate that such a shift could have
been accomplished in the course of a few dozen
generations of selection (Zohary 1969; Hillman and
Davies 1999; Kislev 2002; Tanno and Willcox 2006a;
Fuller 2007). It is our view, however, that much
more research is needed in order to increase our
understanding of the shifts between wild and
domesticated populations, and the speed of their
development.
To date, the most effective (and easy to trace) diag-
nostic indication for domestication in archaeologi-
cal cereal remains is the rough disarticulation scars
in the internodes (Plates 2 and 3). This shift is con-
trolled mostly by a recessive mutation in a single
gene or by two such mutations (Table 7). As Kislev
(1989b) noted, some lower-ear spikelets of wild
cereals, up to 10% of them, do not disarticulate nat-
urally. This factor had to be taken into account while
discussing the domesticated status of a given
archaeobotanical assemblage.
Tanno and Willcox (in press) re-examined recently
the spikelet bases from eleven Turkish-Syrian sites
dated to the PPNA-PPNB period. They draw atten-
tion to the difference between upper and lower
scars, between wheat and barley, and especially
noted that only when the abscission scar is well pre-
served it can be use as a diagnostic tool.
A less reliable diagnostic trait is grain size. Seeds
in cultivars are usually plumper and larger than
those produced by their wild relatives. However,
the increase of grain size evolves slowly from
domestication onward. As Nesbit (2002) noted,
grain’s shape and size differences are clearly visible
in Pottery Neolithic find, but less clear in the Pre-
Pottery Neolithic. Recently, Gegas et al. (2010)
explored the genetic basis of the variation in wheat
grain size and shape. Their work demonstrates that
‘grain size has progressively increased through alter-
ations both in grain width and length, followed at
later stages by modifications in grain shape’.
Therefore, size increase in grains (under cultivation)
is not a fully reliable indication of the early stage of
domestication.
A third major outcome of introducing wild
cereals into the regime of cultivation is the break-
down of the wild mode of seed germination. This
breakdown is manifested in the reduction of germi-
nation inhibition from dormant to synchronous ger-
mination. Most wild grain crops, especially annuals
growing in Mediterranean-type (dry summer, wet
winter) or in semi-desert climates, depend for their
survival on the regulation of germination. A com-
mon adaptation is a delay of germination from the
time of seed maturation until the onset of favoura-
ble conditions in the next growing season. In
the Mediterranean basin this means germination
inhibition—even if the seed is wetted—from the
time of seed maturation in early summer to the start
of the rainy season in the following autumn.
Another adaptation is the delay of germination of
grains from the same dispersal unit over two or
more years. This protects the populations from the
crippling effects of dry or otherwise bad years. In
wild emmer wheat the dispersal unit contains two
kernels. One germinates in the ensuing autumn;
while in the second kernel, germination is fre-
quently delayed until the following year. Wild bar-
ley diaspores contain each only a single seed, but
not all dispersal units germinate in the first winter.
Under cultivation, this wild-type regulation of ger-
mination is no longer advantageous for the farmer,
and mutants in which germination inhibition has
broken down are selected automatically. This type
of selection has evidently been operative in the Old
World domesticated cereals. Most cultivars have
lost the germination-inhibition patterns which char-
acterized their wild progenitors, so that all their
seeds germinate at the same time whenever planted
by the farmer.
Several other traits that characterize domesti-
cated cereals (the ‘domestication syndrome’) seem
to be the outcome of conscious and/or unconscious
selection under domestication (for review see
Zohary 1969; Harlan et al. 1973). They include:
(i) selection towards erect plants, synchronous til-
lering, and uniform ripening;
(ii) increase of seed production by addition of fer-
tile florets and/or increase in the size of the
inflorescence or the number of ears or panicles
produced per individual plant;
(iii) decrease of awns, of glumes’ thickness, and
investment of grains (from hulled to naked
grains).
 CEREALS 23

Wheats: Triticum Chromosomally, wheats comprise a polyploid

Wheats are the universal cereals of Old World
Mediterranean-type agriculture. Together with bar-
ley, they constitute the principal grain stocks that
founded Neolithic agriculture, and the main ele-
ment responsible for its successful spread. Since this
early start, wheats have retained their crucial role in
Old World food production and have given rise to
numerous advanced forms. As of 2008, wheats and
rice rank second in the world’s grain production,
whereas maize rank first (http://faostat.fao.org/
site/339/default.aspx). Wheats are grown exten-
sively throughout the temperate, Mediterranean,
and subtropical parts of both hemispheres.1
Wheats are superior to most other cereals (e.g.
maize, rice, sorghum, or barley) in their nutritive
value. Their grains contain not only starch (60–80%
carbohydrate), but also significant amounts of pro-
tein (8–14%). The gluten proteins present in the
grain give wheat dough its stickiness, and its ability
to rise when leavened. In other words, the gluten
proteins bestow the dough with unique rising and
baking qualities. Wheats were, and still are, the pre-
ferred staple food of traditional farming communi-
ties throughout the Old World. Thus, it is not
surprising that in numerous cultures, the idea of
food has been equated with bread.
Several distinct wild species of the genus Triticum
L. were introduced into domestication. Most mod-
ern wheat cultivars belong to two biological species:
(i) bread wheat, Triticum aestivum (2n = 6x = 42 chro-
mosomes), valued for the baking of high rising
bread; and (ii) hard or durum-type wheat, T. turgi-
dum, (2n = 4x = 28 chromosomes), used for the
preparation of pasta and low-rising bread. Other
Triticum species, as well as more primitive cultivars
of the above-mentioned two species, were impor-
tant in the past and survive today only as relic crops.
All wheats are almost fully self-pollinated.
Therefore, variation in these cereals is moulded in
the form of numerous inbred lines.
series (for review of wheat cytogenetics, see Riley
1965; Sears 1969; Miller 1987; Feldman et al. 1995).
Some domesticated forms have a diploid chromo-
some number (2n = 2x = 14 chromosomes) and con-
tain two sets of a single genome (designated AA).
Other wheats are tetraploid (2n = 4x = 28) and com-
bine two distinct genomes (either BBAA or GGAA).
Others, are hexaploid (2n = 6x = 42) and contain
three different genomes (BBAADD). Domestic
wheats fall into five chromosomal groups: two dip-
loid wheats, two tetraploid wheats, and a single
hexaploid wheat. Forms within each group are inter-
fertile and share the same chromosome constitution.
In contrast, hybrids between groups are highly ster-
ile. The taxonomic classification of domesticated
wheats and their closely related wild types (Van
Slageren 1994) is based on these cytogenetic criteria.
The following five biological species (Table 3) are
recognized today in the genus Triticum:
1. Diploid T. monococcum L. or einkorn wheat
(genomic designation AmAm) comprises both wild
and domesticated forms (Fig. 3). Domesticated
einkorn with its characteristic hulled grains was an
important grain crop in the past. It survives today
only as a relic crop.
2. Diploid T. urartu Tuman. ex Gandiljan.
(genomic designation AA). This diploid wheat com-
prises only wild forms and is largely confined to the
Fertile Crescent belt and extends to Armenia
(Waines 1996; Valkoun et al. 1998). Apparently, it
contributed its diploid set of chromosomes (long
before domestication) to the 4x hard wheat, T. turgi-
dum, and the 6x bread wheat, T. aestivum (Dvořák et
al. 1998; Dvořák et al. 1993, 1988). Morphologically,
T. urartu closely resembles the wild forms of T.
monococcum. However, crossing experiments indi-
cate that T. urartu is reproductively isolated from
both wild and domesticated einkorn wheats. Inter-
specific F1 hybrids between these wheats are male-
sterile and do not set seeds upon self-pollination

1
Excluding the closely related Aegilops L., which comprises some twenty-two species. Some Aegilops species show close
cytogenetic links with the wheats. Indeed, diploid Aegilops species participated in the formation of the polyploid Triticum
species. For this reason several researchers (see, for example, Sears 1969; Morris and Sears 1967) lump together both gen-
era, and include the Aegilops species within Triticum. Others (e.g. van Slageren 1994; Zohary 1965) keep the traditional
classification, but consider Triticum and Aegilops as a single natural group (the ‘wheat group’).
24 DOMESTICATION OF PLANTS IN THE OLD WORLD
(Waines and Barnhart 1992; Waines 1996; Valkoun
et al. 1998). Also isozyme and DNA tests (Jaaska 1997b)
indicate that these two diploid wheats are distinct.
3. Tetraploid T. turgidum L. (genomic designation
BBAA) comprises (see Table 3) wild emmer wheat,
domesticated emmer wheat, durum wheat, and
several other domesticated tetraploid forms (Figs 4
and 5, and Plates 2–4). Cytogenetic and molecular
tests have shown that the genome A of tetraploid T.
turgidum is closely related to the diploid genome
present in T. urartu (Dvořák et al. 1993, 1988; Jaaska
1997b). In contrast, the genome of diploid T. mono-
coccum is more remote. Consequently, the classical
explanation of the polyploid origin of tetraploid T.
turgidum had to be revised. Now it is clear that a T.
urartu-like diploid wheat was involved in the poly-
ploid origin (in the wild) of tetraploid T. turgidum
subsp. dicoccoides; while diploid wild T. monococ-
cum—long considered as the donor of the A genome
to wild emmer wheat—had little to do with its poly-
ploid origin. From the start of agriculture, hulled
(= glumed) emmer wheat emerges as the principal
stock of domesticated wheat. It gave rise to the wide
range of present-day, free-threshing, tetraploid
durum-type wheats. By hybridization with a wild
Aegilops species, domesticated tetraploid T. turgi-
dum formed an additional species: the hexaploid
bread wheat (see below).
4. Tetraploid T. timopheevi Zhuk. (genomic desig-
nation GGAA) includes both wild and domesti-
cated hulled forms. Domesticated Timopheev’s
wheat is endemic to a small area in Georgia, and
seems to represent only a local episode in wheat-
crop evolution. The A genome in T. timopheevi is
most similar to the genome present in diploid
T. urartu indicating that an urartu-like diploid ances-
tor was probably involved in the polyploid forma-
tion (in the wild) of timopheevi’s wheat.
5. Hexaploid T. aestivum L. or bread wheat (genomic
designation BBAADD) originated under cultivation by
the addition of the DD chromosome complement of
the wild grass Aegilops tauschii Coss. [= Ae. squarrosa
L.] to the domesticated tetraploid BBAA turgidum
wheats. The extraordinarily variable T. aestivum group
constitutes the most important wheat crop of today
(Fig. 6). It comprises several primitive, hulled, spelt-
type wheats, and numerous free-threshing forms
(including modern bread wheat).
Traditional wheat classification has suffered from
excessive splitting. Formally, wheat taxonomists
regarded every main morphological type in the
domesticated wheats as an independent species and
recognized more than a dozen distinct species in the
genus Triticum (for details, see the treatments of
Percival 1921; Schiemann 1948; Dorofeev et al. 1979).
However, in view of the accumulated information
on genetic affinities between types, such species
delimitation is no longer justified. Today, wheats are
usually grouped in five biological species. Table 3
(p. 29) lists the various traditional species names in
Triticum together with the modern grouping of the
wheats. Table 4 presents the wild stocks from which
the principal domesticated cereals were derived and
summarizes the main evolutionary events that led
to the formation of the crops.
Domesticated wheats fall into several distinct
classes according to their response to threshing:
1. The more primitive forms, i.e. diploid einkorn,
tetraploid emmer, and hexaploid spelt have hulled
grains. Their grains are enclosed in the spikelet by
toughened glumes that do not break during thresh-
ing. In hulled wheats, the products of threshing are
spikelets, not grains (Fig. 7). More advanced domes-
ticated wheats, i.e. tetraploid durum-type and hex-
aploid bread wheats, are free-threshing. Their glumes
are thinner and do not invest the grains tightly.
Threshing releases the naked kernels (e.g. Fig. 5, 6).
Because of this difference in threshing product, the
handling of hulled wheats by the farmer is different
from that of the free-threshing ones. In the first, the
spikelets are stored and marketed as such, and the
grains have to be freed (usually by pounding) before
they can be used. The utilization of naked wheats is
simpler. After threshing, the free grains are win-
nowed, sieved, and then stored ready for milling.
Because of the different appearance of the marketed
products, hulled and free-threshing wheats were
often regarded in antiquity as different cereals and
they were even called different names. Yet, one has
to bear in mind that hulled and naked wheat forms
can belong to the same biological species (Table 3).
In bread wheat, T. aestivum, the difference between
hulled and free-threshing varieties is governed
mainly by a single mutation (the q gene). In con-
trast, in most free-threshing, T. turgidum wheats, the
 CEREALS 25

A D
upper B lower

C F

Fig. 3 Diploid einkorn wheat, Triticum monococcum. Left: A–ear (1:1), B–shattering spikelet (2:1), upper–upper disarticulation scar, lower–lower
disarticulation scar, and C–grain (3:1) of wild einkorn, T. monococcum subsp. baeoticum. Right: D–ear (1:1), E–non-shattering spikelet (2:1), and
F–grain (3:1) of domesticated einkorn, T. monococcum subsp. monococcum (Schiemann 1948).
26 DOMESTICATION OF PLANTS IN THE OLD WORLD

C E

Fig. 4 Tetraploid hulled emmer wheats, Triticum turgidum. Left: A–ear (1:1), B–spikelet (2:1), and C–grain (3:1) of wild emmer wheat, T. turgidum
subsp dicoccoides. Right: D–ear (1:1), and E–grain (3:1) of domesticated emmer wheat, T. turgidum subsp. dicoccum (Schiemann 1948). Note the
smooth disarticulation scars in B, both below and above the internode, as in Fig. 3B.
 CEREALS 27

A
C

B D

Fig. 5 Tetraploid free-threshing emmer wheats, Triticum turgidum. Left: A–ear (1:1) and B–grain (3:1) of free-threshing durum wheat, T. turgidum
subsp. durum. Right: C–ear (1:1), and D–grain (3:1) of free-threshing rivet wheat, T. turgidum subsp. turgidum (Schiemann 1948).
28 DOMESTICATION OF PLANTS IN THE OLD WORLD

A
B C D

Fig. 6 Hexaploid bread wheats, Triticum aestivum. A–Ear and grain of spelt wheat, T. aestivum subsp. spelta. B–Ear and grain of club wheat, T.
aestivum subsp. compactum. C and D–Ears and grains of awned and awnless varieties of bread wheat, T. aestivum subsp. aestivum. Ears 1:1;
grains 3:1 (Schiemann 1948).
 CEREALS 29

Table 3 Taxonomic classification of wheats, Triticum L. Morphological types (or species) according to traditional classification and their modern
grouping on the basis of cytogenetic and molecular affinities (Van Slageren 1994). Dashed lines separate between ploidy levels.

Modern grouping (biological species) Traditional classification

(i) Diploid (2n = 14) einkorn wheat

Genomic constitution: AA
Both wild and domesticated forms.
Collective name: T. monococcum L
1. T. monococcum L. subsp. aegilopoides (Link) Thell. 1. Wild einkorn T. baeoticum Boiss. emend. Schiem. (brittle, hulled).
Including single-grain forms (subsp. aegilopoides) and two-grain forms (subsp.
thaoudar)
2. T. monococcum L. subsp. monococcum 2. Domesticated einkorn T. monococcum L. (non-brittle, hulled)
(ii) Diploid (2n = 14). Genomic constitution: AA Only wild forms
1. T. urartu Tuman ex Gand. 1. Wild T. urartu Tuman. (brittle, hulled)

(iii) Tetraploid (2n = 28) wild and domesticated emmer wheats,

durum-type wheats, etc.
Genomic constitution: BBAA
Both wild and domesticated forms
Collective name: T. turgidum L.
1. T. turgidum ssp. dicoccoides (Körn. ex Asch. & Graebn.) Thell.
2. T. turgidum L. ssp. dicoccum (Schrank) Thell.*
3. T. turgidum L. ssp. durum (Schrank) Thell.
4. T. turgidum L. ssp. turgidum
5. T. turgidum L. ssp. polonicum (L.) Thell.
6. T. turgidum L. ssp. carthlicum (Nevski) Löve & Löve
7. T. turgidum L. ssp. parvicoccum Kislev
(iv) Tetraploid (2n = 28) Timopheev’s wheat
Genomic constitution: GGAA
Both wild and domesticated forms
Collective name: T. timopheevii Zhuk.
1. T. timopheevii Zhuk. ssp. armeniacum (Jakubz.) van Slageren
2. T. timopheevii Zhuk. ssp. timopheevii
1. Wild emmer T. dicoccoides (Körn. ex. Aschers. & Graebner) Schweinf. (brittle,
hulled)
2. Domesticated emmer T. dicoccum Schübl. (non-brittle, hulled)
3. Macaroni or hard wheat T. durum Desf. (domesticated, free-threshing)
4. Rivet wheat T. turgidum L. (domesticated, free-threshing)
5. Polish wheat T. polonicum L. (domesticated, free-threshing)
6. T. carthlicum Nevski [= T. persicum Vav.] (domesticated, free-threshing)
7. T. parvicoccum Kislev small grained archaeobotanical forms (domesticated,
free-threshing)
1. Wild Timopheev’s wheat T. araraticum Jakubz. (brittle, hulled)
2. Domesticated Timopheev’s wheat T. timopheevii Zhuk. (non-brittle, hulled)
(the uncertain ‘new’ glume wheat might belong to this group)

(v) Hexaploid (2n = 42) bread wheat

Genomic constitution: BBAADD (the D genome contributor is
Aegilops tauschii)
Only domesticated forms
Collective name: T. aestivum L.
1. T. aestivum L. ssp. spelta (L.) Thell. 1. Spelt T. spelta L. (non-brittle, hulled)
2. T. aestivum L. ssp. macha (Dek. & Men.) MK 2. T. macha Dekr. & Men. (non-brittle, hulled)
3. - 3. T. vavilovii Tuman. (non-brittle, hulled)
4. T. aestivum L. ssp. aestivum 4. Bread wheat T. aestivum L. [= T. vulgare Host; T. sativum Lam.] (free-threshing)
5. T. aestivum L. ssp. compactum (Host) MK 5. Club wheat T. compactum Host. [= T. aestivo-compactum Schiem.]
(free-threshing)
6. T. aestivum L. ssp. sphaerococcum (Percival) Mk 6. Indian dwarf wheat T. sphaerococcum Perc. (free-threshing)
(vi) Hexaploid (2n = 42) Zhukovsky’s wheat
Genomic constitution: GGAAAA
Only domesticated forms
1. Triticum zhukovskyi Men. & Er. (non-brittle, hulled)

Note: hulled wheat = glume wheat. * In this book, we use ‘dicoccum’ rather than Van Slageren’s ‘dicoccon’, as many taxonomists believe this is the cor-
rect
30 DOMESTICATION OF PLANTS IN THE OLD WORLD
shift from ‘hulledness’ to ‘nakedness’ was brought
about by a polygenic system.
2. Another important morphological trait in
wheats is the manner in which the ear shatters in
wild forms, or stays intact in the domesticated
forms. Wild wheats are adapted to disseminate their
grains by having brittle ears that disarticulate at
maturity into individual spikelets (dispersal units).
In wild einkorn (Fig. 3), wild emmer (Fig. 4), and
wild Timopheevi’s wheat, the point of disarticulation
is the upper and lower abscission scars (Fig. 3, 4 and
7). Each spikelet with a wedge-shaped rachis inter-
node at its base constitutes an arrow-like device
that inserts the grain into the ground (Zohary and
Brick 1961; Zohary 1969; Harlan et al. 1973). In
Aegilops tauschii, and consequently in spelt wheat,
the disarticulation point is frequently below the
upper abscission scar. The dispersal unit is a ‘barrel-
shaped’ spikelet with a characteristic rachis inter-
node joined to its upper side (Figs 7C and 14, below
and p. 49). In contrast to wild types, all domesti-
cated wheats have non-brittle ears that stay intact
after maturation. Thus, they depend on the farmer
for their reaping, threshing, and sowing. Yet, thresh-
ing fractures the ears of the various domestic, non-
brittle wheats in different ways, and significantly
these morphological differences are also discernible
in archaeobotanical remains.
Fig. 7 The threshing products of the three main types of cultivated hulled wheats: A–Einkorn, Triticum monococcum subsp. monococcum.
B–Emmer, T. turgidum subsp. dicoccum. C–Spelt, T. aestivum subsp. spelta (fresh material). Note that in A and B the rachis internode is attached
below the dispersal unit, while in C it is attached above it.
The following are indications for discerning between
domesticated and wild material wheats in archaeo-
botanical remains (see Table 3):
(i) Usually, the ears of hulled diploid and tetraploid
domesticated wheats (einkorn, emmer, and
Timopheevi’s wheats) break during threshing at the
same place at which their wild counterparts disar-
ticulate spontaneously. In other words, threshing in
these still primitive wheats ‘mimics’ the shattering
pattern of their wild progenitors. The individual
spikelet with the internode at its base is the product
of threshing (Fig. 7, A and B, Plate 3). However,
instead of the smooth abscission scars, which char-
acterize the wild forms, the surface of the breakage
scars in the domestic hulled wheats is rough.
Therefore, prevalence of spikelet forks showing
rough scars (Plate 3, Fig. 8) in wheat archaeobotani-
cal assemblages, serves as a critical indication of
domestication. This criterion fits most other grain
crops as well.
(ii) The breakage point at each spikelet of hulled
hexaploid spelt-type wheats is endowed by the
wild DD contributor. In spelt wheat, this diagnostic
pressure-fracture breaks the internode below the dis-
articulation scar. Therefore, the rachis internode
remains attached to the upper side of the spikelet
(Fig. 7C). In archaeobotanical assemblages, spike-
 CEREALS 31

lets which follow the disarticulation pattern of Ae.
tauschii indicate the presence of hexaploid spelt-
type wheats, though this identification must be con-
firmed using glume shape and texture.
(iii) In free-threshing domesticated wheats (tetra-
ploid durum-type and hexaploid bread wheats) the
rachis of the ear is thickened throughout its length.
Threshing breaks the glumes, lemmas, and paleas
at or near their base, and releases the naked grains.
The thickened rachis breaks irregularly, frequently
into segments of two to five internodes (Fig. 9).
In the hulled wheats, burning frequently destroys
the upper parts of the investing glumes and lemma/
palea, exposing and freeing the grains. In contrast,
the ‘spikelet forks’ (rachis internode and glume
bases attached above it, Fig. 8), or its separated
glume bases, often survive charring as discrete
units. When found in archaeobotanical assem-
blages, such charred units are telltale indicators for
the recognition of hulled wheats.
In free-threshing wheats, the glumes and pales
are thin and papery, and frequently do not survive
burning. Also, the glumes, pales and most of the
rachis fragments are winnowed out as chaff. As a
result, they are rare in archaeobotanical assem-
blages. In contrast, some of the fragments of the
tough rachis (particularly the more basal parts) are

Fig. 8 Charred remains of domesticated einkorn wheat retrieved from a Michelsberger culture grave at Regensburg, Germany. (A): spikelet forks,
showing the rough scars typical of domesticated forms. (B): grains from the same hoard (Photographs kindly provided by D. Kučan).

0 5 mm

Fig. 9A Rachis segments, the diagnostic traits for the recognition of free-threshing wheats in archaeological remains. Carbonized rachis
fragments of 4X free-threshing wheats, Neolithic Tell Ramad, Syria (van Zeist 1976).
32 DOMESTICATION OF PLANTS IN THE OLD WORLD

Free-threshing forms of Free-threshing forms of

4× Triticum turgidum 6× Triticum aestivum

1
1
2
2

3
3

4 4

5
5

5 5

Fig. 9B The structure of the rachis fragment in tetraploid and in hexaploid free-threshing wheats, pointing out five diagnostic traits for
distinguishing between 4X Triticum turgidum cultivars (excluding carthlicum forms) and 6X T. aestivum cultivars (Hillman 2001, and pers. comm.).

Diagnostic traits Tetraploid free-threshing wheats Hexaploid free-threshing wheats

1. Lower part of the glumes
2. Shape of the glume base
3. Shape of the rachis internode
4+5. Structure of the edge of the internode
Usually retained after threshing, including the
lower part of the keel
A prominent lump is present beneath each
glume base
The sides of the internodes are more or less
straight, and converge gradually downwards
The internode is relatively thick, smooth, with
rounded edge
Deciduous and fully removed by threshing
The lump is missing (or is only very
rudimentarily developed)
The sides of the internodes are curved
The edge is thinner, grooved by a fine
longitudinal furrow, with ridges frequently
supporting short hairs

similar to the kernels in their size and weight, and
upon winnowing they occasionally stay with the
grains. Such fragments usually char well and
become critical markers (Fig. 9A) for the presence of
free-threshing wheats in archaeological excavations
(van Zeist 1976; Hillman 1984).
Such rachis segments can also be used for distin-
guishing between tetraploid and hexaploid free-
threshing wheats (Maier 1996; Hillman et al. 1996;
Jacomet 2006). The diagnostic traits separating the two
chromosomal levels are given in some detail in Fig. 9B
(above). Finds of naked forms, belonging to both T.
turgidum and T. aestivum, have been lumped together
in archaeobotanical reports prior to the 1990s either as
T. turgidum-T. aestivum or as ‘aestivo-compactum’.
However, several reports have appeared, in which the
4x turgidum and the 6x aestivum free-threshing remains
have been soundly identified (for review see Maier
1996). In some naked wheat samples obtained from
Neolithic lake dwellings in central Europe, such mor-
phological identification verified the results of ancient
DNA analysis (e.g. Schlumbaum et al. 1998).
Table 4 The wild progenitors of the main domesticated wheats and their principal derivatives under domestication
Wild einkorn wheat (2x) Wild emmer wheat (4x) T. turgidum subsp.
T. monococcum subsp. aegilopoides, diploid (AA), dicoccoides, tetraploid (BBAA), brittle ears, hulled
brittle ears, hulled grain. grain.
domestication domestication
Domesticated einkorn wheat Hard wheats
T. monococcum subsp. monococcum, diploid (AA), Domesticated T. turgidum, tetraploid (BBAA), non-
non-brittle ears, hulled. brittle ears.
Widely cultivated in the past; relic today. 1st stage: hulled emmer wheat T. turgidum subsp.
dicoccum (widely distributed in the past;
relic today).
2nd stage: free-threshing forms, mostly durum wheat.
Derived spontaneously from T. turgidum subsp.
dicoccum (common in Mediterranean-type climates).
Wild Aegilops tauschii, diploid (DD), brittle ears,
hulled grains.
Not domesticated, but contributed its genome to
hexaploid wheats.
addition of the D genome through
spontaneous hybridization and
chromosome doubling
Spelt and bread wheats (6x)
Domestiated T. aestivum, hexaploid (BBAADD),
non-brittle.
1st stage: hulled spelt-type forms (relic today).
2nd stage: free-threshing bread wheat.
Derived spontaneously from spelt-type (the most
common and most variable wheat crop today).
CEREALS
33
34 DOMESTICATION OF PLANTS IN THE OLD WORLD
Einkorn wheat: Triticum monococcum
Einkorn wheat is a diploid (2n = 2x = 14 chromo-
somes), a relatively uniform annual grain crop, with
characteristic hulled grains, and delicate ears and
spikelets (Fig. 3). A single non-shattering domesti-
cated form of this wheat (usually referred to as T.
sinskajae) is apparently a free-threshing mutant.
Most domesticated einkorn varieties produce one
grain per spikelet, hence its name, but cultivars
with two grains exist as well (Fig. 8B). Today einkorn
is a relic crop, although it is still sporadically grown
in western Turkey, the Balkan countries, Germany,
Switzerland, Spain, as well as Caucasia (Nesbitt
and Samuel 1996a). In the past, einkorn cultivation
was much more extensive. This wheat was one of
the founder grain crops of Neolithic agriculture in
the Fertile Crescent and a principal component of
the early crop assortment in Europe, especially in
the Aegean region. Since the Bronze Age, its impor-
tance seems to have declined gradually, very likely
due to the competition from free-threshing wheats.
Einkorn is a small plant, rarely more than 70 cm
high, with a relatively low yield, but it can survive
on poor soils where other wheat types fail. The fine
yellow flour is nutritious, but gives bread of poor
rising qualities. Thus einkorn has been consumed
primarily as porridge or as cooked whole grains.
Since Roman times, a considerable part of the yield
has been fed to animals (Schiemann 1948; Harlan
1981).
Wild ancestry
The wild ancestry of domesticated einkorn wheat is
well established. Domesticated Triticum monococ-
cum subsp. monococcum is closely related to a group
of wild and weedy wheat forms spread over south-
west Asia and adjacent territories, and traditionally
referred to as wild einkorn or T. baeoticum Boiss.
emend. Schiem. (Table 3). Both wild and domesti-
cated einkorns are morphologically similar (Fig. 3).
Both are diploid and contain identical chromosomes
(genomic designation AmAm). Hybrids between
wild aegilopoides and domesticated monococcum are
fully fertile and chromosome pairing in their meio-
sis is normal. The principle distinguishing trait
between wild and domesticated einkorn is the mode
of seed dispersal. Wild forms have brittle ears and
the individual spikelets disarticulate at maturity to
disperse the seed. In domesticated einkorn this
essential adaptation to wild conditions no longer
exists. The mature ear remains intact and breaks
into individual spikelets only upon pressure (dur-
ing threshing and/or flailing). Seed dispersal
depends on reaping and sowing by humans.
Another diagnostic character indicating domestica-
tion is the shape of the grain (van Zeist 1976). In
domesticated forms, the kernels tend to be wider
compared to the wild forms (Fig. 3, p. 25), but this
sign is less reliable (Gegas et al. 2010) (see p. 22).
Most specialists today regard the aegilopoides
wheat not as an independent species but as the wild
progenitor of the domesticated crop. It is taxonomi-
cally placed as a subspecies within the crop com-
plex as T. monococcum L. subsp. aegilopoides (Link)
Thell. [= subsp. baeoticum (Boiss.) A. et D. Löve].
Wild einkorn is widely distributed over western
Asia and can be found also in the southern Balkan
countries (Harlan and Zohary 1966; Zohary 1969).
Its distribution centre lies in the central part of the
Fertile Crescent (i.e. northern Syria, southern
Turkey, northern Iraq and adjacent Iran, as well as
some parts of western Anatolia) (Map 3). In these
areas, wild einkorn is widely distributed as a com-
ponent of oak park-forests and steppe-like forma-
tions. In addition to occupying such primary
habitats, wild einkorn also grows as a weed and
colonizer of secondary habitats, such as edges of
cultivation and roadsides. Sometimes it also
invades fields of domesticated cereals. Aegilopoides
wheats are distributed over a wide ecological range
with respect to both soils and climates. Edaphically,
wild einkorn shows a definite affinity to basaltic
soils, marls, clays, and limestones. Climatically, it
thrives in the summer-dry foothills of the northern
Euphrates basin, as well as on the bitterly cold,
elevated plateau of central and eastern Anatolia
(1400–2000 m altitude), with its summer rains.
However, it does not succeed in hot and very arid
climates. Further away from its distribution centre,
wild einkorn is restricted mainly to segetal or sec-
ondary habitats, i.e. sites that were probably not
available before the opening up of these areas to
agricultural activity.
Several major eco-geographical and morphologi-
cal forms can be recognized in wild einkorn. In the

0 100 200 miles
0 200 400 km
Map 3 Geographical distribution of wild einkorn wheat, Triticum monococcum subsp. baeoticum (= T. baeoticum). The area in which wild einkorn
is widely distributed is shaded. Dots represent additional sites outside the main distribution area, harboring mainly weedy forms (based on Zohary
1989a).
north and north-west parts of its range, plants with
small, single-awned, one-seeded spikelets prevail.
These are sometimes named T. aegilopoides (Link)
Bal. In the summer-dry southern areas, more robust
plants with two-grained, two-awned spikelets,
sometimes referred to as T. thaoudar Reuter, are
common. But in central Anatolia, Transcaucasia,
and adjacent territories of Iran, a series of interme-
diate forms, bridging aegilopoides and thaoudar forms
abound. In fact, many Anatolian populations of
wild einkorn show a wide range of variation in
spikelet morphology and include single-grained
individuals (with a single awn), two-grained plants
(with two well-developed awns) and various
intergradations between the two extreme types.
Heun et al. (1997) examined sixty-eight represent-
ative lines of domesticated einkorn Triticum mono-
coccum subsp. monococcum and 268 lines of wild
einkorn T. monococcum subsp. aegilopoides for ampli-
CEREALS 35
fied fragment length polymorphism (AFLP) DNA
analysis. The latter originated from the ‘Fertile
Crescent’ (194 samples) and from several other parts
of the distribution area of this wild wheat (74 sam-
ples). Among the wild einkorn lines tested, a group
of eleven lines (which came from Karacadağ range
in south-east Turkey) turned out to be distinctively
separated from all other wild einkorn lines. These
were genetically most similar to the tested domesti-
cated einkorn lines, indicating that they belong to
the wild source from which the crop could have
evolved. The data obtained support the assumption
of a monophyletic origin of domesticated einkorn
wheat. Recently, Kilian et al. (2007) conducted a
much larger study of genetic markers of 321 wild
lines and 92 domesticated lines of einkorn. This
study has found that three wild einkorn races arose
prior to domestication without human intervention,
and only one of them became domesticated.
36 DOMESTICATION OF PLANTS IN THE OLD WORLD
According to this study, this race (which was col-
lected from two regions in Turkey, Karacadağ and
the Kartal-Karadağ range) was subjected to multiple
independent domestication events, possibly after
the wild grains from these two areas in south-east
Turkey were dispersed throughout a wider region
by the earliest cultivators.
The Fertile Crescent belt also harbours T. urartu
Tuman. (see above), a diploid wild wheat which
resembles closely, and is easily confused with, the
two-grained forms of wild einkorn (Valkoun et al.
1998). However, T. urartu can be recognized by its
spreading awns, the presence of a small third awn
at the tip of the spikelet, the lack of hairs on the
basal leaves, and by the colour of its grains (Waines
and Barnhart 1992; Waines 1996). It shows a clear
preference for basaltic soils, and frequently forms
mixed stands with wild einkorn wheat.
Archaeological evidence
Einkorn wheat was probably collected extensively
from the wild before its introduction to cultiva-
tion. The earliest reported einkorn finds are the
Fig. 10 Carbonized grains of wild einkorn wheat, Triticum monococcum subsp. baeoticum, and wild rye, Secale sp. Epi-Palaeolithic Mureybit, Syria
(van Zeist and Casparie 1968). Note the elongated shape of the kernels, which characterizes wild forms.
charred, narrow, wild-type kernels (Fig. 10), from
Epi-Palaeolithic pre-agriculture layers, ca. 12,700–
11,100 cal BP, Tell Abu Hureyra (Hillman 1975,
2000a; Hillman et al. 1989) and ca. 11,800–11,300 cal
BP in Mureybit (van Zeist and Bakker-Heeres 1986;
van Zeist and Casparie 1968), northern Syria.
Aegilopoides-type einkorn remains continue to
appear in some Early PPNB (Pre-Pottery Neolithic
B), 10,500–10,100 cal BP, settlements in south-west
Asia where there are already definite signs of
wheat and barley domestication; i.e. in Tell Abu
Hureyra, Syria (Hillman 1975, 2000b; de Moulins
1997, 2000; Hillman et al. 1989), ca. 10,250–9,550 cal
BP Çayönü (van Zeist and de Roller 1991–2, 2003;
van Zeist 1972) and 9,450–8,450 cal BP Can Hasan
III (Hillman 1972), Turkey, and (ca. 10,200–10,150
cal BP) Ali Kosh, and also in later, ca. 8,350–7,750
cal BP Tepe Sabz, Iran (Helbaek 1969). In some of
these sites (as well as other contemporary south-
west Asian settlements) one is faced also with
plumper kernels characteristic of domesticated
einkorn (Fig. 11).
The first definite domesticated einkorn wheat
appears in two Early PPNB sites, namely Çayönü
 CEREALS 37

A 0 5 mm B

Fig. 11 Comparison between charred grains of wild and of domesticated einkorn wheats, Triticum monococcum. A–Wild einkorn, T. monococcum
subsp. baeoticum, from pre-agriculture Mureybit, Syria (van Zeist and Casparie 1968). B–Domesticated einkorn, T. monococcum subsp.
monococcum, from early Neolithic Nea Nikomedeia, Greece (van Zeist and Bottema 1971). The upper and middle domesticated grains are from
one-seeded spikelets (‘spindle shaped’), while the lower grain is from two-seeded spikelet. The grains of two-seeded einkorn spikelets are not
curved in ventral side, but straight and flat, and therefore look similar to those of domesticated emmer (Fig. 12). They can be distinguished by the
fact that they are slenderer, still somewhat spindle shaped in dorsal view, and have a pointed apex.

(van Zeist 1972; van Zeist and de Roller 1991–2,
2003) and Cafer Höyük (de Moulins 1997), southern
Turkey (Map 1). (A third possible site, Nevali Çori
(Pasternak 1998), has been mooted. We await final
confirmation. Tanno and Willcox (in press) re-exam-
ined its domestication status and concluded that
there is no evidence of einkorn domestication.) In
these sites, numerous spikelet forks were found,
showing rough breakage scars—that is, the most
reliable sign of domestication. These finds indicate
that einkorn belongs to the small group of annual
grain plants that founded agriculture in the Fertile
Crescent. It seems, then, that around 10,500–10,100
cal BP domesticated einkorn appeared first in the
Euphrates Valley.
From these localions, which are situated more or
less within the present range of distribution of wild
einkorn (Map 3), this domesticated cereal spreads
further south to Middle PPNB, ca. 10,200–9,550 cal
BP, Tell el Aswad (van Zeist and Bakker-Heeres
1985) near Damascus, and to ca. 9,900–9,550 cal BP
Jericho (Hopf 1983). In both these Early Neolithic
sites, remains of einkorn are, however, relatively
few in number. This wheat appears to have been
less frequently cultivated than the two other
Neolithic founder cereals, namely emmer wheat
and barley (Map 1).
The domesticated status of einkorn wheat from
ca. 10,650–9,550 cal BP Early PPNB Kissonerga-
Mylouthkia and Shillourokambos, Cyprus (Willcox
38 DOMESTICATION OF PLANTS IN THE OLD WORLD
2000; Murray 2003) relies currently on grain shape,
and therefore needs further confirmation by chaff
remain. However, as no wild wheats grow in the
island, it seems that this find represents early intro-
duction of agriculture from elsewhere.
Somewhat later, einkorn emerges as one of the
principal crops in established Neolithic food produc-
tion in south-west Asia. It re-appears together with
emmer wheat and barley but shows a definite prefer-
ence for areas with relatively cool climates. It is rare
in warmer places and totally absent from hot regions
such as Egypt and Lower Mesopotamia; in the Israel-
Jordan region, it has been found only once—in
Middle PPNB Jericho (Hopf 1983). Einkorn remains
continue to be found frequently in Chalcolithic,
Bronze Age, and Iron Age sites in south-west Asia.
Along with emmer (pp. 44–48), it is gradually
replaced by the free-threshing wheats.
Einkorn wheat played an important role in the
early spread of Neolithic agriculture beyond its
Fertile Crescent ‘core area’. Charred remains of
domesticated einkorn are numerous in the first half
of the ninth millennium BP contexts of the Aegean
Belt. Such sites are ca. 8,700–6,800 cal BP Cape
Andreas-Kastros, Cyprus (van Zeist 1981), ca.
8,650–8,400 cal BP, Knossos, Crete (Sarpaki 2009),
and ca. 8,650–8,200 cal BP, Sesklo, Greece (Hopf,
1962; Kroll, 1981a). It was also found in several early
Greek and Macedonian agricultural settlements
which developed during the second half of the
ninth millennium BP (Renfrew 1979; Kroll 1981a),
such as ca. 8,200–6,650 cal BP, Anza, Macedonia
(Renfrew 1979). Recently, Valamoti (2011) concluded
that einkorn dominates many Neolithic assem-
blages in Greece. These settlements extended as far
as ca. 8,150–7,000 cal BP Obre, Bosnia-Hercegovina,
former Yugoslavia (Renfrew 1974). At the same
time, its reach was as far east as ca. 8,200–7,850 cal
BP Jeitun, south Turkmenistan, and served there as
the principal staple (Charles and Hillman 1992;
Harris and Gosden 1996).
Much richer remains of einkorn wheat are avail-
able from sites representing earliest agriculture in
the first half of the eighth millennium BP in Bulgaria,
like ca. 7,950–7,650 cal BP Early Neolithic I Kapitan
Dimitrievo (Marinova 2006, forthcoming), and
contemporaneous Karanovo (Thanheiser 1997;
Marinova 2004, 2006) and Kovacevo (Popova 1992;
Marinova 2006). In these sites, einkorn wheat con-
tinue to be represented in large quantities up until
the Bronze Age. Here (and see details in Chapter 10
and Map 1, p. 2), einkorn often prevails over emmer
in frequency of pure hoards (see Thanheiser 1997;
Marinova 2004, 2006, forthcoming); and retains its
principal role throughout the Neolithic and Bronze
Age (Hopf 1973a; Renfrew 1979). At the same time
einkorn reaches ca. 7,950–7,250 cal BP Starčevo-
Körös culture sites in southern Hungary (Hartyányi
and Nováki 1971; Hartyányi et al. 1968; Füzes 1990),
as well as ca. 7,600–7,500 cal BP Sacarovca, Moldavia
(Januševič 1984; Kuzminova et al. 1998; Monah
2007b).
Einkorn is also one of the main cereal crops of sec-
ond half of the eighth millennium Linearbandkeramik
culture; i.e. the first farming settlements in central
Europe (Willerding 1980; Nesbitt and Samuel 1996a).
During this short period, farming spread across cen-
tral and northern Europe to Spain, as well as to
Egypt (where no einkorn was found). Einkorn usu-
ally occurs in a mixture with emmer wheat or some-
times as a pure crop. In some locations both cereals
are equally common, but occasionally einkorn pre-
vails. This situation persists all over Neolithic cen-
tral and northern Europe, although barley becomes
more frequent, and partly replaces the wheats in the
later sites. Similar dominance of einkorn and emmer
is found in eastern Europe, but the relative propor-
tions of the two wheats vary considerably according
to local conditions and cultures. The numerous data
already available indicate that wheat cultivation in
the Neolithic and Bronze Age of central Europe
was mainly dependent on hulled cultivars, which
do not seem to be replaced here in most cases by
free-threshing wheats, as was the case in the
Mediterranean basin and south-west Asia.
Representative sites for this period are the Brześć
group (Bieniek 2007) and contemporaneous
Gwoździec, Poland (Bieniek and Lityńska-Zając
2001; Lityńska-Zając 2007); Neolithic Vinča culture,
Liubcova, Rumania (Cârciumaru 1996); Zánka-
Vasútállomás, Hungary (Füzes 1990, 1991); Blatné,
Slovakia (Hajnalová 1989); Bietigheim-Bissingen
(Piening 1989) and Hienheim (Bakels 1978), Germany;
Schletz, Austria (Schneider 1994; Kohler-Schneider
2007); Cova de Can Sadurní, Spain (Blasco et al. 1999);
and Aubechies, Belgium (Bakels and Rousselle 1985).

Compared to its principal role in the Balkans and
in the central European Linearbandkeramik cul-
ture, einkorn is much scarcer in plant remains
obtained from the early Impressed Ware culture in
the west Mediterranean basin. Einkorn appears
together with emmer and free-threshing wheat in
the Impressed Ware (Cardial) settlements in Italy,
like ca. 8,000–7,600 cal BP Scamuso (Costantini et al.
1997), ca. 7,750–7,250 cal BP Pienza (Castelletti
1976), and ca. 7,750–7,150 cal BP La Marmotta
(Rottoli 1993, 2002), but rarely as a common crop.
In Early Neolithic Sammardenchia and other adja-
cent northern Italian sites (Rottoli 1993, 2002, 2005;
Rottoli and Pessina 2007), einkorn takes a more
important role. It appears in Impressed Ware Early
Neolithic of southern coastal France, like ca. 7,900–
7,500 cal BP Pont de Roque Haute, (Marinval 2007),
and ca. 7,650–6,700 cal BP Châteauneuf-les-
Martigues (Rottoli 2005; Rottoli and Pessina 2007).
In Spain, einkorn arrives only in the second half of
the eighth millennium, sites like ca. 7,400–7,050 cal
BP Coveta de l’Or (Hopf and Schubart 1965;
Lopez,1980), and adjacent Cova de Cendres (Buxó
1997), and contemporaneous Cova de Can Sadurní
(Blasco et al. 1999; López et al. 2003; Peña-Chocarro
2007) and North Meseta sites (Stika 1999). In most
of these contexts it does not serve as a major cereal,
and it occurs afterwards in these areas only
sporadically.
Some einkorn also appears in early Neolithic sites
in ca. 7,950–7,150 cal BP Aratashen and Aknashen,
Armenia (Hovsepyan and Willcox 2008), and con-
temporaneous Arukhlo 1 and 2, Georgia (Lisitsina
1984; Januševič 1984; Schultze-Motel 1988a). In
Transcaucasia, Chokh, Dagestan, it appears only in
the end of eighth millennium BP (Lisitsina 1984),
again as a member of the south-west Asian crop
ensemble.
The final movements of einkorn, and other
domesticated crops, further into Europe is less
rapid. During the seventh millennium, it reached
Switzerland in ca. 6,450–5,450 cal BP Middle
Neolithic sites in Sion (Martin et al. 2008a, 2008b);
(Lundström-Baudais in press) and later, ca. 6,250–
4,450 cal BP Late Neolithic Egolzwil 3 (Bollinger
1994; Jacomet 2007) and Zürich (Jacomet 1988,
2006; Jacomet et al. 1989; Brombacher 1997; Favre
2002; Brombacher et al. 2005); Early Neolithic, ca.
CEREALS 39
6,300–5,900 cal BP, Swifterbant S3, Netherlands
(van Zeist and Palfenier-Vegter 1981); and in ca.
6,000–4,750 cal BP to Eneolithic settlements in
Ukraine, like Luca Vrublevecaja (Januševič 1976),
Majdaneckoe, Majaki, and Usatovo (Pashkevich
2005). In the sixth millennium it reached early
Neolithic Britain, ca. 5,600–5,300 cal BP The
Stumble (Murphy 1989), and later to Middle
Neolithic, ca. 5,000 cal BP, Alvastra, Sweden
(Göransson et al. 1995; Hjelmqvist 1955) and Funnel
Beaker culture, ca. 4,950–4,850 cal BP Sarup,
Denmark (Jørgensen 1981).
Einkorn wheat persisted in many parts of Europe
until medieval times. Its cultivation almost disap-
peared in the twentieth and the twenty-first
centuries.
Emmer and durum-type wheats:
Triticum turgidum
This is a varied aggregate of domesticated tetra-
ploid (2n = 2x = 28 chromosomes) wheats. All share
BBAA chromosomes, and all are almost fully inter-
fertile with one another. Therefore, these wheats are
included in a single biological species (Table 3).
According to their response to threshing, the domes-
tic turgidum wheats fall into two groups of cultivars
that are frequently recognizable also in archaeologi-
cal remains:
(i) The products of threshing of hulled non-
shattering emmer wheat, T. turgidum L. subsp. dic-
occum (Schrank) Thell. (traditionally called T.
dicoccum Schübl.) are the individual spikelets (Fig.
4B, Plate 3). The grains remain invested by the pales
and glumes. In domestic emmer, as in einkorn,
threshing results in breaking the rachis of the ear in
its weakest points below each spikelet. This paral-
lels the disarticulation pattern found in wild emmer.
Taxonomically, emmer represents the primitive sit-
uation in domesticated turgidum wheats.
(ii) The more advanced free-threshing tetraploids
(Fig. 5) evolved under domestication from hulled
emmer. The common representative of this group
today is durum wheat, T. turgidum subsp. durum
(Desf.) Thell. [=T. durum Desf.]. Less common types
are: rivet wheat, T. turgidum subsp. turgidum L. [=T.
turgidum L.]; Polish wheat, T. turgidum subsp. polon-
40 DOMESTICATION OF PLANTS IN THE OLD WORLD
icum (L.) [=T. polonicum L.]; and T. turgidum subsp.
carthlicum (Nevsky) Löve & Löve [=T. carthlicum
Nevski]. The glumes in all these domesticated tetra-
ploid forms are relatively thin. Threshing breaks the
glumes at their bases and frees the naked grains.
The rachis is usually uniformly tough and thresh-
ing breaks it into irregular fragments.
Hulled emmer was the principal wheat of Old
World agriculture in Neolithic and early Bronze
Age at the Mediterranean basin, south-west Asia,
and temperate Europe. It was used for food and
apparently also for brewing beer. Later, more
advanced, naked free-threshing tetraploid and hex-
aploid domestic types gradually replaced it. At
present, hulled emmer is still a relic crop, sporadi-
cally grown in some parts of Europe and south-west
Asia; i.e. the Balkan countries, eastern Slovakia,
Hungary, Spain, Anatolia, Iran, Caucasia, and India
(Nesbitt and Samuel 1996b; Perrino et al. 1996).
Free-threshing durum-type hard wheats are the
main contemporary representatives of the BBAA
tetraploid wheats. Such naked, free-threshing tetra-
ploids (possibly including the small grain forms
described by Kislev (1980, 2009) as T. parvicoccum)
probably appeared already in Neolithic times and
gradually gained prominence. Since classical times,
free-threshing tetraploid cultivars constituted the
main wheat crop in the summer-dry, relatively
warm Mediterranean basin. In more continental cli-
mates and in temperate areas with summer rains,
traditional wheat production depended heavily on
another free-threshing wheat, namely hexaploid
T. aestivum.
Wild ancestry
Genetic and morphological evidence indicates (for
review see Zohary 1969; Feldman et al. 1995; Feldman
and Kislev 2007) that the domesticated tetraploid
turgidum wheats (both the hulled dicoccum forms
and the free-threshing durum-type varieties) are
closely related to a wild wheat native to south-west
Asia, and traditionally called T. dicoccoides (Körn. ex
Aschers. et Gräbn.) Schweinfort Thell. and currently
classified as T. turgidum subsp. dicoccoides (Körn. ex
Asch. & Graebn.) Thell. (wild emmer wheat, see
Plate 1, Plate 2, Fig. 4, Table 3). This is an annual,
predominantly self-pollinated, tetraploid wheat,
with characteristic large and brittle ears and big
elongated grains that show a striking similarity to
some domesticated emmer (Fig. 5) and durum (Fig.
5) cultivars. It is the only wild stock in the genus
Triticum that is cross-compatible and fully inter-
fertile with the domesticated turgidum wheats.
Hybrids between wild dicoccoides forms and all
domesticated members of T. turgidum aggregate
show normal chromosome pairing in meiosis, indi-
cating that all these tetraploid wheats contain
homologous chromosomes (BBAA chromosome
sets). The close genetic affinities between wild
emmer and the domesticated members of T. turgi-
dum are also indicated by spontaneous hybridiza-
tion that occurs occasionally when these wild and
domesticated wheats grow side by side. On the
basis of these close genetic and morphological rela-
tionships, wheat researchers regard subsp. dicoc-
coides as the wild progenitor of the domesticated
tetraploid emmer and durum wheats. They rank
wild emmer as the wild subspecies of the T. turgi-
dum crop complex (see Tables 3 and 4).
In the tetraploid turgidum wheats, the most con-
spicuous diagnostic difference between wild and
tame is the seed-dispersal mechanism (Zohary and
Brick 1961; Zohary 1969; Elbaum et al. 2007). Wild
dicoccoides wheats have brittle ears that shatter upon
maturity into individual spikelets. Each spikelet
operates as an arrow-like device disseminating the
seed by inserting them into the ground. The ‘wild-
type’ rachis disarticulation and the spikelet mor-
phology reflect specialization in seed dissemination.
This ensures the survival of the wild forms under
wild conditions. Under the manmade system of till-
age, reaping, threshing, and sowing, this adaptation
broke down and non-brittle types were automati-
cally selected. Significantly (Table 7, p. 61), the shift
from a brittle spike (in wild dicoccoides wheat) to a
non-brittle spike (in domesticated dicoccum wheats)
is governed by recessive alleles at two major loci
that are located respectively on the short arm of
chromosomes 3A and 3B (Watanabe et al. 2002, 2006;
Nalam et al. 2006). In addition, wild and domesti-
cated forms differ from one another in kernel
morphology (van Zeist 1976). In domesticated dicoc-
cum and durum forms, the grain tends to be wider
and thicker as well as rounder in cross-section com-

pared to the wild dicoccoides counterpart (compare
grains in Figs 4 and 5). These domestication traits
are quite helpful in analyzing archaeological
remains.
Wild emmer, T. turgidum spp. dicoccoides (Körn. ex
Asch. & Graebn.) Thell. [=T. dicoccoides (Körn.)
Schweinf.] is more restricted in its distribution and
more confined in its ecology than wild einkorn. Its
range covers the Fertile Crescent: Israel, Jordan,
south-western Syria, Lebanon, south-eastern
Turkey, northern Iraq, and western Iran (Map 4). It
is most widespread in the catchment area of the
upper Jordan Valley. Wild emmer was found in 1906
in eastern Galilee and on the slopes of Mt Hermon
in Israel (Aaronsohn 1910; Schiemann 1956). Wild
dicoccoides wheats grow as common annual compo-
nents in the herbaceous cover of the Tabor oak
(Quercus ithaburensis) park-forest belt and related
steppe-like herbaceous plant formations. They are
confined to basaltic and hard limestone bedrocks
and are completely absent on marls and chalks. In
rocky places that have not been severely over-
grazed, dicoccoides wheat often grows in large
stands. With wild barley Hordeum spontaneum and
wild oat Avena sterilis, they form ‘fields of wild
cereals’.
Further north, wild emmer occurs in the Anti-
Lebanon range and again in the oak park-forest belt
of Quercus brantii in south-eastern Turkey, north
Iraq, and west Iran (Map 4). Whereas dicoccoides
wheat occurs alone in the Levant, it grows sympatri-
cally with a second wild tetraploid wheat, T.
timopheevi subsp. armeniacum (Jakubz.) van Slageren
(see p. 51), in the north-eastern part of its distribution
area. Cytogenetically, these two wild wheats have
distinct genomic and cytoplasmic constitutions and
are separated from one another by strong sterility
reproductive barriers (Sachs 1953; Wagenaar 1966;
Maan 1973). However, they are so similar morpho-
logically that it is very hard or even impossible to
separate between ears of the two species without
cytogenetic confirmation or the use of molecular
genetic markers. In the north and the north-east, dic-
occoides wheat is closely associated not only with
Hordeum spontaneum but also with wild einkorn, T.
monococcum subsp. baeoticum and T. urartu.
In north Israel and in south Syria, dicoccoides wheat
shows a multitude of forms and often builds con-
CEREALS 41
spicuously polymorphic populations that are easily
noticed by their variation in glumes’ hairiness, col-
our of the spike, the size of the spikelets, and the
shape of the leaves (Poyarkova 1988; Poyarkova and
Gerechter-Amitai 1991). Climatically too, wild
emmer shows a considerable range of adaptation. It
is distributed over a wide altitudinal range. Robust,
early maturing, types occupy the winter-warm basin
around the Sea of Galilee to altitudes as low as 100 m
below sea level. More slender, late-blooming forms
occur higher up in the Galilee mountains and climb
to elevations of 1600 m on the east- and south-facing
slopes of Mt Hermon. In the Zagros Mountains of
north Iraq and west Iran, dicoccoides wheats occur at
altitudes ranging from 700 to 1600 m.
Molecular studies have placed a probable single
origin of emmer wheat in Turkey. Özkan et al. (2002)
examined twenty-four AFLP loci of ninety-noine
wild emmer lines from all over the Fertile Crescent
belt and several dozens of different cultivars. They
showed that the most likely place of origin for
emmer wheat is in south-eastern Turkey. Recently, a
study based on 131 RFLP loci, Luo et al. (2007)
reached similar results.
Archaeological evidence
(a) Hulled emmer wheats: Similar to einkorn wheat
(p. 34) and barley (p. 51), emmer wheat was col-
lected from the wild long before its domestication.
Brittle, dicoccoides-like remains, with relatively
narrow grains (Fig. 12) appear in several early
Fertile Crescent sites. A clear indication of pre-
agriculture gathering of wild emmer wheat is avail-
able from the south-western shore of the Sea of
Galilee in Israel. Here, remains of subsp. dicoccoides
(as well as wild barley) were recovered from Ohalo
II, a submerged ca. 23,000 cal BP early Epi-
Paleolithic site (Kislev et al. 1992; Simchoni 1998;
Weiss 2002, 2009; Weiss et al. 2004, 2008). Dicoccoides-
type grains were discovered in ca. 11,700–10,550 cal
BP, PPNA Netiv Hagdud and Gilgal, Israel (Kislev
1997; Hartmann 2006; Weiss et al. 2006).
The earliest fully convincing sign to-date of domes-
ticated emmer come from the numerous spikelet
forks retrieved from ca. 10,250–9,550 cal BP Early
PPNB Çayönü (van Zeist 1972; van Zeist and de
42 DOMESTICATION OF PLANTS IN THE OLD WORLD

0 100 200 miles Triticum dicoccoides

0 200 400 km Triticum ararticum

Map 4 Geographical distribution of wild emmer wheat, Triticum turgidum subsp. dicoccoides [= T. dicoccoides] and wild Timopheev’s wheat,
T. timopheevii subsp. araraticum [= T. araraticum], that grow side by side in the same habitats (compiled from Harlan and Zohary 1966; Rao and
Smith 1968; Dagan and Zohary 1970; Maan 1973; Tanaka and Ishii 1973; Tanaka et al. 1979; and unpublished data of D. Zohary).

Roller 1991–2, 2003). Here, hundreds of spikelet
forks were discovered, all showing rough breakage
scars characteristic of domestic emmer. A rough
breakage scar is by far the best indication that we are
dealing with domesticated forms of wheat in archae-
ological deposits. These spikelet forks are accompa-
nied by wild-type narrow grains, which give some
indication that mutant rachises predated the increase
in seed-size in the domestication process. Numerous
spikelet forks, with similar telltale rough scars, are
available also from contemporary contexts at Cafer
Höyük (de Moulins 1997). (A third possible site,
Nevali Çori (Pasternak 1998), has been announced
only preliminarily; we wait for its final publication.)
All these finds indicate that at Early PPNB, emmer
domestication must have been well underway in the
Fertile Crescent.
The domesticated status of emmer wheat from
ca. 10,650–9,550 cal BP Early PPNB Kissonerga-
Mylouthkia and Shillourokambos, Cyprus (Willcox
2000; Murray 2003) was not confirmed yet by chaff
remains. Even so, as no wild wheats grow in the
island, it seems that this find represents early intro-
duction of agriculture from outside.
A less reliable diagnostic trait for early domesti-
cation of wheat is grain size. In cultivars, seeds are
usually plumper and larger than those in their wild
relatives (see grain sizes in Figs 3–4, 11–12).
However, this process takes longer, and therefore
size increase cannot indicate reliably the start of
domestication (see p. 22).
To-date, the earliest conspicuous grain-size
increase is reported from Early PPNB Tell Aswad,
25 km south-east of Damascus (van Zeist and
Bakker-Heeres 1985). Here, dicoccum-like plump
kernels start to appear in the lowest habitation level
Ia, 10,500–10,200 cal BP (van Zeist and Bakker-
Heeres 1985; Willcox 2005), and also in Middle
 CEREALS 43

A B
0 5 mm

Fig. 12 Comparison between carbonized grains of wild and of domesticated emmer wheats, Triticum turgidum, Neolithic Çayönü, Turkey (van
Zeist 1972). A–Wild emmer wheat, Triticum turgidum subsp. dicoccoides [= T. dicoccoides]. B–Domesticated emmer wheat, Triticum turgidum
subsp. dicoccum [= T. dicoccum]. Note that compared to the more ‘spindle-shaped’ grains of one-seeded spikelets of domesticated einkorn (Fig.
13) the grains of cultivated emmer have a wider, blunter apex and a straighter side shape.

PPNB phases II (10,200–9,550 cal BP). In these
phases, however, no rachis segments have been
retrieved. Significantly, no wild dicoccoides-like nar-
row kernels were retrieved from Tell Aswad. The
present climate in the rain-shadowed Damascus
basin is far too dry for wild wheat, and it was prob-
ably arid 10,000 years ago also). As van Zeist and
Bakker-Heeres emphasize, the continuous presence
of morphologically discernible dicoccum kernels, the
total absence (from the very start) of dicoccoides-like
material, and the extreme dryness (less than 200 mm
annual rainfall) suggest that emmer wheat was
introduced into the Damascus basin as a domesti-
cated cereal, not later than thesecond half of the
eleventh millennium BP. From ca. 9,800–8,700 cal
BP onward, charred grains, that morphologically
conform with dicoccum, appear also at Tell Abu
Hureyra, north-east Syria, again with no rachis seg-
ments (Hillman 1975, 2000b; Hillman et al. 1989),
and in contemporary Pre-Pottery Neolithic B ca.
9,450–8,450 cal BP, Can Hasan III and ca. 9,350–8,950
cal BP, Çatalhöyük East, Turkey (Helbaek 1964a;
Fairbairn et al. 2002, 2005, 2007), ca. 9,600–8,750 cal
BP Ali Kosh, Khuzistan (Helbaek 1969), ca. 9,900–
9,550 cal BP, Jericho, Israel (Hopf 1983) and ca.
9,450–9,300 cal BP, Jarmo, Iraq (Braidwood 1960;
Helbaek 1959a, 1960).
From the very beginnings of agriculture in the
Fertile Crescent (some 10,500–10,100 cal BP), emmer
is the principal cereal of the newly established Pre-
Pottery Neolithic B (PPNB) farming settlements
(Map 1). Although remains of domesticated einkorn
and domesticated barley also occur quite regularly
in these contexts, emmer prevails quantitatively in
most cases. At several of the examined sites the
remains of emmer include not only carbonized
44 DOMESTICATION OF PLANTS IN THE OLD WORLD
plump seeds, but also well-preserved spikelets or
spikelet forks showing the effects of pressure frac-
turing at the base of the rachis internode (Figs 7 and
8). The prevalence of rough scars indicates that this
wheat had non-brittle ears. This is unequivocal
proof that there and then, grain cultivation was
already in practice.
Emmer wheat continues to be the principal grain
crop during the later stages of the Neolithic in
south-west Asia, and it was apparently widely
grown in this area also in Chalcolithic and Bronze
Age times, although in the later periods the hulled
cultivars (emmer and einkorn) were replaced by
free-threshing wheats.
Emmer wheat was also the main crop in the
spread of the Neolithic agricultural technology
from the Fertile Crescent ‘core area’ towards the
west, the north, the east, and the south. It is the most
common constituent of the crop assemblage that
started agriculture in the Aegean region in the ninth
millennium BP. Such sites are ca. 9,000–7,600 cal BP,
Dhali Agridhi, Cyprus (Stewart 1974), ca. 9,000–
8,050 cal BP, Franchthi Cave (Hansen 1991b, 1992),
and ca. 8,650–8,200 cal BP, Sesklo, Greece (Hopf
1962; Kroll 1981a), and ca. 8,650–8,400 cal BP,
Knossos, Crete (Sarpaki 2009) (for review, see
Renfrew 1979; Kroll 1981a, 1981b, 1991). In the sec-
ond half of the ninth millennium BP, emmer wheat
is continuing to spread to the Balkan countries, such
as ca. 8,150–7,000 cal BP, Obre, Macedonia (Renfrew
1974, 1979; Kroll 1991). In the early Neolithic, it also
spread to Central Asia ca. 8,200–7,850 cal BP, Jeitun
(Djeitun) (Charles and Hillman 1992; Harris and
Gosden 1996).
Emmer was the principal cereal of the Linear-
bandkeramik farmers that started Neolithic agri-
culture in central Europe in the first and second
half of the eighth millennium BP. Evidence can be
found at sites like ca. 7,950–5,250 cal BP, Körös-
Starčevo, Hungary (Hartyányi et al. 1968; Hartyáni
and Nováki 1971, 1975; Füzes 1990); ca. 7,650–7,400
cal BP Mohelnice, Czech Republic (Opravil 1979,
1981; Kühn 1981); ca. 7,500–6,550 cal BP,
Aldenhovener Platte, Germany (Knörzer 1973,
1974, 1997); ca. 7,450–6,950 cal BP, Brześć, Poland
(Bieniek 2007); ca. 7,150–6,800 cal BP, Schletz,
Austria (Schneider 1994; Kohler-Schneider 2007).
At the numerous Linearbandkeramik sites exam-
ined to-date (for review, see Willerding 1980; Nesbitt
and Samuel 1996b), emmer is usually found side by
side or in admixture with einkorn. Yet in most
places, emmer is the more common wheat. This
relationship persists all over central and northern
Europe during the later Neolithic and the Bronze
Age. Thus in central Europe, wheat production
during the Neolithic and Bronze Age depended
heavily on hulled wheats. Naked wheats do not
seem to have replaced them as quickly as they did
in the Mediterranean basin and in south-west Asia,
though they are present in central Europe since the
Neolithic.
Rich finds of emmer have also been discovered
at places belonging to the Impressed Ware culture
that introduced agriculture into the western
Mediterranean basin during the eighth millennium
BP (Hopf 1991). Such sites include ca. 8,000–7,600
cal BP, Scamuso, Italy, ca. 7,400–7,050 cal BP Coveta
de l’Or and Cova de Cendres (Costantini et al.
1997), and ca. 7,400–6,650 cal BP Cova de Can
Sadurní (Blasco et al. 1999), Spain. In Impressed
Ware contexts, emmer is generally accompanied by
free-threshing wheats, and in many localities the
latter prevail. Emmer is also found in younger sites
of this region, but again, only in minor quantities
compared to the more advanced free-threshing
wheats.
Emmer is, however, the wheat of Neolithic
and Bronze Age Egypt (Fig. 13, Plate 5), such as
ca. 7,500–6,650 cal BP Fayum and 1,325 BC
Tutankhamun tomb (Caton-Thompson and Gardner
1934; Täckholm 1976; Germer 1989a; Hepper 1990;
Wetterstrom 1993; Wendrich and Cappers 2005; de
Vartavan et al. 2010; see also Fig. 8).
As to the diffusion of Neolithic agriculture from
the Fertile Crescent towards the east, there is much
less information at hand. Judging by the rare data
available, emmer wheat was one of the main ele-
ments in the crop assemblage discovered in several
Neolithic sites in Caucasia and Transcaucasia, dated
into the eighth millennium BP, as attested by the
finds from ca. 7,950–7,150 cal BP Aratashen and
Aknashen, Armenia (Januševič 1984; Wasylikowa
et al. 1991; Hovsepyan and Willcox 2008).
Finally, it should be highlighted that among the
three main Neolithic grain crops (einkorn, emmer,
and barley), the wild progenitor of emmer has the

0 1 cm
Fig. 13 Desiccated spikelets of domesticated emmer wheat, Triticum turgidum subsp. dicoccum (G. Schweinfurth collection, fifth-dynasty Abusir,
Egypt).
most limited geographic distribution. It is confined
to the Fertile Crescent ‘arc’—compare Map 3 with
Maps 4 and 6. Therefore, the numerous finds of
domesticated emmer retrieved from sites outside
the Fertile Crescent serve as convincing evidence
for the fertile crescent as the core area where
Neolithic agriculture could have evolved, and sub-
sequently disseminated.
(b) Free-threshing tetraploid wheats: Free-threshing
wheat forms, identifiable by their rachis fragments
(Fig. 9A), made their appearance in south-west Asia
soon after the appearance of domesticated emmer
wheat (for review see Maier 1996). Some remains
start to appear already among the plant remains of
two Middle PPNB sites, ca. 10,200–9,550 cal BP Tell
Aswad, Syria (van Zeist and Bakker-Heeres 1985),
and ca. 10,100–9,450 cal BP Aşikli Höyük, Turkey
(van Zeist and de Roller 1995). More were found
CEREALS 45
somewhat later, in ca. 9,450–8,450 cal BP Late/Final
Neolithic Can Hasan III, Turkey, where numerous
kernels and rachis fragments were found (Hillmam
1972, 1978), and Late PPNB 9,450–8,600 cal BP Tell
Bouqras, Syria (van Zeist and Waterbolk-van Rooijen
1985). They also occur in ca. 9,350–8,950 cal BP
Middle/Late PPNB Çatalhöyük, Turkey (Helbaek
1964a; Fairbairn et al. 2002, 2005, 2007), and Tell
Ramad, Syria (van Zeist and Bakker-Heeres 1985).
In the first stages of research, archaeobotanists
could not distinguish between tetraploid and hexa-
ploid free-threshing wheats. Almost always, they
have referred to them as T. turgidum-T. aestivum or
as ‘aestivo-compactum’ wheats. Yet most researchers,
when faced with remains obtained from the PPNB
farming villages in the Fertile Crescent, assumed
that they represent 4x durum-like forms rather than
6x aestivum wheats. As argued by Zohary (1969b)
46 DOMESTICATION OF PLANTS IN THE OLD WORLD
and by van Zeist (1976), hexaploid bread wheats
must have originated outside the ‘Fertile Crescent
area’; and this development occurred only after
Neolithic wheat agriculture reached the primary
habitats of Aegilops tauschii distribution area (Map
5). This contact very likely happened in the Caspian
belt somewhere between 8,000 and 7,000 BP. Thus,
the early naked wheats in the Fertile Crescent core
area could not possibly have been 6x T. aestivum.
They should be attributed to 4x T. turgidum.
Nesbitt (2001) claim, in contrast, for a different
scenario, based on the hexaploid rachis remains
from ca. 9,450–8,450 cal BP PPNB Can Hasan III
(Hillmam 1972, 1978) and from Cafer Höyük (de
Moulins 1997). According to Nesbitt, these finds
demonstrate either that agriculture reached the
Caspian belt earlier, or that the prehistoric primary
distribution of Ae. tauschii extended further west.
The chromosomal level of the free-threshing
wheats retrieved from later south-west Asian con-
texts, and from sites representing the early stages of
the agricultural expansion to Europe and west Asia,
has been even more problematic. In many parts of
0 100 200 miles
0 200 400 km
Map 5 Geographical distribution of Aegilops tauschii [= Ae. squarrosa]. The areas in which primary habitats occur are shaded. Dots represent
additional sites, mainly of weedy forms (based on Zohary et al. 1969; van Slageren 1994).
this vast area (for details see Maier 1996), naked
wheats continue to appear—in Neolithic and in
Bronze Age times—in progressively increasing pro-
portions. But assigning them to either 4x T. turgidum
or to 6x T. aestivum was until recently practically
impossible. They were almost always referred to as
‘aestivo-compactum’ or T. turgidum-T. aestivum forms.
From the beginning of the seventh millennium
BP onward, free-threshing wheats appear in Greek
sites. Further west, naked wheats abound in the
Impressed Ware (Cardial) sites of Italy, south France,
and Spain; i.e. in settlements which represent the
early stages of agriculture expansion into the west
Mediterranean basin (Hopf 1991; Maier 1996). Free-
threshing wheats also abound in the Middle
Neolithic lake-shore sites of Switzerland and its
neighboring areas (Jacomet et al. 1991; Maier 1996;
Jacomet 2004, 2006, 2007). Thus, in south-west Asia,
the Mediterranean basin and the Alps, we are faced
with a rather early, partial replacement of hulled
wheats by free-threshing types. Emmer and einkorn
continued to occur and they are occasionally
retrieved in large quantities. Yet towards the Late

Bronze Age, south-west Asia and the Mediterranean
basin both commonly show a prevalence of naked
wheats. The large quantities of naked wheat
unearthed in Bronze Age towns in the Levant are
also impressive. In contrast, the wheat of ancient
Egypt was hulled emmer and it retained its domi-
nance in the Nile Valley as late as Hellenistic times.
It was replaced first by naked durum wheat by the
Ptolemaic rulers (Murray 2000b; Wetterstrom
1993).
Our ability to distinguish between the 4x and 6x
free-threshing wheats on the basis of rachis remains
is already established (Jacomet 1987; Hillman et al.
1996; Maier 1996). As noted in Fig. 9B (p. 32), a set of
morphological traits, traceable in rachis segments,
has been found to be diagnostically reliable for such
identification. Several reports have already appeared
in which the remains of naked forms of 4x T. turgi-
dum and of 6x T. aestivum are soundly recognized.
Thus Maier (1996) had convincingly demonstrated
that the early free-threshing wheats grown at the
Middle Neolithic lake-shore site at ca. 5,850–5,450
cal BP Hornstaad, Lake Constance, Germany, were
tetraploid (Maier 2001). Maier also stresses the
fact that some of the rachis segments, whole spikes,
and parts of spikes retrieved from several other
well-investigated, Middle Neolithic lake-shore sites
at the northern foothills of the Alps, manifest 4x T.
turgidum morphology (see also Jacomet et al. 1989,
1991; Brombacher 1997). These finds indicate that at
that time, wheat production was already based, in
part, on growing naked tetraploid forms.
So far, the Alpine Middle Neolithic sites consti-
tute the only area where the identification of the 4x
naked T. turgidum (as well as 6x T. aestivum) has
been carried out on a large scale. But one would
expect that additional finds of ‘aestivo-compactum’
wheats, coming from other parts of Europe and
west Asia, will be re-examined soon. Very likely the
origins and spread of the free-threshing 4x and 6x
wheats will be much better understood in a few
years from now.
Bread wheat: Triticum aestivum
Bread wheat is the most variable aggregate of
domesticated wheats, and economically the most
important wheat species. It accounts for about 90%
CEREALS 47
of the total world wheat production today and
includes numerous and contrasting types. All varie-
ties are hexaploid (2n = 6x = 42 chromosomes) and
inter-fertile when crossed with one another. All
share the BBAADD genomic constitution.
Hexaploid T. aestivum is a new wheat species that
evolved under domestication from the domesti-
cated tetraploid T. turgidum stock. In contrast with
the diploid and tetraploid wheats, it does not have
a wild hexaploid counterpart. For cytogeneticists
and plant evolutionists, bread wheat stands as one
of the classic examples of evolution by polyploidy.
Genome analysis studies have shown (Kihara 1944;
McFadden and Sears 1944, 1946) that T. aestivum is a
hybridization product between tetraploid turgidum
wheat (genomic constitution BBAA) and a diploid
wild grass Aegilops tauschii Coss. [= Ae. squarrosa L.]
(genomic constitution DD). In other words, hexa-
ploid bread wheats originated by the addition of a
third genome to the two genomes already present
in tetraploid T. turgidum. Bread wheat has been syn-
thesized in the laboratory by crossing the two puta-
tive parents and doubling the chromosomes in the
hybrids. Since no BBAADD hexaploid wheat occurs
in the wild (except subsp. tibetanum which is a feral
wheat that presumably escaped from cultivation),
this development could have occurred only under
cultivation; i.e. by the hybridization of the already
domesticated tetraploid wheat with the wild Ae.
tauschii.
Hexaploid wheats fall into two groups according
to their response to threshing:
(i) All the cultivars in this group are hulled (glumed),
and today they survive mainly as relic crops.
Prominent among them is spelt wheat, T. aestivum
subsp. spelta (L.) Thell. (traditionally called T. spelta
L.). An additional type, the macha wheat, T. aesti-
vum subsp. macha (Dekapr. et Menabde) MacKey
[=T. macha Dekapr. et Menabde], endemic to west-
ern Georgia, also belongs to this group. In all hulled
hexaploid wheats, the products of threshing are the
individual spikelets. But while in einkorn and
emmer the rachis segment below the spikelet
remains attached to the unit (Fig. 7 A, B), in hulled
hexaploid wheats, the segment attached is fre-
quently the internode above the base of the spikelet
(Fig. 7 C). In archaeological remains, the upper
48 DOMESTICATION OF PLANTS IN THE OLD WORLD
rachis segments, as well as the characteristic struc-
ture of the glumes, serve as diagnostic traits for
spelt-wheat identification.
(ii) Free-threshing bread wheats (Fig. 6 A, B, C) are
predominantly used today. Most important and
almost universal is bread wheat or common wheat,
T. aestivum subsp. aestivum MacKey [also referred to
as T. aestivum L., T. sativum L., or T. vulgare Host].
Two other free-threshing hexaploid types are: (a)
Club wheat, T. aestivum subsp. compactum (Host)
MacKey [= T. compactum Host], with compact ears.
It is grown today in Afghanistan, as well as in the
north-western United States. It has also been found
in early European cultures; (b) Indian dwarf wheat,
T. aestivum subsp. sphaerococcum (Perc.) MacKey [=
T. sphaerococcum Perc.], with characteristic small
grains, is native to India and Pakistan.
Experimental evidence indicates that the first hexa-
ploid wheats were spelt-like. Artificial synthesis of
T. aestivum (by crossing and fusing tetraploid BBAA
T. turgidum with diploid DD Aegilops tauschii)
always results in hulled products (Kerber and
Rowland 1974), irrespective of whether the turgi-
dum BBAA parent is hulled or naked. It also sug-
gests that the free-threshing condition in hexaploid
bread wheats was brought about by two events: the
appearance of the free-threshing gene Q located on
chromosome 5A, and the mutation from Tg to tg in
the gene responsible for the tenacious glumes trait
on chromosome 2D. All present-day 6x naked
wheats examined carry the tg/tg Q/Q genotype.
As already mentioned (pp. 32, 47), the history of
the 6x free-threshing bread wheats could not be eas-
ily reconstructed first, since their archaeological
remains were difficult to separate from those of 4x T.
turgidum. In the past, the shape of the kernel was
considered a valuable diagnostic trait, because hexa-
ploid forms tend to have relatively plumper grains
with blunt tips. However, grain shape proved to be
problematic because tetraploid and hexaploid
wheats overlap considerably in their seed shape.
This overlap is more prominent in archaeological
remains, because of swelling and other deforma-
tions caused by the charring (Hopf 1955; van Zeist
1976; Harlan 1981). Thus carbonized wheat remains
showing obtuse, plump kernels need not necessarily
be 6x T. aestivum. They may also represent 4x T. tur-
gidum. Similar difficulties have been encountered
with another feature of the grain, namely the shape
of the scutellum (the connecting tissue between the
embryo and the endosperm). It is short, wide, and
shows a straight profile in the bread wheats; it is
elongated, curved, and shows a curved profile in the
durum and einkorn wheats. Yet in this trait, one also
encounters variation and some overlapping. For
these reasons most archaeobotanists in the past did
not attempt to distinguish between 4x and 6x naked
wheats. Instead, they lumped them together as ‘aes-
tivo-compactum’ or T. turgidum-T. aestivum finds.
Not so long ago (Hillman 2001), it has become
clear that free-threshing 4x T. turgidum forms can be
separated from their 6x T. aestivum counterparts by
the morphology of their rachis segments (Fig. 9B).
Parallel to what is happening with the free-threshing
4x wheats (pp. 45–46), a more precise understanding
of the history of the naked 6x wheats was suggested
by Maier (1996), but more research is needed.
Wild ancestry
As already stated (see above), hexaploid bread
wheat originated by the addition of the DD chro-
mosome complement of Aegilops tauschii [=Ae.
squarrosa] to the domesticated, tetraploid BBAA tur-
gidum wheats. Ae. tauschii (Fig. 14) was never
domesticated as such. The examination of its ecol-
ogy and distribution reveals that this wild grass
contributed substantially to the adaptation and the
worldwide success of the bread wheats.
The first significant feature about Ae. tauschii is its
distribution, which also reflects its climatic require-
ments (Zohary 1969). This is the easternmost dip-
loid species in the wheat group (Triticum-Aegilops).
Its centre of distribution does not lie in the
Mediterranean south-west Asia but in continental
or temperate central Asia. It is widespread and very
common in northern Iran and adjacent Transcaucasia,
Transcaspia, and Afghanistan (see Map 5, and Fig 73
in van Slageren 1994). From this geographic centre,
Ae. tauschii spreads west to east Syria, and east to
Pakistan. In central Asia it is recorded as far east as
Kirghizia and adjacent parts of Kazakhstan.
Ae. tauschii is a variable species. It is represented
by a multitude of forms, from slender types with
cylindrical spikes, to more robust plants with thick,
 CEREALS 49

beaded spikes (see Fig. 14). The latter (subsp. stran-

gulata) are confined to the south Caspian basin. Ae.
tauschii is not only widely distributed, it also occurs
over a wide range of ecological conditions. Like
wild einkorn and wild barley, it occupies both pri-
mary and segetal habitats. In the centre of its distri-
bution this plant is a frequent annual component of
open formations. It thrives in areas characterized by
continental climatic conditions, from the dry sage-
brush steppes of the elevated Iranian and Afghan
plateaus to desert margins, as well as in more tem-
perate climates such as the rain-soaked southern
coastal plain of the Caspian Sea. At the same time,
all over this area Ae. tauschii is a successful colonizer
of secondary, manmade habitats and a common
weed in cereal fields. Towards the periphery of its
0 1 2 cm
distribution, it is almost exclusively a weed in culti-
vation. Thus in the case of Ae. tauschii, we are faced
with a colonizer plant which apparently expanded
its distribution over secondary habitats with the Fig. 14 Lower parts of the spike, and individual spikelets, of diploid
Aegilops tauschii [= Ae. squarrosa], the donor of the D genome to
opening up of the land by agriculture.
hexaploid Triticum aestivum wheats (from the reference collection of
These ecological and distributional facts provide D. Fuller, Institute of Archaeology, UCL).
clues to the plausible place of origin of T. aestivum
and explain some of the bread wheat’s ecological
specifications. At the start of agriculture, the two ically placed in subsp. strangulata) are closest to the
contributors that fused to form the hexaploid wheats genome D found in the hexaploid wheats.
were evidently geographically separated. Wild The addition of the D genome greatly extended
emmer was restricted to south-west Asia and Ae. the range of adaptation of domesticated wheats
tauschii did not spread westward from north Iran. (Zohary 1969). As mentioned above, domesticated
Thus contacts between the tetraploid wheats and Ae. tetraploid wheats are fit to the Mediterranean-type
tauschii could have been established only after the environments (with mild winters and warm, rain-
domestication of emmer and the spread of domesti- less summers). The incorporation of the Ae. tauschii
cated tetraploid T. turgidum to north Iran and adja- genome made the hexaploid plants more capable of
cent Transcaucasia. This expansion probably took withstanding continental winters and humid sum-
place between 8,000 and 7,000 BP (van Zeist 1976). mers. This facilitated the spread of hexaploid bread
The most likely area of origin of the hexaploid bread wheats over the continental plateaus of Asia and
wheat, therefore, is the south-western corner of the the colder, temperate areas in eastern, central, and
Caspian belt. Such association between domesti- northern Europe. It also explains the distribution
cated T. turgidum and weedy Ae. tauschii in cultiva- patterns of wheats in traditional agriculture; i.e. the
tion can still be found in this area (Matsuoka et al. centering of the 4x durum races in south-west Asia
2008). This suggestion is supported by molecular and the Mediterranean basin, and the prevalence of
studies which have revealed the range and the struc- 6x aestivum forms in temperate Europe and conti-
turing of isozyme and DNA polymorphism in Ae. nental west Asia.
tauschii and compared it with its counterparts on the Additional explanation for the origin of European
D genome of hexaploid T. aestivum (Dvořák et al. spelt as independent of the Asian spelt was already
1998). The assembled data indicate that populations proposed in the first half of the twentieth century
of Ae. tauschii native to Armenia and the south-west (e.g. Bertsch F. 1943; Bertsch K. 1950; Kuckuck and
part of the Caspian Sea belt (mainly forms taxonom- Schiemann 1957). The proponents of this hypothesis
50 DOMESTICATION OF PLANTS IN THE OLD WORLD
claim that European spelt was formed by secondary
hybridization between T. aestivum and T. dicoccum
(and see discussion in Nesbitt 2001). An important
development for this hypothesis gain Recently,
Blatter, Jacomet and Schlumbaum (2002) supported
the European origin by analyzing of glutenin subu-
nit genes B1-1 and A1–2 in fifty-eight accessions of
hexa- and tetraploid wheat from Europe and Asia.
Their results suggests a polyphyletic origin of the
A- and B-genomes of hexaploid wheat, and that
European spelt does not derive from the hulled pro-
genitors of bread wheat but originated by introgres-
sion of a tetraploid wheat into free-threshing
hexaploid wheat as a secondary evolution after the
development of bread wheat.
Archaeological evidence
(a) Hulled (=glumed)spelt wheat: Spelt has been
reported by Januševič (1984) from ca. 8,000–7,150
cal BP Neolithic contexts of Arukhlo 1, Transcaucasia.
The finds are few, but they unmistakably show the
telltale upper rachis segment attached to the lower
spikelet. These data confirm several earlier, poorly
documented reports by other Russian researchers
(Lisitsina 1978) on the occurrence of spelt wheat in
some sites in the Kura river plain, Transcaucasia,
dating to the seventh millennium BP, and possibly
even to the eighth millennium BP. It is therefore
clear that in the seventh millennium BP, hexaploid
hulled T. aestivum was already grown in the Caspian
belt. Significantly these early finds come from the
area most likely to be the area of formation of hexa-
ploid wheats; i.e. a region in which the domesti-
cated 4x T. turgidum could have first come in contact
with (truly wild) 2x Ae. tauschii.
Another relatively early spelt wheat find comes
from the ca. 7,600–7,400 cal BP Early Neolithic
Sacarovca, Moldavia (Januševič 1984; Kuzminova
et al. 1998; Monah 2007a). Here, only a few charred
spikelets were detected among numerous remains
of emmer wheat. But these clearly display the diag-
nostic arrangement of the rachis segment of spelt.
Sometime later, spelt appears sporadically in east
and central European sites, like Zánka-Vasútállomás,
Hungary (Monah 2007a) and Blatné, Slovakia
(Hajnalová 1989). Similar finds are available from
late Neolithic Poland, e.g. ca. 6,350–5,950 cal BP,
Brześć (Bieniek 2007) and Rumania, e.g. ca. 6,950–
6,600 cal BP, Poduri (Cârciumaru and Monah 1985;
Monah and Monah 2008), Cortaillod sur les
Rochettes Est, Switzerland (Akeret 2005), and
Germany (Blankenhorn and Hopf 1982). In some
Tripolye culture settlements in Moldavia and
Bessarabia, spelt wheat was apparently grown as a
crop by itself (Januševič 1976, 1978). Several well-
preserved spikelets, admixed as an ‘impurity’ with
einkorn and emmer remains, were retrieved from
ca. 5,700 BP Gumelnitza culture Ovčarovo,
north-eastern Bulgaria (Januševič 1978). The well-
preserved spikelets leave little doubt that hexaploid
spelt wheat was already in use at that time. Spelt
wheat remains from Bronze and Iron Age are more
numerous and come from all over east, central, and
north Europe, (Hajnalová 1978; Gyulai 2003) as well
as from Greece (Kroll 1983). Spelt was also well
known to the Romans (Jasny 1944) and in Europe it
survived as a crop until the start of the twentieth
century. (Very small quantities of spelt are still
grown today in south Germany, north Spain, as well
as in several other parts of Europe and west Asia).
It is important to note that the presence of
Neolithic spelt is questioned recently by Nesbitt
(2001 and pers. comm.). In his view, all or most of
these finds are either of Ae. cylindrica (a common
weed), new-type emmer wheat (which has strongly
striated glumes, superficially similar to those of
spelt), or based on grain morphology and are thus
invalid. Detailed examination of published spelt is
under way and awaits publication. This view con-
curs with recent molecular evidence, which sug-
gests spelt arose anew in Europe, rather than move
from south-west Asia to Europe.
(b) Free-threshing bread wheat: Genetic analysis indi-
cates (p. 48) that naked hexaploid wheats could have
evolved rather quickly from their more primitive
spelt relatives since the shift was apparently pro-
duced by only two mutations. Thus it is plausible to
assume that shortly after the formation and estab-
lishment of hexaploid spelt, the naked derivatives
could have appeared as well. In other words, from
the seventh millennium BP onwards, the progres-
sively more common free-threshing wheats encoun-
tered in archaeological digs could represent not

only tetraploid forms but hexaploid cultivars as
well. A large scale re-examination (by the discrimi-
nating rachis morphology) of early remains of ‘aes-
tivo-compactum’ naked wheats in west Asia and
Europe has not yet been attempted. The continental
and temperate parts of west Asia and Europe are,
ecologically, the territories best suited for the early
establishment of the 6x free-threshing bread wheats.
It is therefore likely that from the late Neolithic and
the Bronze Age onward, remains of naked wheats
from places like Caucasia, central Asia, the central
Anatolian plateau, India, and east and central
Europe could represent largely 6x aestivum material
rather than 4x turgidum forms.
Timopheev’s wheat: Triticum
timopheevii
This tetraploid species comprises both domesti-
cated and wild forms that are genomically different
and reproductively isolated from the more common
tetraploid turgidum stock. F1 Hybrids between turgi-
dum and timopheevii wheats are sterile and manifest
a considerable amount of chromosomal irregulari-
ties in meiosis. On the basis of the available cytoge-
netic and molecular evidence, the genomic formula
assigned to T. timopheevii (Zhuk.) Zhuk. is GGAA.
In other words, GGAA T. timopheevii shares a simi-
lar genome A with BBAA T. turgidum and BBAADD
T. aestivum, but does not contain their B genome
(Sears 1969; Maan 1973). Moreover, even the genome
A found in T. timopheevii is different from that
encountered in T. turgidum and T. aestivum. It also
contains a distinct cytoplasm (Maan 1973), which
induces male sterility when combined with BBAA
chromosomes.
Domesticated Timopheev’s wheat is hulled, and
shows close morphological similarities to domesti-
cated emmer. It is an endemic crop restricted to
western Georgia. The wild wheat from which it
could have been derived is well known. The culti-
gens are fully inter-fertile and share identical chro-
mosomal constitution with a group of brittle wild
forms scattered over south-eastern Turkey, north
Iraq, west Iran, and Transcaucasia. These were for-
merly named T. araraticum Jakubz. [=T. armeniacum
(Jakubz.) Makush], but are now regarded as the
CEREALS 51
wild race of the domesticated crop; i.e. T. timopheevii
subsp. armeniacum (Jakubz.) van Slageren. (Please
note that when we use the traditional classification,
we refer to this wheat as T. araraticum). As already
mentioned, armeniacum forms are genetically well
isolated from both the wild and the domesticated
forms of the tetraploid turgidum aggregate.
However, morphologically they show striking simi-
larities to wild emmer, T. turgidum subsp. dicoccoides.
Moreover, in Turkey, Iran, and Iraq, both dicoccoides
and armeniacum wheats are distributed over the
same area (Map 4, p. 42) and it is practically impos-
sible to separate their ripe ears from one another
upon morphological examination (Tanaka and Ishii
1973). This can only be done by crossing and/or by
molecular tests. For these reasons taxonomists deal-
ing with the flora of south-west Asia (e.g. Bor 1968)
frequently lump all Kurdish wild tetraploid wheat
material together in what they call T. dicoccoides,
disregarding the fact that they are actually con-
fronted with two reproductively well-isolated
entities.
From the point of view of domestication and
spread of domesticated wheats, the role of the
tetraploid subsp. armeniacum GGAA stock is appar-
ently negligible. With the exception of the much
localized Georgian T. timopheevii cultivars, the chro-
mosomes of the wild armeniacum stock are absent
in the vast ensembles of cultivars and local land
races of both tetraploid and hexaploid wheats. It
could be argued that in the early days, both wild
dicoccoides and wild armeniacum wheats could have
been taken into cultivation in south-eastern Turkey,
northern Iraq, and western Iran, and that the early
non-brittle, hulled wheat remains from these places
represent both stocks. Yet the question remains: if
such alleged domesticated GGAA wheats were
indeed produced in south-west Asia, why were
they totally replaced by BBAA wheats, even among
the local land races?
Barley: Hordeum vulgare
Domesticated barley, Hordeum vulgare subsp. vul-
gare, is one of the main cereals of Mediterranean
agriculture and a founder crop of Old World
Neolithic food production. All over this vast area,
52 DOMESTICATION OF PLANTS IN THE OLD WORLD
barley is a universal companion of wheat, but in
comparison with the latter it is regarded as an infe-
rior staple and poor person’s bread. However, bar-
ley withstands drier conditions, poorer soils, and
some salinity. Because of these qualities, it has been
the principal grain produced in numerous areas,
and an important element of the human diet.
Indeed, barley has a much wider geographic distri-
bution in the Mediterranean region and south-west
Asia than wheat. Barley is also the main cereal used
for beer fermentation. Preparation of this beverage
seems to be a very old tradition (Hopf 1976; Darby
et al. 1977; von Bothmer et al. 1991; Samuel 1996).
The crop was, and still is, a most important feed
supplement for domestic animals.
Barley is an annual, predominantly self-pollinated
diploid (2n = 2x = 14 chromosomes) grain crop.
Consequently, its variation is structured in true breed-
ing lines. Hundreds of modern varieties and thou-
sands of land races are known. All cultivars have
non-brittle ears; i.e. the spikes stay intact after ripen-
ing and are harvested and threshed by humans. This
is in sharp contrast with wild barley forms, in which
ears are always brittle. Non-brittleness in domesti-
cated barley is governed by a mutation in either one
of two tightly linked ‘brittle’ genes (Bt1, Bt2, Table 7,
p. 61). The brittle wild-type allele in each locus is
dominant; the non-brittle alleles are recessive. Like
domesticated emmer, Btr1 and Btr2 are also located
on the short arm of the homoeologous group 3 chro-
mosome (3H). Many cultivars are homozygous reces-
sive for both mutations. Others carry only one
mutation (Takahashi 1964, 1972; Komatsuda and
Mano 2002). Non-brittle mutations survive only
under domestication, and non-shattering ears serve
as a reliable indicator of domestication.
Barley ears have a unique structure. They contain
triplets of spikelets arranged alternately on the
rachis (Hockett and Nilan 1985; Harlan 1995a).
According to the morphology of the spikelets,
barley under domestication can be divided into two
principal morphological types:
(i) Two-rowed forms—H. vulgare subsp. distichum,
traditionally called Hordeum distichum L.—in which
only the median spikelet in each triplet is both her-
maphrodite and fertile, and usually armed with a
prominent awn. The two lateral spikelets are males,
reduced, borne on longer stalks, and are grainless
and awnless. Each ear thus contains two rows of
grain-producing spikelets, one in each side (Fig.
15D).
(ii) Six-rowed forms—H. vulgare subsp. Vulgare,
traditionally referred to as H. hexastichum L.—in
which the three spikelets in each triplet are her-
maphrodite, bear grain, and usually all are awned.
Ears in these varieties therefore have six rows of fer-
tile spikelets, three in each side (Fig. 10E).
The two-rowed condition is primitive; it is
found in the wild progenitor of the crop as well as
in all other Old World, wild Hordeum species. Six-
rowed types of Hordeum vulgare were derived
under domestication (Zohary 1969). A recessive
mutation in the gene V and a dominant mutation
in the gene i confer fertility to the lateral spikelets
and cause the shift from two-rowed to six-rowed
ears. The genotypic constitution of two-rowed
forms is VVii. That of ordinary six-rowed culti-
vars is vvII (see Gymer 1978; Hockett and Nilan
1985).
Wild barleys, as well as the majority of the
domesticated forms have hulled grains; i.e. the
pales are fused with the grains and cover them,
even after threshing. In some domesticated varie-
ties this hulled trait is lost. The ripe grains are free
and are released, when the ears mature and dry, by
threshing. In traditional farming communities,
naked barleys were frequently favoured for the
preparation of food, whereas hulled forms are pre-
ferred for brewing beer and for animal feed. The
naked grain trait is controlled by a single recessive
gene (n).
Because of the striking differences in ear and
grain morphology, taxonomists in the past fre-
quently considered two-rowed barley forms and
six-rowed barley forms as two separate species: H.
distichum L. and H. hexastichum L. This nomencla-
ture is still used by some botanists. Others divided
the crop even further and called the forms contain-
ing fertile lateral spikelets on lax ears H. tetrastichum
Körn., and those with dense ears H. hexastichum
L. However, today we know (Zohary 1971; von
Bothmer et al. 1991) that all domesticated barleys
contain homologous chromosomes and are fully
inter-fertile. Splitting is, therefore, genetically

unjustified and the main domesticated barley types
represent races of a single variable crop complex,
H. vulgare L. (see Table 5).
Carbonized grains are the most common barley
remains found in archaeological excavations.
Occasionally whole triplets are also retrieved, mak-
ing it possible to determine whether the cereal was
of the two-rowed or six-rowed type. Six-rowed bar-
ley forms can also be distinguished from two-rowed
barleys by the morphology of the grains. In two-
rowed barleys all kernels are straight and symmet-
rical. In six-rowed barleys the lateral grains are
often slightly bent and somewhat asymmetrical (Fig
16). The position and structure of the scars of the
spikelets on the rachis internode provide an addi-
tional diagnostic trait to identify six-rowed barley
in charred remains. The grains of naked barley
(often called var. coeleste or var. nudum) are com-
monly recognized by their somewhat shriveled skin
and by the furrow that stays narrow also near the
apex (Figs 15 and 16).
In summary, remains of two-rowed barley tri-
plets in early archaeological sites can be very use-
ful for the identification of the start of barley
domestication. As with wheats, the disarticulation
scars in the wild, brittle forms are smooth, whereas
in non-brittle domestic varieties, threshing pro-
duces rough breakage scars. However, since in
wild barley the first and second lowermost triplets
(at the base of the ear) do not disarticulate easily,
rare triplets with rough scars can be produced also
upon reaping (and threshing) of wild spontaneum
stands. As argued by Kislev (1997a), claims for
barley domestication cannot be based on finds of
rare breakage scars among numerous smooth ones.
Only a prevalence of triplet remains with rough
scars is a reliable indication of domestication. For
the same reason, another good indication is finds
of non-basal rachis fragment with more than one
triplet.
Wild ancestry
The wild ancestor of the domesticated barley is well
known (Harlan and Zohary 1966; Zohary 1969). The
crop shows close affinities to a group of wild and
weedy barley forms which are traditionally grouped
in Hordeum spontaneum C. Koch, but which are, in
CEREALS 53
fact, the wild race or subspecies of the domesticated
crop. The correct name for this wild type is there-
fore H. vulgare L. subsp. spontaneum (C. Koch) Thell.
(Table 5 and Fig. 15). These are annual, brittle, two-
rowed, diploid (2n = 2x = 14 chromosomes), pre-
dominantly self-pollinated barley forms, and the
only wild Hordeum stock that is cross-compatible
and fully inter-fertile with the domesticated barley.
Vulgare x spontaneum hybrids show normal chromo-
some pairing in meiosis. Morphologically, the simi-
larity between wild spontaneum and domesticated
two-rowed distichum varieties is striking. They dif-
fer mainly in their modes of seed dispersal.
Spontaneum ears are brittle and at maturity they
disarticulate into individual arrow-like triplets.
These are highly specialized devices which ensure
the survival of the plant under wild conditions.
Under cultivation this specialization broke down
and non-brittle mutants were automatically selected
for in the manmade system of reaping, threshing,
and sowing.
The close genetic affinities between the domesti-
cated crop and wild spontaneum barleys are indi-
cated also by spontaneous hybridization that occurs
sporadically when wild and tame forms grow side
by side. Some of these kinds of hybridization prod-
ucts, combining brittle ears and fertile lateral spike-
lets, were in the past erroneously regarded as
genuinely wild types and even given a specific rank
(H. agriocrithon Åberg). Extensive isozyme, seed
storage proteins, and DNA tests have been carried
out in barley (Nevo 1992). The results confirm the
close relationships between the wild and domesti-
cated entities grouped in the H. vulgare crop com-
plex. They also clearly show that genetic
polymorphism in the spontaneum wild populations
are much wider than that present in the domesti-
cated gene pool.
Hordeum vulgare subsp. spontaneum is spread over
the east-Mediterranean basin and the west Asiatic
countries (Map 6), penetrating as far as Turkmenia,
Afghanistan, Ladakh, and Tibet. Wild barley occu-
pies both primary habitats and segetal, manmade,
habitats. Its distribution centre lies in the Fertile
Crescent, starting from Israel and Jordan in the
south-west, stretching north towards south Turkey
and bending south-east towards Iraqi Kurdistan and
south-west Iran. In this area, wild spontaneum barley
54 DOMESTICATION OF PLANTS IN THE OLD WORLD

is continuously and widely distributed. It consti-

tutes an important annual component of open her-
baceous formations, and it is particularly common
in the summer-dry deciduous oak park-forest belt,
east, north, and west of the Syrian Desert and the
Euphrates basin, and on the slopes facing the Jordan
Rift Valley. From here, H. vulgare subsp. spontaneum
spills over the drier steppes and semi-deserts.
In south-west Asian countries, wild barley also
occupies a whole array of secondary habitats; i.e.
opened-up Mediterranean maquis, abandoned
fields, and roadsides. It also infests cereal cultiva-
tion and fruit-tree plantations (Harlan and Zohary
1966). Further west, it is found in the Aegean region,
the Mediterranean shore of Egypt and Cyrenaica,
and extends further west to Algeria and Morocco.
Further east, in north-east Iran, Central Asia, and
Afghanistan, wild spontaneum barley is much more
sporadic in its distribution; it rarely builds large
stands and seems to be restricted, in most localities,
to segetal habitats, ruins, or to sites which have been
drastically churned by human activity. In general,
wild barley does not tolerate extreme cold and it is
only occasionally found above 1500 m above sea
level. It is almost completely absent from the ele-
vated continental plateaus of Turkey and Iran. On
the other hand, it is somewhat more drought resist-
ant than the wild wheats and penetrates relatively
deeply into the warm steppes and deserts.
The origins of domesticated barley are still not
fully understood. Early crossing experiments and
chloroplast DNA typing have suggested that
domesticated barley evolved from one, two, or very
few events (Zohary 1999). Later, Badr et al. (2000)
examined 400 AFLP loci in 317 wild and 57 domes-
ticated lines and found that barley is probably of a
monophyletic origin resting in the Israel-Jordan
area. However, in recent studies that included
sequencing of seven genetic loci, Morell and Clegg
(2007), and also Molina-Cano et al. (2005), Saisho
u
and Purugganan (2007) and Wang and Ding (2009)
suggested two origins: one within the Fertile
Crescent and a second independent domestication A B C 1
farther east, possibly at the eastern edge of the
Iranian Plateau. Apparently, the European and Fig. 15 Continues overleaf
north African barley were largely connected to the
Fertile Crescent while much of Asian barley is con-
nected to the eastern center.
 CEREALS 55

D H E

Fig. 15 Main types of barley, Hordeum vulgare. A–Ear of wild barley, H. vulgare subsp. spontaneum [= H. spontaneum]. The ear is structured by
seed-dispersal units (disarticulation units, spikelets) that disarticulate and shatters upon maturity. B–Seed-dispersal unit in dorsal side. C–Seed-
dispersal unit in ventral side, notice the upper (u) and lower (l) disarticulation scars. Note the ‘arrowhead-shape’ of the seed-dispersal units, with
long and sturdy awns (partially removed in C). D–Ear of domesticated two-rowed barley, H. vulgare subsp. distichum. E–Ear of domesticated
six-rowed barley, H. vulgare subsp. vulgare. F–Grain of hulled barley. G–Grain of naked barley. H–Triplet of six-rowed cultivated barley. Ears 1:1;
grains 3:1 (Schiemann 1948).
56 DOMESTICATION OF PLANTS IN THE OLD WORLD

A B C
0 5 mm

Fig. 16 Comparison between carbonized grains of wild and domesticated barleys. A–Wild barley, Hordeum vulgare subsp. spontaneum [= H.
spontaneum] from final Mesolithic Mureybit, Syria (van Zeist and Casparie 1968). B–Grains of domesticated barley, H. vulgare subsp. vulgare.
C–Grains of naked barley, H. vulgare var. nudum from Medieval Archsum, Germany (Kroll 1975). The arrow indicates lateral, twisted, grain;
characteristic of the six-rowed barley cultivar.

Archaeological evidence of approximately 260,000 grains (and 12,000 wild

oat grains) from ca. 11,700–10,550 cal BP Gilgal,
Barley first appears in several pre-agriculture or
Israel, which most probably represents early, pre-
incipient sites in south-west Asia. The remains are
domestication, farming (Hartmann 2006; Weiss
of brittle, two-rowed forms, morphologically iden-
et al. 2006). Wild barley is available from the ca.
tical with present-day wild spontaneum barley, and
9,600–8,750 cal BP Bus Mordeh phase of Ali Kosh,
apparently collected from the wild (Fig. 16). The
Iran (Helbaek 1969), the earliest layers, ca. 10,250–
earliest records of such wild barley harvest comes
9,550 cal BP, of Çayönü, Turkey (van Zeist 1972;
from ca. 50,000 BP Kebara Cave (Lev et al. 2005) and
van Zeist and de Roller 1991–2, 1995), and from ca.
from ca. 23,000 cal BP Ohalo II, a submerged Early
8,700 BP pre-ceramic Beidha, Jordan (Helbaek
Epi-Palaeolithic site on the south shore of the Sea of
1966c). At the last three sites, brittle spontaneum-
Galilee, Israel (Kislev et al. 1992; Simchoni 1998;
type barley was found in contexts showing definite
Weiss 2002, 2009; Weiss et al. 2004, 2008). In Ohalo
signs of domesticated wheat.
II, the remains of H. spontaneum were found
Remains of non-brittle two-rowed barley—
together with wild emmer wheat. Other early signs
domesticated forms that could survive only under
of H. spontaneum collection from the wild come
cultivation—came from ca. 10,200–9,550 cal BP
from ca.15,500–10,150 cal BP, Franchthi Cave,
Middle PPNB (phase II) in Tell Aswad (van Zeist
Greece, from ca. 11,800–11,300 cal BP Mureybit (van
and Bakker-Heeres 1985), and from ca. 9,450–9,300
Zeist and Casparie 1968; van Zeist and Bakker-
cal BP Neolithic Jarmo, Iraq (Helbaek 1959a, 1960;
Heeres 1986), from ca. 10,500–10,200 cal BP Tell
Braidwood 1960). In the latter site, Helbaek (1959b)
Aswad, east of Damascus, Syria (van Zeist and
was the first to show two-rowed barley remains
Bakker-Heeres 1985), and from ca. 11,700–10,550
still closely resembling wild spontaneum but also
cal BP Pre-Pottery Neolithic A, Netiv Hagdud,
displaying a non-brittle rachis. Similar finds were
north of Jericho, Israel (Kislev 1997; Hartmann
reported by Hopf (1983) in ca. 9,900–9,550 cal BP
2006; Weiss et al. 2006). An unusual find is a hoard
pre-pottery Jericho. Indicative clues come from Ali

Table 5 Taxonomy of the barley crop complex: Species according to traditional classification and their modern ranking on the basis of cytogenetic
affinities
Traditional classification
Section Cerealia Åberg within the genus Hordeum L. containing the
following species:
1. Wild two-rowed barley H. spontaneum C. Koch Brittle, hulled.
2. Domesticated two-rowed barley H. distichum L. Non-brittle, mostly
hulled.
3. Domesticated six-rowed barley H. vulgare L. [= H. hexastichum L.]
Non-brittle, both hulled and naked forms
4. Brittle six-rowed barley H. agriocrithon Åberg.
Kosh (Helbaek 1969), where the brittle spontaneum-
like material characterized the lower layers, dated
to ca. 9,600–8,750 cal BP, and in the upper strata
dated to ca. 9,400–9,250 cal BP, it was replaced by
non-brittle, broad-seeded, domestic forms.
Hulled barley has been reported in ca. 10,650–
9,550 cal BP Early PPNB Kissonerga-Mylouthkia
(Murray 2003) and Shillourokambos (Willcox 2000),
Cyprus. Their domesticated status was based on
grain size, while chaff remains are either wild-type
or badly preserved. The domestic status of these
finds requires further confirmation by chaff
remains. However, as einkorn and emmer were
found alongside with barley, and as no wild wheats
grow in the island, it seems that cereals were intro-
duced to the island from outside already at this
early stage.
Domesticated barley continues to be a principal
grain crop in the south-west Asia throughout the
Neolithic period. Its remains have been recovered
side by side with wheats, in most Neolithic sites in
which rich plant remains were retrieved. Shortly
afterwards, we are faced with more advanced
forms; i.e. six-rowed hulled as well as naked culti-
vars of barley.
Remains of six-rowed barley start to appear in
Turkey already in ca. 9,350–8,950 cal BP Aceramic
Neolithic Çatalhöyük, while in the ca. 8,950–8,350 cal
BP Ceramic Neolithic phase, it became firmly estab-
lished (Helbaek 1964a; Fairbairn et al. 2002, 2005,
2007). In ca. 8,200–7,800 cal BP Late Neolithic Hacilar,
it is represented by both hulled and naked varieties
(Helbaek 1970). In the earliest phases of Ali Kosh,
CEREALS 57
Modern grouping
A single species containing both wild and domesticated forms.
Collective name: H. vulgare L.
1. H. vulgare subsp. Spontaneum
2. H. vulgare subsp. distichum [= H. vulgare convar. distichon]
3. H. vulgare subsp. vulgare [= H. vulgare convar. vulgare]
4. Agriocrithon forms are now known to be secondary hybrid
derivatives between 1 and 3.
Iran (Bus Mordeh and Ali Kosh, ca. 9,600–9,250 cal
BP), only two-rowed barley occurs. But in the
Mohammed Jaffar phase (ca. 8,400–8,350 cal BP),
some six-rowed elements, as well as naked kernels,
appear among the otherwise two-rowed material
(Helbaek 1969). During the eighth millennium BP,
hulled six-rowed barley establishes itself as the main
cereal of the Mesopotamian basin as in ca. 7,300–7,000
cal BP Tell-es-Sawwan, Iraq (Helbaek 1964b). Hulled
barleys are common in south-west Asia also during
the Chalcolithic and Bronze Age. In these periods
they show a tendency to outnumber the wheats.
Being less sensitive to changes in climate and soils,
barley adapted more easily to extreme conditions
and probably replaced wheats on depleted soils, or
in irrigated areas suffering from salinization.
Barley was one of the principal crops that spread
grain agriculture from south-west Asia, first to the
Aegean region and subsequently to the Balkan
countries, Caucasia, west Mediterranean basin, and
central Europe, as well as west Europe, Egypt, and
Transcaucasia (Maps 1 and 2).
During the ninth millennium BP, barley emerges in
the Aegean region and Greece as a constant compan-
ion of emmer and einkorn wheats. It is represented by
both two-rowed and six-rowed forms and also by
naked varieties (practically all Neolithic naked bar-
leys are six-rowed). Such sites are Early Neolithic
Franchthi Cave (Hansen 1991a, 1992), Aceramic
Neolithic Cap Andreas-Kastros (van Zeist 1981) and
Sesklo (Hopf 1962; Kroll 1981a), Greece, Aceramic
Neolithic Dhali Agridhi, Cyprus (Stewart 1974), and
Knossos, Crete (Sarpaki 2009). In Chalcolithic and
58 DOMESTICATION OF PLANTS IN THE OLD WORLD
0 100 200 miles
0 200 400 km
Map 6 Geographical distribution of wild barley, Hordeum vulgare subsp. spontaneum [= H. spontaneum]. The area in which wild barley is widely
distributed is shaded. Dots represent additional sites, mainly of weedy forms. Wild barley extends eastwards beyond the boundaries of this map
as far as Tibet and west China (Harlan and Zohary 1966).
Bronze Age Greece, barley is again gaining in impor-
tance and frequently becomes the prevailing cereal.
At the same time, barley spread to Central Asia
ca. 8,200–7,850 cal BP Jeitun, Turkmenistan, where
the six-rowed form was found (Charles and Hillman
1992; Harris and Gosden 1996).
Together with emmer and einkorn wheats, bar-
leys emerge as one of the principal cereals of the
Linearbandkeramik farmers that started Neolithic
agriculture in central Europe in the first and second
half of the eighth millennium BP.
In eighth millennium BP, Neolithic cultures in the
Balkans (notably in Bulgaria, see Marinova 2004,
2006), central and west Europe (e.g. Araus et al.
2007; Rottoli and Pessina 2007), one encounters
mainly six-rowed forms and mostly naked grains
from early on. Barley is relatively rare in the
Linearbandkeramik culture. In some sites, naked
grains are as common as hulled ones. In central
Europe (as well as northern Europe), its importance
clearly increases in later Neolithic times and espe-
cially in the Bronze Age.
In the Impressed Ware and later Neolithic cul-
tures of the west Mediterranean basin (eighth
through to the sixth millennia BP), barley remains
are plentiful and they consist mainly of hulled and
naked six-rowed forms (Hopf 1991). It remains a
close associate of free-threshing wheats in these ter-
ritories throughout the Bronze Age.
Barley is the main companion of emmer wheat
in the Neolithic settlement of the Nile Valley in
the second half of the eighth millennium BP
(Wetterstrom 1993) and also in this area it main-
tained its important role in food production through
Neolithic and Bronze Age times. Barley also seems
to be a main element in the diffusion of south-west
Asian agriculture towards the east. It is present in
the crop assemblage in several sites in Caucasia and
Transcaucasia (Lisitsina 1984), in the first half of the
eighth millennium BP in Caucasia, and in the sec-
ond half in Transcaucasia.
In conclusion, the archaeological finds show bar-
ley as a founder crop of the south-west Asian
Neolithic agriculture and as a close companion of

emmer and einkorn wheats. It is also clear that wild
spontaneum barley is the ancestral stock from which
domesticated barley was derived. This wild barley
is a common annual plant in the Fertile Crescent
and the area of its distribution largely coincides
with the sites of the earliest finds of barley cultiva-
tion. This coincidence indicates conclusively that
south-west Asia is the place of origin of domesti-
cated barley. The archaeological remains enable us
to trace the main developments of barley under
domestication: first, the fixation of non-brittle
mutations, and subsequently, the emergence of six-
rowed hulled, and naked types. The principal role
of barley in food production in the Old World in
Neolithic and Bronze Age times is presently well
documented.
Rye: Secale cereale
Rye, Secale cereale subsp. cereale, is a characteristic
grain crop of the temperate regions of the Old
World. It is particularly appreciated in northern and
eastern Europe because of its winter hardiness,
resistance to drought, and its ability to grow on
acid, sandy soils. Thus it succeeds under conditions
in which wheat frequently fails (Evans 1995). Rye
grains contain appreciable amounts of proteins and
can be baked into dark-coloured ‘rye-bread’. Much
of the present world production of rye is consumed
in the form of bread, appreciated for its flavour and
distinctive dense texture. The grains are also used
as a high energy animal feed and for the prepara-
tion of rye whisky. The green plants are also com-
monly used for fodder. In contrast to most grain
crops that are self-pollinating, rye is a cross-
pollinated cereal. Yields depend, among other fac-
tors, on effective wind pollination.
Wild ancestry
Domesticated rye belongs to the small genus Secale
L. Its diversity (in the wild) is centered in central
and west Asia. Despite relatively large numbers of
studies performed in this genus, its taxonomy is
still somewhat uncertain. A widely accepted classi-
fication of this genus is the one proposed by Sencer
and Hawkes 1980 (Zohary 1971; Stutz 1972;
Kobylyanskyi 1989) and recently found to agree
CEREALS 59
with rDNA ITS sequence (de Bustos and Jouve 2002)
and microsatellite markers (Shang et al. 2006). The
following four biological species are now recog-
nized. All are diploids (2n = 2x = 14 chromosomes)
and are cold-tolerant.
There is wide variation within this genus, both
in the cultivars and in the wild forms—inside
populations and between them. This variability is
due to the fact that rye is largely cross-pollinated
and a perennial plant. As a result of this variabil-
ity, classification is a complicated task in this small
genus.
These are the current main four taxonomic groups
in the genus Secale (Table 6):
1. The crop complex of S. cereale L. which contains
the domesticated varieties, as well as con-specific
weedy races and wild forms. All are annual, self-
incompatible, chromosomally homologous, and
fully inter-fertile with one another. Hybrids
between the domesticated various forms are fully,
or almost fully, inter-fertile with one another and
show normal formation of seven bivalents in meio-
sis. In the past, rye taxonomists (such as Roshevitz
1947) split this complex into several species. The
domesticated varieties were grouped in S. cereale
L., while the variable weeds and wild types were
treated as separate species (S. segetale (Zhuk.)
Roshev., S. afghanicum (Vav.) Roshev., S. dighoricum
(Vav.) Roshev., S. ancestrale Zhuk., and S. vavilovii
Grossh.). Yet the results of comprehensive cytoge-
netic tests (Nürnberg-Krüger 1960a, 1960b; Khush
1963b; Stutz 1972; Vences et al. 1987) indicated that
such splitting is unjustified. The species described
are now regarded as main races or subspecies of
the crop complex (Sencer and Hawkes 1980;
Kobylyanskyi 1989; Shang et al. 2006; de Bustos and
Jouve 2002). The variable S. cereale complex is cur-
rently roughly divided into the following principal
races:
(i) Domesticated varieties: These are non-shattering
annual, self-incompatible plants with charac-
teristic large and plump grains (Fig. 17C).
(ii) Non-shattering weeds: This is a variable aggregate
of obligatory weeds infesting wheat cultivation
in Turkey, adjacent areas in Syria, Iraq, and Iran,
60 DOMESTICATION OF PLANTS IN THE OLD WORLD
as well as the Balkan countries, Caucasia, and
Transcaucasia. The mature ears of these weeds
do not shatter and they mimic wheat in grain
size and grain weight. Consequently, they are
harvested and threshed together with the wheat.
Since traditional winnowing does not separate
grains of rye from those of wheat, rye seed is
included in the harvest and planted with the
wheat in the subsequent year. Farmers, particu-
larly those in the elevated plateau of Anatolia
and Armenia, tolerate some rye-weed infesta-
tion in their wheat crop, and for a good reason:
in bad years with extreme cold and dry weather
the rye weeds survive when wheat does not—
ensuring the population a supply of what is
sometimes referred to as the ‘wheat of Allah’
(Hillman 1978).
(iii) Semi-shattering weeds: These weedy ryes are
common in north-east Iran, Armenia,
Afghanistan, and adjacent central Asian repub-
lics. Partially shattering ryes may infest wheat
and barley cultivation, particularly irrigated
fields. The rachis is semi-brittle; the upper part
of the mature ear shatters spontaneously while
the lower part stays intact and is reaped together
with the wheat crop. Different populations vary
in the degree of ear shattering. Frequently two-
thirds of the seeds produced undergoes ‘wild-
type’ dissemination and are dispersed in the
domesticated field. The remaining seeds are
taken by the farmer to the threshing floor.
(iv) Fully shattering wild types: In these forms the
rachis is fully fragile and the mature ear disar-
Table 6 Characteristics of the four taxonomic groups in the genus Secale
Species Shattering
S. cereale
(i) Domesticated varieties Non-shattering
(ii) Non-shattering weeds Non-shattering
(iii) Semi-shattering weeds Semi-shattering
(iv) Fully shattering wild types Fully shattering
S. montanum Fully shattering
S. iranicum Fully shattering
S. sylvestre Fully shattering
ticulates spontaneously into individual spike-
lets. The kernels are narrow and fully covered
by the brittle glumes. The individual wedge-
like spikelets serve as seed-dispersal devices.
The first brittle S. cereale forms were discov-
ered near Aydin, western Turkey, by the
Russian botanist P.M. Zhukovsky, and named
by him ‘S. ancestrale’(Fig. 17B). Today, this
robust rye type is known also from several
other localities in the Izmir area. But since this
rye infests fig plantations, roadsides, and vine-
yards, it is obviously a weed and not a genu-
inely wild type. Truly wild-rye forms,
chromosomally homologous with the crop,
occur in Armenia and in eastern Turkey (Map
7). They were first discovered in Armenia,
where they have been named Secale vavilovii
Grossh. Subsequently, large populations of
these brittle, annual, cross-pollinated vavilovii
forms were found also in eastern Turkey (Stutz
1972; Sencer and Hawkes 1980; D. Zohary
unpublished data). This wild rye thrives on
basaltic bedrocks. On the lower slopes of Mt
Ararat, and in similar open terrain in adjacent
volcanic areas, it frequently builds extensive
stands. Compared to the weedy and robust
ancestrale rye, vavilovii plants are somewhat
shorter (usually 50–80 cm tall), but they grow
in primary habitats and are obviously truly
wild. Since they are fully inter-fertile with the
domesticated rye (Kostoff 1937; Stutz 1972;
Sencer and Hawkes 1980) they are regarded as
a wild race (subspecies) of S. cereale.
Wild/domesticated/ Annual/perennial Pollination system
weed
Domesticated Annual Cross-pollinating
Weed Annual Cross-pollinating
Weed Annual Cross-pollinating
Wild Annual Cross-pollinating
Wild Perennial Cross-pollinating
Wild Annual self-pollinating
Wild Annual self-pollinating
 CEREALS 61

Table 7 Recessive mutations which changed wild-type trait into domestic-type trait in south-west Asian founder crops. These mutations are
responsible for the shift from wild type seed dispersal to human-dependent crops. There is no conclusive information yet regarding the situation in
bitter vetch.

Crop Wild-type trait Domestication trait Number of recessive Source

mutations involved

Einkorn wheat Shattering ears Non-shattering ears 1 Love and Craig 1924
Emmer wheat Shattering ears Non-shattering ears 2 Nalam et al. 2006
Barley Shattering ears Non-shattering ears 2 Takahashi 1955; Zohary 1960
Lentil Dehiscent pod Dehiscent pod 1 Ladizinsky 1979
Pea Dehiscent pod Indehiscent pod 1 Waines 1975
Chickpea Dehiscent pod Indehiscent pod 1 Kazan et al. 1993
Bitter vetch Dehiscent pod Indehiscent pod 2 Ladizinsky and van Oss 1984
Flax Dehiscent capsule Indehiscent capsule 1 Gill and Yermanos 1967;
Diederichsen and Hammer 1995

2. Secale montanum Guss. [= S. strictum (Persl.)
Persl.]. This rye species comprises a variable group
of perennial, cross-pollinated forms, native to ele-
vated plateaus and mountain systems in south-west
Asia (especially Turkey), as well as the Caucasus,
north Iran, the south Balkans, south Italy, Sicily, and
Morocco (Sencer and Hawkes 1980). This species
was apparently never domesticated. Perennial mon-
tanum forms have a tufted growth habit and pressed,
shattering ears (Fig. 17A, Plate 7). Both morphologi-
cally and cytogenetically, perennial montanum forms
are more distant from the crop. They differ from the
domestic and wild forms of S. cereale by two chro-
mosomal translocations (Stutz 1972; Sencer and
Hawkes 1980). However, hybrids between annual
S. cereale and perennial S. montanum are easy to
make and they are only semi-sterile. Also, S. monta-
num is morphologically very variable and includes
many eco-geographic races, such as: S. dalmaticum
Vis., S. anatolicum Boiss, S. ciliatoglume (Boiss.)
Grossh., and S. kupriyanowii Grossh.
Over considerable areas on the elevated plateaus
of Anatolia, S. montanum grows side by side with
S. cereale, the first as a common grass in the non-
arable steppe habitats, and the second as a weed
in adjacent domesticated fields. Spontaneous
hybridization between these two cross-pollinated
grasses was found to be rather frequent in such
contact places, particularly at the edges of cultiva-
tion (Stutz 1972; D. Zohary unpublished data).
Introgressive hybridization often characterizes
both species in these areas. Thus, in their main
geographic centre, S. montanum and S. cereale are
not reproductively fully isolated from one another
and they still exchange genes.
3. Secale iranicum Kobyl. This is an insufficiently
known annual wild rye. It is similar, in its general
habit, to the wild and weedy brittle forms of S. cereale.
Yet it is widely divergent from the crop chromosoma-
lly. It is also a self-pollinated species, has relatively
small seeds, and it is isolated from the crop (and other
Secale species) by strong hybrid sterility barriers. Until
now, S. iranicum is known only from two collections
made by H. Kuckuck in 1956 near Hamadan, Iran. He
erroneously identified his plants as S. vavilovii and
under that name sent seed samples to several centres
of rye research (see Kuckuck 1973), causing confusion
among researchers as to the nature of S. vavilovii. In
fact, the genetic tests performed by Nürnberg-Krüger
(1960a, 1960b), Kranz (1961), Khush (1963a, 1963b),
Pérez de la Vega and Allard (1984), and Vences et al.
(1987)—assumed to have been carried out on
S. vavilovii—were actually performed on this totally
different taxon. Only Kostoff (1937), Stutz (1972), and
Sencer and Hawks (1980) used genuine S. vavilovii in
their tests. It seems that Kuckuck’s collections repre-
sent an additional, distinct Secale species. Indeed it
was described as such by Kobylyanskyi (1989).
62 DOMESTICATION OF PLANTS IN THE OLD WORLD
0 100 200 miles
0 200 400 km
Map 7 Geographical distribution of brittle wild rye, Secale cereale subsp. vavilovii [= S. vavilovii] (based on Stutz 1972; Sencer and Hawkes
1980).
4. Secale sylvestre Host. This is an annual, self-
pollinating rye, with characteristic long awns,
native to the Aralo-Caspian basin. Morphologically,
S. sylvestre is well separated from the former three
rye species. It also differs from S. cereale by three
chromosomal translocations, and from S. montanum
by a single one. It is largely inter-sterile with both
these species.
Archaeological evidence
Very few remains of Secale have been discovered in
the Neolithic and Bronze Age settlements in south-
west Asia. This is surprising since Turkey, Armenia,
and Iran harbour a wealth of wild and weedy forms
of S. cereale, and charred grains (Fig. 18) of this cereal
can be easily identified.
The earliest remains come from Epi-Palaeolithic
sites in the Upper Euphrates valley in northern
Syria. Numerous charred grains, later identified as
a mixture of both wild rye and wild einkorn wheat
(Fig. 10), were retrieved in ca. 11,800–11,300 cal BP
Tell Mureybit (van Zeist and Casparie 1968; van
Zeist and Bakker-Heeres 1986; Willcox and Fornite
1999). The narrow shape of the kernels (Fig. 10)
indicates that they represent wild forms. Similar
narrow, wild-type rye grains (either of Secale cereale
subsp. vavilovii or of S. montanum) were found in the
ca. 12,700–11,100 cal BP Epi-Palaeolithic Tell Abu
Hureyra (Hillman 1975, 2000a; Hillman et al. 1989,
2001). On the basis of grain morphology, Hillman
et al. (2001) suggested these were domesticated
form, a view that was later criticized, because of
lack of chaff and problematic dating (e.g. Nesbitt
2002). Wild rye grains also discovered in two PPNA
northern Syrian sites, ca. 11,500–11,000 cal BP, Jerf el
Ahmar (Willcox 2002; Willcox et al. 2008, 2009), and
ca. 10,700–10,400 cal BP Djade el Mughara (Willcox
et al. 2008). Later, domesticated rye was found in ca.
9,450–8,450 cal BP PPNB Can Hasan III (Hillmam
1972, 1978) and in very small quantities at the
nearby ca. 9,350–8,950 cal BP site of Aceramic
Neolithic Çatalhöyük East, Turkey (Helbaek 1964a;
Fairbairn et al. 2002, 2005, 2007). At Can Hasan III
 CEREALS 63

A
Fig. 17 Ears and grains of wild and domesticated ryes. A–Perennial, brittle, Secale montanum. B–Annual, brittle (or semi-brittle), S. cereale subsp.
ancestrale. C–Domesticated, non-brittle, S. cereale subsp. cereale, variety Petkus. Ears 1:1, grains 3:1 (Schiemann 1948).

relatively plump grains were discovered, together
with some non-brittle rachis segments. As argued
by Hillman (1978), these finds suggest that in
Anatolia, rye had already entered cultivation in
early Neolithic times, either as a non-brittle obliga-
tory ‘ryeweed’ infesting wheat fields, or as a full-
fledged domesticated cereal crop. Yet, no additional
rye remains have been discovered in other Neolithic
south-west Asian sites. The next record comes from
ca. 4,000 BP Bronze Age levels of Alaca Höyük in
north-central Anatolia (Hillman 1978). Here a pure
hoard of carbonized large grains of S. cereale was
64 DOMESTICATION OF PLANTS IN THE OLD WORLD

B C

Fig. 17 Continued.
 CEREALS 65

discovered, indicating that at that time rye was

grown as a crop in its own right.
Outside the Fertile Crescent, early information on
rye is fragmentary (for review see Behre 1992). The
earliest convincing finds in Europe are from the fol-
lowing sites: (i) Sammardenchia and adjacent Early
Neolithic sites (ca. 7,550–6,450 cal BP), northern
Italy (Pessina and Rottoli 1996; Rottoli 2005; Rottoli
and Pessina 2007). (ii) Middle Neolithic, Bükk cul-
ture (ca. 6,950–6,650 cal BP) Šarišské Michalˇany-
Fedelemka, Slovakia (Hajnalová 1993; Hajnalová
and Hajnalová 2004). (iii) Several late Neolithic
Funnel Beaker (TRB) culture sites in north, central,
and south Poland (Giżbert 1960; Wasylikowa et al.
1991). Only a limited number of rye grains were
found in the Polish sites compared to the abundance
of wheat or barley grains. (iv) Few contemporary
sites in Rumania, particularly Gumelnitza a culture
(second half of the sixth millennium BP) Mǎgura
Coneşti (Cârciumaru 1996) where a hoard of some
1000 charred kernels was discovered, suggesting
that rye was grown as a crop, and ca. 6,650–6,350 cal
BP Poduri (Cârciumaru and Monah 1985; Monah
and Monah 2008) 0 5 mm
Further evidence on rye cultivation in Europe
comes from several Bronze Age settlements in the Fig. 18 Carbonized grains of domesticated rye, Secale cereale,
Czech Republic and Slovakia where Tempír (1966, Medieval Archsum, Germany (Kroll 1975).
1969) discovered rich remains of carbonized grains.
Bronze Age records of rye are available also from
Rumania (Cârciumaru 1996); Austria (Kohler- In summary, the available evidence from the liv-
Schneider 2001, 2003); and from Moldavia and the ing plants points to the annual, brittle rye [= Secale
Ukraine (Wasylikowa et al. 1991). But in most cases cereale subsp. vavilovii] as the wild ancestor of the
they represent rye grains contaminating other domesticated crop. Eastern Turkey and adjacent
cereals. Somewhat later, rye appears in Iron Age Armenia seem to be the probable place of origin.
settlements in Germany (Hopf 1982, Table 1), The archaeological record supports this notion. The
Denmark (Helbaek 1954), Poland (Willerding earliest sign of rye, associated with agriculture,
1970), and Crimea (Januševič 1978)—usually comes from central Anatolia. Compared to wheat
admixed with barley or wheat. Also in Hasanlu, and barley, remains of rye in archaeological excava-
Iran (Tosi 1975), S. cereale appears at the end of the tions are discouragingly few. Additional finds are
fourth millennium BP and seems to have been a necessary in order to establish the place and pattern
staple crop throughout the Iron Age. Rye was part of mode of origin, and mode of the spread of the rye
of Roman grain agriculture and was grown in the crop.
cooler northern provinces. Carbonized rye grains The available evidence also suggests that rye
have been retrieved from several Roman frontier evolved first as a tolerated weed, and was only later
sites along the Rhine and the Danube (Hillman picked up as a crop. The data also indicate that vari-
1978), as well as from the British Isles (Jessen and ation build-up in this cereal and the impressive
Helbaek 1944). evolvement of weedy ryes could have been consid-
66 DOMESTICATION OF PLANTS IN THE OLD WORLD
erably enhanced by introgressive hybridization
with perennial Secale montanum—the common wild
rye element distributed over the elevated, continen-
tal parts of Anatolia and adjacent areas in south-
west Asia.
The recent discovery of early Neolithic rye in
Italy and Middle Neolithic in Slovakia supports the
view that, despite earlier assumptions, rye arrived
in Europe not through the Caucasus but rather via
the Aegean Basin and the south Balkan.
Common oat: Avena sativa
Avena sativa L., common oat, is a major cereal crop,
in traditional Old World grain agriculture. It is a
close companion of wheats and barley. The crop
succeeds well in moist climates of temperate lati-
tudes as well as in summer-dry Mediterranean con-
ditions. In north-west Europe, oat frequently thrives
better than wheat and is cultivated as a principal
crop (Leggett and Thomas 1995). The nutritive
value of oat grain is high; it contains about 15–16%
protein and 8% fat. The crop serves as a staple in
human diet and as a high energy supplement for
farm animals.
Three cytogenetically distinct stocks of Avena
L. occur under domestication. Each had an inde-
pendent origin (Zohary 1971). Yet, only one species,
common oat, A. sativa (hexaploid, 2n = 6x = 42 chro-
mosomes), established itself as a principal cereal.
Two other oats (A. strigosa Schreb. and A. abyssinica
Hochst.) are minor crops of negligible significance
The main oat crop, A. sativa, is very variable. It
comprises numerous contrasting domesticated
varieties, as well as weedy races and truly wild
forms. All are inter-fertile with one another and
share the same hexaploid genomic constitution
(Loskutov 2001). All are characterized by non-
shattering panicles. According to their response to
threshing, they are placed in three taxonomic ‘spe-
cies’ that are actually only races of the same com-
plex crop. Most cultivars are hulled; i.e. their grains
remain invested by the pales. In some of these
hulled forms (A. sativa in the narrow sense), thresh-
ing results in pressure breakage of the spikelet’s
rachilla at the base of the upper floret. The rachilla
segment remains attached to the lower floret. In
other sativa cultivars (traditionally named A. byzan-
tina C. Koch), the rachilla segment breaks at its base
and remains attached to the upper floret. The basal
floret shows an abscission scar. In several hexaploid
cultigens (frequently called A. nuda L.) the grains
are free and threshing releases the naked kernels.
Nakedness is a derived trait and occurs only under
domestication.
Wild ancestry
Avena L. is a Mediterranean genus comprising some
twenty-five annual species. The majority of the oat
species are diploid (2n = 2x = 14 chromosomes),
seven are tetraploids (2n = 4x = 28 chromosomes),
and six are hexaploid (2n = 6x = 42 chromosomes)
(Leggett and Thomas 1995). The domesticated sativa
oats (subsp. sativa) show tight genetic affinities and
close morphological similarities to a group of wild
and weedy oats, which are widely distributed over
the Mediterranean basin, among them are A. sterilis
L. (Fig. 19), A. ludoviciana Dur., A. occidentalis Dur.,
and A. fatua L. These wild types are also hexaploid
and contain chromosomes homologous to those
present in the cultivars (Loskutov 2008). Moreover,
all wild and domesticated hexaploid oats are inter-
fertile and occasionally cross in nature. Because of
these close affinities, sterilis, occidentalis, and fatua
oats are now recognized as the wild races of the
domesticated crop and are placed within the A.
sativa crop complex (Malzew 1930; Ladizinsky and
Zohary 1971; de Wet 1981). The sativa complex is,
however, chromosomally distinct and reproduc-
tively isolated from all other Avena species.
Molecular studies have also confirmed this. A set of
413 AFLP bands tested in the 25 Avena species indi-
cate that the sativa complex is a distinct group (Fu
and Williams 2008), which is characterized as the
only hexaploid member in this genus.
Zhou et al (1999) examined several A. sterilis
accessions from the Iran–Iraq–Turkey region and
cultivated accessions, and found close association
between them based on 248 polymorphic RAPD
markers. Along with Jellen & Beard (2000), they
divided the hexaploid group using the 7C-17 sepa-
ration translocation and suggested at least two
independent paths of domestication: one from A.

sterilis (with the translocation) to A. sativa, and one
from A. sterilis to A. byzantina (without it). Their
conclusion was that A. sativa and byzantina are dis-
tinct races of the hexaploid biological species and
was domesticated independently of one another.
Just as in wheats and barley (see above), domes-
tication brought about a breakdown of the origi-
nal way of seed dispersal. Wild and weedy oats
produce highly specialized drill-type diaspores,
with characteristic kinked awns, in order to dis-
seminate their seed and to insert them into the
ground. They shed their seed immediately after
maturation. As already noted, domesticated vari-
eties have non-shedding panicles. Reduction of
the awns and the evolution of relatively compact
panicles, are additional conspicuous develop-
ments under domestication.
Two distinct modes of seed dispersal operate in
the wild members of the A. sativa complex, and
serve as a diagnostic trait for their evaluation. Most
widespread are oats in which the spikelet shows a
single disarticulation point at the base of the lower
floret. Consequently, the whole spikelet (minus the
glumes), with two to three invested kernels, consti-
tute a ‘drill-type’ dispersal unit (Plate 8). This fruit-
ing type was traditionally named A. sterilis. In other
forms (conventionally placed under A. fatua) the
rachilla of the spikelet disarticulates at each node.
The florets, each with a single seed, disperse
individually.
Sterilis-type oats widely colonize the Mediterr-
anean basin from the Atlantic coast of Morocco and
Portugal in the west to the Zagros Mountains in the
east. In south-west Asia, they frequently grow
together with wild wheats and barley. In addition to
massive occupation of primary habitats, sterilis oats
colonize abandoned cultivated ground all over the
Mediterranean region aggressively, and they grow
as a noxious weed in wheat and barley fields,
orchards, and roadsides. Most conspicuous is the
variation in the size of the spikelet, its hairiness, and
colour. Forms with smaller spikelets are sometimes
referred to as A. ludoviciana Dur., and those with
larger spikelets and three to four fertile florets are
called A. macrocarpa Mönch.
Fatua oat varieties are distinctively weedy. They
are widely distributed over the whole belt of Old
CEREALS 67
World agriculture where they infest cereal fields
and grow at the edges of cultivation. Only rarely do
they occupy primary habitats. Fatua forms also
thrive in colder, more continental climates. In ele-
vated mountain cultivation and on the northern
fringes of west Asiatic grain agriculture, fatua plants
sometimes replace sterilis oats entirely.
Archaeological evidence
The only sign of oat cultivation in south-west Asia
came from an unusual find of some 12,000 Avena
sterilis grains, no chaff, from ca. 11,700–10,550 cal
BP PPNA Gilgal, Israel. This hoard also contains
about 260,000 grains of wild barley, and probably
represents pre-domestication cultivation as part of
a barley field (Hartmann 2006; Weiss et al. 2006).
Otherwise, there is no sign of oat cultivation or
domestication in Neolithic or Bronze Age sites in
south-west Asia and the Mediterranean basin.
This, in spite of the fact that wild sterilis and fatua
forms are massively distributed over these territo-
ries and even grow together with wild wheats and
wild barley. The few oat remains retrieved from
Neolithic Fertile Crescent and European sites seem
to represent only shattering wild or weedy sterilis
or fatua forms (Fig. 19). Definite indications of
domestication; i.e. remains of non-shattering sativa
or byzantina plants with their characteristic plump
seed (Fig. 20) appear first in Europe, but only in
the fourth and third millennia BP contexts
(Willerding 1970, pp. 345–6; Villaret-von Rochow
1971). The earliest sites to report A. sativa finds,
although rare, are ca. 7,600–7,400 cal BP, Sacarovca,
Moldavia (Januševič 1984; Kuzminova et al. 1998;
Monah 2007b,), and ca. 6,650–6,350 cal BP Poduri,
Rumania (Cârciumaru and Monah 1985; Monah
and Monah 2008). Furthermore, Füzes (1990, as
reported by Gyulai 2007) identified impressions
with some grains of naked oat (Avena cf. nuda)
from ca. 7,000–6,300 BP Aszód-Papi Földek,
Hungry. Later, domesticated oats are present in
fourth millennium BP Nitriansky Hrádok, Czech
Republic (Tempír 1969; Kühn 1981), Slovakia
(Tempír 1966), and ca. 2,900–2,800 cal BP
Forsandmoen and adjacent sites in Norway
(Bakkevig 1982, 1995).
68 DOMESTICATION OF PLANTS IN THE OLD WORLD

A B

0 2 cm

Fig. 19 Wild oat, Avena sativa subsp. sterilis [= A. sterilis]. A–Fruiting panicle. B–Seed-dispersal unit (spikelet without glumes) (Malzew 1930,
plates 86, 88).

0 Tzvelev [= P. spontaneum Lyssov ex Zhuk.]. Recently
5 mm
Fig. 20 Carbonized grains of domesticated oat, Avena sativa,
Medieval Archsum, Germany Kroll 1975 (Kroll 1975).
Domestic oat becomes much abundant in central
and western European contexts from the Roman
Period onward.
In conclusion, the data available from the archae-
ological excavations, as well as the evidence
obtained on the ecology and distribution of the
wild relatives, support the notion that domesti-
cated A. sativa should be regarded as a secondary
crop. Probably, oat started its evolution under
domestication not as a crop but by evolving weedy
types that infested wheat and barley cultivation.
Only later were such weeds picked up and planted
intentionally. In the temperate periphery of Old
World agriculture, oats not only supplemented
wheat and barley, but also evolved into a principal
grain crop.
CEREALS 69
Broomcorn millet: Panicum
miliaceum
Broomcorn millet (common millet, Proso millet),
Panicum miliaceum L. (Plate 9), ranks among the har-
diest cereals. It is a warm-season crop which stands
up well to intense heat, poor soils, and severe
droughts, completing its life cycle in a very short
time (60–90 days) and succeeding in areas with
short rainy seasons. This is the true millet of classic
times (the Romans’ milium and the Hebrews’
dokhan). Today, P. miliaceum is grown mainly in east-
ern and central Asia, in India and (sparsely) in
south-west Asia. The de-husked grains are boiled
and cooked like rice, or ground for the preparation
of porridge. They are quite rich (10–11%) in pro-
teins. The seeds are also used as birdfeed. Broomcorn
millet is tetraploid (2n = 4x = 36 chromosomes). It is
mainly self-pollinated.
The wild ancestor of the domesticated P. mil-
iaceum has not been satisfactorily identified. Weedy
forms of this millet are widespread in central Asia,
from the Aralo-Caspian basin in the west to
Sinkiang and Mongolia in the East. They are
referred to as P. miliaceum subsp. ruderale (Kitag.)
these weeds, with their characteristic shattering
panicles, also spread to central Europe and north
America (Scholz 1983). Probably, the vast semi-dry
areas in central Asia harbour not only weedy, but
also genuinely wild miliaceum forms. Lu et al (2009)
have identified phytoliths and biomolecular com-
ponents of certain P. miliaceum types dated to the
eleventh to ninth millennia BP. However, there are
still large complexities and points of debate regard-
ing the crop/weedy type status of any findings of
P. miliaceum (Hunt et al. 2008). Additional archaeo-
botanical finds and taxonomical studies are neces-
sary in order to establish the origin and dispersal
patterns of this crop.
Archaeological evidence
Identifying Panicum miliaceum remains and differenti-
ating it from those of Setaria italica can be problematic.
Hunt et al. (2008), compiled a comprehensive review
of early, pre-7,000 cal BP, finds of both Panicum and
Setaria finds from archaeological sites across Eurasia.
70 DOMESTICATION OF PLANTS IN THE OLD WORLD

They also referred to issues of taphonomy and identi-

fication criteria offered so far, as well as the differen-
tiation between wild/weed/crop status (see there).
The earliest archaeobotanical P. miliaceum grains are
reported from a number of early Neolithic sites in
north China, e.g. 7,670–7,610 cal BP Xinglongwa
(Zhao 2011) and ca. 8,060–7,750 cal BP, Yuezhuang site
(Crawford et al. 2006). No early finds are available
from central Asia. However, remains of millet have
been reported from several Neolithic sites dated to
the first half of the eighth millennium BP, such as ca.
8,000–7,150 cal BP Arukhlo 1 and Arukhlo 2, Georgia
(Januševič 1984; Lisitsina 1984; Schultze-Motel 1988a), A
ca. 7,600–7,400 cal BP Sacarovca, Moldavia (Januševič
1984; Monah 2007b; Kuzminova et al. 1998), and ca.
7,650–7,400 cal BP Mohelnice, Czech Republic
(Opravil 1979, 1981; Kühn 1981). Broomcorn millet is
also reported from seventh millennium BP VI strata
in Tepe Yahya, Iran (Costantini and Costantini-Biasini
1985).
As Hunt et al. (2008) indicated, it is too early to
determine whether these were locally domesticated
or represent early east–west movement of a domes-
ticated crop. However, the Georgian finds come
B
from sites situated not far away from the geographic
belt where P. miliaceum grows wild.
0 5 mm
Finds of the characteristically small oval seeds
(Fig. 21A) of P. miliaceum continue to appear in late
eighth and from the seventh millennia BP settle- Fig. 21 Carbonized grains of: A–Broomcorn millet, Panicum
miliaceum; B–Foxtail millet, Setaria italica. Bronze Age Kastanas,
ments in east and central Europe. The nature of its
Greece (Kroll 1983).
appearance is geographically sporadic throughout
the Linearbandkeramik and later sites, but became
more common in late Neolithic and in Bronze Age
cultures. Some of these finds are ca. 7,250–6,650 cal Gumelnitsa culture contexts Morteni, Rumania
BP Zánka-Vasútállomás, Hungary (Füzes 1990, (Cârciumaru 1996, p. 91), and in ca. 5,900–5,700 cal
1991), ca. 6,500–6,000 cal BP Rivne, Ukraine BP Gomolava, Macedonia (van Zeist 1975, 2003).
(Pashkevich 2003), ca. 6,150 BP Domica (= Kesevo) Somewhat later, lumps of pure charred grains were
in Czechoslovakia (Fietz 1936; Tempír 1969), and in retrieved from the Funnel Beaker (TRB) strata in
seventh millennium BP Sesklo, Greece (Hopf 1962; Szlachcin, Poland (Wasylikowa et al. 1991). The
Kroll 1981b). earliest evidence for broomcorn millet in Austria
Van Zeist (1980) suggested that P. miliaceum came from ca. 5,050–4,750 cal BP late Neolithic
reached the Mediterranean basin from the north or Anzingerberg/Hundssteig (Kohler-Schneider 2007;
the north-east. The find from ca. 6,300–4,600 cal BP Kohler-Schneider and Caneppele 2009). At about
Monte Còvolo, Italy (Pals and Voorrips 1979), indi- the same time, broomcorn millet arrived to ca.
cates that this was a rather early movement. 4,800–4,200 cal BP Crasto de Palheiros, Portugal
Remains of P. miliaceum also occur in Tripolye cul- (Pinto da Silva 1976).
ture (ca. 6,000–4,750 cal BP) sites such as Soroki in During the Bronze Age, it appears in ca. 1,850–800
the Ukraine (Januševič 1976, 1978), in ca. 5,500 BP cal BC Zürich, Switzerland (Jacomet 1988, 2004;

Jacomet et al. 1989; Brombacher and Jacomet 1997;
Favre 2002; Brombacher et al. 2005), and to ca. 905–
869 BC Grésine (Bouby and Billaud 2001), and later
in Ouroux-Marnay (Hopf 1985), France. In the
north, the earliest finds include ca. 1,400–950 cal BC
Lindebjerg and Voldtofte, Denmark (Rowley-
Conwy 1979, 1983), and ca. 1,150–950 cal BC Borge
Vestre, Norway (Sandvik 2007, 2008).
This crop was also discovered from ca. 2,350–2,100
cal BC Harappan Shortughai, Afghanistan (Willcox
1991), and a large quantity of charred grains was
uncovered in ca. 1,900–1,550 BC Bronze Age
Haftavan, Iran (Nesbitt and Summers 1988). The
first known deposit of identifiable grains in Iraq
comes from ca. 700 BC Nimrud (Helbaek 1966b).
Signs of the cultivation of P. miliaceum in the
Levant appear relatively late – Middle Bronze Age
Tell Mozan, Syria (Riehl 2000), and ca. 1,200–500 BC
Iron Age Deir Alla, Jordan (Neef 1989).
In conclusion, the evidence from the living plants
and the archaeological remains is still fragmentary,
but it does show that broomcorn millet is a rela-
tively old crop, and that it does not belong to the
south-west Asian Neolithic crop assemblage. It
may be that P. miliaceum is an eastern or central
Asian element that was picked up and added to
wheats and barley agriculture soon after its arrival
and establishment in these areas. The scanty data
available do not rule out the possibility that P. mil-
iaceum (together with foxtail millet, below)
represents an independent experiment in domes-
tication, both in East Asia and Caucasian-European
region.
Foxtail millet: Setaria italica
Foxtail or Italian millet, Setaria italica (L.) P. Beauv.
(Plate 9), is a principle founder crop of Chinese
grain agriculture. Genetic evidence suggests it was
domesticated independently in other parts of Asia
(Fukunaga et al. 2005). Italian millet and broomcorn
millet are the major crops in Chinese dry-land agri-
culture of a wide loess area along the Yellow River,
from southern Mongolian Steppe in the north and
the Huai River in the south (Zhao 2011). It is widely
cultivated today in India, China, and Japan, while it
is much rarer in European and south-west Asian
agriculture. This is a diploid (2n = 2x = 18 chromo-
CEREALS 71
somes), predominantly self-pollinated millet with
characteristic dense, bristle-bearing panicles, and
small, oval grains which are tightly enclosed by
their pales (Prasada Rao et al. 1987; de Wet 1995).
Similar to the broomcorn millet (p. 69), Foxtail mil-
let is a warm-season cereal. It survives well under
dry conditions and completes its growth cycle in a
relatively short time.
The wild progenitor of foxtail millet is well iden-
tified. Domesticated S. italica shows close morpho-
logical affinities to, and is inter-fertile with, wild S.
viridis (L.) P. Beauv., a common variable summer
weed widely spread across Eurasia (de Wet et al.
1979). Molecular markers and components have
revealed that the origin of foxtail millet domestica-
tion was probably in China (Le Thierry d’Ennequin
et al. 2000; Lu et al. 2009) with the possibility of a few
additional domestication events elsewhere in
Eurasia (Fukunaga et al. 2005). The spontaneous
and the domesticated foxtail millets differ from one
another mainly in their seed-dispersal biology. Wild
and weedy forms shatter their seed while the culti-
vars retain them. Genetic tests indicate that the shift
is governed by two complementary recessive muta-
tions (Prasada Rao et al. 1987). Under domestica-
tion, foxtail millet underwent changes in plant
habit, including a reduction of the number of
flowering tillers, and enlargement of the flowering
panicles (de Wet 1995).
Archaeological evidence
Setaria italica is the principal grain crop of the
Neolithic agriculture in north China (Ho 1977;
Zhimin 1989; Crawford 1992; Zhao 2011).
Identifying Setaria italica remains, and differentiat-
ing it from those of Panicum miliaceum, can be
problematic. Hunt et al. (2008) compiled a compre-
hensive review of early, pre-7,000 cal BP finds of
both Panicum and Setaria from archaeological sites
across Eurasia. They also referred to issues of
taphonomy and identification criteria offered so
far, as well as the differentiation between wild/
weed/crop statuses. The earliest archaeobotanical
Setarica italica grains are reported from a number
of early Neolithic sites in north China, e.g. 7,670–
7,610 cal BP Xinglongwa (Zhao 2011) and ca.
8,060–7,750 cal BP Yuezhuang site (Crawford et al.
72 DOMESTICATION OF PLANTS IN THE OLD WORLD
2006). Zaho (2011) stresses that broomcorn millet is
always more abundant than foxtail millet in these
early sites (which is true for later sites and regions
as well). However, as indicated by Hunt et al.
(2008), unlike broomcorn millet which occurs in
pre-7,000 cal BC sites both in western Asia and in
Europe, foxtail millet occurs, at this time, only in
northern China.
Soon afterwards, S. italica emerges as the prin-
cipal cereal of the Yang-shao culture (seventh and
fifth millennia BP). The finds demonstrate that in
Yang-shao times, foxtail millet was occasionally
accompanied by broomcorn millet, P. miliaceum
(p. 69). Both millets continue to play a major role
in north China’s food production today (Ho
1977).
In Europe, carbonized seeds (which were defi-
nitely assigned to S. italica) first appear in the sec-
ond millennium BC Bronze Age settlements in
central Europe, like ca. 1,850–800 cal BC Zürich,
Switzerland (Netolitzky 1914; Jacomet 1988, 2004,
Jacomet et al. 1989, Brombacher and Jacomet 1997;
Favre 2002; Brombacher et al. 2005) and in ca. 905–
869 cal BC Grésine, France (Bouby and Billaud
2001). This millet is also reported from Late Bronze
Age Kastanas, Macedonia, Greece (Kroll 1983; see
Fig. 21). Its earliest evidence in Austria came from
Late Bronze Age ca. 1,200–700 cal BC Stillfried
(Kohler-Schneider 2001, 2003). Additional finds are
available from Iron Age Europe. The earliest defi-
nite evidence for foxtail millet cultivation in south-
west Asia comes from Iron Age (ca. 600 BC) Tille
Höyük, south-east Turkey (Nesbitt and Summers
1988). Here a large quantity of pure grains was
uncovered, indicating the use of foxtail millet as a
crop.
The available archaeological evidence indicates
that foxtail millet is a relatively old domesticant. It
was probably first taken into cultivation in north-
east Asia. Together with rice and common millet,
they served as founder crops in the development
of agriculture in north China. Like common mil-
let, S. italica does not belong to the Neolithic
south-west Asian crop assemblage. Moreover, it
appears in Europe rather late. It is still an open
question whether S. italica was taken into cultiva-
tion only in north China or also elsewhere in this
vast area.
Latecomers: sorghum and rice
Two additional principal cereals—sorghum, Sorghum
bicolor (L.) Moench, and rice, Oryza sativa L.—are not
indigenous to south-west Asia and the Mediterranean
basin. They are latecomers that probably arrived in
these areas as fully developed crops only in Greek
and Roman times, or even later.
Sorghum
Sorghum bicolor (L.) Moench is a warm-weather
grain crop grown extensively in Africa, south-west
Asia, and the Indian subcontinent. Both sorghum
and pearl millet are able to tolerate very arid condi-
tions and have a major role in food production is
these areas. Wild forms closely related to the varia-
ble sorghum crop are confined to Africa south of the
Sahara (including Yemen). This indicates that this
grain crop must have been domesticated in this
area, probably in the Savanna belt south of the
Sahara (de Wet et al. 1976; Harlan 1992b; Wetterstrom
1998). Sorghum is a diploid (2n = 2x = 2 chromo-
somes), predominantly self-pollinated crop. During
domestication, it underwent considerable differen-
tiation. Five principal domesticated races have been
recognized in this crop (Harlan and Stemler 1976).
The wild relatives of the crop are very variable.
Four principal eco-geographical wild races have
been recognized in S. bicolor (de Wet et al. 1976, Map
1), as well as segetal forms (including companion
weeds) that frequently infest sorghum cultivation.
The wild and weedy forms are also diploid and pre-
dominantly self-pollinated. All are inter-fertile with
one another as well as with the cultivars. Using
cytological techniques and molecular markers such
as mitochondrial ITS variation across all twenty-
five species of Sorghum, the crop and its wild pro-
genitors were grouped into a distinct phylogenetic
lineage of Sorghum (Dillon et al. 2001; Price et al.
2005). Similar to rice (pp. 73–74), wild and weedy S.
bicolor frequently hybridize with their domesticated
counterparts. For this reason, in Africa, one is con-
fronted with a variable crop complex comprising:
(i) wild forms occupying more stable habitats; (ii)
cultivars grown by the farmers; and (iii) weeds col-
onizing secondary habitats and infesting sorghum
cultivation. The finds in pre-agriculture (ca 8,000

BP) Nabta Playa, south Egypt, (Wasylikowa and
Kubiak-Martens 1995) have shown that seeds of
wild S. bicolor were collected from the wild before
its domestication.
The archaeological exploration of sub-saharan
Africa is in its early stages, and we lack critical
information for determining when and where sor-
ghum could have been taken into cultivation. So far,
remains of African domesticated sorghum are avail-
able only from late contexts. The richest finds, pre-
served by desiccation, came from Qasr Ibrim,
Egyptian Nubia (Rowley-Conwy 1991; Rowly-
Conwy et al. 1999; Clapham and Rowley-Conwy
2007). Remains of local wild forms such as S. bicolor
(L.) Moench subsp. arundinaceum (Desf.) de Wet &
Harlan, occur in contexts dated 800–600 BC, while
domesticated sorghum, conforming in its morphol-
ogy to race bicolor cultivars, abound in contexts
dated from 100 AD to 1,200 AD. A bouquet of ripe
panicles, dated 42–640 AD, intermediate in its mor-
phology between bicolor and durra cultivars, was
also retrieved, and by 1,200 AD the more advanced,
free-threshing durra forms make their appearance.
The earliest archaeological evidence on sorghum
cultivation, available up to date, does not come
from Africa but from the Indian subcontinent.
Reports about the presence of sorghum are availa-
ble from about a dozen of second millennium BC
sites in Pakistan and India (see review by Fuller
2002). As Fuller remarks, in some of these cases, the
identification is problematic. However, the remains
uncovered from several other locations definitely
belong to this crop. Since Sorghum bicolor does not
grow wild in India, these finds have been inter-
preted (see Vishnu-Mittre and Savithri 1982; Harlan
1992b) as indicating: (i) an even earlier domestica-
tion in Africa, and (ii) an early migration of domes-
tic sorghum, from East Africa into the Indian
subcontinent. This interpretation got further sup-
port from the fact that several other African grain
crops, namely: pearl millet Pennisetum glaucum (L.)
R. Br., cow pea Vigna unguiculata (L.) Walp., and
hyacinth bean Lablab purpureus (L.) Sweet, show
similar patterns of migration. Their wild progeni-
tors are restricted to Africa. They emerge (as crops)
in the Indian subcontinent already in the second
millennium BC. Further, this view of early spread
CEREALS 73
was supported by the recent finds of ca. 2,500 BC
domesticated pearl millet in Mali, west Africa
(Manning et al. 2011).
Sorghum seems to have arrived late in south-west
Asia and the Mediterranean basin, even after the
medieval period, in light of its absence from medi-
eval sites in Syria (Samuel 2001: 429). As late as
Roman times, this cereal was almost unknown in
the belt of Mediterranean agriculture. Although
reported (as a crop) from Qasr Ibrim, from 100 AD
onwards (Rowley-Conwy 1991), sorghum cultiva-
tion seems to have stopped in Nubia and did not
establish itself in Lower Egypt. The advanced durra
cultivars which now characterize traditional sum-
mer cropping in south-west Asia, in India and in
Pakistan, are not widespread in Africa. There, they
are largely confined to the fringes of the Sahara and
to Ethiopia. For these reasons Harlan and Stemler
(1976) speculated that the more advanced, free-
threshing durra forms evolved in India from more
primitive bicolor material brought from east Africa,
and that durra type sorghum was introduced to
south-west Asia and north-east Africa rather late.
However, this is just an attractive speculation. As
argued by Rowley-Conwy et al. (1997, 1999), a
sounder appreciation of sorghum domestication
would be possible only when its start in Africa will
be better understood.
Rice
Oryza sativa L. is an east and south Asian element. It
is the domesticated founder crop in this part of the
World (Chang 1995) and is one of the major food
suppliers to the human race. Rice is a diploid
(2n = 4x = 24 chromosomes; genomic constitution
AA), predominately self-pollinated. This is a very
productive crop when grown in paddies, where
fixed nitrogen is naturally provided by co-habiting
blue-green algae. In the course of its long and com-
plicated history of cultivation, Asian rice under-
went considerable differentiation. Thousands of
cultivars evolved, fitting the wide range of environ-
mental conditions into which this crop complex has
been introduced. They fall into two main groups: (i)
short, thick grained ‘japonica’ (= ‘sinica’) forms,
adapted to the relatively cool climate in northern
74 DOMESTICATION OF PLANTS IN THE OLD WORLD
China and south-east Asia; and (ii) long, thin
grained ‘indica’ adapted to hot, tropical climates.
However, numerous other varieties and sub-
varieties are recognized (see Fuller et al. 2010 for a
detail treatment and updated archaeobotanical and
genetic knowledge).
The crop is closely related to, and fully inter-fertile
with, a variable assemblage of wild and weedy rice
forms which are widely distributed over south and
east Asia including the Indian subcontinent, south
China, and Indonesia (Chang 1995; Vaughan et al.
2008). They include both perennial and annual wild
forms (frequently called O. rufipogon Griff. and O.
nivara Sharma and Shastry respectively), as well as
shattering weedy rices (known as spontanea forms of
O. sativa), which frequently infest rice cultivation. In
many places over these vast territories, the domesti-
cated cultivars, the weedy forms, and the wild races
grow side by side, hybridize with one another, and
form a huge crop complex. There is an ongoing
debate as to whether perennial O. rufipogon or annual
O. nivara, or both, were the direct ancestors of O.
sativa (Sweeney and McCouch 2007). The morpho-
logical and genetic separation between and within
these taxa is, therefore, problematic. Recent genetic
studies (Fuller and Sato 2008; Fuller et al. 2010) sug-
gest that japonica rice was domesticated in the
Yangtze River areas in China sometime about 10,000
years ago, and then spread westward, hybridized
with the wild progenitor of indica rice, and this
hybrid eventually domesticated as the indica rice
somewhat around 4,000 BP.
Since the distributional range of the wild rela-
tives of domesticated Oryza sativa is immense, one
can provide only a general orientation as to where
the wild progenitor of rice was introduced into
domestication. However, archaeological finds in
China, India, and several countries in south-east
Asia provide some clues as to where and when
rice domestication originated (Sweeney and
McCouch 2007; Fuller et al. 2010; Hosoya et al.
2010; Nesbitt et al. 2010).
Based on increases in domesticated-type spikelet
bases, in rice grains, and in arable weeds typical of
rice paddies, between strata, Fuller et al. (2009) sug-
gested that rice’s pre-domestication cultivation
began around 7,000 cal BP in the Lower Yangtze
region, China, pre-dating its local domestication in
around 6,000 cal BP. Somewhat later (in the seventh
millennium BP), remains of domesticated rice
appear at Chengtoushan in the middle Yangtze
Valley (Fuller et al. 2007). Domesticated rice appears
in central China in about 5,000–4,000 BP (Crawford
and Shen 1998).
The earliest convincing signs of rice cultivation
in the Indian subcontinent come from contexts
dated to the second millennium BC; when rice
apparently became an important summer crop in
the subcontinent (Nesbitt et al. 2010).The number
of finds increases considerably from the second
millennium BC onwards (Glover and Higham
1996; Meadow 1996; Fuller 2002). Towards the end
of the third millennium BC, imprints of rice stalks
and husks, as well as charred grains, appear in
farming sites in the central and eastern parts of the
Ganga Valley. By about 2,000–1,800 BC, rice seems
to have been added effectively as a summer crop,
to wheat and barley agriculture in the Indus Valley
and adjacent areas, marking the beginning of the
distinctive, two crop system of the Harappan
agriculture.
Asiatic rice was probably introduced into
south-west Asia in Hellenistic times. Both Greek
and Roman writers knew about this cereal and
said that it was grown in Baktria, Mesopotamia,
and Syria (Lenz 1859, pp. 229–30). A large sample
of rice grains was retrieved from an AD first-cen-
tury Parthian grave, Ville Royale II, Susa, Iran
(Miller 1981). In Roman times, highly prized large
kernel rice was produced in Israel (Feliks 1983)
and imported rice is known from ca. 50 BC
Berenike on the Red Sea coat of Egypt (Cappers
2006). At this time, rice was also grown in the Po
Valley in Italy.
 C H A PTER 4
Pulses
Annual legumes (from the Papilionaceae/Fabaceae
family, of the Leguminosae) cultivated for their seed
accompany the cereals in most regions of grain agri-
culture. They are attractive because contrary to
most other flowering plants, legumes are able to fix
atmospheric nitrogen through symbiosis with the
root bacterium Rhizobium. Rather than use nitrogen
up, pulses add it to the soil. By practising rotation
or mixing of legume crops with cereals, the cultiva-
tor is able to maintain higher levels of soil fertility.
Another virtue is that the seeds of pulses are excep-
tionally rich in storage proteins, whereas grass ker-
nels are rich in starch. Therefore they complement
each other as food elements and contribute to a bal-
anced human diet. In traditional agricultural com-
munities, pulses served—and still serve—as a main
meat substitute.
Both the agronomic compensation and the die-
tary complementation between cereals and pulses
were appreciated in the early days of agriculture.
Each major agricultural civilisation developed not
only its staple cereals, but also its characteristic
companion legumes. Wheats and barley agriculture
in west Asia and Europe had pea, lentil, faba bean,
and chickpea. Maize in Meso-America was accom-
panied by several species of Phaseolus beans, and in
South America also by the groundnut. Pearl millet
and sorghum cultivation in the African Savannah
belt were associated with cowpea and Bombara
groundnut. Soybean was added to cereal cultiva-
tion in China. Hyacinth bean, black gram, and green
gram were introduced into agriculture in India.
Pulses seem to have started their role as compan-
ions of wheats and barley very early in the agricul-
tural history of the Old World. The available
archaeological evidence indicates that pea, lentil,
chickpea, and bitter vetch (probably also grass pea)
were introduced into cultivation more or less
together with the principal cereals. Their remains
abound in south-west Asian Neolithic settlements.
Some of them (particularly lentil and pea) are com-
mon in the contexts of the Neolithic sites that
appeared soon afterwards all over the vast area
from the Atlantic coast of Europe to the Indian sub-
continent. In other words, in south-west Asia,
wheats and barley were not domesticated alone;
pulses accompanied them from the very start
(Zohary and Hopf 1973; van Zeist 1980; Smartt 1990;
Zohary 1996; Butler 1998 Butler 2009).
Several other pulses became grain crops in west
Asia and in Europe only after the establishment of
the ‘first wave’ of legume crops. Prominent among
them were the faba bean and fenugreek. They were
followed, apparently later, by the white lupin. The
cowpea, Vigna unguiculata, was introduced into cul-
tivation in Africa south of the Sahara and reached
the Mediterranean basin only in classical times.
Self-pollination seems to have been a major asset
in domestication also in the seed legumes. Similar
to the cereals (pp. 20–23), the majority of Neolithic
‘first wave’ domesticates (pea, lentil, chickpea, and
bitter vetch) are predominantly self-pollinated. So
are their wild progenitors. Thus, in this group of
grain crops, wild self-pollinated candidates were
‘pre-adapted’ for domestication as compared with
cross-pollinated plants. The advantages conferred
by self-pollination are the establishment of a barrier
between wild and cultivated populations, and the
automatic fixation of desired genotypes.
As in the cereals (pp. 22–23), the evolution
under domestication of the seed legumes is char-
acterized by the development of a syndrome of
75
76 DOMESTICATION OF PLANTS IN THE OLD WORLD

domestication traits (Harlan 1975; Hammer 1984; today reached their present large dimensions only
Smartt and Hymowitz 1985; Harlan 1992a). Very in classical times.
likely, many of these domestic traits evolved as a In several legumes, domestication brought about
result of predominant self-pollination and uncon- striking changes also in the plant habit. The wild
scious selection (Zohary 1989b). relatives of the cultivated pea, grass pea, and the
First, in the wild type, there is an automatic selec-
tion for the breakdown of seed dispersal and the
selection for the retention of seeds in the pod. In most
pulses (e.g. pea, lentil, the various members of the A
genus Vicia) seed dispersal in the wild depends on the
bursting of the mature pods. The pods of domesti-
cated forms do not burst or at least do not split open
quickly. They are harvested and threshed by the
farmer. In pea, lentil, chickpea, and bitter vetch, the
change to non-dehiscing or slow-splitting pods is a B
simple genetic event. It is controlled by a single reces-
sive mutation (or two such mutations, Table 7, p. 61).
A second change is the loss of the wild-type seed
dormancy. The seeds of most wild Mediterranean
pulses have a relatively thick and coarse seed coat,
which insulates the seeds from water penetration,
and spreads germination over several years. This C
wild-type adaptation breaks down under domestica-
tion. Seeds of the domestic varieties tend to have thin-
ner seed coats and are permeable to water.
Consequently, in the traditional Mediterranean pulse
crops, practically all the fresh seeds germinate the
same year they are sown. In other words, the intro- D
duction of the wild progenitors into a system of reap-
ing and sowing brought about an automatic selection
of mutations causing a breakdown of the wild-type
germination inhibition. At least in some pulses, the
seed coat structure can be used—as a telltale trait—to E
identify domestication. However, this is only occa-
sionally feasible in archaeobotanical studies. Most of
the time, charring pulverizes the seed coat, leaving
only the cotyledons and the embryo intact. Seed coats
persist only in exceptional cases of preservation.
Another trend under domestication is the change Fig. 22 Archaeological carbonized seed remains (left) and, for
in seed dimensions (Fig. 22). Most pulse crops bear comparison, seeds from modern cultivars (right) of the first five
south-west Asian domesticated pulses. A–Pea, Pisum sativum,
considerably larger seeds than their wild ancestors.
carbonized seeds from Early Bronze Age Arad, Israel. B–Lentil, Lens
In some cases, there is a three- or fourfold increase culinaris, carbonized seeds from Late Bronze Age Manole, Bulgaria.
in the volume and the weight of the seed (Zohary C–Faba-like bean, Vicia faba var. minor, carbonized seeds from
and Hopf 1973; Smartt 1990). This increase has been Copper Age Chibanes, Portugal. D–Bitter vetch, Vicia ervilia,
a gradual process. Seeds retrieved from Neolithic carbonized seeds from Late Bronze Age Manole, Bulgaria.
E–Chickpea, Cicer arietinum, carbonized seeds from Early Bronze Age
contexts are still relatively small and not very differ-
Arad, Israel (Zohary and Hopf 1973). Note conspicuous size difference
ent in size from those of their wild relatives. The between archaeological and modern seeds; caused by the effect of
large-seeded pulses with which we are familiar charring and/or selection under domestication towards larger seeds.

vetches are climbers with tender branches and char-
acteristic tendrils. Wild lentils are small, delicate
plants. Under cultivation, most of these pulses
evolved stiffer free-standing, robust stems, and a
reduced dependence on climbing—traits that make
them better adapted to grow in stands in tilled
fields.
The wild-type chemical defences have also been
selected against. Many wild legumes contain potent
toxins and antimetabolites in their seeds to protect
them against animal predation. Cultivars fre-
quently lack, or contain only reduced amounts of
these toxic compounds. In some others, fermenta-
tion (e.g. in soybean) or cooking (e.g. in common
bean) is necessary to render the seeds safe for
human consumption.
Lentil: Lens culinaris
Lentil ranks among the oldest and the most appreci-
ated grain legumes of the Old World. It is grown
from the Atlantic coast of Spain and Morocco in the
west, to the Indian subcontinent in the east (Smartt
1990; Zohary 1995a). In Mediterranean grain agri-
culture, it is a characteristic companion of wheat
and barley. Compared to the cereals, yields are rela-
tively low, but lentil stands out as one of the most
nutritious and tasty pulses. The protein content is
about 25%, and lentil constitutes an important meat
substitute in peasant communities. Large quantities
of lentils are produced and consumed (in soup,
paste, in mixture with wheat or rice) in south-west
Asia, India, Pakistan, Ethiopia, and, countries bor-
dering the Mediterranean Sea.
The domesticated L. culinaris subsp. culinaris
Medik. [= Lens culinaris Medik.] manifests a wide
range of morphological variation both in its vegeta-
tive and reproductive parts. Like many other annual
grain crops, lentil is predominantly self-pollinated.
Consequently, numerous true breeding lines and
aggregates of land races have evolved in this grain
crop. All cultivated lentil varieties are diploid (2n =
2x = 14 chromosomes) and predominantly inter-
fertile.
Conventionally, lentil cultivars are grouped in
two intergrading clusters of seed sizes: (a) small-
seeded lentils (subsp. microsperma), with small pods
and small 3–6 mm seed; (b) large-seeded lentils
PULSES 77
(subsp. macrosperma), with larger pods and with
seed attaining 6–9 mm in diameter. As in other
pulses, domestication brought about the retention
of the seed in the pod (pod’s indehiscence) and a
gradual increase in seed size. Macrosperma forms
are to be regarded as more advanced; they start to
appear rather late in archaeological sequences—
only in the third millennium BP. Carbonised seed is
the main element in archaeological assemblages.
Occasionally pods, or fragments of pods, occur as
well.
Wild ancestry
Cultivated lentil belongs to Lens Mill., a small legu-
minous genus, now known to contain the following
taxa (van Oss et al. 1997; Ferguson et al. 1998, 2000;
Sonnante et al. 2009): (i) the crop L. culinaris Medik.
subsp. Culinaris; (ii) the well-recognized wild pro-
genitor L. culinaris subsp. orientalis (Boiss.) Ponert
[= L. orientalis (Boiss.) Shmalh.]; (iii) two more sub-
species of L. culinaris – subsp. odemensis (Ladiz.)
M.E. Ferguson et al., and subsp. tomentosus (Ladiz.)
M.E. Ferguson et al.; and (iv) additional three ordi-
nary wild species: L. nigricans (M. Bieb.) Godr.,
L. ervoides (Brign.) Grande, and L. lamottei Czefr. All
members of Lens, both wild and domesticated, are
annual, ephemeral, diploid, predominantly self-
pollinated plants. All the wild taxa seem to be repro-
ductively isolated from one another by various
combinations of reproductive isolation barriers,
such as the genetic system of selfing, the lack of
crossing, the invariability of the F1 inter-taxa
hybrids, chromosome behaviour in meiosis, and
consequently, inter-specific hybrid sterility. The
presence of such barriers indicates that we are con-
fronted with fully developed biological species. In
contrast, the cross-fertility of culinaris lines with ori-
entalis ones, indicate that both taxa should be
lumped together (as subspecies) in the crop’s bio-
logical species (Zohary 1995a).
Geographically, the genus Lens is a Mediterranean-
basin, south-western, and central Asian element
(Map 8) (Zohary and Hopf 1973; van Oss et al. 1997;
Ladizinsky 1993, 1999; Ferguson et al. 1998). The
wild progenitor of the domesticated plant, subsp.
culinaris, shows close morphological, cytogenetic,
and molecular affinities, to wild subsp. orientalis. In
78 DOMESTICATION OF PLANTS IN THE OLD WORLD
0 100 200 miles
0 200 400 km
Map 8 Geographical distribution of wild lentil, Lens culinaris subsp. orientalis [= L. orientalis] (based on Zohary and Hopf 1973; Ladizinsky 1993;
Gabrielian and Zohary 2004). Wild lentil extends eastwards beyond the boundaries of this map into north Afghanistan and adjacent central Asia
(Ladizinsky and Abbo 1993).
fact, subsp. orientalis (Fig. 23) looks like a miniatur-
ized subsp. culinaris, but bears pods that burst open
immediately after maturation. Cytogenetic tests
have shown (Ladizinsky et al. 1984) that in south-
west Asia, subsp. orientalis is chromosomally varia-
ble. In contrast, the populations growing in central
Asia are chromosomally uniform and exhibit the
‘standard’ chromosomal type only. It also contains
five to six additional chromosomal races that differ
from the standard race by one or even two chromo-
somal rearrangements (mostly translocations).
Cytogenetic affinities of the crop with other Lens
species are less tight. Crosses between L. culinanis
(and/or L. orientalis) with L. odemensis Ladiz. result
in viable F1 hybrids. They are totally sterile since
their parents differ by four chromosomal rearrange-
ments. The four other Lens species, namely L. nigri-
cans (M. Bieb.) Gord., L. ervoides (Brign.) Grande,
L. tomentosus Ladiz., and L. lamottei Czefran. turned
out to be genetically more remote. Crosses between
them and the crop ended in hybrid embryo break-
down (Ladizinsky et al. 1984; van Oss et al. 1997).
Finally, cpDNA tests (Mayer and Soltis 1994), as
well as several other molecular comparisons carried
out on the various Lens species (for enumeration,
consult van Oss et al. 1997), indicated close genetic
links between domestic L. culinaris and wild-type
L. orientalis. They also revealed much wider genetic
distances from the other wild taxa.
Ladizinsky (1999) examined cpDNA restriction
patterns. This, and additional information from
crossability and chromosomal architecture, led him
to conclude that Lentil originated from the territo-
ries around Turkey and north Syria. The same areas
were also mentioned in later rDNA studies con-
ducted by Sonnante et al. (2003) and summarized
recently by Sonnante et al. (2009).
The accumulated botanical and genetic evidence
in Lens establishes orientalis as the wild progenitor
of the cultivated lentil. It also indicates from which
orientalis chromosome type the crop was derived.
As argued by Zohary (1999), the rich chromosomal

Fig. 23 Wild lentil, Lens culinaris subsp. orientalis [= L. orientalis]—a flowering and fruiting plant (Zohary and Hopf 1973).
polymorphism found in the wild progenitor, com-
pared with the chromosomal uniformity in the cul-
tivars, suggests that this pulse crop was taken into
cultivation only once or a very few times.
Finally, the botanical naming in Lens needs an
adjustment. Since it is now clear that wild L. orienta-
lis is the progenitor stock from which the crop has
been derived, this wild lentil should be regarded as
part of the crop’s biological species. Its appropriate
taxonomic ranking is therefore L. culinaris subsp.
orientalis.
Like other grain crops, wild lentils differ from the
domesticated varieties in their seed dispersal biol-
ogy. As already mentioned, in orientalis plants the
pods burst open upon maturation. In contrast, in
cultivars the pods do not dehisce immediately and
the seeds are retained. This difference is governed
by a single mutation (see Table 7), the non-dehiscent
condition (the domestic type) being recessive to the
dehiscent one (the wild type).
Geographically, orientalis is a south-western and
Central Asiatic element (Map 8). It is distributed
PULSES 79
over Armenia, Turkey, Syria, Lebanon, Israel,
Jordan, north Iraq, west and north Iran, and reap-
pears in Turkmenistan, Tajikistan, and Uzbekistan.
Subspecies orientalis grows primarily on shallow,
stony soils, and gravelly hillsides in open or
steppe-like habitats. It also enters disturbed locali-
ties such as stony patches or stone heaps bordering
orchards and cereal cultivation. In most parts of its
distribution, L. orientalis is rather inconspicuous or
even rare. It usually forms small, scattered colo-
nies. However, on stony slopes of Mt Hermon, in
the Anti-Lebanon, the oak park-forest belt of
southern Turkey, and the western escarpments of
the Zagros range, L. orientalis is locally common at
1200–1600 m altitude (Ladizinsky and Abbo 1993).
Frequently it grows side by side with bitter vetch
(Vicia ervilia).
Archaeological evidence
Lentils seem to be closely associated with the start
of wheats and barley domestication in south-west
80 DOMESTICATION OF PLANTS IN THE OLD WORLD
Asia. Probably, this legume was introduced into
cultivation in this region together with emmer,
einkorn, and barley; that is, it should be regarded as
a founder crop of Old World Neolithic agriculture
(Zohary and Hopf 1973). Lentils were apparently
collected from the wild in south-west Asia before
the firm establishment of farming villages. Wild
lentils first appeared in Mousterian, ca. 50,000–
60,000 BP, Kebara Cave, Mt. Carmel (Lev et al. 2005),
and Upper Palaeolithic, ca. 23,000 cal BP, Ohalo II
(Kislev et al. 1992; Simchoni 1998; Weiss 2002, 2009;
Weiss et al. 2004, 2008), Israel. Small, carbonized
seeds of lentil have been retrieved from several pre-
domestication sites across the Fertile Crescent:
Mureybit, ca. 11,800–11,300 cal BP (van Zeist and
Casparie 1968; van Zeist and Bakker-Heeres 1986),
Tell Abu Hureyra, ca. 13,400–11,350 cal BP (Hillman
1975, 2000a; Hillman et al. 2001), Djade el Mughara,
ca. 10,700–10,400 cal BP (Willcox et al. 2008) in Syria,
and from Netiv Hagdud, ca. 11,300–11,100 cal BP
(Kislev 1997), Israel. In these early settlements we
find the collections of wild L. orientalis together with
wild wheats and wild barley, and evidently their
pre-domestication cultivation (Weiss et al. 2006;
Willcox et al. 2008).
A few small lentil seeds have also been found in
the Palaeo- and Mesolithic (pre-farming) layers of
Fig. 24 Carbonized seeds of domesticated lentil, Lens culinaris,
cotyledons only, no seed coat survived charring, PPNB Yiftah’el, Israel
(photograph kindly provided by O. Simchoni).
Franchthi Cave (ca. 15,500–8,750 cal BP), Greece
(Hansen 1991a, 1992), and from Early and Middle
Neolithic layers in Grotta dell’Uzzo (ca. 7,650–6,450
cal BP), Sicily (Costantini 1989). It is very likely that
they represent collected local wild L. nigricans.
Somewhat later, at the end of the tenth and in the
ninth millennia BP, charred seeds of lentil appear in
most of the Pre-Pottery Neolithic B (PPNB) early
farming villages in south-west Asia (Map 1). The
seeds are still similar in size to those of wild forms
(2.5–3.00 mm in diameter), and usually do not occur
in quantities. Yet they are always associated with
domesticated wheat and barley. Among the richest
sites are Tell Aswad, ca. 10,200–9,550 cal BP (van
Zeist and Bakker-Heeres 1982 (1985), Tell Abu
Hureyra, ca. 10,200–8,700 cal BP (Hillman 1975,
2000a, 2000b, 2001; de Moulins 1997, 2000; Hillman
et al. 1989), Jericho, ca. 10,250–9,500 cal BP (Hopf
1983), Çayönü, ca. 10,600-9,900 cal BP (van Zeist
1972; van Zeist and de Roller 1991–2; van Zeist et al.
2003), and Ali Kosh, Iran, ca. 9,600–8,800 cal BP
(Helbaek 1969).
A large hoard of carbonized lentils was recovered
from the ca. 10,100–9,700 cal BP Middle Pre-Pottery
Neolithic B Yiftah’el, north Israel (Fig. 24). The size
of the hoard (7.4 kg, ca. 1,400,000 seeds) and its con-
tamination by the fruits of the weed Galium tricor-
nutum (which is a characteristic weed in lentil
cultivation) indicate that there and then, lentil was
already cultivated (Garfinkel et al. 1988).
Large amounts of lentil seeds were discovered
also in somewhat later phases of the Neolithic settle-
ments in the south-west Asia: in ca. 9,450–9,300 cal
BP Jarmo, Iraq (Helbaek 1959a, 1960; Braidwood
1960), in ca. 9,250–9,000 cal BP Tell Ramad, Syria
(van Zeist and Bakker-Heeres 1985), in ca. 8,200–
7,800 cal BP ceramic Hacilar (Helbaek 1970), in ca.
7,800–7,350 cal BP Girikihaciyan (van Zeist 1979–80),
and in ca. 8,350–7,750 cal BP Tepe Sabz, Deh Luran
Valley, Iran (Helbaek 1969). The Tepe Sabz lentils
had already attained 4.2 mm in diameter. This is an
obvious development under domestication.
Lentils appear frequently in Cyprus as early as
Aceramic Neolithic: in ca. 9,000–7,600 cal BP Dhali
Agridhi, Idalion (Stewart 1974), ca. 8,700–6,800 cal
BP Cap Andreas-Kastros (van Zeist, 1981), and ca.
8,400–8,200 cal BP Khirokitia (Waines and Stanley
Price 1975–1977; Hansen 1991b, 1994).

In the eighth millennium BP, frequent seeds
and pods of small-seeded lentil, were found also
in ca. 7,950–7,150 cal BP Aratashen and Aknashen,
Armenia (Hovsepyan and Willcox 2008).
In the eighth and seventh millennia BP, lentils
seem to be closely associated with the spread of
Neolithic agriculture into south-eastern Europe
(Map 2—Plate 6). Here, too, they are found together
with domesticated emmer, einkorn, and barley.
Charred remains of lentil-seed are present in
almost all early Neolithic Greek agriculture settle-
ments such as aceramic Ghediki (Renfrew 1979),
the pre-ceramic basal levels of Argissa-Magula
(Hopf 1962), ca. 8,650—8,200 cal BP Sesklo (Hopf
1962; Kroll 1981a) and Nea Nikomedeia (van Zeist
and Bottema 1971). Lentil remains are also fre-
quent in later Neolithic and Bronze Age contexts
in the Greek world, such as ca. 8,650–8,400 cal BP,
Knossos (Sarpaki 2009) and second half of fifth
millennium BP Kastanas (Kroll 1983, 1984). They
were also found in large amounts in some of the
Early and Middle (ca. 7,500–6,950 cal BP) Neolithic
Bulgarian sites such as ca. 8,000–7,550 cal BP
Karanovo Mogila (Thanheiser 1997; Marinova
2004, 2006), as well as in ca. 8,200–6,650 cal BP
Anza (Starčevo culture) in Macedonia (Renfrew
1976) and in some Impressed Ware sites in south-
ern Italy (Costantini et al. 1997).
In Germany, as well as in several adjacent central
European countries, lentil seeds were uncovered in
several Linearbandkeramik sites (Willerding, 1980;
Körber-Grohne 1987). Such sites are ca. 7,500–6,550
cal BP Aldenhovener Platte (Knörzer 1973, 1974,
1997), ca. 7,150–6,800 cal BP Schletz in Austria
(Schneider 1994; Kohler-Schneider 2007), ca. 7,400–
6,550 cal BP Bylany, in the Czech Republic (Tempír
1979), ca. 7,550–6,700 cal BP Menneville, France
(Bakels 1984, 1991; Ilett et al. 1995) and also in ca.
7,400–7,050 ca BP Coveta de l’Or and Cova de
Cendres, Spain (Hopf and Schubart 1965; Lopez
1980; Buxó 1997), frequently together with pea. In
Hungary, lentil occurs in Middle Neolithic, ca. 6,350
cal BP Dévaványa-Réhelyi dűlő (Hartyányi et al.
1968), and in Rumania in Vinča culture Liubcova
(Cârciumaru 1996). As in the case of the pea
(pp. 85–86) and other pulses, lentils in many Bronze
Age settlements in Europe seem to be sparser than
in Neolithic times.
PULSES 81
Lentils accompany wheats and barley in the
spread of south-west Asian agriculture southwards
to Egypt and eastwards to the Caspian basin and
the Indian subcontinent. The earliest records of Lens
from Egypt come from ca. 7,150–5,950 cal BP
Neolithic Merimde (Wetterstrom 1993). Lentils have
also been found in pre-dynastic, ca. 6,000–5,500 BP
Amratian (Naqada I) culture sites (Wetterstrom
1993), and at sites of all epochs ever since. Further
east, charred lentil seeds appear in ca. 2,350–2,100
cal BP Shortughai, Afghanistan (Willcox 1991), and
in several Harappan culture sites in Pakistan and in
north-west India—this from the older phases of this
culture onward (Fuller 2002). Some of the lentil
remains discovered in Miri Qalat, south-west
Pakistan (Tengberg 1999) might be even older.
In analyzing lentil remains obtained from the
early south-west Asian sites, it is often difficult to
decide whether they represent wild material or
domesticated forms. The seed of wild Lens is mor-
phologically very similar to that of the domesticated
pulse. The only trait indicating domestication in the
archaeological record is the increase in seed size. Yet
this process was slow and gradual. The first signs of
seed size increase appear as late as in the end of the
eighth millennium BP; i.e. some two millennia after
the definite establishment of wheat and barley cul-
tivation in south-west Asia. We have, however,
some circumstantial evidence suggesting that lentil
farming is as old as agriculture itself. As already
pointed out, the hoard discovered in ca. 10,100–
9,700 cal BP Yiftah’el indicates treatment as a crop.
Indicative are also the data on the spread of Neolithic
agriculture into Europe (Zohary and Hopf 1973).
Lentils are repeatedly encountered in the early
European agricultural settlements of the seventh
millennium BP, situated far outside the areas in
which either L. orientalis or L. nigricans grow wild.
This is a strong indication that lentils were already
cultivated in these regions. Such early diffusion,
together with the main south-west Asian cereals,
could only be the outcome of an even earlier domes-
tication, in the area where the wild progenitor
occurs.
In summary, archaeological remains do not pro-
vide us with any direct diagnostic trait for a conclu-
sive determination of the start of lentil domestication.
Moreover, it is very doubtful whether comparative
82 DOMESTICATION OF PLANTS IN THE OLD WORLD
morphology will provide us with such clues in the
future. Yet once Neolithic agriculture is soundly
established, cultivation of lentil is part of it. The
available archaeological information on early
remains of lentil comes from south-west Asia. This
is the very territory over which L. orientalis is geo-
graphically distributed.
Pea: Pisum sativum
The pea ranks among the oldest grain legumes of
the Old World. From its early beginnings, this crop
has been a close companion of wheat and barley
cultivation (Zohary and Hopf 1973). Pea, Pisum sati-
vum L., is well adapted to both warm Mediterranean-
type and cool temperate conditions. In peasant
communities in south-west Asia, the Mediterranean
basin, temperate Europe, Ethiopia, and north-west
India, it constitutes an important source of protein
for human consumption. The protein content of the
seed is about 22%. Today, pea ranks among the
world’s most important pulses (Smartt 1990; Davies
1995). It is grown quite extensively in cool regions
such as northern Europe and north-western USA.
Mature dry seeds were the principal product in
classical times. Today, a substantial proportion of
the seed and/or pods are also is harvested as imma-
ture vegetable crop either for direct consumption or
for canning and freezing.
Pea is a diploid (2n = 2x = 14 chromosomes) and
predominantly self-pollinated crop. As a conse-
quence of the self-pollination system, variation in
pea is moulded in numerous true breeding lines.
These were used by Gregor Mendel for his classic
genetic experiments, some one hundred and sixty
years ago. Cultivated pea shows a wide range of
morphological variation. Hundreds of land races are
recognized and dozens of modern cultivars have
been produced by breeders. All are inter-fertile, or at
least partly inter-fertile. Cultivars differ from one
another in numerous traits such as the height and
habit of the plant (from short field types to tall climb-
ers), the colour of the flower (blue, purple, to white),
the size and form of the seed, the texture and the
colour of the seed coat, and the colour of the cotyle-
dons (yellow or green). Forms with coloured flow-
ers, relatively small seeds, and long vines are
frequently called field peas (var. arvense), while cul-
tivars with white flowers, large seeds, and shorter
branches are known as garden peas (var. sativum).
As in many other grain legumes, the conspicuous
features of evolution under domestication are: (i)
indehiscence of the pod—the retention of seed on
the ripe pod, a trait governed by a single recessive
mutation (Table 7, p. 61); (ii) the gradual increase of
seed size from 3–4 mm to 6–8 mm in diameter; and
(iii) the reduction of the relatively thick texture and
rough surface of the seed coat (testa), which evolved
to be thin and smooth, resulting in the breakdown of
the germination inhibition of wild peas. Seed
remains constitute the bulk of pea material recov-
ered from archaeological excavations. Other plant
parts, such as pods, were hardly ever retrieved.
Wild ancestry
The domesticated pea belongs to a small legumi-
nous genus, Pisum L., restricted, in the wild, to the
Mediterranean basin and south-west Asia. All mem-
bers of this genus are annual, diploid (2n = 2x = 14
chromosomes) predominantly self-pollinated plants.
On the combined basis of morphology, ecology,
cytogenetics, and molecular analysis, most botanists
now recognize two biological species in Pisum:P.
sativum and P. fulvum (Davis 1970; Ben Ze’ev and
Zohary 1973; Waines 1975; Waines and Stanley Price
1975–1977).
P. sativum L. (the crop complex) contains a varia-
ble aggregate of cultivated peas, and numerous
wild races closely related to the crop. All forms
within the complex readily cross with one another
and the hybrids are fully or almost fully fertile. All
show full chromosome homology except for a sin-
gle reciprocal translocation found in some wild
forms and also in very few cultivated varieties.
P. fulvum Sibth. & Sm. is a distinct east-Mediterra-
nean wild species with characteristic yellow-brown-
ish flowers, and chromosomes that are considerably
divergent from those present in P. sativum. The two
species are reproductively well isolated in nature,
and their hybrids are semi-sterile.
The wild forms of P. sativum fall into two main
morphological types:
(i) A tall ‘maquis type’, previously called P. elatius
M. Bieb. [= P. sativum L. subsp. elatius (M. Bieb.)

Representative sites of wild humble peas
Representative sites of wild elatius peas
Map 9 Geographical distribution of the two main wild races of pea, Pisum sativum: (i) ‘steppe-type’ humile forms, and (ii) ‘maquis-type’ elatius
forms (based on Zohary and Hopf 1973). In the west-Mediterranean basin, wild elatius peas extend beyond the boundaries of the map and reach
as far as Spain.
Aschers. & Graebn.] is omni-Mediterranean in
its distribution and it thrives as a sporadic
climber in maquis formations in the relatively
mesic parts of the Mediterranean basin.
Occasionally elatius peas also colonize hedges
and terraces bordering agricultural fields.
(ii) A shorter, more xeric ‘steppe type’ (Fig. 25), for-
mally named P. humile Boiss. & Noë [= P. syria-
cum (Berger) Lehm; P. sativum L. var. pumilio
Meikle], is geographically restricted to south-
west Asia (Map 9). It occurs in the deciduous
oak park-forest belt and in open, steppe-like,
herbaceous vegetation, characteristic to south-
west Asia, i.e. in the same zone that harbours
the wild progenitors of wheats, barley, lentil,
and flax. From such primary habitats humile
peas spill over to secondary habitats and occa-
sionally infest cereal cultivation. In their gen-
eral habit, some humile forms resemble the
cultivated legume closely, and differ from it
mainly by their rough seed coat, and by pods
PULSES 83
0 100 200 miles
0 200 400 km
that burst open and disperse the seed soon after
maturation.
The boundaries between these two principal wild
races are occasionally blurred. Intergrading forms
are particularly common, especially in Turkey.
Spontaneous hybridization between humile or ela-
tius peas and the domestic varieties also occurs
sporadically.
Chromosomally, the wild peas of the P. sativum
complex fall into two chromosomal types (Ben
Ze’ev and Zohary 1973). Two humile collections,
one from Mt Hermon area and the other from the
environs of Ankara, Turkey, were found to have a
chromosome complement identical to that prevail-
ing in the cultivated varieties. Hybrids between
such tame and wild forms show normal chromo-
some pairing and are fully fertile. In contrast, four
other humile collections from south and central
Israel, and elatius material from Israel and Italy,
differ from the crop by a single, chromosomal
84 DOMESTICATION OF PLANTS IN THE OLD WORLD
translocation. Hybrids in such cases show full chro-
mosome pairing but some reduced fertility due to
the translocation heterozygosity.
Nasiri et al. (2009) have examined SSR (microsat-
ellite) markers of wild peas and cultivars. They
have found that these markers effectively differenti-
ate between cultivars and the wild accessions. In
addition, P. sativum subsp. fulvum was found to be
the closest relative of the cultivars, but it should be
noted that subsp. humile was not tested in this
study.
In summary, the available evidence from the liv-
ing plants points to wild humile peas as the progeni-
tor stock for pea domestication. Humile peas show
closer morphological similarities than elatius peas
to the domesticated aggregate and grow in steppes
or steppe-like habitats; i.e. under open conditions
that are not very different from those prevailing
under domestication. The Turkish and the south
Levant forms, having chromosomes identical with
Fig. 25 Wild pea (‘steppe-type’) Pisum sativum subsp. humile [= P. syriacum]—a flowering and fruiting plant (Zohary and Hopf 1973).
those present in the cultivars within humile peas,
should be regarded as the primary ancestral stock.
This is also supported by chloroplast DNA compar-
isons (Palmer et al. 1985). Yet it is very likely that the
humile forms with the chromosome translocation, as
well as the more mesic wild elatius peas, contrib-
uted some genes to the domesticated ensemble
through occasional secondary hybridization.
Finally, it is important to note the occasional testa
remains in archaeological pea finds. These finds
allow us to differentiate between domesticated
peas, those with smooth seed coat, and the wild-
type seeds, those with rough seed coat (Werker et al.
1979; Butler 2009).
Archaeological evidence
Remains of peas are present in many of the Pre-
Pottery Neolithic B farming villages that developed
in the Fertile Crescent in the second half of the tenth

to the ninth millennium BP (Map 1). The first find-
ings of wild-form pea come from Upper Palaeolithic
ca. 23,000 cal BP Ohalo II, Israel (Kislev et al. 1992;
Simchoni 1998; Weiss 2002, 2009; Weiss et al. 2004,
2008). Some of the earliest finds of cultivated peas
come from ca. 10,500–10,200 cal BP Tell Aswad in
south Syria (van Zeist and Bakker-Heeres 1985), ca.
10,250–9,550 cal BP Çayönu in south-east Turkey
(van Zeist and Bakker-Heeres 1985), ca. 9,900–9,550
cal BP PPNB Jericho (Hopf 1983) and ‘Ain Ghazal,
Jordan (Rollefson et al. 1985). Larger quantities of
peas were available from somewhat later Neolithic
phases in south-west Asia—from the eighth millen-
nium BP. Carbonized seed accompanied the domes-
ticated wheats and barley in ca. 9,350-8,950 cal BP
Çatalhöyük (Helbaek 1964a; Fairbairn et al. 2002,
2005, 2007), and a large hoard in final Neolithic, ca.
8,550–8,150 cal BP Erbaba, Turkey (van Zeist and
Buitenhuis 1983). Few seeds have been found in ca.
9,450–9,300 cal BP Jermo, Iraq (Helbaek 1959b, 1960;
Braidwood 1960).
In contrast to the wheats and barley, the earliest
archaeological remains of pea do not provide us
with simple traits for a foolproof recognition of
domestication (Zohary and Hopf 1973). In domesti-
cated peas there is a general trend towards an
increase in seed size and hilum elongation, but such
changes occurred gradually in the course of domes-
tication. In early finds there is considerable overlap-
ping in the dimensions of wild and domesticated
forms. Perhaps the most reliable indication of
domestication in peas is provided by the surface of
the seed coat (testa)—as long as it is preserved. Wild
peas are characterized by a rough or granular sur-
face, while domesticated varieties have smooth
seed coats. However, seed coats in most finds are
missing, which makes it impossible to ascertain
whether they are wild or domesticated. The lower
levels, ca. 10,250–9,550 cal BP, of Çayönü (van Zeist
and Bakker-Heeres 1985) retained some fragments
of seed coats showing a rough surface, thus indicat-
ing a collection from the wild. Wild-type seed coats
occur even much later in Late Neolithic, ca. 8,200–
7,800 cal BP, Hacilar (Helbaek 1970). Significantly,
the remains from upper levels of Çayönü include a
single seed with a smooth seed-coat; and those from
ca. 9,350–8,950 cal BP Çatalhöyük (Helbaek 1964a;
Fairbairn et al. 2002, 2005, 2007), Late PPNB 8,700–
PULSES 85
7,000 cal BP Can Hasan I (Renfrew 1968), and from
ca. 9,450–8,600 cal BP Bouqras (van Zeist and
Waterbolk-van Rooijen 1985) show the smooth seed
coat characteristic of domesticated varieties. This
strongly suggests that domestication of peas in
south-west Asia is as old as the domestication of
wheats and barley.
During the ninth millennium BP, peas spread out-
side the Fertile Crescent into Cyprus, the Aegean
(Fig. 26), and the Balkan spheres. Peas appear in
Aceramic Neolithic Cyprus in: ca. 9,000–7,600 cal
BP Dhali Agridhi, Idalion (Stewart 1974), ca. 8,700–
6,800 cal BP Cap Andreas-Kastros (van Zeist 1981),
and ca. 8,400–8,200 cal BP Khirokitia (Waines and
Stanley Price 1975–1977; Hansen 1991b, 1994). At
early Neolithic Greece they were found in ca. 8,400–
8,100 cal BP Nea Nikomedeia (van Zeist and
Bottema 1971), and at two early Neolithic, Starčevo-
culture, sites in Macedonia: ca. 8,200–6,650 cal BP
Anza, and ca. 8,150–7,000 cal BP Obre (Renfrew
1974, 1976). The Nea Nikomedeia carbonized seeds
are well-preserved and reveal the smooth seed coat
of domesticated forms.
Peas seem to be associated with the spread of
Neolithic agriculture into Europe in the eighth mil-
lennium BP (Zohary and Hopf 1973; van Zeis 1980).
5 mm
0
Fig. 26 Charred seed of domesticated pea, Pisum sativum,
cotyledons only, no seed coat survived charring, Late Neolithic Dimini,
Greece (Kroll 1979).
86 DOMESTICATION OF PLANTS IN THE OLD WORLD
Here again they are closely associated with wheats
and barley production (Map 2—Plate 6). At the first
half of the eighth millennium, peas were found
mainly in the southern European countries border-
ing the Mediterranean, while in the second half they
reached the middle European countries.
Peas are present in Early Neolithic Bulgaria: ca.
8,000–7,700 cal BP Kovacevo (Popova 1992; Marinova
2006), ca. 7,850–7,700 cal BP Kapitan Dimitrievo
(Marinova 2006, forthcoming), and ca. 7,850–7,500
cal BP Ašmaska Mogila (Hopf 1973; Renfrew 1979,
Table 7), and also in early Neolithic, Starčevo-Criş
culture ca. 7,600–7,500 cal BP Sacarovca, Moldavia
(Januševič 1984; Kuzminova et al. 1998). In Italy,
early Neolithic peas appear first in the south and
later in the north, in ca. 8,000–7,600 cal BP Scamuso
(Costantini et al. 1997), ca. 7,750–7,150 cal BP La
Marmotta (Rottoli 1993, 2002), and finally in ca.
7,550–6,450 cal BP Sammardenchia (Pessina and
Rottoli 1996; Rottoli 2005; Rottoli and Pessina
2007). Already in the beginning of the first half of
the eighth millennium BP, peas appear further
east in ca. 8,000–7,150 cal BP Arukhlo 1 and
Arukhlo 2, Georgia (Januševič 1984; Lisitsina 1984;
Schultze-Motel 1988a), and further west in ca.
7,950–5,450 cal BP Balma Margineda, Spain
(Marinval 1995). Later, they reached ca. 7,550–
6,700 cal BP Menneville, France (Bakels 1984, 1991;
Ilett et al. 1995).
In the second half of the eighth millennium
BP, pea is associated with the spread of the
Linearbandkeramik (LBK) culture in Europe
(Willerding 1980) as well as the expansion of agri-
culture to Egypt. In the lower Rhine Valley (Rosdorf),
peas are common in LBK sites dated ca. 7,450-7,150
cal BP (Kirleis and Willerding 2008); and in central
Germany large amounts of charred seeds have been
recovered from similar LBK settlements, such as ca.
6900–6500 cal BP Dresden-Nickern (Baumann and
Schultze-Motel 1968). In these samples, seed coats
are frequently well-preserved and, with one excep-
tion, they show the smooth surfaces characteristic
of the domestic crop. Also in LBK Třtice in Bohemia
(Tempír 1973, 1979), peas appear in quantities; and
they reappear in several early Neolithic sites in
Moldavia, like ca. 7,600–7,400 cal BP Sacarovca
(Januševič 1984; Kuzminova et al. 1998; Monah
2007b). All over this area, from Ukraine (e.g. Rivne,
Pashkevich 2003) to western Germany, peas become
more frequent in late Neolithic times. The earliest
finds from Poland, so far, come from Funnel Beaker
contexts in ca. 5,600–5,300 cal BP Čmielów
(Klichowska 1976). A hoard of pea seeds was found
in a Dudéşti culture context in ca. 6,500–6,000 cal BP
Cîrcea, Rumania (Cârciumaru 1996). In the west
Mediterranean basin, a few seeds have been found
in ca. 7,950–5,450 cal BP Balma Margineda, Spain
(Marinval 1995), and the evidence continues to the
Chalcolithic and Bronze Age layers. Generally, finds
of peas, as well as of other pulses, are sparser
throughout European Bronze Age compared to the
Neolithic finds.
Richer deposits of pulses, including lentil and
broad bean, occur again in the European settle-
ments of later periods (Zohary and Hopf 1973).
Such finds came from ca. 4,150–3,200 cal BP
Canton of Grisons, Switzerland (Jacomet et al.
1998, 1999), ca. 4,950–3,350 cal BP Albertfalva,
Hungary (Gyulai 2003), ca. 3,200–2,700 cal BP
Stillfried, Austria (Kohler-Schneider 2003, 2007),
ca. 3,360-3,700 cal BP Wange and Overhespen,
Belgium (Bakels 1992), and ca. 3,900–3,350 cal BP
Nitriansky Hrádok, Czech Republic (Tempír 1969;
Kühn 1981).
Pea remains appear also in late eighth millen-
nium BP Neolithic settlements of the Nile Valley,
Caucasia and Transcaucasia. In Egypt, early finds
come from ca. 7,150–5,950 cal BP Merimde, Nile
delta (Wetterstrom 1993) and from ca. 5,650–5,300
cal BP pre-dynastic Nagada settlements, Upper
Egypt (Wetterstrom 1993). Further east, the earliest
finds are of a later date. Pea remains were retrieved
from ca. 2,350–2,100 cal BP Shortughai, Afghanistan
(Willcox 1991). They are present in ca. fifth millen-
nium BP Harappa (Vishnu-Mittre and Savithri
1982), and in several other Harappan sites in
Pakistan and north-west India, from the older
phases of this culture onward (Fuller 2000).
In conclusion, the archaeological evidence
establishes pea as one of the founder grain crops
of south-west Asian Neolithic agriculture. Since
this early start, pea seems to be a consistent ele-
ment in Neolithic and Bronze Age food produc-
tion throughout West Asia and Europe and a
common companion of wheats and barley. The
evidence from the living plants complements the

archaeological finds. The wild humile forms, with
chromosomes and cpDNA identical to those pre-
vailing in the cultivated crop, should be regarded
as the closest wild stock from which this pulse
crop evolved.
Chickpea: Cicer arietinum
Chickpea is a valued seed legume of traditional
agriculture in the Mediterranean basin, western
Asia, as well as India and Ethiopia. It is a member of
the grain ensemble found in south-west Asian
Neolithic and Bronze Age remains. Chickpea is
adapted to a subtropical or Mediterranean-type cli-
mate; it grows almost exclusively in the post-rainy
season on moisture stored in the soil. The main
growers today are in the Indian subcontinent, the
countries around the Mediterranean basin and
Ethiopia (http://faostat.fao.org). Like lentil and
pea, chickpea, with a seed protein content of some
2%, constitutes an important meat substitute in
peasant communities.
Domesticated chickpea, Cicer arietinum L. subsp.
arietinum, is a predominantly self-pollinated annual
crop with pods containing one to two seeds. The
cultivars show a wide range of variation in size,
colour, and shape of the seed, and in the size and
form of leaves and flowers. All cultivated varieties
are diploid (2n = 2x = 16 chromosomes) and inter-
fertile. Chickpea land races are grouped into two
inter-connected clusters (Smithson et al. 1985;
Smartt 1990). Large-seeded varieties (known as
‘Kabuli’ type) with relatively smooth, rounded,
light-coloured seed coats and pale cream flowers,
predominate in the Mediterranean countries and
south-west Asia. Varieties producing small, wrin-
kled seed (‘Desi’ type) with dark-coloured seed-
coats and usually purple flowers prevail in the
eastern and southern parts of the distribution area
of the crop; i.e. in India, Afghanistan, Pakistan, and
Ethiopia.
Seed dispersal in wild chickpea is attained in two
ways: (i) shattering of the seeds from the dry, split-
ting pods, still attached to the mother plant; and (ii)
dispersal of whole inflated pods by wind. As in
most other seed legumes, the conspicuous features
of evolution under domestication in chickpea are
the retention of pods and seeds on the plants and
PULSES 87
the gradual increase in seed size from 3.5 to 6.0 mm
and more. Large-seeded forms are obviously more
advanced. Another change is the development of a
smooth seed coat, and the reduction of its thickness.
Seed remains are the only material recovered to-
date in archaeological excavations.
Wild ancestry
The domesticated chickpea, Cicer arietinum subsp.
arietinum, is a member of a leguminous genus com-
prising some forty species, centred in central and
western Asia (van der Maesen 1972, 1987; Cole et al.
1998). The majority of Cicer species are herbaceous
perennials or dwarf shrubs; some are annuals. The
domesticated pulse shows close morphological
affinities (and an almost identical seed-protein pro-
file) to two wild species of chickpea: C. echinosper-
mum Davis and C. reticulatum Ladiz. (Ladizinsky
and Adler 1976). These two wild chickpeas are dip-
loid, self-pollinated annuals, recorded only in
south-eastern Turkey. Externally, they resemble
each other closely, but they can be distinguished
from one another by testa (seed-coat) texture, which
is echinate in C. echinospermum and reticulate in
C. reticulatum. Also, they differ from one another by
their soil preferences; the first grows mainly on
basaltic soils, while the second thrives on limestone
bedrock. Of the two wild chickpeas only C. reticula-
tum (Plate 10) is fully inter-fertile and contains chro-
mosomes homologous to those found in the crop
(Ladizinsky and Adler 1976). Crosses between the
domesticated chickpea and C. echinospermum, and
also those between C. reticulatum and C. echinosper-
mum are more difficult to achieve, and the inter-
specific F1 hybrids obtained proved to be highly
sterile. Attempts to cross the domesticated chickpea
with other closely related wild Cicer species—
C. pinnatifidum Jaub. & Spach, C. judaicum Boiss.,
and C. bijugum Rech.—failed altogether, indicating
that these wild species are not closely related to the
crop. The distribution of C. reticulatum is currently
restricted to south-eastern Turkey (Map 10). Lev-
Yadunet al. (2000) argue for this limited area, where
other wild progenitors grow as well, to be the ‘core
area’ for all founder crops.
Since this distribution is so limited, molecular
research has been conducted on material collected
88 DOMESTICATION OF PLANTS IN THE OLD WORLD
0 100 200 miles
0 200 400 km
Map 10 Geographical distribution of wild chickpea, Cicer arietinum subsp. reticulatum [= C. reticulatum] (based on Ladizinsky and Adler 1976;
Tanno and Willcox 2006b; and unpublished data of D. Zohary).
from this area only. Indeed, the molecular evidence
seems to supports the identification of C. reticulatum
as the wild progenitor. Both Nguyen et al. (2004),
upon examining 214 AFLP loci of 94 different acces-
sions of chickpea and its wild relatives, and Sethy
et al. (2006) upon examining microsatellite markers,
pointed out the same wild progenitor.
In summary, the morphological, cytogenetic, and
molecular evidence currently available, points to
C. reticulatum as the wild progenitor of the domesti-
cated plant. Consequently, it is referred to as C. ari-
etinum subsp. reticulatum. The central part of the
Fertile Crescent seems to be the territory in which
chickpea could have been taken into domestication.
It is noteworthy that unlike wild C. reticulatum,
which is an annual plant that germinates in the
rainy season, south-west Asian domesticated
chickpeas are grown as summer crops. This shift to
summer growing, is attributed by Abbo et al. (2003)
to the interaction with the pathogen Didymella
rabiei, the casual agent of Ascochyta blight. Another
attraction for the grower to turn chickpea into an
early summer crop are the many hours of daylight
during the long, summer days, which enhance
speedy maturation of seeds, even in colder envi-
ronments (ibid). It seems that the available poten-
tial of chickpea as a major grain crop is not yet
fully utilized.
Archaeological evidence
Like lentil and pea, chickpea seems to have been
closely associated with the start of agriculture in
south-west Asia. However, in Pre-Pottery Neolithic

B contexts in this core area, chickpea is much rarer
than the two other pulses. The available evidence
indicates that several closely related annual wild
Cicer species co-exist in the south-west Asia.
A few charred chickpea seeds were recovered
from various phases of aceramic Çayönü, ca.
10,250–9,550 cal BP and later (van Zeist 1972; van
Zeist and de Roller 1991–2, 2003), from contempo-
rary Nevali Çori (Pasternak 1998), and from ca.
10,200–9,500 cal BP Aşikli Höyük (van Zeist and de
Roller 1995) in Turkey. A few more were found in
Middle PPNB, ca. 9,400–7,000 cal BP Tell Abu
Hureyra, northern Syria (Hillman 1975, 1989; de
Moulins 2000). The seeds still correspond in size to
those of C. reticulatum and, because these sites are
situated within or close to the present distribution
area of the wild progenitor, the finds could repre-
sent either wild or domesticated material.
Chickpea seeds were retrieved from PPNA, ca.
10,600–10,250 cal BP, Tell el-Kerkh, north-west Syria
(Tanno and Willcox 2006b). A few of these seeds are
somehow larger and plumper than the wild progen-
itor. Tanno and Willcox (2006b, p. 200) argue that
this might indicate that they are ‘intermediate stage’
between wild and domesticated chickpeas. The
seeds retrieved from Pre-Pottery Neolithic B, ca.
9,900–9,550 cal BP, Jericho (Hopf 1983) and ‘Ain
Ghazal (Rollefson et al. 1985), and those from the
Late PPNB level in Ramad near Damascus (van Zeist
and Bakker-Heeres 1985) probably represent domes-
ticated forms. The latter sites lie far away from the
territory of the wild progenitor, and the specimens
from Jericho seem to have smooth coats. Likewise,
some 500 chickpeas seeds, few of them with smooth
testa, from ca. 8,350–8,050 cal BP pottery Neolithic
site of Höyücek, south-west Turkey, positively rep-
resent domesticated chickpea (Martinoli and Nesbitt
2003).
Early Bronze Age remains of chickpeas are more
abundant. Considerable amounts of charred, well-
preserved, fairly large seeds with smooth coats
were retrieved in Israel and Jordan from Early
Bronze Age Arad (Hopf 1978c), contemporary
Jericho (Hopf, 1983), Bab edh-Dhra (McCreery
1979) and ca. 4,900 cal BP Tell es-Sa’idiyeh
(Cartwright 2003; and persoal communication).
These finds provide a clear indication of chickpea
domestication since the only chickpea growing
PULSES 89
wild in Israel and Jordan, namely Cicer judaicum,
has significantly smaller seed and a characteristic
rough seed coat.
A single, well-preserved chickpea was retrieved
from a pre-Sesklo layer, eighth millennium BP Otzaki,
Greece (Kroll 1981a). A few seeds were also found in
Early Neolithic, ca. 8,000–7,700 cal BP Kovacevo
(Popova 1992; Marinova 2006) and ca. 7,850–7,700 cal
BP Kapitan Dimitrievo (Marinova, forthcoming) in
Bulgaria. Richer finds are available from late
Neolithic, ca. 6,650–6,300 cal BP, Dimini, Greece
(Kroll 1979). They measure 4.24 × 3.77 × 3.51 mm.
The domesticated pulse must therefore have reached
Europe with one of the first waves of migration of
early grain crops from south-west Asia.
The crop is missing in Neolithic layers of former
Yugoslavia, and the western, central, and northern
European countries. From Bronze Age to classical
times, chickpea evidently ranked among the popu-
lar pulses of the Mediterranean basin and the south-
west Asian countries. The only early chickpea find
from Egypt is ca. 1,325 BC Tutankhamun tomb
(Germer 1989a; Hepper 1990; de Vartavan et al.
2010).
In conclusion, the evidence from the living plants
and the plant remains discovered in archaeological
excavations indicate that Cicer arietinum belongs to
the Early Neolithic grain crop assemblage of south-
west Asia. Archaeological data is limited, but in this
legume, the delimitation of the place of origin is
relatively simple: the wild progenitor of the domes-
ticated chickpea is endemic to the central part of the
Fertile Crescent. Very probably, this pulse was first
brought into domestication there.
Faba bean: Vicia faba
Together with lentil, pea, and chickpea, the faba
bean, Vicia faba L. (broad bean, horse bean), belongs
to the principal pulses of Old World agriculture. It
grows well in both warm, summer-dry Mediter-
ranean-type environments, and in the more north-
erly temperate parts of Europe and Asia (Bond et al.
1985). The erect, robust plant readily bears thresha-
ble pods and relatively large seed (Plate 11) with
high (about 2–25%) protein content. In some Asian
and Mediterranean countries, particularly in Egypt,
the dry seeds of the faba bean provide a staple
90 DOMESTICATION OF PLANTS IN THE OLD WORLD
protein source for the poor. Unripe pods are also
used when green. In both Europe and Asia, the seed
is regarded as a valuable animal feed, hence the
name ‘horse bean’. Large quantities of V. faba are
produced in several Mediterranean countries:
Egypt, Morocco, Spain, Italy, Turkey, as well as in
Ethiopia, temperate Europe, south-western Asia,
and especially in China.
Vicia faba is an annual, diploid grain crop
(2n = 2x = 12 chromosomes). In contrast to the major-
ity of the grain crops, the faba bean is not fully a self-
pollinated crop (Lawes 1980). It is self-compatible,
and a considerable amount of cross-pollination takes
place in numerous cultivars. Furthermore, when
subjected to selfing, many domesticated varieties
manifest various degrees of inbreeding depression.
On the other hand, some numerous faba cultigens,
particularly paucijuga forms from India and
Afghanistan, are able to self-pollinate.
Under domestication, faba beans developed a
considerable amount of morphological variation
and different ecological adaptations. Conspicuous
intra-specific differences are displayed in various
traits like vegetative habit, pod structure, pod shat-
tering, and the shape, colour, and size of the seed.
Seed size has served as the principal character for
intra-specific subdivision. Faba bean taxonomists
(Muratova 1931; Hanelt 1972) recognize three or
four intergrading and inter-fertile main types of
cultivar forms in V. faba. Forms with relatively small
rounded seed measuring 6–13 mm are placed in
V. faba var. minor. Some small-seeded forms, grown
mainly in India, Pakistan, and Afghanistan, also
show reduced numbers of leaflets (mostly one pair
per leaf), and have very small flowers. They are fre-
quently regarded as a distinct intra-specific taxon:
var. paucijuga. Forms with medium-sized seed, that
are 15–2 mm long, 12–15 mm wide, and 5–8 mm
thick, are placed in var. major. Large-seeded faba
seem to have evolved under domestication rela-
tively late. Archaeological remains ranging from the
Neolithic period to Roman times are almost all
within the range of var. minor.
Wild ancestry
The wild ancestor of the domestic faba bean has not
yet been discovered despite intensive research in
the last fifty years. In its general morphological
appearance, this domestic pulse resembles another
group of large-seeded south-west Asian and
Mediterranean wild vetches, namely: V. narbonensis
L., V. serratifolia Jacq., V. johannis Tamamsch., V. gali-
lea Plitm. & Zoh., V. hyaenis-ciamus Mouterde, and
V. kalakhensis Khattab, Maxted & Bisby. The first
three taxa are widely distributed over south-west
Asia and the Mediterranean basin, V. galilea is a
south-west Asian element, and the last two taxa are
strict endemics confined each to a small territory in
Syria (Maxted 1995). The morphological similarities
between the domestic faba bean and this group of
wild vetches are considerable. Consequently, tax-
onomists dealing with the genus Vicia, used to place
both the crop and these wild taxa in a common sec-
tion (Sect. Faba Aschers. & Graebn.). The ancestry of
the crop was sought among these wild taxa.
However, extensive cytogenetic tests proved that
the faba crop is unique both chromosomally and
reproductively. The first surprise was that while V.
faba has 2n = 2x = 12 chromosomes, all the above-
mentioned wild vetches were found to have
2n = 2x = 14 chromosomes. Furthermore, large-
scale attempts to cross the wild vetches with the
faba crop (Schäfer 1973; Ladizinsky 1975) failed
altogether, indicating that V. faba is reproductively
strongly isolated from these vetches. These finds
ruled out the candidacy of the above-mentioned
faba-resembling wild vetches for the ancestry of the
crop. They are now regarded as a separate aggre-
gate of species ‘Vicia narbonensis complex’, (see
Schäfer 1973; Bennett and Maxted 1997). Maxted
(1995) went a step further. He separated this
2n = 2x = 14 aggregate from V. faba and considers it
as an independent taxonomic section (Sect.
Narbonensis (Radzhi) Maxted). Finally, molecular
comparisons (van de Ven et al. 1993; Jaaska 1997a;
Potokina et al. 1999) also indicate that V. faba is not
close to the V. narbonensis complex. In summary, in
the case of the faba bean, we still need to discover
an elusive 12-chromosomed wild progenitor that, in
all likelihood, is very limited in its geographic dis-
tribution. There is also the possibility that this wild
ancestor is extinct.
While these faba-resembling vetches have been
ruled out as candidates for the wild ancestry of the
crop, their common occurrence in south-west Asia

and the Mediterranean basin affects our ability to
identify V. faba remains in archaeological excava-
tions. In many places today, these wild vetches are
locally common. They constitute an attractive ele-
ment for collection from the wild, and as Enneking
and Maxted (1995) argue, V. narbonensis even show
some signs of independent domestication. Therefore,
one can expect that seed remains of members of the
narbonensis group could occasionally appear in
archaeological contexts, particularly in the early
sites. Distinguishing between the faba crop and
these vetches is another story. Considerable similar-
ity exists—both in shape and size—between the
seeds of V. narbonensis (or some of its close rela-
tives), and those produced by small-seeded faba cul-
tivars. For these reasons, the separation of V. faba
from V. narbonensis complex, in archaeological
remains, might be difficult; more so, since charring
usually destroys the seed coats. Identification
becomes very unreliable when only few seeds (or
seed fragments) are available. Unfortunately such
paucity characterizes almost all finds retrieved from
early archaeological contexts.
As a result, archaeobotanical faba remains should
not be regarded as ‘V. faba’ but have to be consid-
ered as V. faba-like’.
Archaeological evidence
As already noted, the identification of V. faba-like
remains from early sites is complicated. All wild
progenitors closely resemble (morphologically, as
well as at the chromosome and molecular levels)
their domestic forbears, but the situation is even
more problematic in V. faba, as we do not know its
wild progenitor. Earliest charred V. faba-like seeds
appear in ca. 13,150–11,250 cal BP Iraq ed Dubb,
Jordan (Colledge 2001). However, it is more likely
these are either wild or intrusions. Later, during the
PPNA, faba-like seeds were found in ca. 10,700–
10,400 cal BP, Djade el Mughara (Willcox et al. 2008),
and the rich (437 seeds) find in ca. 10,600–10,250 cal
BP, Tell el-Kerkh, north-west Syria (Tanno and
Willcox 2006b). A few of these seeds are somehow
flat and wedge-shaped, and probably closely resem-
ble the wild progenitor (Tanno and Willcox 2006b,
p. 203). The only rich find available so far comes
from Middle PPNB, ca. 10,100–9,700 cal BP, Yiftah’el,
PULSES 91
Israel. Here, hoards of some 2,600 charred seeds
(without seed coats) were retrieved (Kislev 1985).
The seeds are small (5.5 × 4.7 × 4.0 mm), conform-
ing in size with those of wild V. narbonensis or
V. galilea—which are common plants in Yiftah’el
area. But compared to the more globular shape of
the seeds of these wild vetches, they are flatter and
somewhat thicker at the hilum’s side. The last two
features suggest that Yiftah’el’s remains belong to
V. faba. Yet, this evidence is not conclusive. Apart
from this single find, all other early records of
V. faba-like remains are indefinite. A few faba-like
seeds are available from several Pre-Pottery
Neolithic B farming sites in south-west Asia, such
as ca. 9,900–9,550 cal BP Jericho (Hopf 1983), Tell
Abu Hureyra (Hillman 1975; de Moulins 2000), Cap
Andreas-Kastros (van Zeist 1981), or Nevali Çori
(Pasternak 1998).
Already during the Neolithic, faba-like beans
appear also in several southern European countries:
in ca. 7,500–6,550 cal BP Scamuso, Italy (Costantini
et al. 1997), and in south-eastern Spain, like ca.
7,400–7,050 cal BP Coveta de l’Or (Hopf and
Schubart 1965; Lopez 1980), and adjacent Cova de
10 mm
5
0
Fig. 27 Carbonized seeds of faba-like bean, Vicia faba var. minor,
cotyledons only, no seed coat survived charring, Early Bronze Age
Zambujal, Portugal (Hopf 1981).
92 DOMESTICATION OF PLANTS IN THE OLD WORLD
Cendres (Buxó 1997). During the seventh millen-
nium it appears in ca. 6,500–5,500 cal BP Kaf That
el-Ghar, Morocco (Ballouche and Marinval 2003).
But these few, testa-less remains do not permit us to
decide whether they represent faba bean or mem-
bers of the V. narbonensis group. Similarly, the rare,
small, globular V. faba-type seeds reported from
Greek Late Neolithic, ca. 6,650–6,300 cal BP, Dimini
(Kroll 1979) and Sesklo (Hopf 1962, 1981a), also con-
form well with V. narbonensis group. V. faba appears
in several European Bronze Age sites (Fig. 27): ca.
4,950–3,350 cal BP Albertfalva, Hungary (Gyulai
2003), ca. 4,150–3,200 cal BP Canton of Grisons
(Graubünden), Switzerland (Jacomet et al. 1998,
1999), ca. 3,200–2,700 cal BP Late Bronze Age
Stillfried, Austria (Kohler-Schneider 2001, 2003),
and Late Bronze Age, ca. 2,905–2,869 cal BP, Grésine,
France (Bouby and Billaud 2001).
Significantly, many of the sites north of the Alps
are outside (or almost outside) the distribution
range of the members of the V. narbonensis group,
and this fact minimizes the possibility that the
excavated seeds belong to them. Furthermore, seed
size in several locations is already larger than that
found in the wild vetches. This strongly suggests
that we are confronted with domesticated V. faba.
Numerous finds of faba beans are available from
Iron Age and classical times in Europe and west
Asia, indicating that V. faba had been used as a
major food source.
When the information from the living plants and
from the archaeological remains is put together we
are led to the following conclusions:
(i) The wild progenitor of the domesticated faba
bean has not been satisfactorily identified. We
still need to discover the elusive 12-chromo-
some ancestor.
(ii) We still know very little about the beginnings
of V. faba domestication. A main reason being
that the early (PPNA/Early PPNB—twelfth–
eleventh millennia BP) finds identified as V.
faba seed might actually belong either to the
crop or to members of the V. narbonensis species
complex. More definite conclusions can only be
reached after additional well-preserved archae-
ological evidence is recovered or when the wild
progenitor of the crop is found.
Bitter vetch: Vicia ervilia
Bitter vetch, Vicia ervilia (L.) Willd., is a small, annual
diploid (2n = 2x = 14 chromosomes), self-pollinated
pulse with beaded pods and characteristic angular
seeds. At present it is grown as a minor crop in the
Mediterranean basin and south-west Asia. The
principal producer is Turkey. As its name implies,
its seeds are bitter and toxic to humans. However,
soaking, leaching, and steaming in water can
remove the poisonous substances, making them
palatable (van Zeist 1988; Enneking 1995; Miller
2002). Roman literature reports that in Classical
times, this vetch was utilized primarily as an ani-
mal feed since it is regarded as inferior for human
consumption and is only eaten by the very poor, or
in times of famine. Pliny the Elder (Natural History:
18.38) states that bitter vetch (‘ervi’) has medicinal
properties, like vetch (‘vicia’). He mentions that bit-
ter vetch ‘cured’ the Emperor Augustus.
The wild ancestry of the domesticated bitter vetch
is satisfactorily established (Zohary and Hopf 1973;
Ladizinsky and van Oss 1984). Truly wild forms of
this vetch, growing in primary habitats, are
restricted in their distribution and known from
Anatolia, Armenia, north Iraq, the Anti-Lebanon
(including Mt Hermon), and from Jebel Druz (Map
11). They show a striking morphological resem-
blance to the domesticated crop but differ from it by
their dehiscent pods and slightly smaller seeds.
Hybrids between the cultivars and the wild forms
are fully fertile. Weedy races and feral forms of
V. ervilia occasionally infest grain crops and edges
of cultivated fields throughout south-west Asia and
Greece.
Unfortunately, we did not find molecular research
aiming to support the identification of the wild pro-
genitor and whether it had a monophyletic or
polyphyletic origin.
Archaeological evidence
The earliest finds of bitter vetch appear in
Mousterian, ca. 65,000–48,000 cal BP, Kebara Cave,
Israel (Lev et al. 2005). Afterwards, it was found in
several Syrian sites. First in Epi-Paleolithic, ca.
13,400–11,350 cal BP, Tell Abu Hureyra (Hillman
1975; Hillman et al., 1989, 2001), and ca. 11,800–

0 100 200 miles
0 200 400 km
Map 11 Geographical distribution of wild bitter vetch, Vicia ervilia (based on Zohary and Hopf 1973; Townsend 1974; Ladizinsky and van Oss
1984; Gabrielian and Zohary 2004).
11,300 cal BP Mureybit (van Zeist and Casparie
1968; van Zeist and Bakker-Heeres 1986), and later
in PPNA, ca. 11,500–11,000 cal BP, Jerf el-ahmar
(Willcox 2002; Willcox et al. 2008, 2009), and ca.
10,700–10,400 cal BP Djade el Mughara (Willcox
et al. 2008). Later, seeds of bitter vetch first appear
in several agricultural settlements in Turkey.
Thousands of carbonized seed of V. ervilia were
found in various phases of Aceramic, ca. 10,250–
9,550 cal BP, Çayönü (van Zeist 1972; van Zeist and
de Roller 1991–2, 2003), and in ca. 10,100–9,450 cal
BP Aşikli Höyük (van Zeist and de Roller,1995).
Bitter vetch is also common in aceramic, ca. 9,450–
8,450 cal BP, Can Hasan III (Hillmam 1972, 1978). In
addition, it appears in contexts of ca. 9,350–8,950 cal
BP Çatalhöyük (Helbaek 1964a; Fairbairn et al. 2002,
2005, 2007), ca. 8,550–8,150 cal BP Erbaba (van Zeist
and Buitenhuis 1983), and ca. 8,200–7,800 cal BP
Hacilar (Helbaek 1970). In addition V. ervilla was
very important in the early phase, ca. 8,500–8,000
cal BP, of Gritille (Miller 2002), and ca. 7,800–7,350
cal BP Girikihaciyan (van Zeist 1979–80). A few
seeds, and even a single pod remain, were found in
PULSES 93
ca. 7,950–7,150 cal BP Aratashen and Aknashen,
Armenia (Hovsepyan and Willcox 2008). However,
it is impossible to determine definitely whether
these remains represent wild or domesticated mate-
rial. Later finds, such as Middle Bronze Age Shiloh
(Kislev 1993) and Beit She’an (Simchoni et al. 2007),
Israel, are also relatively few.
Considerable amounts of charred seed of V. ervilia
have been discovered in late Neolithic and Bronze
Age Greece (Fig. 28). The earliest rich (we refer to
the large quantity as indication for the domestica-
tion status of the find) and well-identified bitter
vetch finds come from ca. 8,400–8,100 cal BP Nea
Nikomedeia (van Zeist and Bottema 1971), followed
by several late Neolithic and early Bronze Age finds
(Renfrew 1979; Kroll 1983). Bulgarian sites show
similar situation. Large amounts of bitter vetch
were found in early Neolithic, ca. 8,000–7,550 cal BP
Karanovo, followed by storages of it in Middle
Neolithic, late Neolithic, Late Eneolithic, and Early
Bronze Age strata (Thanheiser 1997; Marinova 2004,
2006). Huge, pure hoards of bitter vetch grains were
discovered in Middle Neolithic, ca. 7,450–7,300 cal
94 DOMESTICATION OF PLANTS IN THE OLD WORLD
Fig. 28 Carbonized seeds of bitter vetch, Vicia ervilia, cotyledons only, no seed coat survived charring, Late Neolithic Dimini,
Greece (Kroll 1979).
BP, Azmaška Mogila (Hopf 1973a; Renfrew 1979).
Carbonized remains of this pulse are also very fre-
quent in Bulgaria throughout the Eneolithic and the
Bronze Age (Januševič 1978; Renfrew 1979) and
sometimes constitute the main plant material
retrieved from the sites.
Outside Turkey, Greece, and Bulgaria, remnants
of the crop are much less common. Some seeds were
found in ca. 7,550–6,450 cal BP Middle Neolithic
Sammardenchia, Italy (Pessina and Rottoli 1996;
Rottoli 2005; Rottoli and Pessina 2007), and contem-
poraneous Grotta dell’Uzzo, Sicily (Costantini
1989). However, over large areas in West Europe
and the Mediterranean Basin they are missing alto-
gether. A few grains of bitter vetch were retrieved
from Late Neolithic Gomolava, in Serbia (van Zeist
1975). Several finds are reported from Rumania, like
ca. 6,350–6,150 cal BP Tell Hârşova (Cârciumaru
1996; Monah 2002; Monah and Monah 2008), includ-
ing an almost pure hoard from Gumelnitsa culture
in Cǎscioarele (Cârciumaru 1996). A few seeds were
uncovered in Tripolye culture sites in Moldavia and
the Ukraine (Wasylikowa et al. 1991). Some rare
finds were also discovered in ca. 5,650–5,300 cal BP
5 mm
Naqada, Egypt (Wetterstrom 1993). Later, small
amounts of bitter vetch have been found in ca.
3,350–3,250 cal BP Bölcske-Vörösgyír, Hungary
(Berzsényi and Gyulai 1998), ca. 3,200–2,700 cal BP
Stillfried, Austria (Kohler-Schneider 2001, 2003),
and ca. 2,905–2,869 cal BP Grésine, France (Bouby
and Billaud 2001). Yet on the whole, remains of
V. ervilia in Neolithic and Bronze Age contexts show
a distinct regional pattern. The cultivation of this
pulse in the past seems to have been heavily cen-
tered in Anatolia and the Balkans.
The combined evidence from the living plants
and from the archaeological remains indicates that
bitter vetch was a founder crop, part of south-west
Asian Early Neolithic crop assemblage. This pulse
was probably taken into domestication in Anatolia
or the Levant; i.e. in the general area in which it still
grows wild today. However, since there are no reli-
able diagnostic traits by which wild and domesti-
cated forms of bitter vetch (in archaeological
remains) can be distinguished from each other, the
early archaeological finds could be either. The size
and purity of the numerous samples of this pulse
retrieved from the Neolithic and Bronze Age sites in

the Balkans and Turkey, strongly suggests that
V. ervilia was domesticated at that time and place.
We know very little about the mode of utilization of
this bitter seeded legume by the Neolithic and
Bronze Age farmers.
Common vetch: Vicia sativa
Common vetch, Vicia sativa L., is another member of
the genus Vicia that is frequent in Mediterranean
grain agriculture. This vetch is very variable, includ-
ing wild forms, weeds, and cultivars. In modern
times, it constitutes a minor crop cultivated for hay
and for seeds, grown almost exclusively for animal
feed. Similar to bitter vetch, the seeds are not attrac-
tive for human consumption. Common vetch is also
a frequent contaminant of lentil and bitter vetch
cultivation. Seeds of the latter pulses sold in local
markets frequently contain scattered V. sativa seed.
The domesticated V. sativa is a self-pollinated,
diploid plant (2n = 2x =12 chromosomes) with
straggling or ascending growth habit and rounded,
somewhat compressed, smooth seeds (4.5–7.0 mm
in diameter). The cultivars are closely related to an
extraordinarily variable (and chromosomally com-
plex) aggregate of wild types and weedy forms, the
distribution of which is centred in the Mediterranean
basin (Zohary and Plitman 1979; Maxted 1995). All
are now grouped, together with the cultivars, in the
V. sativa complex. Most cultivars, together with
morphologically closely related weeds and escap-
ees are placed in V. sativa subsp. sativa.
Archaeological evidence
Carbonized seeds of V. sativa have been reported
from several Neolithic and Bronze Age sites in
south-west Asia and Europe. Since seed sizes of
weedy forms and wild types overlap those found in
the cultivars, it is difficult to conclude whether the
remains represent domesticated forms, weedy con-
taminants or collection from the wild. Definite indi-
cations of sativa cultivars are available only from
Roman times onward.
The earliest archaeological records of V. sativa
comes from the Natufian and Neolithic Tell Abu
Hureyra, Syria (Hillman 1975, 2001; Hillman et al.
1989; de Moulins 2000) and from pre-ceramic
PULSES 95
Neolithic Can Hasan III, Turkey (Hillman 1972).
They are followed by several records from Neolithic
and Eneolithic Bulgaria (Renfrew 1973), Italy
(Rottoli 1993; Pessina and Rottoli, 1996; Rottoli 2002,
2005; Rottoli and Pessina 2007), Hungary (Hartyáni
and Nováki 1975), and Slovakia (Hajnalová, 1989).
Common vetch is also reported from Predynastic
Maadi, Egypt (van Zeist and de Roller 1993, 2003),
and from several Bronze Age contexts such as the
second half of the fifth millennium BP Ak-Tepe near
Ashkabad, Turkmenia (Lisitsina and Prishchepenko
1977, as cited by Schultze-Motel 1974b), and from
Slovakia (Kühn 1981).
Grass pea: Lathyrus sativus
Grass pea or chickling vetch, Lathyrus sativus L., is
a minor pulse crop of traditional agriculture in the
Mediterranean basin, south-west Asia, Ethiopia,
and the north-western parts of the Indian subcon-
tinent (Kearney and Smartt, 1995). India is the
main producer today, and there L. sativus is appre-
ciated for its ability to grow in dry places and on
poor soils. Grass pea is a diploid (2n = 2x = 14
chromosomes), predominantly self-pollinated
annual with branched, straggling, or climbing
habit, blue (sometimes violet or white) flowers and
characteristic smooth seeds with pressed sides.
Recent cytogenetic studies found out a mixed mat-
ing system with an outcrossing rate of 36%
(Gutiérrez-Marcos et al. 2006). At present the pulse
is used mainly as animal feed, though in India and
Ethiopia it serves also as an article of human diet
and is consumed by the very poor, and in times of
famine. For this reason, as well as its palatability,
L. sativus is currently examined for serving as a
crop replacement, especially in depleted fields
(Polignano et al. 2009). Consuming large amounts
of grass pea seed can be dangerous. The seed con-
tains a water-soluble non-protein neurotoxin
amino acid that causes Lathyrism, a crippling neu-
rological disorder. Because the neurotoxin is a
water-soluble, one hour boiling of the seeds
removes some 70% of it (Jha 1987).
Domesticated grass pea shows close morphologi-
cal resemblance to a group of wild Lathyrus species
distributed over the Mediterranean basin and south-
west Asia. It seems closest to L. cicera L., a wild grass
96 DOMESTICATION OF PLANTS IN THE OLD WORLD
pea growing in several eastern Mediterranean and
south-west Asian countries (Greece, Turkey, North
Iraq, North Iran, Transcaucasia, and Egypt). In these
countries, L. cicera abounds as a weed in cereal culti-
vation. L. cicera can be crossed with L. sativus, and
the hybrids produced are at least partly fertile
(Kearney and Smartt 1995). Therefore, L. cicera could
be considered as the wild progenitor from which the
domesticated grass pea was derived. But L. cicera is
very variable, and is cytogenetically complex.
Moreover, several other Lathyrus species also show
close morphological affinities with L. sativus.
Therefore, the final determination of ancestry in this
crop has to depend on critical cytogenetic and
molecular tests. These are as yet unavailable. As in
many other grain legumes, domestication of the
grass pea resulted in retention of the seeds in the
pods, loss of seed dormancy, and an increase in seed
size. Seeds of L. sativus cultivars are somewhat larger
(6–8 mm in diameter) than those of their wild rela-
tives (5–6 mm).
Unfortunately, we did not find any molecular
research seeking to support the identification of the
wild progenitor and whether it was a monophyletic
or polyphyletic mode of origin.
Archaeological evidence
Carbonized seeds of grass pea appear in several
south-western Asian, Aegean, and west Mediter-
ranean Neolithic settlements. To-date, the oldest
remains come from ca. 10,250–9,550 cal BP Çayönü,
Turkey (van Zeist 1972; van Zeist and de Roller
1991-2, 2003) and ca. 9,450–9,300 cal BP Jarmo, Iraq
(Helbaek 1960). A few grass pea seeds were also dis-
covered in Aceramic Neolithic, ca. 8,350–7,750 cal
BP, Tepe Sabz, Iran (Helbaek 1970), and Nevali Çori,
Turkey (Pasternak 1998). It is impossible to decide
whether they represent domesticated forms or col-
lection from the wild. Over 800 charred seeds were
found in flotation samples from ca. 8,500–7,500 cal
BP Final PPNB Gritille, on the Euphrates, south-east
Turkey (Miller 1991, 2002). Such a large quantity is
likely to be a crop. The bulk of Neolithic grass pea
finds comes, however, from Greece and Bulgaria
(Kislev 1989a). About 200 seeds were discovered in
ca.8,000 BP Prodromus (Halstead and Jones 1980). A
seven-litre hoard was uncovered in ca. 6,500–5,000
cal BP Late Neolithic Servia, where Hubbard
(Hubbard 2000), claim by ‘circumstantial evidence’
that toxins were leached by soaking, and some more
in Late Neolithic, ca. 6,650–6,300 cal BP, Dimini
(Kroll 1979). Numerous seeds were also retrieved
from early Neolithic, ca. 7,850–7,500 cal BP, Azmaška
Mogila (Hopf 1973a), as well as in contemporane-
ous storage from Kapitan Dimitrievo, Bulgaria
(Marinova 2006, forthcoming). The large stores of
seeds found at Late Chalcolithic Kuruçay (3,700–
3,200 cal BC) in south-west Turkey are consistent
with an Aegean and south Balkan centre of impor-
tance for this crop (Nesbitt 1996).
A few Lathyrus-type carbonized seeds were
uncovered also in several Impressed Ware and
Chasséen sites in south France (Courtin and Erroux
1974). In the Bronze Age, remains of grass pea con-
tinue to appear in south-west Asia and in south-
eastern Europe (for a compilation of data see Kislev
1989b). They were discovered, for example, in Early
Bronze Age Lachish, Israel (Helbaek 1958), in Tell
Basmosian, Iraq (Helbaek 1963), in ca. 4,500–4,000
cal BP Early Bronze Age Kastanas, Greece (Kroll
1983; see also Fig. 29), and in Middle Bronze Age
Tiszaalpár Várdomb, Hungary (Hartyáni 1982). In
most of these finds the seeds are small and the seed
coats are missing. It is therefore impossible to decide
whether they represent a collection from the wild,
an infestation of L. cicera-type weeds in fields of
other crops, or domesticated forms. Yet the sheer
quantities of this pulse in Gritille, Prodromus,
Servia, Azmaška Mogila, Kapitan Dimitrievo, and
Dimini, seem to indicate domestication. Finally,
grass pea is also reported from several Harappa cul-
ture sites in Pakistan and north-west India (Fuller
2000).
The grass pea might belong to a group of founder
crops that started Early Neolithic crop assemblage,
or it may have been added soon after the establish-
ment of grain agriculture. When, where, and how
this happened is still difficult to say, more so since
the wild relatives of this pulse are widely distrib-
uted over the entire Mediterranean basin. The early
finds from Çayönü and Jarmo, as well the large
number in Gritille, suggest that L. sativus may have
been taken into domestication in south-west Asia,
yet the proposal that it is south Balkan in origin
(Kislev 1989b) is another valid option.

0 10 mm
Fig. 29 Carbonized seeds of grass pea, Lathyrus sativus, cotyledons
only, no seed coat survived charring, Bronze Age Kastanas, Greece
(Kroll 1983).
Spanish vechling: Lathyrus clymenum
Lathyrus clymenum L. is a Mediterranean grain crop
of restricted distribution, still grown today in sev-
eral Aegean islands such as Thera (Santorini), Anafi,
and Karpathos (Sarpaki and Jones 1990). Like other
legumes, the seeds are protein-rich and highly
nutritious. L. clymenum is known as a fodder crop,
as well as a staple food. Like Lathyrus, the seeds are
toxic and might cause lathyrism (Melamed et al.
2009). Wild and weedy forms of L. clymenum are
widely distributed in the western and central parts
of the Mediterranean basin, from west Turkey to the
Iberian Peninsula, and from Cyrenaica to Morocco.
Previously, the living plant was unknown in the
Levant (Kislev, 1993), but recently Melamed et al.
(2009) reported four wild populations in Israel.
PULSES 97
Archaeological evidence
Relative to other legumes, finds of L. clymenum are
rather rare. The earliest and largest find came from
ca. 4,850–4,550 cal BP Early Bronze Age II levels of
Yenibademli Höyük on the Gökçeada island, Turkey
(Oybak-Dönmez 2005). Seeds of L. clymenum have
been discovered in a storage room in ca. 3,900–3,700
cal BP Middle Bronze Age IIA Tel Nami, a coastal
site in Israel, suggesting the transport of this pulse
from the Aegean basin into the Levant by maritime
traders (Kislev 1993). Large quantities of charred
seed of L. clymenum, placed also in storage jars, were
discovered in Akrotiri, Thera Island, in a house
destroyed by the volcanic eruption that devastated
this island in ca. 3,578 cal BP (Sarpaki and Jones
1990). These authors also reported seeds of this
pulse among plant remains retrieved from Late
Minoan II Knossos, Crete, and contemporary
Phylakopi, Melos. These finds establishes L. cly-
menum as a local, Aegean, Bronze Age domestic
plant, which survives today only as a relic.
Fenugreek: Trigonella
foenum-graecum
Fenugreek, Trigonella foenum-graecum L., (Legumi-
nosae) is a minor pulse crop of Mediterranean
agriculture grown in southern Europe, south-west
Asia, Ethiopia, and the northern parts of the Indian
subcontinent.
The seed is widely used as a condiment and as an
important flavoring ingredient for the preparation
of curry powder and soups. Fenugreek is also used
today as an effective nitrogen fixer and effective soil
renovator (Small 1997). It is an annual, diploid
(2n = 2x = 16 chromosomes), self-pollinated, leg-
ume with long curved, linear-lanceolate, stiff pods
(mostly growing to a height of 30–60 cm), and char-
acteristic roughly quadrangular seeds which have
prominent radicles (‘beaks’). The plant has a long
taproot with numerous fine laterals bearing
nitrogen-fixing nodules.
The crop shows very close morphological resem-
blance to a group of wild Trigonella species distrib-
uted over the east-Mediterranean basin and
south-west Asia (Širjaev 1932; Ladizinsky and
Vosa 1986; Small 1997). It is not yet clear which
98 DOMESTICATION OF PLANTS IN THE OLD WORLD
member of this group (Section Foenum graecum
Širjaev) gave rise to the domesticated fenugreek.
Yet since all these wild relatives have small distri-
bution areas and are restricted to south-west Asia
and the eastern parts of the Mediterranean basin, it
seems likely that T. foenum-graecum was brought
into domestication somewhere within this geo-
graphic region.
Archaeological evidence
Charred fenugreek seeds are quite rare in archaeo-
botanical assemblages. Some were retrieved from
south-west Asian sites such as ca. 6,000 BP Tell
Halaf, Iraq (Neuweiler 1935), Iron Age Lachish,
Israel (Helbaek, 1958), and Iron Age Deir Alla,
Jordan (Neef 1989). In addition, desiccated seeds
were found in ca. 5,450–5,650 cal BP Ma’adi
(Neuweiler 1946), and in several baskets, mixed
with coriander, in eighteenth dynasty, ca. 1,325 BC
Tutankhamun tomb (Neuweiler 1946; Hepper 1990;
de Vartavan et al. 2010) in Egypt. The available
archaeological evidence on this pulse is still frag-
mentary, yet it indicates that fenugreek was part of
south-west Asian agriculture already in the Early
Bronze Age.
0 100 200 miles
0 200 400 km
Map 12 Geographical distribution of wild white lupin, Lupinus albus subsp. graecus [= L. graecus] (based on Gladstones 1974).
Lupins: Lupinus
Several annual members of the genus Lupinus L., dis-
tributed over the Mediterranean basin, have been intro-
duced into domestication and are grown today either
as grain crop or as forage plants and green manure.
Prominent among them are the white lupin, L. albus
L., the yellow lupin, L. luteus L., and the narrow-leaved
lupin, L. angustifolius L. (Gladstones 1974; Hanelt 1986;
Hill 1995; Cowling et al. 1998). All are vigorous growers
and produce large, attractive seed. However, their use
is complicated by the fact that lupins generally contain
bitter alkaloids which are difficult to remove.
Domestication of these grain pulses required an initial
selection for non-bitter seed and non-dehiscent pods.
The white lupin, L. albus [= L. termis Forssk.], is a
traditional Mediterranean pulse crop, which has
been grown in this area since antiquity. Even today
the white lupin is an appreciated food crop and it is
still commonly cultivated in some Mediterranean
countries,particularly in Egypt.
The wild progenitor of the domesticated white
lupin is well identified. The cultivars are inter-fer-
tile and chromosomally identical (2n = 2x = 50 chro-
mosomes) with wild forms, which are native to the
southern part of the Balkans, the Aegean islands,

and the Aegean belt in west Turkey (Map 12). They
also extend to the Sila Mountains in south Italy (but
are very rare there). These were traditionally called
L. graecus Boiss. and Spruner. More recently they
were taxonomically included in the crop complex as
L. albus subsp. graecus (Boiss. and Spruner) Franco
and Silva. Wild forms are characterized by dehis-
cent pods, smaller, more bitter seed, and thicker,
impermeable, pigmented seed coats.
Archaeological evidence
Earliest L. albus find is reported from ca. 5,450–5,650
cal BP, Late Neolithic, Predynastic Maadi, Egypt,
but this single seed cannot serve as evidence for its
domestication status and cultivation in Predynastic
Egypt (van Zeist and de Roller 2003). Bronze Age
remains of white lupin were found in Thera island,
PULSES 99
Greece (Sarpaki 1992b), and a single find is availa-
ble from Late Bronze Age Hala Sultan Tekke,
Cyprus (Hjelmqvist 1979a). Desiccated seed of L.
albus—with their characteristic smooth, white
coats—were reported from several Roman sites in
Egypt (Germer 1985). They were substantiated by
additional finds in Qasr Ibrim (Rowley-Conwy
1994), Mons Claudianus (van der Veen 1998, 2001a),
and Berenike and Shenshef (Vermeeren and
Cappers 2002; Cappers 2006).
In conclusion, the distribution of the wild forms
of L. albus seems to indicate that this pulse was
taken into domestication somewhere in the Aegean
basin. When this happened is hard to say since pre-
Roman finds are very scanty. However, the total
absence of this crop in Neolithic sites in the
Mediterranean basin and Anatolia suggests a late
domestication.
 C H A PTER 5
Oil- and fibre-producing crops
Indigenous plants producing oil and/or fibre were
also taken into cultivation at the start of agriculture
in south-west Asia and the Mediterranean basin
during the middle-late PPNB, ca. tenth-ninth mil-
lennia BP. Oil and fat are an important component
of human’s diet, and most societies have a source of
fat/oil from plants or animals. Thus, these sources
are always an important component of agricultural
systems.
Flax is apparently the earliest and the best-docu-
mented oil and fibre crop. Remains of linseed and
fragments of linen indicate that flax belongs to the
first wave founder crops of south-west Asia and
that it maintained its leading role in the Old World’s
fibre and oil production crop from Neolithic times
to the early years of the twentieth century. Two
other important fibre crops are hemp and cotton.
The evidence for the beginnings of their domestica-
tion is still insufficient, but it is clear that they were
introduced into cultivation outside south-west Asia.
However, both hemp (in East Asia) and cotton (in
the Indian subcontinent) were already under culti-
vation in the fourth millennium BP.
Several oil plants seem to have entered cultiva-
tion not directly, but first by evolving weedy forms.
They were added to the crop assembly only after
the firm establishment of the principal founder
crops such as wheat, barley, lentil, pea, and flax. The
case of the gold of pleasure, Camelina sativa (p. 111)
is well documented. Several other cruciferous plants
(e.g. Sinapis and members of the genus Brassica)
seem to have followed a similar mode of evolution
under domestication.
In a similar way to the cereals and the pulses,
the introduction of oil-bearing seed plants into
agriculture triggered the evolution of character-
100
istic domestication traits. Under the new system
of sowing, reaping, and threshing, selection
brought about the breakdown of the wild mode
of seed dispersal and the retention of the seed on
the mother plant. Indeed, in most domesticated
oil crops the fruits do not dehisce. In others there
is a delay in fruit opening. Another conspicuous
evolutionary trend is yield increase. Compared
with their wild relatives, most cultivars of oil-
bearing seed crops have frequently larger inflo-
rescences and/or bigger fruits (with larger or
more numerous seeds). The wild mode of germi-
nation inhibition has also broken down and as a
result, seed coats have become thinner. Finally,
cultivated varieties frequently show increased
oil content. Yet this is not a general rule. Seeds of
some wild types are as rich in oil as their culti-
vated counterparts. In fibre-producing plants the
accent is on long, strong fibres. If stem fibres are
used (as in flax and hemp), long and straight
plants evolve. If lint is the goal (as in cotton),
mutations determining long lint (not found in
the wild) are selected.
We still know very little about how oil and fibres
were extracted from plants in the early phases of
agriculture. However, it seems reasonable to assume
that:
(i) Oil could be obtained by decantation; i.e. by
crushing the seed, pouring hot water on the
meal, and scooping the oil after setting. In some
cases, before crushing, the seeds were softened
by allowing them to absorb water.
(ii) Flax and hemp stems were retted and broken in
order to free the fibres. Possibly both technolo-
gies antedate agriculture.

Flax: Linum usitatissimum
Flax, Linum usitatissimum L. (Lineaceae), is an
annual, self-pollinated crop with characteristic slen-
der, strong stems and rounded capsules which (in
cultivated forms) do not dehisce, but retain the oval,
compressed, shining seed. The crop is diploid
(2n = 2x = 30 chromosomes) and predominantly
self-pollinated. Consequently, in this crop, variation
has been moulded in the form of numerous true
breeding lines and aggregates of land races. Two
specializations are apparent: (i) oil varieties that are
relatively short (30–70 cm), branched, and usually
bear large seed. They are grown for high yield of
linseed; (ii) fibre varieties that are taller, sparsely
branched, and usually bear small seed. Transitional
forms cultivated for both oil and fibre occur as
well.
Flax was a principal oil and fibre source in the
Old World and probably the earliest cultivated
plant used for weaving clothes. Until recently,
flax was extensively grown all over the area from
the Atlantic coast of Europe in the west, to Russia
and India in the east, and Ethiopia in the south
(Durrant 1976). In antiquity, flax fibres (which are
stronger than cotton or wool) were the principal
vegetable fibre used for weaving textiles in
Europe and western Asia (Plate 12). However,
from the industrial revolution onwards, flax was
gradually replaced by cotton imports, and cur-
rently it has been almost completely replaced by
cotton and synthetic fibres. The seed contains
about 40% oil and in peasant communities, lin-
seed was used as a source for edible oil and high-
grade lighting oil.
Cooking oil is obtained from linseed by cold
pressing. Drying oil (used for paints and varnishes)
is produced by hot treatment of the seed before
pressing (hot pressing).
The fibres for spinning are obtained from fibre-
cell bundles running the length of the stem and
forming a ring in the cortex. The stems are har-
vested before the maturation of the seed.
Traditionally they were first dried, and then
immersed (‘wetted’) in water to allow the microbial
decomposition (‘retting’) of the pectin connecting
the fibres with other cells and tissues of the stem.
After retting, the stems were dried and the fibres
OIL- AND FIBRE-PRODUCING CROPS 101
(averaging 4 cm in length) were separated by
pounding (‘breaking’) and combing.
Flax is represented in archaeological excavations
both by seed and, occasionally, capsules, and by
remnants of stems or textiles. In the latter, fibres can
be identified microscopically if they are not carbon-
ized (Körber-Grohne 1991).
Wild ancestry
Linum L. is a relatively large genus comprising some
200 species spread over the temperate, Mediter-
ranean, and steppe belts of the northern hemi-
sphere. Domesticated flax, Linum usitatissimum L.,
is most closely related to wild L. bienne Mill. [= L.
angustifolium Huds.]. These two flaxes have the
same chromosome number (2n = 2x = 30 chromo-
somes), inter-cross readily, and are fully inter-fertile
(Gill and Yermanos 1967). L. bienne (Fig. 30), with its
characteristic strong branches, blue flowers, and
dehiscent capsules is widely distributed over west
Europe, the Mediterranean basin, North Africa,
south-west Asia, and Caucasia (Map 13). Some wild
forms are biennial or perennial, others are annual,
all are predominantly self-pollinated. L. bienne
grows mainly in wet places such as moist grassy
areas, springs, seepage areas on rocky slopes, moist
clay soils, and marshy lands. Occasionally it is also
found as a weed in fields and at the edges of culti-
vation (including flax cultivation). In such places,
hybridization between wild and domesticated
flaxes occurs sporadically. On the basis of its close
morphological and genetic affinities to the culti-
vated crop, L. bienne is identified as the wild pro-
genitor of L. usitatissimum (Diederichsen and
Hammer 1995). It is therefore correct to regard this
wild flax as part of the crop species; i.e. as the wild
subspecies of the crop. As in many other grain
crops, the main changes under domestication are
the shift to non-splitting capsules, the increase of
seed size, and the selection for higher oil yield, or
longer stems with a high amount of long fibres.
Fu (2005) studied 2727 worldwide accessions of
flax using RAPD markers, and Allaby et al. (2005)
have studied the sad2 locus in both wild and domes-
ticated accessions of flax. In addition, Fu and Allaby
(2010) assessed phylogenetic relationships among
twenty-nine Linum accessions from sixteen wild
102 DOMESTICATION OF PLANTS IN THE OLD WORLD

5 cm

A B

0 5 mm

Fig. 30 Wild flax, Linum usitatissimum subsp. bienne [= L. bienne]. A–Flowering and fruiting stem; B–Capsule (Zohary 1972, plate 374).

0 100 200 miles
0 200 400 km
Map 13 Geographical distribution of wild flax, Linum usitatissimum subsp. bienne [= L. bienne].
and domesticated species, based on four non-cod-
ing regions of chloroplast DNA sequences. The dif-
ferences in genetic diversity between and within
accessions revealed in these studies suggest that
domesticated flax is of a monophyletic origin, it has
probably evolved from a single domesticating
event, although the geographic location of this
event is still obscure. In addition, these data indi-
cate that flax domestication selected for the larger,
oil-rich, seeds of the oil producing variety, rather
than its fibers (Allaby et al. 2005; Fu and Allaby
2010). In other words, these studies suggest that flax
was first domesticated as an oil plant.
Archaeological evidence
Flax was apparently used by humans already before
its domestication. Archaeological finds of seeds are
more likely to result from cultivation (or gathering)
for seed consumption (as McCorrison 1997), since
harvesting for fibers often occurs before seed matu-
ration. Recently, twisted and dyed flax fibres were
reported in Upper Palaeolithic, ca. 30,000 year-old,
Dzudzuana Cave, Georgia (Kvavadze et al. 2009,
OIL- AND FIBRE-PRODUCING CROPS 103
but see discussion by Bergfjord et al. 2010; Kvavadze
et al. 2010). The oldest wild linseed remains retrieved
from archaeological sites in south-west Asia come
from ca. 11,800–11,300 cal BP Tell Mureybit (van
Zeist and Casparie 1968; van Zeist and Bakker-
Heeres 1986). Soon after, seeds of flax were found in
many of the Pre-Pottery Neolithic B farming vil-
lages that appeared in the Fertile Crescent from
10,500 cal BP onward (Map 1). Some of the earlier
finds come from Çayönü, Turkey (van Zeist 1972;
van Zeist and de Roller 1991-2, 2003), Tell Aswad,
near Damascus, Syria (van Zeist and Bakker-Heeres
1985), Ali Kosh, Iran, (Helbaek 1969), Jericho, Israel
(Hopf 1983), and ‘Ain Ghazal, Jordan (Rollefson
et al. 1985). The seeds are still small, similar in size
to those of wild bienne forms. Yet they are almost
always associated with domesticated wheats and
barley.
Fragments of a capsule from 9,900–9,550 cal BP
Middle PPNB Jericho (Hopf 1983, Fig. 31) are prob-
ably the earliest indication we have today of domes-
ticated flax. Another indication of early flax
domestication comes from linseed remains recov-
ered from Late PPNB, 8,700–8,600 cal BP, levels of
104 DOMESTICATION OF PLANTS IN THE OLD WORLD
Tell Ramad, Syria (Fig. 32). The calculated size of
the seeds from this site, corrected for charring
shrinkage, ranges from 3.2 to 4.1 mm in length. This
is already within the size class of the L. usitatissi-
mum seed, the lower limit of which lies at 3.0 mm. It
is therefore an attractive indication for flax domesti-
cation under rain-dependent conditions before ca.
8,600 cal BP (van Zeist and Bakker-Heeres 1975).
Such domesticated linseeds were found in Pottery
Neolithic Nahal Zehora, Mount Carmel, Israel, ca.
8,150–7,850 cal BP (Kislev and Hartmann forthcom-
ing). In the Mesopotamian basin, linseed sizes in ca.
8,350–7,750 cal BP Tell Sabz, Iran (Helbaek 1969)
and in Halafian, ca. 7,750–7,250 cal BP Arpachiya
(Helbaek 1959a, 1959b) are even bigger (4.7–4.8 mm
long). As argued by Helbaek, these large seeds indi-
cate advanced domestication and demonstrate that
flax was part of the irrigated grain agriculture sys-
tem that evolved in this region. In Choga Mami,
Iraq, Helbaek (1972) reports a rare find in the earlier,
ca. 7,800–7,750 cal BP, stratum, which became fre-
quent at the following stratum, dated to the second
half of eighth millennium. Linseed also appears in
several later south-west Asian Neolithic and Bronze
Age sites. A complete capsule was retrieved from
Early Bronze Age, ca. 3,500–3,300 cal BP, Jericho
(Fig. 31).
Flax is associated with the spread of Neolithic
agriculture from the Fertile Crescent ‘core area’ into
Europe and the Nile Valley (Map 2—Plate 6) from
the early Neolithic (eighth millennium BP) onward.
Linseed has been recovered from sites dating from
the first half of the eighth millennium BP in the
0 4
Fig. 31 Carbonized capsule of domesticated flax, Linum usitatissimum subsp. usitatissimum, Bronze Age Jericho (Hopf 1983).
Mediterranean Basin and southern parts of Europe.
Such sites are Knossos, Crete (Sarpaki 2009), a few
sites in Thessaly, Greece (Kroll 1981b, 1991), La
Marmotta, Italy (Rottoli 1993, 2002), and as far as
Mohelnice, Moravia (Opravil 1979, 1981; Kühn
1981).
Further north, in central and western Europe, the
earliest cultivated flax finds are from the second
half of the eighth millennium BP. They belong to the
earliest Linearbandkeramik, food production cul-
ture in the area. Sites include those in the
Aldenhovener Platte (Knörzer 1973, 1974, 1997) and
Rosdorf (Kirleis and Willerding 2008) in Germany,
Brześć in Poland (Bieniek 2007), and North Meseta
sites (including Cueva de La Vaquera, La Lámpara,
and Revilla del Campo) (López et al. 2003; Stika
2005; Peña-Chocarro 2007), Spain. The earliest find
in Austria, however came from Late Linear-
bandkeramik, ca. 7,150–6,800 cal BP, Schletz
(Schneider 1994; Kohler-Schneider 2007). In
Switzerland it occurs among plant remains in many
lake-shore settlements (Jacomet 2007) but arrives
only in the second half of the seventh millennium
BP. It is still rare in the relatively early (ca. 6,250 cal
BP) Egolzwil 3 culture (Bollinger 1994; Jacomet
2007), but becomes frequent in the later Neolithic
and Bronze Age times, in the settlements at Lake
Zürich or at Late Neolithic Lake Biel (Ammann et al.
1981; Brombacher 1997, 2000; Brombacher and
Jacomet 2003). At the second half of the sixth mil-
lennium BP, a few seeds, capsules, and textile were
found in Station III de Clairvaux, France (Lundström-
Baudais 1984). At Late Bronze Age, 2,905–2,869
8 mm

Ramad
Ras Shamra
0 3 mm
Fig. 32 Carbonized seeds of domesticated flax, Linum usitatissimum subsp. usitatissimum, from Neolithic Ramad, and Ras Shamra, Syria (van
Zeist and Bakker-Heeres 1975).
(dendrodated) BP, Grésine, France, linseed and cap-
sules were frequently found (Bouby and Billaud
2001). Flax is similarly common in lake-shore settle-
ments in fifth millennium BP south Germany, such
as Ödenahlen at the Federseeried (Maier 1995). At
the same time, linseed was found for the first time
in Late Neolithic Aartswoud, The Netherlands (Pals
1984). Linseed was also retrieved from late Neolithic
and Early Bronze Age northern Italy; that is, from
Lagozza (Buschan 1895), from Valeggio (Villaret-
von Rochow 1958), and Frioli (Pessina and Rottoli
1996). In Rumania, lumps of charred linseed were
discovered in Sucidava-Celei (Cârciumaru 1996), in
contexts dated to the transition between the
Eneolithic and the Bronze Age (ca. 4,850–4,150 cal
BP). The earliest linseed, including fibers, in Spain
and Portugal were uncovered in Chalcolithic (ca.
4,800–4,200 cal BP) Buraco da Pala, Portugal and
several El Argar culture, ca. 4,150–3,350 cal BP, Early
Bronze Age sites in Spain.
The earliest documentation of flax cultivation in
Scandinavia appears in Bronze Age, ca. 3,150–2,950
cal BP, Borge vestre, Norway (Sandvik 2007, 2008),
while in Denmark strong indication for flax agricul-
ture and industry revealed first in Late Bonze Age,
ca. 2,800 cal BP, Frydenlund (Henriksen and Runge
2009).
OIL- AND FIBRE-PRODUCING CROPS 105
In Egypt, flax was retrieved (together with emmer
wheat and barley) in the earliest Neolithic farming
sites discovered in the Nile Valley during the eighth
millennium BP, Fayum (Caton-Thompson and
Gardner 1934; Wetterstrom 1993; Wendrich and
Cappers 2005) and Merimde (Wetterstrom 1993,
1998). Remains of flax seeds were also found in ca.
5,200 BP El Omari (Wetterstrom 1993), and they
continue to appear in several other pre-dynastic
sites. From the start of dynastic times onwards, flax
emerges as one of the founder principal grain crops
of Egypt. It kept this role until very recently.
Remains of flax textiles also make an early appear-
ance, attesting that fiber varieties developed not
much later than the oil-producing varieties. The
best examples come from the drier parts of south-
west Asia, where due to low humidity, woven mate-
rial survived without carbonization. Pieces of
exquisitely woven linen were discovered among
Pre-Pottery Neolithic B (beginning of the ninth mil-
lennium BP) remains in Nahal Hemar Cave near the
southern tip of Dead Sea, Israel (Schick 1988). A sin-
gle piece of linen was found in Neolithic (eighth
millennium BP) Fayum, Egypt (Caton-Thompson
and Gardner 1934). Linen fragments were retrieved
from the Chalcolithic, ca. 6,200–6,000 cal BP, ‘Cave
of the Treasure’ near the Dead Sea, Israel (Bar-Adon
106 DOMESTICATION OF PLANTS IN THE OLD WORLD
1980). The most remarkable linen find was discov-
ered in contemporaneous ‘Cave of the Warrior’ near
Jericho. It includes a beautifully preserved, colored,
and fringed kilt and sash, and large (seven meters
long and two meters wide) shroud, wrapping the
body of a dead nobleman. These finds indicate a
high level of familiarity and expertise in domesti-
cated linen production, already extant in this period
(Schick 1998, 2003). Flax textiles were common in
the Old, Middle, and New Kingdoms of Egypt,
where linen was extensively used for wrapping
mummies (Täckholm 1976; Germer 1985) Egyptian
retting, spinning, and weaving of flax is beautifully
recorded in the twelfth-dynasty Beni Hasan grave
paintings (Täckholm 1976).
In summary, the available archaeological evi-
dence clearly suggests that flax belongs to the first
group of grain crops that started agriculture in the
Levant during the Middle PPNB. As for distinguish-
ing the domesticated from the wild flax, the situa-
tion here is parallel to the cereals and pulses; it is
manifested in the function of the seed-dispersal
apparatus (diaspore). In wild populations, the
diaspores split and disperse the seeds immediately
after maturation. In contrast, breakage of the wild
type seed-dispersal mechanism does not occur in
the domesticated type and the capsule stays intact
and waits for the reaper. Hence, separated seed-
dispersal units in the archaeobotanical assemblage
indicate the wild type, while whole capsule, or
fused diaspores, indicate domesticated type. In
addition, the gradual increase in seed size and the
use of linen indicate that flax domestication was,
very probably, already practised in the PPNB
Levant, as attested first by Middle PPNB capsules
and then by Late PPNB seeds and linen textile. The
evidence from the living plants shows that wild
bienne forms are widespread in the ‘arc’, and fully
support the contention that domestication occurred
in the Fertile Crescent. Molecular studies also sug-
gest single domesticated events for flax, of the oil-
producing variety.
Hemp: Cannabis sativa
Hemp, Cannabis sativa L., is a tall (2–3.6 m) dioe-
cious, wind-pollinated herb, and a member of a
special small family Cannabaceae. Hemp is an
extraordinarily variable taxon that contains wild
forms, weedy types, and domestic cultivars. It is
grown for three main purposes: (i) for fibres
obtained from the bast of the stems; (ii) for the
seeds that are used either for extraction of oil or as
an animal feed; and (iii) as a source of a psychoac-
tive drug produced by the glandular hairs of the
plant. Special cultivar groups have been devel-
oped for the different uses (Small 1995). Fibre vari-
eties are tall, succeed in both temperate and
tropical climates, and contain negligible quantities
of the drug. Hemp textiles are strong but coarse
and in traditional communities they were used for
cheap clothing, sacks, rags, or sails. Hemp oil is
used mainly for technical purposes such as var-
nish or soap. The drug is obtained from special
varieties (frequently referred to as the indica group)
grown in warm climates. Only the tops of the
female plants contain appreciable quantities of the
Cannabis drug and are harvested for the prepara-
tion of marijuana (or hashish). Most taxonomists
consider C. sativa and C. indica to be one species,
but Hillig (2005), based on allozyme variation at 17
gene loci from 157 Cannabis accessions, suggest
that Cannabis has derived from two major gene
pools. On the basis of this data, he recognized
C. sativa and C. indica as separate species, both
with seeds and fibre landraces.
Domesticated hemp is closely related to, and fully
inter-fertile with, an extraordinarily variable aggre-
gate of wild and weedy forms. Except for traits asso-
ciated with the economic use, wild forms differ from
the cultivars by relatively smaller achenes (‘seed’),
which also possess adhering perianths and elongated
bases. The crop is mostly a central Asiatic element.
Temperate territories in this vast area such as the
Caspian Basin, parts of Afghanistan, central Asia,
and the Himalayas, harbour spontaneous C. sativa
plants that seem to be wild. Very likely, populations
that appear to grow wild are not fully indigenous,
but have been introgressed extensively with the cul-
tivated varieties and weedy types which also abound
in the same areas (Small and Cronquist 1976). The
picture of Cannabis is even further complicated by
the marked tendency of this cross-pollinated crop to
revert to wild. Naturalized derivatives and weedy
races of C. sativa are now distributed not only in cen-
tral Asia, but also all over the world.

Archaeological records on the early establish-
ment of hemp are not available so far. However, this
plant must have been taken into domestication
quite early, somewhere in temperate Asia. On the
basis of linguistic and cultural evidence Li (1974)
concluded that hemp was probably grown in China
by at least 4500 BP and that it was the only fibre
available to the ancient peoples of northern and
north-eastern China. Merlin (2003) summarizes the
available evidence, archaeological and otherwise,
of hemp, as well as other psychoactive plants in the
old world.
The recent expansion of hemp to south-west Asia,
the Mediterranean basin and Europe is now well
documented. Remains of hemp fabrics are available
from ca. eighth century BC Gordion, Anatolia
(Bellinger 1962). It also seems certain that the plant
was known to the Sarmatians and Scythians who
occupied the southern part of Russia between
approximately 700 and 300 BC (Godwin 1967).
Herodotus (ca. 446 BC), describes a Scythians puri-
fication ritual: ‘On a framework of three sticks meet-
ing at the top, they stretch pieces of woolen cloth,
taking care to get the joins as perfect as they can,
and inside this little tent they put a dish with red-
hot stones on it. Then they take some hemp seed,
creep into the tent, and throw the seeds on the hot
stones. At once it began to smoke, giving off vapor
unsurpassed by any vapor-bath one could find in
Greece. The Scythians enjoy it so much that they
would howl with pleasure’ (Herodotus, IV, 75, after
Merlin 2003). Such a practice is attested by a special
find in a series of fifth-century BC frozen kurgans
(burial mounds) at Pazyryk, Siberia. A 1.2 m-high
wooden frame tent was found within each burial,
and a bronze vessel inside it. Stones and hemp seeds
were found inside each vessel. ‘A leather pouch
with hemp seeds provided supplies, and scattered
hemp, coriander and melilot seeds were also recov-
ered’ (Rudenko 1970, as cited by Merlin 2003;
Sherratt 1995). Despite the appeal of such a find,
hemp seeds can be confused with those of Panicum
miliaceum (Bakels 2003), for example, re-identifica-
tion is therefore advised. At contemporaneous sites
in Ukraine, Pashkevich (1999) found Cannabis seeds
in several Scythian sites.
At that time hemp became a well-known fibre
crop in south-west Asia and Greece. It spread to
OIL- AND FIBRE-PRODUCING CROPS 107
Italy and Sicily in about 100 BC. The fibres were
used, among other things, to make ropes and sails.
An interesting find of both fruits and fibres came
from the Iron Age, ca. 2,400–2,050 cal BP, salt-min-
ing site and settlement Dürrnberg/Hallein, Austria
(Werneck 1949; Swidrak 1999; Boenke 2007). The
spread of C. sativa to central and north Europe from
Roman times on is now documented by a wealth of
macro-remains, palynological evidence, and writ-
ten sources (Dörfler 1990). The narcotic properties
of C. sativa were recognized in India by ca. 1000
BC.
Old World cottons: Gossypium
arboreum and G. herbaceum
Gossypium is a large genus of the family Malvaceae
containing some fifty species, distributed over large
landmasses in tropical and subtropical warm envi-
ronments. There are four different cotton crops, two
species in the Old World (G. arboreum L. and G. her-
baceum L.), and another two in the New World
(G. hirsutum L. and G. barbadense L.). An important
difference between the domestic species of the Old
World and the New World is that Old World cotton
crops are all diploid (2n = 2x = 26 chromosomes,
genomic designation AA), whereas the New World
crops are tetraploid (2n = 4x = 52 chromosomes,
genomic designation AADD). In this book we are
dealing with the Old World material only.
There is a revival of cotton as one of the most
important commercial plants for the present-day
textile industry. Both G. herbaceum and G. arboreum
bear spinnable but relatively short (less than 22 mm
long) seed fibres or lint (Lee 1984). These were the
only Gossypium crops grown in the warmer parts of
Asia and Africa before the discovery of America.
According to Fryxell (1984, p. 30) the widest crop
diversity of G. arboreum was in the Indian subconti-
nent, while the center of G. herbaceum’s diversity
was in East Africa and in the Levant. In post-Colum-
bus times, Old World cottons have been almost fully
replaced by their American counterparts (G. hirsu-
tum and G. barbadense), which produce longer (24-
34 mm long) lint. Today G. arboreum and G. herbaceum
survive as relic crops, and account for less than 1%
of the world cotton production. For a survey of both
domesticated and wild cottons consult Lee (1984),
108 DOMESTICATION OF PLANTS IN THE OLD WORLD
Fryxell (1979, 1984), Wendel (1995), and Wendel and
Cronn (2003). Butterworth et al. (2009) described
recently the morphological transformation under-
gone by cotton fibers in the course of domestication;
from short, coarse fibers in the progenitor wild spe-
cies, to the long, fine fibers in the domesticated
species.
Both G. arboreum and G. herbaceum are plants
growing in warm climates, and basically are peren-
nial crops (the first can reach the size of a small tree).
However, under domestication some annual culti-
vars evolved as well, in both crops, and rendered
cotton cultivation possible in areas with relatively
cold winters. The two species, G. arboreum and G.
herbaceum, are obviously closely related genetically.
They are similar morphologically, cross readily, and
their F1 hybrids are almost fully fertile. Both carry
homologous chromosomes (genome A) and an
identical chloroplast genome. When grown side by
side, they tend to hybridize spontaneously. In such
cases, their morphological boundaries are fre-
quently blurred. This suggests that both cultigens
belong to the same crop species complex and could
have evolved from a single wild ancestral stock. On
the other hand, domestic G. arboreum and G. her-
baceum were found to differ from one another by a
chromosomal translocation, and to manifest hybrid
breakdown at the second (F2) and the third (F3)
hybrid generations (Phillips 1961). Moreover, iso-
zyme tests showed that the two crops are character-
ized by numerous distinct alleles, indicating that
the genetic similarity between them is lower than
the values known to occur in most crop-ancestor
species pairs (Wendel et al. 1989, 2003). As argued
by these authors, such divergence does not support
the notion of two cultigens derived from a common
wild ancestor. Instead, it suggests that each evolved
from a distinct wild stock. Moreover, they indicate
that cotton is unique among crop plants in that four
separate species (the two Old World species and the
two New World species) were independently
domesticated.
The present knowledge of the close wild relatives
of the Old World cultivated cottons (wild diploid
taxa containing genome A) is still insufficient, and
of little help in solving the problems of how and
where G. arboreum and G. herbaceum were domesti-
cated. Wild forms of G. herbaceum have already been
discovered and identified both taxonomically and
cytogenetically. They are referred to today as G. her-
baceum L. subsp. africanum (Watt) Hutch. These wild
taxa are known to grow only in southern Africa
(Vollesen 1987), a region where agriculture arrived
at a very late date, and geographically a place
widely separated from the traditional territory of G.
herbaceum cultivation. The situation in G. arboreum
is even more perplexing. Gossypium taxonomists
(Fryxell 1979, 1984; Vollensen 1987) do not mention
any wild arboreum forms in their studies. It is not
clear whether this is a real absence or only a reflec-
tion of the paucity of field findings. All in all, wild
diploid cottons—containing genome A—have not
yet been conclusively detected in the geographic
areas in which they should be expected—Africa
north of the equator, south Arabia and/or the Indian
subcontinent.
Archaeological evidence
The archaeobotanical documentation of cotton is far
from being satisfactory. The earliest evidence of cot-
ton fibers came from ca. 8,000–6,500 BP Ceramic
Neolithic Mehrgarh, Baluchistan (Moulherat et al.
2002). The fibers were identified as Gossypium sp.,
most probably represent wild origin. Later, reliable
signs of cotton use, as attested by fibers and seeds
finds, come from Harappan sites (ca. 4,250–3,750
BP) in the Indian subcontinent. Here fragments of
cotton textiles and strings, preserved by copper and
silver oxides, were uncovered in Mohenjo-Daro,
Pakistan (Gullati and Turner 1929). In addition, cot-
ton remains are reported from contemporary
Harappa and several other Indian and Pakistani
sites (Fuller 2000, 2008). As stressed by Meadow
(1996), these finds indicate that Gossypium fibers
were exploited in the Indus valley as early as the
end of the fifth millennium BP. However, it is
unclear whether these early remains represent
arboreum or herbaceum cottons.
A more problematic early find comes from ca.
4,500 BP Afyea, Egyptian Nubia. Cotton seed and
lint hairs (intermediate between those borne by
wild forms and those produced by herbaceum culti-
vars) were discovered in goat coprolites (Chowdhury
and Buth 1970, 1971). It is hard to imagine that the
Afyea find represents domestic cotton, since it is not
accompanied by any other contemporary signs of
cotton cultivation or cotton use in the Nile Valley,

which is known so well archaeo-botanically. As
Moulherat et al. (2002) and Fuller (2008) noted, if the
identification of sixth Millennium BP cotton fibers
from Dhuweila, Jordan (Betts et al. 1994) will be con-
firmed, it was most likely imported, perhaps from
the Indian subcontinent.
Cotton seems to have moved into south-west
Asia and the Mediterranean basin fairly late. The
earliest sign comes from Assyria where we are
informed that ‘trees bearing wool’ were grown by
Sennacherib at about 694 BC (Thompson 1949).
There are no indications that this early introduction
succeeded, but the mentioning of a tree suggests
that G. arboreum was involved. Cotton is already
referred to in Greek, Roman, and Jewish literary
sources, from Hellenistic times onwards. Watson
(1983, pp. 32, 34) surveyed the literary information
and concluded that already in Greek and Roman
times, Indian cotton goods were imported into
south-west Asia and the Mediterranean basin, and
that they were rare and expensive commodities.
The diffusion of cotton cultivation (and of its
processing technologies) is another matter. There is
no sound literary support for cotton cultivation in
south-west Asia at that time. The Jewish sources
Mishna and Talmud (particularly the late ones dat-
ing to the AD fifth and sixth centuries) refer to sev-
eral traits of the cotton plant itself, indicating that
cotton might already have been grown in the Levant
countries (Amar 2000). By compiling current data,
Wild (1997) and Decker (2009) demonstrated that
cotton cultivation was widespread in Roman Egypt.
To-date, the earliest unequivocal archaeobotanical
evidence on cotton cultivation outside the Indian
subcontinent, comes from late Sassanian (AD sixth
and early seventh centuries) Merv in Turkmenistan
(Nesbitt 1993, 1994). Fully developed cotton culti-
vation and sophisticated cotton industry appear in
the Levant and the Mediterranean basin only in
early Islamic times (Watson 1983; Amar 2000).
Poppy: Papaver somniferum
Poppy, Papaver somniferum L. (Papaveraceae), is
grown for two purposes (Duke 1973). First, it is
famous as a source of opium, which is obtained
from the latex released by the plants after gashing
their unripe capsules. It is also cultivated for its
tasty seeds, which are rich in oil. The seeds are con-
OIL- AND FIBRE-PRODUCING CROPS 109
sumed as such or used to extract poppy oil. Because
of this dual use, two series of cultivars have evolved
in P. somniferum. The opium forms are grouped in
subsp. somniferum Corb., while the oil varieties are
collectively known as subsp. hortensis (Hussenot)
Corb. The oil is used both for eating and for indus-
trial purposes. The opium is a powerful medicinal
and narcotic element. Its effects, including pain kill-
ing, were already appreciated in antiquity (Merlin
2003).
Domesticated poppy, Papaver somniferum, is pre-
dominantly self-pollinated. Most cultivars are dip-
loid (2n = 2x = 22 chromosomes), while wild
setigerum DC poppies comprise both diploid and
tetraploid (2n = 4x = 44 chromosomes) types.
The crop is closely related to wild and weedy
forms of poppies that grow, mainly in coastal areas,
in the western part of the Mediterranean basin,
including most of the islands (Map 14).
The diploid setigerum forms were found to be fully
inter-fertile with the somniferum cultivars (Hammer
and Fritsch 1977). They are therefore assumed to be
the progenitor stock from which the cultivated
poppy evolved. This necessitated a taxonomic recon-
sidering of the rank of P. setigerum, which is placed
as the wild race, subsp. setigerum (DC.) Corb., of the
P. somniferum crop complex (Kadereit 1986).
The most conspicuous differences between the
tame and the wild plants are the considerable
increase in capsule size, and the retention of the
seed in the capsule in the poppies under domestica-
tion—in domesticated forms the pores beneath the
stigmatic lobes do not open. Domestic forms also
show a tendency to increased seed size. However,
some wild populations bear fairly large seeds which
considerably overlap the dimensions (and seed-coat
reticulate sculpture, see Fig. 33) found in the culti-
vated poppy. Seed size is therefore an unreliable
trait for recognition of domestication in this crop
(Fritsch 1979); which necessitates capsule remains
for properly identified domestic form in archaeo-
logical assemblages.
Archaeological evidence
The earliest wild poppy seed came from ca. 8,000–
7,500 cal BP PPNC Atlit Yam, Carmel Coast, Israel
(Kislev et al. 2004). This find might indicate that its
distribution area (see Map 14) included Israel dur-
110 DOMESTICATION OF PLANTS IN THE OLD WORLD

0 100 200 miles

0 200 400 km

Map 14 Geographical distribution of wild poppy, Papaver somniferum subsp. setigerum [= P. setigerum] (based on La Valva et al. 1985; Kadereit
1986). Setigerum forms extend westwards beyond the boundaries of this map into Madeira, the Canary, and the Azores Islands.

ing the early Neolithic period. Abundant water-
logged seeds of domesticated poppy came from ca.
Fig. 33 The reticulate sculpture of the seed coat in poppy, Papaver
somniferum (Schoch et al. 1988).
7,750–7,150 cal BP Early Neolithic La Marmotta,
Italy (Rottoli 1993, 2002). A few carbonized domes-
ticated poppy seeds were found in the
Linearbandkeramik settlements of the Aldenhovener
Platte, north-western Germany (Knörzer 1971,
1997), Meindling, south Germany (Bakels 1996),
and a few more in Zeslawice, Poland (Giżbert 1960),
and north of the Alps (Kreuz 2007).
Numerous remains of charred seed, as well as
occasional capsules, are available from middle and
late Neolithic central Europe (for review see
Schultze-Motel 1979a; Merlin 2003). They were also
retrieved from middle Neolithic Menneville, north
France (Bakels 1984), and became common in Late
Neolithic (fifth millennium BP) and in Bronze Age
lake-shore settlements in Switzerland (Jacomet et al.
1989; Brombacher 1997), in south Germany (Maier
1995; Rösch 1998), and in Lagozza, north Italy
(Buschan 1895). Four desiccated, beautifully pre-
served, wild poppy capsules were reported by
Neuweiler (1935) from Cueva de los Murciélagos

near Albuñol, Granada, south Spain (despite the
same name, this is a different cave from that of
Córdoba mentioned above).
Significantly, poppies were not discovered in
Neolithic sites in southeastern Europe and in south-
west Asia. They appear, however, in several Bronze
Age sites in Greece, Bulgaria, and the former
Yugoslavia (Kroll 1991). As stressed by van Zeist
(1980) and by Bakels (1982) this fact, combined with
the distribution area of wild setigerum poppies (Map
14), suggests a west Mediterranean domestication.
In other words, P. somniferum does not belong to the
primary ‘first circle’ Fertile Crescent crops that
started food production in Europe. It is a represent-
ative of the ‘second circle’ domesticants, (i.e. crops
that were added to the original assemblage outside
the Fertile Crescent core area).
Gold of pleasure: Camelina sativa
Gold of pleasure or false flax, Camelina sativa (L.)
Crantz., of the mustard family Cruciferae (Brassi-
caceae) is a relic oil plant rapidly disappearing from
cultivation. Yet until the 1940s, Camelina was an
important oil crop in eastern and central Europe.
Camelina seed, with their characteristic protruding
embryos, appear repeatedly in European archaeo-
logical contexts.
The crop, subsp. sativa (Mill.) E. Schmid, is closely
related to, and inter-fertile with, a variable aggre-
gate of wild and weedy forms distributed over
Europe and south-west Asia (for review, see
Markgraf 1975). Truly wild, late-flowering forms,
now recognized as subsp. microcarpa (Andrz.) E.
Schmid, but formerly referred to as C. microcarpa
Andrz., grow in east European steppes and adjacent
Asiatic territories. They have characteristic small
fruits. Closely related to them are early flowering
hairy forms, known as subsp. pilosa (DC.) E. Schmid,
which thrive in fields of winter cereals over most of
Europe. These are obviously recently evolved
weeds, which spread from east Europe westward,
over cultivated lands. An additional distinct
Camelina weedy race, namely subsp. alyssum (Mill.)
E. Schmid [= C. sativa var. linicola Prusch.] evolved
in association with flax cultivation in Europe. These
linicola forms, with their long erect stems and hard
fruits, serve as well-documented examples of evo-
OIL- AND FIBRE-PRODUCING CROPS 111
lution of weed mimicry to a specific cultivated crop.
The cultivated varieties of C. sativa differ from the
wild and weedy forms by their non-dehiscent, large,
and pyriform fruits, and by bigger (1.5–2.0 mm
long) seed, which contain an appreciable amount
(27–31%) of edible oil.
Archaeological evidence
Camelina seeds (as well as occasional pods) appear
repeatedly in central and eastern European archae-
ological contexts. However, they start late and the
earliest finds come from fifth and fourth millennia
BP contexts. Large samples, as well as pure sam-
ples; i.e. material that can be confidently regarded
as confident domestication appear even later.
In central Europe the oldest remains come from
final Neolithic contexts (ca 4,000 BC) of Auvernier,
Switzerland, and these are followed by several
Bronze Age (1,800–1,200 BC) finds from Poland,
Hungary, Germany, and north Italy (Schultze-Motel
1979b; Wasylikowa et al. 2001). Iron Age finds are
more numerous. Camelina remains become particu-
larly common in the coastal areas of the Baltic and
the North Sea. At least some of these finds (particu-
larly the large samples with bigger seeds) seem to
represent Camelina cultivars.
Further south-east, a single seed of Camelina was
found in Chalcolithic (ca. 5,000 BP) Pefkakia,
Thessaly (Kroll 1991), and many more in ca. 4,200
BP Sucidava-Celei, Rumania (Cârciumaru 1996).
Camelina seeds were also retrieved as a weed of flax
from Late Chalcolithic Kuruçay in south-west
Anatolia (Nesbitt 1996), from Early Bronze Age
Demircihüyük in north-west Anatolia (Schlichtherle
1977/1978), and in Iron Age Yoncatepe in Van Lake
province (Dönmez and Belli 2007). They reappear in
several Late Bronze Age sites in Greece, Bulgaria,
and the former Yugoslavia (Kroll 1991), and also in
Late Bronze Age Hadidi on the Syrian-Turkish bor-
der (Miller 1991). In the latter site their concentra-
tion again suggests cultivation.
The combined evidence from the living plants
and from archaeology indicates that C. sativa is a
secondary crop. Probably this crucifer entered agri-
culture first by evolving weedy races that infested
flax and cereal cultivation. Only later was the weed
picked up as an oil crop.
112 DOMESTICATION OF PLANTS IN THE OLD WORLD
Other cruciferous oil crops
The cultivation of several other members of the
mustard family (Cruciferae/Brassicaceae) is amply
recorded in classical times. They have been appreci-
ated as oil or mustard sources (extracted from their
seed), and/or for their vegetable parts (see Chapter
7). Foremost among these crucifers are: radish
Raphanus sativus L., turnip Brassica rapa L. [= B.
campestris L.], rape or swede B. napus L. (tetraploid,
2n = 4x = 38 chromosomes), and the various mus-
tards; i.e. the white mustard Sinapis alba L., (diploid,
2n = 2x = 16 chromosomes), black mustard B. nigra
(L.) Koch, (tetraploid, 2n = 4x = 36 chromosomes),
and brown mustard B. juncea (L.) Czern. (diploid,
2n = 2x = 2 chromosomes). All these crops have wild
forms distributed over west Asia and Europe. They
include variable, aggressive races of weeds, which
infest agricultural land far beyond these territories.
It is very likely that seed of wild forms (Brassica,
Sinapis, and also Camelina and Descurainia) has been
collected or even grown as oil sources in Switzerland
and Germany in the late Neolithic times
(Schlichtherle 1981).
These crucifers were already well-established oil
crops in Hellenistic and Roman times. Therefore, it
is safe to assume that they were taken into cultiva-
tion earlier. Yet, there are almost no archaeological
records available for any of these crops. (For the
scant information present, see Renfrew 1973, pp.
166–167, and the annual reviews of Schultze-Motel
1968–1994, and Kroll 1995–2000, as well as the
online databases http://www.cuminum.de/
archaeobotany/database/; http://www.archaeo-
botany.de/database.html). Suggestions as to the
origin of these plants are necessarily based on lin-
guistic considerations.
One rich find of cruciferous seed is described
from ca. 5,000 BP Khafadje, Iraq. Here, numerous
carbonized seeds were discovered in several loca-
tions at the Temple Oval (Bedigian and Harlan
1986). They were studied by E. Schiemann (1933)
who concluded that they belonged to Brassica or
Sinapis. It is very difficult to distinguish between
the seeds of these two genera in charred material.
Two cakes with crushed seed of Brassica/Sinapis
were found by Willcox (2002) in tenth millenium BP
PPNA Jerf el Ahmar, Syria. The presence of Brassica
rapa remains in several late Neolithic and Bronze
Age lake-shore sites in Switzerland (e.g. Seeberg,
Villaret-von Rochow 1967), suggests a similar use.
In other words, turnip may already have been an
oil-bearing ‘tolerated weed’ at this relatively early
time.
Sesame: Sesamum indicum
Sesame, Sesamum indicum L. [= S. orientale L.]
(Pedaliaceae), is a traditional, warm-season, annual
oil crop whose genus contains some two wild spe-
cies. Cultivars of this crop have been grown in trop-
ical and subtropical environments, such as
south-west Asia and the Mediterranean basin.
Sesamum is highly appreciated for its oil that keeps
fresh for a long time without turning rancid (Nayar
1995). It was extensively cultivated in the Graeco-
Roman World (Lenz 1859) both for its attractive oil
and for its edible seed. Theophrastus categorizes
sesame, along with the millets, as one of the main
summer crops of his time. Yet despite such wide use
in Roman times, sesame is not an indigenous crop
in the Mediterranean and south-west Asian
agriculture.
The genus Sesamum includes tropical and sub-
tropical wild species, grouped in four sections
(Ihlenfeldt and Grabow-Seidensticker 1979). The
majority of these species occur in Africa. A few
occur in the Indian peninsula and in south-east
Asia. All these taxa (except for the crop varieties) do
not occur in south-west Asia or in the Mediterranean
basin. In fact, this genus is geographically restricted
to tropical and subtropical Africa south of the
Sahara (the majority of the wild species), and only a
few species in the Indian subcontinent and the Far
East.
A group of wild-growing and weedy forms occur
in northwest India, as reported by Bedigian and
Harlan (1986) and Bedigian (1998, 2004, 2010) as S.
orientale L. var. malabaricum Nar. A second similar
wild group was discovered on the southern and
western shores of the Indian peninsula, and named
S. mulayanum Nair (Hiremath and Patil 1999). Both
groups show close morphological and cytogenetic
affinities with the crop. Very likely they are ele-
ments of the wild stock from which cultivated ses-
ame could have been derived.
 OIL- AND FIBRE-PRODUCING CROPS 113

Archaeological evidence They are followed by four large finds of Iron Age
jars containing carbonized sesame seed that were
Sesame seeds are very fragile when charred (even
excavated in Karmir-blur on the outskirts of
more so than other oil-rich seeds). In part, this
Yerevan, Armenia, dated to 900 and 600 BC. The site
explains their scarcity in the archaeobotanical
also contains elaborate installations for the extrac-
record. The oldest record of S. indicum cultivation
tion of oil from the seed (Bedigian 1985). Another
comes from ca 4,250–3,750 BP Harappa in the Indus
find of the same period comes from the Urartu
Valley (for review see Bedigian 1985, 1998, 2010;
Kingdom of Bastam, north-west Iran (Hopf and
Fuller 2003). This was augmented by the find of a
Willerding 1989). In addition, some 200 sesame
small quantity of well-preserved, charred sesame
seeds were uncovered in Iron Age, ca. 800 BC, Deir
seed in Miri Qalat and Shahi Tump, Markan region,
Alla, Jordan (Neef 1989).
Pakistan, in contexts dated to the second half of the
It seems that S. indicum arrived to south-west
fifth millennium BP (Tengberg 1999).
Asia and the Mediterranean basin from the east in
The earliest definite signs of sesame seeds in south-
the Early Bronze Age (Bennett and Maxted 1997).
west Asia come from ca. middle of the fifth millen-
The relative small number of finds probably indi-
nium BP, Early Bronze Age Abu Salabikh,
cates small-scale cultivation for culinary purposes
Mesopotamia (Charles 1993). Somewhat later, it
rather than large-scale oil production (Neef 1989). It
appears in ca. 3,200 BP, Late Bronze Age, Tell Sabi
was apparently taken into cultivation in the Indian
Abyad, northern Syria (van Zeist 1994). At adjacent
subcontinent. It is still undecided when and where
Tell Schech Hamad, van Zeist (2001, 2003) reported
sesame was domesticated. However, Fuller (2003)
sesame seeds from two contexts, a single seed from
and Bedigian (2004, 2010) felt that the combined
ca thirty-third century BP and a few more seeds from
evidence obtained both from the living plants and
ca. twenty-seventh century BP. Also in the same
from archaeological sites strongly supports the
period, baskets of sesame seeds were found in
notion that domestication of sesame started in the
Tutankhamun’s tomb in Egypt, ca. 1,325 BC (de
Indian sub-continent and subsequently migrated
Vartavan et al. 2010) The unique circumstances of
westward. It appears that only during the Greek
King Tut perfectly-preserved un-charred funerary
period did sesame became a major oil-producing
offerings most probably represent a rare import, fit
crop in south-west Asia.
for the king’s table—certainly not a local cultivation.
 C H A PTER 6
Fruit trees and nuts
Fruit-bearing trees constitute an important element
of food production in the countries bordering the
Mediterranean Sea. Their long-standing economic
importance is amply reflected in classical traditions
(Stager 1985). Five of the Biblical ‘seven species’ are
founder crops of Mediterranean agriculture. Olive
oil, wine, raisins, dates, and common figs were (and
still are) staple agricultural products in south-west
Asia and the Mediterranean basin.
Like the Neolithic grain crops, but significantly
later, the first indigenous fruit trees were brought
into domestication in the Fertile Crescent area
(Zohary and Spiegel-Roy 1975). In comparison to
grain agriculture, horticulture evolved much later in
the history of food production in Europe and south-
west Asia. The first definite signs of fruit tree domes-
ticants appear in south-west Asia in Chalcolithic
contexts (seventh millennium BP), that is, several
millennia after the firm establishment of grain agri-
culture in this region. In Early Bronze Age, olives,
grapes, and figs emerge as important additions to
cereals and pulses throughout the eastern
Mediterranean basin, including the Aegean belt,
while date palms have been grown at that time in the
warmer and drier southern niches of this region.
Horticulture is very different from grain crop
agriculture. Cereals and pulses are ‘short invest-
ment’ annual crops. Their mature grains are ready
to be harvested several months after sowing. The
grain grower can move from place to place after the
harvest, and practice shifting farming. In contrast,
fruit trees are perennials. Orchards start to bear fruit
three to eight years after planting, and reach full
productivity several years later. Horticulture, there-
fore, needs protection from intruders the year
round, indicating a settled way of life.
114
Genetically, domestication of fruit trees means
changing the reproductive biology of the plants
involved (Zohary and Spiegel-Roy 1975; Zohary
1984) by shifting from sexual reproduction (in the
wild) to vegetative propagation (under cultivation).
As a rule, domesticated varieties of fruit trees are
maintained vegetatively by the farmer (as clones)
by rooting of twigs, use of suckers, or by the more
sophisticated technique of scion grafting. They are
seldom raised from seed. This is in sharp contrast
with their wild relatives, which reproduce mainly
from seed. In other words, wild fruit tree pop-
ulations maintain themselves through sexual
reproduction and are distinctly allogamous. Cross-
pollination is brought about either by self-incom-
patibility or by dioecy (separate male and female
individuals). Spontaneous populations manifest
wide variability and maintain high levels of hetero-
zygosity. Consequently, their seedlings segregate
widely in numerous traits, including the size, shape,
and palatability of the fruits.
In the hands of the farmer, vegetative propaga-
tion has been a powerful device to prevent genetic
segregation and to ‘fix’ desired types. By discarding
sexual reproduction and inventing clonal propaga-
tion, the farmer can: (i) select and maintain excep-
tional individuals with desirable fruit traits from
among large numbers of variable inferior trees, and
(ii) duplicate (clone) the chosen types to obtain
genetically identical saplings. In the case of fruit
trees, this is no small achievement. Since fruit trees
are generally cross-pollinated and widely hetero-
zygous, most progeny obtained from seed (even
progeny derived from the superior cultivars) do not
bear the desired traits and therefore are econom-
ically worthless. The shift from seed planting to

vegetative propagation has been the practical solu-
tion to assure a dependable supply of desired geno-
types. In most fruit trees, this development made
domestication possible. Only several nut trees (such
as almond, walnut, and carob) were traditionally
maintained by seed planting.
Plant remains retrieved from archaeological exca-
vations indicate that the olive, grapevine, fig, date
palm, pomegranate, and the sycamore fig were the
first fruit trees introduced into domestication in
south-west Asia and Europe. Significantly, all these
‘first-wave’ fruits lend themselves to simple vegeta-
tive propagation by cutting and rooting of twigs (in
the grapevine, fig, and sycamore fig), digging out of
suckers (in pomegranate), planting basal knobs (in
olive) or by transplanting offshoots (in date palm).
In all these founder fruit crops the early growers
did not have to resort to more sophisticated tech-
niques of vegetative propagation (such as grafting).
Wild olives, grapes, figs, dates, and pomegranates
were thus ‘pre-adapted’ for early domestication.
This adaptation was probably vital for their success
in becoming the first domesticated fruit trees in
south-west Asia and the Mediterranean basin.
Several other fruit trees, such as apple, European
pear, plum, sweet cherry, carob, and pistachio were
introduced into cultivation much later. Definite evi-
dence for their domestication appears only in the
third millennium BP and their extensive incorpora-
tion into horticulture seems to have taken place
only in Greek and Roman times. A plausible expla-
nation for the late appearance of these ‘second-
wave’ domesticated fruits is that they do not lend
themselves to simple vegetative propagation. Their
maintenance is based almost entirely on grafting.
The Greeks and the Romans were already familiar
with this art, and we have ample documentation
that in classical times, apples and pears were main-
tained by this sophisticated method of vegetative
reproduction (for review, see White 1970, pp. 248,
257–8). It is not clear when and where detached
scion-grafting was invented. The earliest reliable
description of grafting in the Mediterranean basin
was given by Theophrastus (II, v. 3) who lived in
Greece in the fourth century BC. Obviously, the
introduction of this sophisticated method of propa-
gation enabled the domestication of a new range of
fruit trees. Grafting was probably invented outside
FRUIT TREES AND NUTS 115
the region of Mediterranean horticulture, and intro-
duced into this region from the east. The earliest
information on the use of grafting comes from
China in connection with citrus-fruit domestication,
and one of these texts is dated to ca. 139 BC (Cooper
and Chapot 1977; Métailié (2007).
The adoption of clonal cultivation means that
most fruit trees have undergone very few sexual
cycles in the five or six millennia since their intro-
duction into cultivation. Domesticated clones per-
sisted for hundreds of years. From the standpoint of
evolution under domestication, this means a severe
restriction on selection. In other words, selection
(both conscious and unconscious) could have oper-
ated only during a limited number of generations,
and we have to expect that the cultivars have not
diverged considerably from their progenitors’ gene
pools. This is in sharp contrast to the annual self-
pollinated grain crops, where selection could have
operated effectively during thousands of genera-
tions. Indeed, the domesticated varieties of fruit
trees can be regarded as exceptional, highly hetero-
zygous individuals of their biological species—
clones that excel primarily in fruit size and fruit
quality. The absence of profound genetic changes in
the fruit trees under domestication is also apparent
in their ecology. The climatic requirements of the
cultivars closely resemble those of their wild rela-
tives. Unlike the cereals and pulses, the fruit crops
have not been pushed much beyond the climatic
requirements of their wild ancestors.
As already stated, most fruit trees under domesti-
cation are derived from wild progenitors in which
cross-pollination is maintained by one of the fol-
lowing genetic systems: self-incompatibility, dioecy,
or dichogamy. Because of this background, the shift
from sexual reproduction to planting of vegeta-
tively propagated clones introduced serious limita-
tions on fruiting. Planting of a single self-incompatible
clone, or alternatively female clones, would not
bring about fruit set. Several agronomic devices,
assuring fruit set in the orchard, have been empiri-
cally adopted. They were accompanied by uncon-
scious selection for several types of mutations that
resolved the restrictions set by self-incompatibility
and sex determination.
In hermaphroditic, self-incompatible species such
as olive, apple, or pear, the early planters very likely
116 DOMESTICATION OF PLANTS IN THE OLD WORLD
realized that to obtain satisfactory fruit set, it was
necessary to plant together two or more synchro-
nously-flowering clones (Table 8). The traditional
cultivation of such fruit trees is based on mixed
planting, a practice which brings about pollination
between different genotypes. Two additional solu-
tions are the outcome of unconscious selection: (i) in
the peach, apricot, sour cherry, European plum, as
well as in several cultivars of almonds or olives, we
find mutations that caused the breakdown of self-
incompatibility, (or at least rendered the system
‘leaky’), so that self-pollination also results in con-
siderable fruit set in such clones; and (ii) in other
self-incompatible taxa, pollination has been dis-
pensed with altogether by incorporation of muta-
tions conferring parthenocarpy (fruit development
without fertilization and without seed set). Several
clones of cultivated pear show this adaptation.
In dioecious species, fruit set is safeguarded in
parallel ways. In the pistachio, ‘mixed planting’ is
employed, and some male individuals are planted
together with the female clones. In date palms and
Smyrna-type figs, natural pollination is frequently
augmented by artificial pollination. Also in dioe-
cious fruit trees, the early planters must have uncon-
sciously selected two types of mutations that
rendered cross-pollination unnecessary: (i) a genetic
Table 8 Horticultural safeguarding fruit set in fruit trees (compiled
after Zohary 1984)
The Genetic System The Agricultural Solution
Self-incompatible fruit 1. Planting together a few synchronously
trees flowering clones.
2. Mutations toward breakdown (some
almonds, 4x cherries) or reduction
(e.g. some olives, plume, apples, and
pears) of self-incompatibility.
3. Parthenocarpic-induced mutations
(e.g. some pears).
Dioecious fruit trees 1. Planting additional male plants
(e.g. pistachio).
2. Artificial pollination (e.g. date palm,
Smyrna-type figs).
3. Unconscious selection toward:
(i) hermaphrodite plants (e.g. grape)
(ii) parthenocarpy (e.g. common fig,
Corinth-type grapes).
change from dioecy (in the wild) to hermaphro-
ditism (under domestication) evolved in the grape-
vine; and (ii) a replacement of sexual reproduction
by parthenocarpy occurred in many clones of the
fig and the sycamore fig.
In most fruit trees, population variation in seed
morphology is quite large. This phenomenon makes
definite differentiation between wild and domesti-
cated forms in early archaeological contexts prob-
lematic. Clear-cut signs, equal to the smooth/rough
disarticulation scar in the cereals (p. 22) are currently
unknown. Like cereals, such differences, especially
in seed-size, can be seen in later phases. As a result,
we cannot be certain whether the earliest finds are
wild or domesticated. Recent adoption of morpho-
metric tools (such as Terral et al. 2010) might lead to
better identification in the future.
Olive: Olea europaea
The olive, Olea europaea L. (Oleaceae), is the most
prominent, and economically the most important
classical fruit of the Mediterranean basin (Zohary
1995a). It comprises the oldest group of plants that
founded horticulture in the Old World, together
with grapevine, fig, and date palm (Zohary and
Spiegel-Roy 1975; Boardman 1976; Stager 1985).
Since Bronze Age, the wealth of many Mediterranean
peoples centred on the cultivation of olives, which
provided valuable storable oil as well as edible
fruits. Olive oil has been used in eating and cook-
ing, as well as for ointment and lighting. Because of
its excellent durability, it served as a principal arti-
cle of commerce. The whole fruit was also preserved
and consumed. Bread and olives were, and still are,
a staple diet in peasant communities throughout
the Mediterranean basin.
Olives grow in typical Mediterranean climates.
This fruit crop, and its closely related wild oleaster
forms, are considered reliable indicators of
Mediterranean environments. The olive is a rela-
tively slow-growing tree, diploid (2n = 4x = 46
chromosomes), and fruit production starts five to
six years after planting. If well managed, the long-
living olive trees can keep fruiting for hundreds of
years. Olives, under domestication, manifest con-
siderable variation in the size, shape, and oil con-
tent of their fruits. Hundreds of distinct varieties

are recognized. Different parts of the Mediterranean
basin are frequently characterized by specific local
forms. The olive crops can be roughly subdivided
into two main types: (a) oil varieties, the ripe fruits
of which contain at least 2% oil (the oil is mono-
unsaturated with a high percentage of Oleic acid
and used for cooking, salt dressing, and food pres-
ervation); and (b) table olives: less oily forms used
for the preservation of whole fruits, by pickling or
salting. There are also dual-purpose cultivars (see
Lavee and Zohary 2011). Carbonized kernels
(stones) and charred wood of pruned twigs, used as
firewood, constitute the bulk of olive remains in
archaeological excavations.
Wild ancestry
The domesticated olive, O. europaea L., shows close
affinities to a group of wild and feral olives distrib-
uted over the Mediterranean basin and tradition-
ally referred to as ‘oleaster’ olives (Plate 13). These
wild forms are fully inter-fertile with the domesti-
cated varieties, and interconnected with them by
sporadic, spontaneous hybridization (Zohary and
Spiegel-Roy 1975; Zohary 1995b; Lavee and Zohary
2011). Oleaster olives and the domesticated clones
have similar climatic and soil requirements.
Previously, many botanists regarded the wild
oleaster forms as representing an independent spe-
cies, O. oleaster Hoffm. & Link. More recently,
because of its close morphological and genetic affin-
ities to the domesticated fruit tree, most researchers
dealing with Mediterranean plants regard the
oleaster forms as the wild progenitor of the crop.
They place it within O. europaea L. species complex,
either as a subspecies [subsp. oleaster (Hoffm. &
Link) Hegi] or as a variety [var. sylvestris (Mill.)
Lehr. = var. oleaster (Hoffm. & Link) DC].
Oleaster olives differ from the domesticated clones
mainly by their smaller fruits and usually by spines-
cent juvenile branches. These wild fruits have less
fleshy mesocarp and contain less oil. However, fre-
quently the stones are not much smaller than those
in the cultivars. Wild olives are almost fully self-
incompatible, and as with many other trees main-
taining such a genetic system, oleaster populations
often show a wide range of electrophoretically dis-
cernible allozyme variation. In comparison, the
FRUIT TREES AND NUTS 117
domesticated varieties (of the domesticated gene
pool) are much less variable (Quazzini et al. 1993;
Zech-Matterne and Leconte 2010). Under cultiva-
tion, the reproductive biology of the tree changed.
While wild olives reproduce by sexual reproduc-
tion from seed, domestic varieties are maintained
by vegetative propagation and are, in fact, clones.
Traditional vegetative propagation depends prima-
rily on the utilization of knobs (= uovuli) and trun-
cheons that develop at the base of the trunk, and by
grafting. Today, vegetative propagation in horticul-
ture relies mainly on tissue culture.
In Olea (as with numerous other fruit trees) the
shift to vegetative propagation is the cultivator’s
countermeasure to the problems of wide genetic
segregation that characterize reproduction in cross-
pollinated plants. Domesticated olive clones are
considerably heterozygous. When their sexually
produced seeds are planted, the ensuing progeny
segregate widely. In fact, most sexual seedlings
resemble the wild forms in their morphology and
are poor or useless in terms of fruit quality.
Consequently, propagation from seed is impractical
in oleoculture. In order to ‘fix’ useful genotypes, the
grower has to resort to clonal propagation. Seedlings
can only be used as a variable raw material for
selection of new clones. At present, such selection of
seedlings is performed in some olive-breeding pro-
grams. Spontaneous seedlings have accompanied
olive growing from its very start, and rare individu-
als showing superior qualities may have caught the
attention of early cultivators and were then picked
up as the new clones.
Over large areas in the Mediterranean basin, wild
oleaster-type, olives thrive as common constituents
of maquis and garrigue formations (Map 15). In addi-
tion, these shrubs and small trees frequently colo-
nize secondary habitats, such as the edges of
cultivation or abandoned orchards. In such places,
particularly in areas of olive plantations, spontane-
ous olives seem frequently to be ‘feral’. Such ‘feral’
plants can derive from hybridization (i) between
different domestic clones, or (ii) between domesti-
cated clones and adjacent wild oleaster plants.
The Mediterranean wild olives are associated
with olive cultivation also in another way. They
often served as stock material for grafting. Oleaster
suckers or knobs are being dug out (in the wild),
118 DOMESTICATION OF PLANTS IN THE OLD WORLD
Fig. 34 Charred stones of olives, Olea europaea, Chalcolithic Tuleilat Ghassul, Jordan (Zohary and Spiegel-Roy 1975).
and planted in orchards as hardy stock material for
grafting of domestic scions. In some localities, for
example in western Turkey, one still encounters
another old tradition, namely the grafting of wild
olives growing in non-arable maquis and/or gar-
rigue vegetation. The grafted wild olives in such
niches are protected by the farmers, while other
trees and shrubs are cut for firewood and further
suppressed by intensive grazing.
Olea europaea is the only Mediterranean repre-
sentative of the genus Olea L., which includes some
two to twenty-five species distributed over tropical
and southern Africa (the main center), and southern
Asia and China (as well as eastern Australia, New
Caledonia, and New Zealand). Both the olive crop
and the Mediterranean oleaster forms are genetically
closely related to (and probably inter-fertile with)
several non-Mediterranean wild olive taxa. Most
widespread among the latter are various east
African, south Arabian, south Iranian, and Afghan
wild subspecies of the O. europea taxonomic com-
plex (subsp. sylvestris, subsp. cerasiformis, subsp.

0 100 200 miles
0 200 400 km
Map 15 Geographical distribution of wild olive, Olea europaea subsp. oleaster (based on Zohary and Spiegel-Roy 1975; Lavee and Zohary
2011). Additional wild forms of the olive species-complex occur in: (i) southern and east Africa (including west and south Arabian Peninsula); (ii)
Afghanistan; and (iii) west China (Zohary 1994; Lavee and Zohary 2011). Note that only the wild Mediterranean oleaster stock seems to be the
progenitor of the domesticated olive.
laperrinei, subsp. cuspidata, subsp. ferruginea, subsp.
chrysophylla, subsp. africana). The morphological
differences between these more tropical wild olives
and their Mediterranean counterparts are relatively
small. Consequently, Green and Wickens (1989)
regarded some of them only as an additional sub-
species of the European olive and named them
O. europaea L. subsp. cuspidata (Wall.) Ciferri.
However, these south Asian and east African olives
are geographically separated from their
Mediterranean relatives by wide morphological
and molecular discontinuities. They are also
adapted to different environments. Such isolation
may justify their ranking as independent species.
Using multiple types of molecular markers (e.g.
mitotypes, SSR, and RAPD data), Breton et al.
(2003, 2006) concluded that eastern and western
Mediterranean oleaster populations differ, with cul-
tivars of eastern origin (populations from Israel,
Syria, and Turkey). Also, they suggest at least two
simultaneous domestications occurred on opposite
ends of the Mediterranean Basin, possibly the sec-
ond one in the Spain/Corsica region.
Other non-Mediterranean wild types closely
related to the crop are O. europaea subsp. laperrinei
FRUIT TREES AND NUTS 119
(Batt. & Trab.) Cif. and O. europaea L. subsp. cerasi-
formis (Webb & Berth.) Kunkel & Sunding [= O.
europaea L. var. maderiensis Hart.]. The first is a
Saharo-Montane relic that bridges the Mediterranean
forms with their African savannah relatives. It is
confined to a few inner mountains in the Sahara
Desert and does not come close to the Mediterranean
forms, except perhaps in the southern Atlas
Mountains. The second is the wild olive of the
Macronesian Islands. (For details on the taxonomy
and distribution of all these non-Mediterranean
wild olives consult Green and Wickens 1989; Zohary
1994; Lavee and Zohary 2011).
The evidence from living plants therefore shows
that the domesticated olive is closely related to the
Mediterranean oleaster wild forms. These could be
regarded as the general stock from which the
domesticated fruit tree had been derived.
Archaeological evidence
It is likely that olives were collected from the wild
long before their domestication. The olive find in
Epi-Palaeolithic Ohalo II, Sea of Galilee (Kislev et al.
1992; Simchoni 1998; Weiss 2002, 2009; Weiss
120 DOMESTICATION OF PLANTS IN THE OLD WORLD
et al. 2004, 2008) undoubtedly represent gathering
from the wild. Some olive stones were found in ca.
8,150–7,850 cal BP Pottery Neolithic Nahal Zehora,
Mount Carmel, Israel (Kislev and Hartmann forth-
coming). Also the thousands of waterlogged stones
(and stone fragments) uncovered in the submerged,
late Neolithic (ca. seventh millennium BP) sites off
Mt Carmel (Galili et al. 1989; Galili and Shavit 1994–
5) seem to represent gathering from the wild as well
(see Kislev 1994–5). The numerous charred stones
found in contemporary Dhali Agridhi, Cyprus
(Stewart 1974), are very likely of a similar nature. So
are the stones and/or charcoal recovered from the
Mesolithic and Neolithic Grotta dell’ Uzzo, Sicily
(Costantini 1989) or from Neolithic and Chalcolithic
sites in Spain (Buxó 1997), and in Natufian and early
Neolithic Nahal Oren, Mt Carmel (Noy et al. 1973).
Definite signs of olive domestication come from
several Chalcolithic sites in Israel and Jordan.
Numerous well-preserved, carbonized olive stones
(see Fig. 34), as well as charred olive wood, were
uncovered in ca. 6,800–5,800 cal BP Chalcolithic
Tuleilat Ghassul, north of the Dead Sea, together
with cereal grains, dates, and pulses (Zohary and
Spiegel-Roy 1975; Kislev 1987; Neef 1990). This site
lies far outside the natural range of wild olives. No
oleaster olives occur today in the lower Jordan Valley,
or below the sea-level escarpments. The region is
too dry for it, which was probably the case in
Chalcolithic times. In Israel and Jordan, the areas
nearest to the lower Jordan Valley that support wild
olives are the western flanks of the Judean Hills, Mt
Carmel, and the Gilead. This indicates that the
Tuleilat Ghassul olives may have been products of
cultivation. They were probably raised under irri-
gation in a similar manner in which olives are
grown today in the Jordan Valley.
The finds of rich olive remains in Tuleilat Ghassul
are supplemented by olive stones and charcoal
remains from: (i) three additional Chalcolithic sites
in the Jordan Valley, namely Tell Saf (Gophna and
Kislev 1979), Tell Shuna North, and Tell Abu Hamid
(Neef 1990)—they contain masses of fragments of
crushed stones; i.e. waste products of olive press-
ing; and (ii) contemporary sites in the Golan
Heights (Epstein 1978, 1993). Additional support is
given by a simultaneous marked rise in olive pol-
len values around 6,500 cal BP in the pollen dia-
gram of Birkat-Ram, Golan Heights (Neumann
et al. 2007).
Later, olive remains abound in ca. 4,900–4,700 cal
BP Early Bronze Age Arad (Hopf 1978c), Bab edh-
Drah (McCreery 1979), ca. 4,900 cal BP Tell es-
Sa’idiyeh, Jordan (Cartwright 2003; and pers.
comm.), and several other Early Bronze Age sites
(Liphschitz et al. 1991; Liphschitz 2008). Outside
Israel and Jordan, Chalcolithic and Early Bronze
Age finds of olives have been few thus far. Olive
remains were found in fifth millennium BP Tell
Soukas, Syria (Helbaek 1962), in ca. 4,850–4,200 cal
BP Troy, Turkey (Riehl 1997), and in Early and
Middle Bronze Age Marki Alonia (Adams and
Simmons 1996), and ca. 3,450–3,350 cal BP Hala
Sultan Tekke (Hjelmqvist 1979a), Cyprus. A few are
also present in several early Minoan sites such as
Myrtos (Renfrew 1972), and Knossos, Crete (Evans
1928, p. 135). Some stones were found even from ca.
5,400–4,200 cal BP Zambujal, Portugal (Hopf 1981),
where other sources of evidence (Blitzer 1993) indi-
cate olive cultivation at this time. Sarpaki (Sarpaki
2009) indicates, however, that olives are totally
missing from Knossos. In the Middle and Late
Bronze Age, olive cultivation and olive-oil produc-
tion seem already to have been well established
throughout the countries bordering the east shore
of the Mediterranean Sea (Stager 1985) and in Late
Bronze Age, also in mainland Greece (Boardman
1976; Hansen 1988). The successful establishment
and large scale utilization of Olea is indicated also
by the increase, from early Bronze Age on, of olive
pollen grains in cores obtained from the Sea of
Galilee (Baruch 1990), as well as by the appearance
of numerous presses, olive oil vessels, and depic-
tion of olives in Bronze Age art.
Apparently, olive horticulture did not play a
major role in Bronze Age Egypt. Instead, the export
of olive oil from the southern Levant to Egypt is
well documented in the Bronze Age archaeology
(Stager 1985). Although charred olive stones were
found already in ca. eighteenth to the sixteenth cen-
turies BC Kom el-Rabi’a, Memphis (Murray 2000b)
and Tell el-Dab’a, Nile Delta (Thanheiser 2004), as
cited by (Newton et al. 2006), it is hard to determine
their origin. They were most probably imported.
Likewise, the unique finds of olive leaves and twigs
in garlands and jars of olive oil from ca. 1,325 BC

Tutankhamun tomb (Germer 1989b; Hepper 1990;
de Vartavan et al. 2010). Olive cultivation was prob-
ably introduced into the west Mediterranean basin
in the early part of the third millennium BP by the
Phoenician and Greek colonists (Boardman 1976).
This trade is well demonstrated by thousands of
olive stones, including some 2,500 in a single
Canaanite Jar, from late fourteenth century BC Late
Bronze Age Uluburun shipwreck off the south-
western Turkish coast (Haldane 1993).
In conclusion, in the Mediterranean basin olives
constitute a complex of wild forms, weedy types,
and domesticated clones. All are genetically loosely
interconnected and show similar climatic and
edaphic preferences. Information from the living
plants corresponds well with the available archaeo-
logical evidence. The earliest archaeological records
of olive cultivation come from the lands bordering
the eastern shores of the Mediterranean Sea—a ter-
ritory where genuinely wild oleaster olives thrive
today. Hence, on the basis of the combined evi-
dence, one is led to the conclusion that the olive was
probably first brought into cultivation in the
Levant.
Grapevine: Vitis vinifera
Grapevine, Vitis vinifera L. (Vitaceae), is one of the
classical fruits of the Old World. Together with the
olive, common fig, pomegranate, and date palm, it
comprises the oldest group of fruit trees around
which horticulture evolved in the Mediterranean
basin (Zohary and Spiegel-Roy 1975). Since Early
Bronze Age, grapes have contributed significantly
to food production in this area, providing fresh
fruits rich in sugar (the berries contain 15–25%
sugar), easily storable dried raisins, and juice for
fermentation of wine. The latter became an impor-
tant trade element in the countries around the
Mediterranean Sea.
The grapevine thrives in Mediterranean-type
environments, but, compared to the olive, it toler-
ates cooler and more humid conditions. For this
reason, viticulture extends northward beyond the
Mediterranean basin and succeeds in areas with a
relatively mild climate in western and central
Europe, and in western and central Asia. Vitis is a
perennial, diploid (2n = 2x = 38 chromosomes)
FRUIT TREES AND NUTS 121
climber. Wild populations of Vitis are dioecious and
reproduce from seeds. Under domestication, grape-
vine cultivars are propagated vegetatively by root-
ing winter dormant twigs or by grafting. The
grapevine has to be pruned yearly in order to
remain confined to a manageable size, and for regu-
lation of fruit production. As with most other fruit
trees, viticulture is based on the ‘fixation’ and main-
tenance of vegetative clones. Almost all grape culti-
vars bear hermaphrodite flowers and set fruit by
self-pollination. Traditional Old World viticulture
was based on thousands of distinct clones (Alleweldt
et al. 1991; Mullins et al. 1992; Olmo 1995b). These
vary widely in their habit, climate, and soil prefer-
ences, as well as in the shape, size, colour, taste, and
sweetness of their fruits. Juicy, small-berried varie-
ties with a rather acid taste are commonly used for
wine production, especially in Europe. Types with
sweet, large fruits prevail as table grapes. Some
clones, such as the traditional Black Corinth and
Sultanina, bear small, seedless berries appreciated
in raisin production. The berries of many of table-
bred varieties are seedless. The grapevine is a fast
growing fruit crop. Production usually starts three
years after planting.
Charred pips and wood constitute the bulk of
grape wine remains in archaeological excavation,
although some sites also yield whole fruits as well.
Wild ancestry
The genus Vitis contains some sixty-five species
native to northern and south-east Asia and central
America. All species are diploid, and are largely
self-pollinated.
The domesticated grapevine, V. vinifera L., is
closely related to an aggregate of wild and feral
forms (Fig. 35a) distributed over Europe and west-
ern Asia. Classical botanists regarded these wild
forms as an independent species, V. sylvestris
(C.C. Gmelin) Hegi. However, since these wild
forms show close morphological and genetic affini-
ties with the domesticated stock, most botanists
now regard sylvestris as the wild race of the domes-
ticated crop. Currently, they are classified as sub-
species sylvestris (C.C. Gmelin) Berger, together
with subspecies vinifera, within the V. vinifera crop
complex, and are considered as the source from
122 DOMESTICATION OF PLANTS IN THE OLD WORLD
which the cultivars could have been derived (see,
for example, Hegi 1935; Webb 1968; Mullins et al.
1992). These wild forms are largely dioceous, dip-
loid, and their population contains 50% male and
50% female individuals. In such wild populations,
cross-pollination is the rule.
Sylvestris grapes are widely distributed from the
Atlantic coast to Tadzhikistan and Kazakhstan (Map
16). A unique community of wild grapes grows in
southern Kazakhstanthe most northern area of wild
grapes in central Asia (Dzhangaliev et al. 2003).
They are primarily forest climbers, suited to the
humid and climatically mild deciduous forest area
south of the Caspian Sea and along the southern
coast of the Black Sea. Sylvestris grapevines also
abound in the relatively cooler and more mesic
northern parts of the Mediterranean sclerophyllous
vegetation belt—from Turkey and Crimea, through
Greece and former Yugoslavia, to Italy, France,
Spain, and north-west Africa. Along the Rhine and
the Danube, wild grapes used to penetrate deeply
into central Europe, yet these populations are
largely extinct. Scattered stands of wild grapes
occur also in more xeric territories and in less woody
places in the Fertile Crescent. However here, sylves-
tris grapes are mostly confined to gorges and to the
vicinity of springs and streams.
The boundary between the domesticated grape
clones and the wild forms is blurred by the presence
of escapees and secondary derivatives of hybridiza-
tion. Spontaneous crossing between wild plants
and cultivars have been found repeatedly where
sylvestris grapes grow in close proximity to vine-
yards, as the F1 hybrids are fully fertile. To summa-
rize, in V. vinifera we are faced, (in the Mediterranean
basin and in south-east Asia) with a variable com-
plex of wild forms (growing in primary habitats),
escapees and seed-propagated weedy types (that
occur mainly in disturbed surroundings), and
orchards of cultivars. The picture of the pre-agricul-
ture distribution of the wild grapevine has probably
been blurred by introgression between tame and
wild types, and by ‘weedy’ forms occupying human
disturbed environments. However, there can be lit-
tle doubt that sylvestris grapes are indigenous ele-
ments in southern Europe, the Euxinian and the
Hyrcanian vegetation belts south of the Black Sea
and the Caspian Sea, and relatively wet places in
the Mediterranean vegetation belt in south-west
Asia.
Sylvestris grapes differ from the domesticated
varieties by their relatively small and usually acid
berries (Fig. 35a), which are quite suitable for the
preparation of wine (for details see Olmo 1995a,
Table 3.1). Wild grapes may also be recognized by
more numerous (three to four) pips per berry, and
by somewhat more globular pips, generally with
stalks or ‘beaks’ constricted at the attachment to the
main body of the pip (Fig. 35b). Among the thou-
sands of domesticated grapevine clones and among
the variable wild sylvestris populations, the range
of variation in pip morphology is very wide.
Furthermore, the variation in the domesticated
assemblage and in the wild forms overlaps consid-
erably. For this reason, pip morphology cannot be
regarded as a completely safe diagnostic trait for
distinguishing between wild and domesticated Vitis
remains in archaeological excavations (but see dif-
ferent view in Kislev 1988).
Kroll (1999b), however, points to a difference in
pip production between wild and domestic grape-
vines. As wild Vitis are usually dioecious, an unpol-
linated female flower will fail to produce seeds
and will drop. In contrast (see below) nearly all
(a)
0 20 40 mm
Fig. 35a Fruiting wild grapevine, Vitis vinifera subsp. sylvestris
(Zaprjagaeva 1964, plate 283).
 FRUIT TREES AND NUTS 123

0 100 200 miles

0 200 400 km

Map 16 Geographical distribution of wild grapevine, Vitis vinifera subsp. sylvestris (based on Zohary and Spiegel-Roy 1975). Wild grape
extends toward the east beyond the boundaries of this map and reappears in a few locations in Turkmenistan, Tadzhikistan, and Kazakhstan.

domesticated grapes are hermaphrodites, and self-
pollination in their flowers is possible. Such a flower
may develop in several directions. It can drop unde-
veloped, develop into a berry with underdeveloped
pips, or develop into a berry with no pips (‘seed-
less’ varieties). For this reason, berries of wild forms
contain three to four fully developed pips, but never
underdeveloped pips. On the other hand, the ber-
ries of (traditional) domestic varieties frequently
contain two normal sized pips and a partly devel-
oped third pip. The presence of such reduced pips
may indicate domestication.

(b)

0 5 10 15 mm

Fig. 35b Variation in pip morphology in wild forms of the grapevine, Vitis vinifera subsp. sylvestris (Zaprjagaeva 1964, Plate 286). Note the
wide variation in the size and shape of the pips and their beaks.
124 DOMESTICATION OF PLANTS IN THE OLD WORLD
Domestication has brought about considerable
changes in the reproductive biology of the grape.
The first obvious change is the shift from the wild-
type sexual reproduction (by seed) to vegetative
propagation of clones (by cuttings or by grafting).
Just as in the olive, this is the cultivator’s way of
overcoming wide segregation in the seedlings and
achieving ‘fixation’ of desired types. The second
conspicuous development is the breakdown of the
wild-type system of sex determination. Usually,
wild sylvestris plants are dioecious, and their popu-
lations contain an equal proportion of male and
female individuals. Fruit setting in the wild depends
on cross-pollination. In contrast, nearly all domesti-
cated grape varieties are hermaphrodites (self-fer-
tile). Their flowers contain both a pistil and anthers.
The hermaphroditic condition ensures self-pollina-
tion and fruit setting without the need of male pol-
len donors. Sex determination in wild sylvestris
populations is governed by a single gene (Olmo
1995b). Female individuals are homogametic—they
carry a homozygotic recessive genotype Sum Sum
that suppresses the development of anthers. Male
individuals are heterozygous for a dominant, female
suppressing SuF allele, and have a SuF Sum geno-
type. The change, under domestication, from bisex-
uality (= dioecism) to hermaphroditism was attained
by a mutation shift to allele Su+ that is dominant
over Sum and brings about the development of both
the pistil and the anthers. Many hermaphroditic
cultivars are still heterozygous and posses a Su+ Sum
genotype. Other hermaphroditic clones contain a
homozygous Su+ Su+ genotype.
Vitis vinifera is the sole Mediterranean representa-
tive of the genus Vitis. This is a rather large and
variable genus comprising some sixty species (de
Lattin 1939; Levadaux 1956; Mullins et al. 1992).
About two-thirds of these species occur in temper-
ate North America, and one-third is distributed
over temperate and subtropical parts of East Asia.
All wild members of the genus Vitis are dioecious,
perennial woody climbers with coiled tendrils. All
known Vitis species, (excluding the Muscadinia
group which is now regarded as an independent
genus) have 2n = 2x = 38 chromosomes, and can be
easily crossed experimentally. Their F1 hybrids are
vigorous and fertile. In nature, the principal taxa
(‘species’) are often not fully reproductively isolated
from one another by various geographical and eco-
logical barriers.
Consequently, the boundaries between the main
morphological forms are frequently blurred, mak-
ing a clear-cut delimitation of species in Vitis a dif-
ficult or even an impossible task.
Before the colonization of North America, viticul-
ture in Europe and western Asia was restricted to
the gene pool present in the V. vinifera complex.
However, in modern times, several wild species
native to America have been used either as addi-
tional genetic sources for breeding of new varieties
of grapes, or as hardy wild sources of stocks confer-
ring resistance against the devastating attack of the
Phylloxera root aphid.
Archaeological evidence
Berries of wild Vitis were gathered from the wild
long before the domestication of the V. vinifera.
Charred pips and occasionally also entire berries or
raisins have been discovered in numerous prehis-
toric sites in Europe and south-west Asia, particu-
larly in northern Greece, former Yugoslavia, Italy,
Switzerland, Germany, France, and Spain (for enu-
meration of finds, see Riviera Núñez and Walker
1989; Kroll 1991; Colledge and Conolly, 2007).
Several dozens pips discovered in late Neolithic (ca.
9,000–7,600 cal BP) Dhali Agridhi, Cyprus (Stewart
1974) also seem to represent wild grape. In some of
these prehistoric sites, like ca. 8,200–6,650 cal BP
Anza, Macedonia (Renfrew 1976), ca. 7,800–7,600
cal BP Knossos, Crete (Sarpaki 2009), and ca. 6,350–
6,150 cal BP Tell Hârşova, Rumania (Cârciumaru
1996; Monah 2002; Monah and Monah 2008), such
pips were found together with domesticated grain
crops.
As already mentioned (see p. 122) the range of
variation in pip morphology in domesticated and in
wild forms overlaps considerably. For this reason
pip morphology cannot be regarded as a fully safe
diagnostic trait for distinguishing between wild
and domesticated Vitis remains in archaeological
excavations (and see different view in Kislev 1988).
However, the appearance of a crop in an ecological
environment where its wild relative does not grow
can support the argument for its domestication

status (Zohary and Spiegel-Roy 1975, and recent
review in Miller 2008).
The earliest probable domesticated finds of V. vin-
fera come from several Early Bronze Age sites in the
Levant. Pips and charred berries containing two to
three seeds were uncovered in Early Bronze Age
(ca. 3,500–3,300 cal BP) Jericho, Israel (Hopf, 1983).
These finds have been complemented by remains
found in Early Bronze Age Lachish (Helbaek 1958)
and Arad (Hopf 1978c), Israel, Numeira, Bab edh-
Dhra (McCreery 1979), and ca. 4,900 cal BP Tell es-
Sa’idiyeh (Cartwright 2003; and pers. comm.),
Jordan, and Kurban Höyük, south-eastern Turkey
(Miller 1986). Arad yielded two samples of charred
wood, and in Numeira numerous pips and hun-
dreds of whole berries were excavated. The fruits
are small, and the pips roundish and relatively
short-beaked. But since wild Vitis is absent today in
the Jordan Valley and Judea, and is unlikely to have
grown wild in these areas in the second half of the
sixth and the fifth millennia BP, the combination of
pips and charred wood provides a solid proof that
Vitis was already domesticated at that time.
Additional archaeological finds are available from
sites in south-east Turkey and in North Syria (for
enumeration of sites see Zettler and Miller 1995;
Miller 2008). The remains retrieved from Kurban
Höyük, Urfa district, south Turkey (Miller 1986;
Miller 1991, 2008) are instructive since pips of grapes
were recorded from 5% of the samples taken from
late Chalcolithic layers in this site. Their occurrence
increased to 10% of the samples taken from Early
Bronze Age levels. They became frequent (66%) and
appear in masses in mid-late Early Bronze Age. As
reported by Miller (1982), mineralized pips, pre-
sumably from cess deposits, are the most direct evi-
dence of fruit consumption in late fifth millennium
latrines at the site. This profile seems to reflect a
rapid growth of viticulture in the north Levant at
the start of the Bronze Age.
If confirmed, the charred pips from Chalcolithic
Tell Shuna North in the Jordan Valley (Neef 1998),
might represent an even earlier domestication.
Finally, it was suggested that infrared spectros-
copy detection of tartaric acid in residues on archae-
ological vessels is a marker for wine (Singleton
1994). As Miller (2008) notes, the chemical signature
of tartaric acid can result from various other related
FRUIT TREES AND NUTS 125
products: vinegar, raisins, grape juice, pekmez
(grape ‘molasses’), or one of few other plants that
produce it (see Cavalieri et al. 2003). At present, the
earliest evidence of tartaric acid comes from the
Neolithic site, ca. eighth millennium BP, Hajji Firuz,
Turkey (McGovern et al. 1996; McGovern 2003). As
this site lies at the edge of the current range for wild
grape (Map 16), it is hard to determine whether
these were wild or domesticated grapes. Such resi-
dues were found on the inner surface of a large jar
recovered from sixth millennium BP Godin Tepe,
western Iran, some 400 km distant from wild grape-
vine territory (Badler 1995; Badler et al. 1990;
McGovern and Michel 1995).
From the second half of the fifth millennium
BP on, fresh grapes, raisins, and wine are also
repeatedly recorded in Mesopotamian cuneiform
sources (Postgate 1987).
Grape growing areas in the south Levant were
exporting wine and raisins to Egypt already in the
Early Bronze Age (Stager 1985; James 1995). The
dimensions of this trade were demonstrated further
recently by a discovery of hundreds of vessels at the
tomb of one of Egypt’s earliest kings, buried around
5,100 BP at Abydos (McGovern 1998). Neutron acti-
vation analysis of jars showed that they were made
in south Levant. The infrared spectrometry tests of
sediments left inside the jars identified tartaric acid,
and confirmed the suspicion that they were wine
vessels. This conclusion was further supported by a
few pips found inside some of the jars. Somewhat
later, during the first and second dynasties (ca.
5,000–4,700 BP), signs appear that viticulture and
wine making were introduced into the Nile Valley.
Remains of grapes, depiction of grapevine growing,
wine production, and evidence on import of wine
and raisins continue to appear in Egypt from the
Old Kingdom times onwards (Germer 1989b; James
1995; Murray 2000c). Among the earliest finds are
from Pre-dynastic Tel Ibrahim Awad (de Roller
1992; Thanheiser 1992) and Tell el-Fara’in (Buto),
the Nile Delta (Thanheiser 1991), and a rich find of
fragments of desiccated raisins in third dynasty,
sometime around 4,600 cal BP, Djoser pyramid at
Saqqara (Lauer et al. 1950; Germer 1985). To the east
of Egypt, the earliest evidence currently comes
from ca. 2,200–1,600 cal BP Jerma (Garama), Libya
(Pelling 2008).
126 DOMESTICATION OF PLANTS IN THE OLD WORLD
Viticulture does not flourish in most parts of the
hot Nile Valley. As argued by Stager (1985), the most
plausible source for such introduction is the Levant.
Climatically, Egypt lies outside the optimal range
for Vitis. In dynastic times, raisins and wine were
largely imported into Egypt, grapes were grown
only as a luxury crop, and were restricted mainly to
the cooler delta area.
In the Aegean area signs of grapevine cultivation
appear somewhat later. In Thessaly and Macedonia
pips become so common at several late Neolithic
(late second half of seventh Millennium onward
BP) sites that Kroll (1991) remarks that their sheer
numbers could suggest cultivation. However, con-
vincing evidence comes only from Early Helladic
IV (fifth millennium BP) Lerna in south Greece,
where numerous pips were uncovered (Hopf
1961b). Additional remains were uncovered from
Middle and from Late Bronze Age sites (Hansen
1988). They include several hundreds of pips from
Late Helladic, second half of fourth millennium BP,
Kastanas (Kroll 1983). Parallel to the situation in the
Levant, the development of grapevine cultivation
in the Helladic and Minoan cultures is indicated
also by the presence of presses, and the appearance
of specific wine jars and wine cups.
Archaeobotanical documentation of Bronze Age
V. vinifera is also available from Transcaucasia
(Wasylikowa et al. 1991). Pips appear in Georgia in
Early Bronze Age. They become common in Middle
and Late Bronze Age (second and first millennia
BC). Also in Armenia, pips were frequently retrieved
from the second millennium BC onwards, while
the earliest central Asian finds came from ca.
5,000–3,700 cal BP Anau, Turkmenistan (Miller 1999;
Miller 2003).
From the end of the fifth millennium BP, there are
convincing signs of grapevine cultivation in
Baluchistan. Grape pips were uncovered among
food offerings placed in graves at ca. 4,000 BP Shahr-
i-Sokhta, south-east Iran (Costantini 1977). Charred
wood of Vitis (cf. V. vinifera) was retrieved from the
latest strata (Period VII, ca. 4,000 BP) in Mehrgarh,
Pakistan (Thiebault 1989), and pips were found in
contemporary, neighbouring Nausharo (Costantini
and Costantini-Biasini 1986), and in Miri Qalat
(Tengberg 1999). Similar to Egypt, these territories
lie far away from the wild habitats of V. vinifera.
These finds (particularly the charcoal) indicate
imported cultivation.
Viticulture was apparently introduced to the west
Mediterranean basin by Phoenician and Greek colo-
nists (Stager 1985; Buxó 1997). The Romans brought
this crop to temperate Europe (Loeschke 1933;
König 1989).
In summary, the situation in Vitis parallels that
found in Olea. The earliest archaeological indica-
tions of viticulture come from areas close to the
eastern shore of the Mediterranean Sea. In this terri-
tory (Map 16), we still find wild sylvestris-type pop-
ulations that might have been used for developing
the cultivated grape. While oleaster olives are con-
fined to the typical Mediterranean warm and sum-
mer-dry climate, sylvestris vines thrive in more
temperate and mesic conditions. These adaptations
are paralleled under cultivation. On the basis of the
available information (both from the living plants
and from archaeobotanical remains) we assume
that the Levant is the probable area in which Vitis
vinifera domestication could have been initiated.
Fig: Ficus carica
The Mediterranean fig, Ficus carica L. (Moraceae), is
the third classical fruit crop associated with the
beginning of horticulture in the Mediterranean
basin and south-west Asia (Zohary and Spiegel-Roy
1975). This common fig (on the basis of available
evidence) has been part of food production in this
area since Early Bronze Age, providing fresh fruit in
summer and storable, sugar-rich, dry figs all year
round. It is a relatively fast-growing fruit crop.
Production starts three to four years after planting.
Early fig cultivation was centred on typical
Mediterranean environments, in close association
with the olive and grapevine. The fig tree is diploid
(2n = 2x = 26 chromosomes), functionally dioecious,
cross-pollinated, and spontaneous populations
show half-male and half-female individuals
(Kjellberg et al. 1987). They depend entirely on sex-
ual reproduction from seed. Under domestication
their propagation is vegetative. The grower main-
tains desired genotypes by rooting of winter-dor-
mant twigs and occasionally by grafting. Since
F. carica is bisexual, female fruit-bearing clones
are vegetatively propagated. Such cultivars are

D E
Fig. 36 Common fig, Ficus carica. A–a twig with a flowering syconium (‘fruit’) on top; B–Mature syconium; C–Cross-section in a syconium;
D–Staminate flower with four anthers; E–Modified short styled female flower; F–Long styled female flower (with kind permission of J. Galil).
considerably heterozygous and when progeny-
tested, they manifest wide segregation. Most of the
seedlings are economically useless. The majority of
present-day cultivars, known as ‘common figs’, are
parthenocarpic. They were selected for their ability
to produce sweet, fleshy, large ‘fruits’ (syconia)
without pollination and fertilization. A second
group of highly appreciated female cultivars, the
‘Smyrna-type’ figs, still require pollination and fer-
tilization for fruit development (Condit, 1947;
Zohary 1995a; Flaishman et al. 2008).
A single dominant mutation determines the shift
(under domestication) to parthenocarpy.
The reproductive biology of F. carica is complex
(Storey 1976; Galil and Neeman 1977; Valdeyron
and Lloyd 1979; Beck and Lord 1988) and based on:
(i) a highly specialized inflorescence (the syconium,
Fig. 36) that the growers refer to as the fig’s ‘fruit’;
FRUIT TREES AND NUTS 127
3 mm
A B C
3 mm
F
(ii) the presence of two sex forms of which the
female morph is known as the ‘true’ or common fig,
and the male as caprifig; and (iii) an elaborate sym-
biosis between the plant and its pollinator, the fig
wasp Blastophaga psenes.
The syconium (unique to the genus Ficus), is a
fleshy flowering branch transformed into a hollow
receptacle that bears numerous minute flowers on
its inner surface, and is open to the outside by a nar-
row orifice (ostiole). The ‘true’ fruits are small dru-
plets or ‘seeds’ each developing in a female flower
inside the syconium.
Young syconia on the ‘true figs’ contain only pis-
tillate, female, long-styled flowers. In contrast male
‘caprifigs’ produce spongy, non-palatable syconia
containing both staminate male flowers (Fig. 36D)
and modified short-styled female flowers (Fig. 36E).
Female flowers nourish the Blastophaga larvae by
128 DOMESTICATION OF PLANTS IN THE OLD WORLD
turning into galls when eggs are laid into them.
True figs produce their main crops of ‘fruits’ in late
summer. Caprifigs usually bear three crops of syco-
nia during the year: over-wintering mamme, numer-
ous profichi that develop during spring, and
mammoni which ripen in autumn. In all three of
them, the Blastophaga wasps develop synchronously
with the syconia. Many of the pollen-carrying
female wasps emerging from mature syconia do not
land only on new caprifig syconia, but are attracted
also to the numerous young female syconia borne at
this time by true figs (Galil and Neeman 1977). They
enter the syconia through the orifice, become
trapped in them, bring about pollination, and per-
ish. Since the wasps are unable to insert their eggs
into the long-styled female flowers, the maturing
‘fruits’ of the true fig do not harbour Blastophaga lar-
vae even after having been visited by the insects.
The cultivator growing Smyrna-type figs makes
use of the symbiosis between the fig and the fig-
wasp by artificial pollination known as caprifica-
tion. Twigs with mature profichi are collected from
wild caprifig trees in early summer and suspended
on the true fig trees in the plantation. The pollinat-
ing wasps are thus brought near to the female syco-
nia. Caprification is an ancient procedure. It was
practised in Greek and Roman times (White 1970);
probably even earlier.
Carbonized small seed constitute the bulk of fig
tree remains retrieved from archaeological excava-
tion. Some sites have also yielded charred dry syco-
nia. The archaeological record of figs is sparse. It is
likely to be biased since the ‘seeds’ of F. carica are
small and may be overlooked.
Wild ancestry
The domesticated fig tree shows a close morpho-
logical resemblance, striking similarities in climatic
requirements, and tight genetic interconnections
with an aggregate of wild and weedy forms that are
widely distributed over the Mediterranean basin
(Map 17). Botanists (Tutin 1993; Browicz 1986;
Zohary 1973, p. 631) regard these spontaneous figs
as the wild progenitor of the cultivated fruit trees
and place them within F. carica L. Wild populations
of figs (Fig. 36) grow mainly in the low altitudes of
the Mediterranean maquis and garrigue formations.
They occupy rock crevices, gorges, streamsides, and
similar primary habitats. They are often comple-
mented by a wide range of feral types occupying
secondary, manmade habitats such as edges of plan-
tations, terrace walls in cultivation, ruins, collapsed
cisterns, cave entrances, etc. Frequently these
‘weedy’ types seem to have been derived from seed
produced by local domesticated clones that were
pollinated by the adjacent wild-growing caprifigs
(Zohary and Spiegel-Roy 1975; Flaishman et al.
2008) Wild females usually bear relatively small,
barely edible sycons.
The Mediterranean wild-feral-domesticated
F. carica species-complex is closely related also to a
group of non-Mediterranean, wild, deciduous Ficus
taxa distributed south and east of the Mediterranean
region and adapted to semi-arid and higher tem-
perature environments (Warburg, 1905; Zohary
1973, p. 630; Browicz 1986). Taxonomically, all of
them form a single natural group (series Carica in
Section Eusyce) within the genus Ficus. The latter is
an enormous genus comprising some 800 species
distributed mostly in the tropics. Tall, large figs
grow in the lower zone of the mesic, deciduous for-
ests of the Colchic (Black Sea) district of northern
Turkey and the Hyrcanic (south Caspian Sea) dis-
trict of Iran and adjacent Caucasia. These forest
types intergrade with the typical Mediterranean F.
carica. Most authors place these mesic, wild forms
within F. carica L. However, several Russian bota-
nists (see, for example, Zhukovsky 1964), treat them
as two independent species: F. colchica Grossh. and
F. hyrcanica Grossh.
Other members of the series Carica are warm-
climate, xeric, shrubby types distributed outside
the traditional area of fig cultivation: F. johannis
Boiss. [syn. F. geraniifolia Miq.] in the Zagros
Mountains and in southern Iran; F. virgata Roxb.
in Afghanistan; F. pseudosycomorus Decne. in south
Jordan, the Israeli Negev, Sinai, and Egypt; and F.
palmata Forssk. in Yemen, Somalia, and Ethiopia.
Some of these wild figs are interconnected by
intermediate forms, and taxonomically should
perhaps be considered only as eco-geographic
subspecies. But none of them, with the exception
of the tall, Colchic forest type, have estab-
lished noteworthy contacts with the F. carica crop
complex.

0 100 200 miles
0 200 400 km
Map 17 Geographical distribution of wild fig, Ficus carica. The distribution in the eastern half of the map is based mainly on data from Browicz
1986.
Several of these non-Mediterranean wild figs
were genetically tested (Storey and Condit 1969)
and found to have a chromosome complement
identical to that of the domesticated fig (2n = 2x =26
chromosomes). All members of series Carica seem
to be fully inter-fertile with the crop complex but
adapted to different ecological regimes. The close
affinities between the members of the Carica series
are also indicated by the behaviour of the symbiotic
wasp Blastophaga psenes L. At least in experimental
plots, this insect moves freely between the
Mediterranean and non-Mediterranean types.
Due to the complex situation in fig reproductive
biology, attempts to reach molecular identification
and classification of fig cultivars and their wild rela-
tives in the last decade or so were not yet fully suc-
cess (see Flaishman et al. 2008; Aradhya et al. 2010).
Archaeological evidence
The oldest known fig pips came from ca. 800,000 BP
Mousterian Gesher Benot Ya’akov, Israel (Melamed
et al. 2011). Later, charred fig pips were retrieved
from numerous Early Neolithic sites in south-west
Asia (Miller 1991, Table 2). The earliest Neolithic
finds came from ca. 11,700–10,550 cal BP PPNA
Netiv Hagdud (Kislev 1997) and ca. 11,400–11,200
Gilgal (Kislev et al. 2006a), Israel (where several
FRUIT TREES AND NUTS 129
fruits were found as well). Kislev, Hartmann and
Bar-Yosef (2006a), even suggest these find represent
first domestication (Lev-Yadun et al. 2006, and see
discussion in Kislev et al. 2006b). Additional
Neolithic finds include ca. 10,500–10,200 cal BP
Aceramic Neolithic Tell Aswad, Syria (van Zeist
and Bakker-Heeres 1985), ca. 8,150–7,850 cal BP
Nahal Zehora II, Israel (Kislev and Hartmann forth-
coming) and ca. 9,300–8,500 BP PPNB ‘Ain Ghazal,
Jordan (Rollefson et al. 1985). They continue to
appear in this area during later Neolithic times,
such as ca. 8,000–7,500 BP PPNC Atlit-Yam (Kislev
et al. 2004) and the first half of the seventh millen-
nium BP Ceramic Neolithic Jericho (Hopf 1983),
Israel. In addition, fig pips were discovered in ca.
8,650–8,400 cal BP Aceramic Neolithic Knossos,
Crete (Sarpaki 2009), and ceramic Neolithic Sesklo
(Kroll 1981a) and Toumba Balomenou (Sarpaki
1995), Greece. Fig remains were found in some west
Mediterranean early sites such as middle Neolithic
Grotta dell’ Uzzo, Sicily (Costantini 1989) and
Bronze Age Valeggio, north Italy (Villaret-von
Rochow 1958). It is impossible to distinguish
between pips of wild and cultivated figs; the exca-
vated material can be interpreted as either. However,
we are inclined to regard all these finds as repre-
senting collections from the wild. Most likely, the
Carica fig was introduced into cultivation at the
130 DOMESTICATION OF PLANTS IN THE OLD WORLD
same time as its two horticultural companions: olive
and grapevine.
As to later sites, carbonized pips of figs were
uncovered in Chalcolithic Tell Shuna North, Tell
Abu Hamid, and Tuleilat Ghassul (Neef 1990), in
Chalcolithic and ca. 3,500–3,300 cal BP Bronze Age
Jericho (Hopf 1983), in Early Bronze Age Bab edh-
Dhra (McCreery 1979) in the Dead Sea basin and ca.
3,450–3,350 cal BP Hala Sultan Tekke, Cyprus
(Hjelmqvist 1979b). Very small pips, as well as
several whole fruits, were retrieved from late
Neolithic and from younger strata of fifth millen-
nium BP Lerna Greece (Hopf 1961b). Cuneiform
sources indicate that figs were grown in
Mesopotamia from the second half of the fifth mil-
lennium BP on (Postgate 1987). In Egypt, the earli-
est archaeobotanical record of figs is from the ca.
5,450–5,650 cal. BP Maadi, and Pre-dynastic Tell el-
Fara’in (Buto), the Nile Delta (Thanheiser 1991),
and a beautiful drawing of baboons harvesting figs
in twelfth dynasty Khnumhotop III’s grave (ca.
3,900 BP) in Beni Hasan (Darby et al. 1977).
Contemporary indications of fig cultivation are
available also from Syria (Stager 1985). They show
that since the Bronze Age, figs accompanied olives
and grapes as main horticultural elements of the
rain-dependent agriculture in the Mediterranean
basin.
When the archaeobotanical data and the available
information extracted from living plants are com-
bined, one is led to the conclusion that in this fruit
crop too, the earliest signs of cultivation appear in
the eastern part of the Mediterranean basin and in
south-west Asia. This territory harbours the closest
wild relatives of the cultivated fig. In other words,
the wild F. carica forms of the Levant, southern
Turkey, and the Aegean belt can be regarded as the
most likely ancestral wild stock from which early
fig domesticates were derived.
In summary, the main changes in this fruit crop
under domestication were the shift to vegetative
propagation of female clones as follows: (i) the
increase of the size, amount of flesh, and sugar con-
tent of the syconia; (ii) the introduction of artificial
pollination and caprification; and (iii) the effective
selection for parthenocarpy where pollination is not
necessary for fruit set. Such developments were
already part of fig horticulture in classical times.
Sycamore fig: Ficus sycomorus
The cultivation of the sycamore fig, Ficus sycomorus
L. (Moraceae), has been almost exclusively an
Egyptian speciality. Compared with the common
fig, F. carica, it is a much taller and larger tree, but it
produces smaller, inferior syconia. Since early
dynastic times, ca. 6,000 years BP, the sycamore fig
was (and still is) a commonly used fruit crop and a
valued timber source in the lower Nile Valley
(Täckholm 1976). Outside Egypt, the sycamore fig
was less esteemed, though it was introduced into
the warmer parts of Israel and to a lesser extent to
several other locations on the shores of the
Mediterranean Sea (e.g. Lebanon, Cyprus, Tunisia).
As with other members of the genus Ficus, the
reproduction of the wild trees is from seed, and
seed setting depends on pollination by a specific
symbiotic wasp, Ceratosolen arabicus Mayr. In con-
trast, cultivation depends on clonal propagation
(rooting of twigs), and fruit production no longer
depends on the natural pollinator. Some clones are
parthenocarpic, (set fruit without fertilization). In
others, fruit maturation is artificially induced by
gashing the surface of the young syconia. This is an
old tradition in Egypt and in Israel (Galil et al.
1976).
Wild F. sycomorus is widely distributed in eastern
Africa from Sudan to South Africa, with an exten-
sion to Yemen. The tall, big trees, growing near
streams and in beds of ephemeral watercourses,
constitute a conspicuous component of savannah
landscapes. Throughout this area, F. sycomorus
reproduces sexually. As previously mentioned, in
this fig a specific symbiotic Ceratosolen wasp brings
about pollination. As far as we know (Galil et al.
1976), spontaneously reproducing sycamore figs, as
well as their pollinating wasp, do not occur today in
Egypt, but they still thrive in nearby Sudan.
Archaeological evidence
Remains of F. sycomorus start to appear in Egypt in
pre-dynastic times. The earliest records come from
ca. 6,000–5,500 BP Amratian sites, and from ca.
4,600–4,400 BP El Omari (Wetterstrom 1993, 1998).
Shortly after, sycamore fig remains appear in quan-
tity in Egyptian excavations from the start of the

fifth millennium BP onward (Galil et al. 1976;
Germer 1985; Murray 2000b). The fruit and the
wood, and sometimes even the twigs, are richly
represented in the tombs of the Early, Middle, and
Late Kingdoms. In some cases, the parched sycons
bear characteristic gashing marks indicating that
this art was practised in Egypt already in ancient
times.
There is no doubt that Egypt was the principal
area of sycamore fig development. In spite of the
absence of spontaneous trees and their pollinating
wasp in Egypt at present, it seems that F. sycomorus
was brought into domestication in Egypt (Galil et al.
1976).
Date palm: Phoenix dactylifera
The date palm, Phoenix dactylifera L. (Palmae), ranks
among of the first fruit trees that were brought into
domestication in the Old World. It was already part
of south-west Asian food production in the
Chalcolithic period (Zohary and Spiegel-Roy 1975).
Compared with the three other classic fruit trees
(olive, grapevine, fig), the date palm requires a
much warmer and dry-summer climate. High tem-
peratures, rainless summers, and very low humid-
ity are particularly important for fruit setting and
fruit ripening. Date horticulture is therefore centred
in the deserts south of the Mediterranean Sea and in
the southern fringe of south-west Asia (from south
Iran to the Atlantic coast of north-west Africa). This
is an almost rainless belt, in which date horticulture
depends on mild winters, hot and dry summers,
and a steady water supply, either by the presence of
a high level of ground water or by artificial irriga-
tion. Date palms can withstand considerable salin-
ity and they grow well even when watered by
brackish water. Arab folklore has cleverly summed
up the ecology of the date palm: ‘Its feet are in the
water and its head is in the fire of the sky’. Date
palms are very productive and the annual fruit
yield may be as high as 100–200 kg per tree. All over
these warm Arabian and Saharan deserts, the sugar-
rich fruits (some 60–70% sugar content) serve as an
important staple food for local people. Other parts
of the palm tree are also intensely used. In addition,
the worldwide date-palm production has grown
from 1.75 million metric tons in 1962 to 7.2 million
FRUIT TREES AND NUTS 131
metric tons in 2009 (http://faostat.fao.org). At
present, the largest producer of dates is Egypt (Chao
and Krueger 2007). The trunks are split longitudi-
nally into halves or quarters, and serve as beams for
building, the leaves are usedwidely for roofing,
matting, and basketry, and the fibres of the bark are
used to prepare ropes. Date palms start to bear
fruits four to five years after planting, and reach full
fruit production at eight to ten years. For details on
date palm cultivation and on its reproductive biol-
ogy, consult Nixon (1951), Dowson (1982), Reuveni
(1986), Chao and Krueger (2007).
Phoenix dactylifera is a diploid (2n = 2x = 36 chro-
mosomes) dioecious palm characterized by its tall
trunk (up to 25 m), its large, leathery, feather-shaped
leaves, and its ability (when young) to produce
basal suckers. The vegetative propagation of clones,
practised by the farmer, depends on these suckers.
Wild populations, as well as seedlings derived from
domesticated clones, consist of an equal proportion
of female and male individuals. This is brought
about by a single gene, or block of genes, determin-
ing sexuality. The male morph is heterozygous
and the female homozygous recessive. Date palms
are wind-pollinated. In domesticated palm-groves,
artificial pollination is applied by the farmer, and
the fraction of male individuals required for effec-
tive pollination is reduced to one male per twenty-
five to fifty female trees. This tradition is very old
(Schiel 1913; Pruessner 1920). Apparently it was
practiced in Mesopotamia already in Hammurabi’s
time (1,792–1,750 BC).
Wild ancestry
The wild stock from which the domesticated date
palm could have been derived has been satisfacto-
rily identified (Zohary and Spiegel-Roy 1975). The
crop is closely related to a variable aggregate of
wild and feral palms distributed over the hot and
dry southern parts of south-west Asia as well as the
north-eastern Saharan and North Arabian deserts
(Map 18). These spontaneous dates show close mor-
phological similarities and parallel climatic require-
ments with the domesticated clones. In addition,
they are fully inter-fertile with the cultivars and are
interconnected with them by occasional hybridiza-
tion. Botanists place these wild-growing date palm
132 DOMESTICATION OF PLANTS IN THE OLD WORLD
populations within the species complex of P. dac-
tylifera L. The wild-growing forms produce basal
suckers just as the domesticated varieties, but differ
from the domesticated clones by their smaller fruits.
These wild dates contain relatively little pulp and
are frequently unpalatable (or even indigestible).
Thus, in the date palm, domestication has led to the
increase in fruit size and pulp quality. Wild and
domesticated date palms differ in their mode of
reproduction. Sexual reproduction is the rule in
wild stands, while domestication has brought a
shift to vegetative propagation—to the ‘fixation’ of
desired highly heterozygous female clones. P. dac-
tylifera was well suited for this shift since it pro-
duces easily transferable suckers.
Spontaneously growing dates occur in almost the
entire area of date cultivation (Map 18) and fre-
quently represent secondary escapees. However, in
some areas in south-west Asia and very probably
also in the north-eastern Sahara, Arabia, and
Baluchistan, dates are genuinely wild and occupy
primary niches. Prominent among these locations
are lowland Khuzistan—the southern base of the
Map 18 Geographical distribution of wild and feral forms of date palm, Phoenix dactylifera (based on Fischer 1881). In south Arabia, Sudan,
Eritrea, Somalia, and Senegal they sometimes cross with the native P. reclinata; and in the Indus basin with P. sylvestris.
Zagros Range facing the Persian Gulf—and the
southern part of the Dead Sea basin (Zohary and
Spiegel-Roy 1975). In these exceptionally hot, dry
territories, wild-type dactylifera palms, with their
characteristic small and mostly inedible fruits,
thrive in gorges, wet rocky escarpments, seepage
areas in wadi beds, and brackish springs, where
they constitute a conspicuous element in the vege-
tation. In P. dactylifera we are therefore faced with a
variable complex of wild forms, segregating escap-
ees, and domesticated clones, which are all geneti-
cally interconnected by occasional hybridization. It
is often difficult to decide whether non-domesticated
material is genuinely wild, or whether it represents
weedy forms or secondary seedlings derived from
domesticated clones. However, it is impossible to
delimit the pre-agriculture distribution of the wild
date. There can be little doubt that the wild dactylif-
era forms are indigenous to the hot and dry parts of
south and west Asia. Date palms thrived in these
territories long before the initiation of agriculture.
This is attested by the finds of charred pieces of P.
dactylifera wood in 23,000 BP retrieved from Ohalo
0 200 400 miles
0 200 400 km

Fig. 37 Carbonized stones of date palm, Phoenix dactylifera, Chalcolithic Tuleilat Ghassul, Jordan (Zohary and Spiegel-Roy 1975).
II (Liphschitz and Nadel 1997), and the presence of
numerous date-pollen grains in Mousterian and
Baradostian strata in Shanidar Cave in the Zagros
foothills (Solecki 1975).
Phoenix dactylifera is the main south-west Asian
wild representative of its genus that comprises
some thirteen species distributed over Africa and
south Asia (Jones 1995; Barrow 1998). The only
other wild date that occurs in the east Mediterranean
basin is P. theophrastii Greuter, a narrow endemic
date palm confined to Crete and several locations in
coastal south-west Turkey (Barrow 1998). Several
species of Phoenix have been tested cytogenetically.
All show a constant chromosome complement
(2n = 2x = 36 chromosomes) and are fully inter-fer-
tile with one another. As in the case of Vitis, species
in Phoenix are reproductively isolated from one
another not by inter-sterility but by geographical
and ecological barriers. Two additional wild Phoenix
species grow on the fringe of traditional date culti-
vation in the Old World and, in these areas, they
may have enriched the gene pool of the domesti-
FRUIT TREES AND NUTS 133
cated fruit trees through spontaneous hybridiza-
tion. In south Arabia, and in Africa south of the
Sahara, date cultivation comes in contact with the
bushy P. reclinata Jack., whereas in the Indus Valley
contact is achieved with the tall, non-suckering,
more tropical P. sylvestris Roxb. In both territories,
hybridization between the cultivated date and the
native wild species has been detected (Munier
1973).
Recently, Rhouma et al. (2008) employed random
amplified microsatellite polymorphism markers
(RAMPOs) among thirty date-palm cultivars and
ten male trees to assess the genetic diversity and
phylogenic relationships in date palms. Their results
support the Mesopotamian origin of the date-palm
domestication.
Archaeological evidence
Archaeological sites with date palm remains are not
abundant so far, but this may reflect the fact that
most excavations in south-west Asia and the
134 DOMESTICATION OF PLANTS IN THE OLD WORLD
Mediterranean basin were conducted in areas too
cold for date-palm cultivation.
The few stones of date palm reported from Egypt,
Israel, Iran, and Pakistan, dated eighth and seventh
millennia BP, probably represent material collected
from the wild. The earliest remains of what seem to
be domesticated dates have been found by Seton
Lloyd in the ca 6,000 BP Ubaidian horizon at Eridu,
Lower Mesopotamia (Gillett 1981). R.J. Braidwood
(pers. comm.) remarked that ‘buckets of date stones’
were uncovered in this site. In Jordan, charred ker-
nels (Fig. 37) were uncovered in the ca. 6,800–5,800
cal BP classic Chalcolithic Tuleilat Ghassul in the
Jordan valley (Zohary and Spiegel-Roy 1975). Date
kernels were also retrieved from ca. 5.500–5,200 BP
Chalcolithic of the ‘Cave of the Treasure’ in Nahal
Mishmar, Israel (Bar-Adon 1980), and a single date
kernel is available from Early Bronze Age Jericho
(Hopf 1983). Date kernels were also found among
offerings deposited in a tomb in the ‘Royal Cemetery’
at Ur, lower Mesopotamia and dated to the late fifth
millennium BP (Ellison et al. 1978). Fruits and ker-
nels from numerous archaeological sites from the
early sixth millennium BP across eastern Arabia
indicate its important role ever since the Early
Bronze Age (Cleuziou and Costantini 1982; Nesbitt
1993; Méry and Tengberg 2009; Beech 2001, 2003).
Indeed, the occurrence of these finds near current
and past habitats of wild date, emphasize the need
to establish their domestication status, beyond the
sheer quantity of kernels in some of the finds.
From the Bronze Age on, date cultivation seems
well established in the warmer sections of south-west
Asia. Dates are frequently mentioned in Sumerian
and Akkadian cuneiform sources from the second
half of the fifth millennium BP onwards (Postgate
1987; Nesbitt 1993). They are depicted also on seal
engravings. The written documentation shows that
by the Late Uruk period (ca. 5,400–5,100 BP) the cities
of Sumer possessed flourishing date plantations.
Remains of dates appear rarely in Pre-dynastic
Egypt, such as Hierakonpolis and El Omari, and
prior to the Middle Kingdom much of the date
remains consist of leaves, fibre, and wood, rather
than fruits (Murray 2000b). Täckholm (1976) and
Murray (2000b) stresses that most finds are from the
New Kingdom and post-Pharaonic periods.
Täckholm (1976) noted that in the New Kingdom
remains of date fruits occur ‘in almost every second
tomb; in all forms, all sizes, and all colours’. Since
the second half of the fifth millennium BP, date-
palm cultivation seems to have been practised also
in the dry and warm parts of Baluchistan and in
adjacent parts of the Indian subcontinent (Costantini
and Costantini-Biasini 1985). In classical times,
dates appear to be an important food element in
lower Mesopotamia, Oman, the lower Jordan Valley,
the Nile Valley, and the desert oases of North Africa,
Arabia, and Baluchistan.
The available archaeological data, as well as the
information on the living date palm, seem to focus
on south-west Asia as the initial place of P. dactylif-
era domestication. This area which furnished the
earliest indications on date palm cultivation also
harbours wild forms from which the first cultivated
clones could have been obtained. The date palm
was probably brought into cultivation somewhere
in the lower Mesopotamian basin, or in some oases
in the southern fringe of the Fertile Crescent.
Pomegranate: Punica granatum
The pomegranate, Punica granatum L. (Punicaceae),
is an appreciated minor crop in traditional
Mediterranean horticulture. It is a deciduous bush
or small tree with conspicuous red flowers and
large (6–12 cm across) fruits characterized by a
leathery rind, persistent crown-like calyx, and
numerous seed covered with a juicy flesh which can
be eaten fresh, or whose juice can be extracted, and
also be fermented into wine.
The wild ancestor of the domesticated pomegran-
ate has been satisfactorily identified (Zohary and
Spiegel-Roy 1975). Wild forms of P. granatum
grow in masses in the south Caspian belt, in north-
eastern Turkey, and in Albania and Montenegro.
Domestication brought about an increase in fruit
size and a shift from sexual reproduction to clonal
propagation. Desired genotypes are easily main-
tained by rooting of winter-dormant suckers.
Cultivars set fruit by self-pollination.
Archaeological evidence
The pomegranate seems to belong to the early
ensemble of domesticated fruit trees in the Old

World. In addition to supplying fresh fruit, they
were also considered to be a symbol of fertility. The
earliest carbonized pips and fragments of pome-
granate rind have been obtained from Early Bronze
Age Jericho (Hopf 1983), Arad (Hopf 1978c), and in
the burned Palace’s kitchen in ca. 4,900 cal. BP Tell
es-Sa’idiyeh (ancient Zaretan), Jordan (Cartwright
2003, and pers. comm.). They also appear in Late
Bronze Age ca. 3,450–3,350 cal BP Hala Sultan Tekke,
Cyprus (Hjelmqvist 1979a), and Tiryns, Greece
(Kroll 1982). An interesting find is the numerous
seeds and fruit fragments in a large jar, and hun-
dreds of seeds across the site, from the late twenty-
fourth century BP Late Bronze Age Uluburun
shipwreck (Haldane 1993). As Ward (2003) indi-
cates, this is an indication of the pomegranate’s sta-
tus as a luxury item in Bronze Age eastern
Mediterranean culture. Pomegranates are recorded
in several ancient Mesopotamian cuneiform
sources—from the second half of the fifth millen-
nium BP on (Postgate 1987). In fact, pomegranate is
depicted in artefacts and paintings before its appear-
ance as plant remains, first occurring in Mesopotamia
(see details and discussion in Ward 2003). In Egypt,
Schweinfurth (1891) identified pomegranates
among the vegetable remains of the twelfth dynasty
(ca. 1990–1800 BC) Drah Abu el Naga, Egypt.
A large, dry pomegranate was found in the tomb of
Djehuty, the butler of Queen Hatshepsut, dating
from about 1470 BC (Hepper 1990). Pomegranates
do not occur in a wild state in south Levant.
Therefore, the occurrence of Punica remains in Early
Bronze Age in Israel and Jordan suggest well estab-
lished cultivation.
Apple: Malus domestica
The domestic European apple, Malus domestica
Borkh. (= M. pumila Mill.) of the rose family
(Rosaceae), is the most important (63 million tons
for 2008) fruit crop of the temperate, cooler parts of
the Old World (Way et al. 1991; Watkins 1995;
http://faostat.fao.org). Since Greek and Roman
times, apples have been extensively grown in the
temperate parts of Europe, south-west Asia, and
probably also in the Caucasus and in central Asia.
They thrive in regions where winters are sufficiently
cold to provide this fruit crop with a chilling phase
FRUIT TREES AND NUTS 135
necessary for breaking bud-dormancy. The fruit is a
pome, a ‘false fruit’, the greater part of which is
formed not by the ovary but by the receptacle of the
flower. In the core of the pomes of most cultivars
there are five leathery chambers or loculi, each nor-
mally containing two seeds.
Apples under domestication are very variable.
Some 2000 distinct cultivars have already been
described. They vary in fruit size (3–12 cm in
diameter), colour (red, yellow, green), shape, and
in the texture and taste of the pulp (sweet to acid).
The majority of present-day cultivars belong to
the group of dessert apples that bear relatively
sweet fruits that are consumed fresh, or are used
to prepare compotes or pies. Cooking apple vari-
eties produce harder and often larger pomes.
Other cultivars are cider apples, selected for pro-
duction of vintage quality cider. Less fancy types
of this alcoholic beverage can be prepared by fer-
menting ordinary dessert apples or even wild
‘crab apples’. Finally, some apples are grown for
their ornamental features, while other ones serve
as rootstocks on which superior scions are
grafted.
Wild ancestry
The apple, Malus Mill., contains some fifteen indig-
enous, ‘primary’ wild species. Most of them are
self-incompatible, contain 2n = 2x = 34 chromo-
somes, can be crossed with one another, and their
interspecific hybrids are fertile or at least partially
fertile (Korban 1986; Janick et al. 2003; Richards et al.
2009). However, some cultivars and wild forms are
triploid (2n = 3x = 51 chromosomes) and a few are
tetraploid (2n = 4x = 68 chromosomes). The domes-
ticated apple varieties (usually referred to as
M. domestica Borkh. or as M. pumila Mill.) are closely
related to a variable group of wild ‘crab apples’
(series Pumilae in section Malus of the genus Malus
(Rehder 1940; Way et al. 1991). It is a variable
domestic crop species complex that contains also a
large number of ‘secondary’ inter-specific hybrids,
introgressive hybridization products, and feral
populations.
These wild apples (Plate 14) are widely distrib-
uted over the temperate forest zone of Europe,
south-west and central Asia, and in Siberia. Closest
136 DOMESTICATION OF PLANTS IN THE OLD WORLD
to the crop (and freely crossing with it) are the
European, Caucasian, and Siberian wild crab apple
forms, which are placed by some apple taxonomists
in M. sylvestris (L.) Miller. In other taxonomic treat-
ments, the western populations of these crab apples
are kept as M. sylvestris and M. orientalis, while
Siberian forms are very variable and regarded as a
third subspecies, M. sieversii (Ledeb.) M. Roem.
(Janick et al. 2003).
Taxonomically, Malus is relatively a large genus,
containing several sections, subsections and series.
The crop (together with its close wild relatives) is
placed in section Malus subsection Malus series
Pumilae, of the genus Malus (Rehder 1940; Way et al.
1991, p. 30; Watkins 1995; Janick et al. 2003). This is
the largest section in this genus. Its members are
widely distributed over the temperate and cool for-
est areas of Europe and Asia from the Atlantic coast
of Europe to China, and expand to North America.
As stressed by Korban (1986), by Way et al. (1991)
and by Watkins (1995) most of the wild species, as
well as the domestica cultivars, cross readily with
one another. Their hybrids and ‘hybrid swarms’ are
fertile or at least partly fertile. This is particularly
the case with the domesticated apple varieties. If
they are introduced and planted close to their wild
relatives, they readily cross with them. Before apple
domestication, wild apple populations were appar-
ently isolated from one another mainly by geo-
graphic and ecological reproductive barriers.
However, this type of isolation broke down when
apple-culture moved in and tied these variable taxa
together.
Wild-growing crab apples reproduce from seed,
and are self-incompatible. They are characterized
by small fruits (1.5–3.0 cm in diameter), and are
very variable in their shape, colour, and taste. The
majority of plant taxonomists rank all these forms
as eco-subspecific units of the European wild ‘crab
apple’ M. sylvestris (L.) Mill. (Map 19). However,
because they show a marked differentiation from
eco-geographical races, some taxonomists regard
them as independent, vicarious species. Forms
growing in Europe are usually referred to as M. syl-
vestris (L.) Mill. subsp. sylvestris. Forms growing in
Anatolia, the Caucasus and Transcaucasus, and
northern Iran are regarded (Browicz 1972) as a dis-
tinct regional subspecies, M. sylvestris subsp. orien-
talis (Uglitzkich) Browicz. Further east M. sylvestris
is replaced in central Asia and Siberia, by a group of
central Asiatic wild crab apples that are either
grouped in M. sieversii (Ledeb.) M. Roem. or split
even further by Russian botanists who recognize
several additional local species: M. kirghizorum A. &
Fed. and M. turkmenorum Juz. & M. Pop. The east-
ern Siberian and north Chinese forms are placed in
M. prunifolia (Willd.) Borkh. Variation in apples is
further complicated by the fact that in numerous
places in Europe, south-west Asia, the Caucasus,
and in central Asia, apple cultivation has been prac-
tised in areas also supporting wild apple popula-
tions. Furthermore, in some regions—for example,
in Anatolia and the Balkans—wild apples were
commonly used, as in situ stock material for graft-
ing. Under such conditions, spontaneous hybridi-
zation between tame and wild apples occurs quite
frequently, resulting in the formation of swarms of
secondary hybrid derivatives. Recently, crossing of
the European apples has further complicated varia-
tion patterns in apples with numerous wild and
cultivated forms introduced from other geographi-
cal origins (particularly from central and Eastern
Asia). This development has already produced a
whole array of new hybrid combinations.
Based on morphological traits and several
genetic tools of species relationships, most
researchers (e.g. Wagner and Weeden 2000; Forte
et al. 2002; Harris and Juniper. 2002; Richards et al.
2009), support the central Asian apple M. sieversii
as the ancestor of domesticated apple. These stud-
ies point to the wild M. sieversii forests of Tian Shan
Mountains of central Asia as probable center of
origin. In particular, the recent full genome
sequencing of Velasco et al. (2010) validates M. siev-
ersii as the progenitor of the cultivated apple. It
also rejected the view of Coart et al. (2006) who
concluded that the European crab apple M. sylves-
tris is the ancestor of domesticated apples.
Taxonomically, if M. sieversii is the wild progenitor
of the domesticated crop, domesticated apple
should be regarded as its subspecies.
Archaeological evidence
Apples no doubt were collected from the wild
long before their domestication and before the

0 100 200 miles
0 200 400 km
Map 19 Geographical distribution of the European crab apple, Malus sylvestris (including subsp. orientalis) (based mainly on Browicz 1972,
and pers. comm.). Other wild apple forms, closely related to the domesticated apple, are common in central and east Asia (Forsline et al. 2003).
establishment of vegetatively propagated clones.
Numerous carbonized remains of small apples
(2–27 mm in diameter), often cut into halves for
parching, were uncovered in Neolithic and
Bronze Age Switzerland (Schweingruber 1979).
These remains correspond morphologically very
well to the wild crab apples that grow today it
that area. Similar remains of wild, small apples
were retrieved in numerous Neolithic and Bronze
Age sites in Europe; for example in former
Yugoslavia, Hungary, Poland, Czechoslovakia,
Austria, Germany, and Denmark (for an enu-
meration of finds, see Hopf 1973b; Schultze-
Motel 1968–1994; Kroll 1995–2000, as well as the
online databases http://www.cuminum.de/archae-
obotany/database/; http://www.archaeobotany.de/
database.html).
Small apples, cut transversely into halves,
threaded and dried on strings, were found among
offerings deposited in a tomb in the ‘Royal Cemetry’
at Ur, lower Mesopotamia, and dated to the Sargonid
period, late fifth millennium BP (Ellison et al. 1978).
FRUIT TREES AND NUTS 137
This discovery confirmed descriptions of strings of
apples in cuneiform texts from the same time. Since
apples do not grow wild in warm and dry
Mesopotamia, such small apples could represent a
long-distance import of wild apples or early
attempts of apple cultivation. Because Malus very
rarely lends itself to simple vegetative propagation,
apple cultivation in these early times would have
been necessarily based on seed planting; in other
words, on handling of variable segregating individ-
uals resembling ‘crab apples’. This is indeed what
the small, stringed apples look like. Yet even such
low-grade apples seem to have been appreciated in
ancient Babylonia, and were used for offering. A
similar early indication of apple cultivation comes
from ca. tenth century BC Iron Age Kadesh Barne’a
on the border between the Negev and Sinai. Here
several dozens of well-preserved charred fruits
have been discovered (Rudolph Cohen, pers.
comm.). Wild apples do not occur in this area and
their southernmost limit today is Turkey. These
138 DOMESTICATION OF PLANTS IN THE OLD WORLD
remains, therefore, strongly suggest that apples
were cultivated in the oasis of Kadesh Barne’a.
Whatever the nature of the Ur and Kadesh Barne’a
finds, one fact is clear: the apple did not evolve into
a major fruit crop in Bronze Age times. While the
cultivation of the olive, the grapevine, and the date
palm accelerated dramatically in the fifth and fourth
millennia BP, there are no signs of a similar develop-
ment in the apple. This tree seems to have obtained
its significant role in the Old World horticulture only
in classical times. This development is apparently
linked to the introduction of grafting. The Greeks
were already familiar with the art of grafting (White
1970, p. 248) and Theophrastus writes about the
maintenance of apple clones by this sophisticated
method of vegetative propagation.
In conclusion, we still know very little about the
time and place of apple domestication except that in
classical times apples were already extensively
grown in the Old World. Wild apples are widely
distributed over Europe and west Asia, where they
have diverged into numerous types and closely
related species that still retain full inter-fertility.
Apple cultivation is superimposed on this wild
background. It is therefore futile to try to delimit the
area of initial domestication on the basis of the evi-
dence available from the living plants. Most
(although not all) genetic evidence indicate that the
apple was domesticated in the Tian Shan Mountains
of central Asia, and reached Europe with remarka-
bly little later hybridization. Even so, exceptional
Malus individuals may have been picked up not
once and in a single place, but many times and in
several areas. Furthermore, many cultivars are
hybridization products combining genes from sev-
eral distinct geographical sources. Nevertheless,
most of the traditional varieties in Europe and
south-west Asia have their closest morphological
affinities with the wild M. sylvestris and this
undoubtedly is the principal, but not the only, wild
source from which the old cultigens in these regions
have evolved.
The archaeological finds make it clear that apples
have been extensively collected from the wild dur-
ing Neolithic and Bronze Age times. In the Bronze
Age and the Iron Age they may have been occasion-
ally planted from seed (as a luxury fruit). However,
the first signs of substantial apple cultivation appear
much later than those of olive, date-palm, fig, or
grapevine horticulture. The introduction of grafting
is apparently a key element in this development
and actually defines domestication. Because apple
cultivation depends almost entirely on this method
of vegetative propagation, it could only have been
practised after this technology became known in
south-west Asia and Europe. Where and when
grafting was invented is not clear, although its ini-
tiation was probably not in the area of Mediterranean
horticulture (see p. 115).
Pear: Pyrus communis
The domesticated pear of Europe and west Asia
Pyrus communis L. [= P. domestica Med.] (Rosaceae),
is second to the apple in its contribution to fruit pro-
duction in the cool and temperate parts of Old
World agriculture (Watkins 1986). Frequently, pears
are close companions of apples, and both fruit crops
thrive in similar environments. Pears, too, require
sufficient winter chilling to ensure normal flower-
ing and fruit setting. As in apples, the fruits are
pomes in which the larger part is formed by the
receptacle of the flower and only the inner ‘core’
containing the seed, develops from the ovary.
Usually the pomes have the distinctive pyriform
shape of Pyrus and their pulp contains stone cells
that give pears their characteristic gritty texture.
Domesticated European pears are extremely vari-
able. About one thousand distinct cultivars are
known by their varietal names. Like most other
fruit trees of the rose family, pears are self-incom-
patible (Zielinski et al. 1965). Wild pears are diploid
(2n = 2x = 34 chromosomes), and the majority of
domestic clones are also diploid. However, some of
the latter are triploid (2n = 3x = 51 chromosomes) or
tetraploid (2n = 4x = 68 chromosomes). Pears segre-
gate widely when grown from seed, and most of the
progeny is worthless. Like apples, the maintenance
of desired clones depends almost entirely on
grafting.
Wild ancestry
The genus Pyrus L. comprises some thirty wild spe-
cies distributed over Europe and Asia from the
Atlantic shore of west Europe to China (Rehder

0 100 200 miles
0 200 400 km
Map 20 Geographical distribution of wild forms of the European pear, Pyrus communis (based on Browicz 1992; additional information kindly
provided by this author). A few other wild pear forms, closely related to the domesticated pear, are growing throughout Asia (Dzhangaliev et al. 2003).
1940; Browicz 1993). Different wild species of pears
were domesticated in Europe, and independently in
east Asia. The domesticated pear of Europe (P. com-
munis) is closely related to a variable group of wild
pears distributed over the temperate parts of Europe,
northern Turkey and the Caucasus (Map 20).
The wild populations that thrive in temperate
Europe are frequently called P. pyraster Burgstd.
Those growing in the eastern parts of Asia are
referred to as P. caucasica Fed. Because of their
tight morphological and genetic affinities and full
inter-fertility with the crop, these wild pears are
now considered as the two eco-geographic wild
subspecies of the crop’s complex from which the
European domestic pear P. communis could have
been derived. Most of these wild pears are rather
spinescent and bear very gritty small fruits (1.5–
3.0 cm in diameter). In summary, all wild popula-
tions are diploid (2n = 2x = 34 chromosomes)
self-incompatible trees, and all are inter-fertile with
the European domesticated crop.
FRUIT TREES AND NUTS 139
Spontaneous crosses between these wild pears
and domesticated clones are quite frequent. In
numerous localities in Europe and western Asia,
pyraster or caucasica wild forms and communis culti-
vars are interconnected by feral forms and hybridi-
zation derivatives that thrive best at edges of
cultivation and in clearings of forests adjacent to
cultivation. Armenia and the Caucasus are extremely
rich in wild pear forms.
Because of their tight morphological and
genetic affinities with the crop, Browicz (1992,
1993) in his taxonomic treatment of Pyrus, placed
pyraster and caucasica pears as wild subspecies, in
P. communis.
Several other wild pears that grow in the general
area of traditional pear cultivation are inter-fertile
with the domesticated crop. Some or even all of
these species have enriched the genetic variation of
domesticated pears through hybridization and
introgression (Rubtzov 1944; Watkins 1986).
Prominent among these wild pears are P. spinosa
140 DOMESTICATION OF PLANTS IN THE OLD WORLD
Forssk. [= P. amygdaliformis Vill.], native to west
Turkey, the Aegean basin, and the south Balkans;
P. elaeagrifolia Pallas distributed over Turkey and
east Bulgaria; P. salicifolia Pallas in Caucasia and
adjacent areas in Turkey; the more arid P. syriaca
Boiss. of the Fertile Crescent (Plates 15 and 16); and
its closely related P. korshinskyi Litw. in central Asia
(for details on the distribution of these species see
Browicz 1982, pp. 49–52 and maps 79–83.) In earlier
times, wild pears were frequently grafted in situ.
This practice is still common in Anatolia, where
farmers frequently spare individuals of wild P. com-
munis, P. syriaca, P. spinosa, and P. elaeagrifolia at the
edges of cultivation and inside grain crop fields.
These trees are grafted with domesticated clones.
Often only one or a few main branches bear scions;
the others are left as they are. In such situations,
cross-pollination between the two types is almost
inevitable.
Finally, it should be pointed out that several wild
pears native to northern China and eastern Siberia
were apparently brought into domestication inde-
pendently and gave rise to the Far East pear culti-
vars (Watkins, 1986). These include: (i) the Chinese
sand pear, P. pyrifolia (Burm.) Nakai, (ii) the Ussurian
pear, P. ussuriensis Maxim, and (iii) the Chinese
white pear P. bretschneideri Rehd. Crosses between
P. communis and these three east Asian pears resulted
in the selection of new commercial pear varieties
and have caused further extensive genetic fusion in
the genus Pyrus.
Archaeological evidence
Pears were collected from the wild long before their
introduction into cultivation. Charred remains of
small fruits, sometimes halved, and probably dried,
were found in several Neolithic and Bronze Age
sites in Europe, e.g. north Italy, Switzerland,
Yugoslavia, and Germany (for enumeration of sites,
see Hopf 1978a; Schultze-Motel 1968–1994; Kroll
1995–2000; as well as the online databases http://
www.cuminum.de/archaeobotany/database/ ;
http://www.archaeobotany.de/database.html) .
Similar finds are available from late Neolithic
Dimini and Bronze Age Kastanas, Greece (Kroll
1983), and from some Tripolye culture settlements
in Moldavia and the Ukraine (Januševič 1978).
The archaeological evidence until now does not
provide definite clues about the beginning of pear
domestication. Reliable information on pear culti-
vation first appears in the works of the Greek and
the Roman writers (Hedrick et al. 1921; White 1970).
Fruit growers learned by experience that a grafted
branch gave fruit true to type, whereas seedlings
tended to revert to the wild state or to produce fruit
of poor quality. Theophrastus (ca. 371–287 BC)
describes three cultivated varieties in Greece and
mentions that they were propagated by grafting
since propagation by seeds resulted in inferior
progeny. Cato (235–150 BC) described methods for
pear cultivation and reported on six pear cultivars
grown in his time. Somewhat later, Pliny (23–79
AD) recognized thirty-five cultivars. The range of
fruit characters described by these authors covers
many types of today’s pears.
In conclusion, the patterns of variation in Old
World pears closely resemble those of apples.
Wild pears are widely distributed over Europe
and western Asia and have diverged into numer-
ous local ecogeographical races and species,
though they retain their inter-fertility. Pear culti-
vation is superimposed on wild populations and
hybridization between wild and tame pears has
repeatedly produced secondary hybrid deriva-
tives that tend to colonize disturbed grounds.
Faced with a crop complex of such a vast distribu-
tion and extensive genetic fusion, one can scarcely
determine an area of origin of pear cultivation on
the basis of the living plants. Various regions in
west Asia and in Europe, within the distribution
range of pyraster and caucasica pears, are equal
possibilities.
Plum: Prunus domestica
Plums rank third to apples and pears in terms of
their role in fruit production in the cooler and tem-
perate parts of the world. They are eaten fresh,
cooked, or dried as prunes. The principal cultivated
plums of Europe and south-west Asia are hexaploid
(2n = 6x = 48 chromosomes) Prunus domestica
L. (Watkins 1986; Ramming and Coicu 1995; Körber-
Grohne 1996). This is a variable fruit crop in the
Rosaceae, comprising two fully inter-fertile groups
of cultivars: (i) the relatively small fruited damsons,

bullaces, and greengages, P. domestica L. subsp. insi-
titia (L.) C. K. Schneider [= P. insititia L.]; and (ii) the
common European plums, P. domestica L. subsp.
domestica C. K. Schneider [= P. domestica L. sensu
stricto], with relatively large fruits. Also grown in
this part of the world is the closely related cherry
plum or myroblan, P. cerasifera Ehrh. [= P. divaricata
Ledeb.], with fruits resembling those of the dam-
sons. P. cerasifera is mainly diploid (2n = 2x = 16
chromosomes), but contains also tetraploid
(2n = 4x = 32 chromosomes) and hexaploid
(2n = 6x = 48 chromosomes) forms. Over most of
Europe, the cherry plum is used mainly as an orna-
mental, or as root-stock for grafting, and cultivars
planted for their fruits are rather rare (Stika and
Frank 1988). However, in the Caucasus, fruit bear-
ing cherry plums (Alyčia cultivars) constitute an
important fruit crop (Hanelt 1997). In central
Europe, the tetraploid (2n = 4x = 32 chromosomes)
sloe, P. spinosa L., have been taken into cultivation
locally. Clones of this spiny plum, with relatively
larger and less astringent fruits, have been selected.
They still survive (frequently in hedges) particu-
larly in the Czech Republic, Slovakia, Austria and
Armenia (Hanelt 1997; Gabrielian and Zohary
2004).
Other plums, belonging to the diploid
(2n = 2x = 16 chromosomes) P. salicina Lindl. have
been taken into cultivation in east Asia and are
known as Japanese plums. Until about 130 years
ago, the European plums and the Japanese plums
were separate crops, but in recent years, P. domes-
tica cultivars constitute an important fruit crop
(Hanelt 1997). In central Europe, tetraploid culti-
vars were crossed with the Japanese plum. Some of
the modern plum cultivars are derivatives of these
crosses.
Plum cultivation depends almost entirely on
grafting. Traditional methods of vegetative propa-
gation (rooting of shoots, etc.) do not succeed in this
fruit tree, except in the case of some suckering insi-
titia forms. Reproduction from seed is also impracti-
cal since cultivars are highly heterozygous and their
progeny segregate widely. Most of the seedlings are
economically worthless. The majority of hexaploid
P. domestica clones are self-compatible or at least
partly self-compatible. In contrast the diploid plums
are self-incompatible.
FRUIT TREES AND NUTS 141
Wild ancestry
The plums comprise a distinct taxonomic group (of
about two species), distributed over the temperate
parts of Eurasia and north America. Some botanists
follow the traditional classification and regard the
plums as an independent genus (Prunus L. sensu
stricto). Others, taking into consideration the fact
that plums can be crossed to several other stone-
fruit trees, regard them only as a subgenus within
an extended Prunus L. (which also includes the
cherries, almonds, peaches, apricots, laurel cherries,
etc.).
The closest wild relatives of the hexaploid com-
mon plum are the spontaneous, somewhat spines-
cent, insititia-like populations of P. domestica. As
argued by Werneck and Bertsch (1959), these
populations suggest that P. domestica could have
preceded agriculture and should be regarded as
an indigenous element in middle Europe. They
might be native also in the Balkans and Turkey.
They bear small (2–3 cm) subglobose fruits, which
are quite common in many parts of temperate
Europe and Turkey. They thrive in woods, cleared
hillsides, edges of cultivation, and hedges.
European botanists (see, for example, Webb 1968)
regard them as feral or naturalized elements.
However, pre-Neolithic remains of carbonized
plum stones discovered in the Upper Rhine and
the Danube regions closely resemble the stones of
present day.
In addition, the hexaploid European plums
(P. domestica subsp. domestica) and the hexaploid
damsons, bullaces, and greengages (P. domestica
subsp. insititia) are closely related to a variable
aggregate of wild and domesticated plums grouped
in P. cerasifera Ehrh. [= P. divaricata Ledeb.]. The wild
forms of this aggregate are distributed over the
Balkans, Turkey, the Levant, Caucasia, the south
Caspian coast, and central Asia (Browicz, 1996, Map
21). They are self-incompatible, reproduce from seed
and bear roundish, small (ca. 2 cm in diameter)
green, purple, or dark violet fruits, which taste very
much like the domesticated P. domestica plums.
P. cerasifera [= P. divaricata] wild forms fall into sev-
eral eco-geographical subspecies (Browicz 1996): (i)
subsp. cerasifera, with glabrous leaves and twigs,
grows in the more temperate northern territories; (ii)
142 DOMESTICATION OF PLANTS IN THE OLD WORLD
subsp. caspica (Kovalev & V. Ekim) Luneva [= P.
caspica Kovalev & V. Ekim], with pubescent petioles,
pedicels and leaves with velutinous lower surface,
occur in the Caspian coast of Caucasia and adjacent
Iran; (iii) subsp. ursina (Kotschy) Browicz [= P. ursina
Kotschy], with short pedicels and more or less
pubescent shoots and leaves, is native to the Levant
and south-east Turkey; and (iv) the smaller subsp.
sogdiana (Vassilez.) Cinovskis [= P. sogdiana Vassilez.]
native to central Asia (Tadzhikistan, Uzbekistan,
and southern Kirgizia). Chromosomally, wild
P. cerasifera is not yet adequately studied. Yet, it is
clear that it comprises diploid (2n = 2x = 16 chromo-
somes) forms (apparently the common chromosome
level) as well as tetraploid (2n = 4x = 32 chromo-
somes) and hexaploid (2n = 6x = 48 chromosomes)
chromosome races (Beridze and Kvatchadze 1981;
Watkins 1986) which are morphologically very simi-
lar to their diploid relatives.
Closely related to P. domestica (and to P. cerasifera)
are two additional wild plums: (i) P. cocomilia Ten., a
diploid (2n = 2x = 16 chromosomes) small tree,
native to the southern Balkan and western Turkey
areas, and extending to Calabria in Italy, with yel-
low (sometimes reddish-yellow), almost sessile
round fruits that ripen much later than those of the
wild cherry plum. (ii) P. brigantina Vill., a bush (ca.
2 m. high) with similar yellow, late-ripening fruits
restricted to a small mountainous area near
Briançon, south-east France. Both wild plums are
quite similar morphologically. They might repre-
sent only geographical races of a single species.
Domesticated hexaploid P. domestica had been
thought to be a polyploid product of a cross between
diploid P. cerasifera [= P. divaricata] plums and tetra-
ploid sloe, P. spinosa L. (Crane and Lawrence 1952,
p. 237; Watkins 1995). The latter is a wild, thorny
bush widely distributed over central and northern
Europe and the cooler and wetter areas in western
Asia. However, the available cytogenetic evidence
in support of such polyploid origin of the hexaploid
domesticated domestica plums is far from convinc-
ing. Comparative morphology does not confirm
this supposition. The domestica plums look strik-
ingly similar to the cerasifera forms. Both groups
intergrade and apparently cross with one another.
In contrast, P. spinosa, with its small and very astrin-
gent fruits, is morphologically distinct and seems to
be isolated reproductively from the domestica-ceras-
ifera plums. Hybrids between these two groups are
almost fully sterile. The hypothesis that hexaploid
P. domestica has a CC SS S1S1 genomic constitution,
(i.e. contains two genomes (S,S1) contributed by P.
spinosa), does not comply with the above facts. As
argued by Zohary (1992), it is unlikely that the sloe
contributed two genomes (or even a single genome)
to the domesticated hexaploid domestica plums. In
conclusion, the 6X P. domestica plums seem to be
closely related to the 2X, 4X, and 6X P. cerasifera.
Together they form what seems to be a P. cerasifera-
P. domestica polyploid crop complex. If wild forms
of 6X P. domestica existed in south-east Europe and/
or south-west Asia prior to domestication (and they
probably did), they should be regarded as the ances-
tral stock for the development of this fruit crop.
However, if as some botanists believe, domestica
plums evolved only under domestication, the plau-
sible principal progenitor is the P. cerasifera aggre-
gate. In the Balkans and west Turkey, crosses with
2X P. cocomilia could have enriched variation under
domestication. Also the 4X sloe, P. spinosa, might
have contributed to the domestic gene pool, not by
allopolyploidy and donation of entire genomes, but
through secondary hybridization—by rare forma-
tion of 5X domestica X spinosa spontaneous hybrids
and subsequent introgression.
Archaeological evidence
Plum stones appear in several Neolithic and Bronze
Age sites in Europe, in countries such as Italy,
Switzerland, Austria, and Germany. They are vari-
able in shape, yet they fall within the morphological
range of present-day cerasifera and insititia plums
(Bertsch and Bertsch 1949; Körber-Grohne 1996). In
addition, a large quantity of stones reported as
P. domestica subsp. insititia was uncovered in Bronze
Age Tell Mudrets (Gudzhova Mogila), Bulgaria
(Popova 1995). All these finds seem to represent
fruit collected from the wild. In contrast with the
paucity of plum remains in Neolithic and Bronze
Age contexts, domestica-type stones were repeatedly
uncovered in numerous Roman sites in Germany
and neighbouring countries (Körber-Grohne 1996;
Stika 1996; Willerding 1996). While no archaeobo-
tanical finds preserved remains in 79 BC Pompeii,

Fig. 38 Plum, Prunus domestica, Roman Period Masada, Israel
(photograph kindly provided by M. Kislev, Bar Ilan University).
wall painting there depicts blue, purple, and yellow
plums. Feemster and Meyer (2002) identified the
blue and purple plums as P. domestica, while the yel-
low one remains unidentified. They continue to be
present also in later periods (Fig. 38). Such appear-
ance suggests cultivation.
In summation, we still know very little about the
origin of cultivated P. domestica and the beginnings
of plum domestication, but since its culture depends
on grafting, this fruit tree was probably taken into
cultivation together with apples and pears. The ear-
liest literary records of plum planting and grafting
are from Roman times.
Cherries: Prunus avium and P. cerasus
Cherries are characteristic fruit trees of the cooler,
temperate parts of the Old World (Iezzoni et al. 1991;
Watkins 1995). Two species of cherries are widely
grown: (i) the diploid (2n = 2x = 16 chromosomes)
FRUIT TREES AND NUTS 143
sweet cherry Prunus avium L., and (ii) the tetraploid
(2n = 4x = 32 chromosomes) sour or morello cherry,
P. cerasus L.
The sweet cherry, P. avium L., is a rather tall
tree (up to 2 m high) with sweet, round, red-black
berries. Both the wild forms and the cultivars are
largely self-incompatible and require cross-polli-
nation in order to set fruits. Maintenance is by
grafting. According to Pliny, this fruit crop was
brought to Rome from the Pontus area (south-
eastern part of the Black Sea basin) by Locullus
in 73 BC upon his return from the East (White
1970).
The domesticated clones of the sweet cherry are
closely related to a group of wild and feral forms of
P. avium, which are widely distributed over
temperate Europe, northern Turkey, Caucasia, and
Transcaucasia (Meusel et al. 1965, Vol. 2, p. 227;
Webb 1968; Browicz 1982, Vol. 1, Map 72). The ripe
fruits of these spontaneous cherries are smaller than
those of the domesticated varieties and usually
reach only 10 mm in diameter. They show the same
black-red colour of P. avium and have either a sweet
or a bitter taste, but are never acid. Also the stones
of the wild forms closely resemble those of the cul-
tivars but are usually smaller (7–9 mm in the wild
as compared to 9–13 mm in most of the domesti-
cated forms). Both the cultivated and the wild
P. avium are diploid (2n = 4x = 32 chromosomes)
and largely self-incompatible.
The domesticated sour cherry, P. cerasus [=
Cerasus vulgaris Miller] is a smaller tree, rarely
exceeding 8 m in height, with bright red berries
and a characteristic acid taste (Hanelt 2001, vol 1.
pp. 505–508). Most cultivars are self-compatible.
The crop is closely related to, and fully inter-fertile
with, the ground cherry, P. fruticosa Pallas., a wild
tetraploid (2n = 4x = 32 chromosomes) self-polli-
nated shrub or small tree, native to parts of eastern
Europe and adjacent south-west Siberia. Both the
wild and cultivated are tetraploid, fully inter-fer-
tile, largely self-compatible and known to hybrid-
ize with one another (Hanelt 1997). P. fruticosa is
likely the direct wild progenitor of the domesti-
cated sour cherry, or at least a major contributor to
its formation through crosses with P. avium. Its
small fruits have a cherry-like taste but are too
astringent to be palatable.
144 DOMESTICATION OF PLANTS IN THE OLD WORLD
Archaeological Evidence
Cherries were collected from the wild long before
their cultivation. The numerous finds of Neolithic
and Bronze Age stones in central European settle-
ments (Bertsch and Bertsch 1949) seem to repre-
sent wild forms. Cultivated cherries make their
appearance rather late. The earliest reliable report
of cherry cultivation comes from classical times.
Pliny tells that Lucullus, in the first century BC
introduced to Rome a superior cherry variety which
he had obtained in the Pontus region of the Black
Sea (White 1970). Large quantities of cherry stones
were found in both Roman and medieval contexts
in many European countries (Baas 1980; Schultze-
Motel 1968–1994; Kroll 1995–2000).
Latecomers: apricot, peach, and quince
Three other members of the rose family, Rosaceae,
namely (i) apricot, Armeniaca vulgaris Lam.
[= Prunus armeniaca L.], (ii) peach, Persica vulgaris
Miller [= Amygdalus persica L., Prunus persica (L.)
Barsch.] and (iii) quince Cydonia oblonga Miller
[= C. vulgaris Pers.] are not Mediterranean ele-
ments. They seem to have arrived to the temperate
parts of Europe and the Mediterranean basin from
the east, and very probably already as domesti-
cated horticultural crops. Wild forms of apricot,
peach, and quince, fully inter-fertile with their
respective cultivars, grow in central and eastern
Asia. Smaller stands of wild apricot also extend to
the Transcaucasia (including Armenia and
Dagestan). The wild quince is much more restricted
in its geographic distribution. Wild forms of quince
thrive in the belt of the Hyrcanian vegetation in
the southern part of the Caspian Sea basin.
Probably, they arrived in south-west Asia and in
Europe via Persia and/or Armenia as crops at a
late date. Reliable archaeobotanical and literary
evidence on their arrival appears only in classic
period (fifth century BC).
Apricot Armeniaca vulgaris Lam. (= Prunus arme-
niaca L.) is a diploid (2n = 2x = 16 chromosomes)
fruit-tree, grown for its large, fleshy fruits. Some
cultivars are self-compatible, and a few others are
self-incompatible. Wild-growing populations of
apricot, are fully inter-fertile with the crop. Wild
forms of A. vulgaris, with their relatively small
leaves and fruits, are widely distributed over cen-
tral Asia, (particularly in Tien Shan mountain sys-
tem and several other mountain ranges, and in
north-eastern China, Korea, eastern Siberia, and
Ussuria). There, an additional two or three species
of Armeniaca occur (Mehlenbacher et al. 1990;
Watkins 1995). Smaller stands of wild apricot extend
to Armenia (Gabrielian and Zohary 2004) and to
Dagestan. Indeed, apricot stones were reported
from several ca. 6,000–4,750 cal. BP Eneolithic settle-
ments in Ukraine (Januševič 1978; Pashkevich
2005).
In the wild forms, and in many cultivars, the large
seeds (enclosed in hard shells) are bitter. Similar to
almonds, this bitterness protects the seeds from
predation. In some other cultivars this wild-type
adaptation broke down. The seeds are palatable
and frequently used as nuts.
The apricot was apparently grown in China some
3,000 years ago. It is widely assumed that it was
introduced into domestication in northern China or
eastern Asia (Bailey and Hough 1975; Watkins 1986;
Buttner 2001). However, the evidence supporting a
Chinese origin is still scanty. We still know very lit-
tle about the early history of apricot domestication
in central Asia, or Armenia—territories where wild
forms apricot occur as well.
The apricot reached the Mediterranean region
relatively late. Very likely, it was introduced into
south-west Asia from Iran or Armenia around the
third century BP (Hopf 1973b). A few hundred years
later apricot cultivation was well-established in
Syria, Turkey, Greece, and Italy.
Peach, Persica vulgaris Miller [= Amygdalus persica
L.; Prunus persica (L.) Barsch.] is a diploid
(2n = 2x = 16 chromosomes) fruit crop. Its wild
forms occur in the mountainous areas of Tibet and
western China. There are records pointing to peach
domestication in China as early as 4,000 years old
(Hesse 1975; Huang et al. 2008). On the basis of
available classical literature, Hedrick (1919, p. 463)
concluded that the peach reached Greece from
Persia in classical times (at about 300–400 BC).
However, a discovery of peach remains in ca. sev-
enth century BC Heraion, Samos (Kučan 1995) indi-
cates that its arrival might have been several
hundreds of years earlier. The Romans did not

cultivate it until AD first century (Huang et al. 2008).
Soon after, the peach was established as a valued
fruit crop in the Mediterranean basin. Bakels and
Jacomet (2003) surveyed the archaeobotanical finds
of ‘luxury food’ in Roman Period Europe. They
report peach remains in thirty-two sites, mostly in
the south; the few peaches in the north are all from
military settlements—apparently imported. Roman
Period finds are reported also from Israel and
Egypt.
Quince Cydonia oblonga Miller [= C. vulgaris Pers.]
is a large shrub or a small tree, largely confined to
the Hyrcanian vegetation belt in the southern
coastal areas of the Caspian Sea (Browicz 1996).
Quince is a diploid (2n = 2x = 34 chromosomes)
monotypic genus (only a single species in its genus)
in this restricted territory. Wild forms of C. oblonga
are frequent in Azerbaijan, especially in Talish and
in Dagestan districts, as well as along the Caspian
Sea coastal belt in northern Iran. Wild growing
quince populations are reported also from the Kopet
Dagh range in South Turkmenia, but they might be
feral. Apparently, domestication took place within
this restricted area. The relatively large and gritty
fruits (poms) of domestic quince clones are widely
used in south-west Asia and in the Mediterranean
basin. Usually, the fruits are not eaten raw, but
cooked to prepare jams, compotes and beverages.
So far, there is no archaeobotanical documentation
on this fruit crop. However, literary sources indicate
that the quince reached the Mediterranean region in
classical times.
When used as a stock for grafting (for apples,
pears, etc.) quince frequently produce a dwarfing
effect in the grafted scions, resulting in smaller trees
which make it easier to harvest the fruits.
Some archaeological find were retrieved mainly
in medieval contexts in Germany and other central
European countries (Schultze-Motel 1968–1994; and
Kroll 1995–2000).
Carob: Ceratonia siliqua
The carob, Ceratonia siliqua L. (Caesalpiniaceae), is a
characteristic constituent of the evergreen, maquis-
type vegetation in low-elevation areas in the
Mediterranean basin. All over these territories, this
evergreen dioecious tree has been extensively culti-
FRUIT TREES AND NUTS 145
vated for its pods, which are used both for human
consumption and as a sugar-rich animal feed (Batlle
and Tous 1997; Zohary 2002). Under domestication,
clones bearing fleshier, sweeter pods have been
selected. In superior cultivars, the pod’s sugar con-
tent can reach 50% or more. In plantations, the
majority of the trees are female. This requires the
addition of male pollen donors to the plantation
in attempt for satisfactory production of pods.
Hermaphrodite mutants that do not need cross-
pollination for fruit set occurred as well, and were
protected by growers. With the advent of mecha-
nized agriculture and the availability of relatively
low-priced sugar, carob cultivation in the
Mediterranean basin shrank considerably. However,
the endosperm of C. siliqua seed contains a unique,
highly viscose gum (galactomannan carob gum),
which is increasingly used as a stabilizer in com-
mercial food products. This new development
might cause a comeback of carob cultivation.
Similar to most other fruit crops, the domestica-
tion of C. siliqua is based on a shift from sexual
reproduction (in the wild) to vegetative propaga-
tion (under domestication). The carob does not
lend itself to simple methods of vegetative propa-
gation. Therefore, the growing of domestic carob
clones depends entirely on scion grafting. For this
reason, it is likely that this fruit tree was taken into
domestication only after the arrival of this propa-
gation technology into the Mediterranean basin.
Domestication probably, happened only in the later
part of the third millennium BP.
The available archaeobotanical documentation is
too scanty to suggest when and where this fruit tree
could have been taken into domestication (Zohary
2002). Carob pollen grains were detected in a pollen
core in the Hula Valley, north Israel (Weinstein-
Evron 1983), in sediments belonging to the Early
Würm period (ca. 40,000 BP). C. siliqua pollen grains
were found also in ca. 14,000 BP Hayonim Cave
(West Galilee) in contexts dated ca. 14,000 BP (Leroi-
Gourhan 1981). Two charred carob seeds were
uncovered in Pre-Pottery Neolithic contexts at
Nahal Oren (Noy et al. 1973), and pieces of charred
wood were discovered in ca. 10,000-9,300 BP Pre-
Pottery Neolithic A and in ca. 9,300-8,500 BP Pre-
Pottery Neolithic B levels in Jericho (Western 1971).
Finally, both seeds and wood charcoal of carob were
146 DOMESTICATION OF PLANTS IN THE OLD WORLD
uncovered at ca. 8,000–7,500 BP late Pre-Pottery
Neolithic Atlit-Yam (Galili et al. 1993). All these
finds antedate the beginnings of horticulture in
south-west Asia and show that C. siliqua was part of
Levantine flora at least since early Paleolithic times.
Carob remains continue to appear later, in Pottery
Neolithic, Chalcolithic, and Bronze Age contexts
(Liphschitz 1987; Kislev 1988; Zohary 2002;
Liphschitz, 2008). A few charred seeds were found
in Chalcolithic Abior Cave near Jericho (Kislev
1992), Early Bronze Age II Arad (Hopf 1978b) and
from Iron Age Ashkelon (Weiss and Kislev 2004).
Yet most of its remains (recovered mostly in Israel)
come from Roman times or later. Jewish and Roman
literary sources provide clearer clues. The tree is not
even mentioned in the Old Testament. In contrast,
carob cultivation is soundly documented in later
Jewish sources—the Mishna (first century BC to AD
second century) and the Talmud (third to the fifth
centuries AD). During the Mishna time, carob culti-
vars were extensively grown in Israel and their
pods were commonly used as human food (Feliks
1994). The Mishna specifies that the maintenance of
these clones depend on grafting. Furthermore, a
clear distinction is made between: (i) the superior
pods produced by grafted cultivars, which are fit
for human consumption, and (ii) the inferior pods
borne by non-grafted carobs, which are suited only
for animal feeding. In Rome, two contemporary
writers, Columella (4 BC to AD 65) and Pliny the
elder (AD 23–79), describe the carob and its cultiva-
tion at some length. In addition, numerous charred
carob pods were uncovered in AD 79 Herculaneum
and Pompeii (Meyer 1980).
The situation in the western part of the
Mediterranean basin is less clear. Wild growing
C. siliqua trees abound here. Yet some researchers
(see Batlle and Tous 1997) suspect that in the west-
Mediterranean basin the carob is naturalized.
Citrus fruits
Domesticated citrus fruits (the genus Citrus L.,
Rutaceae), had their origin in south-east Asia and in
the Indian subcontinent (Davies and Albrigo 1994;
Roose et al. 1995). In these parts of the world one
still finds the wild relatives from which they could
have been derived. Citrus cultigens comprise some
eight main types (citron, lime, lemon, pummelo,
bitter orange, orange, mandarin, and grapefruit).
All are largely inter-fertile, and when crossed pro-
duce a wide range of recombinant and intermediate
progeny. Citrus specialists usually refer to each of
these main types as a distinct domestic species.
However, the majority of them are probably only
hybrid products that originated, only under domes-
tication, from much fewer wild stocks; probably
from wild forms of citron, mandarin, and pummelo
(Scora 1988).
Numerous cultivars of citrus are apomictic. They
set seed containing asexual, nucellar embryos,
which retain the genetic constitution of the mother
plant. Such a mode of reproduction is exceptional
among fruit crops and it allows the maintenance of
desired genotypes by the simple planting of seeds.
Apomixis was apparently of major advantage in the
early stages of domestication of citrus fruits. By this
means, vegetative propagation was possible before
the introduction of grafting in the Mediterranean
area.
The citron Citrus medica L. is the only citrus crop
that was grown in south-west Asia and the
Mediterranean basin in Greek times. It apparently
arrived from India (via Persia), and its fruits, with
their characteristic thick aromatic rind, were appre-
ciated medicinally. Jewish literary sources (and
archaeological depictions in mosaics and coins)
indicate that in Hellenistic times, the citron acquired
a religious symbolic role (especially in Sukkot, the
Feast of Tabernacles). Theophrastus’ detailed
description of its cultivation and propagation leaves
little doubt that by the end of the fourth century BC,
C. medica was already well established in the east
Mediterranean basin. A single archaeobotanical find
suggests an even earlier arrival: Hjelmqvist (1979a)
uncovered a few charred citrus seeds in ca. 3,200 BP
Bronze Age Hala Sultan Tekke, Cyprus. But these
remains have not yet been directly dated (by AMS
radiocarbon method) to confirm their antiquity.
Better identified and dated finds came from Roman
sites in Egypt, Mons Claudianus, where entire fruits
including seeds were found (van der Veen 2001a),
Mons Porphyrites (van der Veen and Tabinor 2007),
and Quseir al-Qadim (van der Veen 2003).
Several other citrus crops: lemon, C. limon (L.)
Osbeck; Seville or bitter orange C. aurantium L.; lime

C. aurantiifolia (Chistm. & Panz) Swingle; and pum-
melo C. maxima (Burm.) Merrill., arrived in the
Mediterranean basin, apparently in early Islamic
times (Amar 2000; Ramón-Laca 2003). Ramón Laca
(2003) studied literary sources and concluded that
these crops entered Europe via the Iberian Peninsula
and Sicily. As for grapefruit (C. paradisi Macfad.),
mandarin (C. reticulata Blanco), and sweet orange
(C. aurantium L.), she concluded that they arrived to
the West only during the last 500–200 years as a
result of the trade with the British and Portuguese
colonies.
Almond: Amygdalus communis
The domesticated almond, Amygdalus communis L.
(Rosaceae) [= Prunus amygdalus Batsch., P. dulcis
(Miller) D.A. Webb], is a widely grown nut tree in
the Mediterranean basin (Watkins 1986), and prob-
ably one of the earliest, fruit tree domesticants in
Old World agriculture. Among the Rosaceous fruit
crops, almonds are the earliest to flower (Plate
17). They thrive best in the relatively warm
Mediterranean-type climate. Compared with olives
and grapes, almonds can endure somewhat drier
conditions. Most domesticated almonds, as well as
the wild forms, are diploid (2n = 2x = 16 chromo-
somes), self-incompatible, and cross-fertile. Some
cultivars, such as the Apulia or the Bari groups,
carry a mutation that renders them self-compatible.
Wild almonds are characterized by the amygdaline
contents of their seeds, which render them bitter
and poisonous. The main trends of evolution under
domestication were selection for mutant types with
non-poisonous, sweet seed, larger drupes, and
softer, thinner shells. Almonds cannot be propa-
gated by rooting, and its modern cultivation
depends mainly on grafting. In contrast with most
other fruit trees, almonds are also planted from
seeds. These seedlings produce considerably var-
ied fruits, in shape and size of the fruits, in hard-
ness and thickness of the shells, and in seed
bitterness. Most of the non-bitter individuals are
very tasty. The non-bitter condition is governed by
a single dominant mutation. Therefore, 75% or
more of the progeny raised from seed harvested in
non-bitter almond plantations produce sweet fruits
(Heppner 1923; Heppner 1926; Spiegel-Roy and
FRUIT TREES AND NUTS 147
Kochba 1981). For this purpose, farmers tradition-
ally collected seeds from trees producing sweet
seeds. Seed planting necessitates rogueing of
unwanted bitter individuals. This has been a tradi-
tional way of almond cultivation in numerous
localities in the Mediterranean basin and in south-
west Asia.
Wild ancestry
The genus Amygdalus L. comprises some twenty-six
vicarious species distributed over south-west and
central Asia, as well as south-east Europe (Browicz
and Zohary 1996). The almond is closely related to
an aggregate of wild forms native to the Levant
countries. These wild almonds, which are placed
taxonomically within A. communis complex, fall into
two intergrading eco-geographical races:
(i) relatively large wild and weedy forms, A. com-
munis subsp. spontanea, thrive in Mediterranean
environments (350–800 mm annual rainfall)
and resemble closely the domesticated varieties
in flower and leaf morphology, in early bloom-
ing, and in their climatic requirements; and
(ii) more xeric, smaller wild forms, A. communis
subsp. microphylla (Post) Browicz & D. Zohary
[= A. korschinskyi (Hand.-Mazz.) Bornm.], which
occupy drier ‘steppe forests’ or steppe-like
environments. These almonds have smaller
leaves and fruits compared to their more robust
Mediterranean counterparts.
Wild forms, placed taxonomically in subsp. sponta-
nea, are identified as the main progenitor of the
domesticated varieties (Browicz and Zohary 1996).
They differ from the cultivars mainly by their
smaller fruits, harder shells with fewer pits and
their intensely bitter seed. The bitterness represents
a chemical defence. It is brought about by the pres-
ence of the glycoside amygdalin, which becomes
transformed into deadly prussic acid (hydrogen
cyanide) after crushing, chewing, or any other
injury to the seed. The consumption of a few dozen
bitter nuts yields enough prussic acid to prove fatal
for human beings. Domestication of the almond
focused upon selection of non-bitter almonds. Sweet
148 DOMESTICATION OF PLANTS IN THE OLD WORLD
cultivars, known as var. dulcis, contain only traces
of amygdalin. Some of them are homozygous, oth-
ers are heterozygous for the dominant mutation.
Like numerous other fruit trees, A. communis
comprises a complex of interconnected wild forms,
weedy derivatives, and cultivars in its main area of
distribution. In the Levant, wild forms of subsp.
spontanea grow in Mediterranean maquis and oak
park-forest formations, thriving best on rocky
slopes and escarpments facing south. ‘Weedy’ forms
often colonize secondary habitats such as neglected
plantations, edges of terrace cultivation, and road-
sides. The latter are frequently feral derivatives, or
products of spontaneous hybridization between
cultivated and wild forms.
Several other Amygdalus species, growing outside
the Levant, are quite close morphologically to the
crop and are interconnected to it by spontaneous
hybridization (Browicz and Zohary 1996). Most
prominent among them are: A. webbii Spach, native
to the Aegean basin and south Italy; A. fenzliana
(Fritsch) Lipsky in north-east Turkey, adjacent
Armenia, and Caucasia, A. kuramica Korsh. in north-
east Afghanistan, and A. bucharica Korsh. in
Tadzhikistan and Uzbekistan. All these wild
almonds were very likely isolated geographically
from A. communis in the past. However, the spread
of almond cultivation into their territories brought
them in contact with the crop. As evident from
spontaneous hybrids encountered in places of con-
tacts, this superimposition initiated hybridization.
The extent of the gene-flow between the domesti-
cated crop and these wild almonds is hard to assess.
Similar to the apple or pear, the presence of inter-
mediates and recombinants indicates that introgres-
sion from the local wild species could have
facilitated the development of locally adapted
A. communis cultivars and/or helped in the estab-
lishment of local feral A. communis stands.
Archaeological evidence
Almonds were apparently collected from the wild
long before the plant’s domestication. It is unclear
how to differentiate between wild and domesti-
cated almond shells in archaeological finds (endo-
carp). Charred wild almond shells were uncovered
in Epi-Palaeolithic Ohalo II, Israel (Kislev et al.
1992; Simchoni 1998; Weiss 2002, 2009; Weiss et al.
2004, 2008), and in Epi-Palaeolithic Öküzini cave,
Turkey (Martinoli and Jacomet 2004), and in
Mesolithic and Neolithic levels at the Franchthi
Cave, Greece (Hansen 1991a). Charred PPNA
almonds were found in Netiv Hagdud, Jordan
Valley, Israel (Kislev 1997; Hartmann 2006), and in
Jerf al Ahmar on the Euphratus river, Syria (Willcox
2002; Willcox et al. 2008, 2009), and Early PPNB
Çayönü, Turkey (van Zeist 1972; van Zeist and de
Roller 1991–2, 2003). A rich find is available from
ca. 10,000 cal BP Late Natufian Hallan Çemi Tepesi,
South Turkey (Rosenberg et al. 1995). Here a large
quantity of charred wild fruits was uncovered,
suggesting the use of wild almond seeds. (Perhaps
latent toxicity was mitigated, either by leaching
the water-soluble prussic acid, or by extraction
and use of almond oil). In Knossos, Crete, almonds
played an important role during the Neolithic
Period, from Aceramic Neolithic, ca. 8,650–8,400
cal BP on (Sarpaki 2009).
In later periods, almond remains come from late
Neolithic Dhali Agridhi, Cyprus (Stewart 1974).
Aceramic Neolithic Çatalhöyük, Turkey (Helbaek
1964a; Fairbairn et al. 2002, 2005, 2007) Tepe
Musiyan, Iran (Helbaek 1969), as well as Late
Neolithic Dimini (Kroll 1979), Sesklo (Kroll 1981a),
and Sitagroi (Renfrew 1979, Table 3) in northern
Greece. They also occur in late Neolithic/
Chalcolithic Hacilar, Turkey (Helbaek 1970).
Recently, Martinoli and Jacomet (2004) differenti-
ated between wild Amygdalus and Prunus on the
basis of their morphology.
Almonds are also reported from Early Bronze
Age Bab edh-Dhra, Jordan (McCreery 1979). Here
they appear with numerous remains of grapevine
and olive, which probably represent domesticated
trees. These could have been grown either under
irrigation (near to the site) or under rain-depend-
ent conditions (at higher elevations nearby).
Remains of almond shells continue to appear in
Bronze Age sites in south-west Asia, Egypt, and
Greece. They were found in ca. 1,325 BC
Tutankhamun tomb in Egypt (Germer, 1989a;
Hepper 1990; de Vartavan et al. 2010; see also
Fig. 39). From classical times on, A. communis
is recorded as a characteristic element of
Mediterranean horticulture (White 1970).

Fig. 39 Desiccated almond shells, Tutankhamun’s tomb, New Kingdom, Egypt (Photo: A. McRobb, with kind permission of F. Nigel Hepper,
The Royal Botanic Gardens, Kew, UK).
In conclusion, almonds appear to have been
members of the earliest domesticated fruit-tree
assemblage in the Old World. They were probably
introduced into domestication in the eastern part of
the Mediterranean basin, apparently at about the
same time that the olive, grapevine, and date palm
were domesticated, i.e. in the Chalcolithic Period, in
about 6,300–5,300 cal BP.
The almond does not lend itself to vegetative
propagation from suckers, cuttings, or rooting.
However, unlike most Mediterranean fruit trees,
attractive almonds can be raised from seeds. This
indicates that the almond could have been domesti-
cated even before the introduction of grafting.
Walnut: Juglans regia
The common or Persian walnut tree, Juglans regia L.
(Juglandaceae), is a traditional nut of Old World
agriculture. This large tree produces beautiful hard
timber in addition to its fruits. The fruit is a drupe,
having a fleshy, green, outer layer that encloses the
‘nut’, which dries and falls off at maturity. The hard
shell of the nut (the endocarp) encloses a single
large, edible seed, which is rich in oil. Juglans regia is
FRUIT TREES AND NUTS 149
diploid (2n = 2x = 32 chromosomes) and is a wind-
pollinated, monoecious tree.
Juglans regia grows wild in mesic, temperate,
deciduous forests of the Balkans, eastern Turkey,
Armenia, Azerbaijan, north Iran, the south Caspian
region, the Caucasus, and central Asia. It reappears
in the Tien Shan province of western China. Wild
trees produce variable small fruits (2–3 cm in diam-
eter), which have relatively thick shells. Their seed
is as tasty as that of the domesticated tree. The
domesticated varieties have larger fruits, which are
3–6 cm in diameter. They are grown today in the
native distribution range of J. regia as well as in cen-
tral and western Europe and in the more xeric envi-
ronments in the Mediterranean basin and western
Asia. In the latter, the walnut thrives best in cool,
hilly areas and its cultivation usually benefits from
supplementary irrigation in summer.
Like the almond, the walnut does not lend itself
to rooting by cuttings or to propagation by suckers.
Cultivation depends mainly on grafting of selected
clones but, like almond, this nut tree can also be
raised from seed. However, by this method trees
vary considerably in the size and shape of their fruit
and in the thickness of the shells.
150 DOMESTICATION OF PLANTS IN THE OLD WORLD
Information pertaining to the time and place of
walnut domestication is still inadequate. Earliest
remains of wild J. regia appear in ca. 10,650–10,250
cal BP Mesolithic Balma Margineda, Spain (Marinval
1995), ca. 9,450–8,450 cal BP Late/Final Neolithic
Can Hasan III, Turkey (Hillman 1972; Hillman
1978), and ca. 7,550–6,450 cal BP Early Neolithic
Sammardenchia and adjacent sites, northern Italy
(Pessina and Rottoli 1996; Rottoli 2005; Rottoli and
Pessina 2007). Such remains continue to appear in
several European and south-west Asian countries
in the later Bronze and Iron Ages. As for the Levant,
Liphschitz (2008) reports wood remains from ca.
1,550–1,150 cal BC Middle Bronze onward in north-
ern Israel.
Eventually, J. regia became more common from
Roman time onwards. Bakels and Jacomet (Bakels
and Jacomet 2003) surveyed the archaeobotanical
finds of ‘luxury food’ in Roman Europe, and report
walnut remains in seventy-two sites. Roman period
finds reported also from Israel and Egypt. Such sites
are: the Cave of the Pool and Abior, the Sandal and
the Spear caves, Judean Desert, Israel (Zaitschek
1962; Kislev 1992; Simchoni and Kislev 2009), and
Bernike, Quasir al-Qadim, Mons Claudianus, and
Mons Porphyrites, Egypt (Cappers 1996, 2006; van
der Veen 1999, 2001a; Vermeeren and Cappers
1997/2002a). Greek, Roman, and Jewish literary
sources indicate that the walnut was already widely
grown in Classical times in the Mediterranean Basin
and south-west Asia. J. regia cultivation started
probably before Roman times, particularly in south-
west Asia.
Palynological data (Bottema 1980; Bottema and
Woldring 1984) indicated that J. regia disappeared
from south-east Europe and south-west Turkey
during the last glaciation (Würm glaciation), and
reappeared in the Balkans and west Turkey not
before the middle of the fourth millennium BP. This
late reappearance suggests that J. regia did not
return to these areas as a post-glacial wild element
but was reintroduced by humans. If this is true, the
spontaneously growing walnuts in the Balkans and
central Europe represent feral derivatives of culti-
vated walnuts introduced by humans as recently as
the Bronze Age. This point to north-eastern Turkey,
the Caucasus, and north Iran as the most plausible
area of walnut domestication.
A large quantity of desiccated and charred wal-
nut fruits were found in Roman Masada, Israel.
They include both thick (majority) and thin (minor-
ity) shells (Kislev pers. comm.). This indicates that
the mutant of ‘thin shell’ was already present some
2,000 years ago.
Chestnut: Castanea sativa
The sweet chestnut, Castanea sativa L. (Fagaceae), is
a large tree and a valued nut crop in the humid
parts of northern Turkey and southern Europe.
Chestnuts trees have been utilized extensively in
various south European countries and in Asia Minor
for managed coppicing and pollarding stands, as
well as for their nuts. The nuts formed an important
part of the diet in traditional farming communi-
ties and were also used as animal feed. Chestnut
production in southern Europe and south-west Asia
suffered debilitating damage from the attacks of
two fungal diseases (chestnut blight and ink root
disease), introduced from America in the 1890s. As
a result of these diseases, chestnut stands in many
places were decimated and production fell to 10.2%
of the former level.
Chestnut domestication is based primarily on the
selection of clones producing large and tasty nuts,
and their maintenance by grafting. Wild or natural-
ized stands which thrive on steep slopes and acid
soils are frequently grafted in situ with scions pro-
ducing superior nuts.
Castanea L. comprises some 10–12 species spread
over the northern hemisphere (Richardson 1986).
The domesticated chestnut is closely related to a
variable aggregate of wild and feral forms which
occur in the northern parts of the Mediterranean
basin, north Turkey, and Caucasia (for details, see
Meusel et al. 1965, Vol. 2, p. 121; Browicz 1982, Map
48). Such wild forms also penetrate deeply into the
climatically milder areas in central and western
Europe. Because of their close taxonomic affinity
with the domesticated clones, botanists include
these spontaneous chestnuts within C. sativa.
As with the walnut, our knowledge of the place
of origin and time of domestication of C. sativa is
still inadequate. Charred remains of this lovely nut
tree start to appear in several European countries in
the later Iron Age. Remains become more common

from Roman time onwards (for enumeration of the
finds see Schultze-Motel 1968–1994; Kroll 1995–
2000, as well as the online databases http://www.
cuminum.de/archaeobotany/database/; http://
www.archaeobotany.de/database.html). The paly-
nological record indicates that C. sativa disappeared
(or almost disappeared) from southern Europe dur-
ing the Würm glaciation, and that its pollen reap-
peared in quantity in Anatolia and Greece only at
about 1,500–1,300 BC (van Zeist 1980). An even
more recent increase in pollen frequency was found
in Italy and other west Mediterranean sites (van
Zeist 1980). This suggests that the comeback of
C. sativa to Europe was assisted by humans, and
again points to north Turkey and the Caucasus as
the most probable area for the initial chestnut
domestication. However, as van Zeist (1991) noted,
chestnut remains were retrieved from ca. eleventh
century BC Greifensee-Bochen near Zürich; and its
presence was attested (by charcoal remains) in ca.
1,500 BC Monte Leoni, north Italy. These finds sug-
gest that C. sativa might have survived the last gla-
ciation in some local refugia in Europe, and that its
reoccupation might not necessarily have been exclu-
sively from the east.
Hazelnut: Corylus avellana
Hazelnut (also known as filbert), Corylus avellana L.,
belong to Corylus L. (Betulaceae)—a genus of about
fifteen species of deciduous shrubs and trees.
(Wright 1986). This common hazel of Europe and
west Asia, appears as a shrub or small tree. Its nuts,
measuring 10–25 mm in length, are covered with an
involucre. C. avellana is a common component of the
broad-leaved oak and beech forest belt of temperate
Europe, Caucasia, north Turkey, and the Caspian
belt of Iran (Meusel et al. 1965, Vol. 2, p. 118; Browicz
1982, Map 48).
The tasty, rounded or ovoid nuts are easily col-
lected and shelled. Its remains have been repeatedly
retrieved from many Neolithic, Bronze Age,
Classical, and Medieval contexts all over Europe
and south-east Asia. The European hazel was also
taken into cultivation and planted both for its nuts
and for a supply of branches used for the prepara-
tion of hurdles and walking sticks. Superior clones
were traditionally kept by the layering of branches.
FRUIT TREES AND NUTS 151
Like almonds, hazels were also grown from seed
obtained from superior individuals. Although sex-
ual reproduction results in a wide variation in nut
size and shape, all nuts, even the small ones, are
tasty. It is not clear where and when the domestica-
tion of C. avellana was started, but apparently clones
of this shrub were already planted by the Romans
(White 1970, p. 259).
Corylus maxima Mill. is fully inter-fertile with and
frequently included taxonomically within C. avel-
lana. It is native to the Balkans, north Turkey, and
Caucasia. The fruit is somewhat larger than in typi-
cal C. avellana. C. maxima has very attractive nuts
(Lagerstedt 1975) which were, and still are, exten-
sively collected from the wild, and superior plants
were taken into cultivation (as clones). Because of
the bigger fruit size and better yields, planting of
maxima-type clones spread considerably in recent
times and often replaced C. avellana in west and cen-
tral Europe and Turkey. Today the world produc-
tion of hazel nut is primarily based on C. maxima
cultivars, (which are called the true filberts), or on
modern hybrid derivatives obtained from crosses
between C. maxima, C. avellana, and several addi-
tional Corylus species.
Pistachio: Pistacia vera
The pistachio nut, Pistacia vera L. (Anacardiaceae),
ranks among the most drought resistant of nut trees.
Domesticated varieties of this crop are extensively
grown in Iran, south-eastern Turkey, central and
south-west Asia, for its oval (ca. 1–2 cm long), tasty
nuts. The shell encloses a single oil-rich seed and
splits longitudinally, along its lateral suture, when
the nut is ripe. Pistacia vera is a dioecious, diploid
(2n = 2x = 30 chromosomes) wind pollinated nut
tree. Fruit-bearing female clones have been tradi-
tionally planted, intermixed with male individuals.
Domestication depends on grafting. Frequently, the
P. vera cultivars are grafted in situ on stocks of other
wild Pistacia species (P. palaestina, P. atlantica, P. khin-
juk, or P. terebinthus). They may be pollinated by
several other indigenous pistachio species.
Pistacia vera grows wild in north-east Iran, north
Afghanistan, and in the middle Asian republics—
Uzbekistan, Tadzhikistan, Kirgizia, and the south-
ern most parts of Turkmenia and Kazakhstan—where
152 DOMESTICATION OF PLANTS IN THE OLD WORLD
this xeric tree constitutes a conspicuous element in
the local semi-arid, ‘steppe forest’ vegetation belt
(Browicz 1988, map 5; Zohary 1996, Fig. 5). Wild
forms have smaller (but edible) fruits. They are also
collected and consumed by the local people. Recent
molecular research revealed the occurrence of
genetic diversity and affinity among wild forms
and cultivars in the genus Pistacia (e.g. Shanjani
et al. 2009; Pazouki et al. 2010).
The distribution area of the wild forms of P. vera
indicates that this tree could have been brought into
cultivation in central Asia. Indeed, the earliest
archaeological documentation of its use comes from
Bronze Age Djarkutan, Uzbekistan (Miller 1999),
and from late Neolithic and Bronze Age Tepe Yahya,
Iran (Costantini and Costantini-Biasini 1985).
Whether these finds represent collection from the
wild or domestication is hard to say. But the total
dependence of pistachio on grafting today suggests
relatively late domestication (the earliest literary
source of grafting technology is Theofrastus fourth
century AD). P. vera was introduced into the
Mediterranean region under the reign of Emperor
Tiberius (Buttner 2001, p. 1057). There are no clear
signs of P. vera cultivation in south-west Asia or in
southern Europe before classical times, although a
single P. vera half-shell was reported by Renfrew
(1973) in late Neolithic Sesklo, Greece. Another sin-
gle shell was found by Kroll (1983) in Bronze Age
Kastanas, Greece. In our view both these finds are
questionable. They probably represent more mod-
ern intrusions into the reported layers.
 C H A PTER 7
Vegetables and tubers
Compared to cereal grains and stones of fruits, veg-
etables and tuber crops are highly perishable. In
archaeological contexts, their soft parts are only
rarely charred and preserved. It is therefore not sur-
prising that very few remains of vegetables and
tubers have been found in archaeological excava-
tions. The only exceptions in south-west Asia are
Egypt, and to some extent caves in the Judean
Desert and the Dead Sea Rift Valley, where numer-
ous desiccated vegetable and tuber remains sur-
vived because of the extreme dryness.
The available archaeobotanical data on the early
phases of vegetable domestication in the Levant are
therefore fragmentary. However, this is partly com-
pensated for by evidence from Mesopotamian
Bronze Age literary sources, and by drawings and
descriptions found in Egyptian tombs. The com-
bined evidence shows that, since the start of the
fourth millennium BP, vegetable gardens consti-
tuted an integral element of food production both
in Mesopotamia and in the Nile Valley. Furthermore,
most of the vegetables grown at these early times
have been satisfactorily identified: watermelon,
melon, leek, garlic, onion, lettuce, and chufa appear
to be major constituents of Bronze Age vegetable
production.
The number of vegetable crops grown in human-
made gardens in South-west Asia, Egypt, and
Europe, seem to have increased considerably in the
third millennium BP (= first millennium BC). An
early written source is the list of plants grown in the
garden of Merodach-Baladan in Babylonia (ca. 2,720
BP). The list enumerates some of the earlier vegeta-
bles (lettuce and garlic), as well as beet, turnip, and
cress (Körber-Grohne 1987). Towards the end of the
third millennium BP the number of vegetable crops
seems to have increased considerably. Greek, Roman,
and Jewish classic writers were familiar with a large
number of vegetables (for reviews see Lenz 1859;
White 1970; Körber-Grohne 1987; Small 2006). At that
time cabbage, turnip, beet, celery, carrot, parsnip,
asparagus, and several other herbs, had already
become part of the vegetable garden. Significantly,
almost all these new cultigens have had closely
related wild forms growing in the same geographical
regions. Therefore, it seems they could have been
taken into cultivation in that part of the world.
Ethnographic data shows (see, for example, Ertuğ,
2004, 2009) that wild-growing vegetables take part in
the diet of traditional farming communities, even at
the present time. There is no doubt that this tradition
of collecting wild vegetables predates agriculture.
Recently, seeds of several Mediterranean herbs
and vegetables (such as celery, dill, and parsnip)
have been found in Neolithic layers in Europe (see
below and in Chapter 8). It is uncertain whether
these are wild forms or domesticated ones. They
might have been local wild-growing plants, garden
planting, or imported from the Mediterranean
region.
Watermelon: Citrullus lanatus
The watermelon, Citrullus lanatus (Thunb.) Matsum.
& Nakai [= C. vulgaris Schrad.], is belongs to the
gourd family (Cucurbitaceae), having crawling
liana. It is an annual summer crop, widely seed
planted in warm regions, and greatly appreciated
for its large sweet fruit with a useful source of water
(90% or more). It is diploid (2n = 2x = 22 chromo-
somes) and largely cross-pollinated. In addition to
the pulp, the oil-rich seeds are also consumed.
153
154 DOMESTICATION OF PLANTS IN THE OLD WORLD

The crop is closely related to four diploid wild Archaeological evidence

taxa, placed in the genus Citrullus (Jeffrey 1975:
Significantly, the characteristically small seed of C.
Singh 1990): (i) C. colocynthis (L.) Schrad., or colo-
colocynthis appear in several early Egyptian, Lybian,
cynth, a perennial wild watermelon bearing rela-
and south-west Asian sites, including Neolithic
tively small (5–8 cm), bitter fruits, which is widely
Armant (Lityńska-Zając 1993), ca. 5,650–5,850
distributed over the deserts and semi-deserts of
BP Nagada (Wetterstrom 1993, 1998), and ca. 5,500–
Africa, the Mediterranean basin, and south-west
5,300 BP Hierakonpolis (El Hadidi et al. 1996) in
Asia; (ii) C. ecirrhosus Cogn., a similar perennial
Egypt, Pre-Pottery Neolithic B Nahal Hemar Cave,
watermelon, restricted to the Kalahari Desert; (iii)
Israel (Kislev 1988), and a series of sites ranging
C. lanatus, wild growing, annual watermelon
from ca. 5,800 BP to Roman times in Libya (van der
forms, with somewhat larger fruits, which also
Veen et al. 1996). These finds indicate that humans
thrive in south-west Africa; and (iv) C. rehmii De
probably used the wild colocynth prior to its
Winter, an additional annual watermelon species
domestication.
reported from Namibia (de Winter 1990). All these
Watermelon was grown in the Nile Valley at least
wild types are inter-fertile with the crop
since the start of the fourth millennium BP. Several
(Shimotsuma 1963; Navot and Zamir 1987; de
finds of its characteristically large seeds (5–8 mm in
Winter 1990). Yet C. lanatus and C. rehmii stand
diameter), as well as a single find of leaves, are
somewhat closer to the cultigens in morphology
reported by Keimer (1924, pp. 17–18), the oldest
and molecular composition. On the basis of this
of them dating from the twelfth dynasty (ca.
evidence, some botanists concluded that the lana-
3,985–3,773 BP). Numerous desiccated watermelon
tus-type annual forms constitute the wild stock
seeds were found in Tutankhamun tomb, ca. 1,325
from which the crop evolved, and that south-west
BC (Germer 1989b; Hepper 1990; de Vartavan et al.
Africa probably was the place where watermelon
2010). The Agricultural Museum of Dokki, Cairo,
domestication took place.
has a sample of watermelon seed retrieved from the
Currently, the available archaeological informa-
New Kingdom Thebes (Darby et al. 1977). Van Zeist
tion does not seem to support south-west Africa as
(1983) confirmed these early finds and reported on
the locale of watermelon domestication. Soundly
the presence of watermelon remains in the founda-
identified remains of domestic watermelon appear
tion deposits of the walls of two eighteenth-dynasty
in Egypt already at the start of the fourth millen-
(ca. 3,500 BP) temples near Semna, Nubia.
nium BP. As far as we know, at this time farming
Wasylikova and van der Veen (2004) describe water-
was not yet practiced in south-west Africa (see chap-
melon remains as a funeral gift (together with
ter 9). In other words, when watermelon was
domestic grapes, olives, figs, dates, and rushnuts)
already a crop in Egypt, the cultural environment
in the same temples. Finally, Cox and van der Veen
for its domestication did not yet exist in the area
(2008) reported a significant size increase of water-
where we find its closest wild relatives today. This
melon seeds from the Roman to the Islamic period
contradiction is hard to explain.
at Quseir al-Qadim, Egypt. They also suggest that
The intensely bitter fruits of the colocynth have
the site dwellers had eaten the seeds during the
long been valued for their strong purgative
Islamic period, but not during the Roman period.
effects. The fruits are still collected by nomads in
the Sahara and south-west Asia and sold to phar-
macies. After special preparation, the seeds are
Melon: Cucumis melo
sometimes used as human food (Hedrick 1919;
Osborn 1968). The domesticated melon, Cucumis melo L., is a sec-
Crosses between wild C. colocynthis and the ond cucurbit crop, which was probably brought
domesticated watermelon have shown that the shift into cultivation in south-west Asia or in Egypt at a
from bitter fruits to non-bitter ones is caused by a relatively early date. It is a very variable crop that
recessive mutation in a single gene (Navot et al. includes both: (i) sweet fruited varieties (melons or
1990). muskmelons), and (ii) non-sweet green-fruited

forms (chate-melons, orfakus in Arabic). The latter
are rare today, frequently bear bent fruits, and are
consumed like cucumbers. They were traditionally
referred to as C. chate Hasselqu., or C. melo L. var
chate (Hasselqu.) Naud.
The wild ancestry of the domesticated crop
(2n = 2x = 14 chromosomes) is well established
(Jeffrey 1980; Sebastian et al. 2010). The domesti-
cated varieties show close morphological resem-
blance and full inter-fertility with a variable group
of wild and weedy annual and perennial melons
distributed over the subtropical and tropical parts
of Asia, Africa, and Australia (Kirkbride 1993). The
more xeric type (frequently referred to as C. callosus
(Röttl.) Cong.), with ovaries covered with dense
long hairs, is native to central and south-west Asia.
It is closest to the melon cultivars raised in western
Asia and the Mediterranean basin. The Indian and
east Asiatic cultigens show closer affinities to a sec-
ond, more tropical, wild melon race in which the
young ovaries are covered by a short, close
pubescence.
Archaeological evidence
There are few archaeological remains of melons.
However, they seem to indicate that C. melo was
cultivated in Egypt in the Bronze Age. Melon
seeds were discovered in Predynastic, ca. 5,650–
5,450 cal BP Hierakonpolis (Fahmy 1995, 2003,
2005; El Hadidi et al. 1996; Fahmy et al. 2008), and
ca. 5,450–5,300 cal BP Maadi (van Zeist and de
Roller 1993, 2003). Illustrations of offerings, as
well as faience models of what are clearly the bent
fruits of the green-fruited melons (var. chate) deco-
rate several ancient Egyptian tombs from the Old
Kingdom period on (Keimer 1924). Three carbon-
ized seeds are available from Late Bronze Age
Tiryns, Greece (Kroll 1982) and a few others from
ca. 4,000 BP Shahr-i Sokhta, east Iran (Costantini
1977).
The melon, C. melo was the principal (and the
oldest) cucurbitaceous vegetable of ancient
Mesopotamia, at least from the beginning of the
fourth millennium BP on (van Zeist and Waterbolk-
Rooijen 2003, p.74). The ‘small’ and ‘finger’ varie-
ties mentioned by Stol (1987b) were probably
cucumber-like, non-sweet forms. The ‘large’ and
VEGETABLES AND TUBERS 155
the ‘ripe’ forms could represent more advanced
non-sweet or even sweet melons. The sweet melons
are the melopepo of the Greeks. Indeed, numerous
C. melo seeds were uncovered in twenty-seventhth
century BP Heraion, Samos (Kučan 1995).
A third Old World principal cucurbit, namely the
cucumber, Cucumis sativus L. (diploid, 2n = 2x = 24
chromosomes), does not belong to the indigenous
vegetable ensemble of south-east Asia and the
Mediterranean basin. Wild forms of this vegetable,
namely C. sativum L. var. hardwickii (Royle) Alef.
occur in the Himalayas and in adjacent territories
east of this mountain belt (Jeffrey 1980; Kirkbride
1993; Sebastian et al. 2010). These wild forms were
reported also from the province of western Ghats,
Peninsular India (Kuriachan and Beevy 1992). Most
likely, C. sativus was taken into cultivation in India
and arrived in the Mediterranean basin rather late.
Exactly when this happened is still hard to say,
especially since the seeds of C. sativus and C. melo
are quite similar in their external features. They do
differ, however, in their seed coat anatomy (Kučan
1995). In the future, this might help to trace the
arrival of the cucumber into south-west Asia,
although typically cucumbers are eaten unripe
before the seeds mature.
Recently, Janick, Paris, and Parrish (2007) reviewed
the occurrence of cucurbits in Classical and Jewish
literature, as well as in artistic evidence from around
the Mediterranean basin. They described the use
and growing of melon, bottle gourd (Lagenaria sicer-
aria, used mainly as vessels or utensils), and water-
melon as the most important cucurbits in this area
during the Roman Period. However they found no
convincing evidence for the existence of C. sativus
around the Mediterranean in this time period, and
they conclude that the cucumis of Columella and
Pliny was not actually a cucumber, but C. melo subsp.
melo Flexuosus Group (snake/chate melons, or
fakus). It seems, therefore, that the cucumber arrived
to the area only after the classical times.
Leek: Allium porrum
The garden leeks (kurrats in Arabic), placed in
Allium porrum L. (Liliaceae), ranked as one of the
popular vegetables in the old world. They are
mostly tetraploid (2n = 4x = 32 chromosomes) and
156 DOMESTICATION OF PLANTS IN THE OLD WORLD

seed planted. Ancient Egyptian wall carvings and
drawings, as well as several finds of dried speci-
mens (Täckholm and Drar 1954; Keimer 1984) show
that A. porrum was part of the Egyptian food pro-
duction from the fourth millennium BP onwards.
Outside Egypt, there are few remains. These include
wild or domesticated material from Early and
Middle Bronze Age Jericho (Hopf 1983). Available
texts indicated that leek was grown in Mesopotamia
at the beginning of the fourth millennium BP (Stol
1987a).
Leek is certainly a Mediterranean or south-west
Asian element and its wild progenitor is well iden-
tified (McCollum 1976). The more robust varieties
grown for their thick ‘pseudostem’ (A. porrum L.
senu stricto) and the slender leafy forms (sometimes
referred to as A. kurrat Schweinf.), are all closely
related to, and inter-fertile with, the wild and weedy
tetraploid forms of wild Allium ampeloprasum L.,
which is widely distributed in the Mediterranean
basin.
millennium BP) and later tombs (Täckholm and
Drar 1954; Germer 1989b). Outside of Egypt, there
are few ancient garlic remains. A large number of
carbonized garlic cloves were uncovered from the
fourth millennium BP Tell ed-Der, Iraq (van Zeist
and Vynckler 1984). Also, garlic is well recognized
linguistically in numerous cuneiform records that
indicate garlic was cultivated in Mesopotamia since
at least the beginning of the fourth millennium BP
(Stol 1987a). In the last decade, garlic finds were
reported from some European sites as well.
The wild ancestry of domesticated garlic has not
been definitely established so far. The common ste-
rility of the cultivars makes it difficult to identify of
its wild progenitor. Because of this feature, the gar-
lic crop has not yet been fully cross-tested with its
wild relatives. The cultivars bear the closest mor-

Garlic: Allium sativum

Garlic, Allium sativum L. (Liliaceae), was also an
early constituent of the south-west Asian vegetable
garden. Garlic is a diploid (2n = 2x = 16 chromo-
somes) bulb crop (and see Hirschegger et al. 2010 for
molecular and taxonomic treatment). In contrast to
the leek, garlic is maintained almost totally by veg-
etative propagation (planting of individual cloves).
Up until now, the prevailing view was that most
garlic cultivars are totally sterile, and their flowers
do not set seeds. However, a few recent collections
obtained in central Asia and the Caucasus
(Kamenetsky 2007) was found to be almost seed-
fertile or semi-fertile.
The earliest, excellently preserved, desiccated
garlic remains were discovered in the ‘Cave of the
Treasure’ near Ein Gedi, Israel, dated the ca. 5,970
BP Middle Chalcolithic period (Zaitschek 1961;
M. Broshi pers. comm.). Egypt provides early archae-
ological evidence for this crop as well. Similarly
excellently preserved garlic (Fig. 40) was found in
ca. 1,325 BC Tutankhamun’s tomb (Germer 1989a;
Hepper 1990; de Vartavan et al. 2010). Several well- Fig. 40 Desiccated bulbs of garlic, Tutankhamun’s tomb, New
preserved dry remains of garlic are available Kingdom, Egypt (Photo: H. Barton, with kind permission of Griffith
from other Egyptian eighteenth dynasty (mid-fourth Institute, Ashmolean Museum, Oxford).

phological, as well as molecular resemblance to
Allium longicuspis Regel, known from several scat-
tered locations in east Turkey, Iran, and central Asia.
This taxon has been regarded (Stearn 1978) as the
most probable candidate for the ancestry of domes-
ticated garlic clones. However, similar to the culti-
vars, A. longicuspis inflorescence is bulbiferous and
mostly sterile. As argued by Mathew (1996) this
raises serious doubts whether it could be regarded
as a wild ancestor of domesticated A. sativum.
Alternatively, it might represent cultivated garlic
planted (and abandoned) by nomads. For this rea-
son, other fully fertile close relatives, namely
A. tuncelianum (Kollmann) N. Özhatay, B. Mathew
& Siraneci (native to south-east Turkey), and A.
macrochaetum Boiss. & Hauskn. (native to south-
west Asia) were suggested by Mathew (1996) as
additional candidates for the ancestry of domesti-
cated garlic. Also, A. truncatum (Feinbr.) Kollman &
D. Zohary (native to the Levant) is morphologically
close enough to be suspected. All these three wild
Alliums smell like garlic.
Onion: Allium cepa
Onion, A. cepa L., is the third member of its genus
that was apparently highly appreciated as a garden
crop in south-west Asia and the Mediterranean
basin. It is a diploid (2n = 2x = 16 chromosomes),
cross-pollinated, seed-planted bulb crop. Wall carv-
ings and drawings depicting offerings of onions
and illustrations showing how onions are planted
and watered appear in numerous Egyptian tombs
from the Old Kingdom onwards (Germer 1989b;
Hepper 1990). These include the Unas (ca. 4,420 BP)
and Pepi II (ca. 4,200 BP) pyramids. Well-preserved
onion remains are available from the eighteenth-
dynasty tombs in Egypt (Täckholm and Drar 1954;
Germer 1989b). They are complemented by some-
what later finds, including bulbs placed in mum-
mies (and see detailed survey in Murray 2000b).
Egypt continues to be the main locale for onions in
later periods, like Roman Iron Age Berenike and
Shenshef (Vermeeren and Cappers 2002; Cappers,
2006), Kellis (Thanheiser et al. 2002), Mons
Claudianus (van der Veen 2001a), and Quseir al-
Qadim (van der Veen 2003; van der Veen et al., 2009).
Outside of Egypt, undisputed early archaeobotani-
VEGETABLES AND TUBERS 157
cal records of onions are not yet available. Cuneiform
sources indicate (Stol 1987a) that this vegetable has
been grown in Mesopotamia since the fourth mil-
lennium BP.
Like garlic, wild onion is apparently not a
Mediterranean element (Brewster, 1994). Its progen-
itor has not been convincingly identified. The crop
shows close morphological and molecular affinities
with two central Asiatic wild onions: A. oschaninii
O. Fedtsch., which grows in north Afghanistan,
Tadzhikistan, and neighbouring Uzbekistan, and
A. vavilovii M. Pop. & Vved., native to the Kopet-
dag mountains, Turkmenia (Hanelt 1990). Crosses
between A. oschaninii and domesticated A. cepa gave
rise to sterile F1 hybrids (Hanelt 1985; Hanelt et al.
1992), indicating that this wild onion is not a direct
source from which the vegetable has been derived.
In contrast, A. vavilovii is reported to be almost fully
inter-fertile with the crop (Raamsdonk et al. 1992).
There are doubts whether these vavilovii collections
represent genuine wild material or only feral deriv-
atives of the crop.
Lettuce: Lactuca sativa
Lettuce, Lactuca sativa L., from the Compositae
(Asteraceae) family, is an annual, diploid
(2n = 2x = 18 chromosomes), self- pollinated vege-
table crop. It is definitely a west-Asiatic and
Mediterranean element. The main use of lettuce is
for salads. Its wild ancestor is fully identified
(Zohary 1991; de Vries 1997; Koopman et al. 1998).
The crop shows close morphological resemblance
to a series of six to seven wild lactuca species native
to south-west Asia. Its closest wild relative is weedy
L. serriola L., which abounds in the Mediterranean
basin and south-west Asia. It is an aggressive sum-
mer weed in much of the subtropical and temperate
climate zones of the world.
Wall illustrations of rosettes of a tall, large vegeta-
ble with subulate leaves appear in numerous Old
Kingdom and Middle Kingdom tombs and monu-
ments in Egypt (Keimer 1924; Körber-Grohne 1987,
plates 81–82). Most archaeobotanists agree that
these leafy vegetables depict the garden lettuce.
Unfortunately, the available illustrations are not
yet complemented (in Egypt) by dry remains of let-
tuce leaves or lettuce rosettes, although seeds of
158 DOMESTICATION OF PLANTS IN THE OLD WORLD
lettuce have been reported (Germer 1985). Outside
Egypt, indisputable records of L. sativa cultivation
appear only in Greek and Roman literature (Pliny,
for example, mentions eleven distinct forms).
Chufa or rush nut: Cyperus esculentus
Chufa or rush nut, Cyperus esculentus L. (Cyperaceae),
ranks among the oldest domesticated plants in
Egypt. It is a vegetatively propagated sedge crop.
Its small (1.5–2.5 cm) ellipsoid-globose tubers
appear in large quantities in Egyptian archaeologi-
cal contexts from the pre-dynastic Amratian period
(ca. 5,900–5,650 BP) onward (Täckholm and Drar
1950; Wetterstrom 1993, 1998). Dry remains of
Cyperus esculentus tubers were discovered as foun-
dation deposits in two temples near Semna, Nubia,
ca. 3,500 BP. They indicate that Chufa was already a
crop by the fourth millennium BP (van Zeist 1983).
In addition, chufa tubers are frequently illustrated
in tombs (Darby et al. 1977). Cyperus esculentus rep-
resents a local domestication. The wild and weedy
forms of this sedge (var. aureus Richt.) with its
smaller and more fibrous tubers, grow in the Nile
Valley, in wet places, and is collected or grown by
the local people. While Chufa was an important
food element in ancient Egypt throughout dynastic
times, it did not spread much beyond this area.
There are almost no contemporary records of this
tuber-bearing sedge from other parts of the Old
World.
Cabbage: Brassica oleracea
European cabbage, Brassica oleracea L., from the
Cruciferae (Brassicaceae) family, was a well-
established Mediterranean garden vegetable and
oil crop already during Greek and Roman times
(Körber-Grohne 1987). This extraordinarily poly-
morphic crop includes (U 1935; Hodgkin 1995) leafy
kales, hearting cabbages, broccoli, cauliflower,
Brussels sprouts, kohlrabi, and marrowstem kale
(the latter is an animal feed). In classical times leafy,
cauline forms prevailed. Theophrastus (372–287
BC) mentions three kinds of cabbage: curly leaved,
smooth leaved, and a wild-type. Pliny the elder (AD
23–79) reported on a wider variety of domesticated
forms, which might also have included hearting
types and broccoli-like forms. Types like cauliflower
and Brussels sprouts appeared much later.
The B. oleracea cultivars are closely related to, and
fully (or almost fully) inter-fertile with a varied
aggregate of perennial, herbaceous, largely self-
incompatible, diploid (2n = 2x = 18 chromosomes;
genomic constitution CC), wild cabbage races.
These wild varieties (the B. oleracea ‘cytodeme’)
grow mainly on winter-rain coastal cliffs and simi-
lar maritime habitats along the Atlantic coast of
Europe, the Mediterranean basin, and the Canary
Islands (Snogerup et al. 1990; Gustafsson and
Lannér-Herrera 1997). The following closely related
ten wild oleracea taxa have been recognized in this
complex; many of them are narrow endemics: (i)
Wild B. oleracea subsp. oleracea, native to the Atlantic
coast of Spain, France, UK, and Helgoland; (ii) B.
montana Pourret [= B. oleracea subsp. robertiana (Gay)
Bonnier & Layens] native to the coast of north-east
Spain and the French and the Italian Rivieras; (iii)
B. rupestris Rafin. and (iv) B. villosa Biv. (sensu lato),
both in west Sicily; (v) B. incana Ten. in east Sicily,
the west coast of Italy, and several Adriatic islands;
(vi) B. macrocarpa Guss., endemic to Isole Egadi near
Sicily; (vii) B. insularis Moris in Corsica, Sardinia,
and Tunisia; (viii) B. cretica Lam. in Greece, the
Aegean islands, Crete, and south-west Turkey; (ix)
B. hilarionis Post in Cyprus; and (x) B. bourgeaui
(Webb) Kuntze in the Canary islands.
As to the location of B. oleracea initial domestica-
tion, two possible ancestral populations were sug-
gested, but no conclusive evidence can be offered
currently. Based on morphological similarities,
cytogenetic affinities, and molecular comparisons,
the Atlantic B. oleracea subsp. oleracea seems to be
the closest wild relative of the domesticated varie-
ties of cabbage (Song et al. 1990; Hodgkin 1995;
Gustafsson and Lannér-Herrera 1997). The other
wild vicarious taxa are more distantly related. Yet
some of them (e.g. B. cretica) could also have been
involved in the evolution of the crop. On the other
hand, (Maggioni et al. 2010) gathered linguistic, lit-
erary, and historical information, and found out
that they indicate the domestication of B. oleracea in
the ancient Greek-speaking area of the central and
east Mediterranean. The molecular survey of
Christensen et al. (2011) characterizes diversity and
asseses the genetic structure among seventeen

accessions of kale landraces, cultivars, and wild
populations from Europe. The results of this
research, however, were inconclusive regarding the
ability to indicate a single geographical location for
kale’s centre of origin.
Turnip: Brassica rapa
Turnip, Brassica rapa L. [= B. campestris L.], of the
cabbage family (Cruciferae/Brassicaceae), is a
cross-pollinated vegetable widely cultivated in
China and south-west Asia for its swollen hypocot-
yle (root turnip), its leaves, and for its oil (turnip
rape or oil-seed rape, subsp. rapa). Other groups of
turnip cultivars, such as Chinese cabbage and
Peking cabbage (subsp. pekinensis) have evolved in
east Asia. The wild progenitor of this crop complex
is well identified. Wild forms, and particularly
weedy races of B. rapa are distributed all over the
Mediterranean basin, temperate Europe, Siberia,
and in south-west and east Asia. They are fully
inter-fertile, diploid (2n = 2x = 2 chromosomes),
and chromosomally identical with the cultivars.
Because of the vast distribution of the wild turnip
forms, and the differences between the western and
eastern cultivars, it seems possible that B. rapa was
domesticated independently both in the belt of
Mediterranean agriculture and in China. (Note that
the Chinese finds are not treated in this book).
Archaeological evidence
Similar to many other Old World vegetables, turnip
is poorly represented in archaeological contexts. It
is, however, soundly identified linguistically. Thus,
written sources partly compensate for this defi-
ciency. Seeds of B. rapa were found in several
Neolithic and Bronze Age lake-dwelling sites in
Switzerland and neighbouring countries. They indi-
cate that at these early times, wild turnip was an
element of the weed flora of arable lands in this part
of Europe (Körber-Grohne 1987; Jacomet et al. 1991;
Brombacher 1995; Jacomet 2007). Probably, the
seeds of these weedy forms were collected for their
oil, but B. rapa does not seem to have been a crop at
that time. In contrast, root turnip, grown for its
swollen hypocotyle, was part of the Old World
vegetable garden. The earliest sign of its cultivation
VEGETABLES AND TUBERS 159
is found in the list of plants grown in the garden of
King Merodach Baladan II (722–711 BC) in Babylonia
(Körber-Grohne 1987). From Hellenistic times
onwards, turnip is treated by several Greek and
Roman authors who wrote about agriculture (for
review see Lenz 1859). It is also referred to in the
Jewish Mishna (Feliks 1983). From Roman Iron Age
onward, B. rapa is present in few archaeological
sites across Europe.
Swede, Brassica napus L., is a root and oil-seed
cruciferous crop similar looking and closely related
to the turnip. Yet swede is a tetraploid cultigen
(2n = 4x = 38 chromosomes). It probably originated
under domestication, by hybridization (and chro-
mosome doubling) between the diploid cabbage B.
oleracea and the diploid turnip B. rapa. This root
occurs in archaeological finds came from Roman
Period onward.
Beet: Beta vulgaris
Beet, Beta vulgaris L. (Chenopodiaceae) is a variable,
multi-purpose crop, comprising four principal
groups of cultivars (Körber-Grohn, 1987; Frese
2003). The oldest cultigens are the leaf beet (with
edible leaves) and the garden beetroot (with swol-
len hypocotyle). Both are used as food. Their domes-
tication was followed by the development in Europe
of large-rooted fodder beet and, at the start of the
nineteenth century, by breeding of sugar beet. The
crop is closely related to and fully interfertile with a
group of chromosomally homologous, diploid
(2n = 2x = 18 chromosomes), wild taxa, placed in
section Beta of the genus Beta (Ford-Lloyd and
Williams 1975; Ford-Lloyd 1995). Closest to the
cultigens is a variable aggregate of wild and weedy
beets regarded by these authors as wild races of the
crop complex, namely: (i) B. vulgaris subsp. maritima
(L.) Arcangeli [= B. maritima L.] which is widely dis-
tributed in the Mediterranean basin, south-west
Asia, and the Atlantic coastal belt of Europe; (ii) B.
vulgaris subsp. macrocarpa (Guss.) Thell. [= B. mac-
ropcarpa Guss.], native to the warmer, more arid
parts in the Mediterranean basin; and (iii) B. vulgaris
subsp. adanensis (Pamukç.) Ford-Lloyds & Williams
[= B. adaninsis Pamukç.], which thrives in the
Mediterranean parts of Turkey. These wild beets
constitute the general stock from which the domestic
160 DOMESTICATION OF PLANTS IN THE OLD WORLD
beet could have been derived. Subsp. maritima is
most probably the principal progenitor.
Beet is poorly represented in archaeological con-
texts, and only a few of its remains have been
uncovered (Knörzer 1991; Kreuz 1994–5; Kučan
1995). Moreover, charred fruit remains of domestic
forms are very similar to those of wild forms.
However, beet is linguistically well identified.
Early written sources mentioning this vegetable
compensate, to some extent, for the paucity of the
archaeological evidence. The earliest written docu-
mentation on beet cultivation comes from eighth
century BC Babylonia (Körber-Grohne 1987).
Theophrastus (ca. 372–287 BC) describes this vege-
table as having a thick taproot similar to that of the
radish. Roman and Jewish literary sources indicate
that in the first century BC domestic beet was rep-
resented in the Mediterranean basin by leafy forms
(chards, mangold) and probably also by beetroot
cultivars.
Beet remains are available also from a few sites in
Egypt: a desiccated part of a flowering stem was
found in the thirrd-dynasty Saqqara pyramid at
Memphis (Lauer et al. 1950; Germer 1985). Four
charred beet fruits were retrieved from Neolithic
Aartswoud, north Netherlands (Pals 1984). Since
wild forms of B. vulgaris abound in these places, it is
hard to say whether the finds represent domestic
forms or wild material. The majority of the scanty
remains of charred beet fruits come from Roman
sites in Germany (Knörzer 1991; Stika 1996).
Significantly, at least some of these German sites lie
outside the distribution area of wild beets, and sug-
gests they were domesticated. The appearance of
domesticated beet in Roman Germany corroborates
the contemporary literary sources.
Carrot: Daucus carota
The carrot, Daucus carota L. (Umbelliferae/Apiaceae),
is another vegetable which was grown in the
Mediterranean basin already in classical times.
There are both annual and biannual forms of this
crop. Two main types of domesticated carrots are
recognized: (i) ‘eastern domestic forms’ with dark
purple, anthocyanin-coloured roots, and (ii) ‘west-
ern forms’ with orange, carotene-coloured roots.
Members of the first group, grown mainly in
Afghanistan, Iran, and Turkey, often have branched
roots and are considered the more primitive culti-
vars. Carotene-coloured cultivars characterize car-
rot cultivation in the Mediterranean basin and
temperate Europe. All commercial cultivars belong
to this group.
The available archaeological information on
carrot is deplorably fragmentary. Consequently,
the tracing of its history under domestication
relies, to a large extent, on written sources. The
earliest satisfactory evidence on carrot domestica-
tion comes from Greek and Roman writers
(Körber-Grohne 1987). The plant named by
Dioscorides (60 AD) as ‘staphylinos’ is apparently
D. carota. The telltale dark spot in the centre of the
inflorescence, mentioned by Dioscorides identi-
fies it as such. We are told that the staphylinos’
aromatic root is being used both as food and for
medicine, and that garden-grown plants are tastier
than the wild ones, but less potent medicinally.
The illustration in Dioscorides codex, drawn in
Constantinople ca. AD 500, shows a carrot plant
with a thick, orange-coloured root, indicating that
carotene-containing cultivars already existed at
that time. Carrot mericarps became more abun-
dant in Middle Ages archaeological contexts
throughout Europe.
The domesticated carrot is closely related to, and
fully inter-fertile with, a variable assemblage of
wild and weedy carrot races, all included by taxon-
omists in the D. carota species complex (Heywood
1983). All are diploid (2n = 2x = 18 chromosomes)
and largely cross-pollinated herbs. About a dozen
eco-geographic wild subspecies have been recog-
nized in D. carota, most of them in the Mediterranean
basin and south-western Asia. Weedy forms have
an even wider distribution and extend over most of
temperate Europe. They often grow close (or even
within) carrot cultivation. Under such conditions
spontaneous hybridization between the domesti-
cated carrot and its weedy relatives is quite
frequently.
Celery: Apium graveolens
Celery, Apium graveolens L. (Umbelliferae/Apiaceae),
is presently grown for its fleshy petioles (var. dulce)
or for its swollen hypocotyle (var. rapaceum).

However, less advanced cultivars grown for their
leaves and/or thick aromatic hypocotyle have had
a much longer history under domestication. They
were part of the Mediterranean vegetable garden
assemblage in classical times, appreciated both as
medicinal and garnishing herbs. The wild ancestor
of celery is well identified (2n = 2x = 22 chromo-
somes). Wild forms of A. graveolens, fully inter-
fertile with the cultivars, thrive in marshy places
all over the Mediterranean basin and south-
western Asia. They extend north, along saline coastal
marshes, as far as the British Isles and Denmark.
Celery leaves and inflorescences were part of the
garlands placed in ca. 1,325 BC Tutankhamun
tomb, and with the mummy of Kent (second
dynasty, ca. 1,000 BC) in Egypt (Germer 1985;
Hepper 1990; de Vartavan et al. 2010), and celery
mericarps were discovered in seventh century BC
Heraion, Samos (Kučan 1995). However, since A.
graveolens grows wild in these areas it is hard to
decide whether these remains represent wild or
cultivated forms. The information available from
Switzerland is even more puzzling. Here, celery
mericarps were found in several late Neolithic
lake-shore settlements (e.g. Jacomet 1988, 2004;
Jacomet et al. 1989; Brombacher 1997; Favre 2002;
Brombacher et al. 2005); i.e. places in which A. gra-
veolens does not grow wild today. This suggests
transport and planting by humans. Yet it is hard to
believe that this plant was under domestication in
such early times. Convincing evidence on celery
domestication is available only from classical times
(Körber-Grohne 1987). By Roman times A. graveo-
lens was part of the vegetable garden in the
Mediterranean basin. Moreover, at that time its use
seems to have expanded north of the Alps. Celery
were retrieved from several Roman sites in Italy,
Switzerland, and Germany (Stika 1996; Willerding
1996). Evidence of celery also arrives from ca.
2,700–2,400 cal BP Zinchecra, Libya (van der Veen,
1992a, 1992b).
Parsnip: Pastinaca sativa
Parsnip Pastinaca sativa L. is another umbelliferous
(Umbelliferae/Apiaceae) root vegetable grown in
the Mediterranean basin and in temperate Europe
since classical times (Körber-Grohne 1987; Riggs
VEGETABLES AND TUBERS 161
1995). Its white, thick taproot has been used as a
cooked vegetable and also for animal fodder.
Parsnip is a biennial (2n = 2x = 22 chromosomes)
crop, grown from seed. It is a European and west
Asian element. Wild and weedy forms of this
vegetable (bearing much thinner and less succu-
lent roots) are widely distributed over temperate
Europe and north Turkey. They are also found in
some of the relatively moister parts of the
Mediterranean basin itself. In some areas (espe-
cially in the north) wild-looking P. sativa might be
feral.
Remains of parsnip have been retrieved from
Roman sites in Germany and several other
European countries (Körber-Grohne 1987; Kreuz
1994–5; Stika 1996). Altogether the archaeological
evidence on parsnip is limited. Similar to several
other vegetables, Greek and Roman literary
sources constitute a major source of information
on its early use. There are some difficulties in dis-
tinguishing between parsnip and carrot in classi-
cal writings since both vegetables seem to have
been sometimes called ‘pastinaca’, yet each vegeta-
ble appears to be well under domestication in
Roman times.
Asparagus: Asparagus officinalis
Asparagus officinalis L. (Liliaceae) is a rhizomatous,
perennial vegetable cultivated for its tender, young
shoots (‘spears’). The crop is dioecious, and diploid
(2n = 2x = 2 chromosomes). It is a Mediterranean
and European element. Wild forms of A. officinalis
are widely distributed, particularly in slightly
moist habitats in the Mediterranean basin, in
adjacent parts of south-western Asia, and extend
to numerous parts of temperate Europe. Cytogenet-
ically, the variable wild forms of A. officinalis have
not been studied extensively, but they are known to
contain both diploid (2n = 2x = 2 chromosomes)
and tetraploid (2n = 4x = 40 chromosomes) forms.
The crop is most likely derived from a diploid wild
stock.
Asparagus is a relatively large genus containing
some sixty species. Only A. officinalis is cultivated
as a vegetable, while several other members of
this genus are used as ornamentals. Traditionally,
the young shoots of wild A. officinalis (as well as
162 DOMESTICATION OF PLANTS IN THE OLD WORLD
those of several other Mediterranean Asparagus
species) have been collected from the wild (Ertuğ
2009). This practice survives today in Turkey, and
some other countries. Greek and Roman literary
sources indicate (Körber-Grohne 1987) that by 200
BC, A. officinalis was grown as a garden vegetable
in Greece. It was appreciated for its medicinal
qualities and for its delicious taste. Evidence on
its use in Greece is soon followed by substantial
written information on its cultivation in Rome.
Evidence of older asparagus cultivation is almost
nonexistent.
 C H A PTER 8
Condiments
Like the vegetables, several condiments indigenous
to west Asia and the Mediterranean basin entered
domestication rather early. The tradition of their
planting and usage is well documented in classical
Greek, Roman, and Jewish sources. Only few
remains of this group of crops have been discovered
in archaeological contexts. Many condiments were
used not only for their seeds but also for their green
leaves, which like vegetables rarely survived
archaeologically. Because of the rarity of condiment
finds, our knowledge of the beginning of their
domestication is insufficient. Many of the available
records come from Egypt, where the combination
between the use of a wide range of plants and desic-
cation in desert environment, leads to the survival
of a wide range of material (see Murray 2000a, b, c,
as an excellent review of the Egyptian evidence).
Frequent finds of several condiments and herbs
of Mediterranean origin, e.g. parsley, dill, celery,
and lemon balm (Melissa officinalis), start to appear
in lake-shore settlements in Switzerland already in
Late Neolithic, ca. 6,000 cal. BP. It is unknown yet
whether these species were locally grown or were
imported from regions closest to the Mediterranean
coast (Jacomet 2007).
Interestingly, the remains of four condiments
(coriander, cumin, dill, and black cumin) were dis-
covered in several Hellenistic and Roman sites in
Egypt, like the remote desert quarry of Mons
Claudianus, Egypt (Van der Veen 1998, 2001b), the
Red Sea harbour Berenike (Vermeeren and Cappers
2002; Cappers 2006), and Quseir al-Qadim (Van der
Veen 2003, 2009). These finds indicate that during
the Roman Period they were considered essen-
tial for Romans’ diet. As Livarda and van der Veen
(2008) noted, the availability of condiments
increased significantly with the Roman Empire, and
to some extent characterizes this period. The fol-
lowing sections review the available evidence on
what seem to have been the first condiments of the
Old World.
Coriander: Coriandrum sativum
Coriander, Coriandrum sativum L. (Umbelliferae/
Apiaceae), is an annual, largely diploid (2n = 2x = 22
chromosomes), self-pollinated crop, grown for its
aromatic seeds and foliage (Diederichsen 1996;
Small 1997). It is one of the earliest condiment crops
of the Old World (Gabrielian and Zohary 2004), and
it was known in classical times. Plant remains and
linguistic evidence indicate that its use started much
earlier.
Today C. sativum occurs spontaneously over wide
areas of Old World agriculture and it is hard to
define exactly where this plant is wild and where it
only recently established itself as a weed or as a
naturalized element. In south-west Asia and
Armenia, C. sativum seems to grow wild in rocky
openings, in oak park-forests, oak scrub, and steppe-
like formations. Probably, these territories were the
source of both the cultivated forms and the weedy
races.
Archaeological evidence
Fifteen desiccated coriander mericarps were retrieved
from Pre-Pottery Neolithic B Nahal Hemar Cave
(Kislev 1988), and eleven such mericarps from ca.
8,000–7,500 BP PPNC Atlit-Yam (Kislev et al. 2004),
Israel. If they are not intrusive, they represent the
earliest archaeological find of this condiment. Early
163
164 DOMESTICATION OF PLANTS IN THE OLD WORLD
finds of coriander came also from several Late
Neolithic sites: Tell Hammam et-Turkman, Syria
(van Zeist 2003), Kapitan Dimitrievo, Bulgaria
(Marinova 2006), and Poduri, Rumania (Cârciumaru
and Monah, 1985; Monah and Monah 2008). In the
latter, the ca. 6,650–6,350 cal. BP Eneolithic stratum,
coriander fruits were found mixed with those of
Sambucus nigra, inside a painted amphora and
wooded cask. A large sample (about half a litre, the
main item in several baskets) of round, small, meri-
carps of C. sativum was found in ca. 1,325 BC
Tutankhamun’s tomb, Egypt (Renfrew 1973; Germer
1989a; Hepper 1990; de Vartavan et al. 2010). Because
coriander does not grow wild in Egypt, its presence
in this tomb, as well as in several other Egyptian
sites, suggest that this condiment was under culti-
vation at that time. Coriander remains were also
discovered in the fourth millennium BP Tell ed-Der,
Syria (van Zeist and Vynckler 1984), and in several
Bronze Age sites: Sitagroi (Renfrew 1973) and
Akrotiri, Thera (Sarpaki 1992a), Greece, Ulu Burun
shipwreck, Turkey (Haldane 1993; Ward 2003), and
Umm el-Mara, Syria (Schwartz et al. 2000). A single
half-fruit of C. sativum was found in Late Bronze
Age Apliki, Cyprus (Renfrew 1973). Coriander was
also known from ca. 1,200–500 B.C. Iron Age Deir
Alla, Jordan (Neef 1989), and late Assyrian Nimrud
(Helbaek 1966b). It was also found in several Iron
Age and Roman sites in Egypt: Berenike and
Shenshef (Vermeeren and Cappers 2002), Kellis
(Thanheiser et al. 2002), Mons Claudianus (Van der
Veen 2001b), and Quseir al-Qadim (Van der Veen
2003, 2009).
Cumin and dill: Cuminum cyminum and
Anethum graveolens
Two additional aromatic umbellifers (Umbelliferae/
Apiaceae)—cumin, Cuminum cyminum L., and dill,
Anethum graveolens L.—were part of Greek and
Roman agriculture. Wild forms of cumin are
unknown in south-west Asia, but occur in central
Asia. Wild and weedy types of dill are widespread
in the Mediterranean basin and in West Asia.
Earliest cumin seeds were uncovered in ca. 8,000–
7,500 BP PPNC Atlit-Yam, Israel (Kislev et al. 2004),
and later in the fourth millennium BP Tell ed-Der,
Syria (van Zeist and Vynckler 1984). Later, it was
found as well in Iron Age Deir Alla, Jordan (Neef
1989). In Egypt, a few seeds of cumin were found in
New Kingdom Deir el Medineh (Germer 1985), and
a basket full of cumin was included in the burial of
Kha, architect of Amenophis III, eighteenth dynasty
(Wilson 1988). Cumin was found also in several
later Iron Age and Roman Age sites in Egypt:
Berenike and Shenshef (Vermeeren and Cappers
2002; Cappers 2006), Kellis (Thanheiser et al. 2002),
and Mons Claudianus (Van der Veen 2001b),
Egypt.
Earliest remains of dill was found in several late
Neolithic lake-shore settlements in Switzerland
(Jacomet and Brombacher 2005; Jacomet 2007).
Several twigs of dill were found in the tomb of
Amenophis II, eighteenth dynasty, ca 1,550–1,292
BC (Germer 1985) and this condiment is also
reported from seventh century BC Heraion, Samos
(Kučan 1995). Later it was found in ca 700–400 BC
Early Garamantian Zinchecra, Libya (van der Veen
1992a), in Roman Iron Age Berenike and Shenshef
(Vermeeren and Cappers 2002; Cappers 2006), and
Iron Age Mons Claudianus (Van der Veen 2001b),
Egypt.
This evidence suggests that already in the
fourth millennium BP, both cumin and dill were
under cultivation. However, it is difficult to con-
clude where and when domestication of this con-
diment began.
Black cumin: Nigella sativa
Black cumin, Nigella sativa L., of the buttercup
family Ranunculaceae, has been another tradi-
tional condiment of the Old World during classi-
cal times (Small 1997). Its black seeds were
extensively used to flavour food. Archaeological
information on the early cultivation of black
cumin is scanty, but seeds of N. sativa were
reported from ancient Egypt, including ca. 1,325
BC Tutankhamun tomb (Germer 1989a; Hepper
1990; de Vartavan et al. 2010), and Iron Age Mons
Claudianus (Van der Veen 2001b). They also
appear in Iron Age Deir Alla, Jordan (Neef 1989).
Linguistic considerations suggest that this plant
was also grown in ancient Mesopotamia
(Thompson 1949). Small (1997) identified the use
of Black Cumin in Sanskrit, in India.

Wild and weedy forms of N. sativa grow in grain
and fallow fields and occur in south Turkey, Syria,
north Iraq, as well as in several adjacent territories.
This indicates that south-west Asia could have been
the place of domestication of this condiment.
Saffron: Crocus sativus
Saffron, Crocus sativus L., of the Iris family Iridaceae,
is another member of the early group of condiments.
The long, scarlet lobes harvested from its flowers
were highly valued for flavouring foods and for col-
ouring them golden-yellow. Saffron was extensively
grown in south-west Asia and the Mediterranean
basin in classical times and it maintained its role as a
very, expensive condiment until the beginning of the
second century. Recently, production became increas-
ingly impractical because the collection of the styles
requires a vast amount of labour. Today saffron culti-
vation is declining and Crocus survives only as a relic
CONDIMENTS 165
crop, despite its position as one of the most expen-
sive spices (Winterhalter and Straubinger 2000).
No remains of C. sativus have been traced in
archaeological excavations and indeed the chances
for the preservation of the delicate stigmas seem
fairly low. However, Minoan (ca 1645 BC) pottery
and frescoes (e.g. the renowned ‘The Gathering of
the Crocus’ from Akrotiri, Santorini) depict Crocus
flowers with long exserted, red, stigmatic branches
(Mathew 1977; Negbi 1999), and these probably
represent C. sativus. Also linguistic evidence sug-
gests that saffron was brought into cultivation
before classical times.
The cultivated saffron is an autumn-flowering,
sterile triploid (3n = 3x = 24 chromosomes) main-
tained by vegetative propagation. It is morphologi-
cally closest to the Greek species C. cartwrightianus
Herbert, and is possibly a clonal selection from this
wild diploid (2n = 2x = 16) species (Mathew 1977,
1999; Rashed-Mohassel 2006).
 C H A PTER 9
Dye crops
A group of dye plants, native to south-western Asia
and the Mediterranean basin had probably entered
domestication already before classical times. They
are amply documented in Greek, Roman, and
Jewish literary sources. Woad (Isatis tinctoria L.),
dyer’s rocket (Reseda luteola L.), Madder (Rubia tinc-
torum L.), Safflower (Carthamus tinctorius L.), and
indigo (Indigofera tinctoria L.), were characteristic
dye crops of Old World agriculture, and their dye
stuffs were extensively used to colour textiles and
leather. They continued to be highly appreciated
crops until the end of the nineteenth century. Then,
the invention of low-priced synthetic dyes caused
their collapse. In a matter of a few decades, the tra-
ditional dye crops of the Old World became com-
mercially redundant and their cultivation died out.
Today they represent agricultural drop-outs. Most
of the cultivars are extinct. Only a few forms sur-
vive in remote farming communities, in botanical
gardens, or in the hands of hobbyists.
To-date, only few remnants of dye crops have been
uncovered in archaeological excavations. This rarity
apparently stems from the use of perishable vegeta-
tive parts which are rarely preserved, rather than
seeds and fruits, for dye production. Consequently,
we know very little about the beginnings of dye-plant
domestication. Early literary sources, and some anal-
yses of dye elements colouring ancient Egyptian tex-
tiles (Eastwood 1984; Germer 1992) make it clear that
in the fourth millennium BP, dye crops were used
extensively to colour textiles and leather (Krochmal
and Krochmal 1974). Some twisted and dyed flax fib-
ers were recently reported in Upper Palaeolithic, ca.
30,000 year-old, Dzudzuana Cave, Georgia (Kvavadze
et al. 2009), but raised severe criticism of their validity
(Bergfjord et al. 2010; Kvavadze et al. 2010).
166
Woad: Isatis tinctoria
Woad, a member of the mustard family (Cruciferae/
Brassicaceae), was a principal dye crop in Europe
and in south-west Asia. Until the end of the nine-
teenth century, woad was cultivated extensively (by
seed planting) for its blue indigo dye, which was
widely used to colour textiles. The pigment was
extracted from the leaves of the vegetative parts of
the plants. To obtain the dye, the leaves were dried,
powdered, and fermented in alkaline medium—a
tedious and very smelly process (Guarino et al.
2000). Today, natural indigo is a commercial rarity.
In the textile industry it was mostly replaced by
synthetic aniline dyes.
Woad—Isatis tinctoria L.—is usually a biennial,
cross pollinating, tetraploid (2n = 4x = 28 chromo-
somes, Darlington and Wyle 1955) plant, with char-
acteristic pale yellow flowers and winged fruits. It
is native to south-west Asia and the Aegean area,
but it also extends to temperate Europe. Some of the
present weedy populations of I. tinctoria (particu-
larly those of temperate Europe) may be feral deriv-
atives of the former crop. Recent research on the
genetic variation and population structure of
Eurasian collections revealed that the origin of the
domesticated crop might have been western and
central Asia (Spataro et al. 2007).
Similar to madder (see below), classical literary
sources show that in the third millennium BP, woad
was extensively used in south-west Asia, in the
Mediterranean basin, and in temperate Europe
(Körber-Grohne 1987). In Neolithic Çatalhöyük
(Fairbairn et al. 2002), few seeds of cf. Isatis sp. were
recovered, although their identification and use
is uncertain. Zech-Matterne and Leconte (2010)

compiled recent archaeobotanical and other evi-
dence from north-eastern Europe and raised the
possibility of local cultivation of woad in the fifth to
the fourth century BC in northern Gaul. In addition,
indigo dye (indigotin) was detected in several blue-
coloured ancient Egyptian textiles found in eight-
eenth dynasty (1,370 BC) Tell el Amarna (Eastwood
1984; Germer 1992). While it is impossible to distin-
guish chemically between indigo extracted from
Isatis or from Indigofera (p. 168), only the first seems
to have been present in the Mediterranean basin in
classical times. Indigofera arrived later.
Dyer’s rocket: Reseda luteola
Dyer’s rocket or weld Reseda luteola L. (Resedaceae)
is the source of a very intense flavon-type yellow
pigment extracted from its roots, stems, foliage, and
flowers. This plant was extensively cultivated (by
seed planting) until the beginning of the twentieth
century when the natural dye was replaced by
cheaper synthetic yellow dyes.
Dyer’s rocket is an annual or a biennial plant
with erect stems and with yellow inflorescences.
The plant is apparently indigenous to the east
Mediterranean basin and to south-west Asia and
became naturalized far beyond its original native
geographic range.
Historical sources report that R. luteola was har-
vested in the middle of its flowering season, dried,
and sold to textile dyers for dye extraction. It is one
of the better natural sources of yellow colour, valu-
able because it can be used with woad to produce
green, an uncommon colour in dyes (Shewry et al.
1997).
Classical literary sources show that dyer’s
rocket was domesticated and extensively used
already in the Old World in the third millennium
BP (Körber-Grohne 1987). Like woad and madder,
it was probably taken into cultivation even earlier.
From Late Bronze Age on, it was found in various
sites in west Europe: Belgium, Netherlands,
Germany, France, and UK (see Schultze-Motel
1968–1994; Kroll 1995–2000, as well as the online
databases http://www.cuminum.de/archaeobot-
any/database/; http://www.archaeobotany.de/
database.html
DYE CROPS 167
Madder: Rubia tinctorum
Madder, Rubia tinctorum L. (Rubiaceae) was, until
the end of the nineteenth century, one of the most
appreciated dye plants of south-west Asia and
Europe. It was extensively grown (by vegetative
propagation) all over this area for its rhizomes from
which a brilliant red pigment (alizarin) was
extracted. Alizarin was widely used to colour linen,
wool, cotton, and leather. The dye becomes fixed to
the textile fibres only after their treatment with a
mordant (alum salts). Furthermore, mordanting
with different metals produces different hues (alu-
minium alum induces dark red coloration; iron
alum results in brown-red colour; chromium alum
produces a red-violet colour).
Pliny (Natural History XXXV xlii) described the
technique of madder dyeing in Egypt eloquently:
‘They employ a very remarkable process of colouring tex-
tiles. After rubbing the cloth, which is white at first, they
saturate it not with the dye—but with mordants that are
calculated to absorb the colour. This done, the textiles still
unchanged in their appearance are plunged into a caul-
dron of boiling dye and are removed the next minute fully
coloured. It is a remarkable fact, too, that although the
cauldron contains a uniform dye, the material taken out is
of various colours—according to the nature of the mor-
dants that have been respectively added to the cloth. These
colours will never wash out.’
Madder is a perennial herb with characteristic
whorls of lanceolate leaves and climbing or stagger-
ing scabrous stems that can reach the length of
1–2 m. Its wild forms are native to south-west
and central Asia (F. Ehrendorfer, pers. comm.).
Spontaneous populations of R. tinctorum thrive also
in the Mediterranean basin, mostly in hedgerows,
waste ground, and margins of cultivated fields.
Possibly, they represent feral derivatives of former
cultivation.
It is still impossible to conclude where and when
madder was taken into domestication or when the
alizarin colouring technology was developed.
Chemical analysis detected madder dye in red-
coloured flax textiles retrieved from eighteenth
dynasty (1,370 BC) Tell el Amarna, Egypt (Eastwood
1984; Germer 1992). Various Greek, Roman, and
Jewish literary sources show that at the end of the
third millennium BP, R. tinctorum was extensively
168 DOMESTICATION OF PLANTS IN THE OLD WORLD
cultivated in Persia, Anatolia, and the Mediterranean
basin (Körber-Grohne 1987). Evidently, domestica-
tion of madder started long before these times.
Synthetic alizarin was invented in Germany in
1869. In a matter of only a few decades, the low-cost
synthetic equivalent replaced the much more expen-
sive natural dyestuff and caused a collapse of mad-
der cultivation.
True indigo: Indigofera tinctoria
In India, as well as in several other tropical and sub-
tropical parts of south Asia and Africa south of the
Sahara, the traditional source for indigo blue dye
was not woad but several members of the genus
Indigofera, from the Papilionaceae/Fabaceae family
of the Leguminosae. Prior to the invention of syn-
thetic indigo, a common cultigen was I. tinctoria L.
Wild forms of this dye crop occur in India, and this
plant was probably domesticated (propagated from
seeds) in this subcontinent (Lemmens and Wessel-
Riemens 1991). Several other species of Indigofera,
among them I. arrecta Hochst., native to eastern and
southern Africa, were also extensively cultivated. In
the Indian subcontinent, the pigment extracted
from the leaves of Indigofera crops was commer-
cially so valuable, that it was referred to as the ‘king
of dye-stuffs’. Sound documentation on its cultiva-
tion is available only after the Arab conquest of
Egypt.
Safflower: Carthamus tinctorius
Safflower, Carthamus tinctorius L. (Compositae/
Asteraceae), was a traditional dye plant of the Old
World. Its yellow-red flowers were used extensively
to dye textiles and to colour foods. Carthamus is
basically an Irano-Turanian genus comprising thir-
teen to fourteen annual species of thistles (Hanelt
1963). With the recent development of synthetic
dyes, the importance of this crop declined consider-
ably. Since the 1950s, safflower acquired a new use
and has emerged as a modern industrial oil crop
(Knowles and Ashri 1995).
The crop is taxonomically closely related to a
group of 3–4 diploid (2n = 2x = 24 chromosomes)
wild species, placed in section Carthamus, native to
south-west and central Asia (Hanelt 1963; Knowles
and Ashri 1995). C. tinctorius was cross-tested with
two wild and weedy members of this section,
namely: (i) C. persicus Willd.[=C. flavescens Willd.]
that occurs in the arid and semi-arid regions of the
Levant countries, the Syrian desert, south Anatolia
and upper Mesopotamia; and (ii) C. oxyacanthus M.
Bieb., that grows in the Trans-Caucasus, Iran,
Afghanistan, and Central Asia. Both species were
found to be inter-fertile and chromosomally homol-
ogous with the crop. C. tinctorius is probably also
inter-fertile with a third diploid wild member of this
section, namely C. gypsicola Iljin, a geographically
much more restricted species, which grows on gyp-
sum-rich substrates in the southern Trans-Caucasus,
the southern Caspian basin, and in Middle Asia. In
contrast, members of other taxonomic sections in
Carthamus are far more distant, and hard or impos-
sible to cross with the domesticated safflower. These
three taxa, therefore, can be the progenitor stock of
the domesticated plant. Chapman et al. (2010, 2007)
conducted recent DNA sequence studies in an
attempt to trace the variation and the origin of
domesticated safflower. They suggest that C. tincto-
rius is most likely derived from wild C. palaestinus.
However, C. palaestinus is apparently a synonym for
C. persicus (Greuter 2006–2009; Flann 2009).
Marinova and Riehl (2009) reviewed recently the
rich find of safflower crop in archaeological assem-
blages. Carthamus sp. occurs in the region in several
Neolithic and Chalcolithic sites. C. tinctorius achenes
appear first in ca. 5,000 cal. BP Early Bronze Age
sites in Syria, from where it spread later to Turkey,
Bulgaria, and Serbia, to Egypt, the Aegean, and
south-eastern Europe. They suggest that the use of
safflower for oil production started almost from the
beginning of its cultivation.
In Egypt, seeds of safflower were found in Late
Bronze Age, ca. 1,325 BC Tutankhamun tomb
(Germer 1989a; Hepper 1990; de Vartavan et al.
2010). Well-preserved garlands composed of C. tinc-
torius flowers were found adorning eighteenth-
dynasty (middle fourth millennium BP) mummies
in Egypt (Keimer 1924; Täckholm 1976). Chemical
analysis of Egyptian textiles dated from the twelfth
dynasty showed safflower to be one of the dyes
used (Darby et al. 1977).This dye plant is also
well identified in early Mesopotamian cuneiform
records.
 C H A PTER 1 0
Plant remains in representative
archaeological sites
This chapter summarizes the information on plant
remains retrieved from representative Neolithic
and Bronze Age sites in west Asia, Europe, and
north Africa. It presents a selected list which was
compiled in order to answer the question, if one has
to sketch the origin and the early spread of domes-
ticated plants in the Old World, what would be the
minimum number of archaeological locations that
could provide an adequate account on the present
state of our knowledge?
The information is arranged country by country.
Representative sites had to be selected from coun-
tries which are still very poorly studied, as well as
from areas that had a long tradition of archaeologi-
cal excavation. In order to attain this goal, numerous
well-analyzed locations in intensively researched
countries were, not included in this chapter, while
poorer sites in less thoroughly surveyed countries
do appear on the list. Altogether some 191 sites (or
groups of sites) were chosen to represent the archae-
ological evidence as it stands today.
Most sites are radiocarbon-dated, some were den-
dro-dated; for others we have only estimated dates.
Many of the radiocarbon dates are AMS dating, but
certainly not all of them. When available, both
uncalibrated and calibrated radiocarbon dates were
given, using OxCal Program v3.10. In order to com-
pare between sites and strata (since we did not
intend to cover the broad issue of dating in full) all
dates were rounded to the nearest fifty years.
Traditional dating has been given where radiocar-
bon analyses were not available. If needed, the
reader is advised to seek accurate dates and the type
of material dated in the cited references. More dates
are available in various web-based databases and
online journals, such as ‘C14 radiocarbon CONTEXT’
(http://context-database.uni-koeln.de/index.php),
‘Radiocarbon’ (http://www.radiocarbon.org), or
‘Archaeometry’ (http://www.wiley.com/bw/jour-
nal.asp?ref=0003-813x). A general chronological
chart (Appendix B) for the different regions is given
on pp. 198–199.
This book presents archaeological data from sites
across vast geographical area and long chronologi-
cal periods. In the literature, one can find different
time periods and geographical boundaries for each
region covered. It is beyond the scope of this book
to create one sytem that will cover it all. Therefore,
hereafter we rely on the terminology used in the
archaeobotanical literature cited. However, to make
the first stages of agriculture easier to understand,
we used one chronological framework for south-
west Asia, as Table 9 presents.
For information on the geographic location of the
sites, consult Appendix A (Maps 21–24, pp. 194–196)
and the general references given for each country.
Iran
(General reference: Miller 1991; Charles 2007)
1. Ali Kosh, Deh Luran Plain, Khuzistan
(Helbaek 1969). (i) Middle PPNB period, Bus
Mordeh phase (8450±90 to 8000±50 uncal BP = ca.
9,600–8,750 cal BP). Rich remains: wild-type einkorn
wheat (few); einkorn wheat (rare); emmer wheat
(prevailing); brittle two-rowed barley (frequent);
naked barley (few). Wild: Linum (rare); Prosopis
farcta, Pistacia atlantica, and Capparis spinosa (rare).
(ii) Middle PPNB period, Ali Kosh phase (ca. 8,290
uncal BP = ca. 9,400–9,250 cal BP). Rich remains:
169
170 DOMESTICATION OF PLANTS IN THE OLD WORLD
Table 9 Chronology of cultural entities for southern Levantine Pre-Pottery
Neolithic Periods (after Goring-Morris and Belfer-Cohen 2011).
Period Entity/phase
Late Epipaleolithic Final Natufian
Pre-Pottery Neolithic A PPNA
Pre-Pottery Neolithic B Early PPNB
Middle PPNB
Late PPNB
Final PPNB/PPNC
Early Pottery Neolithic Yarmukian
brittle einkorn wheat (rare); einkorn wheat (rare);
emmer wheat (prevailing); two-rowed barley (few);
naked barley (rare); lentil (rare). Wild: Linum (rare);
Avena (few); Prosopis farcta (frequent); Pistacia atlan-
tica and Capparis spinosa (rare). (iii) Pottery Neolithic,
Mohammed Jaffar phase (ca. 7,550 uncal BP = ca.
8,400–8,350 cal BP). Rich remains: emmer wheat
(frequent); two-rowed barley (prevailing); six-
rowed barley (rare); lentil (rare). Wild: Linum (rare);
Avena (frequent); Pistacia atlantica and Capparis
spinosa (rare).
2. Tepe Sabz, Deh Luran Plain, Khuzistan
(Helbaek 1969). Chalcolithic, Sabz phase and
Khazineh phase (ca. 7,450–6,950 uncal BP = ca.
8,350–7,750 cal BP). Scarce remains: wild-type
einkorn wheat (few); einkorn wheat (few); emmer
wheat (few); free-threshing wheat (frequent); two-
rowed barley (prevailing); six-rowed barley (fre-
quent); naked barley (few); lentil (frequent); grass
pea (few); flax (frequent); Wild: Triticum boeoticum
(few); Avena (few); Capparis (few); Prosopis (fre-
quent); Amygdalus (few); Pistacia (few); legumes
(frequent); other wild grasses (few).
3. Tepe Hasanlu, Solduz Valley (Tosi 1975).
Pottery Neolithic. (i) Hajji Firuz Tepe, periods
X–VIII (ca. eighth to the sixth millennia BP).
Unspecified quantities: emmer wheat (frequent);
two-rowed barley (prevailing). (ii) Pisdeli Tepe,
period VIII (ca. sixth millennium BP). Rich remains:
emmer wheat (frequent); free-threshing wheat (fre-
quent); two and six-rowed barley (prevailing).
4. Tepe Yahya and adjacent sites, Dowlatabad
Plain 200 km south of Kerman (Costantini and
Costantini-Biasini 1985). Pottery Neolithic, periods
VII and VI (late eighth millennium BP to the sev-
enth millennium BP). Rich remains, unspecified
Calibrated 14C years BP
12,500–11,750
12,175–11,000
10,950–10,300
10,150–9,725
9,400–8,900
9,050–8,450
8,400–7,700
quantities. The main crops are einkorn wheat,
emmer wheat, two and six-rowed barley. Free-
threshing wheat and a rounded, small-grained
‘sphaerococcum’ like barley, and broomcorn millet,
are also present.
Iraq
(General references: Braidwood 1960; Renfrew 1984;
Miller 1991; Charles 2007)
1. Jarmo, Kurdistan (Helbaek 1959a, 1960;
Braidwood 1960). PPNB (ca. 8,350 uncal BP = ca.
9,450–9,300 cal BP). Scarce remains (not yet fully
published, both imprints and carbonized remains):
brittle and non-brittle einkorn wheat (rare); brittle
and non-brittle emmer wheat (frequent); brittle
two-rowed barley (frequent); non-brittle two-rowed
barley (few); lentil (rare); pea (rare). Wild: Pistacia;
Prosopis; Aegilops; Lathyrus.
2. Tell es-Sawwan, Samarra (Helbaek 1964b).
Pottery Neolithic (ca. 7,300–7,000 BP). Numerous
remains: einkorn wheat (few); emmer wheat (fre-
quent); free-threshing wheat (few); two-rowed and
six-rowed barley (prevailing); naked barley (fre-
quent); flax (rare). Wild: Prosopis farcta (frequent);
Capparis spinosa (frequent).
3. Yarym Tepe, northern Iraq (Bakhteyev and
Yanushevich 1980). (i) Yarym Tepe I, Pottery
Neolithic (eighth millennium BP). Numerous
remains: emmer wheat (few); free-threshing wheat
(few); spelt wheat? (rare); hulled two-rowed barley
(rare); hulled six-rowed barley (prevailing); naked
barley (few). (ii) Yarym Tepe II Chalcolithic (sixth
millennium BP). Rich remains: emmer wheat (few);
free-threshing wheat (rare); spelt wheat? (few);
hulled six-rowed barley (prevailing); naked barley

(few). No other plants mentioned. (Note that the
identification of spelt wheat is based on kernel
morphology only. It cannot be regarded as
definite.)
4. Choga Mami, Mandali (Helbaek 1972). (i)
Samarra phase, Pottery Neolithic (second half of
eighth millennium BP). Rich remains: brittle einkorn
wheat (rare); einkorn wheat (few); emmer wheat
(frequent); free-threshing wheat (frequent); brittle
two-rowed barley (frequent); two-rowed and six-
rowed barley (rare); naked barley (frequent); lentil
(frequent); pea (few); flax (frequent). Wild: Avena
sp. (few); Lolium sp. (prevailing); other grasses (fre-
quent); Pistacia atlantica (few). (ii) Post-Samarra
phase (ca. 6950 uncal BP = ca. 7800–7750 cal BP).
Numerous remains: brittle einkorn and einkorn
wheat (rare); emmer wheat (few); free-threshing
wheat (rare); six-rowed barley (few); naked barley
(rare); lentil (few); flax (rare). Wild: Lolium sp. (pre-
vailing); Pistacia atlantica (few).
Turkey
(General references: Nesbitt and Samuel 1996b;
Miller 1991)
1. Çayönü, near Ergani, Diyarbakir Province
(Van Zeist 1972, 143–66; Van Zeist and De Roller
1991–2, 2003). PPNB (ca. 9,150–8,650 uncal BP = ca.
10,250–9,550 cal BP). Rich charred remains (includ-
ing chaff): emmer wheat (frequent), the kernels are
still narrow resembling those of wild forms, but
hundreds of spikelet forks and of glume bases, all
have the domestic-type disarticulation scars; pea
(frequent), including very few seeds with rough
and with smooth seed coat; lentil (frequent); bitter
vetch (very frequent); chickpea (rare); flax (few).
Wild: Hordeum spontaneum (rare); Secale sp. (rare);
Vicia sp. (frequent); Lathyrus cicera/sativus (rare);
Linum sp. (few); Ficus carica (rare); Vitis vinifera
(few); Pistacia (frequent); Amygdalus sp. (few);
Quercus sp. (rare); Celtis sp. (few); several weeds.
2. Can Hasan III, Konya plain (Hillmam 1972,
1978). PPNB (ca. 8,400–7,700 uncal BP = ca. 9,450–
8,450 cal BP). Numerous remains, both charred
grains and chaff: einkorn wheat (few), both wild
and domestic forms; emmer wheat (frequent); free-
threshing Triticum aestivum-like wheat (prevailing),
represented by numerous kernels and rachis frag-
PLANT REMAINS IN REPRESENTATIVE ARCHAEOLOGICAL SITES 171
ments; two-rowed hulled barley (frequent); several
plump rye kernels; lentil (few); bitter vetch (fre-
quent). Wild: Vicia sativa-like (frequent); Juglans
regia (rare); Celtis tournefortii (frequent); Vitis vinifera
subsp. sylvestris (rare); Prunus sp. and Crataegus sp.
(rare); Lithospermum arvense (very common).
3. Hacilar, Konya Plain (Helbaek 1970). Pottery
Neolithic (ca. 7,350–7,000 uncal BP = ca. 8,200–7,800
cal BP). Rich remains: brittle einkorn wheat (fre-
quent); einkorn wheat (frequent); emmer wheat
(few); free-threshing wheat (few); two-rowed bar-
ley (rare); six-rowed barley (few); naked barley
(prevailing); lentil (few); bitter vetch (few). Wild:
Pisum sp. (frequent); Aegilops umbellulata (rare);
Pistacia atlantica (rare); Celtis australis (frequent);
Capparis spinosa (few); Malus sp. (rare); Amygdalus
sp. (rare).
4. Çatalhöyük East, Konya Plain (Helbaek 1964a;
Fairbairn et al. 2002, 2005, 2007). (i) PPNB, layers
Pre-XIIA–D (8,240±55 uncal BP to 8,090±55 uncal
BP = ca. 9,350–8,950 cal BP). Rich remains: einkorn
wheat (frequent); emmer wheat (prevailing); free-
threshing wheat (frequent); naked six-rowed barley
(rare); rye (few); lentil (prevailing); pea (few); bitter
vetch (frequent). Wild einkorn, probable weed
(few). Wild fruits and seeds: Celtis (frequent);
Pistacia (frequent); Quercus acorns (frequent);
Amygdalus sp. (frequent); Malus sp. (rare); Pisum ela-
tius (rare); Ficus (rare); Prunus (rare); Rubus (rare);
Capparis (rare); Rhus coriaria (rare). Tubers of
Bolboschoenus maritimus (frequent). (ii) Neolithic,
Level XII–VI (8,090±55 uncal BP to 7,521±77 uncal
BP = ca. 8,950–8,350 cal BP). Rich remains including
storage contexts, most assemblage equivalent to
Aceramic Neolithic, except: naked six-rowed barley
(frequent). Wild fruits and seeds: Celtis (predomi-
nant); Juniperus sp. (rare); wild crucifers in storage
contexts Capsella bursa-pastoris and Descurainia (fre-
quent); Helianthemum (rare); Taeniatherum caput-
medusae (rare); Eremopyrum (rare). Tubers of
Bolboschoenus maritimus (few).
5. Aşikli Höyük, Central Anatolia (van Zeist and
de Roller 1995). PPNB (8,920±50 to 8,515±40 BP =
ca. 10,200–9,500 cal BP). Numerous charred remains
(including chaff): einkorn wheat (few); emmer
wheat (prevailing); free-threshing wheat (few);
hulled two-rowed barley (frequent); naked barley
(few); lentil (few); pea (rare); bitter vetch (frequent).
172 DOMESTICATION OF PLANTS IN THE OLD WORLD
Wild: Celtis tournefortii (frequent); Pistacia sp. (few);
Amygdalus sp. (rare).
6. Erbaba, Beysehir, south-central Anatolia (van
Zeist and Buitenhuis 1983). Pottery Neolithic (ca.
5,800–5,400 uncal BC = ca. 8,550–8,150 cal BP). Rich
remains: einkorn wheat (frequent); emmer wheat
and free-threshing wheat (prevailing); spelt wheat?
(rare); hulled two-rowed barley (frequent); naked
barley (frequent); lentil (frequent); pea (co-prevail-
ing); bitter vetch (frequent). Wild: Lathyrus cf. cicera;
Triticum boeoticum and several herbaceous plants
and weeds.
7. Girikihaciyan, near Diyarbakir (van Zeist
1979–80). Halafian (Chalcolithic, ca. 6,950–6,450
uncal BP = ca. 7,800–7,350 cal BP). Numerous
remains: einkorn wheat (rare); emmer wheat (pre-
vailing); free-threshing wheat (rare); two-rowed
barley (few); lentil (frequent); chickpea (few); bitter
vetch (frequent); flax (few). Wild: Pistacia atlantica
(rare); Amygdalus sp. and Crataegus sp. (rare); Vicia
sp. (few); Lolium sp. (few).
8. Troy and adjacent Kumtepe, the Troad, north-
west Turkey (Riehl 1997). (i) Kumtepe, Neolithic
Period (ca. 4,805–4,370 cal BC = ca. 6,750–6,300 cal
BP). Few charred remains: einkorn wheat, grains
and chaff (rare); emmer wheat, chaff (rare); six-
rowed hulled barley, grains and chaff (rare); lentil
(frequent); bitter vetch (prevailing); flax (rare); fig
(frequent). (ii) Kumtepe, Chalcolithic Period (ca.
3,370–2,910 cal BC = ca. 5,300–4,850 cal BP). Few
charred remains: einkorn wheat, chaff and grains
(frequent); emmer wheat, chaff, and grains (prevail-
ing); six-rowed hulled barley, grains, and chaff
(few); lentil (rare); faba bean (rare); bitter vetch
(few); Lathyrus cicera/sativus (rare); fig (few); grape-
vine (few). (iii) Troy, Early Bronze Age (ca. 2,900–
2,200 BC = ca. 4,850–4,200 cal BP). Few charred
remains: einkorn wheat, chaff, and grains (fre-
quent); emmer wheat, grains, and chaff (prevail-
ing); six-rowed hulled barley, grains, and chaff
(few); lentil (rare); bitter vetch (few); Lathyrus
cicera/sativus (rare); olive (rare); fig (few); grapevine
(few). (iv) Troy, Middle Bronze Age (ca. 2,200–1,700
BC = ca. 4,150–3,650 cal BP). Very rich charred
remains: einkorn wheat, chaff, and grains (few);
emmer wheat, chaff, and grains (frequent); free-
threshing hexaploid wheat, chaff, and grains (few);
six-rowed hulled barley, grains, and chaff (fre-
quent); lentil (rare); pea (frequent); bitter vetch
(few); Lathyrus cicera/sativus (rare); faba bean (rare);
flax (prevailing); fig (few); grapevine (few); gold of
pleasure (few). (v) Troy, Late Bronze Age (ca. 1,700–
1,200 BC = ca. 3,650–3,200 cal BP). Rich charred
remains: einkorn wheat, grains, and chaff (few);
emmer wheat, chaff, and grains (few); free-thresh-
ing hexaploid wheat, chaff, and grains (few); six-
rowed hulled barley, grains, and chaff (frequent);
broomcorn millet (rare); lentil (rare); chickpea (pre-
vailing); bitter vetch (frequent); Lathyrus
cicera/sativus (rare); faba chickpea (rare); flax (pre-
vailing); fig (rare); grapevine (rare); gold of pleasure
(rare).
Syria
(General reference: Miller 1991; Willcox 2007;
Willcox et al. 2008, 2009)
1. Tell Abu Hureyra, Northern Syria, Euphrates
valley (Hillman 1975, 1989, 2000a, 2000b; De
Moulins 1997, 2000; Hillman et al. 2001). (i) Epi-
Palaeolithic (ca. 12,700–11,100 cal BP). Rich remains
of charred seeds belonging to a wide range of spe-
cies, all conforming to wild forms in their morphol-
ogy, and no chaff remains: narrow grains of wild
two-grained einkorn wheat (rare) and wild rye
(prevailing); small seeded lentil (few); bitter vetch
(rare); Pistacia sp. (frequent); Celtis sp. (rare);
Capparis spinosa (rare); large numbers of Polygonum
and Bolboschoenus maritimus nutlets; seeds of
numerous grasses (Stipa prevailing), and herbs
(Brassicaceae prevailing). (ii) PPNB culture, four
excavation trenches of various strata (from 8640±100
uncal BP, from trench B phase 2 = ca. 9,800–9,500 cal
BP, to 802±100 uncal. BP from trench G phase 6 = ca.
9,050–8,700 cal BP). Numerous, various charred
remains from the four trenches in moderate quanti-
ties in each sample: wild and domesticated einkorn,
grain and chaff; hexaploid or tetraploid wheat; wild
and domesticated barley; wild and domesticated
rye; lentil; chickpea; large vetch; Prosopis farcta;
small legumes (frequent); Capparis spinosa; grape
pips (few). Many other grass and herb seeds and
uncharred Boraginaceae.
2. Tell Mureybit, northern Syria, Euphrates val-
ley (van Zeist and Casparie 1968; van Zeist and
Bakker-Heeres 1986). Epi-Palaeolithic (ca. 10,200–

9,900 uncal BP = ca. 11,800–11,300 cal BP). Rich
remains of charred seeds—all conforming in their
morphology to wild forms: narrow grains of wild
two-grained einkorn wheat and of wild rye (pre-
vailing); glumes of rye (few); narrow, wild-type
grains of barley (few); small-seeded lentil (few); bit-
ter vetch (rare); wild flax (rare); Pistacia atlantica
(rare); numerous grass and herb seeds.
3. Jerf el Ahmar, northern Syria, Euphrates val-
ley (Willcox 2000; Willcox et al. 2008, 2009). (i) PPNA
early phase (ca. 9,800–9,500 uncal BP = ca. 11,500–
11,000 cal BP). Rich charred remains, chaff indicat-
ing that all cereals conform to wild forms in
their morphology: two-grained wild einkorn, T.
boeoticum/T. urartu (rare); wild rye, Secale sp. (pre-
vailing); wild barley (prevailing); Lens sp.
(frequent); Vicia ervilia (rare); wild small grasses (H.
murinum type, Stipa sp., Taeniatherum) (frequent).
Pistacia sp. (frequent); Amygdalus sp. (frequent);
Polygonum/Rumex (rare).1 (ii) PPNA late phase (ca.
9500–9300 uncal BP = ca. 11,000–10,600 cal BP).
Similar to the early phase, only wild barley increases
and wild rye declines.
4. Djade el Mughara, northern Syria, Euphrates
valley (Willcox et al. 2008). PPNA (ca. 9,300–9,000
uncal BP = ca. 10,700–10,400 cal BP). Similar to Jerf
el Ahmar, except that wild-type founder species,
emmer, single-grained einkorn, chickpea, and faba
bean appear for the first time in northern Syria.
5. Tell Aswad, south-east of Damascus (van Zeist
and Bakker-Heeres 1985). (i) Early PPNB (ca. 9,300–
9,000 uncal BP = ca. 10,500–10,200 cal BP). The dates
of these levels (considered initially to be PPNA) have
been recently revised based on AMS dating of emmer
grains (Willcox 2005, Table 1) retrieved from the early
excavations studied by van Zeist, as well as more
recent excavations. Numerous remains of charred
seeds: plump domestic-type grains of emmer wheat
(frequent); either wild or domestic two-rowed barley
(few); small-seeded lentil (rare); pea (few); Pistacia sp.
(prevailing); Ficus carica (frequent); Capparis spinosa
(rare); Amygdalus sp. shells (rare); Trigonella sp. (fre-
quent); seeds of numerous grass and herb species. (ii)
Middle PPNB (ca. 10,200–9,550 cal BP). Numerous
remains of charred seeds: plump, domestic-type
grains of emmer wheat (frequent); free-threshing
1
Willcox argues for pre-domestic agriculture.
PLANT REMAINS IN REPRESENTATIVE ARCHAEOLOGICAL SITES 173
wheat (few); either wild or domestic two-rowed bar-
ley (rare); naked barley (rare); small-seeded lentil
(rare); pea (few); early finds of flax (frequent). Wild:
Pistacia sp. (frequent); Ficus carica (frequent); Capparis
spinosa (rare); Vitis vinifera (rare); Trigonella sp. (fre-
quent); Cyperus sp., Carex sp., Lolium sp., and Phalaris
sp. (frequent); numerous grass and herb seeds.
Israel and Jordan
(General references: Miller 1991; Neef 1998)
1. Ohalo II, Sea of Galilee (Kislev et al. 1992;
Simchoni 1998; Weiss 2002, 2009; Weiss et al. 2004,
2008). Upper Palaeolithic (ca. 23,000 cal BP). Rich
charred remains, ca. 140 wild taxa: wild emmer
wheat, T. dicoccoides (frequent); wild barley, H. spon-
taneum (prevailing); wild lentil, L. orientalis (rare);
wild pea, Pisum sativum subsp. humile (rare). Nine
edible wild fruits: wild almond, Amygdalus commu-
nis (rare); Crataegus aaronia/azarolus (few); Nitraria
schoberi (frequent); Olea europaea (rare); Pistacia
atlantica (rare); Pyrus syriaca (rare); Quercus sp. (few);
Vitis vinifera (rare); Zizyphus spina-christi (rare).
Grains of several wild grasses such as Aegilops,
Avena, and Bromus; seeds of several wild herbs.
2. Netiv Hagdud and Gilgal, Lower Jordan Valley
(Kislev 1997; Hartmann 2006; Weiss et al. 2006). Pre-
Pottery Neolithic A (ca. 10,000–9,400 uncal BP = ca.
11,700–10,550 cal BP). Rich remains of charred wild
plant remains: dicoccoides-type emmer wheat (fre-
quent); spontaneum-type barley (prevailing, storage);
other wild Horduem species (few); Avena sterilis (fre-
quent, storage); A. wiestii (rare); Aegilops sp. (rare);
Alopecurus utriculatus (few); Lens sp. (frequent); other
legume species (rare); Ficus carica (very common);
Vitis vinifera (rare); Pistacia atlantica (few); Quercus ith-
aburensis (rare); Amygdalus communis (rare).
3. Nahal Zehora II , Mount Carmel , Israel
(Kislev and Hartmann forthcoming). Pottery
Neolithic (ca. 7,300–7,100 uncal BP = ca. 8,150–
7,850 cal BP). Rich charred remains: emmer
wheat (prevailing); parvicoccum-type wheat (fre-
quent); barley (few); lentil (few); faba bean (rare);
flax (rare). Fruit trees: fig (rare); olive (rare);
Pistacia atlantica/palaestina (rare). Wild/weedy
grasses, including Lolium temulentum (rare).
174 DOMESTICATION OF PLANTS IN THE OLD WORLD
4. Jericho, Lower Jordan Valley (Hopf 1983). (i)
Middle Pre-Pottery Neolithic B (ca. 9,900–9,550 cal
BP). Rich remains: einkorn wheat and emmer wheat
(prevailing); two-rowed barley (frequent); lentil
(frequent); pea (frequent); Vicia faba-type (few);
chickpea (rare); flax (an imprint of a single capsule).
(ii) Ceramic Neolithic (first half of the seventh mil-
lennium BP). Rich remains: einkorn wheat and
emmer wheat (frequent); two-rowed barley (pre-
vailing). Wild: some grass and herb seeds. (iii)
Chalcolithic (early sixth millennium BP). Numerous
remains: wheat (rare); two-rowed barley (prevail-
ing). Wild: Ficus carica (rare). (iv) Early Bronze Age
(ca. 3,500–3,300 BPBP). Rich remains: einkorn wheat
(frequent); emmer wheat (prevailing); free-thresh-
ing wheat (frequent); two-rowed and six-rowed
barley (frequent); lentil (frequent); chickpea (few);
flax, a capsule and seed (rare); grapevine (frequent,
both pips and berries); date palm (rare); fig (rare).
Wild: Allium cf. ampeloprasum (few); Pistacia atlantica
(rare); numerous grass and herb seeds. (v) Middle
Bronze Age. Rich remains: emmer wheat (frequent);
free-threshing wheat (frequent); six-rowed barley
(prevailing); lentil (rare); pomegranate (rare). Wild:
Allium cf. ampeloprasum (rare); some grass and herb
seeds.
5. Atlit Yam, Carmel Coast (Kislev et al. 2004; and
pers. comm.). Pre-Pottery Neolithic C (ca. 8,000–
7,500 cal BPBP). Rich charred and waterlogged
remains in a well: emmer wheat (prevailing); naked
(T. parvicoccum) wheat (frequent); barley (few); lentil
(few); chickpea (rare); flax. Fruits and nuts: Ficus
carica (frequent); Vitis sylvestris; Amygdalus
communis/korschinskii; Pistacia atlantica; P. lentiscus;
P. Palaestina; Phoenix theophrasti. Large group of wild
and weedy plants: Papaver setigerum; Cuminum cymi-
num; Pinus halepensis; Lolium temulentum; Phalaris
paradoxa; Mercurialis annua; Malva parviflora; Rumex
pulcher; Daucus carota; Coriandrum sativum; Murinum
sylibum; Capparis spinosa; Rubus sanguineus.
6. Tuleilat Ghassul, Lower Jordan Valley and
Shiqmim, Negev (Zohary and Spiegel-Roy 1975;
Kislev 1987). Late Chalcolithic (ca. 6,800–5,800 cal
BP). Rich remains: emmer wheat (few); two and six-
rowed barley (frequent); lentil (few); olive (frequent,
in Ghassul only); date palm (few, in Ghassul only).
7. Tell es-Sa’idiyeh, Jordan (Cartwright 2003;
and pers. comm.). Early Bronze Age (ca. 4,900 BP).
Very rich charred remains, in a palace’s kitchen:
wheat; barley; lentil; chickpea; faba bean-type;
grapes; figs; pomegranate; olive; Wild: Cratagus sp.;
Ziziphus spina-christi; Pistasia sp.; Quercus sp.;
Capparis spinosa, flower buds.
8. Bab edh-Dhra, south-east of Dead Sea
(McCreery 1979). Early Bronze Age. Rich remains:
emmer wheat (probably also einkorn and free-
threshing wheat) (few); two-rowed barley (few);
six-rowed barley (prevailing); lentil (few); chickpea
(frequent); linseed (few); olive (frequent); grapevine
(frequent, whole berries and pips); fig (frequent);
almond (rare). Wild: Pistacia atlantica (few); Prunus
insititia-like (rare); Lolium temulentum and herb
seeds. Textiles made of flax fibres.
Egypt
(General references: Täckholm 1976; Germer 1985;
Wetterstrom 1993, 1998; Murray 2000a, 2000b, 2000c;
de Vartavan et al. 2010).
1. Nabta Playa, Western Desert, south Egypt
(Wasylikowa 1997; Wasylikowa et al. 1997;
Wasylikowa and Dahlberg 1999). Site E-75–6. Pre-
agriculture site (8,095±12 to 7,950±90 uncal BP = ca.
9,250–8,650 cal BP). Site E-75–6. Rich charred
remains of wild plants: Sorghum bicolor subsp.
arundinaceum (very frequent); Echinochloa colona
(very frequent); Panicum turgidum (frequent); sev-
eral other grass species (few); Arnebia sp. (frequent);
Capparis cf. decidua (few); Schouwia purpurea (very
frequent); Boerhavia sp. (frequent); several legumes
(very frequent); Zizyphus cf. spina-christi (frequent);
tubers of Cyperus cf. rotundus (frequent).
2. Farafra Oasis, Hidden Valley, Western Desert
(Barakat and Fahmy 1999; Fahmy 2001). Neolithic
period (6,028±150 to 5,163±374 uncal BC = ca. 7,050–
5,450 cal BP). Few charred remains of wild plants,
grasses predominant: Sorghum sp. (prevailing);
Setaria verticillata (frequent); Panicum repens (fre-
quent); Echninochloa colona (frequent); Brachiaria sp.
(few); Cenchrus (rare); Digitaria sp. (rare); Citrullus
colocynthis (rare).
3. Maadi, near Cairo (van Zeist and de Roller
1993; van Zeist et al. 2003). Late Neolithic, Predynastic
(4,680±70 to 4,900±70 uncal BP = ca. 5,650–5,450 cal
BP). Rich charred and desiccated remains: emmer
wheat, grain and chaff (prevailing); two-rowed and

six-rowed barley (frequent); lentil (rare); linseed
(rare); Ficus sp. (rare); Cucumis cf. melo (rare).
4. Fayum, Kom K site (Caton-Thompson and
Gardner 1934; Wetterstrom 1993; Wendrich and
Cappers 2005). Early Neolithic (6,095±250 to
6,391±180 uncal BP = ca. 7,500–6,650 cal BP). Large
quantities of parched and of charred grains in
underground silos: mainly emmer wheat, two-
rowed, and six-rowed barley; also flax textile.
5. Merimde, Beni Salâme, western Nile Delta. (i)
Neolithic (6,130±110 BP to 5,260±90 uncal BP = ca.
7,150–5,950 cal BP) (Wetterstrom 1993). Rich
remains: emmer wheat (prevailing); free-threshing
wheat? (rare); hulled six-rowed barley (frequent);
lentil (few); pea (few); flax (rare). Wild: Lolium and
several other weeds, sedges, and legumes (fre-
quent). (ii) Middle Neolithic (Helbaek 1955). Rich
remains: emmer wheat (prevailing); free-threshing
wheat (rare and later absent); hulled six-rowed bar-
ley (frequent).
6. Naqada (Nagada) area, Upper Egypt
(Wetterstrom 1993). (i) Site KH3, Nagada I (ca.
5,900–5,650 BP). Numerous remains: emmer wheat
(prevailing); hulled six-rowed barley (frequent);
pea (rare); flax (rare). Wild: Citrullus colocynthis
(rare); Zizyphus spina-christi (rare). (ii) South Town,
Nagada II (ca. 5,650–5,300 BP). Numerous remains:
emmer wheat (prevailing); hulled six-rowed barley
(frequent); pea (rare); bitter vetch (rare); flax (few).
Wild: Citrullus colocynthis (few); Zizyphus spina-
christi (rare); some weeds.
7. Hierakonpolis, Upper Egypt (Fahmy 1995,
2003, 2005; Fahmy et al. 2008). (i) Site HK11C, trash
mound, Naqada IIAB (ca. 5,700–5,600 BP). Rich
remains: emmer wheat (prevailing); free-threshing
wheat (rare); hulled six-rowed barley (frequent);
hulled two-rowed barley (rare); naked barley (rare);
Cucumis melo (frequent); flax (frequent). Wild:
Balanites aegyptiaca (rare); Ceruana pratensis (fre-
quent); Fimbristylis bisumbellata (rare); Setaria verti-
cillata (rare). Weeds: Lolium temulentum (few);
Phalaris minor (frequent); as well as other field
weeds. (ii) Site HK43, Workers’ Cemetery, Naqada
IIBC (ca. 5,450–5,300 BP). Numerous remains:
emmer wheat (prevailing); free-threshing wheat
(rare); hulled six rowed barley (frequent); hulled
two-rowed barley (rare); naked barley (rare);
Cucumis melo (rare). Wild: large quantities of culms
PLANT REMAINS IN REPRESENTATIVE ARCHAEOLOGICAL SITES 175
of Juncus, which have been used for making of mat-
ting; Anethum graveolens (frequent); Balanites aegyp-
tiaca (rare); Ceruana pratensis (rare); Cyperus
esculentus (rare); Acacia (frequent); Ziziphus spina-
christi (frequent); also flax textile.
8. Kom el-Hisn, west part of the Nile delta
(Moens and Wetterstrom 1988). Old Kingdom
deposits (ca. 4,700–4,250 BP). Numerous remains:
emmer wheat (prevailing); barley (frequent); pea
(rare); flax (rare); clover seeds, probably Trifolium
alexandrinum (co-prevailing). Wild: Lolium temulen-
tum and Phalaris paradoxa (frequent); numerous
herbs, reeds, and sedges.
9. Saqqara, Memphis. (i) Djoser (Zoser) pyra-
mid, thirdrd dynasty, ca. 4,630–4,611 BP (Lauer et
al. 1950). Rich remains (desiccated material):
emmer wheat and hulled six-rowed barley (the
bulk of the find); lentil (rare); sycamore fig
(numerous dry sycons). Apparently imported:
fragments of grapevine raisins (few); Mimusops
schimperi (rare); Juniperus oxycedrus (few); Wild:
Lolium temulentum (the commonest weed); Phalaris
paradoxa (rare); Vicia sativa (few); V. lutea (few); V.
narbonensis (rare); Lathyrus aphaca (few); L. mar-
moratus (rare); L. hirsutus (few); Scorpiurus muri-
cata (few); Trigonella hamosa (rare); Medicago
hispida (rare); Rumex dentatus (few); Anthemis
pseudocotula (rare); Beta vulgaris (rare); Zizyphus
spina-christi (numerous); Acacia nilotica (rare);
Balanites aegyptiaca (rare). (ii) Queen Icheti’s tomb,
sixth dynasty, ca. 4,550 BP (Helbaek 1953). Rich
remains (mummified material): emmer wheat
prevailing.
10. Tutankhamun tomb, Valley of the Kings, near
Thebes (Germer 1989a; Hepper 1990; de Vartavan
et al. 2010) New Kingdom, nineteenth dynasty (ca.
1,325 BC). An extremely rich find of desiccated,
excellently preserved plants and plant products:
emmer wheat (main element in the model granary);
six-rowed barley (large quantities, in baskets, includ-
ing germinated seeds in ‘Osiris bed’); lentil (few);
chickpea (few); fenugreek (in several baskets, mixed
with coriander); flax (linen, bow strings, few seeds);
olives (leaves and twigs in garlands, jars of olive oil);
grape wine (shrivelled berries and their pips, wine
amphoras with records on production sites and
dates); sycamore fig (fruits and timber); date palm
(fruits in basket, ropes); almond (basket with fruits);
176 DOMESTICATION OF PLANTS IN THE OLD WORLD
watermelon (numerous seeds in basket); garlic
(numerous bulbs, some with leaves); black cumin
(numerous seeds); coriander seeds (main item in
several baskets); safflower (numerous achens, tex-
tiles dyed with safflower pigment). Remains of sev-
eral ornamental plants and plants collected from the
wild (Grewia sp.; Ziziphus spina-christi; Juniperus sp.).
Timber (in furniture and implements) of numerous
local and foreign trees, including imported cedar
(Cedrus libani) and ebony (Dalbergia melanoxylon).
Libya
(General reference: van der Veen 1995, 2006)
1. Zinchecra, Hilltop settlement, Fazzan, south-
ern Libya (van der Veen 1992a, 1992b). Early
Garamantian (ca. 2,700–2,400 cal BP). Rich charred
remains: emmer wheat (frequent); bread wheat
(rare); six-rowed barley (frequent); date palm (fre-
quent); grapevine (rare); fig. Herbs: celery (Apium
graveolens); purslane (Portulaca oleracea); dill
(Anethum graveolens); fennel (Foeniculum vulgare).
Wild: Citrullus colocynthis; Rhus tripartita.
2. Jerma (Garama), Wadi settlement, Fazzan,
southern Libya (Pelling 2008). (i) ca. twenty-fourth
to the twenty-first century BP, Scarce remains:
emmer wheat (rare); barley (rare); fig (rare); grape-
vine (rare); date palm (rare). Wild: Pennesitum glau-
cum. (ii) ca. 2,200–1,600 cal BP, rich charred remains:
durum wheat; bread wheat; barley; sorghum
(Sorghum bicolor); olive; grapevine; fig; date palm;
almond; pomegranate; cotton (Gossypium sp.). Wild:
Pennisetum glaucum.
Morocco
Kaf That el-Ghar, NW Morocco (Ballouche and
Marinval 2003). Early Neolithic, Cardial culture (ca.
6,500–5,500 cal BP). Few charred remains: emmer
wheat (prevailing); bread/durum wheat (few);
naked barley (very rare); broad bean (rare).
Caucasia and Transcaucasia
(General reference: Wasylikowa et al. 1991)
1. Chokh, Dagestan (Lisitsina 1984). Neolithic
(end of eighth millennium BP to the seventh millen-
nium BP). Unspecified quantities: einkorn wheat;
emmer wheat; free-threshing wheat; hulled barley;
naked barley; several wild grasses and trees.
2. Arukhlo 1 and Arukhlo 2, Bolnisskiy region,
Georgia (Januševič 1984; Lisitsina 1984; Schultze-
Motel 1988). Neolithic, Aruchlo 1 Period (7,135±70
uncal BP to 6,365±140 uncal BP = ca. 8,000–7,150 cal
BP). Unspecified quantities (except for the wheats),
mostly imprints: einkorn wheat (few); emmer wheat
(numerous); free-threshing wheat (prevailing); spelt
wheat (rare); two-rowed and six-rowed hulled bar-
ley; naked barley; broomcorn millet; lentil; pea; bit-
ter vetch.
3. Imiris-Gora, Marnsulskij region, Georgia
(Lisitsina and Prishchepenko 1977; Schultze-Motel
1988a). Eneolithic (ca. 6,300±12 uncal BP = ca. 7,400–
7,000 cal BP). Imprints, unspecified quantities:
emmer wheat; free-threshing wheat; spelt wheat(?);
six-rowed hulled and naked barley; broomcorn mil-
let(?); Avena sp.
4. Aratashen and Aknashen, Ararat Valley,
Armenia (Hovsepyan and Willcox 2008). Neolithic
culture (ca. 7,035±69 uncal BP to ca. 6,350±70 uncal
BP = ca. 7,950–7,150 cal BP). Numerous charred
remains and imprints: einkorn wheat (rare); emmer
wheat (few); free-threshing wheat (frequent); hulled
barley (rare); naked six-row barley (frequent); small-
seeded lentil, seeds and pods (frequent); bitter vetch
(few). Wild: Alyssum desertorum (prevailing, capsule
imprints); Camelina microcarpa (frequent, capsule
imprints); Vitis sylvestris; Celtis sp.; Elaeagnus sp.;
Buglossoides arvensis (frequent); Rumex cf. crispus;
Bolboschoenus maritimus.
Central Asia
(General reference: Harris and Gosden 1996)
1. Jeitun (Djeitun), south Turkmenistan (Charles
and Hillman 1992; Harris et al. 1993; Harris et al.
1996). Neolithic Jeitun culture (7,270±100 to 7,100±90
uncal BP = ca. 8,200–7,850 cal BP). Numerous
charred remains: einkorn wheat (prevailing); emmer
wheat (few); six-row barley (few). Rare trace of free-
threshing wheat (?).
2. Anau, south Turkmenistan (Miller 1999; Miller,
2003). Chalcolithic (Anau North, ca. 6,500–5,000 cal
BP) and Bronze Age (Anau South, ca. 5,000–3,700
cal BP). Few charred remains: bread wheat (fre-
quent); six-row barley (prevailing); pea, in Bronze

Age only (rare); grape, in Bronze Age only (rare).
Wild: Celtis sp.; Capparis sp.; as well as genera from
several families notably the Chenopodiaceae,
Fabaceae, and Poaceae.
3. Shortugai, Oxus River Valley, Afghanistan
(Willcox 1991). Mature Harappan culture (2,245±100
uncal B.C. = 2,350–2,100 cal BP) Numerous remains:
free-threshing wheat (frequent); barley (prevailing);
broomcorn millet (frequent); lentil (few); pea (few);
flax (rare); grapevine (frequent, probably culti-
vated). Wild: Pistacia vera; Eleagnus angustifolia;
Amygdalus sp.; Prosopis sp.; several grasses, includ-
ing Aegilops tauschii and herbs.
Cyprus
(General references: Hansen 1991a, 1994; Kroll 1991;
Miller 1991)
1. Kissonerga-Mylouthkia and Shillourokam
bos, Cyprus (Willcox 2000; Murray, 2003). Cypro-
Early PPNB (9,310±80 to 8,670±80 uncal BP = ca.
10,650–9,550 cal BP). Numerous charred and impres-
sion remains: einkorn wheat (?), grains, and chaff
(rare); emmer wheat (?), grains, and chaff (rare);
hulled barley (?) (frequent). Most other taxa were
not identified to species level and probably wild-
growing: legumes, fruits, and numerous wild/
weeds taxa. The domesticated status of the cereals
requires further support.
2. Cap Andreas-Kastros (van Zeist 1981).
Aceramic Neolithic (6,140±200 to 7,775±125 uncal
BP = 8,700–6,800 cal BP). Numerous remains:
einkorn wheat (frequent); emmer wheat (prevail-
ing); six-rowed barley (frequent); lentil (frequent);
pea (?) (rare); flax (rare). Wild: Vicia faba/narbonensis
(rare); Olea europaea (rare); Pistacia atlantica/terebin-
thus (rare); Ficus carica (rare); Lolium sp. (frequent);
Vicia sp. (few); Malva sp. (few).
3. Khirokitia (Waines and Stanley Price 1975–
1977; Hansen 1991a, 1994). Aceramic Neolithic (ca.
7500 uncal BP = ca. 8,400–8,200 cal BP). Rich remains
(including chaff): einkorn wheat and emmer wheat
(prevailing); six-rowed barley (few); lentil (frequent);
pea (few). Wild: Lathyrus cf. sativus (rare); Vicia faba/
narbonensis (rare); Ficus carica (frequent); Olea euro-
paea (rare); Prunus insititia (few); Pistacia sp. (rare).
4. Dhali Agridhi, Idalion (Stewart 1974). Aceramic
Neolithic (7,990±80 to 7,290±465 uncal BP = ca.
PLANT REMAINS IN REPRESENTATIVE ARCHAEOLOGICAL SITES 177
9,000–7,600 cal BP). Numerous remains: emmer
wheat (few); free-threshing wheat (rare); two- and
six-rowed hulled barley (frequent); naked barley
(few); lentil (frequent); pea (rare). Wild: Olea euro-
paea (frequent); Vitis vinifera (few); Ficus carica (fre-
quent); Amygdalus sp. (rare); Pistacia sp. (rare);
Anchusa sp. (frequent).
5. Hala Sultan Tekke, near Larnaca (Hjelmqvist
1979a). Late Bronze Age (ca. 3,200 uncal BP = ca.
3,450–3,350 cal BP). Rich remains: emmer wheat
(few imprints); free-threshing wheat (rare); hulled
and naked barley (prevailing-imprints); lentil (rare);
olive (few); grapevine pips (frequent); fig pips (fre-
quent); pomegranate (rare); citron (rare). Wild:
Pistacia terebinthus (rare); Capparis spinosa (rare);
Zizyphus spina-christi (rare); Lupinus albus (rare);
Chrozophora verbascifolia (few); Onopordon cf. illyri-
cum (few).
Greece
(General references: Renfrew 1979; Kroll 1991;
Valamoti and Kotsakis 2007).
1. Franchthi Cave, Argolis (Hansen 1991a, 1992).
(i) Upper Palaeolithic (ca. 13,000–9,000 uncal BP =
ca. 15,500–10,150 cal BP). Scarce remains: Wild: brit-
tle two-rowed barley (few); Lens sp. (few) Avena sp.
(rare); Vicia sp. (few); Pistacia sp. (few); Amygdalus
sp. (few); Lithospermum, Alkanna, Anchusa (fre-
quent). (ii) Mesolithic (ca. 9,300–8,000 uncal BP = ca.
10,600–8,750 cal BP). Scarce remains: Wild: brittle
two-rowed barley (few); Lens sp. (few); Pisum sp.
(few); Avena sp. (few); Pistacia sp., Amygdalus sp.
Pyrus sp. (few); Lithospermum, Alkanna and Anchusa
(frequent). (iii) Early Neolithic (7,794±140 to
6,940±90 uncal BP = ca. 8,800–7,700 cal BP). Scarce
remains: emmer wheat (few); two-rowed barley
(few). Wild: Pistacia sp. (few); Amygdalus sp. (rare);
Lithospermum, Alkanna, Anchusa, Lens sp., Vicia sp.
(few). (iv) Middle Neolithic (ca. 7,000–6,300 uncal
BP = 7,950–7,200 cal BP). Scarce remains: einkorn
wheat (rare); emmer wheat (frequent); two-rowed
barley (frequent); lentil (few); bitter vetch (rare).
Wild: Pistacia sp. and Amygdalus sp. and Pyrus sp.
(rare); Lithospermum, Alkanna, Anchusa (rare). (v)
Late Neolithic (ca. 6,300–4,800 uncal BP = 7,300–
5,450 cal BP). Scarce remains: einkorn wheat (fre-
quent); two-rowed barley (frequent); lentil (few);
178 DOMESTICATION OF PLANTS IN THE OLD WORLD
bitter vetch (rare). Wild: Pistacia sp. Pyrus sp.,
Amygdalus sp., Vitis sylvestris (rare); Lithospermum,
Alkanna, and Anchusa (rare).
2. Sesklo (including Argissa and Otzaki),
Thessaly (Hopf 1962; Kroll 1981a). (i) Aceramic and
Early Neolithic (7,755±97 to 7,483±72 uncal BP = ca.
8,650–8,200 cal BP). Numerous remains: einkorn
wheat (frequent); emmer wheat (prevailing); six-
rowed barley (frequent); lentil (frequent); bitter
vetch (few). Wild: Ficus carica (frequent); Vitis vinif-
era (rare); Sambucus ebulus (rare); Pistacia atlantica
(frequent); Lithospermum arvense (frequent); Avena
sp. (frequent). (ii) Proto- and pre-Sesklo phases
(eighth millennium BP). Rich remains: einkorn and
emmer wheat (prevailing); free-threshing wheat
(rare); two-rowed barley (few); six-rowed hulled
and naked barley (frequent); lentil (frequent); pea
(few); bitter vetch (few); chickpea (rare); flax (few).
Wild: Ficus carica (frequent); Sambucus ebulus (rare);
Pistacia atlantica (rare); Lithospermum arvense (fre-
quent); Avena sp. (few). (iii) Sesklo phase (seventh
millennium BP). Numerous remains: einkorn wheat
(frequent); emmer wheat (prevailing); six-rowed
barley (few); broomcorn millet (rare); lentil (fre-
quent); pea (rare); bitter vetch (rare); flax (rare).
Wild: Ficus carica (frequent); Vitis vinifera (rare);
Pistacia atlantica (rare); Lithospermum arvense (fre-
quent); Avena sp. (few); (iv) Dimini phase, Late
Neolithic (early to middle of sixth millennium BP).
Numerous remains; einkorn wheat (frequent);
emmer wheat (prevailing); free-threshing wheat
(rare); six-rowed barley (frequent); lentil (few); faba-
like bean (rare); bitter vetch (rare); flax (rare). Wild:
Ficus carica (frequent); Vitis vinifera (frequent);
Amygdalus sp. (rare); Lithospermum arvernse (rare);
Avena sp. (few). (v) Rachmani phase, Late Neolithic
(late sixth to early third millennium BP). Rich
remains: einkorn wheat, emmer wheat, six-rowed
barley, and lentil (equally frequent); free-threshing
wheat (rare); naked barley (rare); pea (rare); grass
pea (few); flax (rare). Wild: Ficus carica (frequent);
Vitis vinifera (rare); Quercus acorns (rare); Amygdalus
sp. (rare); Camelina sativa (a single seed); Lithospermum
arvense (few); Avena sp. (few).
3. Nea Nikomedeia, Macedonia (van Zeist and
Bottema 1971). Early Neolithic (ca. 7,470 uncal BP =
ca. 8,400–8,100 cal BP). Rich remains: einkorn wheat
(frequent); emmer wheat (prevailing); naked barley
(frequent); lentil (frequent); pea (frequent); bitter
vetch (frequent). Wild: Quercus acorns (few); Cornus
mas (rare); Prunus cf. spinosa (rare).
4. Toumba Balomenou, Chaeronia, Boiotia
(Sarpaki 1995). Numerous remains, mostly frag-
mented seeds, and chaff. (i) Early Neolithic (ca.
6,800–6,400 BP = ca. 7,700–7,250 cal BP): einkorn
wheat, mainly chaff (prevailing); barley (few); lentil
(frequent); pea (rare); bitter vetch (rare). Wild:
Lolium temulentum; Lathyrus cicera/sativus; Rubus sp.;
Pistacia terebinthus; Vitis vinifera; Ficus carica; numer-
ous herbs. (ii) Early-Middle Neolithic (ca. 6,400–
5,600 BP = ca. 7,400–6,300 cal BP): einkorn wheat,
mainly chaff (prevailing); barley (few); lentil (fre-
quent); pea (rare); bitter vetch (rare). Wild: Lolium
temulentum; Avena sp.; Lathyrus cicera/sativus; Rubus
sp.; Pistacia terebinthus; Vitis vinifera; Ficus carica;
numerous herbs.
5. Dimini, Thessaly (Kroll 1979). Late Neolithic,
Classic phase (5,630±150 uncal BP = ca. 6,650–6,300
cal BP). Rich remains: einkorn wheat (rare); emmer
wheat (frequent); six-rowed barley (rare); naked
barley (frequent); lentil (frequent); pea (frequent);
faba bean (few); bitter vetch (few); chickpea (few);
grass pea (frequent). Wild: Vitis vinifera (rare);
Amygdalus sp. (few).
6. Lerna, Argolis (Hopf 1961a). (i) Final Neolithic:
Ficus carica, Arbutus. (ii) Early Bronze Age (fifth mil-
lennium BP). Numerous remains: einkorn and
emmer wheats (few); six-rowed barley and naked
barley (few); lentil (frequent); pea (few); faba bean
(frequent); bitter vetch (frequent); grass pea (?)
(few); flax (frequent); fig (frequent, both fruits and
pips); grapevine (few).
7. Kastanas, Macedonia (Kroll 1983, 1984). (i)
Early Bronze Age (second half of fifth millennium
BP). Rich remains: einkorn wheat (frequent); emmer
wheat (frequent); free-threshing wheat (rare); spelt
wheat (rare); six-rowed barley (prevailing); lentil
(frequent); pea (rare); bitter vetch (frequent); faba
bean (rare); grass pea (rare); flax (few); poppy (rare);
grapevine (few); fig (frequent). Wild: Quercus acorns
(few); Pyrus sp. (few); Rubus fruticosus (rare);
Sambucus ebulus (few); Cornus mas (few); Lolium
temulentum (few); other grass and herb seeds. (ii)
Late Bronze Age (second half of fourth millennium
BP). Rich remains: einkorn and emmer wheats (fre-
quent); spelt wheat (rare); free-threshing wheat

(rare); six-rowed barley (few); lentil (frequent); bitter
vetch (prevailing); broomcorn millet (frequent); fox-
tail millet (few); poppy (rare); grapevine (frequent);
fig (few). Wild: Quercus acorns (rare); Sambucus ebu-
lus (rare); Secale cereale and Avena sp. (contaminating
wheat and barley); Camelina sativa (rare); Fragaria sp.
(rare); numerous grass and herb seeds.
Crete
Knossos, Crete (Sarpaki 2009). (i) Aceramic
Neolithic (7,740±130 uncal BP = ca. 8,650–8,400 cal
BP). Rich charred remains: einkorn wheat (rare);
emmer wheat (rare); bread wheat (prevailing);
hulled two-rowed and six-rowed barley (rare);
naked six-rowed barley (rare); lentil (few); fig
(rare); almond (rare). Wild: Avena sp.; Lolium sp.
(ii) Early Neolithic I (ca. 7,200–7,000 uncal BP = ca.
8,050–7,800 cal BP). Few remains, mainly charred:
einkorn wheat (rare); ‘eastivo-compactum’ wheat
(prevailing); barley (frequent); lentil ? (rare); flax
(rare); fig, nutlets and fruits (frequent); almond
(frequent). Wild condiments/aromatic: Raphanus
raphanistrum (wild radish), pods (frequent);
Satureja thymbra (few). (iii) Early Neolithic II (ca.
7,000–6,800 uncal BP = ca. 7,800–7,600 cal BP). Few
charred remains: ‘eastivo-compactum’ wheat (fre-
quent); barley (few); lentil (rare); flax? (rare); fig,
nutlets and fruits (prevailing); almond (frequent);
grapevine (rare). Wild condiments/aromatic:
Raphanus raphanistrum (wild radish), pods and
seeds (frequent); Satureja thymbra (rare); Thymelaea
hirsuta (rare). (iv) Middle and Late Neolithic. Rare
remains, mainly charred: wheat/barley (rare);
lentil (rare); pea? (rare); fig (frequent); almond
(prevailing).
Former Yugoslavia
(General references: Renfrew 1979; Kroll 1991)
1. Starčevo, Serbia (Renfrew 1979, Table 5). Early
Neolithic (ca. 5,000 BC = ca. 7,000 BP). Numerous
remains, all from impression: einkorn wheat (few);
emmer wheat (prevailing); naked wheat (few); six-
rowed barley (few); pea (few). Wild: Malus sp. and
Cornus mas (few).
2. Anza, Macedonia (Renfrew 1976). (i) Early
Neolithic (7,270±140 to 6,070±200 uncal BP = ca.
PLANT REMAINS IN REPRESENTATIVE ARCHAEOLOGICAL SITES 179
8,200–6,650 cal BP). Numerous finds: einkorn wheat
(few); emmer wheat (frequent); free-threshing
wheat (few); six-rowed barley (few); lentil (rare);
pea (rare). Wild: Malus sp. Corylus avellana, Cornus
mas, and Vitis vinifera (few). (ii) Late Neolithic (ca.
6,100 BP). Scarce remains: einkorn and emmer
wheat (frequent); six-rowed barley (rare); lentil
(rare); pea (rare).
3. Obre, Bosnia-Hercegovina (Renfrew 1974). (i)
Early Neolithic, Starčevo group (7,240±60 to
6,150±60 uncal BP = ca. 8,150–7,000 cal BP). Scarce
remains: einkorn wheat (rare); emmer wheat (pre-
vailing); free-threshing wheat (rare); lentil (few);
pea (frequent, in a single context). Wild: Cornus mas
(rare). (ii) Late Neolithic, Butmir culture (4,000–
3,860 BC = ca. 6,000–5,900 BP). Numerous remains:
einkorn wheat (frequent); emmer wheat (frequent);
free-threshing wheat (rare); six-rowed barley (fre-
quent); naked barley (frequent); lentil (frequent).
4. Vršnik, Štip, Macedonia (Hopf 1961a). Early
Neolithic (Starčevo group, 4,900 BC = ca. 6,900 BP).
Numerous remains: einkorn wheat (prevailing);
emmer wheat (frequent); free-threshing wheat
(rare); barley (rare).
5. Pokrovnik, Dalmatia (Karg and Müller 1990).
Middle Neolithic, Danilo culture (6,290±65 BP = ca.
7,300–7,200 cal BP). Numerous remains in a storage
context: einkorn wheat (few); emmer wheat
(prevailing).
6. Gomolava, Serbia (van Zeist 1975, 2003). Late
Neolithic, Vinča culture (ca. 5,715±75 uncal BP = ca.
5,900–5,700 cal BP). Rich remains: einkorn wheat
(prevailing); emmer wheat (frequent); six-rowed
barley (frequent); broomcorn millet (frequent); len-
til (few); pea (rare); Vicia sp. (rare); flax (rare). Wild:
Malus sp.; Cornus mas; Vitis vinifera; Physalis alkekengi
(all few); Fragaria vesca (rare); Avena sp. (rare).
Bulgaria
(General references: Renfrew 1979; Kroll 1991;
Popova 1995; Marinova 2007).
1. Karanovo, Sliven (Thanheiser 1997; Marinova
2004, 2006). (i) Early Neolithic, Phase I–II/III (ca.
8,000–7,550 cal BP). Few remains: einkorn wheat
(frequent); emmer wheat (prevailing); free-thresh-
ing wheats (few); hulled two/six-rowed barley (fre-
quent); lentil (few); bitter vetch (frequent); grass pea
180 DOMESTICATION OF PLANTS IN THE OLD WORLD
(few). Possible weeds: additional ca. 18 taxa. (ii)
Middle Neolithic, Phase III (6,510±60 to 6,130±60
uncal BP = ca. 7,500–6,950 cal BP). Rich remains:
einkorn wheat (few); emmer wheat (prevailing);
free-threshing wheat (few); naked six-rowed barley
(storages); grass pea (storages); bitter vetch (stor-
ages); lentil (storages). Possible weeds: additional
ca. two taxa. (iii) Late Neolithic, Phase III/IV and
IV (ca. 6,850–6,400 cal BP). Rich charred remains:
einkorn wheat (few); emmer wheat (prevailing);
free-threshing wheats (few); naked six-rowed bar-
ley (storages); grass pea (storages); bitter vetch
(storages); lentil (storages). Possible weeds: addi-
tional ca. thirty taxa. (iv) Early Eneolithic, Phase V.
Rich remains: einkorn wheat (storages); emmer
wheat (few); naked six-rowed barley (prevailing);
lentil (rare); bitter vetch (storages). Possible weeds:
additional ca. thirty-five taxa. (v) Late Eneolithic,
Phase VI (ca. 6,650 cal BP).Rich remains: einkorn
wheat (frequent); emmer wheat (frequent); naked
six-rowed barley (few); bitter vetch (storages). (vi)
Early Bronze Age, Phase VII (ca. 4,650–4,450 cal BP).
Rich remains: einkorn wheat (frequent); emmer
wheat (few); hulled six-rowed barley (prevailing);
lentil (rare); bitter vetch (storages); grass pea (rare);
safflower (few).Weeds: Galium sp.; Polygonum
convolvulus.
Wild fruits have been found in almost all phases:
Cornus mas (frequent); Vitis vinifera subsp. sylvestris;
Prunus avium/cerasus; Rubus ulmifolius/idaeus;
Sambucus nigra; Ficus carica; Corylus avellana; Fragaria
vesca; Quercus sp. (acorns); Physalis sp.
2. Kovacevo, Blagoevgrad (Popova 1992;
Marinova 2006). (i) Early Neolithic, Kovacevo
Ia-Id (7,245±36 to 7,028±35 uncal BP = ca. 8,150–
7,850 cal BP). Rich charred remains: einkorn
wheat (frequent); emmer wheat (frequent); hulled
barley (very frequent); lentil (frequent); grass pea
(very frequent); pea (few); bitter vetch (few);
chickpea (few). Wild: Cornus mas (very frequent);
Vitis vinifera subsp. sylvestris (frequent); Corylus
avellana; Prunus avium/cerasus (frequent);
Sambucus nigra; Rubus ulmifolius/idaeus (fre-
quent); Pistacia terebinthus. Possible weeds: addi-
tional ca. twenty-seven taxa. (ii) Bronze Age.
Rich charred remains: einkorn wheat (prevail-
ing); barley (frequent); millet (rare); grass pea
(few). Weed: Galium sp.
3. Azmaška (Tell Azmak), Sliven (Hopf 1973a;
Renfrew 1979, Table 7). (i) Early Neolithic, Phase I
(ca. 7,850–7,500 cal BP). Rich remains: einkorn
wheat (few); emmer wheat (prevailing); free-thresh-
ing wheat (frequent); naked six-rowed barley (rare);
lentil (frequent); pea (few); grass pea (few). (ii)
Middle Neolithic, Phase II (6,476±100 uncal BP = ca.
7,450–7,300 cal BP). Rich remains: bitter vetch (pure
storage). Wild: Sambucus sp. (iii) Eneolithic (ca.
6,400–6,300 cal. BP). Rich remains: einkorn wheat
(frequent); emmer wheat (frequent); naked six-
rowed barley (frequent); hulled six-rowed barley
(rare); pea (few); lentil (frequent); bitter vetch (pre-
vailing). Wild: Vicia sp.
4. Kapitan Dimitrievo, Pazardzhik (Marinova
2006, forthcoming). (i) Early Neolithic I (ca. 7,950–
7,650 cal BP). Rich charred remains: einkorn wheat
(storages); emmer wheat (storages); grass pea (stor-
ages). (ii) Early Neolithic II (ca. 7,850–7,700 cal BP).
Rich charred remains: einkorn wheat (prevailing,
storages); emmer wheat (frequent, storages); hulled
barley (storages); naked barley (storages); lentil
(frequent); grass pea (storages); bitter vetch (rare);
pea (rare); chickpea (rare). Wild: Cornus mas (very
frequent); Vitis vinifera subsp. sylvestris; Pistacia tere-
binthus; Corylus avellana; Prunus avium/cerasus;
Sambucus nigra (frequent); Rubus ulmifolius/idaeus
(frequent); Physalis sp. Possible weeds: additional
ca. thirty-six taxa. (iii) Late Neolithic: einkorn wheat
(prevailing); emmer wheat (frequent); naked barley
(rare); lentil (frequent); pea (few); bitter vetch (rare);
grass pea (prevailing); coriander (few). Wild: Cornus
mas (frequent); Corylus avellana; Prunus avium/cerasus
(frequent); Sambucus nigra (frequent); Rubus ulmifo-
lius/idaeus (frequent); Physalis sp.; Fragaria vesca.
Possible weeds: additional ca. two taxa. (iv) Early
Eneolithic. Rich charred remains: einkorn wheat
(prevailing); emmer wheat (few); naked wheat
(rare); bitter vetch (rare); grass pea (few).
5. Galabovo, south Bulgaria (Popova 1995b;
Popova 2001). (i) Chalcolithic culture. Rich remains:
einkorn wheat (very frequent); emmer wheat (fre-
quent); spelt wheat (few); free-threshing compactum-
type wheat (rare); hulled and naked barley
(frequent); bitter vetch (very frequent). (ii) Early
Bronze Age culture. Rich remains: einkorn wheat
(prevailing); emmer wheat (very frequent); spelt
wheat (rare); free-threshing compactum-type wheat

(rare); hulled and naked barley (frequent); lentil
(very frequent, storages); bitter vetch (very frequent,
storages). Weeds: Agrostemma githago; Chenopodium
album; Polygonum lapatifolium; Rumex sp.
6. Yunatzite, Pazardzhik (Popova and Pavlova
1994; Marinova and Popova 2008). (i) Late
Chalcolithic. Rich charred remains: einkorn wheat
(storages); emmer wheat (frequent); free-threshing
wheat (rare); naked barley (storages); millet (few);
lentil (frequent); bitter vetch (very frequent); grass
pea (storages); chickpea (few). Weeds: Agrostemma
githago; Chenopodium album; Polygonum convolvulus;
Rumex sp. (ii) Bronze Age. Rich charred remains:
einkorn wheat (very frequent, storages); emmer
wheat (frequent); hulled barley (prevailing, stor-
ages); lentil (frequent); bitter vetch (storages); grass
pea (storages). Wild: Quercus sp. acorns (frequent);
Vitis vinifera subsp. Sylvestris. Weeds: Agrostemma
githago; Bromus sp.; Centaurea sp.; Polygonum avicu-
lare; Brassicaceae.
Rumania
(General references: Wasylikowa et al. 1991;
Cârciumaru 1996; Monah 2007)
1. Liubcova, Caraş Severin district (Cârciumaru
1996). Neolithic Vinča culture B2 (second half of the
eighth millennium BP). Rich charred remains:
einkorn wheat (frequent); emmer wheat (frequent);
free-threshing wheat (frequent); hulled and naked
barley (few); lentil (prevailing). Wild: Galium spu-
rium; Rumex acetosa; Vicia sp.
2. Cârcea, Dolj district (Cârciumaru 1996).
Neolithic Dudeşti culture (ca. 6,500–6,000 BP). Rich
charred remains: einkorn wheat (few); emmer
wheat (few); free-threshing wheat (frequent); barley
(few); pea (pure hoard); rye (few).
3. Poduri, Bacău district (Cârciumaru and
Monah 1985; Monah and Monah 2008). (i) Late
Neolithic, Precucuteni culture (ca. 6,950–6,600 cal
BP). Vessels and adobe silos containing large
amount (tens of kilograms) of charred grains:
einkorn wheat (frequent); emmer wheat (frequent);
durum wheat? (rare); spelt wheat (rare); bread
wheat (frequent); club wheat (rare); domestic bar-
ley (frequent); Coriander (rare). Weeds: Setaria
viridis; Vicia cracca; Polygonum convolvulus; Rumex
acetosa; Galium sp.; Chenopodium album. Wild:
PLANT REMAINS IN REPRESENTATIVE ARCHAEOLOGICAL SITES 181
Corylus avellana; Tilia platyphyllos. (ii) Eneolithic
Cucuteni A culture (ca. 6,650–6,350 cal BP). Einkorn
wheat (frequent); emmer wheat (frequent); club
wheat (rare); bread wheat (frequent); domestic
barley (frequent); oat (rare); rye (rare); pea (rare);
grapevine (rare); Prunus domestica (rare); coriander
(rare). Weeds: Setaria pumila; Agrostemma githago;
Chenopodium album; Rumex acetosa; Brassica nigra;
Thlaspi arvense; Galium sp. Wild: Cerasus avium;
Rubus idaeus; Malus/Pyrus; Corylus avellana; Rosa
sp.; Cornus mas; Sambucus nigra. (iii) Eneolithic
Cucuteni B culture (sixth millennium BP). Einkorn
wheat (frequent); emmer wheat (frequent); domes-
tic barley (frequent); rye (rare); coriander (rare).
Wild: Sambucus nigra; Viburnum lantana; Cornus
mas; Rubus sp.; Vicia sativa.
4. Celei (Sucidava), Olt district (Cârciumaru
1996). Transition from Eneolithic to Bronze Age (ca.
4,200 BP). Large quantities of charred grains:
einkorn wheat (few); emmer wheat (frequent); spelt
wheat (frequent); free-threshing wheat (frequent);
six-rowed hulled barley (frequent); lentil (few);
lumps of linseed. Wild: Rumex acetosa; Carex
rostrata.
5. Tell Hârşova, Constanţa district (Cârciumaru
1996; Monah 2002; Monah and Monah 2008). (i)
Late Neolithic Boian culture (seventh millennium
BP). Einkorn wheat (frequent); wild emmer wheat?
(frequent?); emmer wheat (frequent); spelt wheat
(frequent); pea (frequent). (ii) Late Neolithic/
Eneolithic Gumelniţa Culture (ca. 6,350–6,150 cal
BP). Einkorn wheat (few); emmer wheat (few);
bread wheat (few); domestic barley (few); rye (few);
lentil (frequent); pea (frequent); bitter vetch (pre-
vailing); grapevine (rare). Wild: Vicia sp.; Sambucus
nigra; Vitis sylvestris.
Moldavia and Ukraine
(General references: Januševič 1984; Wasylikowa
et al. 1991; Pashkevich 2003)
1. Sacarovca, Sângerei district, Moldavia
(Januševič 1984; Kuzminova et al. 1998). Early
Neolithic, Starčevo-Criş culture (6,650±50 uncal. BP
= ca. 7,600–7,500 cal BP). Numerous remains, both
imprints and charred grains: einkorn wheat (fre-
quent); emmer wheat (prevailing); spelt wheat
(few); hard wheat, T. aestivum/compactum (rare); oat
182 DOMESTICATION OF PLANTS IN THE OLD WORLD
(rare); domestic barley (frequent); broomcorn mil-
let (frequent); pea (few); cherry plum (rare); plum
(rare). Weeds: Alyssum sp.; Agrostemma sp.; Lathyrus
sp.; Setaria viridis; Setaria glauca; Galium sp. Wild:
Coryllus avelana; Quercus robur; Cornus mas; Malus
sp.; Vitis sylvestris.
2. Ruseştii Noi (Novye Rusešty), Chişinău dis-
trict, Moldavia (Januševič 1976, 1986).(i)
Linearbandkeramik (ca. 7500 BP), ear impressions
in pottery: emmer wheat; spelt wheat. (ii)
Precucuteni (Tripolye A, early phase) culture
(5,615±100 uncal BP = 6,500–6,300 cal BP): impres-
sions and charred grains: einkorn wheat; emmer
wheat; spelt wheat; hard wheat; naked barley. (iii)
Cucuteni A (Tripolye B, middle phase), impressions
and charred grains: emmer wheat; spelt wheat;
naked barley; grapevine.
3. Putineşti, Soroca district, Moldavia (Januševič
1976). Eneolithic, Precucuteni (ca. 6,000 BP).
Impressions and charred grains: einkorn wheat;
emmer wheat; spelt wheat; hard wheat; barley;
naked barley; oat; broomcorn millet? Wild: Cornus
mas; Pyrus elaegrifolia; Cerasus avium.
4. Cuconeştii Vechi (Starye Kukoneshti),
Moldavia (Januševič 1978, 1984; Kuzminova 1988).
Eneolithic, Cucuteni A–B (Tripolye culture, middle
phase). Rich remains (mainly carbonized, few
imprints): einkorn wheat (few); emmer wheat (pre-
vailing); naked barley (few). Weeds: Rumex ace-
tosella; R. acetosa; Chenopodium album; Saponaria
officinalis; Sinapis arvensis; Polygonum aviculare;
Echinochloa crus-galli; Medicago falcata; Solanum
nigrum; Stellaria media; Bromus secalinus; Datura stra-
monium; Glaucium corniculatus.
5. Rivne (Rovno), Rivne Oblast, Ukraine (Pash-
kevich 2003). Linearbandkeramik (ca. 6,500–6,000
BP). Rare imprints: emmer wheat; bread wheat;
naked barley; broomcorn millet; pea.
6. Eneolithic settlements in Ukraine, Tripolye
cultures (ca. 6,000–4,750 BP). Luca Vrublevecaja
(Januševič 1976), Majdaneckoe, Cherkas’ka
Oblast, Majaki and Usatovo, Odessa Oblast
(Pashkevich 2005). Few remains, mostly imprints,
some charred: einkorn wheat; emmer wheat;
spelt wheat; bread wheat; hard wheat; naked
barley; broomcorn millet; pea; plum; cherry
plum; apricot. Wild: Vitis sp.; Pyrus/Malus; Corylus
avellana.
Hungary
(General references: Tempír 1964; Hartyáni and
Nováki 1975; Füzes 1990; Wasylikowa et al. 1991;
Gyulai 2007).
1. Körös-Starčevo culture sites (Szeged-Gyálarét,
Röszke-Lúdvár, Battonya-Basarága) (Hartyányi
et al. 1968; Hartyányi and Nováki 1971; Hartyáni
and Nováki 1975; Füzes 1990). Early Neolithic (ca.
7,950–7,250 cal BP). Imprints: einkorn wheat (pre-
vailing); emmer wheat (few). Probable weeds:
Lathyrus sp.; Bromus sp. Wild: Cornus mas; Corylus
avellana.
2. Füzesabony-Gubakút, Northern Great Plain
(Gyulai 2007). Middle Neolithic LBK culture, Great
Hungarian Plain Group (ca. 7,550–7,150 cal BP).
Scarce charred remains: emmer wheat (rare); barley
(frequent). Weed: Avena fatua.
3. Zánka-Vasútállomás (Füzes 1990, 1991).
Middle Neolithic LBK culture, Trans-Danubian
Group (ca. 7,250–6,650 cal BP). Rich charred remains
and imprints: einkorn wheat (including wild forms),
grains, and rachis fragments (very common); emmer
wheat, grains, and rachis fragments (very common);
bread wheat; club wheat (Triticum aestivum subsp.
compactum); spelt wheat, spikelets; two-rowed bar-
ley grains; common millet; pea. Weeds: Agrostemma
githago; Bromus cf. sterilis; Galium sp.; Fallopia convol-
vulus; Chenopodium sp.; rye brome; T. baeoticum?
4. Dévaványa-Réhelyi dűlő (Hartyányi et al.
1968). Middle Neolithic LBK culture, Eastern
region, Szakálhát-Szilmeg Group (ca. 6,350 cal BP).
Rich charred remains: einkorn wheat (few); emmer
wheat (frequent); barley (prevailing); lentil (few).
Weeds: Fallopia convolvulus; Chenopodium sp.
5. Berettyóújfalu-Szilhalom (Hartyányi et al.
1968). Late Neolithic, Tisza-Herpály-Berettyóvölgy
site. Rich charred remains: einkorn wheat (fre-
quent); emmer wheat (few); bread wheat (rare);
two-rowed naked barley (frequent); common mil-
let; field pea (frequent); bitter vetch (rare). Wild:
Malus sylvestris fruits.
6. Rákoskeresztúr-Újmajor (Gyulai 1999).
Eneolithic (Middle Copper Age), Ludanice culture
(ca. 6,250–5,450 cal BP). Rich remains: emmer wheat
(rare); six-rowed barley (few). Wild: Rumex conglom-
eratus; Schoenoplectus lacustris; Thalicrum flavum;
Chenopodium sp.

7. Budapest. (i) Csepel-Vízmű (Endrődi and
Gyulai 2000). Eneolithic (Late Copper Age), Baden
culture (ca. 5,450–4,950 cal BP). Rich charred
remains: einkorn wheat (few); emmer wheat (rare);
club wheat (rare); barley (rare). Weeds: Bromus
arvensis; B. secalinus; Chenopodium sp.; Fallopia con-
volvulus. Wild: Crataegus monogyna; Fagus silvatica;
Quercus robur/petraea. (ii) Albertfalva (Gyulai 2003).
Early Bronze Age (ca. 4,950–3,550 cal BP), Bell
Beaker culture Csepel Group. Rich charred remains:
einkorn wheat (frequent); emmer wheat (frequent);
bread wheat (rare); spelt wheat (rare); six-rowed
naked barley (few); two-rowed barley (frequent);
pea (rare); small seed fava bean (rare). Wild: crab
apple; many weeds.
8. Bölcske-Vörösgyír (Berzsényi and Gyulai
1998). Middle Bronze Age, Nagyrév and Vatya cul-
ture (ca. 3,550–3,250 cal BP). Rich charred remains:
einkorn wheat (prevailing); emmer wheat (fre-
quent); spelt wheat (few); bread wheat (rare); six-
rowed barley (frequent); two-rowed barley
(frequent); naked barley (few); common millet
(few); small seed lentil (frequent); bitter vetch (fre-
quent); pea (few); small seed fava bean (few).
Weeds: Avena fatua; Bromus arvensis; B. mollis; B. sec-
alinus; B. tectorum; Chenopodium sp.; Fallopia convol-
vulus; Setaria viridis. Wild: Cornus mas; Crataegus
monogyna; Prunus spinosa; Quercus sp.
Austria
(General references: Küster 1991; Kohler-Schneider
2007)
1. Schletz near Asparn, Lower Austria (Schneider
1994; Kohler-Schneider 2007). Late Linearbandkera-
mik culture (6,175±65–6,025±55 uncal BP=ca. 7,150–
6,800 cal BP). Major fortified site of the Early Neolithic.
Rich charred remains: einkorn wheat, mainly two-
grained (very frequent); emmer wheat (very fre-
quent); naked wheat (few); hulled barley (few); pea
(few); lentil (rare); flax (linseed); cf. Camelina sativa;
Papaver somniferum spp. setigerum. Wild: Cornus mas;
Sambucus nigra; Sambucus ebulus; Solanum dulcamara;
Physalis alkekengi. Numerous weeds and herbs:
Chenopodium album; Ch. hybridum; Fallopia convolvulus;
Polygonum aviculare; Bromus secalinus; wild Paniceae.
2. Mondsee, lake-shore settlements, east of
Salzburg, Upper Austria (Hofmann 1924; Kohler-
PLANT REMAINS IN REPRESENTATIVE ARCHAEOLOGICAL SITES 183
Schneider 2007). Late Neolithic (4,910±130–4,750±90
uncal BP=ca. 5,750–5,450 cal BP). Rich charred and
waterlogged remains: einkorn wheat (few); emmer
wheat (very frequent); hulled six-rowed barley
(very frequent); pea (rare); flax (few); poppy (few).
Wild: Corylus avellana;Malus sylvestris (entire and
halves fruits); Rosa canina; Fagus sylvatica; Quercus
robur; Tilia platyphyllos; Rubus idaeus; R. fruticosus;
Fragaria vesca; Daucus carota; Pastinaca sativa.
3. Anzingerberg/Hundssteig, near Krems,
Lower Austria (Kohler-Schneider 2007; Kohler-
Schneider and Caneppele 2009). Late Neolithic
Jevišovice culture (ca. 5,050–4,750 cal BP). Rich
charred remains: einkorn wheat; emmer wheat;
spelt wheat (few, very early); free-threshing wheat
(rare); hulled barley; broomcorn millet (frequent);
foxtail millet (few); lentil (rare); pea (rare); flax
(rare); poppy (very frequant). Wild: Vitis vinifera
subsp. sylvestris; Prunus spinosa; Rubus fruticosus; R.;
Fragaria vesca; Viburnum opulus; Camelina microcarpa;
Daucus carota; Atropa belladonna; Hyoscyamus niger.
Additional ca. 100 wild taxa from various habitats.
4. Stillfried, Lower Austria (Kohler-Schneider
2001; Kohler-Schneider 2003). Late Bronze Age hill-
fort site (ca. 3,200–2,700 cal BP). Rich charred
remains from storage pits and other domestic con-
texts: einkorn wheat (very frequent); emmer wheat
(frequent); spelt wheat (very frequent); free-thresh-
ing wheat (rare); hulled barley (very frequent); rye
(few); broomcorn millet (very frequent); foxtail mil-
let (frequent); lentil (frequent); pea (rare); fava bean
(few); bitter vetch (rare); gold of pleasure (frequent);
poppy (rare); grapevine (few). Wild: Prunus avium/
cerasus; Malus/Pyrus; Fragaria sp.; Digitaria sanguina-
lis; Daucus carota; Physalis alkekengi. Additional ca.
100 wild taxa from various habitats.
5. Dürrnberg/Hallein, Salzburg (Werneck 1949;
Swidrak 1999; Boenke 2007). Iron Age salt mining
site and settlement (ca. 2,400–2,050 cal BP). Rich
remains from human feces (salt preservation),
charred and waterlogged material: emmer wheat
(few); spelt wheat (few); free-threshing wheat (rare),
hulled and naked barley (frequent); broomcorn mil-
let (very frequent); foxtail millet (few); lentil (few);
pea (few); broad bean (few); poppy (frequent); gold
of pleasure (few); hemp, fruits, and fibres (frequent);
flax, seeds, and fibres (few). Wild: Corylus avellana;
Prunus domestica subsp. insititia; P. spinosa; P. avium;
184 DOMESTICATION OF PLANTS IN THE OLD WORLD
Malus sylvestris; Pyrus sp.; Rubus fruticosus; R. cae-
sius; R. idaeus; Fragaria vesca; Rosa sp.; Crataegus
laevigata;Vitis vinifera cf. subsp. sylvestris; Sambucus
nigra; Viburnum lantana; Daucus carota; Carum carvi.
Italy
(General references: Rottoli and Pessina 2007;
Rottoli and Castiglioni 2009)
1. Grotta dell’Uzzo, Sicily (Costantini, 1989). (i)
Mesolithic (ca. 9,950–8,950 uncal BP = ca. 11,400–
10,150 cal BP). Scarce remains, all wild: Arbutus
unedo; legume seeds (probably Lathyrus or Pisum);
Quercus acorns; two pips of Vitis. (ii) Early Neolithic
(6,750±70 uncal BP = ca. 7,650–7,550 cal BP). Scarce
remains (unspecified quantities): einkorn wheat;
emmer wheat; lentil; barley (somewhat later). (iii)
Middle Neolithic (ca. 6,750–6,450 uncal BP = ca.
7,600–7,350 cal BP). Few remains: einkorn wheat;
emmer wheat (prevailing); free-threshing wheat;
barley (relatively common); lentil; pea; bitter vetch;
faba-like bean. Wild: Amygdalus sp; Olea europaea;
Vitis vinifera; Ficus carica.
2. Scamuso, Torre a Mare, Bari (Costantini et al.
1997). Numerous remains including chaff: (i)
Impressed Ware culture (ca. 7,150–6,750 uncal BP =
ca. 8,000–7,600 cal BP): einkorn wheat (few); emmer
wheat (frequent); hulled barley (prevailing); lentil
(few); pea (few). Wild: Vitis vinifera (few); Lathyrus
sp. (frequent). (ii) Painted Ware culture (ca. 6,600–
5,800 uncal BP = ca. 7,500–6,550 cal BP): einkorn
wheat (rare); emmer wheat (frequent); free-thresh-
ing wheat (few); barley (prevailing); lentil (rare);
pea (rare); faba-like bean (a single seed). Wild: Avena
sp. (rare); Vitis vinifera (few); Lathyrus sp. (few); few
herbs.
3. Pienza, Siena (Castelletti 1976). (i) Impressed
Ware (Cardial) culture (ca. 7,750–7,250 cal BP).
Scarce remains: einkorn wheat (rare); emmer wheat
(prevailing); free-threshing wheat (frequent);
hulled/naked barley (few). Wild: Avena sp.; Vitis
sylvestris. (ii) Middle Neolithic (ca. 6,950–6,450
uncal. BP = ca. 7,800–7,350 cal BP). Scarce remains:
emmer wheat (rare); free-threshing wheat (rare);
barley (rare). Wild: cf. Avena. (iii) Late Neolithic (ca.
6,450–5,950 uncal. BP = ca. 7,400–6,750 cal BP).
Scarce remains: emmer wheat (rare); free-threshing
wheat (rare), barley (rare). Wild: Avena sp. (iv)
Bronze Age: Scarce remains: einkorn wheat (few);
emmer wheat (prevailing); hulled barley (few);
broomcorn millet (few); faba bean (few). Wild:
Avena sp.; Cornus mas.
4. Sammardenchia, and other adjacent Early
Neolithic sites in Friuli, northern Italy (Pessina and
Rottoli 1996; Rottoli 2005; Rottoli and Pessina 2007).
Early north Italian Neolithic (6,570±74 uncal BP to
5,684±58 uncal BP= ca. 7,550–6,450 cal BP). Rich
remains: einkorn wheat (frequent); emmer wheat
(frequent); free-threshing wheat (rare); six-rowed
hulled barley (frequent); lentil (frequent); pea (fre-
quent); vetches (Vicia sativa agg.) (frequent); bitter
vetch (few); grass pea (few). Rye, spelt, and millets
apparently infested grain agriculture. Wild: Vitis
vinifera; Corylus avellana; Cornus mas; Malus sylves-
tris; Quercus sp.; Rubus fruticosus agg.; Crataegus sp.;
Prunus insititia/spinosa; Juglans regia; Bromus spp.
(frequent). Weeds: Galium aparine; Rumex sp.; Fallopia
convolvulus.
5. San Marco, Gubbio, Perugia (Costantini and
Stancanelli 1994). Early-Middle Neolithic (6,300–
5,750 uncal BP = ca. 7,250–6,500 cal BP). Rich
remains: einkorn wheat (frequent); emmer wheat
(frequent); free-threshing wheat (prevailing); com-
pactum-type wheat (frequent); hulled barley (few);
naked barley (rare); pea (rare). Wild: Corylus avel-
lana (frequent); Ficus carica; Prunus insititia; Vitis
vinifera.
6. Monte Còvolo, Villa Nuova sul Clisi, Brescia
(Pals and Voorrips 1979). (i) Late Neolithic (ca.
5,850–5,450 uncal BP = ca. 6,700–6,200 cal BP). Scarce
remains: einkorn wheat (rare); emmer wheat (pre-
vailing); six-rowed barley (few). Wild: Cornus mas;
Malus sylvestris; Prunus avium; P. spinosa; Vitis sylves-
tris; Quercus sp.; Rosa sp.; Rubus sp.; Physalis
alkekengi. (ii) White Ware phase (ca. 5,500–4,150
uncal BP = ca. 6,300–4,600 cal BP). Scarce remains:
einkorn wheat (few); emmer whet (prevailing); six-
rowed barley (frequent); broomcorn millet (rare).
Wild: Prunus avium; Quercus sp.; Vitis sylvestris; Rosa
sp; Rubus sp.; Physalis alkekengi. (iii) Bell Beaker
phase (ca. 4,150–3,900 uncal BP = ca. 4,800–4,300 cal
BP). Numerous remains: einkorn wheat (few);
emmer wheat (prevailing); six-rowed barley (fre-
quent). Wild: Cornus mas; Malus sylvestris; Prunus
avium; P. spinosa; Quercus sp.; Vitis sylvestris; Physalis
alkekengi. (iv) Early Bronze Age (ca. 3,900–3,350

uncal BP = ca. 4,400–3,550 cal BP). Scarce remains:
emmer wheat (prevailing); six-rowed barley (few).
Wild: Malus sylvestris.
7. La Marmotta, Lake Bracciano, Rome (Rottoli
1993, 2002). Rich Early Neolithic (6,855±65 uncal BP
to 6,310±75 uncal BP = ca. 7,750–7,150 cal BP) water-
logged remains: einkorn wheat; emmer wheat; free-
threshing turgidum-type wheat; hulled two-rowed
barley; lentil; pea; grass pea; vetches; flax (rare). Wild:
Vitis vinifera (frequent); Papaver somniferum subsp.
setigerum (very frequent); Corylus avellana; Malus syl-
vestris.; Pyrus sp.; Quercus sp.; Ficus carica; Prunus
spinosa/domestica; Fragaria vesca; Cornus mas; Carthamus
lanatus; Silybum marianum; Crataegus sp.; Rubus fruti-
cosus agg.; Cornus sanguinea; Laurus nobilis.
Poland
(General references: Wasylikowa et al. 1991; Bieniek
2007; Lityńska-Zając 2007)
1. Strachów, Silesia (Lityńska-Zając 1997). (i)
Linearbandkeramik culture (6,535–6,170 uncal BP =
ca. 7,350–7,050 cal BP). Numerous remains (imprints
and some charred grains): einkorn wheat (few);
emmer wheat (prevailing); barley (few). Wild: vari-
ous grasses and herbs such as Bromus sp.; Setaria
viridis/verticillata; Echinochloa crus-galli; Polygonum
sp.; Rumex sp.; Chenopodium sp.; Bromus cf. tectorum;
Chenopodium album; Polygonum convolvulus; Rumex
sp. (ii) Lengyel-Polgár culture (ca. 5,730 uncal BP =
ca. 6,450 cal BP). Scarce remains (charred and
imprints): einkorn wheat (few); emmer wheat (few);
barley (very few). (iii) Funnel Beaker culture (ca.
4,800–4,480 uncal BP = ca. 5,550–5,050 cal BP). Some
remains, mostly imprints: einkorn wheat (rare);
emmer wheat (prevailing); barley (rare). Wild: vari-
ous grasses and herbs; among others, Bromus ssp.
and Chenopodium album.
2. Nowa Huta-Mogila, Kraków district (Gluza
1984). Neolithic, Lengyel culture (ca. 5,430 uncal BP
= ca. 6,300–6,200 cal BP). Rich remains: einkorn
wheat (frequent); emmer wheat (prevailing); naked
six-rowed barley (frequent). Large variety of herba-
ceous weedy species: Bromus arvensis (very abound-
ant); Bromus racemosus (abundant); several species
of Chenopodium, Polygonum and Galium; Fallopia con-
volvulus (abundant); Digitaria ssp.; Echinochloaa crus-
galli; Setaria viridis/verticillata; Solanum nigrum;
PLANT REMAINS IN REPRESENTATIVE ARCHAEOLOGICAL SITES 185
Trifolium ssp.; Wild: Corylus avellana; Sambucus
ebulus.
3. Gniechowice and Stary Zamek (Gluza 1994).
Linearbandkeramik culture (ca. 6,950 uncal BP =
7,800–7,750 cal BP). Mostly imprints: einkorn wheat
(few); emmer wheat (prevailing); barley (few);
Bromus sp. (few).
4. Ćmielów, Tarnobrzeg district (Klichowska
1976). Funnel Beaker culture (ca. 5,600–5,300 cal
BP). Rich remains: emmer wheat (prevailing); pea
(few); lentil (few); linseed (frequent). Wild:
Chenopodium album; Bromus secalinus; Rumex acetosa;
Setaria glauca; Berberis vulgaris; Taxus baccata.
5. Gwoździec, Małopolska (Bieniek and Litynska-
Zajac 2001; Lityńska-Zając 2007). Linear Pottery cul-
ture (ca. 7,400–6,600 cal BP). Numerous charred and
imprinted remains: einkorn wheat (frequent);
emmer wheat (prevailing); naked barley (rare).
Wild: Chenopodium album;Bromus sp.; Echinochloa
crus-galli; Setaria pumila; Malus sylvestris (pips and
fruit fragments).
6. Brześć group, Kujawy (Bieniek 2007). (i)
Linearbandkeramik culture (ca. 7,450–6,950 cal BP).
Numerous charred remains: einkorn wheat, mainly
chaff (prevailing); emmer wheat, mainly chaff (fre-
quent); hulled barley (rare); flax (rare); poppy (very
rare). Wild: Chenopodium album; Fallopia convolvulus;
Echinochloa crus-galli; several species of Polygonum;
Physalis alkegengi; Solanum nigrum;Schoenoplectus
tabernaemonatani; and other species. (ii) Lengyel cul-
ture (ca. 6,350–5,950 cal BP). Rich charred remains:
einkorn wheat (frequent); emmer wheat (prevail-
ing); bread wheat (rare); hulled barley (rare). Wild:
several species of Bromus and Galium; Chenopodium
album; Fallopia convolvulus; Hierochloe cf. australis;
Stipa pennata; and other species. (iii) Funnel Beaker
culture (ca. half of the sixth millennium BP).
Numerous charred remains: einkorn wheat, mainly
chaff (prevailing); emmer wheat, chaff (rare); poppy
(rare). Wild: Chenopodium album; Fallopia convolvu-
lus; several species of Polygonum; and others.
Czech Republic and Slovakia
(General references: Hajnalová 1989; Wasylikowa
et al. 1991; Hajnalová 2007)
1. Mohelnice, Litovel, Moravia (Opravil 1979;
Kühn 1981; Opravil 1981). Linearbandkeramik
186 DOMESTICATION OF PLANTS IN THE OLD WORLD
culture (6345±100 to 622±80 uncal BP = ca. 7,400–
7,000 cal BP). Numerous remains: einkorn wheat
(rare); emmer wheat (prevailing); free-threshing
wheat (frequent); two-rowed barley (few); broom-
corn millet (rare); pea or vetch (rare); flax cord (few).
Wild: Malus sylvestris; Corylus avellana.
2. Blatné near Štrky, Bratislava district, Slovakia
(Hajnalová 1989). (i) Linearbandkeramik culture
(ca. 7,250–7,000 cal BP). Numerous remains: einkorn
wheat (frequent); emmer wheat (prevailing); spelt
wheat (few); pea (rare). (ii) Middle Neolithic,
Zeliezovce group (ca. 6,950–6,650 cal BP). Numerous
remains: einkorn wheat (frequent); emmer wheat
(prevailing); spelt wheat (few); hulled six-rowed
barley (rare); pea (rare); lentil (few).
3. Bylany, Kutná Hora, Bohemia (Tempír 1979).
(i) Linearbandkeramik culture (ca. 7,400–7,000 cal
BP), Stroked pottery culture (ca. 7,000–6,550 cal BP).
Numerous remains: einkorn wheat (few); emmer
wheat (prevailing); broomcorn millet (few); pea
(few); lentil (few). (ii) Funnel Beaker culture (ca.
6,950–5,150 cal BP). Rich remains: einkorn wheat
(prevailing); emmer wheat (few); free-threshing
wheat (few); six-rowed hulled barley (frequent).
Wild: several weeds.
4. Šarišské Michalˇany-Fedelemka, Sabinov,
Prešov, Slovakia (Hajnalová 1993; Hajnalová and
Hajnalová 2004). Middle Neolithic (Bükk culture,
ca. 6,950–6,650 cal BP). Numerous charred remains:
einkorn wheat (frequent); emmer wheat (frequent);
spelt wheat (few); free-threshing wheat (few);
hulled and naked barley (frequent); rye (few);
broomcorn millet (few); pea (few); lentil (few); flax
(few). Wild: Malus sp.; cf. Cerasus avium; Prunus sp.;
Rubus sp.; Trapa natans.
5. Bajč-Medzi kanálmi, Komárno, Nitra,
Slovakia (Cheben and Hajnalová 1997). Middle
Neolithic (Želiezovce group, ca. 6,950–4,700 cal BP).
Rich charred and imprints, mostly from in situ con-
tent of an oven: one- and two-seeded einkorn (pre-
vailing); emmer wheat (few); spelt wheat (few);
free-threshing wheat (rare); naked six-row barley
(rare). Weed seeds.
6. Nitriansky Hrádok, Nové Zámky, Slovakia
(Tempír 1969; Kühn 1981). (i) Early Bronze Age
(Maďarovce culture, ca. 1,950–1,400 cal BC = ca.
3,900–3,350 cal BP). Rich remains: einkorn wheat
(frequent); emmer wheat (prevailing); free-thresh-
ing wheat (frequent); six-rowed and two-rowed
barley (frequent); oat (rare); rye (rare); pea (few/
pure); lentil (rare). Wild: Avena fatua; Bromus secali-
nus; Lolium sp.; Vicia sp.; Agrostemma githago; Quercus
robur. (ii) Bronze Age. Numerous remains: einkorn
wheat (rare); emmer wheat (rare); six-rowed barley
(prevailing); pea? Wild: Bromus secalinus; B. arvensis;
Lolium temulentum; Agrostemma githago; various
Cruciferae.
Switzerland
(General references: Jacomet 2006, 2007, 2008;
Jacomet and Brombacher 2005; Jacomet and Behre
2009).
1. Egolzwil 3, Wauwiler Moos, Canton of Lucerne
(Bollinger 1994; Jacomet 2007). Early Late Neolithic
Egolzwil culture (ca. 5,650 uncal BP = ca. 6,250 cal
BP). Lake-shore settlement at a small shallow lake.
Rich remains, mostly waterlogged: einkorn wheat
(rare); emmer wheat (rare); tetraploid free-threshing
wheat (frequent); six-rowed barley (frequent); pea
(frequent); poppy seeds (very frequent); flax, cap-
sules and seeds (few). Wild: hazelnut; crab apple;
blackberry and raspberry; wild strawberry (Fragaria
vesca); rose hips; Brassica rapa (frequent); additional
some 100 taxa—the majority could have been of
use.
2. Zürich (town and lake), numerous lake-shore
settlements (Jacomet 1988, 2004; Jacomet et al. 1989;
Brombacher and Jacomet 1997; Favre 2002;
Brombacher et al. 2005). Very rich (several hundreds
of thousands) charred or excellently preserved
waterlogged plant remains, from several sites cov-
ering the various phases of the lake-shore agricul-
tural settlements. (i) Late Neolithic cultures
Egolzwil, Early to classical Cortaillod, Pfyn, Horgen,
Schnurkeramik (Corded ware) (ca. 6,250–4,450 cal
BP, mostly dendrodates). Einkorn wheat (rare);
emmer wheat (rare in the earlier phases, prevailing
in the later phases); durum/turgidum-type tetraploid
wheat (prevailing in the earlier phases, frequent in
the later phases); six-rowed, mostly naked barley
(frequent); pea (rare, present from Egolzwil culture
(ca. 6,350 cal BP) onward; flax, seeds, and capsules
(becomes frequent from ca. 5,750 cal BP onwards);
poppy (frequent until ca. 4,750 cal BP, then becomes
rare). Mediterranean aromatic plants (from ca. 5,900

cal BP onwards): celery Apium graveolens (rare); dill
Anethum graveolens (rare); lemon balm Melissa offici-
nalis (rare). Large quantities of many plants col-
lected from the wild, particularly: Malus sylvestris;
Corylus avellana; Prunus spinosa; Rubus idaeus; R. fru-
ticosus; Fragaria vesca; Brassica rapa; as well as
remains of numerous weeds and local herbs. (ii)
Bronze Age sites (Mozartstrasse, Alpenquai, and
Wädenswil-Vorder-Au). Different phases of the
Early Bronze Age from ca. 3,850 cal BP onwards,
and Late Bronze Age, ca. 3,000–2,800 cal BP, dendro-
dates). Same crop species as in the Late Neolithic
sites, with the addition of spelt wheat (Triticum
spelta) from Early Bronze Age onwards, and broom-
corn millet (Panicum milieaceum), foxtail millet
(Setaria italica), and lentil (Lens culinaris). At
Alpenquai also Camelina was frequent.
3. Toos-Waldi, Canton of Thurgau (Jacomet and
Behre 2009). Dryland site on a hilltop, Middle
Bronze Age. Rich charred remains: einkorn wheat
(rare); emmer wheat (frequent); naked tetra/hexa-
ploid wheat (rare); spelt wheat (frequent); six-rowed
hulled barley; whole stocks (frequent). Wild: Avena
sp.; Quercus sp., many acorn halves.
4. Canton of Grisons (Graubünden), south-east
Switzerland (Jacomet et al. 1998, 1999). Rich plant
remains from Early Bronze Age onwards from six
sites, the most important one is Savognin-Padnal.
(i) Early to Middle Bronze Age (ca. 4,150–3,200 cal
BP). Einkorn wheat (few); emmer wheat (frequent);
spelt wheat (very frequent at one site); hulled six-
rowed barley (prevailing); pea (frequent); faba bean
(many at one site); flax (few). (ii) Late Bronze Age
(ca. 3,200–2,700 cal BP): einkorn wheat (rare); spelt
wheat (rare); hulled six-rowed barley (prevailing);
broomcorn millet (rare); pea (few). Wild: Corylus
avellana; Quercus sp.; Rosa sp.; Avena fatua; and seeds
of other weeds.
5. Lake Biel, Lake-shore settlements, particularly
Twann, Nidau-BKW, and Port-Stüdeli, Sutz, Lattrigen,
and Lüscherz (Ammann et al. 1981; Brombacher 1997,
2000; Brombacher and Jacomet 2003). (i) Late
Neolithic, different phases of the Cortaillod culture
(ca. 5,800–5,550 BP, dendro-dates). Rich waterlogged
and charred remains, often pure hoards: einkorn
wheat (rare to frequent, also two-grained cultivars
present); emmer wheat (rare); tetraploid free-thresh-
ing wheat (often prevailing); six-rowed barley, mostly
PLANT REMAINS IN REPRESENTATIVE ARCHAEOLOGICAL SITES 187
naked (very frequent); pea, seeds, and pod fragments
(in some layers of Twann frequent); flax (frequent,
stems in Port-Stüdeli); poppy (frequent); dill (rare, at
two sites). Wild: many gathered plants such as Malus
sylvestris; Corylus avellana; Rubus idaeus; R. fruticosus;
Fragaria vesca; Physalis alkekengi; Brassica rapa; Rosa sp.;
Quercus sp. (ii) Different Late Neolithic phases after
5,450 cal BP (Lattrigen, Horgen culture, ca. 5,410–5,176
BP, dendro-dates). Rich waterlogged and charred
remains: einkorn wheat (very rare); emmer wheat
(prevailing at one site); free-threshing tetraploid
wheat (very frequent); six-rowed barley, apparently
naked (very frequent); flax (frequent); poppy (fre-
quent); celery and dill (rare, at two sites). Wild: same
assemblage as in (i).
6. Lake Neuchatel, shore and surroundings
(Villaret-Von Rochow 1971a, 1971b; Baudais-
Lundström 1978; Schlichtherle 1985; Märkle 2000;
Karg and Märkle 2002; Mermod 2007). (i) Late
Neolithic to Early Bronze Age (ca. 5,659–4,400 BP,
dendrodates). Dryland and lake-shore sites, partic-
ularly St Aubin Derrière La Croix, Concise-sous-
Colachoz, Yverdon Avenue des Sports, and St Blaise
Bains des Dames. Rich waterlogged and charred
plant remains, including stocks: einkorn wheat (few
to prevailing); emmer wheat (few to prevailing);
tetraploid naked wheat (usually many); six-rowed
barley (usually many); celery and dill (rare); pea
(rare); flax (frequent); poppy (frequent). (ii)
Cortaillod sur les Rochettes Est (Akeret, 2005). Final
Neolithic, Bell Beaker culture (ca. 4,250 uncal BP =
ca. 4,850 cal BP). Rich remains: einkorn wheat (few);
emmer wheat (few); spelt wheat, grains, and chaff
(frequent); barley (rare); lentil (rare); bitter vetch;
flax. Wild: rich spectrum, including Corylus avellana;
Malus sylvestris; Prunus spinosa; Rosa sp.; Rubus sp.
7. Canton of Valais, valley of Valais (Wallis). (i)
Sites in Sion (Martin et al. 2008b; Lundström-Baudais
in press). Middle Neolithic (ca. 6,450–5,450 cal BP).
Few charred remains: einkorn wheat (prevailing);
emmer wheat (frequent); naked wheat (few); barley
(rare); pea (rare). (ii) Brig-Glis, Waldmatte
(Lundström-Baudais in Curdy et al. 1993). Early
Iron Age (2550–2450 uncal BP = ca. 2750–2450 cal
BP). Rich charred material from a burnt storehouse:
barley (frequent); broomcorn millet (frequent); fox-
tail millet (few); lentil (few); pea (few); bitter vetch
(frequent); faba bean (few). No wheat remains.
188 DOMESTICATION OF PLANTS IN THE OLD WORLD
Germany
(General references: Kreuz 1990; Knörzer 1991;
Küster 1991; Kreuz and Boenke 2002; Bogaard 2004;
Kreuz et al. 2005; Jacomet 2007, 2009; Bittmann pers.
comm.)
1. Hienheim, Kr. Kelheim (Bakels 1978).
Linearbandkeramik culture (6,155±45 to 5,910±50
uncal BP = ca. 7,150–6,650 cal BP). Rich remains:
einkorn wheat and emmer wheat (prevailing,
apparently in equal proportions); pea (frequent);
lentil (few); flax (rare). Wild: Corylus avellana;
numerous weeds.
2. Heilbronn, Württemberg, including Heilbronn-
Böckingen, H.-Gross-Gartach, H.-Willsbach (Bertsch
and Bertsch 1949); and H.-Klingenberg (Stika 1996a).
(i) Linearbandkeramik culture. Rich remains:
einkorn wheat (frequent); emmer wheat (frequent);
hulled six-rowed barley (frequent); naked six-rowed
barley (few); lentil (few); pea (few, except for being
very frequent in H.-Klingenberg); flax (rare). (ii)
Late Neolithic (ca. 4,750–4,650 uncal BP=ca. 5,600–
5,400 cal BP). Rich remains (both grain and chaff):
einkorn wheat (frequent); emmer wheat (frequent);
free-threshing wheat (very frequent); naked barley
(very frequent); hulled barley (few); lentil (few); pea
(very frequent); poppy (rare); flax (rare).
3. Aldenhovener Platte, Langweiler, Lamersdorf,
Bedburg-Garsdorf. Meckenheim and Rödingen
(Knörzer 1973, 1974, 1997). Linearbandkeramik cul-
ture (5,950±140 to 6,370±210 = ca. 7,500–6,550 cal
BP). Rich remains: einkorn wheat and emmer wheat
(frequent); pea (few); lentil (rare); flax (few; in one
place very frequent); poppy (few). Wild: Bromus sec-
alinus (abundant); Vicia sp.; Quercus sp.; Prunus insi-
titia; P. spinosa; Malus sp.; Corylus avellana; Sambucus
sp.
4. Kückhoven, Erkelenz, Rheinland (Knörzer
1998). Linearbandkeramik well (7,040 cal BP, den-
drodates). Rich remains, including chaff: einkorn
wheat (frequent); emmer wheat (prevailing); free-
threshing wheat (few); pea (rare); flax (frequent);
poppy (frequent). Wild: Corylus avellana; Malus syl-
vestris; Prunus spinosa; Rubus caesius; R. fruticosus;
Sambucus nigra, and numerous herbs.
5. Dresden-Nickern (Baumann and Schultze-
Motel 1968). Linearbandkeramik culture (5,945±100
and 5,815±100 uncal BP = ca. 6,900–6,500 cal BP).
Rich remains: emmer wheat (few); pea (a pure
hoard).
6. Dannau, Oldenburg (Kroll 1981b). Middle
Neolithic. Rich remains: einkorn wheat (rare);
emmer wheat (frequent); naked six-rowed barley
(prevailing). Wild: Corylus avellana; Rubus idaeus;
Sambucus nigra.
7. Kr. Ludwigsburg sites. (i) Bietigheim-
Bissingen (Piening 1989). Early Neolithic,
Linearbandkeramik culture (ca. 7,450 cal BP). Rich
remains: einkorn (prevailing); emmer wheat (fre-
quent); naked wheat (rare); barley (rare); pea (rare).
(ii) Eberdingen-Hochdorf (Küster 1983, 1985;
Breuning 1987). Middle Neolithic. Schussenried
group (ca. 6,150–ca. 5,750 uncal BP=ca.7,150–6,500
cal BP). Rich remains: einkorn wheat (almost as fre-
quent as naked barley); emmer wheat (few); free-
threshing wheat (few); naked six-rowed barley
(prevailing); pea (few); linseed (frequent); poppy
(frequent). Possibly cultivated: Brassica rapa;
Petroselinum crispum. Wild: Corylus avellana; Fragaria
vesca; Malus sp.; Sambucus.
8. Federseeried, Württemberg, Lake-shore set-
tlements. (i) Late Neolithic, Aichbühl-Schussenried
group (ca. 5,900 cal BP) (Blankenhorn and Hopf
1982). Very rich remains: einkorn wheat (few);
emmer wheat (frequent); free-threshing wheat (pre-
vailing); spelt wheat (few); hulled six-rowed barley
(few); naked six-rowed barley (rare); poppy (rare).
Wild: Corylus avellana; Fagus sylvatica; Malus sp.;
Fragaria vesca; Rubus idaeus; R. fruticosus. (ii)
Ödenahlen (Maier 1995). Late Neolithic (first half of
the fifth millennium BP). Rich remains, both charred
and waterlogged, including chaff: einkorn wheat
(few); emmer wheat (prevailing); free-threshing
wheat (frequent); barley (few); flax (frequent);
poppy (rare). Wild: fruits of Malus sp. cut to halves
or quarters (numerous); Fragaria vesca; Rubus
idaeus.
9. Lake Constancesites (i) Hilzingen (Stika
1991). Early Neolithic, Linearbandkeramik culture
(ca. 7,250–7,050 cal BP). Rich remains: einkorn
wheat (frequent); emmer wheat (frequent); barley
(rare); pea (prevailing, in storage); flax (rare). Wild:
Corylus avellana; Malus sylvestris; Prunus spinosa. (ii)
Hornstaad, lake-shore or peat bog settlements
(Maier 2001). Late Neolithic, Pfyn (Hornstaader Gr.)
culture (ca. 5,850–5,450 cal BP). Very rich water-

logged remains: einkorn wheat (frequent); emmer
wheat (few); naked wheat (prevailing); barley (fre-
quent); pea (few); flax (few); opium poppy (few).
Wild: Corylus avellana; Malus sylvestris; Prunus
spinosa; Fragaria vesca; Rosa sp.; Quercus sp.; Rubus
idaeus/R. fruticosus; Camelina sativa. Aromaric/oil
plants: Apium sp.; Anethum graveolens; Petroselinum
crispum; Descurainia sophia; Brassica rapa; Galeopsis
tetrahit-type.
10. Rosdorf, Lower Saxony (Kirleis and Willerding
2008) Linearbandkeramik culture (6,350±70 uncal BP=
ca. 7,450–7,150 cal BP). Einkorn wheat and emmer
wheat (co-prevailing); barley (rare); pea (rare); flax
(rare); opium poppy (rare). Wild: Corylus avellana.
The Netherlands
(General references: Van Zeist 1968–70; Bakels
1991)
1. Graetheide, South Limburg; as well as Sittard,
Beek, Elsloo, and Geleen (Bakels 1979, 2001; Bakels
and Rousselle 1985). Linearbandkeramik culture
(ca. 5,300–5,000 cal BC = ca. 7,250–6,950 cal BP).
Numerous remains: einkorn wheat (few); emmer
wheat (frequent); flax/linseed (rare); poppy (rare).
2. Maastricht-Randwijck, South Limburg
(Bakels et al. 1993). Rössen culture (ca. 4,900–4,600
cal BC = ca. 6,850–6,550 cal BP). Numerous remains
(both grains and chaff): einkorn wheat (few); emmer
wheat (few); free-threshing wheat (prevailing);
naked barely (frequent). Wild: Corylus avellana;
Prunus spinosa, and Rosa sp.
3. Swifterbant S3, Flevoland, Western coast, and
adjacent sites (van Zeist and Palfenier-Vegter 1981;
Cappers and Raemaekers 2008). Early Neolithic (ca.
5,000–3,500 cal BC = ca. 6,950–5,450 cal BP).
Numerous remains: einkorn wheat (few); emmer
wheat (rare); naked six-rowed and two-rowed bar-
ley (prevailing). Wild: Crataegus monogyna; Malus
sylvestris; Rubus fruticosus; Corylus avellana; Trapa
natans.
4. Aartswoud, North Holland (Pals 1984). Late
Neolithic (ca. 2,950–2,750 cal BC = ca. 4,900–4,700
cal BP). Rich remains: einkorn wheat (few); emmer
wheat (frequent); free-threshing wheat (few); naked
six-rowed barley (prevailing); linseed (few). Wild:
Malus sp.; Rubus fruticosus; Corylus avellana; Spergula
arvensis; Polygonum convolvulus; P. lapathifolium.
PLANT REMAINS IN REPRESENTATIVE ARCHAEOLOGICAL SITES 189
5. Twisk and Bovenkarspel, West Friesland
(Buurman 1987; Buurman et al. 1995). Middle and
Late Bronze Age (ca. 1,800–800 cal BC = ca. 3,750–
2,750 cal BP). Numerous remains (both grain and
chaff): emmer wheat (frequent); naked barley (pre-
vailing); hulled barley (frequent). Wild: numerous
weeds.
Belgium
(General references: Bakels 1991)
1. Aubechies, and other contemporary sites
(Bakels and Rousselle 1985). Linearbandkeramik
culture (ca. 5,400–4,900 cal BC = ca. 7,350–6,850 cal
BP). Numerous remains: einkorn wheat (frequent);
emmer wheat (prevailing); flax (rare). Wild: numer-
ous herbs.
2. Wange and Overhespen, near St Truiden
(Bakels 1992). Linearbandkeramik culture (ca.
5,400–5,000 cal BC = ca. 7,350–6,850 cal BP). Few
remains: einkorn wheat (few); emmer wheat (pre-
vailing); naked barley (few); pea (few). Wild: numer-
ous herbs.
3. ‘La Bosse de la Tombe’ à Givry (Heim 1979).
Rössen culture (ca. 4,900–4,700 cal BC = ca. 6,850–
6,650 cal BP). Scarce remains: emmer wheat (few);
free-threshing wheat (prevailing). Wild: Polygonum
hydropiper; Stellaria media; Corylus avellana.
Denmark
(General references: Jensen 1991; Robinson 1994,
2003, 2007)
1. Sarup, south-west Funen (Jørgensen 1981).
Funnel Beaker culture (ca. 4,350 uncal BP=4,950–
4,850 cal BP). Numerous charred remains: einkorn
wheat (rare); emmer wheat (prevailing); free-thresh-
ing wheat (rare); naked six-rowed barley (few). Wild:
Malus sylvestris; Corylus avellana; Chenopodium album.
2. Norsminde, Krabbesholm, and Visborg,
Jutland (Andersen 1991, 2000, 2005; Robinson 2007).
Earliest Funnel Beaker culture (ca. 5,850–5,650 cal
BP). Few charred remains: emmer wheat (few);
naked barley (few). Wild: Pyrus/Malus; Corylus avel-
lana; Glyceria plicata; range of arable weeds, ruderal,
and grassland species.
3. Brødrene Gram, Vojens, southern Jutland
(Robinson 2000, 2003). (i) Late Neolithic (4,410–
190 DOMESTICATION OF PLANTS IN THE OLD WORLD
4,150 uncal BP=ca. 5,050–4,600 cal. BP). Rich
charred remains: emmer wheat (frequent); spelt
wheat (few); naked barley (prevailing). Wild: ara-
ble weed species (few). (ii) Early Bronze Age (ca.
3850–3400 cal BP). Rich charred remains: emmer
wheat (few); spelt wheat (few); naked barley (pre-
vailing); hulled six-rowed barley (few). Wild:
range of arable weeds, ruderal, and grassland spe-
cies (abundant).
4. Enkehøj, Gilmosevej, Herning, central Jutland
(Møbjerg et al. 2007). Late Neolithic (ca. 3,950–3,650
cal BP). Rich charred remains: emmer wheat (fre-
quent); spelt wheat (few); naked barley (prevailing).
Wild: Quercus acorns (abundant); range of arable
weeds.
5. Lindebjerg and Voldtofte, Fyn Island (Rowley-
Conwy 1979, 1983). Middle-Late Bronze Age (3,150–
2,850 uncal BP = ca. 3400–2950 cal BP). Rich charred
remains: emmer wheat (rare); free-threshing wheat
(frequent); hulled barley (frequent); naked barley
(prevailing); broomcorn millet (few).
Sweden
(General references: Jensen 1991)
1. Settlements in Scania, southern Sweden
(Engelmark 1992; Larsson 1992). Early Neolithic
(4,925±115 and 4,915±100 uncal BP = ca. 5,900–
5,600 cal BP). Imprints: spelt-type wheats (pre-
vailing); naked barley (frequent); dwarf wheat
(few).
2. Settlements in Bjästamon, Ångermanland,
northern Sweden (Runesson 2007). Middle Neolithic
(3,798 ± 28 uncal BP = ca. 4,650–4,450 cal BP).
Hordeum vulgare.
3. Alvastra, Östergötland (Hjelmqvist 1955;
Göransson et al. 1995). Middle Neolithic culture
(ca. 5,000 cal BP). Numerous carbonized remains
and imprints: einkorn wheat (rare); hulled six-
rowed barley (rare); naked six-rowed barley
(prevailing). Wild: Malus sp. (fruits and pips);
Corylus avellana.
4. Eker, Hjulberga, Rosenland, Närke (Hjelmqvist
1979b). Neolithic culture. Scarce charred remains
and imprints: emmer wheat (rare); free-threshing
wheat (few); naked six-rowed barley (prevailing);
pea (rare). Wild: Corylus avellana; Sinapis arvensis.
Norway
1. Hjelle and Gossen, north-west Norway
(Soltvedt 2000; Hjelle and Solem 2008). Late
Neolithic (Hjelle: 3,760±70 uncal BP = ca. 4,250–
2,000 cal BP; Gossen: 3,72±45 uncal BP = ca. 4,250–
3,950 cal BC). Rich charred remains: naked barley
(prevailing, in Hjelle); hulled barley (frequent, in
Gossen).
2. Borge vestre, Østfold, south-east Norway
(Sandvik 2007, 2008). (i) Late Neolithic (3,680±50
uncal BP = ca. 4,150–3,900 cal BP). Charred grains of
barley. (ii) Bronze Age (2,900±30 uncal BP = ca.
3,150–2,950 cal BP). Charred seeds: barley; Panicum
miliaceum; flax.
3. Kvåle, south-western Norway (Soltvedt et al.
2007). Late Neolithic (3555±65 uncal BP = ca. 3,900–
3,700 cal BP). Charred remains: emmer wheat (fre-
quent); naked barley (frequent).
4. Forsandmoen, and three adjacent smaller sites
(Bakkevig 1982 1995). Bronze Age (ca. 2,900–2,800
cal BP). Rich charred remains: emmer wheat (rare);
free-threshing wheat (rare); naked barley (prevail-
ing, and the only identified cereal at two of the sites);
hulled barley (rare); oat (rare). Wild: Corylus avellana;
Rubus idaeus; Polygonum persicaria; P. tomentosum.
Finland
(General references: Jensen 1991; Häkkinen and
Lempiäinen 1996)
Niuskala, Turku, south-west Finland (Vuorela
and Lempiäinen 1988). Late Neolithic Kiukainen
culture (3,200±170 uncal BP = 3,600–3,250 cal BP).
Numerous remains of charred six-rowed naked
barley (recently confirmed by the presence of
Hordeum-type pollen grains).
Britain and Ireland
(General references: Moffett et al. 1989; Greig 1991;
Jones and Rowley-Conwy 2007)
1. Hambledon Hill, Dorset (Jones and Legge
1987, 2008). Early Neolithic (ca. 4,670±80 uncal BP =
ca. 5,600–5,050 cal BP). Rich charred remains: emmer
wheat, mainly chaff (prevailing); barley, including
six-row naked barley (frequent); and a single grape
pip. Wild: Corylus avellana.

2. Balbridie, Kincardineshire, Grampian, Scotland
(Fairweather and Ralston 1993). Neolithic (ca. 4,82±80
uncal BP = ca. 5,700–5,300 cal BP). Numerous charred
remains, apparently storage: emmer wheat (prevail-
ing); free-threshing wheat (few, but a pure hoard was
found in a posthole); naked barley (frequent); hulled
barley (rare); flax (few).
3. The Stumble, Blackwater Estuary, Essex
(Murphy 1989). Early Neolithic (ca. 4,675±70 uncal
BP = ca. 5,600–5,300 cal BP). Rich, submerged,
charred remains: einkorn wheat (few); emmer
wheat (prevailing); naked barley (few). Wild:
Corylus avellana; Prunus spinosa; Crataegus monogyna;
Rubus sp.; Malus sp.; Tilia sp.; Vicia/Lathyrus sp.;
Rumex sp.; Polygonum sp.; Galium aparine.
4. Tankardstown, Ireland (Monk 1988). Early
Neolithic (4,840±80 uncal BP = ca. 5,650–5,450 cal
BP). Numerous charred remains: emmer wheat
(prevailing). Wild: Malus sylvestris; Corylus avellana.
France
(General references: Marinval 1988; Bakels 1991;
Hopf 1991a; Ruas and Marinval 1991; Bouby 2003;
Zech-Matterne et al. 2009).
1. La Baume Fontbrégoua, Salernes, Var (Courtin
and Erroux 1974). (i) Mesolithic. Few remains: Wild:
Vicia sp.; Vitis sylvestris. (ii) Impressed Ware
(Cardial) culture (6,180±12 uncal BP; 5,690±130
uncal BP = ca. 7,250–6,250 cal BP). Scarce remains:
free-threshing wheat (prevailing). Wild: Lathyrus cf.
cicera; Vicia sp.; Lathyrus cf. cicera; Quercus sp. (iii)
Middle Neolithic (5680±12 uncal BP; 5,330±12 uncal
BP = ca. 6,800–5,800 cal BP). Rich remains: einkorn
wheat (few); emmer wheat (prevailing); free-thresh-
ing wheat (frequent); six-rowed barley (few); naked
six-rowed barley (few). Wild: Vicia sp.; Lathyrus cf.
cicera.
2. Pont de Roque Haute, Portiragnes, Hérault
(Marinval 2007). Early Neolithic, Impressa culture
(6,850±65 uncal BP; 6,745±70 uncal BP= ca. 7,900–
7,500 cal BP). Few carbonized remains: einkorn
wheat (few); emmer wheat (few); free-threshing
wheat (rare).
3. Châteauneuf-les-Martigues, Font des Pigeons,
Bouches-du-Rhône (Courtin et al. 1976). Impressed
Ware (Cardial) culture (6,550±100 uncal BP;
6,050±100 uncal BP = ca. 7650–6,700 cal BP). Rich
PLANT REMAINS IN REPRESENTATIVE ARCHAEOLOGICAL SITES 191
remains: free-threshing wheat and naked six-rowed
barley in equal proportions. Wild: Prunus cerasus;
Pinus sp.
4. Menneville and several other sites in the Valley
of the Aisne (Bakels 1984, 1991; Ilett et al. 1995).
Numerous charred remains. (i) Linearbandkeramik
culture (6,200±190 uncal BP = ca. 7,550–6,700 cal BP):
einkorn wheat (few); emmer wheat (prevailing);
naked barley (frequent); pea (rare); lentil (rare).
Wild: Corylus avellana; Galium aparine; Bromus sp.;
Echinochloa crus-galli; Lapsana communis; Polygonum
convolvulus, Rumex sanguinea. (ii) Middle Neolithic,
Michelsberg culture: emmer wheat (rare); hulled
barley (few); a large number of poppy seeds.
5. Bercy (Paris) (Dietsch 1996, 2001). Middle
Neolithic, Chasséen culture (5,130±80 uncal BP;
4,530±12 uncal BP = ca. 6,250–4,900 cal BP). Rich
waterlogged and few carbonized remains: emmer
wheat (few); free-threshing wheat (frequent); six-
rowed barley (few). Wild: Corylus avellana; Vitis syl-
vestris; Cornus sanguinea; Rubus fruticosus; Quercus
sp.; Crataegus monogyna; Rubus idaeus; Prunus
spinosa; Sambucus spp.; Malus sylvestris.
6. Grotte G, Baudinard, Var (Courtin and Erroux
1974). Middle Neolithic, Chasséen culture. Numerous
remains: emmer wheat (frequent); free-threshing
wheat (frequent); six-rowed barley (prevailing);
chickpea-like seed (rare); faba bean (rare). Wild:
Lathyrus cicera; Vicia sp.; Quercus sp.
7. Station III de Clairvaux, Jura (Lundström-
Baudais 1984). Late Neolithic (4,412±56 uncal BP;
4,272±70 uncal BP = ca. 5,350–4,650 cal BP) lakeside
settlement. Numerous mostly waterlogged remains
of threshing and of wild plants: emmer wheat (fre-
quent); free-threshing wheat (few); six-rowed bar-
ley (prevailing); flax: seed, capsules, and textile
(few); pea (rare); poppy (few). Wild: numerous spe-
cies including Corylus avellana; Malus sylvestris;
Prumus spinosa; Rubus idaeus; R. fruticosus; Fragaria
vesca.
8. Grésine, Brison Saint Innocent, Savoie (Bouby
and Billaud 2001). Late Bronze Age (Dendrochrono-
logy 905–869 BC). Rich waterlogged and carbonized
remains: einkorn wheat (few); emmer wheat (fre-
quent); spelt wheat (frequent); free-threshing wheat
(few); hulled six-rowed barley (frequent); broom-
corn millet (frequent); foxtail millet (prevailing);
pea (rare); lentil (rare); faba bean (rare); grass pea
192 DOMESTICATION OF PLANTS IN THE OLD WORLD
(rare); bitter vetch (rare); poppy (frequent); linseed,
including capsules (frequent); gold of pleasure,
including pods (frequent). Wild: Avena sp.; Brassica
campestris; Vitis sylvestris; Quercus sp.; Corylus avel-
lana; Fragaria vesca; Prunus spinosa; Rosa sp.; Rubus
spp.; Sambucus spp.
Spain
(General references: Hopf 1991a; Zapata et al. 2004;
Buxó and Piqué 2008)
1. Balma Margineda, Andorra (Marinval 1995).
(i) Mesolithic (9,250±160 uncal BP= ca. 10,650–10,250
cal BP). Wild: Prunus spinosa; Corylus avellana;
Quercus sp.; Juglans regia; Pinus pinea. (ii) Early
Neolithic (6,850±160 uncal BP = ca. 7,950–5,450 cal
BP). Rich remains: emmer wheat (few); free-thresh-
ing wheat (frequent); compact type of free-thresh-
ing wheat (rare); naked six-rowed barley (few); pea
(few). Wild: Quercus sp.; Corylus sp.
2. Coveta de l’Or, Beniarrés, Alicante (Hopf and
Schubart 1965; Lopez 1980), and adjacent Cova de
Cendres (Buxó, 1997). Early Neolithic Impressed
Ware (Cardial) culture (6,340±70 uncal BP = ca.
7,400–7,050 cal BP). Rich remains: einkorn wheat
(few); emmer wheat (frequent); free-threshing
wheat (frequent); hulled six-rowed barley (fre-
quent); naked six-rowed barley (prevailing); lentil
(rare); pea (rare); grass pea (rare); faba bean (rare).
Wild: Quercus sp.; Olea europaea var. oleaster.
3. Sites of south-east Spain: (i) Cueva de los
Murciélagos, Zuheros, Córdoba (Hopf and Muñoz
1974; Peña-Chocarro 1999). Early Neolithic
(6,430±130 to 5,660±12 uncal BP = ca. 7,500–5,660
cal BP). Rich remains: emmer wheat (frequent); free-
threshing wheat (prevailing); hulled six-rowed bar-
ley (frequent); naked six-rowed barley (frequent);
Papaver somniferum seeds. (ii) Cueva del Toro, El
Torcal, Antequera, Málaga (Buxó 2004; Hopf unpub-
lished data). Middle Neolithic (5,380±45 uncal BP =
ca. 6,300–6,000 cal BP). Rich remains: emmer wheat
(few); free-threshing wheat (prevailing); hulled six-
rowed barley (few); naked six-rowed barley (fre-
quent); lentil (few); grass pea (few); pea (few); faba
bean (frequent); Papaver somniferum seeds. Wild:
Quercus acorns; Celtis australis. (iii) Cueva de Nerja,
Málaga (Hopf and Pellicer Catalán 1970). Late neo-
lithic (5,065±140 uncal BP = ca. 6,200–5,500 cal BP).
Rich remains: free-threshing wheat (frequent);
hulled six-rowed barley (frequent). Wild: Quercus
sp.; Olea europaea.
4. Cova de Can Sadurní, Begues, Barcelona
(Blasco et al. 1999). (i) Early Neolithic Impressed
Ware (Cardial), Late Cardial/Epicardial culture
(6405±55 uncal BP to 6,050±110 uncal BP = ca. 7,400–
6,650 cal BP). Rich remains: einkorn wheat (fre-
quent); emmer wheat (frequent); free-threshing
wheat (frequent); hulled and naked barley (fre-
quent). Wild: Pistacia lentiscus. (ii) Post-Cardial
phases (5,700±110 uncal BP = ca. 6,850–5,800 cal
BP). Rich remains: einkorn wheat (few); emmer
wheat (frequent); free-threshing wheat (frequent);
hulled and naked barley (frequent); pea; lentil; grass
pea. Wild: Pistacia lentiscus; Vitis vinifera var. sylves-
tris; Ficus carica; Quercus sp.; Arbutus unedo; Vicia
sativa.
5. North Meseta sites including Cueva de La
Vaquera, La Lámpara, and Revilla del Campo
(López et al. 2003; Stika 2005; Peña-Chocarro 2007).
Early Neolithic (6,12±160 uncal BP = ca. 7,400–
6,650 cal BP). Rich remains: einkorn wheat (rare);
emmer wheat (frequent); free-threshing wheat
(rare); hulled and naked barley (frequent); Lens
sp.; Vicia sativa; Vicia sp. Possibly cultivated: Linum
usitatissimum; Papaver somniferum/setigerum. Wild:
Vitis vinifera spp. sylvestris; Quercus sp. Weeds from
arable fields.
6. Kobaederra, Bizkaia, (Zapata 2002) and El
Mirón, Cantabria (Peña-Chocarro et al. 2005). Early
Neolithic (5,500±90 uncal BP to 5,375±90 uncal BP =
6,450–5,950 cal BP). Scarce charred remains: einkorn
wheat (few); emmer wheat (few); naked wheat
(few); barley (few). Wild: Quercus subg. Quercus;
Corylus sp.; Rosaceae pomes.
7. Cerro de la Virgen, Orce, Galera, Granada
(Buxó 1997). Chalcolitic Period (5,250–4,350 cal BP).
(i) Pre-Bell Beaker culture. Few remains: free-
threshing wheat (few); hulled barley (few); naked
barley (prevailing); faba bean (rare). Wild: rhizome
of Stipa tenacissima. (ii) Bell Beaker culture. Rich
remains: emmer wheat (few); free-threshing wheat
(frequent); compact type of free-threshing wheat
(rare); hulled barley (few); naked barley (prevail-
ing); faba bean (few); pea (rare).
8. El Argar culture sites, south-east Spain (Stika
1988; Hopf 1991b). Early Bronze Age (ca. 4,150–3,350

cal BP). Rich remains: einkorn wheat (few); emmer
wheat (few); free-threshing wheat (frequent); hulled
six-rowed barley (prevailing); naked six-rowed bar-
ley (frequent); pea (few); faba bean (few); flax, seed,
and fibers (frequent). Wild: Olea europaea; Vitis vinif-
era; Ficus carica; Pistacia sp.; Quercus acorns; Celtis
australis; Stipa tenacissima.
Portugal
(General references: Pinto da Silva 1988; Hopf
1991)
1. Buraco da Pala, Bragança (Ramil Rego and Aira
Rodríguez 1993). (i) Late Neolithic (ca. 5,500–4,800
cal BP). Rich remains: free-threshing wheat (fre-
quent); naked six-rowed barley (prevailing); hulled
six-rowed barley (rare); faba bean (rare); pea (rare).
Wild: Quercus sp. acorns. (ii) Chalcolithic (ca. 4,800–
4200 cal BP). Rich remains: free-threshing wheat (fre-
quent); naked six-rowed barley (prevailing); hulled
six-rowed barley (few); pea (rare); lentil (rare); faba
PLANT REMAINS IN REPRESENTATIVE ARCHAEOLOGICAL SITES 193
bean (few); flax (few); Papaver seeds (few). Wild: Vitis;
Quercus sp. acorns; Pinus pinaster, kernel.
2. Zambujal, Torres Vedras (Ramil Rego and Aira
Rodríguez 1993). (i) Chalcolithic (ca. 5,400–4,200 cal
BP). Scarce remains: einkorn/emmer wheat (rare);
free-threshing wheat (few); hulled and naked six-
rowed barley (few); faba bean (prevailing); olive
(wild? few). Wild: Pinus sp.; Quercus sp. (ii) Bronze
Age (ca. 4,000–3,600 cal BP). Numerous remains;
einkorn/emmer wheat (rare); free-threshing wheat
(frequent); hulled and naked six-rowed barley (fre-
quent); faba bean (prevailing); flax (rare); olive
(wild? frequent). Wild: Pinus sp.; Quercus sp. acorns;
cork of Quercus suber.
3. Crasto de Palheiros, Murça, Vila Real (Figueiral
and Sanches 2003). Chalcolithic (ca. 4,800–4,200 cal
BP). Scarce remains, including chaff: emmer/spelt
wheat (prevailing); spelt wheat (few); hulled and
naked six-rowed barley (few); broomcorn millet
(few); faba bean (rare). Wild: Arbutus unedo; Juniperus
sp.; Olea europaea.
 Appendix A: Site orientation maps

Chokh

Imiris-Gora

Arukhlo

Aratashen and Aknashen

Asikli Höyük Çayönü

Erbaba Cafer Höyük
Hacilar Girikihaciyan
Çatalhöyük Can Hasan
Yarym
Jerf el Ahmar Tepe
Mureybit
Andreas Kastros
Agridhi Tell Abu Jarmo
Kissonerga-Mylouthkia Hureyra
Shillourokambos Khirokitia
Tell es-Sawwan
Tell Ramad Tell Aswad

Ohalo Choga
Mami
Netiv Hagdud-Gilgal Ali Kosh
Jericho Ghassul
'Ain Ghazal
Tepe Sabz
Bab Edh-Dhra
Merimde
Saqqara
Fayum

0 100 200 miles

0 200 400 km

Map 21 South-west Asia, showing the location of the main archaeological sites mentioned in this book.

194
 Mohelnice
Bietigheim-
Heilbronn Bissingen
Hilzingen Schletz Kukoneshti
Füzesabony-
Zürich Sion Mondsee Gubakút
Sacarovca
Egolzwil Zánka- Szilhalom
Monte Vasútállomás Dévaványa
Còvolo Poduri
Sammardenchia and Szeged-Gyálarét,
Friuli district sites Röszke-Lúdvár and
Battonya-Basarága
Starcevo Liubcova
Gomolava
Cîrcea
Obre
Pienza Pokrovnik Sucidava-Delei
Karanovo Azmaška Mogila
Kapitan
La Marmotta Dimitrievo
Vršnik Kovacevo
Scamuso Anza
Kastanas
Nea
Nikomedeia
Dimini Sesklo
Grotta
dell’Uzzo Balomenou

Lerna
Franchthi
Cave

Knossos
0 100 200 miles
0 200 400 km

Map 22 South and south-east Europe, showing the location of the main archaeological sites mentioned in this book.

Niuskala

Borge vestre
Rogaland
Balbridie
Alvastra
Norsminde,
Krabbesholm
and Visborg
Tankardstown Scania region
Lindebjerg
Sarup Eker
The Stumble
Twisk Danau
Aartswoud
Hambledon Hill Swifterbant S3
Brześć
Kückhoven Rosdorf
Graetheide Gniechowice and
Aubechies
Wange Dresden Stary Zamek
Aldenhovener Ćmielów
Menneville and Platte Gniechowice Strachów
Aisne Valley LBK sites Nowa Huta-Mogila
HeilbronnBylany
Mohelnice Gw oździec
Bietigheim-Bissingen
Hilzingen Hienheim Schletz
0 100 200 miles Federseeried Nitriansky Hrádok
Egolzwil 3 Zürich Blatné Füzesabony-Gubakút
0 200 400 km

Map 23 Central and north Europe, showing the location of the main archaeological sites mentioned in this book.
196 APPENDIX A

Aubechies

Bietigheim-
Menneville and Aisne Heilbronn Bissingen
Valley LBK site
Hilzingen

Zürich Egolzwil 3

Sion

Châteauneuf-
les-Martigues
El Mirón
Kobaederra Pont de Roque Haute

Buraco da Pala North Meseta sites Balma Margineda

Cova de Can Sadurní

Cueva de Vaquera
La Lámpara&
Revilla del Campo
Pedra de Ouro
Villa Nova de S. Pedro
Zambujal Coveta de l’Or
Cova de Cendres
Cueva de los
Murciélagos
Cerro de la Virgen
Cueva del Toro El Argar culture sites
Cueva de Nerja
0 100 200 miles
Kaf Taht El-Ghar
0 200 400 km

Map 24 France, Spain, and Portugal, showing the location of the main archaeological sites mentioned in this book.
Appendix B: Chronological chart for
the main geographical regions
mentioned in the book

197
 South-west West
BP Aegean belt Balkan Egypt BC
Asia Mediterranean
2,000 0
Iron Age

Iron Age Iron Age Late

Bronze Age
3,000 New
1,000
Late Middle/Late Kingdom
Bronze Age Bronze Age Early/Middle Early/Middle
(Minoan, Bronze Age Bronze Age
Mycenaean) (Otoman) (El Argar)
Middle
4,000 2,000
Middle Early Kingdom
Bronze Age Bronze Age

Old
Kingdom
5,000 Early Final 3,000
Chalcolithic
Bronze Age Neolithic
Eneolithic
(Gumelnitsa,
Vanča)
Archaic
6,000 4,000
Late Middle/Late
Neolithic Neolithic
Chalcolithic

7,000 Neolithic 5,000

Middle Early
Neolithic Early Neolithic Neolithic
(Karanovo, (Impressed
Starčevo) ware)

8,000 Pottery 6,000

(Ceramic) Early
Neolithic Neolithic

9,000
thic) 7,000
i
Pre-Pottery esol
Neolithic B (M
(PPNB)
tures
10,000 l 8,000
cu
ming
r
e - fa
Pre-Pottery Pr
11,000 Neolithic A 9,000
(PPNA)

Fig. 41 Chronological chart for the main geographical regions mentioned in the book. The broken line in each column indicates when definite
signs of agriculture start to appear. BP (Before Present) indicates years before 1950, the year 0 of conventional radiocarbon dating.
 BP Central Alpine Scandinavia BC
Ukraine
Europe Belt
2,000 0
Iron Age

Iron Age Iron Age

3,000 Bronze Age 1,000

Bronze Age Bronze Age Bronze Age

Late
Neolithic
4,000 2,000

Early Neolithic
Late (Funnel
Middle/Late Neolithic Beaker)
5,000 Neolithic 3,000
(Rössen,
Lengyel, Eneolithic
Funnel Beaker) (Tripolye)
Early
Neolithic
6,000 Early Early Lake Shore 4,000
Neolithic Neolithic Settlements
(Linear
Bandkeramik)

7,000 5,000

8,000 6,000

)
ithic
l
9,000 eso 7,000
(M
es
ltur
g cu
in
10,000
farm 8,000
e -
Pr

11,000 9,000

Fig. 41 Continued.
 Appendix C: Information on
archaeological sites which appear
on Map 2
1: Jeitun (Charles and Hillman 1992; Harris et al. 1993, 1996).
2: Ali Kosh (Helbaek 1969). 3: Jarmo (Helbaek 1959b, 1960,
1966a; Braidwood 1960). 4: Chokh (Lisitsina 1984). 5:
Aratashen and Aknashen (Hovsepyan and Willcox 2008). 6:
Arukhlo I and II (Januševič 1984; Lisitsina 1984; Schultze-
Motel 1988). 7: Çayönü (Van Zeist 1972, 143–66; Van Zeist
and De Roller 1991–2, 2003). 8: Aşikli Höyük (van Zeist and
de Roller 1995). 9: Tell Abu Hureyra (Hillman 1975, 1989,
2000a; De Moulins 1997, 2000; Hillman et al. 2001). 10: Tell
Aswad (van Zeist and Bakker-Heeres 1985). 11: Ain Ghazal
(Rollefson et al. 1985). 12: Jericho (Hopf 1983). 13: Yiftah’el
(Garfinkel et al. 1988). 14: Kissonerga-Mylouthkia and
Shillourokambos (Willcox 2000; Murray 2003). 15: Fayum
(Caton-Thompson and Gardner 1934; Wetterstrom 1993;
Wendrich and Cappers 2005). 16: Merimde (Wetterstrom
1993). 17: Franchthi Cave (Hansen 1991a, 1992). 18: Nea
Nikomdeia (van Zeist and Bottema 1971). 19: Sesklo
(Hopf 1962; Kroll 1981a). 2: Knossos (Sarpaki 2009). 21:
Rivne (Pashkevich 2003). 22: Sacarovca (Januševič 1984;
Kuzminova et al. 1998). 23: Poduri (Cârciumaru and Monah
1985; Monah and Monah 2008). 24: Liubcova (Cârciumaru
1996). 25: Cîrcea (Cârciumaru 1996). 26: Kovacevo (Popova
1992; Marinova 2006). 27: Kapitan Dimitrievo (Marinova
2006, forthcoming). 28: Karanovo (Thanheiser 1997;
Marinova 2004, 2006). 29: Azmaška Mogila (Hopf 1973a;
Renfrew 1979, Table 7). 30: Anza (Renfrew 1976). 31: Obre
(Renfrew 1974). 32: Szeged-Gyálarét, Röszke-Lúdvár and
Battonya-Basarága (Hartyányi et al. 1968; Hartyányi and
Nováki 1971, 1975; Füzes 1990). 33: Füzesabony-Gubakút
(Gyulai 2007). 34: Zánka-Vasútállomás (Füzes 1990, 1991).
35: Bylany (Tempír 1979). 36: Mohelnice (Opravil 1979,
1981; Kühn 1981). 37: Blatné (Hajnalová 1989). 38:
Gniechowice and Stary Zamek (Gluza 1994). 39: Strachów
(Lityńska-Zając 1997). 40: Brześć (Bieniek 2007). 41:
Gwoździec (Bieniek and Litynska-Zajac 2001; Lityńska-
Zając 2007). 42: Pienza (Castelletti 1976). 43: La Marmotta
(Rottoli 1993, 2002). 44: Sammardenchia and Friuli district
200
sites (Pessina and Rottoli 1996; Rottoli 2005; Rottoli and
Pessina 2007). 45: Scamuso (Costantini et al. 1997). 46: Grotta
dell’Uzzo (Costantini 1989). 47: Schletz (Kohler-Schneider
2007; Schneider 1994). 48: Egolzwil 3 (Bollinger 1994;
Jacomet 2007). 49: Zürich (Jacomet 1988, 2004; Jacomet et al.
1989; Brombacher and Jacomet 1997; Favre 2002; Brombacher
et al. 2005). 50: Sion (Martin et al. 2008b; Lundström-Baudais
in press). 51: Hilzingen (Stika 1991). 52: Heilbronn (Bertsch
and Bertsch 1949; Stika 1996a). 53: Rosdorf (Kirleis and
Willerding 2008). 54: Hienheim (Bakels 1978). 55: Bietigheim-
Bissingen (Piening 1989). 56: Kückhoven (Knörzer 1998). 57:
Aldenhovener Platte (Knörzer 1973, 1974, 1997). 58:
Niuskala (Vuorela and Lempiäinen 1988). 59: Scania region
(Engelmark 1992; Larsson 1992). 60: Bjästamon (Runesson
2007). 61: Hjelle and Gossen (Soltvedt 2000; Hjelle and
Solem 2008). 62: Borge vestre (Sandvik 2007, 2008). 63:
Norsminde, Krabbesholm and Visborg (Andersen 1991,
2000, 2005; Robinson 2007). 64: Pont de Roque Haute
(Marinval 2007). 65: Graetheide (Bakels 1979, 2001; Bakels
and Rousselle 1985). 66: Swifterbant S3 (van Zeist and
Palfenier-Vegter 1981; Cappers and Raemaekers 2008). 67:
Wange and Overhespen (Bakels 1992). 68: Aubechies (Bakels
and Rousselle 1985). 69: The Stumble (Murphy 1989). 70:
Hambledon Hil (Jones and Legge 1987, 2008). 71: Balbridie
(Fairweather and Ralston 1993). 72: Tankardstown (Monk
1988). 73: Menneville and Aisne Valley LBK sites (Bakels
1984, 1991; Ilett et al. 1995). 74: Châteauneuf-les-Martigues
(Courtin et al. 1976). 75: Balma Margineda (Marinval 1995).
76: Coveta de l’Or and Cova de Cendres (Hopf and Schubart
1965; Lopez 1980; Buxó 1997). 77: Cova de Can Sadurní
(Blasco et al. 1999). 78: North Meseta sites ( López et al. 2003;
Stika 2005; Peña-Chocarro 2007). 79: Cueva de los
Murciélagos (Hopf and Muñoz 1974; Peña-Chocarro 1999).
80: Kaf Taht El-Ghar (Ballouche and Marinval 2003). 81:
Kobaederra and El Mirón (Zapata 2002; Peña-Chocarro et
al. 2005). 82: Buraco da Pala (Ramil Rego and Aira Rodríguez
1993). 83: Zinchecra (van der Veen 1992a, 1992b).
 References
Aaronsohn, A. 1910. Agricultural and botanical explorations
in Palestine. Washington DC, United States Deptartment
of Agriculture, Bureau of Plant Industry, Bulletin No.
180.
Abbo, S., Shtienberg, D., Lichtenzveig, J., Lev-Yadun, S.
and Gopher, A. 2003. The chickpea, summer cropping,
and a new model for pulse domestication in the ancient
Near East. Quarterly Review of Biology, 78, 435–48.
Adams, R. and Simmons, D. 1996. Archaeobotanical
remains. In: Frankel, D. and Webb, J. M. (eds.) Marki
Alonia, an early and middle Bronze Age town in Cyprus,
excavations 1990–1994. pp. 223–6, 321–332 (tables).
Jonsered, Paul Åström.
Akeret, Ö. 2005. Plant remains from a Bell Baker site in
Switzerland, and the beginnings of Triticum spelta
(spelt) cultivation in Europe. Vegetation History and
Archaeobotany, 14, 279–86.
Allaby, R. 2010. Integrating the processes in the evolution-
ary system of domestication. Journal of Experimental
Botany, 61, 935–44.
Allaby, R., Peterson, G., Merriwether, D. and Fu, Y.-B. 2005.
Evidence of the domestication history of flax (Linum usi-
tatissimum L.) from genetic diversity of the sad2 locus.
Theoretical and Applied Genetics, 112, 58–65.
Alleweldt, G., Spiegel-Roy, P. and Reisch, B. 1991. Grapes
(Vitis). In: Moore, J. N. and Ballington, J. R. (eds.) Genetic
resources of temperate fruit and nut crops. Pp. 291–327.
Wageningen, The Netherlands International Society for
Horticultural Sciences.
Amar, Z. 2000. Agricultural produce in the Land of Israel dur-
ing the Middle Ages. Jerusalem, Yad Izhak Ben-Zvi (in
Hebrew).
Ammann, B., Bollinger, T., Jacomet, S., Liese-Kleiber, H.
and Piening, U. 1981. Die Neolithischen ufersiedlungen von
Twann, Bd. 14: botanische untersuchungen. Bern, Staatlicher
Lehrmittelverlag.
Aradhya, M., Stover, E., Velasco, D. and Koehmstedt, A.
2010. Genetic structure and differentiation in cultivated
fig (Ficus carica L.). Genetica, 138, 681–94.
Araus, J. L., Ferrio, J. P., Buxo, R. and Voltas, J. 2007. The
historical perspective of dryland agriculture: lessons
learned from 10,000 years of wheat cultivation. Journal
of Experimental Botany, 58, 131–45.
Baas, J. 1980. Bedeutsame kultur- und nutzpflanzen aus
einer Römischen grube. Natur und Museum, 110,
257–62.
Badler, V. R. 1995. The archaeological evidence for wine
making, distribution and consumption at protohistoric
Godin Tepe, Iran. In: McGovern, P. E., Fleming, S. T. and
Katz, S. H. (eds.) The origins and ancient history of wine.
Pp. 45–65. Amsterdam, Gordon and Breach.
Badler, V. R., McGovern, P. E. and Michel, R. H. 1990.
Drink and be merry! Infrared spectroscopy and ancient
Near Eastern wine. In: Biers, W. R and McGovern, P. E.
(eds.) Organic contents of ancient vessels: materials
analysis and archaeological investigation. Philadelphia,
University of Pennsylvania Museum. Museum Applied
Science Center for Archaeology (MASCA) Research Papers,
7, 25–36.
Badr, A., Müller, K., Schäfer-Pregl, R., Rabey, H. E. L.,
Effgen, S., Ibrahim, H. H., Pozzi, C., Rohde, W. and
Salamini, F. 2000. On the origin and domestication his-
tory of barley (Hordeum vulgare). Molecular Biology and
Evolution, 17, 499–510.
Bailey, C. H. and Hough, L. F. 1975. Apricots. In: Janick, J.
and Moore, J. N. (eds.) Advances in fruit breeding. Pp.
367–83. West Lafayette, Indiana: Purdue University
Press.
Baillie, M. G. L. 1995. A slice through time: dendrochronology
and precision dating. London, Routledge.
Bakels, C. C. 1978. Four Linearbandkeramik settlements
and their environment, a paleoecological study of
Sittard, Stein, Elsloo, and Hienheim. Analecta Praehistorica
Leidensia, 11. Leiden, Leiden University Press.
Bakels, C. C. 1982. Der Mohn, die Linearbandkeramik,
und das westliche Mittelmeergebiet. Archäologisches
Korrespondenzblatt, 12, 11–13.
Bakels, C. C. 1984. Carbonized seeds from Northern
France. Analecta Praehistorica Leidensia, 17, 1–27.
Bakels, C. C. 1991. Western continental Europe. In: van Zeist,
W., Wasylikowa, K. and Behre, K.-E. (eds.) Progress in Old
World palaeoethnobotany. Pp. 279–98. Rotterdam, Balkema.
201
202 REFERENCES
Bakels, C. C. 1992. The botanical shadow of two early
Neolithic settlements in Belgium: carbonized seeds and
disturbances in pollen record. In: Pals, J. P., Buurman, J.
and van der Veen, M. (eds.) Festschrift for Professor
W. van Zeist. Pp. 1–19. Review of Palaeobotany and
Palynology, 73, 1–19.
Bakels, C. C. 1996. Fruits and seeds from Linearbandkeramik
settlement at Meindling, Germany, with special reference
to Papaver somniferum. Analecta Praehistorica Leidensia, 25,
55–68.
Bakels, C. C. 2003. Report concerning the contents of a
ceramic vessel found in the ‘white room’ of the Gonur
Temenos, Merv Oasis, Turkmenistan. Electronic Journal
of Vedic Studies (EJVS), 9:1c. (http://www.ejvs.laurasi-
anacademy.com/ejvs0901/ejvs0901c.txt)
Bakels, C. C. and Jacomet, S. 2003. Access to luxury foods
in Central Europe during the Roman period: the
archaeobotanical evidence. World Archaeology, 34,
542–57.
Bakels, C. C. and Rousselle, R. 1985. Restes botaniques et
agriculture du Néolithique ancien en Belgique et aux
Pays-Bas. Helinium, 25, 37–57.
Bakels, C. C., Alkemade, M. J. and Vermeeren, C. E. 1993.
Botanische untersuchungen in der Rössener siedlung
Maastricht- Randwijck. In: Kalis, A. J. and Meurers-
Balke, J. (eds.) 7000 jahre bäuerliche landschaft: entste-
hung, erforschung, erhaltung. Bonn, Rheinland Verlag.
Archaeo-Physica, 13, 35–48.
Bakhteyev, F. K. and Yanushevich, Z. V. 1980. Discoveries
of cultivated plants in the early farming settlements of
Yarym-Tepe I and Yarym-Tepe II in northern Iraq.
Journal of Archaeological Science, 7, 167–78.
Bakkevig, S. 1982. Økologi og økonomi for deler av Sør-
Jaeren i senneolitikum. Del 2. Makrofossil analyse.
Saltvannsflotasjon av materiale fra Rugland på Jaeren.
AmS-Skrifter, 9, 33–40.
Bakkevig, S. 1995. Neue makrofossile Beiträge zur Kenntnis
des vorhistorischen Getreideanbaus in Südwestnorwegen.
In: Kroll, H. and Pasternak, R. (eds.) Res Archaeobotanicae,
International Workgroup for Palaeoethnobotany, Proceedings
of the 9th Symposium. Pp. 5–15. Kiel, Oetker-Voges.
Ballouche, A. and Marinval, P. 2003. Données paly-
nologiques et carpologiques sur la domestication des
plantes et l’agriculture dans le néolithique ancien du
Maroc septentrional (site de Kaf Taht El-Ghar). Revue
d’Archéométrie, 27, 49–54.
Bar-Adon, P. 1980. The cave of the treasure. The finds from the
caves in Nahal Mishmar. Jerusalem, Israel Exploration
Society.
Barakat, H. and Fahmy, A. G. E.-D. 1999. Wild grasses as
‘Neolithic’ food in the eastern Sahara, a review of the
evidence from Egypt. In: van der Veen, M. (ed.) The
Exploitation of Plant Resources in Ancient Africa. Pp. 33–46.
New York, Kluwer-Plenum Press.
Barrow, S. C. 1998. A monograph of Phoenix L. (Palmae:
Coryphoideae). Kew Bulletin, 53, 513–75.
Baruch, U. 1990. Palynological evidence of human impact
on the vegetation as recorded in Late Holocene lake
sediments in Israel. In: Bottema, S., Entjes-Nieborg, G.
and van Zeist, W. (eds.) Man’s role in the shaping of the
eastern Mediterranean landscape. Pp. 283–93. Rotterdam,
Balkema.
Batlle, I. and Tous, J. 1997. Carob tree Ceratonia siliqua L.
Promoting the conservation and use of underutilized and
neglected crops, 17. Rome, International Plant Genetic
Resources Institute.
Baumann, W. and Schultze-Motel, J. 1968. Neolithische kul-
turpflanzenreste aus Sachsen. Arbeits- und Forschungsberichte
der sächsischen Bodendenk-malpflege, 18, 9–28.
Beck, N. G. and Lord, E. M. 1988. Breeding system in Ficus
carica, the common fig. I. Floral diversity. American
Journal of Botany, 75, 1904–12.
Bedigian, D. 1985. Is še-giš-í sesame or flax? Bulletin on
Sumerian agriculture, 2, 159–78.
Bedigian, D. 1998. Early history of sesame cultivation in
the Near East and beyond. In: Damania, A. B., Valkoun, J.
and Qualset, C. O. (eds.) Origins of agriculture and crop
domestication. Pp. 93–101. Aleppo, Syria: ICARDA.
Bedigian, D. 2004. History and lore of sesame in Southwest
Asia. Economic Botany, 58, 329–53.
Bedigian, D. 2010. Cultivated sesame, and wild relatives
in the genus Sesamum L. In: Bedigian, D. (ed.) Sesame: the
genus Sesamum. Medicinal and Aromatic Plants - Industrial
Profiles. Boca Raton, FL: CRC Press.
Bedigian, D. and Harlan, J. R. 1986. Evidence for cultiva-
tion of sesame in the ancient world. Economic Botany, 40,
137–54.
Beech, M. 2003. Archaeobotanical evidence for early date
consumption in the Arabian Gulf. In: ECSSR (ed.) The
date palm – from traditional resource to green wealth. Pp.
11–31. Abu Dhabi, Emirates center for strategic studies
and research.
Beech, M. and Shepherd, E. (2001). Archaeobotanical evi-
dence for early date consumption on Dalma Island,
United Arab Emirates. Antiquity, 75, 83–9.
Behre, K.-E. 1992. The history of rye cultivation in Europe.
Vegetation history and Archaeobotany, 1, 141–56.
Bellinger, L. 1962. Textiles from Gordion. Bulletin of the
Needle and Bobbin Club, 46, 5–33.
Ben Ze’ev, N. and Zohary, D. 1973. Species relationships in
the genus Pisum L. Israel Journal of Botany, 22, 73–91.
Bennett, S. J. and Maxted, N. 1997. An ecogeographic anal-
ysis of the Vicia narbonensis complex. Genetic Resources
and Crop Evolution, 44, 411–28.

Bergfjord, C., Karg, S., Rast-Eicher, A., Nosch, M. L.,
Mannering, U., Allaby, R. G., Murphy, B. M. and Holst,
B. 2010. Comment on ‘30,000-Year-Old Wild Flax Fibers’.
Science, 328, 1634-b.
Beridze, R. K. and Kvatchadze, M. V. 1981. Origin and evo-
lution of cultivated plums of Georgia. Kulturpflanze, 29,
147–50.
Bertsch, F. 1943. Der dinkel. Landw Jahrbuch, 92, 241–52.
Bertsch, K. 1950. Vom ursprung der hexaploiden weizen.
Züchter, 20, 24–7.
Bertsch, K. and Bertsch, F. 1949. Geschichte unserer
Kulturpflanzen, Stuttgart, Wissenschaftliche Verlagsges.
Berzsényi, B. and Gyulai, F. 1998. The archaeobotanical anal-
ysis of the Middle Bronze Age settlement at Bölcske-Vörösgyír.
In: 31st International Symposium on Archaeometry, 27
April-1 May 1998 Budapest. Programme abstracts, Pp.
26–7.
Betts, A., van der Borg, K., de Jong, A., McClintock, C. and
van Strydonck, M. 1994. Early Cotton in North Arabia.
Journal of Archaeological Science, 21, 489–99.
Bieniek, A. 2007. Neolithic plant husbandry in the Kujawy
region of central Poland. In: Colledge, S. and Conolly, J.
(eds.) The origins and spread of domestic plants in southwest
Asia and Europe. Pp. 315–26. Walnut Creek, CA: Left
Coast Press.
Bieniek, A. and Lityńska-Zając, M. 2001. New finds of
Malus sylvestris Mill. (wild apple) from Neolithic sites in
Poland. Vegetation History and Archaeobotany, 10, 105–6.
Blankenhorn, B. and Hopf, M. 1982. Pflanzenreste aus spät-
neolithischen Moorsiedlungen des Federseerieds.
Jahrbuch des Römisch-Germanischen Zentralmuseums Mainz,
29, 74–99.
Blasco, A., Edo, M., Villalba, M. J., Buxó, R., Juan-Tresserras,
J. and Saña Seguí, M. 1999. Del cardial al postcardial en
la cueva de Can Sadurní (Begues, Barcelona). Primeros
datos sobre su secuencia estratigráfica, paleoeconómica
y ambiental. In: Bernabeu, J. and Orozco, T. (eds) Actes
del II Congrés del Neolític a la Península Ibèrica, Saguntum
Supplement 2. Pp. 59–68. València, Universitat de
València.
Blatter, R. H. E., Jacomet, S. and Schlumbaum, A. 2002.
Spelt-specific alleles in HMW glutenin genes from mod-
ern and historical European spelt (Triticum spelta L.).
Theoretical and Applied Genetics, 104(2–3), 329–37.
Blatter, R. H. E., Jacomet, S. Schlumbaum, A. 2004. About
the origin of European spelt (Triticum spelta L.): allelic
differentiation of the HMW Glutenin B1–1 and A1–2
subunit genes. Theoretical and Applied Genetics, 108(2),
360–7.
Blitzer, H. 1993. Olive cultivation and olive production in
Minoan Crete. In: Amouretti, M. C. and Brun, J. P. (eds.)
La production du vin et l’huile en Mediterranée de l’Age du
REFERENCES 203
Bronze à la fin du XVIeme siècle. Symposium International,
Aix-en-Provence 1991. Pp. 163–75. Paris, Bulletin de
Correspondence Hellénique Supplement, 26.
Boardman, J. 1976. The olive in the Mediterranean: its cul-
ture and use. Philosophical Transactions of the Royal Society
of London, Biological Sciences, 275, 187–96.
Boardman, S. and Jones, G. 1989. Experiments on the
effects of charring on cereal plant components. Journal of
Archaeological Science, 17, 1–11.
Boaretto, E. 2007. Determining the chronology of an
archaeological site using radiocarbon: minimizing
uncertainty. Israel Journal of Earth Science, 56, 207–16.
Boaretto, E. 2009. Dating materials in good archaeological
contexts: the next challenge for radiocarbon analysis.
Radiocarbon, 51, 275–81.
Boenke, N. 2007. Analyse botanischer Großreste aus dem
Salzbergbau am Dürrnberg bei Hallein. Rahden/Westfalen,
Universität Marburg/Lahn: Verlag Marie Leidorf
GmbH.
Bollinger, T. 1994. Samenanalytische untersuchung der Früh-
jungsteinzeit-lichen Seeufersiedlung Egolzwil 3 (Kanton
Luzern, Schweiz). Dissertationes Botanicae 221.
Bond, D. A., Lawes, D. A., Hawting, G. C., Saxana, M. C.
and Stephans, J. H. 1985. Faba bean. In: Summerfield,
R. J. and Roberts, E. H. (eds.) Grain legume crops.
Pp. 199–265. London, Collins.
Bor, N. L. 1968. Triticum. In: Townsend, C. C., Guest, E. and
Al-Rawi, A. (eds.) Flora of Iraq. Pp. 194–208. Baghdad,
Iraq: Ministry of Agriculture.
Bottema, S. 1980. On the history of the walnut in south
eastern Europe. Acta Botanica Neerlandica, 29, 343–9.
Bottema, S. and Woldring, H. 1984. Late Quaternary veg-
etation and climate of southwestern Turkey. Part II.
Palaeohistoria, 26, 123–49.
Bouby, L. and Billaud, Y. 2001. Économie agraire à la fin de
l’âge du Bronze sur les bords du lac du Bourget (Savoie,
France). Comptes Rendus de l’Académie des Sciences, Paris,
Sciences de la Terre et des planètes, 333, 749–56.
Braadbaart, F. and Van Bergen, P. F. 2005. Digital imag-
ing analysis of size and shape of wheat and pea upon
heating under anoxic conditions as a function of the
temperature. Vegetation History and Archaeobotany,
14, 67–75.
Braidwood, R. J. 1960. The agricultural revolution. Scientific
American, 203, 130–48.
Breton, C., Besnard, G. and Bervillé, A. A. 2003. Using
multiple types of molecular markers to understand
olive phylogeography. In: Zeder, M. A., Bradley, D. G.,
Emshwiller, E. and Smith, B. D. (eds.) Documenting
domestication: new genetic and archaeological paradigms.
Berkeley and Los Angeles, California: University of
California Press.
204 REFERENCES
Breton, C., Tersac, M. and Bervillé, A. A. 2006. Genetic
diversity and gene flow between the wild olive (oleaster,
Olea europaea L.) and the olive: several Plio-Pleistocene
refuge zones in the Mediterranean basin suggested by
simple sequence repeats analysis. Journal of Biogeography,
33, 1916–28.
Brewster, J. L. 1994. Onions and other vegetable alliums.
Wallingford, Oxon: CAB International.
Brombacher, C. 1995. Wirtschaftliche entwicklung aufgr-
und archäo-botanischer daten. In: Stöckli, W. E., Niffeler, U.
and Gross-Klee, E. (eds.) Die Schweiz vom Paläolithikum
bis zum frühen Mittelalter, Band II: Neolitikum. Pp. 86–96.
Basel: Verlag Schweizerische Gesellschaft für Ur- und
Frühgeschichte.
Brombacher, C. 1997. Archaeobotanical investigations of
Late Neolithic lakeshore settlements (Lake Biel,
Switzerland). Vegetation History and Archaeobotany, 6,
167–86.
Brombacher, C. 2000. Archäobotanische untersuchun-
gen. In: Hafner, A. and Suter, P. (eds.) 3400 v. Chr. - Die
Entwicklung der Bauerngesellschaften im 4. Jahrtausend v.
Chr. am Bielersee aufgrund der Rettungsgrabungen von
Nidau und Sutz-Lattrigen. Pp. 155–68. Bern: Paul Haupt
Verlag.
Brombacher, C. and Jacomet, S. 2003. Ackerbau, sammel-
wirtschaft und umwelt. In: Zwahlen, H. (ed.) Die jung-
neolithische Siedlung Port-Stüdeli. Pp. 66–86. Bern: Paul
Haupt Verlag.
Brombacher, C., Ghiggi, D. and Jacomet, S. 2005.
Untersuchungen der pflanzlichen makroreste. In:
Conscience, A.-C. (ed.) Wädenswil—Vorder Au. Eine
Seeufersiedlung am Übergang vom 17. zum 16. Jh. v.
Chr. im Rahmen der Frühbronzezeit am Zürichsee.
Pp. 25–32. Zürich und Egg: Baudirektion Kanton
Zürich, Hochbauamt, Kantonsarchäologie.
Bronk-Ramsey, C. 1995. Radiocarbon calibration and anal-
ysis of stratigraphy: the OxCal program. Radiocarbon,
37, 425–30.
Bronk-Ramsey, C. 2001. Development of the radiocarbon
program OxCal. Radiocarbon, 43, 355–63.
Browicz, K. 1972. Prunus, Amygdalus, Malus, Pyrus. In:
Davis, P. H. (ed.) Flora of Turkey. Pp. 8–12, 21–8, 157–9,
160–8. Edinburgh, Edinburgh University Press.
Browicz, K. 1982. Chorology of trees and shrubs in south-
west Asia and adjacent regions. Institute of Dendrology,
Polish Academy of Sciences, Kórnik, Poland, 1, 172.
Browicz, K. 1986. Chorology of trees and shrubs in south-
west Asia and adjacent regions. Institute of Dendrology,
Polish Academy of Sciences, Kórnik, Poland, 5, 88.
Browicz, K. 1988. Chorology of trees and shrubs in south-
west Asia and adjacent regions. Institute of Dendrology,
Polish Academy of Sciences, Kórnik, Poland, 6, 86.
Browicz, K. 1992. Chorology of trees and shrubs in south-
west Asia and adjacent regions. Institute of Dendrology,
Polish Academy of Sciences, Kórnik, Poland, 9, 83.
Browicz, K. 1993. Conspect and chorology of the genus
Pyrus L. Arboretum Kórnikie, 38, 17–33.
Browicz, K. 1996. Chorology of trees and shrubs in south-
west Asia and adjacent regions. Institute of Dendrology,
Polish Academy of Sciences, Kórnik, Poland, 11, 16–17.
Browicz, K. and Zohary, D. 1996. The genus Amygdalus L.
(Rosaceae): Species relationships, distribution and evo-
lution under domestication. Genetic Resources and Crop
Evolution, 43, 229–47.
Buschan, G. 1895. Vorgeschichtliche botanik der cultur- und
nutzpflanzen der alten Welt auf grund prähistorischer funde.
Breslau, J. U. Kern Verlag.
Butler, A. 1998. Grain legumes: evidence of these impor-
tant ancient food resources from early pre-agrarian and
agrarian sites in southwest Asia. In: Damania, A. B.,
Valkoum, J., Willcox, G. and Qualset, C. O. (eds.) The
origins of agriculture and crop domestiction. Pp. 102–17.
Aleppo, Syria: ICARDA.
Butler, A. 2009. Evidance of domestication in the Old
World grain legumes. In: Fairbairn, A. and Weiss, E.
(eds.) From foragers to farmers: papers in honour of Gordon
C. Hillman. Pp. 98–102. Oxford, Oxbow Books.
Butterworth, K. M., Adams, D. C., Horner, H. T. and
Wendel, J. F. 2009. Initiation and early development of
fibers in wild and cultivated cotton. International Journal
of Plant Sciences, 170, 561–74.
Buttner, R. 2001. Armeniaca. In: Hanelt, P. (ed.) Mansfeld’s
Encyclopedia of Agricultural and Horticultural Crops.
Pp. 523–7. Berlin, Institute of Plant Genetics and Crop
Plant Researches.
Buurman, J. 1987. A middle Bronze Age corn-stack at
Twisk, province of North Holland. Berichten van de
Rijksdienst voor het Oudheidkundig Bodemonderzoek, 37,
7–37.
Buurman, J., van Geel, B. and Reenen, G. B. A. 1995.
Palaeoecological investigations of a Late Bronze Age
watering place at Bovenkarspel, The Netherlands. In:
Herngreen, G. F. W. and van der Valk, L. (eds.) Neogene
and Quaternary Geology. Pp. 249–70. Mededelingen Rqks
Geologische Dienst.
Buxó, R. 1997. Presence of Olea europaea and Vitis vinifera in
archaeological sites from the Iberian Peninsula.
Lagascalia, 19, 271–82.
Buxó, R. 2004. La explotación de los recursos vegetales de
la Cueva del Toro: aproximación arqueobotánica a la
agricultura en época neolítica. In: Socas, M. D., Camálich,
M. D. and González, P. (eds.) La Cueva del Toro (Sierra de
El Torcal-Antequera-Málaga). Un modelo de Ocupación
Ganadera en el Territorio Andaluz, entre el VI y el II milenios

A.N.E. Pp. 267–84. Sevilla: Junta de Andalucía,
Consejería de Cultura.
Buxó, R. and Piqué, R. 2008. Arqueobotánica, los usos de las
plantas en la Península Ibérica. Barcelona, Ariel.
Cappers, R. T. J. 1996. Chapter 19: Archaeobotanical
remains. In: Sidebotham, S. E. and Wendrich, W. Z.
(eds.) Berenike 1995, preliminary report. Pp. 319–36.
Leiden: Centre for non-Western Studies.
Cappers, R. T. J. 2006. Roman Foodprints at Berenike:
Archaeobotanical Evidence of Subsistence and Trade in the
Eastern Desert of Egypt. Los Angeles, Cotsen Institute of
Archaeology, UCLA.
Cappers, R. T. J. and Raemaekers, D. C. M. 2008. Cereal
cultivation at Swifterbant? Neolithic wetland farming
on the North European Plain. Current Anthropology, 49,
385–402.
Cârciumaru, M. 1996. Paleoetnobotanica: studii în preisto-
ria şi protoistoria României. Iasi, Glasul Bucovinei
Helios.
Cârciumaru, M. and Monah, F. 1985. Raport preliminar
privind semintele carbonizate de la Poduri-Dealul
Ghindaru, judetul Bacau. Memoria Antiquitatis, 19–21,
699–708.
Cartwright, C. R. 2003. Grapes or raisins? An early Bronze
Age larder under the microscope. Antiquity, 77, 345.
Castelletti, L. 1976. Rapporto preliminare sui resti vegetali
macroscopici della serie neoliticobronzo di Pienza
(Siena). Rivista Archeologica dell’Antica Provincia e Diocesi
di Como, 156–157, 243–51.
Caton-Thompson, G. and Gardner, E. W. 1934. The desert
Fayum. London, Royal Anthropological Institute of
Great Britain and Ireland.
Cavalieri, D., McGovern, P. E., Hartl, D. L., Mortimer, R.
and Polsinelli, M. 2003. Evidence for S. cerevisiae fer-
mentation in ancient wine. Journal of Molecular Evolution,
57, S226–32.
Chang, T. T. 1995. Rice. In: Smartt, J. and Simmonds, N. W.
(eds.) Evolution of crop plants. 2 ed. Pp. 147–55. London,
Longman.
Chao, C. T. and Krueger, R. 2007. The date palm (Phoenix
dactylifera L.); overview of biology, uses, varieties, and
genetics. HortScience, 42, 1077–82.
Chapman, M. A., Hvala, J., Strever, J. and Burke, J. M.
2010. Population genetic analysis of safflower (Carthamus
tinctorius; Asteraceae) reveals a Near Eastern origin and
five centers of diversity. American Journal of Botany, 97,
831–40.
Chapman, M. and Bruke, J. 2007. DNA sequence diversity
and the origin of cultivated safflower (Carthamus tincto-
rius L.; Asteraceae). BMC Plant Biology, 7: 60.
Charles, M. 1993. Botanical remains. In: Green, A. (ed.)
Abu Salabikh excavations. Vol. 4. The 6G Ash-tip and its
REFERENCES 205
contents. Pp. 203–7. London, British School of
Archaeology in Iraq.
Charles, M. 2007. East of Eden? A consideration of
Neolithic crop spectra in the eastern Fertile Crescent
and beyond. In: Colledge, S. and Conolly, J. (eds.) The
origins and spread of domestic plants in Southwest Asia and
Europe. Pp. 37–52. Walnut Creek, CA: Left Coast Press.
Charles, M. and Hillman, G. C. 1992. Crop husbandry in
a desert environment: evidence from the charred plant
macroremains. In: Mason, V. M. (ed.) New research at the
Jeitun settlement: preliminary report on the work of the
Soviet-British expedition. Pp. 83–94. Ashgabad, Academy
of Sciences of Turkmenistan.
Cheben, I. and Hajnalová, E. 1997. Neolitische und
Äneolitische öfen in der Slowakei aus der sicht des
archäologen und archäobotanikers. Archaeologia Austriaca,
81, 41–52.
Chowdhury, K. A. and Buth, G. M. 1970. 4500-year-old
seed suggest that true cotton is indigenous to Nubia.
Nature, 227, 85–6.
Chowdhury, K. A. and Buth, G. M. 1971. Cotton seeds
from the Neolithic in Egyptian Nubia and the origin of
the Old World cotton. Biological Journal of the Linnean
Society, 3, 303–13.
Christensen, S., Von Bothmer, R., Poulsen, G., Maggioni, L.,
Phillip, M., Andersen, B. and Jørgensen, R. 2011. AFLP
analysis of genetic diversity in leafy kale (Brassica olera-
cea L. convar. acephala (DC.) Alef.) landraces, cultivars
and wild populations in Europe. Genetic Resources and
Crop Evolution, 58, 657–666.
Morphology, geography and infraspecific taxonomics of
TriticumClapham, A. J. and Rowley-Conwy, P. 2007. New
discoveries at Qasr Ibrim, Lower Nubia. In: Cappers, R.
(ed.) Fields of Change, Progress in African Archaeobotany.
Pp. 157–64. Groningen: Barkhuis Publishing.
Cleuziou, S. and Costantini, L. 1982. A l’origine des oasis.
Recherche, 13, 1180–2.
Coart, E., van Glabeke, S., de Loose, M., Larsen, A. S. and
Roldán-Ruiz, I. 2006. Chloroplast diversity in the genus
Malus: new insights into the relationship between the
European wild apple (Malus sylvestris (L.) Mill.) and the
domesticated apple (Malus domestica Borkh.). Molecular
Ecology, 15, 2171–82.
Cole, S., Maxted, N. and van der Maesen, L. J. G. 1998.
Identification aids for Cicer (Leguminosae, Cicereae)
taxa. Edinburgh Journal of Botany, 55, 243–65.
Colledge, S. 2001 Plant Exploitation on Epipalaeolithic and
Early Neolithic Sites in the Levant. Oxford: BAR
International Series 986.
Colledge, S. and Conolly, J. (eds.) 2007. The origins and
spread of domestic plants in Southwest Asia and Europe,
Walnut Creek, CA: Left Coast Press.
206 REFERENCES
Condit, I. J. 1947. The fig. Waltham, MA, Chronica
Botanica.
Cooper, W. C. and Chapot, H. 1977. Fruit production with
special emphasis on fruit for processing. In: Nagi, S.,
Shur, P. E. and Valdhuis, M. K. (eds.) Citrus science and
technology. Pp. 1–127. Westport, Conn: Avi Publications.
Costantini, L. 1977. Le piante. In: Tucci, G. (ed.) La citta
bruciata del deserto salato. The burnt city in the salt desert.
Pp. 159–71. Venice-Mestre, Erizzo.
Costantini, L. 1989. Plant exploitation at Grotta dell’ Uzzo,
Sicily: new evidence for the transition from Mesolithic
to Neolithic subsistance in southern Europe. In: Harris,
D. R. and Hillman, G. C. (eds.) Foraging and farming: the
evolution of plant exploitation. Pp. 197–206. London,
Unwin and Hyman.
Costantini, L. and Costantini-Biasini, L. 1985. Agriculture
in Baluchistan between the 7th and 3rd millennium bc.
Newsletter of Baluchistan Studies (Istituto Universitario
Orientale, Naples), 2, 16–30.
Costantini, L. and Costantini-Biasini, L. 1986. Palaoethnobo-
tanical investigations in the Middle East and Arabian
Peninsula, 1986. East and West, 36(4), 354–65.
Costantini, L. and Stancanelli, M. 1994. La Prehistoria agri-
cola dell’ Italia centro-meridionale: Il contributo delle
Indogeni Archeobotaniche. Origini, Prehistoria e
Protohistoria delle Civilta Antiche, 17, 199–244.
Costantini, L., Costantini-Biasini, L. and Lentini, A. 1997.
Agricoltura e note sull’ ambiente del’ abitato Neolitico
di Scamuso. In: Biancofiore, F. and Coppola, D. (eds.)
Scamuso. Pp. 199–209. Rome, Dipartimento di Storia
dell’Universita di Roma.
Courtin, J. and Erroux, J. 1974. Aperçu sur l’agriculture
préhistorique dans le Sud-Est de la France. Bulletin de la
Société Préhistorique Française, 71, 321–34.
Courtin, J., Erroux, J. and Thommert, J. H. 1976. Les
céréales du Néolithqiue ancien de Chateauneuf-Les-
Martigues (Bouches-du-Rhône). Bulletin du Muséum
d’Histoire Naturelle de Marseille, 36, 11–15.
Cowling, W. A., Buirchell, B. J. and Tapia, M. E. 1998.
Lupin, Lupinus spp. Promoting the conservation and use of
underutilized and neglected crops, 23, Gatersleben. Rome,
Institute of Plant Genetics and Crop Plant Research,
International Plant Genetic Resources Institute.
Crawford, G. W. 1992. Prehistoric plant domestication in
East Asia. In: Cowen, C. W. and Watson, P. J. (eds.) The
origins of agriculture: an international perspective. Pp. 7–38.
Washington DC, Smithsonian Institution Press.
Crawford, G. W. and Shen, C. 1998. The origins of rice agri-
culture: recent progress in East Asia. Antiquity, 72, 858–66.
Crawford, G. W., Chen, X. and Wang, J. 2006. Houli Culture
rice from the Yuezhuang site, Jinan. Orient Archaeology,
3, 247–51.
Curdy, P., Mottet, M., Nicoud, C., Baudais, D., Lundström-
Baudais, K. and Moulin, B. 1993. Brig-Glis/Waldmatte,
un habitat alpin de l’âge du Fer. Fouilles archéologiques
N9 en Valais. Archäologie der Schweiz, 16, 138–51.
Dagan, J. and Zohary, D. 1970. Wild tetraploid wheat from
West Iran cytogenetically identical with Israeli T. dicoc-
coides. Wheat Information Service, 31, 15–17.
Darby, W. J., Ghalioungui, P. and Grivetti, L. 1977. Food: the
gift of Osiris. London, New York: Academic Press.
Darlington, C. D. and Wyle, A. P. 1955. Chromosome
atlas of flowering plants. London, George Allen and
Unwin Ltd.
Davies, D. R. 1995. Peas. In: Smartt, J. and Simmonds, N. W.
(eds.) Evolution of crop plants. 2 ed, Pp. 294–6. London:
Longman, U.K.
Davies, F. S. and Albrigo, L. G. 1994. Citrus; crop production
science in horticulture. Wallingford, U.K.: CAB
International.
Davis, P. H. 1970. Pisum L. In: Davis, P. H. (ed.) Flora of
Turkey. Vol. 3. Pp. 370–3. Edinburgh, Edinburgh
University Press.
de Bustos, A. and Jouve, N. 2002. Phylogenetic relation-
ships of the genus Secale based on the characterisation of
rDNA ITS sequences. Plant Systematics and Evolution,
235, 147–54.
de Candolle, A. 1886. Origin of cultivated plants. 2 ed.
London, Paul Trench. Reprinted 1964 by Hafner Publ.
Company, New York.
de Lattin, B. 1939. Über den ursprung und die verbreitung
der reben. Züchter, 11, 217–25.
de Moulins, D. 1997. Agriculture changes at Euphrates
and steppe sites in the mid-8th to 6th millennium b.c.
Oxford, BAR International Series 683.
de Moulins, D. 2000. Abu Hureyra 2: plant remains from
the Neolithic. In: Moore, A. M. T., Hillman, G. C. and
Legge, A. J. (eds.) Village on the Euphrates: from foraging to
farming at Abu Hureyra. Pp. 399–416. New York, Oxford
University Press.
de Roller, G.-J. 1992. Archaeobotanical remains from Tell
Ibrahim Awad, seasons 1988 and 1989. In: van den
Brink, E. C. M. (ed.) The Nile delta in transition; 4th - 3rd
millennium B.C. Pp. 111–15. Jerusalem, Israel Exploration
Society.
de Vartavan, C., Arakelyan, A. and Amorós, V. A. 2010.
Codex of ancient Egyptian plant remains. London, SAIS
Academic Books.
de Vries, I. M. 1997. Origin and domestication of Lactuca
sativa L. Genetic Resources and Crop Evolution, 44,
165–74.
de Wet, J. M. J. 1975. Evolutionary dynamics of cereal
domestication. Bulletin of the Torrey Botanical Club, 102,
307–12.

de Wet, J. M. J. 1981. Species concepts and systematics of
domesticated cereals. Kulturpflanze, 29, 177–98.
de Wet, J. M. J. 1995. Foxtail millet. In: Smartt, J. and
Simmonds, N. W. (eds.) Evolution of crop plants. 2 ed,
Pp. 170–2. London, Longman.
de Wet, J. M. J., Harlan, J. R. and Price, E. G. 1976. Variability
in Sorghum bicolor. In: Harlan, J. R., de Wet, J. M. J. and
Stemler, A. B. L. (eds.) The origins of African plant domes-
tication. Pp. 453–63. The Hague, Mouton.
de Wet, J. M. J., Oestry-Stidd, L. L. and Cubero, J. I. 1979.
Origins and evolution of foxtail millets Setaria italica.
Journal d’Agriculture Traditionnelle et de Botanique Appliquée,
26, 53–64.
de Winter, B. 1990. A new species of Citrullus (Benincaseae)
from the Namib Desert, Namibia. Bothalia, 20, 209–11.
Decker, Michael. 2009. Plants and progress: rethinking the
Islamic agricultural revolution. Journal of World History,
20(2), 187–206.
Dickson, J. H., Oeggl, K., Holden, T. G., Handley, L. L.,
O’Connell, T. C. and Preston, T. 2000. The omnivorous
Tyrolean Iceman: colon contents (meat, cereals, pollen,
moss and whipworm) and stable isotope analyses.
Philosophical Transactions of the Royal Society of London,
Biological Science, 355, 1843–9.
Diederichsen, A. 1996. Coriander: Coriandrum sativum L.
Promoting the conservation and use of underutilized
and neglected crops 3. Rome, International Plant Genetic
Resources Institute.
Diederichsen, A. and Hammer, K. 1995. Variation of culti-
vated flax (Linum usitatissimum L. subsp. usitatissimum)
and its wild progenitor pale flax (subsp. angustifolium
(Huds.) Thell.). Genetic Resources and Crop Evolution, 42,
263–72.
Dietsch, M. F. 1996. Gathered fruits and cultivated plants
at Bercy (Paris), a Neolithic village in a fluvial context.
Vegetation History and Archaeobotany, 5, 89–97.
Dietsch, M. F. 2001. Milieux humides pré- et protohisto-riques
dans le Bassin Parisien: L’étude des diaspores, Villeneuve
d’Ascq, Presses Universitaires du Septentrion.
Dillon, S. L., Lawrence, P. K. and Henry, R. 2001. The use
of ribosomal ITS to determine phylogenetic relation-
ships within Sorghum. Plant Systematics and Evolution,
230, 97–110.
Dönmez, E. O. and Belli, O. 2007. Urartian plant cultiva-
tion at Yoncatepe (Van), eastern Turkey. Economic Botany,
61, 290–8.
Dörfler, W. 1990. Die geschichte des hanfanbaus in
Mitteleuropa auf grund palynologischer untersuchungen
und von grossrestnachweisen. Praehistorische Zeitschrift,
65, 218–44.
Dorofeev, V. F., Filatenko, A. A., Migushova, E. F., Udashin,
R. A. and Iakubziner, R. R. 1979. Wheat. In: Dorofeev, V.
REFERENCES 207
F. and Korovina, O. N. (eds.) Flora of Cultivated Plants I.
Leningrad, Kolos. [in Russian].
Dowson, V. H. W. 1982. Date production and protection,
Technical Bulletin No. 35. Rome, FAO.
Duc, G., Bao, S., Baum, M., Redden, B., Sadiki, M., Suso,
M. J., Vishniakova, M. and Zong, X. 2010. Diversity
maintenance and use of Vicia faba L. genetic resources.
Field Crops Research, 115, 270–8.
Duke, J. A. 1973. Utilization of Papaver. Economic Botany,
27, 390–400.
Durrant, A. 1976. Flax and linseed. In: Simmonds, N. W.
(ed.) Evolution of crop plants. Pp. 190–3. London,
Longman.
Dvořák, J., di Terlizzi, P., Zhang, H. B. and Resta, P. 1993.
The evolution of polyploid wheats: identification of the
A genome donor species. Genome, 36, 21–31.
Dvořák, J., Luo, M. C., Yang, Z. L. and Zhang, H. B. 1998.
The structure of the Aegilops tauschii genepool and the
evolution of hexaploid wheat. Theoretical and Applied
Genetics, 67, 657–70.
Dvořák, J., McGuire, P. E. and Cassidy, B. 1988. Apparent
sources of the A genomes of wheats inferred from pol-
ymorphism in abundance and restriction fragment
length of repeated nucleotid sequences. Genome, 30,
680–9.
Dzhangaliev, A. D., Salova, T. N. and Turekhanova, P. M.
2003. The wild fruit and nut plants of Kazakhstan.
Horticultural Reviews, 29, 305–72.
Eastwood, G. M. 1984. Egyptian dyes and colours. In:
Walton, P. (ed.) Dyes on historical and archaeological tex-
tiles. Pp. 9–19. York, Textile Research Associates.
El Hadidi, M. N., Fahmy, A. G. and Willerding, U. 1996.
The paleoethnobotany of locality 11C, Hierakonpolis
(3800–3500 ca. BC); Egypt 1, cultivated crops and wild
plants of potential value. Taekholmia, 16, 31–44.
Elbaum, R., Zaltzman, L., Burgert, I. and Fratzl, P. 2007.
The role of wheat awns in the seed dispersal unit.
Science, 316, 884–6.
Ellison, R., Renfrew, J., Brothwell, D. and Seeley, N. 1978.
Some food offerings from Ur, excavated by Sir Leonard
Woolley, and previously unpublished. Journal of
Archaeological Science, 5, 167–77.
Endrődi, A. and Gyulai, F. 2000. Hearths and other finds of
the Late Copper Age Baden culture at Budapest-Csepel
Island (gynaecomorphic vessels, archaeobotanical r
emains). Archaeologiai Értesítő, 125, 9–44.
Engelmark, R. 1992. A review of the farming economy in
South Scania based on botanical evidence. In: Larsson,
L., Callmer, J. and Stjernquist, B. (eds.) The archaeology of
the cultural landscape. field work and research in a South
Swedish rural region. Pp. 369–75. Stockholm, Acta
Archaeologica Lundensia 4 (19).
208 REFERENCES
Enneking, D. 1995. The toxicity of Vicia species and their uti-
lisation as grain legumes. Nedlands W.A.: University of
Western Australia.
Enneking, D. and Maxted, N. 1995. Narbon bean. In:
Smartt, J. and Simmonds, N. W. (eds.) Evolution of crop
plants. 2 ed, Pp. 316–21. London, Longman.
Epstein, C. 1978. A new aspect of Chalcolithic culture. Bulletin
of the American Schools of Oriental Research, 229, 27–45.
Epstein, C. 1993. Oil production in the Golan Heights dur-
ing the Chalcolithic period. Journal of the Institute of
Archaeology, Tel Aviv University, 20, 133–46.
Ertuğ, F. 2004. Wild edible plants of the Bodrum area
(Muğla, Turkey). Turkish Journal of Botany, 28,
161–74.
Ertuğ, F. 2009. Wild plant foods: routine dietary supple-
ments of famine food? In: Fairbain, A. S. and Weiss, E.
(eds.) From foragers to farmers: papers in honour of Gordon
C. Hillman. Pp. 64–70. Oxford, Oxbow Books.
Evans, A. J. 1928. The Palace of Minos at Knossos. London,
Macmillan.
Evans, G. M. 1995. Rye. In: Smartt, J. and Simmonds, N. W.
(eds.) Evolution of crop plants. 2 ed, Pp. 108–11. London,
Longman.
Evershed, R. P. 2008. Organic residue analysis in archaeol-
ogy: the archaeological biomarker revolution. Archaeo-
metry, 50, 895–924.
Fahmy, A. G. 1995. A historical flora of Egypt. Ph.D.,
University of Cairo.
Fahmy, A. G. 2001. Palaeoethnobotanical studies of the
Neolithic settlement in Hidden Valley, Farafra Oasis,
Egypt. Vegetation History and Archaeobotany, 10, 235–46.
Fahmy, A. G. 2003. Palaeoethnobotanical studies of Egyptian
Predynastic cemeteries: new dimensions and contribu-
tions. In: Neumann, K., Butler, A. and Kahlheber, S. (eds.)
Food, fuel and fields: progress in African archaeobotany.
Pp. 95–106. Köln, Germany: Africa Praehistorica.
Fahmy, A. G. 2005. Missing plant macro remains as indica-
tors of plant exploitation in Predynastic Egypt. Vegetation
History and Archaeobotany, 14, 287–94.
Fahmy, A. G., Friedman, R. and Fadl, M. 2008.
Archaeobotanical studies at Hierakonpolis locality HK6:
The Pre and Early Dynastic elite cemetery. Archeo-Nil,
18, 169–83.
Fairbairn, A. 2005. A history of agricultural production at
Neolithic Çatalhöyük East, Turkey. World Archaeology,
37, 197–210.
Fairbairn, A., Asouti, E., Near, J. and Martinoli, D. 2002.
Macro-botanical evidence for plant use at Neolithic
Çatalhöyük south-central Anatolia, Turkey. Vegetation
History and Archaeobotany, 11, 41–54.
Fairbairn, A., Martinoli, D., Butler, A. and Hillman, G. C.
2007. Wild plant seed storage at Neolithic Çatalhöyük
East, Turkey. Vegetation History and Archaeobotany, 16,
467–79.
Fairbairn, A., Near, J. and Martinoli, D. 2005. Macrobotanical
investigations of the North, South and KOPAL areas at
Çatalhöyük. In: Hodder, I. (ed.) Inhabiting Çatalhöyük: reports
from the 1995–1999 seasons. Pp. 137–201. Cambridge/
Ankara, McDonald Institute for Archaeological Research/
British Institute of Archaeology at Ankara.
Fairweather, A. D. and Ralston, I. B. M. 1993. The Neolithic tim-
ber hall at Balbridie, Grampian Region, Scotland: the build-
ing, the date, the plant macrofossils. Antiquity, 67, 313–23.
Favre, P. 2002. Archäobotanik. In: Achour-Uster, C., Eberli, U.,
Ebersbach, R. and Favre, P. (eds.) Die seeufersiedlungen in
Horgen, die Neolithischen und Bronzezeitlichen fundstellen
dampfschiffsteg und scheller. Pp. 150–80. Zürich, Fotorotar.
Feemster, J. W. M. and Meyer, F. G. (eds.) 2002. The Natural
History of Pompeii, Cambridge, Cambridge University
Press.
Feldman, M. and Kislev, M. E. 2007. Domestication of
emmer wheat and evolution of free-threshing tetraploid
wheat. Israel Journal of Plant Sciences, 55, 207–21.
Feldman, M., Lupton, F. and Miller, T. E. 1995. Wheats. In:
Smartt, J. and Simmonds, N. W. (eds.) Evolution of crop
plants, 2nd edn. Pp. 184–92. London, Longman.
Feliks, Y. 1983. Plants and animals of the Mishna. Jerusalem,
Institute of Mishna Research.
Feliks, Y. 1994. Fruit trees in the Bible and Talmudic literature.
Jerusalem, Rubin Mass Press.
Ferguson, M. E., For-Lloyd, B. V., Robertson, L. D., Maxted, N.
and Nenbury, H. J. 1998. Mapping the geographical distri-
bution of genetic variation in the genus Lens for the
enhanced conservation of plant genetic diversity. Molecular
Ecology, 7, 1743–55.
Ferguson, M. E., Maxted, N., van Slageren, M. and
Robertson, L. D. 2000. A re-assessment of the taxonomy
of Lens Mill. (Leguminosae, Papilionoideae, Vicieae).
Botanical Journal of the Linnean Society, 133, 41–59.
Fietz, A. 1936. Prähistorische pflanzenreste aus der
Slowakei. Verhandlungen des Naturforschenden Vereins in
Brünn, 67, 149–51.
Figueiral, I. and Sanches, M. J. 2003. Eastern Trás-os-
Montes (NE Portugal) from the late prehistory to the
Iron Age: the land and the people. In: Fouache, E. (ed.)
The Mediterranean World, Environment and History.
Pp. 315–32. Paris, Elsevier.
Fischer, T. 1881. Die Dattelpalme, ihre geographische Verbreitung
und culturhistorische Bedeutung, Gotha, J. Perthes.
Flaishman, M. A., Rodov, V. and Stover, E. 2008. The fig: botany,
horticulture and breeding. Horticultural Reviews, 34, 132–96.
Flann, C. (ed.) 2009 + Global Compositae Checklist
(http://compositae.landcareresearch.co.nz), Accessed
28/3/2011.

Ford-Lloyd, B. V. 1995. Sugarbeet and other cultivated
beets. In: Smartt, J. and Simmonds, N. W. (eds.) Evolution
of crop plants. 2 ed, Pp. 35–40. London, Longman.
Ford-Lloyd, B. V. and Williams, J. J. 1975. A revision of Beta
section vulgares (Chenopodiaceae), with new light on
the origin of cultivated beets. Botanical Journal of the
Linnean Society, 71, 89–102.
Forsline, P. L., Aldwinckle, H. S., Dickson, E. E., Luby, J. J.
and Hokanson, S. 2003. Collection, maintenance, char-
acterization and utilization of wild apples of Central
Asia. In: Janick, J., Forsline, P. L., Dickscon, E. E., Way, R.
and Thompson, M. (eds.) Wild apple and fruit trees of
Central Asia. Pp. 1–61. New Jersey, John Wiley and
Sons.
Forte, A. V., Ignatov, A. N., Ponomarenko, V. V. Dorokhov,
D. B., and Savelyev, N. I. 2002. Phylogeny of the Malus
(apple tree) species, inferred from the morphological
traits and molecular DNA analysis. Russian Journal of
Genetics, 38, 1150–60.
Frese, L. 2003. Sugar beets and related wild species - from
collecting to utilization. In: Knüpffer, H. and Ochsmann, J.
(Eds.), Rudolf Mansfeld and Plant Genetic Resources.
Schriften zu Genetischen Ressourcen, 22, 170–81.
Fritsch, R. 1979. Zur Samenmorphologie des Kulturmohns
(Papaver somniferum L.). Kulturpflanze, 27, 217–27.
Fryxell, P. A. 1979. The natural history of the cotton tribe
(Malvaceae, tribe Gossypieae). College Station, Texas:
Texas A and M University Press,.
Fryxell, P. A. 1984. Taxonomy and germplasm resources.
In: Kohel, R. J. and Lewis, C. F. (eds.) Cotton. Pp.
27–57. Madison, Wisconsin: American Society of
Agronomy.
Fu, Y.-B. 2005. Geographic Patterns of RAPD Variation in
Cultivated Flax. Crop Science, 45, 1084–91.
Fu, Y.-B. and Allaby, R. 2010. Phylogenetic network of
Linum species as revealed by non-coding chloroplast
DNA sequences. Genetic Resources and Crop Evolution,
57, 667–77.
Fu, Y.-B. and Williams, D. 2008. AFLP variation in 25 Avena
species. Theoretical and Applied Genetics, 117, 333–42.
Fu, Y.-B., Peterson, G., Diederichsen, A. and Richards, K. W.
2002. RAPD analysis of genetic relationships of seven flax
species in the genus Linum L. Genetic Resources and Crop
Evolution, 49, 253–9.
Fukunaga, K., Ichitani, K., Taura, S., Sato, M. and Kawase,
M. 2005. Ribosomal DNA intergenic spacer sequence in
foxtail millet, Setaria italica (L.) P. Beauv. and its charac-
terization and application to typing of foxtail millet lan-
draces. Hereditas, 142, 38–44.
Fuller, D. Q. 2002. Fifty years of archaeobotanical studies
in India: laying a solid foundation. In: Settar, S. and
Korisettar, R. (eds.) Indian archaeology in retrospect, vol.
REFERENCES 209
III. Archaeology and interactive disciplines. Pp. 247–363.
New Delhi, Manohar.
Fuller, D. Q. 2003. Further evidence on the prehistory of
sesame. Asian Agri-History, 7, 127–37.
Fuller, D. Q. 2007. Contrasting patterns in crop domestica-
tion and domestication rates: recent archaeobotanical
insights from the Old World. Annals of Botany, 100,
903–24.
Fuller, D. Q. 2008. The spread of textile production and
textile crops in India beyond the Harappan Zone: an
aspect of the emergence of craft specialization and sys-
tematic trade. In: Osada, T. and Uessugi, A. (eds.)
Linguistics, Archaeology and the Human Past. Pp. 1–26.
Kyoto, Japan: Research Institute for Humanity and
Nature.
Fuller, D. Q. and Sato, Y.-I. 2008. Japonica rice carried to,
not from, Southeast Asia. Nature Genetics, 40, 1264–5.
Fuller, D. Q., Harvey, E. and Qin, L. 2007. Presumed
domestication? Evidence for wild rice cultivation and
domestication in the fifth millennium BC of the Lower
Yangtze region. Antiquity 81, 316–31.
Fuller, D. Q., Qin, L., Zheng, Y., Zhao, Z., Chen, X., Hosoya,
L. A. and Sun, G.-P. 2009. The domestication process
and domestication rate in rice: spikelet bases from the
Lower Yangtze. Science, 323, 1607–10.
Fuller, D. Q., Sato, Y.-I., Castillo, C., Qin, L., Weisskopf, A.,
Kingwell-Banham, E., Song, J., Ahn, S.-M. and van
Etten, J. 2010. Consilience of genetics and archaeobot-
any in the entangled history of rice. Archaeological and
Anthropological Sciences, 2, 115–31.
Füzes, M. 1990. A földmívelés kezdeti szakaszának (neoli-
tikum és rézkor) növényleletei Magyarországon
(Archaeobotanikai vázlat) (Plant remains from the early
phase of plant cultivation—Neolithic and Copper Age
—in Hungary. An archaeobotanical outline). Tapolcai
Városi Múzeum közleményei, 1, 139–238.
Füzes, M. 1991. Dunántúl korai növénytermesztése és
növényleletei. A Starčevo kultúra és a ‘Tapolcai csoport’
(Die frühe pflanzenzucht und die pflanzenfunde in
Transdanubien—die Starčevo-Kultur und die ‘Tapolca-
Gruppe’). Bibliotheca Musei Tapolcensis 1, 267–362.
Gabrielian, E. and Zohary, D. 2004. Wild relatives of food
crops native to Armenia and Nakhichivan. Flora
Mediterranea, 14, 5–80.
Galil, J. and Neeman, G. 1977. Pollen transfer and pollination in
the common fig (Ficus carica L.). New Phytologist, 79, 163–71.
Galil, J., Stein, M. and Horovitz, A. 1976. On the origin of
the sycamore fig (Ficus sycomorus L.) in the Middle East.
Gardens’ Bulletin, 29, 191–205.
Galili, E. and Shavit, J. 1994–5. Evidence of olive oil pro-
duction from the submerged site at Kfar Samir, Israel.
Journal of the Israel Prehistoric Society, 26, 122–33.
210 REFERENCES
Galili, E., Weinstein-Evron, M. and Zohary, D. 1989.
Appearance of olives in submerged Neolithic sites along
the Carmel coast. Journal of the Israel Prehistoric Society,
22, 95–7.
Galili, E., Weinstein-Evron, M., Hershkovitz, I., Gopher, A.,
Kislev, M., Lernau, O., Kolska-Horwitz, L. and Lernau,
H. 1993. Atlit-Yam—a prehistoric site on the sea-floor off
the Israeli coast. Journal of Field Archaeology, 20, 133–57.
Garfinkel, Y., Kislev, M. E. and Zohary, D. 1988. Lentil in
the Pre-Pottery Neolithic B Yiftah’el: additional evi-
dence of its early domestication. Israel Journal of Botany,
37, 49–51.
Gegas, V. C., Nazari, A., Griffiths, S., Simmonds, J., Fish,
L., Orford, S., Sayers, L., Doonan, J. H. and Snape, J. W.
2010. A genetic framework for grain size and shape vari-
ation in wheat. Plant Cell, 22, 1046–56.
Georges Métailié 2007. Grafting as an agricultural and cul-
tural practice in ancient China. In: Conan, M. and Kress,
W. J. (eds.) Botanical progress, horticultural innovation and
cultural change. Pp. 147–58. Washington DC, Dumbarton
Oaks.
Germer, R. 1985. Flora des Pharaonischen Ägypten, Mainz,
Verlag Philip von Zabern.
Germer, R. 1989a. Ägyptens auswärtige Beziehungen hin-
sichtlich der Kulturgewächse. Dissertationes Botanicae,
133, 57–66.
Germer, R. 1989b. Die Pflanzenmaterialien aus dem Grab des
Tutankhamun, Hildesheim, Gerstenberg.
Germer, R. 1992. Die Textilfärberei und die Verwendung
gefärbter Textilien im alten Ägypten, Wiesbaden,
Harrassowitz.
Gill, K. S. and Yermanos, D. M. 1967. Cytogenetic studies on
the genus Linum. I. Hybrids among the taxa with 15 as
the haploid chromosome number. Crop Science, 7, 27–31.
Gillett, J. B. 1981. Botanical samples. In: Safar, F., Mustafa,
M. A. and Lloyd, S. (eds.) Eridu. Pp. 317–18. Baghdad,
State organization of antiquities and heritage.
Giżbert, W. 1960. Studium porównawcze nad ziarnami
żyta kopalnego (A comparative study on excavated
grains of rye). Archaeologia Polski, 5, 81–90.
Gladstones, J. S. 1974. Lupins of the Mediterranean region
and Africa. Technical Bulletin, 26. Perth, Department of
Agriculture of Western Australia. This is a booklet from a
series.
Glover, I. C. and Higham, F. W. 1996. New evidence for
early rice cultivation in South, Southeast and East Asia.
In: Harris, D. R. (ed.) The origins and spread of agriculture
and pastoralism in Eurasia. Pp. 413–41. London, University
College London Press.
Gluza, I. 1984. Neolithic cereals and weeds from the local-
ity of the Lengyel culture at Nowa Huta-Mogila near
Cracow. Acta Palaeobotanica Polonica, 23, 123–84.
Gluza, I. 1994. Wstepne wyniki badań odcisków roslin-
nych ze stanowisk kultury ceramiki wstegowej rytej w
Gniechowicach i Starym Zamku, woj. wrocławski
(Preliminary results of the investigation of plant
imprints from the Linear Pottery Culture sites at
Gniechowice and Stary Zamek, Wrocław province).
Polish Botanical Studies, Guidebook Series, 11, 55–69.
Godwin, H. 1967. The ancient cultivation of hemp.
Antiquity, 41, 42–8.
Gophna, R. and Kislev, M. E. 1979. Tell Saf (1977–1978).
Revue Biblique, 86, 112–14.
Göransson, H., Lemdahl, G. and Johansson, B. M. 1995.
Alvastra pile dwelling: Palaeoethnobotanical studies.Theses
and Papers in Archaeology, 6. Lund, Lund University
Press.
Goring-Morris, A. N. and Belfer-Cohen, A. 2011 Neolithization
processes in the Levant, the outer envelope. Current
Anthropology, 52(S4), pp. S195–S208.
Green, P. S. and Wickens, G. E. 1989. The Olea europaea
complex. In: Tan, K. (ed.) The P. H. Davis and I. C. Hedge
Festschrift. Pp. 287–99. Edinburgh, Edinburgh University
Press.
Greuter, W. 2006–2009. Compositae (pro parte majore).
In: Greuter, W. and von Raab-Straube, E. (eds.) Euro +
Med Plantbase—the information resource for Euro-
Mediterranean plant diversity, http://www.emplantbase.
org/home.html.
Guarino, C., Casoria, P. and Menale, B. 2000. Cultivation
and use of Isatis tinctoria L. (Brassicaceae) in Southern
Italy. Economic Botany, 54, 395–400.
Gullati, A. N. and Turner, A. J. 1929. A note on early his-
tory of cotton. Bulletin No. 17; Technological series 12.
Bombay, Indian central cotton committee.
Gustafsson, M. and Lannér-Herrera, C. 1997. Overview of
the Brassica oleracea complex: their distribution and eco-
logical specifications. In: Valdés, B., Heywood, V. H.,
Ramondo, F. M. and Zohary, D. (eds.) Proceedings of the
workshops on conservation of the wild relatives of European
cultivated plants. Pp. 27–37. Palermo, Bocconea,
Herbarium Mediterraneum Panoramitanum.
Gutiérrez-Marcos, J. F., Vaquero, F., de Miera, L. E. S. and
Vences, F. J. 2006. High genetic diversity in a world-
wide collection of Lathyrus sativus L. revealed by isozy-
matic analysis. Plant Genetic Resources, 4, 159–71.
Gymer, P. T. 1978. The genetics of the six-row/two-row
character. Barley Genetics Newsletter, 8, 44–6.
Gyulai, F. 2003. Archaeobotanical remains and environ-
ment of Bell Beaker Csepel-Group. In: Czebreszuk, J.
and Szmyt, M. (eds.) The Northeast Frontier of Bell Beakers.
Pp. 277–82. Oxford, Archaeopress.
Gyulai, F. 2007. Seed and fruit remains associated with
Neolithic origins in the Carpathian Basin. In: Colledge,

S. and Conolly, J. (eds.) The origins and spread of domestic
plants in Southwest Asia and Europe. Pp. 125–40. Walnut
Creek, CA: Left Coast Press.
Hajnalová, E. 1978. Funde von Triticum-resten aus einer
Hallstattzeitlichen getreide-speichergrube in Bratislava-
Devin/CSSR. Berichte der Deutschen Botanischen Gesellschaft,
91, 85–96.
Hajnalová, E. 1989. Katalóg zyvskov semien a plodov v arche-
ologických nálezoch na Slovensku. Acta Interdisciplinaria
Archaeologica (Nitra, Czechoslovakia), 6, 3–192.
Hajnalová, E. 1993. Praveké osídenie lokality Sarišské
Michalany dokumentované rastlinnými zvyškami.
Východoslovenský pravek, 4, 49–65.
Hajnalová, E. and Hajnalová, M. 2004. Zbierané rastliny
ako zdroj potravy v praveku strednej Európy a ich
archeobotanické nálezy na Slovensku. In: Janák, V. and
Stuchlik, S. (eds.) Otázky Neolitu a Eneolitu našich zemí,
Acta Archaeologica Opaviensia. Pp. 33–47. Opava, University
of Opava.
Häkkinen, K. and Lempiäinen, T. 1996. Die ältesten getrei-
depflanzen der Finnen und ihre namen. Finnisch-
Ugrische Forschungen, 53, 115–82.
Haldane, C. 1993. Direct evidence for organic cargoes in
the Late Bronze Age. World Archaeology, 24, 348–60.
Halstead, P. and Jones, G. 1980. Early economy in Thessaly.
Anthropologika, 1, 93–117.
Hammer, K. 1984. The domestication syndrome.
Kulturpflanze 32, 11–34.
Hammer, K. and Fritsch, R. 1977. Zur frage nach der
Ursprungsart des kulturmohns (Papaver somniferum L.).
Kulturpflanze, 25, 113–24.
Hanelt, P. (ed.) 2001. Mansfeld’s rncyclopedia of agricultural
and horticultural crops. Berlin, Institute of plant genetics
and crop plant researches.
Hanelt, P. 1963. Monographische Uebersicht der Gattung
Carthamus (Compositae). Feddes Repertorium Specierum
Novarum Regni Vegetabilis, 67, 41–180.
Hanelt, P. 1972. Die infraspezifische variabilität von Vicia
faba und ihre gliederung. Kulturpflanze, 20, 75–128.
Hanelt, P. 1985. Zur taxonomie, chorologie und ökologie
der wildarten von Allium L. sect. Cepa (Mill.) Prokh.
Flora, 176, 99–116.
Hanelt, P. 1986. Pathways of domestication with regard to
crop types (grains, legumes, vegetables). In: Barigozzi, C.
(ed.) The origin and domestication of cultivated plants.
Pp. 179–99. Amsterdam, Elsevier.
Hanelt, P. 1990. Taxonomy, evolution and history. In:
Rabinowitz, H. D. and Brester, J. D. L. (eds.) Onions
and their allied crops. Pp. 2–26. Boca Raton, Florida:
CRC Press.
Hanelt, P. 1997. European wild relatives of Prunus fruit
crops. In: Valdés, B., Heywood, V. H., Raimondo, F. M.
REFERENCES 211
and Zohary, D. (eds.) Proceedings of the workshops on con-
servation of European cultivated plants. Pp. 409–16.
Palermo, Bocconea.
Hanelt, P., Schultze-Motel, J., Fritsch, R., Kruse, J.,
Maass, H., Ohle, H. and Pistrick, K. 1992. Infrageneric
grouping of Allium—the Gatersleben approach. In:
Hanelt, P., Hammer, K. and Knüpfer, H. (eds.) The
genus Allium—Taxonomic problems and genetic resources.
Pp. 107–23. Gatersleben, Proceedings of an interna-
tional symposium held at Gatersleben, Germany,
11–13 June 1991.
Hansen, J. M. 1988. Agriculture in the prehistoric Aegean:
data versus speculation. American Journal of Archaeology,
92, 39–52.
Hansen, J. M. 1991a. Palaeobotany in Cyprus: recent
research. In: Renfrew, J. M. (ed.) New light on early farm-
ing: recent developments in palaeoethnobotany. Pp. 225–36.
Edinburgh, Edinburgh University Press.
Hansen, J. M. 1991b. The Palaeoethnobotany of Franchthi
Cave, Bloomington and Indianapolis, Indiana University
Press.
Hansen, J. M. 1992. Franchthi Cave and the beginnings of
agriculture in Greece and the Aegean. In: Anderson-
Gerfaud, P. C. (ed.) Préhistoire de l’agriculture: nouvelles
approches expérimentales et ethnographiques. Pp. 231–47.
Paris, Monographie du Centre de Recherches
Archéologiques, CNRS.
Hansen, J. M. 1994. Khirokitia plant remains: preliminary
report (1986, 1988–1990). In: Brun, A. L. (ed.) Fouilles
récentes à Khirokitia (Chypre) 1988–1991. Pp. 393–409.
Paris, Éditions Recherche sur les Civilisations.
Harlan, J. R. 1975. Crop and man. Madison, American
Society of Agronomy.
Harlan, J. R. 1981. The early history of wheat: earliest
traces to the sack of Rome. In: Evans, L. T. and
Peacock, W. J. (eds.) Wheat science - today and tomor-
row. Pp. 1–19. Cambridge, UK: Cambridge University
Press.
Harlan, J. R. 1992a. Crops and man. Madison, Wis.,
American Society of Agronomy, Crop Science Society
of America.
Harlan, J. R. 1992b. Indigenous African agriculture. In:
Cowen, C. W. and Watson, P. J. (eds.) The origins of agri-
culture: an international perspective. Pp. 59–70. Washington
DC, Smithsonian Institution Press.
Harlan, J. R. 1995a. Barley. In: Smartt, J. and Simmonds, N.
W. (eds.) Evolution of crop plants. 2 ed, Pp. 140–6. London,
Longman.
Harlan, J. R. 1995b. The Living Fields. Cambridge, UK:
Cambridge University Press.
Harlan, J. R. and de Wet, J. M. J. 1971. Towards a rational
classification of cultivated plants. Taxon, 20, 509–17.
212 REFERENCES
Harlan, J. R. and de Wet, J. M. J. 1973. On the quality of the
evidence for origin and dispersal of cultivated plants.
Current Anthropology, 14, 51–62.
Harlan, J. R. and Stemler, A. B. L. 1976. The races of
Sorghum in Africa. In: Harlan, J. R., De Wet, J. M. J. and
Stemler, A. B. L. (eds.) The origins of African plant domes-
tication. Pp. 465–78. Mouton, The Hague.
Harlan, J. R. and Zohary, D. 1966. Distribution of wild
wheats and barley. Science, 153, 1074–80.
Harlan, J. R., de Wet, J. M. J. and Price, E. G. 1973.
Comparative evolution of cereals. Evolution, 27,
311–25.
Harris, D. R. 1986. Plant and animal domestication and the
origin of agriculture; the contribution of radiocarbon
accelerator dating. In: Gowlett, J. a. J. and Hedge, R. E.
M. (eds.) Archaeological results from accelerator dating.
Pp. 5–21. Oxford University Committee for Archaeology
Monograph II.
Harris, D. R. and Gosden, C. 1996. The beginning of agri-
culture in western Central Asia. In: Harris, D. R. (ed.)
The origins and spread of agriculture and pastoralism in
Eurasia. Pp. 370–89. London, University College London
Press.
Harris, S. A., Robinson, J. P. and Juniper, B. E. 2002. Genetic
clues to the origin of the apple. TRENDS in Genetics
18(8), 426–30.
Hartmann, A. 2006. Reconstruction of the human vegetarian
diet and landscape at Gilgal I—an Early Neolithic site in the
Jordan Valley: archaeobotanical research. Ph.D Thesis, Bar-
Ilan University.
Hartyáni, B. P. 1982. A Tiszaalpár-Várdomb bronzkori
lakóteleprol származó mag-és termésletek. (Die von
der bronzeseitlichen Wohnsiedlung Tiszaalpár-Várdomb
stammenden Korn- und Fruchtfunde). Cumania Archae-
ologia, 7, 133–286.
Hartyáni, B. P. and Nováki, G. 1975. Samen- und
fruchtfunde in Ungarn von der Neusteinzeit bis zum 18.
jahrhundert. Agrátörténeti Szemle: Historia Rerum
Rusticarum, 17, 1–88.
Hartyányi, B. P. and Nováki, G. 1971. Gabonalenyomatok
a Körös-csoport edényein (Getreideabdrücke auf den
Gefässen der Körös-Gruppe). Móra Ferenc Múzeum
Évkönyve, 2, 5–8.
Hartyányi, B., Nováki, G. and Patay, Á. 1968. Növényi
mag- és termésleletek Magyarországon az újkőkortól
a XVIII. sz.-ig I (Samen- und fruchtfunde in Ungarn
von der jungsteinzeit bis zum XVIII Jahrhundert I).
Magyar Mezőgazdasági Múzeum közleményei 1968,
5–85.
Havey, M. J. and Muehlbauer, F. J. 1989. Variability for
restriction fragment lengths and phylogenies in lentil.
Theoretical Applied Genetics, 77, 839–43.
Hedrick, U. P. 1919. Sturtevant’s edible plants of the World.
Report of the New York Agricultural Experimental Station
for the year 1919. New York, Dover Publications.
Hedrick, U. P., Howe, G. H., Taylor, O. M., Francis, E. H.
and Tukey, H. B. 1921. The pears of New York. NY, New
York State Department of Agriculture 29th annual
report.
Hegi, G. 1935. Illustrierte Flora von Mittel-Europa, Munich,
Hanser Verlag.
Heim, J. 1979. Recherche paléobotanique au site néo-
lithique (Roessen) de la ‘Bosse de la Tombe’ á Givry.
Bulletin de la Société Royale belge d’Anthropologie et de
Préhistoire, 90, 65–78.
Helbaek, H. 1953. Queen Ichetis’ wheat: a contribution to
the study of early dynastic emmer of Egypt. Kongelige
Danske Videnskabernes Selskab. Biologiske meddelelser,
21, 1–17.
Helbaek, H. 1954. Prehistoric food plants and weeds in
Denmark. A survey of archaeobotanical research 1923–
1954. Danmarks Geologisk Undersøgelse, 11, 250–61.
Helbaek, H. 1955. Ancient Egyptian wheats. Proceedings of
the Prehistory Society, 21, 93–5.
Helbaek, H. 1958. Plant economy in ancient Lachish. In:
Tufnell, O. (ed.) Lachish (Tell ed-Duweir) IV: the Bronze
Age. Pp. 309–17. Oxford, UK: Oxford University Press.
Helbaek, H. 1959a. Domestication of food plants in the old
world. Science, 130, 365–72.
Helbaek, H. 1959b. Notes on the evolution and history of
Linum. KUML: Årbog for Jysk Arkaeologisk Selskab, 1959,
103–29.
Helbaek, H. 1960. The paeoethnobotany of the Near East
and Europe. In: Braidwood, R. J. and Howe, B. (eds.)
Prehistoric investivations in Iraqi Kurdistan. Pp. 99–118.
Chicago, IL: The University of Chicago Press.
Helbaek, H. 1962. Les grains carbonisés de la 48ème
couche des fouilles de Tell Soukas. Annales Archéologiques
de Syrie, 11–12, 185–6.
Helbaek, H. 1963. Isin Larsan and Horian food remains of
Tell Basmosian in the Dokan Valley. Sumer, 19, 27–35.
Helbaek, H. 1964. First impressions of the Çatal Hüyük
plant husbandry. Anatolian Studies, 14, 121–3.
Helbaek, H. 1966a. Commentary on the phylogenesis
of Triticum and Hordeum. Economic Botany, 20,
350–660.
Helbaek, H. 1966b. The plant remains from Nimrud. In:
Mallowan, M. E. L. (ed.) Nimrud and its remains, vol. II.
Pp. 613–20. London, Collins.
Helbaek, H. 1966c. Pre-pottery Neolithic farming at Beidha
—a preliminary report. Palestine Exploration Quarterly,
98, 61–6.
Helbaek, H. 1969. Plant collecting, dry-farming, and irri-
gation agriculture in prehistoric Deh Luran. In: Hole, F.,

Flannery, K. V. and Neely, J. A. (eds.) Prehistory and
human ecology of the Deh Luran plain, early village sequence
from Khuzistan, Iran. Pp. 383–426. Ann Arbor, University
of Michigan.
Helbaek, H. 1970. The plant husbandry in Hacilar—a
study of cultivation and domestication. In: Mellaart, J.
(ed.) Excavation at Hacilar. Pp. 189–244. Edinburgh,
Edinburgh University Press.
Helbaek, H. 1972. Samarran irrigation agriculture at
Choga Mami in Iraq. Iraq, 34, 35–48.
Henriksen, M. B. and Runge, M. T. 2009. Archaeological
evidence of flax production 500 BC-1000 AD—examples
from Funen, Denmark. In: Karg, S. (ed.) International
FLAX network, Copenhagen. Communicating Culture,
Workshop Info 1, 2009.
Hepper, F. N. 1990. Pharaoh’s flowers, the botanical treasures
of Tutan-khamun. London, Her Majesty’s Stationary
Office.
Heppner, M. J. 1923. The factor for bitterness in the sweet
almond. Genetics, 8, 390–2.
Heppner, M. J. 1926. Further evidence on the factor for bit-
terness in the sweet almond. Genetics, 11, 605–6.
Hesse, C. O. 1975. Peaches. In: Janick, J. and Moore, J. N.
(eds.) Advances in fruit breeding. Pp. 285–347. West
Lafayette, Indiana: Purdue University Press.
Heun, M., Schafer-Pregl, R., Klawan, D., Castagna, R.,
Accerbi, M., Borghi, B. and Salamini, F. 1997. Site of
Einkorn Wheat Domestication Identified by DNA
Fingerprinting. Science, 278, 1312–14.
Heywood, V. H. 1983. Relationships and evolution in the
Daucus carota complex. Israel Journal of Botany, 32,
52–65.
Hill, G. D. 1995. Lupins. In: Smartt, J. and Simmonds, N. W.
(eds.) Evolution of crop plants. 2 ed, Pp. 277–82. London,
Longman.
Hillig, K. W. 2005. Genetic evidence for speciation in
Cannabis (Cannabaceae). Genetic Resources and Crop
Evolution, 52, 161–80.
Hillman, G. C. 1972. Plant remains, in French, D. H. et al.
Report on excavations at Can Hasan III 1969–1970. In:
Higgs, E. S. (ed.) Papers in economic prehistory. Pp. 182–8.
Cambridge, UK: Cambridge University press.
Hillman, G. C. 1975. The plant remains from Tell Abu
Hureyra: a preliminary report. Proceedings of the
Prehistory Society, 41, 70–3.
Hillman, G. C. 1978. On the origin of domestic rye—Secale
cereale: the finds from aceramic Can Hasan III in Turkey.
Anatolian Studies, 28, 157–74.
Hillman, G. C. 1984. Interpretation of archaeological plant
remains: The application of ethnographic models from
Turkey. In: van Zeist, W. and Casparie, W. A. (eds.)
Plants and ancient man. Pp. 1–41. Rotterdam, Balkema.
REFERENCES 213
Hillman, G. C. 1986. Plant foods in ancient diet: The role of
palaeofaeces in general and in Lindau’s man’s gut con-
tents in particular. In: Brothwell, I. M. and Bourke, J.
(eds.) Lindau Man: the body in the bog. Pp. 99–115. London,
British Museum.
Hillman, G. C. 1989. Late Palaeolithic plant foods from
Wadi Kubbaniya in Upper Egypt: dietary diversity,
infant weaning, and seasonality in a riverine environ-
ment. In: Harris, D. R. and Hillman, G. C. (eds.) Foraging
and farming: the evolution of plant exploitation. Pp. 207–39.
London, Unwin Hyman.
Hillman, G. C. 2000a. Abu Hureyra 1: the Epipalaeolithic.
In: Moore, A. M. T., Hillman, G. C. and Legge, A. J. (eds.)
Village on the Euphrates: from foraging to farming at Abu
Hureyra. Pp. 327–99. Oxford, Oxford University Press.
Hillman, G. C. 2000b. The plant food economy of Abu
Hureyra 1 and 2. In: Moore, A. M. T., Hillman, G. C. and
Legg, A. J. (eds.) Village on the Euphrates: from foraging to
farming at Abu Hureyra. Pp. 327–99. Oxford, Oxford
University Press.
Hillman, G. C. 2001. Archaeology, Percival, and the prob-
lems of identifying wheat remains. In: Caligari, P. D. S.
and Brandham, P. E. (eds.) Wheat taxonomy: the legacy of
John Percival. Pp. 27–36. London, Linnean Society.
Hillman, G. C. and Davies, M. S. 1999. Domestication rates
in wild-type wheats and barley under primitive cultiva-
tion. In: Anderson, P. (ed.) Prehistory of agriculture, new
experimental and ethnographic approaches. Pp. 70–102. Los
Angeles, The Institute of Archaeology, University of
California.
Hillman, G. C., Colledge, S. M. and Harris, D. R. 1989.
Plant-food economy during the Epipalaeolithic period
at Tell Abu Hureyra, Syria: dietary diversity, seasonal-
ity, and modes of exploitation. In: Harris, D. R. and
Hillman, G. C. (eds.) Foraging and farming: the evolution of
plant exploitation. Pp. 240–68. London, Unwin Hyman.
Hillman, G. C., Hedges, R., Moore, A. M. T., Colledge,
S. M. and Pettitt, P. 2001. New evidence of Lateglacial
cereal cultivation at Abu Hureyra on the Euphrates. The
Holocene, 11, 383–93.
Hillman, G. C., Mason, S., de Moulins, D. and Nesbitt, M.
1996. Identification of archaeological remains of wheat,
the 1992 London workshop. Circea, 12, 195–209.
Hiremath, S. C. and Patil, C. G. 1999. Genome homology
and putative progenitor of sesame (Sesamum indicum L.).
Journal of Cytology and genetics, 34, 69–71.
Hirschegger, P., Jakse, J., Trontelj, P. and Bohanec, B. 2010.
Origins of Allium ampeloprasum horticultural groups and a
molecular phylogeny of the section Allium (Allium:
Alliaceae). Molecular Phylogenetics and Evolution, 54, 488–97.
Hjelle, K. L. and Solem, T. 2008. Botaniske undersøkelser
—Ormen Lange Nyhamna. In: Bjerck, B. (ed.) NTNU
214 REFERENCES
Vitenskapamuseets arkeologiske undersøkelser Ormen Lange
Nyhamna. Pp. 477–545. Trondheim, Tapir forlag.
Hjelmqvist, H. 1955. Die älteste Geschichte der Kulturp-
flanzen in Schweden. Opera Botanica, 1, 1–186.
Hjelmqvist, H. 1979a. Beiträge zur Kenntnis der prähistor-
ischen Nutzpflanzen in Schweden. Opera Botanica, 17, 3–58.
Hjelmqvist, H. 1979b. Some economic plants and weeds
from the Bronze Age of Cyprus. Studies in Mediterranean
Archaeology, 45, 110–13.
Ho, P. T. 1977. The indigenous origins of Chinese agricul-
ture. In: Reed, C. A. (ed.) Origins of agriculture. Pp. 413–
83. Mouton, The Hague.
Hockett, E. A. and Nilan, R. H. 1985. Genetics. In:
Rasmusson, D. C. (ed.) Barley. Pp. 187–230. Madison,
WI: American Society of Agronomy.
Hodgkin, T. 1995. Cabbages, kales, etc. In: Smartt, J. and
Simmonds, N. W. (eds.) Evolution of crop plants. 2 ed,
Pp. 76–82. London, Longman, UK.
Hofmann, E. 1924. Pflanzenreste der Mondseer pfahlbau-
ten. Siizungsbtrichte der Akademie der Wissenschaften in
Wien (Mathematisch-Naturwissenschaftliche, 1, 379–409.
Hopf, M. 1955. Formveranderungen von Getreidekornern
beim Verkohlen. Berichte der Deutschen Botanischen
Gesellschaft, 68, 191–3.
Hopf, M. 1961a. Pflanzenfunde aus Lerna/Argolis.
Züchter, 31, 239–47.
Hopf, M. 1961b. Untersuchungsbericht über Kornfunde
aus Vršnik. Zbornik na Stipskot Naroden Muzej, 2, 41–5.
Hopf, M. 1962. Bericht über die untersuchungen von
samen und holzkohleresten von der Argissa-Magula
aus den präkeramischen bis mittel-bronze-zeitlichen
schichten. In: Milojčič, V., Boessneck, J. and Hopf, M.
(eds.) Die deutschen Ausgrabungen auf der Argissa-Magula
in Thessalien. Pp. 101–10. Bonn, Rudolf Habelt Verlag.
Hopf, M. 1973a. Frühe kulturpflanzen aus Bulgarien.
Jahrbuch Römisch-Germanisches Zentralmuseum, Mainz,
20, 1–47.
Hopf, M. 1973b. Pflanzenfunde aus Nordspanien: Cortes
de Navarra - El Soto de Medinilla. Madrider Mitteilungen,
14, 133–42.
Hopf, M. 1976. Bier. In: Beck, H. Jankuhn, H., Ranke, K.
and Wenskus, R. (eds.) Reallexikon der Germanischen
Altertumskunde. Pp. 530–3. Berlin, Walter de Gruyter.
Hopf, M. 1978a. Birne (Pyrus communis L.), Bohne (Vicia
faba L.). In: Beck, H., Jankuhn, H., Ranke, K. and
Wenskus, R. (eds.) Reallexikon der Germanischen
Altertumskunde. Pp. 29–32; 183–7. Berlin, Walter de
Gruyter.
Hopf, M. 1978b. Plant remains. In: Amiran, R. (ed.)
Early Arad I. The chalcolithic settlement and Early Bronze
Age city. Pp. 64–82. Jerusalem, Israel Exploration
Society.
Hopf, M. 1978c. Plant remains, strata V-I. In: Amiran, R. (ed.)
Early Arad I, the Chalcolithic Settlement and Early Bronze
City. Pp. 64–82. Jerusalem, The Israel Exploration Society.
Hopf, M. 1981. Pflanzliche reste aus Zambujal. In:
Sangmeister, E. and Schubart, H. (eds.) Zambujal, die
grabungen 1964 bis 1973. Pp. 315–40. Madrid, Mainz:
Madrider Beiträge Band 5, Deutsches Archäologisches
Institut, Verlag Philipp von Zabern.
Hopf, M. 1982. Vor- und frühgeschichtliche kulturpflanzen aus
dem nördlichen Deutschland, Mainz, Zentralmus.
Hopf, M. 1983. Jericho plant remains. In: Kenyon, K. M. and
Holland, T. A. (eds.) Excavations at Jericho. Pp. 576–621.
London, British School of Archaeology in Jerusalem.
Hopf, M. 1985. Bronzezeitliche sämereien aus Ouroux-
Marnay, Dép. Saône-et-Loire. Jahrbuch Römisch-German-
isches Zentralmuseum, Mainz, 32, 255–64.
Hopf, M. 1991. Kulturpflanzenreste aus der sammlung
Siret in Brüssel. In: Schubart, H. and Ulreich, H. (eds.)
Die funde der Südostspanischen Bronzezeit aus der sammlung
Siret. Pp. 397–413. Mainz, Madrider Beiträge Band 17.
Philipp von Zabern.
Hopf, M. and Muñoz, A. M. 1974. Neolithische pflanzen-
reste aus der Höhle Los Murciélagos bei Zuheros (Prov.
Córdoba). Madrider Mitteilungen, 15, 9–27.
Hopf, M. and Pellicer Catalán, M. 1970. Neolithische
getreidefunde in der Höhle von Nerja (Prov. Málaga).
Madrider Mitteilungen, 11, 18–34.
Hopf, M. and Schubart, H. 1965. Getreidefunde aus der
Coveta de l’Or (Prov. Alicante). Madrider Mitteilungen, 6,
20–38.
Hopf, M. and Willerding, U. 1989. Pflanzenreste. In: Kleiss,
W. (ed.) Bastam II. Pp. 263–318. Berlin, Mann.
Hosoya, L., Sato, Y.-I. and Fuller, D. 2010. Editorial: the
archaeobotany of early rice agriculture in Asia.
Archaeological and Anthropological Sciences, 2, 57–9.
Hovsepyan, R. and Willcox, G. 2008. The earliest finds of
cultivated plants in Armenia: evidence from charred
remains and crop processing residues in pisé from the
Neolithic settlements of Aratashen and Aknashen.
Vegetation History and Archaeobotany, 17, 63–71.
Huang, H., Cheng, Z., Zhang, Z. and Wang, Y. 2008.
History of cultivation and trends in China. In: Layne, D.
R. and Bassi, D. (eds.) The Peach: botany, production and
uses. Pp. 37–60. Wallingford, U.K.: CAB International.
Hubbard, R. N. L. 2000. Agriculture and environment at
Servia. In: Ridley, C., Wardle, K. A. and Mould, C. A.
(eds.) Servia: Anglo-Hellenic rescue excavations 1971–73
directed by Katerina Rhomiopoulou and Cressida Ridley.
Athens, Greece: British School at Athens, British
Academy, Institute for Aegean Prehistory.
Hunt, H. V., Linden, M. V., Liu, X., Motuzaite-
Matuzeviciute, G., Colledge, S. and Jones, M. K. 2008.

Millets across Eurasia: chronology context early Panicum
and Setaria Old World sites. Vegetation history and archae-
obotany, 17, S5–18.
Iezzoni, A., Schmidt, H. and Albertini, A. 1991. Cherries
(Prunus). In: Moore, J. N. and Ballington, J. R. (eds.) Genetic
resources of temperate fruit and nut crops. Pp. 111–73.
Wageningen, International Society for Horticultural Science.
Ihlenfeldt, H. D. and Grabow-Seidensticker, U. 1979. The
genus Sesamum L. and the origin of the cultivated ses-
ame. In: Kunkel, G. (ed.) IX Plenary Meeting of AETFAT:
Taxonomic aspects of African economic botany, 1978.
Pp. 53–60. Las Palmas de Gran Canaria, Excmo.
Ilett, M., Constantin, C., Farruggia, J. P. and Bakels, C.
1995. Bâtiments voisins de Rubané et du Groupe de
Villeneuve-St Germain sur le site Bucy-le-Long ‘La
Fosse Tounise’ (Aisne). Revue archéologique de Pacardie, 9:
spécial 9, 17–39.
Jaaska, V. 1997a. Isoenzyme differences between the wild
diploid and tetraploid wheats. Genetic Resources and
Crop Evolution, 44, 137–46.
Jaaska, V. 1997b. Isozyme diversity and phylopgenetic
affinities in Vicia sub-genus Vicia (Fabaceae). Genetic
Resources and Crop Evolution, 44, 557–74.
Jacomet, S. 1987. Prähistorische getreidefunde: eine anleitung
zur bestimmung prähistorischer gersten- und weizenfunde.
Basel, Botanisches Institut der Universität.
Jacomet, S. 1988. Pflanzen Mediterraner herkunft in
Neolithischen see-ufersiedlungen der Schweiz. In:
Küster, H. (ed.) Der prähistorische mensch und seine
umwelt, Festschrift für Udelgard Körber-Grohne. Pp. 205–
12. Stuttgart, Forschungen und Berichte zur Vor- and
Frühgeschichte in Baden-Württemberg, Band 31,
Konrad Theiss Verlag.
Jacomet, S. 2004. Archäobotanische grobuntersuchung
verkohlter getreideklumpen. In: Hügin, U. and Michel-
Tobler, C. (eds.) Oberrieden ZH-Riet—eine frühhorgen-
zeitliche siedlung. Jahrbuch der Schweizerischen gesellschaft
für Ur- und Frühgeschichte, 87, 21–3.
Jacomet, S. 2006. Plant economy of the Northern Alpine
lake dwelling area—3500–2400 BC cal. In: Karg, S.,
Baumeister, R., Schlichtherle, H. and Robinson, D.E.
(Eds.) Economic and environmental changes during the
4th and 3rd Millenia BC. Proceedings of the 25th sym-
posium of the AEA, Sept. 2004 in Bad Buchau, Germany.
Environmental Archaeology, 11, 64–83.
Jacomet, S. 2007. Neolithic plant economies in the north-
ern alpine foreland from 5500–3500 BC cal. In: Colledge,
S. and Conolly, J. (eds.) The origins and spread of domestic
plants in Southwest Asia and Europe. Pp. 221–58. Walnut
Creek CA, Left Coast Press.
Jacomet, S. 2008. Subsistenz und landnutzung während des
3. Jahrtausends v. Chr. aufgrund von archäobotanischen
REFERENCES 215
daten aus dem südwestlichen Mitteleuropa. In: Dörfler,
W. and Müller, J. (eds.) Umwelt—wirtschaft—siedlungen im
dritten vorchristlichen jahrtausend Mitteleuropas und
Südskandinaviens. Pp. 355–77. Neumünster, Wachholz.
Jacomet, S. and Behre, K.-E. 2009. Der mittelbronzezeitli-
che getreidefund. In: Lanzrein, A. N. (ed.) Die befestigte
Höhensiedlung Toos-Waldi von der Frühbronzezeit bis in die
Spätantike. Pp. 71–9. Frauenfeld, Amt für Archäologie
des Kantons Thurgau.
Jacomet, S. and Brombacher, C. 2005. Reconstructing intra-
site patterns in Neolithic lakeshore settlements: the state
of archaeobotanical research and future prospects. In:
Della Casa, P. and Trachsel, M. (eds.) WES’04—Wetland
Economies and Societies. Proceedings of the International
Conference in Zurich, 10–13 March 2004. Pp. 69–94.
Zürich, Chronos.
Jacomet, S., Brombacher, C. and Dick, M. 1989. Archäobotanik
am Zürichsee. ackerbau, sammelwirtschaft und umwelt von
Neolithischen und Bronzezeitlichen seeufersiedlungen im raum
Zürich, ergebnisse von untersuchungen pflanzlicher makroreste
der jahre 1979–1988. Zürich, Orell Füssli Verlag.
Jacomet, S., Brombacher, C. and Dick, M. 1991.
Palaeoethnobotanical work on Swiss Neolithic and
Bronze Age lake dwellings over the past ten years. In:
Renfrew, J. M. (ed.) New light on early farming: recent
developments in palaeoethnobotany. Pp. 257–76. Edinburgh,
Edinburgh University Press.
Jacomet, S., Brombacher, C. and Schraner, E. 1999.
Ackerbau und sammelwirtschaft während der Bronze-
und Eisenzeit in den östlichen Schweizer Alpen-
vorläufuge ergebnisse. In: Della Casa, P. (ed.) Prehistoric
alpine environment, society, and economy. Papers of the
international colloquium PAESE ‘97 in Zürich. Pp. 231–44.
Bonn, Rudolf Habelt.
Jacomet, S., Rachoud-Schneider, A.-M., Zoller, H. and
Burga, C. A. 1998. Vegetationsentwicklung, vegeta-
tionsveränderung durch menschlichen einfluss, acker-
bau und sammelwirtschaft. In: Hochuli, S., Niffeler, U.
and Rychner, V. (eds.) Bronzezeit. Pp. 141–70. Basel,
Verlag Schweizerische Gesellschaft für Ur- und
Frühgeschichte.
James, T. G. H. 1995. The earliest history of wine and its
importance in ancient Egypt. In: McGovern, P. T., Fleming,
S. J. and Katz, S. H. (eds.) The origins and ancient history of
wine. Pp. 197–213. Luxembourg, Gordon and Breach.
Janick, J., Forsline, P. L., Dickscon, E. E., Way, R. and
Thompson, M. (eds.) 2003. Collection, maintenance, char-
acterization and utilization of wild apples of Central Asia.
New Jersey, John Wiley and Sons.
Januševič, Z. V. 1986. Kul’turnye rastenija severnogo
pričernomor’ja. paleoetnobotaničeskie issedovanija. Kišinev,
Moldova: Ştiinţa.
216 REFERENCES
Januševič, Z. V. 1976. Kul’turnye rastenija Iugo-Zapada SSSR
po paleobotaničeskim issledovanijam. Kišinev, Moldova:
Ştiinţa.
Januševič, Z. V. 1978. Prehistoric food plants in the south-
west of the Soviet Union. Berichte der Deutschen
Botanischen Gesellschaft, 91, 59–66.
Januševič, Z. V. 1984. The specific composition of wheat
finds from ancient agricultural centres in the USSR. In:
van Zeist, W. and Casparie, W. A. (eds.) Plants and
ancient man. Pp. 267–76. Rotterdam, Balkema.
Jasny, N. 1944. The wheats of classical antiquity. Baltimore,
Johns Hopkins Press.
Jeffrey, C. 1975. Further notes on Cucurbitaceae: III. Some
Southern African taxa. Kew Bulletin, 30, 475–93.
Jeffrey, C. 1980. A review of Cucurbitaceae. Botanical
Journal of the Linnean Society, 81, 233–47.
Jensen, H. A. 1991. The Nordic countries. In: van Zeist, W.,
Wasylikowa, K. and Behre, K. -E. (eds.) Progress in Old
World palaeoethnobotany. Pp. 335–50. Rotterdam, Balkema.
Jessen, K. and Helbaek, H. 1944. Cereals of Great Britain
and Ireland in prehistoric and early historic times. Det
Kongelige Danske Videnskabernes Selskab. Biologiske
Skrifter, 3, 1–68.
Jha, K. 1987. Effect of the boiling and decanting method of
Khesari (Lathyrus sativus) detoxification on changes in
selected nutrients. Archivos Latinoamericanos de Nutrición,
37, 101–7.
Jones, D. L. 1995. Palms throughout the world. Washington
DC, Smithsonian Institution Press.
Jones, G. and Legge, A. J. 1987. The grape (Vitis vinifera L.)
in the Neolithic of Britain. Antiquity 61, 452–5.
Jones, G. and Legge, A. J. 2008. Evaluating the role of
cereal cultivation in the Neolithic: charred plant remains
from Hambledon Hill. In: Mercer, R. and Healy, F. (eds.)
Hambledon Hill, Dorset, England. Excavation and survey of
a Neolithic monument complex and its surrounding land-
scape. Pp. 469–76. Swindon, English Heritage.
Jørgensen, G. 1981. Korn fra Sarup; Med nogle bemae-
rkinnger om ager-bruget i yngre stenalder i Danmark.
KUML: Årbog for Jysk Arkaeologisk Selskab, 1981, 221–31.
Juniper, B. E. 2007. The mysterious origin of the sweet
apple. American Scientist, 95, 44–51.
Kadereit, J. W. 1986. A revision of Papaver L. sect. Papaver
(Papaveraceae). Botanische jahrbücher für systematik,
pflanzengeschichte und pflanzengeographie, 108, 1–16.
Kamenetsky, R. 2007. Garlic: botany and horticulture.
Horticultural Reviews, 33, 123–72.
Kanazin, V., Talbert, H., See, D., Decamp, P., Nevo, E. and
Blake, T. 2002. Discovery and assay of single-nucleotide
polymorphisms in barley (Hordeum vulgare). Plant molec-
ular biology, 48, 529–37.
Karg, S. and Müller, J. 1990. Neolithische getreidefunde aus
Pokrovnik, Dalmatien. Archäologisches Korrespondenzblatt,
20, 373–86.
Kazan, K., Muehlabuer, F. J., Weeden, N. W. and Ladizinsky,
G. 1993. Inheritance and linkage relationships of mor-
phological and isozyme loci in chickpea (Cicer arietinum
L.). Theoretical and Applied Genetics, 86, 417–26.
Kearney, J. and Smartt, J. 1995. The grasspea. In: Smartt, J.
and Simmonds, N. W. (eds.) Evolution of crop plants. 2 ed,
Pp. 266–70. London, Longman, UK.
Keimer, L. 1924. Die Gartenpflanzen im alten Ägypten,
Hamburg/Berlin, Hoffman u. Campe.
Keimer, L. 1984. Die Gartenpflanzen im alten Ägypten. Mainz,
Verlag Philipp von Zabern.
Kerber, E. R. and Rowland, G. G. 1974. Origin of the free-
threshing character in hexaploid wheat. Canadian Journal
of Genetics and Cytoloty, 16, 145–54.
Khush, G. S. 1963a. Cytogenetic and evolutionary studies
in Secale. III. Cytogenetics of weedy ryes and origin of
cultivated rye. Economic Botany, 17, 60–71.
Khush, G. S. 1963b. Cytogenetic and evolutionary studies
in Secale. IV. Secale vavilovii and its biosystematic status.
Zeitschrift für Pflanzenzüchtung, 50, 34–43.
Kihara, H. 1944. Die Entdeckung des DD-analysators beim
Weizen. Agriculture and Horticulture(Tokyo), 19, 889–90.
Kilian, B., Özkan, H., Walther, A., Kohl, J., Dagan, T.,
Salamini, F. and Martin, W. 2007. Molecular diversity at 18
Loci in 321 wild and 92 domesticate lines reveal no reduc-
tion of nucleotide diversity during Triticum monococcum
(einkorn) domestication: implications for the origin of
agriculture. Molecular Biology and Evolution, 24, 2657–68.
Kirkbride, J. H. 1993. Biosystematic monograph of the genus
Cucumis (Cucurbitaceae). Boone, North Carolina Parkway
Publishers.
Kirleis, W. and Willerding, U. 2008. Die pflanzenreste aus
der Linienbandkeramischen siedlung von Rosdorf-
Mühlengrund, Ldkr. Göttingen, im südöstlichen
Niedersachsen. Praehistorische Zeitschrift, 83, 133–78.
Kislev, M. E. 1980. Triticum parvicoccum sp. nov., the oldest
naked wheat. Israel Journal of Botany, 28, 95–107.
Kislev, M. E. 1985. Early Neolithic Horsebean from
Yiftah’el, Israel. Science, 228, 319–20.
Kislev, M. E. 1987. Chalcolithic plant husbandry and
ancient vegetation at Shiqmim. In: Levy, T. E. (ed.)
Shiqmim I, Studies Concerning Chalcolithic Societies in the
Northern Negev Desert, Israel (1982–1984). Pp. 251–79,
549–63. Oxford, BAR International Series.
Kislev, M. E. 1988. Nahal Hemar cave: desiccated plant
remains, an interim report. Atiqot, 18, 76–81.
Kislev, M. E. 1989a. Origin and cultivation of Lathyrus sati-
vus and L. cicera (Fabaceae). Economic Botany, 43, 262–70.

Kislev, M. E. 1989b. Pre-Domesticated Cereals in the Pre-
Pottery Neolithic A Period. In: Hershkovitz, I. (ed.)
People and Culture in Change. Pp. 147–51. Oxford, BAR
International Series.
Kislev, M. E. 1992. Vegetal food of Bar Kokhba rebels at
Abi’or Cave near Jericho. Review of Palaeobotany and
Palynology, 73, 153–60.
Kislev, M. E. 1994–5. Wild olive stones at submerged, chal-
colithic Kfar Samir, Haifa, Israel. Journal of the Israel
Prehistoric Society, 26, 134–45.
Kislev, M. E. 1997. Early agriculture and palaeoecology of
Netiv Hagdud. In: Bar-Yosef, O. and Gopher, A. (eds.)
An early Neolithic village in the Jordan Valley. Part 1: The
archaeology of Netiv Hagdud. Pp. 201–36. Cambridge,
MA: Peabody Museum of Archaeology and Ethnology,
Harvard University.
Kislev, M. E. 2002. Origin of Annual crops by agro-evolu-
tion. Israel Journal of Plant Sciences, 50, S85–8.
Kislev, M. E. 2009. Reconstructing the ear morphology of
ancient small-grain wheat (Triticum turgidum ssp. parvic-
occum). In: Fairbairn, A. and Weiss, E. (eds.) From foragers
to farmers: papers in honour of Gordon C. Hillman. Pp. 235–
8. Oxford, Oxbow Books.
Kislev, M. E. and Hartmann, A. forthcoming. Food crops
from Nahal Zehora II. In: Gopher, A. (ed.) Village xom-
munities of the Pottery Neolithic Period in the Menashe
Hills, Israel; archaeological investigations at the sites of
Nahal Zehora. Tel Aviv, Tel Aviv University, Sonia and
Marco Nadler Institute of Archaeology, Monograph
Series.
Kislev, M. E. and Rosenzweig, S. 1991. Influence of experi-
mental charring of seed dimensions of pulses. Acta
Interdisciplinaria Archaeologica, 7, 143–57.
Kislev, M. E., Artzy, M. And Marcus, E. 1993. Import of an
Aegean food plant to a Middle Bronze IIA coastal site in
Israel. Levant, 25, 145–54.
Kislev, M. E., Hartmann, A. and Bar-Yosef, O. 2006a. Early
domesticated fig in the Jordan Valley. Science, 312,
1372–4.
Kislev, M. E., Hartmann, A. and Bar-Yosef, O. 2006b.
Response to comment on ‘Early domesticated fig in the
Jordan Valley’. Science, 314, 1683.
Kislev, M. E., Hartmann, A. and Galili, E. 2004. Archaeo-
botanical and archaeoentomological evidence from a
well at Atlit-Yam indicates colder, more humid cli-
mate on the Israeli coast during the PPNC period.
Journal of Archaeological Science , 31 , 1301–10.
Kislev, M. E., Nadel, D. and Carmi, I. 1992. Epipalaeolithic
(19,000 BP) cereal and fruit diet at Ohalo II, Sea of
Galilee, Israel. Review of Palaeobotany and Palynology, 73,
161–6.
REFERENCES 217
Kjellberg, F., Gouyon, P. H., Ibrahim, M., Raymond, M. and
Valdeyron, G. 1987. The stability of the symbiosis between
dioecious figs and their pollinators: a study of Ficus carica
L. and Blastophaga psenes L. Evolution, 41, 693–704.
Klichowska, M. 1976. Aus palaeoethnobotanischen stud-
ien über pflanzen-funde aus dem Neolithikum und der
Bronzezeit auf polnischem boden. Archaeologia Polona,
17, 27–67.
Knörzer, K.-H. 1971. Prähistorische mohnsamen im
Rheinland. Bonner Jahrbücher, 171, 34–9.
Knörzer, K.-H. 1973. Der bandkeramische siedlungsplatz
Langweiler 2: pflanzliche grossreste. Rheinische
Ausgrabungen, 13, 139–52.
Knörzer, K.-H. 1974. Bandkeramische pflanzenfunde von
Bedburg-Garsdorf, Kreis Bergheim/Erft. Rheinische
Ausgrabungen, 15, 173–92.
Knörzer, K.-H. 1991. Geschichte der rübe (Beta vulgaris L.)
mit beiträgen durch grossrestfunde vom Niederrhein. In:
Hajnalová, E. (ed.) Palaeoethnobotany and archaeology:
8th symposium of the international work-group for pal-
aeoethnobotany, Nitra-Nové Vozokany 1989. Acta
Interdiciplinaria Archaeologica, 7, 159–64.
Knörzer, K.-H. 1997. Botanische untersuchungen von 16
Neolithischen siedlungsplätzen im bereich der
Aldenhovener Platte, Kr. Düren und Aachen. In:
Lüning, J. (ed.) Studien zur Neolithischen besiedlung der
Aldenhovener Platte und ihrer umgebung. Pp. 647–84. Köln,
Rheinische Ausgrabungen.
Knörzer, K.-H. 1998. Botanische untersuchungen am
Bandkeramischen brunnen von Erkelenz-Kückhoven. In:
Koschik, H. (ed.) Brunnen der Jungsteinzeit. Pp. 229–46.
Köln-Bonn, Materialien zur Bodendenkmalpflege im
Rheinland 11. Köln und Bonn, Rheinisches Amt für
Bodendenkmalpflege.
Knowles, P. F. and Ashri, A. 1995. Safflower. In: Smartt, J.
and Simmonds, N. W. (eds.) Evolution of crop plants. 2 ed,
Pp. 358–66. London, Longman.
Kobylyanskyi, V. D. 1989. Rye. In: Shishkin, B. K. (ed.)
Flora of Cultivated plants of the USSR. Leningrad,
Agropromizdat.
Kohler-Schneider, M. 2001. Verkohlte kultur- und wildpflanzen-
reste aus Stillfried an der March als spiegel spätbronzezeitlicher
landwirtschaft im weinviertel, Niederösterreich, Wien, Verlag
der Österreichischen Akademie der Wissenschaften.
Kohler-Schneider, M. 2003. Contents of a storage pit from
Late Bronze Age Stillfried, Austria: another record of the
‘new’glume wheat. Vegetation History and Archaeobotany,
12, 105–11.
Kohler-Schneider, M. 2007. Early agriculture and subsist-
ence in Austria: a review of neolithic plant records. In:
Colledge, S. and Conolly, J. (eds.) The origins and spread of
218 REFERENCES
domestic plants in Southwest Asia and Europe. Pp. 209–20.
Walnut Creek, CA: Left Coast Press.
Kohler-Schneider, M. and Caneppele, A. 2009. Late
Neolithic agriculture in eastern Austria: archaeobotani-
cal results from sites of the Baden and Jevišovice cul-
tures (3600–2800 B.C.). Vegetation History and Archaeobotany,
18, 61–74.
Komatsuda, T. and Mano, Y. 2002. Molecular mapping of
the intermedium spike-c (int-c) and non-brittle rachis 1
(btr1) loci in barley(Hordeum vulgare L.). Theoretical and
applied genetics, 105, 85–90.
Komatsuda, T., Pourkheirandish, M., He, C., Azhaguvel,
P., Kanamori, H., Perovic, D., Stein, N., Graner, A.,
Wicker, T., Tagiri, A., Lundqvist, U., Fujimura, T.,
Matsuoka, M., Matsumoto, T. and Yano, M. 2007. Six-
rowed barley originated from a mutation in a homeodo-
main-leucine zipper I-class homeobox gene. Proceedings
of the National Academy of Sciences, 104, 1424–9.
König, M. 1989. Ein fund Römerzeitlicher traubenkerne in
Piesport/Mosel. Dissertationes Botanicae, 133, 107–16.
Koopman, W. J. M., Guetta, E., van der Weil, C. C. M.,
Vosman, B. and van den Berg, R. G. 1998. Phylogenetic
relationships among Lactuca (Asteraceae) species and
related genera based on ITS-1 DNA sequences. American
Journal of Botany, 85, 1517–30.
Korban, S. S. 1986. Interspecific hybridization in Malus.
HortScience, 21, 41–8.
Körber-Grohne, U. 1987. Nutzpflanzen in Deutschland.
Stuttgart, Konrad Theiss.
Körber-Grohne, U. 1991. The determination of fibre plants
in textiles, cordage and wickerwork. In: Renfrew, J. (ed.)
New light on early farming: recent developments in pal-
aeoethnobotany. Pp. 93–104. Edinburgh, Edinburgh
University Press.
Körber-Grohne, U. 1996. Pflaumen, Kirschpflaumen,
Schlehen, heutige pflanzen und ihre geschichte seit der
frühzeit. Stuttgart, Konrad Theiss.
Kostoff, D. 1937. Interspecific hybrids in Secale (rye) I:
Secale cereale x Secale ancestrale, Secale cereale x Secale
vavilovii, Secale cereale x Secale montanum, Secale ancestrale
x Secale vavilovii hybrids. Compte rendu de l’académie des
sciences d l’U.R.S.S, 14, 213–14.
Kranz, A. R. 1961. Cytologische untersuchungen und
genetische beobachtungen an den bastarden zwischen
Secale cereale L. und Secale vavilovii Grossh. Züchter, 31,
219–25.
Kreuz, A. 1994–5. Landwirtschaft und ihre ökologischen
grundlagen in den jahrhunderten um Christi geburt:
uum stand der naturwissenschaftlichen untersuchungen
in Hessen. Berichte der Kommission für Archäologische
Landesforschung in Hessen, 3, 59–91.
Kreuz, A. 2007. Archaeobotanical perspectives on the begin-
ning of agriculture north of the Alps. In: Colledge, S. and
Connoly, J. (eds.) The origins and spread of domestic plants
in Southwest Asia and Europe. Pp. 259–94. Walnut Creek,
CA: Left Coast Press.
Kreuz, A. and Boenke, N. 2002. The presence of two-
grained einkorn at the time of the Bandkeramik cul-
ture. Vegetation History and Archaeobotany, 11, 233–40.
Krochmal, A. and Krochmal, C. 1974. The complete illus-
trated book of dyes from natural sources. New York,
Doubleday and Co.
Kroll, H. (1995–2000). Literature on archaeological remains
of cultivated plants. For 1992/93: Vegetation History and
Archaeobotany, 4, 51–66 (1995). For 1994–5: Vegetation
History and Archaeobotany, 5, 169–200 (1996). For 1995–6:
Vegetation History and Archaeobotany, 6, 25–67 (1997). For
1996–7: Vegetation History and Archaeobotany, 7, 23–56
(1998). For 1997–8: Vegetation History and Archaeobotany,
8, 129–163 (1999a). For 1998–9: Vegetation History and
Archaeobotany, 9, 31–68, 2000.
Kroll, H. 1975. Pflanzlische reste aus subfossilen acker-
böden der Bronze-und Wikingerzeitlichen siedlungen
in Archsum auf Sylt, Studies in Palaeoethnobotany
(Schleswig-Holstein). Folia Quaternaria, 46, 31–4.
Kroll, H. 1979. Kulturpflanzen aus Dimini. Archaeo-Physika,
8, 173–89.
Kroll, H. 1980. Einige vorgeschichtliche vorratsfunde von
kulturpflanzen aus Norddeutschland. Offa, 37, 372–83.w
Kroll, H. 1981a. Mittelneolithisches getreide aus Dannau.
Offa, 38, 85–90.
Kroll, H. 1981b. Thessalische kulturpflanzen. Zeitschrift für
Archäologie, 15, 97–103.
Kroll, H. 1982. Kulturpflanzen von Tiryns. Archäologischer
Anzeiger, 4, 467–85.
Kroll, H. 1983. Kastanas: die pflanzenfunde ausgrabungen
in einem siedlungshügel der Bronze- und Eisenzeit
Makedoniens 1975–1979. In: Hänsel, B. (ed.) Práhistorische
archäologie in südosteuropa. Pp. 1–176. Berlin, Verlag
Volker Spiess.
Kroll, H. 1984. Bronze Age and Iron Age agriculture in
Kastanas, Macedonia. In: van Zeist, W. and Casparie, W. A.
(eds.) Plants and ancient man. Pp. 243–6. Rotterdam,
Balkema.
Kroll, H. 1991. Südosteuropa. In: van Zeist, W., Wasylikowa,
K. and Behre, K. -E. (eds.) Progress in Old World palaeoeth-
nobotany. Pp. 161–77. Rotterdam, Balkema.
Kroll, H. 1999b. Vor- und frühgeschichtliche weinreben -
wild oder angebaut? Eine abschliesende bemerkung.
Trierer Zeitschrift, 62, 151–3.
Kučan, D. 1995. Zur ernährung und dem gebrauch von
pflanzen im Heraion von Samos im 7. Jahrhundert v.
Chr. Jahrbuch des Deutschen Archäologischen Instituts,
110, 1–64.
Kuckuck, H. 1973. Utilization of Iranian genetic material.
Wheat Information Service, 36, 21–6.

Kuckuck. H. and Schiemann, E. 1957. Über das vorkom-
men von spelz und emmer (Triticum spelta und T. dicoc-
cum Schübl.) im Iran. Zeit für Pflanzenzüchtg, 38, 383–96.
Kühn, F. 1981. Rozbory nálezu polních plodin. Pp.75–9.
Prehled Výzkumu 1979. Archeologicky ústav. Brně, CSAV.
Kuriachan, P. and Beevy, S. S. 1992. Occurrence and chro-
mosome number of Cucumis sativus var. hardwickii
(Royle) Alef. in South India and its bearing on the origin
of cultivated cucumber. Euphytica, 61, 131–3.
Küster, H. 1991. Mitteleuropa südlich der Donau, ein-
schliesslich Alpenraum. In: van Zeist, W., Wasylikowa,
K. and Behre, K.-E. (eds.) Progress in Old World palaeoeth-
nobotany. Pp. 179–87. Rotterdam, Balkema.
Kuzminova, N., Dergačev, V. and Larina, O. 1998.
Paleoetnobotaničeskie issledovanija na poselenii
Sakarovka I. Revista Arheologică, 2, 166–82.
Kvavadze, E., Bar-Yosef, O., Belfer-Cohen, A., Boaretto, E.,
Jakeli, N., Matskevich, Z. and Meshvelani, T. 2009.
30,000-year-old wild flax fibres. Science, 325, 13–59.
Kvavadze, E., Bar-Yosef, O., Belfer-Cohen, A., Boaretto, E.,
Jakeli, N., Matskevich, Z. and Meshveliani, T. 2010.
Response to Comment on ‘30,000-Year-Old wild flax
fibres‘. Science, 328, 1634-c.
La Valva, V., Sabato, S. and Gigliano, G. S. 1985.
Morphology and alkaloid chemistry of Papaver setigerum
DC. (Papaveraceae). Taxon, 34, 191–6.
Ladizinsky, G. 1975. On the origin of the broad bean, Vicia
faba L. Israel Journal of Botany, 24, 80–8.
Ladizinsky, G. 1979. The genetics of several morphological
traits in the lentil. The Journal of Heredity, 70, 135–7.
Ladizinsky, G. 1993. Wild lentils. Critical Reviews in Plant
Sciences, 12, 169–84.
Ladizinsky, G. 1999. Identification of the lentil’s wild genetic
stock. Genetic Resources and Crop Evolution, 46, 115–18.
Ladizinsky, G. and Abbo, S. 1993. Wild lentils of central Asia.
FAO/IBPGR Plant Genetic Resources Newsletter, 93, 5–8.
Ladizinsky, G. and Adler, A. 1976. The origin of chickpea
Cicer arietinum L. Euphytica, 25, 211–17.
Ladizinsky, G. and van Oss, H. 1984. Genetic relationship
between wild and cultivated Vicia ervilia. Botanical
Journal of the Linnean Society, 89, 97–100.
Ladizinsky, G. and Vosa, C. G. 1986. C-banding in Trigonella
section Foenum-graecum (Fabaceae). Plant Systematics
and Evolution, 153, 1–5.
Ladizinsky, G. and Zohary, D. 1971. Notes on species
delimitation, species relationships and polyploidy in
Avena L. Euphytica, 20, 380–95.
Ladizinsky, G., Braun, D., Goshen, D. and Muehlbauer, F.
J. 1984. The biological species of the genus Lens L.
Botanical Gazette, 145, 253–61.
Lagerstedt, H. B. 1975. Filberts. In: Janick, J. and Moore,
J. N. (eds.) Advances in fruit breeding. Pp. 456–89. West
Lafayette, Indiana: Purdue University Press.
REFERENCES 219
Larsson, M. 1992. The Early and Middle Neolithic Funnel
Beaker culture in the Ysatd area (Southern Scandinavia),
economic and social change 3100–2300 BC. In: Larsson,
L., Callmer, J. and Stjernquist, B. (eds.) The archaeology of
the cultural landscape. field work and research in a South
Swedish rural region. Pp. 17–91. Stockholm, Acta
Archaeologica Lundensia.
Lauer, J. P., Laurent-Tackholm, V. and Åberg, E. 1950. Les
plantes découvertes dans les souterrains de l’enceinte
du roi Zoser à Saqqarah (IIIe dyn.). Bulletin de l’Institut
d’Égypte, 32, 121–47.
Lavee, S. and Zohary, D. 2011-In Press. Potential of genetic
diversity and effect of geographic isolated resources in
olive breeding. Israel Journal of Plant Science.
Lawes, D. A. 1980. Recent developments in understand-
ing, improvement, and use of Vicia faba. In: Summerfield,
R. J. and Bunting, A. H. (eds.) Advances in legume science.
Pp. 625–36. Richmond, UK: Royal Botanic Gardens,
Kew.
Le Thierry D’ennequin, M., Panaud, O., Toupance, B. and
Sarr, A. 2000. Assessment of genetic relationships
between Setaria italica and its wild relative S. viridis
using AFLP markers. Theoretical and Applied Genetics,
100, 1061–6.
Lee, J. 1984. Cotton as a world crop. In: Kohel, R. J. and
Lewis, C. F. (eds.) Cotton. Pp. 1–25. Madison, Wisconsin:
American Society of Agronomy.
Leggett, J. M. and Thomas, H. 1995. Oat evolution and
cytogenetics. In: Welch, W. (ed.) The oat crop: production
and utilization. Pp. 121–49. London, Chapman and Hall.
Lemmens, R. H. M. J. and Wessel-Riemens, P. C. 1991.
Indigofera L. In: Lemmens, R. H. M. J. and Wulijarni-
Soetjipto, N. (eds.) Plant resources of South-East Asia No. 3:
dye and tanin-producing plants. Pp. 81–3. Wageningen,
Pudoc/Prosea.
Lenz, H. O. 1859. Botanik der alten Griechen und Römer.
Wiesbaden, Sändig oHG.
Leroi-Gourhan, A. 1981. Diagrammes polliniques des sites
archéologiques au Moyen Orient. In: Frey, W. and
Ürpmann, H.-P. (eds.) Beiträge zum Umweltgeschichte des
Vorderen Orients. Pp. 121–33. Wiesbaden, Germany:
Ludwig Reichert.
Lev, E., Kislev, M. E. and Bar-Yosef, O. 2005. Mousterian
vegetal food in Kebara Cave, Mt. Carmel. Journal of
Archaeological Science, 32, 475–84.
Levadaux, L. 1956. Les populations sauvages et cultivées
des Vitis vinifera L. Annales de l’Amélioration des
Plantes, 1, 59–118.
Lev-Yadun, S., Gopher, A. and Abbo, S. 2000. The cradle of
agriculture. Science, 288, 1602–3.
Lev-Yadun, S., Ne’eman, G., Abbo, S. and Flaishman, M. A.
2006. Comment on ‘Early Domesticated Fig in the Jordan
Valley’. Science, 314, 1683.
220 REFERENCES
Li, H. L. 1974. An archaeological and historical account of
Cannabis in China. Economic Botany, 28, 437–48.
Libby, W. F., Anderson, E. C. and Arnold, J. R. 1949. Age
determination by radiocarbon content: world-wide
assay of natural radiocarbon. Science, 109, 227–8.
Liphschitz, N. 1987. Ceratonia siliqua in Israel: an ancient
element or a newcomer? Israel Journal of Botany, 36,
191–7.
Liphschitz, N. 2008. Timber in ancient Israel: dendroarchaeol-
ogy and dendrochronology. Tel Aviv University, Tel Aviv:
Emery and Claire Yass Publications in Archaeology,
Institute of Archaeology.
Liphschitz, N. and Nadel, D. 1997. Charred wood remains
from Ohalo II (19000 B.P.), Sea of Galilee, Israel. Journal
of Israel Prehistorical Society, 27, 5–18.
Liphschitz, N., Gophna, R., M., H. and Biger, G. 1991. The
beginning of olive (Olea europaea) cultivation in the Old
World: a reassessment. Journal of Archaeological Science,
18, 441–53.
Lisitsina, G. N. 1978. Main types of ancient farming on the
Causcasus, on the basis of a palaeo-ethnobotanical
research. Berichte der Deutschen Botanischen Gesellschaft,
91, 47–57.
Lisitsina, G. N. 1984. The Caucasus—a centre of ancient
farming in Eurasia. In: van Zeist, W. and Casparie, W. A.
(eds.) Plants and ancient man. Pp. 285–92. Rotterdam,
Balkema.
Lisitsina, G. N. and Prishchepenko, L. V. 1977. Paleoetno-
botanicheskiye nakhodki Kavkaza i Blizhnego Vostoka.
Moskva, ‘Nauka’.
Lityńska-Zając, M. 1993. Polish archaeobotanical studies
in north Africa: Armant (Egypt). Wiadomosci Botaniczne,
37, 171–2.
Lityńska-Zając, M. 1997. Wyniki badan odcisków
roślinnych z neolitycznego stanowiska w Strachowie,
woj. Wrocław (Investigation results of plant imprints
from the Neolithic site at Strachow, Wrocław province).
In: Kulczycka-Leciejewiczowa, A. (ed.) ‘Strachów. Osiedla
neolitycznych rolników na Ślasku’. Pp. 267–77. Wrocław,
Instytut Archeologii i Etnologii PAN.
Lityńska-Zając, M. 2007. Early Neolithic agriculture in
south Poland as reconstructed from archaeobotanical
plant remains. In: Colledge, S. and Conolly, J. (eds.) The
origins and spread of domestic plants in Southwest Asia
and Europe. Pp. 315–26. Walnut Creek, CA: Left Coast
Press.
Livarda, A. and van der Veen, M. 2008. Social access and
dispersal of condiments in North-West Europe from the
Roman to the medieval period. Vegetation History and
Archaeobotany, 17, 201–9.
Loeschke, S. 1933. Denkmäler vom weinbau aus der zeit
der Römerherrschaft an Mosel, Saar, und Ruwer. Trier,
Selbstverlag der Römischen Abteilung des Deutschen
Weinmuseums.
Lopez, P. 1980. Estudio des semillas prehistoricas algunos
yacimientos Españoles. Trabajos de Prehistoria, 37, 419–32.
López, P., Arnanz, A. M., Macías, R., Uzquiano, P. and Gil,
P. 2003. Arqueobotánica de la Cueva de La Vaquera. In:
Estremera, M. S. (ed.) Primeros agricultores y ganaderos en
la Meseta Norte: el Neolítico de la Cueva de La Vaquera
(Torreiglesias, Segovia). Pp. 247–55. Zamora, Junta de
Castilla y León.
Loskutov, I. G. 2001. Interspecific Crosses in the Genus
Avena L. Russian Journal of Genetics, 37, 467–75.
Loskutov, I. G. 2008. On evolutionary pathways of Avena
species. Genetic Resources and Crop Evolution, 55, 211–20.
Love, H. H. and Craig, W. T. 1924. The Genetic relation
between Triticum dicoccum dicoccoides and a similar mor-
phological type produced synthetically. Journal of
Agriculture Research, 28, 515–19.
Lu, H., Zhang, J., Liu, K.-B., Wu, N., Li, Y., Zhou, K., Ye, M.,
Zhang, T., Zhang, H., Yang, X., Shen, L., Xu, D. and Li,
Q. 2009. Earliest domestication of common millet
(Panicum miliaceum) in East Asia extended to 10,000
years ago. Proceedings of the National Academy of Sciences,
106, 7367–72.
Lundström-Baudais, K. 1984. Palaeoethnobotanical inves-
tigation of plant remains from a Neolithic lakeshore site
in France: Clairvaux, Station III. In: van Zeist, W. and
Casparie, W. A. (eds.) Plants and ancient man. Pp. 293–
305. Rotterdam, Balkema.
Lundström-Baudais, K. (In press). Les paléosemences des
fosses néolithiques du Petit Chasseur IV. In: Besse, M.
and Piguet, M. (eds.) Le site du Petit Chasseur à Sion
(Valais, Suisse): un hameau du Néolithique moyen.
Lausanne, Archeologia Vallesiana.
Luo, M. C., Yang, Z. L., You, F. M., Kawahara, T., Waines, J.
G. and Dvorak, J. 2007. The structure of wild and domes-
ticated emmer wheat populations, gene flow between
them, and the site of emmer domestication. Theoretical
and Applied Genetics, 114, 347–959.
Maan, S. S. 1973. Cytoplasmic and cytogenetic relation-
ships among tetraploid Triticum species. Euphytica, 22,
287–300.
Mackey, J. 1966. Species relationships in Triticum. Proceeding
of the 2nd international Wheat genetics symposium, Lund,
Sweden 1963. Pp. 237–76. Lund, Sweden: Berlingska
Boktryckeriet Lund.
Maggioni, L., Von Bothmer, R., Poulsen, G. and Branca, F.
2010. Origin and domestication of Cole crops (Brassica
oleracea L.): linguistic and literary considerations.
Economic Botany, 64, 109–23.
Maier, U. 1995. Moorstratigraphische und paläobota-
nische Untersuchungen in der jungsteinzeitlichen

Moorsiedlung Ödenahlen am Federsee. Forschungen
und Berichte zur Vor- und Frühgeschichte in Baden-
Württemberg, 46, 143–253.
Maier, U. 1996. Morphological studies of free-threshing
wheat ears from a Neolithic site in southwest Germany,
and the history of naked wheats. Vegetation History and
Archaeobotany, 5, 39–55.
Maier, U. 2001. Archäobotanische untersuchungen in der
Neolithischen ufersiedlung Hornstaad-Hörnle IA am
Bodensee. In: Maier, U. and Vogt, R. (eds.) Siedlun-
gsarchäologie im Alpenvorland VI. Botanische und pedolo-
gische Untersuchungen zur Ufersiedlung Hornstaad-Hörnle
IA. Pp. 9–384. Stuttgart, Konrad Theiss Verlag.
Malzew, A. I. 1930. Ovsyugi i ovsy. Sectio Euavena Griseb
(Wild and cultivated oats. Sectio Euavena Griseb.). Works
of applied botany and plant breeding, Supplement 38. VIR,
Leningrad (Russian).
Manning, K., Pelling, R., Higham, T., Schwenniger, J.-L.
and Fuller, D. Q. 2011. 4500-Year old domesticated pearl
millet (Pennisetum glaucum) from the Tilemsi Valley,
Mali: new insights into an alternative cereal domestica-
tion pathway. Journal of Archaeological Science, 38,
312–22.
Marinova, E. 2004. Archaeobotanische ergebnisse aus der
Bronzezeit von Tell Karanovo und ihr regionaler kon-
text. Diomedes, 3, 53–8.
Marinova, E. 2006. Vergleichende palaeoethnobotanische
Untersuchung zur Vegetationsgeschichte und praehis-
torischen Landnutzung in Bulgarien. Dissertationes
Botanicae 401. Stuttgart, E. Schweizerbart science
publishers.
Marinova, E. 2007. Archaeobotanical data from the early
Neolithic of Bulgaria. In: Connolly, J. and Colledge, S.
(eds.) Early Neolithic in Southwest Asia and Europe: archae-
obotanical perspectives in Neolithic plant economies. Pp.
93–109. Walnut Creek, CA: University College London,
Institute of Archaeology, Left Coast Press.
Marinova, E. and Riehl, S. 2009. Carthamus species in the
ancient Near East and south-eastern Europe: archaeobo-
tanical evidence for their distribution and use as a source
of oil. Vegetation History and Archaeobotany, 18, 341–9.
Marinova, E. forthcoming. Archaeobotanical Studies. In:
Nikolov, V. (ed.) Kapitan Dimitrievo—the excavations of
2001–2004. Sofia, BAN.
Marinval, P. 1995. Recollecció i agricultura de l’epipaleolític
al neolític antic: Anàlisi carpològica de la Balma de la
Margineda. In: Guilaine, J. and Martzluff, M. (eds.) Las
excavacionsa la Balma de la Margineda. Pp. 65–77. Andorra,
Edicions del Govern d’Andorra.
Marinval, P. 2007. Premières données sur l’économie
végétale. In: J., G., C., M. and J.D., V. (eds.) Pont de
Roque-Haute. Nouveaux regards sur la Néolithisation de la
REFERENCES 221
France méditerranéenne. Pp. 215–17. Toulouse, Archives
d’Ecologie Préhistorique.
Markgraf, F. 1975. Camelina sativa (L.) Crantz. In: Hegi, G.
(ed.) Illustrierte Flora von Mitteleuropa. 2 ed, Pp. 342–5.
Berlin, Verlag Paul Parey.
Märkle, T. and Rösch, M. 2008. Experiments on the effects
of carbonization on some cultivated plant seeds.
Vegetation History and Archaeobotany, 17, 257–63.
Martin, L., Jacomet, S. and Thiébault, S. 2008. Plant econ-
omy during the Neolithic in a mountain context: the
case of ‘Le Chenet des Pierres’ in the French Alps (Bozel-
Savoie, France). Vegetation History and Archaeobotany, 17,
113–22.
Martinoli, D. and Jacomet, S. 2004. Identifying endocarp
remains and exploring their use at Epipalaeolithic
Öküzini in southwest Anatolia, Turkey. Vegetation
History and Archaeobotany, 13, 45–54.
Martinoli, D. and Nesbitt, M. 2003. Plant stores at Pottery
Neolithic Höyücek, southwest Turkey. Anatolian Studies,
53, 17–32.
Mathew, B. 1977. Crocus sativus and its allies Iridaceae.
Plant systematics and evolution, 128, 89–103.
Mathew, B. 1996. A review of Allium section Allium,
Richmond, Royal Botanic gardens, Kew.
Mathew, B. 1999. Botany, taxonomy and cytology of C.
sativus L. and its allies. In: Negbi, M. (ed.) Saffron: Crocus
sativus L. Pp. 19–30. Netherlands, Harwood Academic
Publishers.
Matsuoka, Y., Jaffar Aghaei, M., Abbasi, M., Totiaei, A.,
Mozafari, J. and Ohta, S. 2008. Durum wheat cultivation
associated with Aegilops tauschii in northern Iran. Genetic
Resources and Crop Evolution, 55, 861–8.
Maxted, N. 1995. An ecogeographical study of Vicia subgenus
Vicia. Systematic and ecogeopgraphic studies on crop
genepools. 8. Rome, International Plant Genetic
Resources Institute.
Mayer, M. and Soltis, P. S. 1994. Chloroplast DNA phylog-
eny of Lens (Leguminosae): origin and diversity of
cultivated lentil. Theoretical and Applied Genetics, 87,
773–81.
McCollum, G. D. 1976. Onion and allies. In: Simmonds,
N. W. (ed.) Evolution of crop plants. Pp. 186–90. London,
Longman.
McCorrison, J. 1997. The fiber revolution, textile extensifi-
cation, alienatication, and social stratification in ancient
Mesopotamia. Current Anthropology, 38, 517–49.
McCreery, D. W. 1979. Flotation of the Bab edh-Dhra and
Numeira plant remains. Annual of the American Schools of
Oriental Research, 46, 165–9.
McFadden, E. S. and Sears, E. R. 1946. The origin of
Triticum spelta and its free-threshing hexaploid relatives.
Journal of Heredity, 37, 81–90; 107–16.
222 REFERENCES
McGovern, P. E. 1998. Wine for eternity. Archaeology, 51,
28–32.
McGovern, P. E. 2003. Ancient wine: the search for the origins
of viniculture. Princeton, New Jersey: Princeton
University Press.
McGovern, P. E. and Michel, R. H. 1995. The analytical and
archaeological challenge of detecting ancient wine: two
case studies from the ancient Near East. In: McGovern,
P. E., Fleming, S. T. and Katz, S. M. (eds.) The origins and
ancient history of wine. Pp. 57–65. Luxembourg, Gordon
and Breach.
McGovern, P. E., Glusker, D. L., Exner, L. J. and Voigt, M.
M. 1996. Neolithic resinated wine. Nature, 381, 480–1.
Meadow, R. H. 1996. The origins and spread of agriculture
and pastoralism in north western South Asia. In: Harris,
D. R. (ed.) The origins and spread of agriculture and pasto-
ralism in Eurasia. Pp. 390–412. London, University
College London Press.
Mehlenbacher, S. A., Cociu, V. and Hough, L. F. 1990.
Apricots (Prunus). In: Moorem, J.N. and Ballington, Jr
J.R. (eds) Genetic resources of temperate fruit and nut
crops. Acta Horticulturae, 290, 65–107.
Melamed, Y., Kislev, M., Weiss, E. and Simchoni, O. 2011.
Extinction of water plants in the Hula Valley: Evidence for
climate change. Journal of Human Evolution, 60(4), 320–7.
Melamed, Y., Plitmann, U., Shmida, A. and Golan, O. 2009.
Lathyrus clymenum L. in Israel: A ‘revival’ of an ancient
species. Israel Journal of Plant Sciences, 57, 125–30.
Merlin, M. D. 2003. Archaeological evidence for the tradi-
tion of psychoactive plant use in the old world. Economic
Botany, 57, 295–323.
Méry, S. and Tengberg, M. 2009. Food for eternity? The
analysis of a date offering from a 3rd millennium BC
grave at Hili N, Abu Dhabi (United Arab Emirates).
Journal of Archaeological Science, 36, 2012–17.
Meusel, H., Jäger, E. and Weinert, E. 1965. Vergleichende
chorologie der zentraleuropäschen flora. Jena, Verlag Gustav
Fischer.
Meyer, F. G. 1980. Carbonised food plants of Pompeii,
Herculaneum and the villa at Torre Annunziata.
Economic Botany, 34, 401–37.
Miller, N. F. 1981. Plant remains from Ville Royale II, Susa.
Cahiers de la Délégation Archéologique Française en Iran, 12,
137–42.
Miller, N. F. 1986. Vegetation and landuse, archaeological
analysis. In: Algaze, G. et al. (eds.) The Chicago
Euphrates archaeological project 1980–1984: an interim
report. Anatolica, 13, 84–123.
Miller, N. F. 1991. The Near East. In: van Zeist, W.,
Wasylikowa, K. and Behre, K. -E. (eds.) Progress in Old
World palaeoethnobotany. Pp. 133–60. Rotterdam,
Balkema.
Miller, N. F. 1999. Agricultural development in western
Central Asia in the Chalcolithic and Bronze Ages.
Vegetation History and Archaeobotany, 8, 13–19.
Miller, N. F. 2002. Tracing the development of the agropas-
toral economy in Southeastern Anatolia. In: Cappers, R.
and Bottema, S. (eds.) The dawn of farming in the Near
East, production, subsistence and environment. Pp. 85–94.
Berlin, ex oriente.
Miller, N. F. 2003. The use of plants at Anau North. In:
Hiebert, F. T. (ed.) A Central Asian village at the dawn of
civilization, excavations at Anau, Turkmenistan. Pp. 127–38.
Philadelphia, University of Pennsylvania Museum of
Archaeology and Anthropology.
Miller, N. F. 2008. Sweeter than wine? The use of the grape
in Early Western Asia. Antiquity 82, 937–46.
Miller, S. A. and Mitchell, G. V. 1982. Optimisation of
human protein requirements. In: Fox, P. F. and Condon,
J. J. (eds.) Food Proteins. Pp. 105–20. London, Applied
Science Publishers.
Miller, T. E. 1987. Systematics and evolution. In: Lupton, F.
G. H. (ed.) Wheat breeding. Pp. 1–30. London, Chapman
and Hall.
Møbjerg, T., Jensen, P. M. and Mikkelsen, P. H. 2007.
Enkehøj, En boplads med klokkebgerkeramik og korn.
KUML: Årbog for Jysk Arkaeologisk Selskab, 2007, 9–41.
Moens, M. F. and Wetterstrom, W. 1988. The agricultural
economy of an Old Kingdom town in Egypt’s west
delta: insights from the plant remains. Journal of Near
Eastern Studies, 47, 159–73.
Molina-Cano, J. L., Russell, J. R., Moralejo, M. A.,
Escacena, J. L., Arias, G. and Powell, W. 2005.
Chloroplast DNA microsatellite analysis supports a
polyphyletic origin for barley. Theoretical and Applied
Genetics, 110, 613–19.
Monah, F. 2002. Analiza arheobotanică a unui complex
menajer gumelniţean din tell-ul de la Hârşova. Arheologia
Moldovei, 22, 274–81.
Monah, F. 2007a. Nouvelles determinations archéobota-
niques pour la Roumanie. Arheologia Moldovei. 30,
343–52.
Monah, F. 2007b. The spread of cultivated plants in the
regin between the Carpathians and Dniester, 6th/4th
millennia BC. In: Colledge, S. and Conolly, J. (eds.) The
origins and spread of domestic plants in Southwest Asia and
Europe. Pp. Walnut Creek, CA: Left Coast Press.
Monah, F. and Monah, D. 2008. Cercetări arheobotanice în
tell-ul calcolitic Poduri- Dealul Ghindaru. Piatra Neamţ,
Constantin Mătasă.
Monk, M. 1988. Archaeobotanical study of samples from
pipeline sites. In: Gowan, M. (ed.) Three Irish gas pipe-
lines: new archaeological evidence in Munster. Pp. 185–91.
Dublin, Wordwell.

Morrell, P. L. and Clegg, M. T. 2007. Genetic evidence for a
second domestication of barley (Hordeum vulgare) east
of the Fertile Crescent. Proceedings of the National Academy
of Sciences, 104, 3289–94.
Morris, R. and Sears, E. R. 1967. The cytogenetics of wheat
and its relatives. In: Quisenberry, K. S. and Reitz, L. P.
(eds.) Wheat and wheat improvements. Pp. 19–87. Madison,
American Society for Agronomy.
Moulherat, C., Tengberg, M., Haquet, J.-F. and Mille, B.
2002. First evidence of cotton at Neolithic Mehrgarh,
Pakistan: analysis of mineralized fibres from a copper
bead. Journal of Archaeological Science, 29, 1393–401.
Mueller, U. G. and Wolfenbarger, L. L. 1999. AFLP geno-
typing and fingerprinting. Trends in Ecology and
Evolution, 14, 389–94.
Mullins, M. G., Bouquest, A. and Williams, L. E. 1992.
Biology of grapevine. Cambridge, UK: Cambridge
University Press.
Munier, P. 1973. Le Palmier-Dattier. Paris, Maisonneuve et
Larose.
Muratova, V. 1931. Common beans, Vicia faba L.—a
botanical-agronomical monograph. Bulletin of Applied
Botany of Genetics and Plant-Breeding, 50 (Supplement),
1–295.
Murphy, P. 1989. Carbonised Neolithic plant remains from
The Stumble, an intertidal site in Blackwater estuary,
Essex, England. Circea, 6, 21–38.
Murray, M.-A. 2003. Archaeobotanical remains from the
Aceramic Neolithic wells (ca. 9000 BP). In: Peltenburg,
E. (ed.) The colonisation and settlement of Cyprus: investiga-
tions at Kissonerga-Mylouthkia 1977–1996 (Lemba archaeo-
logical project, Cyprus III.1: studies in Mediterranean
archaeology 70:4). Pp. 59–72. Sävedalen, Åström Verlag.
Murray, M.-A. 2000a. Cereal production and processing.
In: Nicholson, P. T. and Shaw, I. (eds.) Ancient Egyptian
Materials and Technology. Pp. 505–36. Cambridge, UK:
Cambridge University Press.
Murray, M.-A. 2000b. Fruits, vegetables, pulses and condi-
ments. In: Nicholson, P. T. and Shaw, I. (eds.) Ancient
Egyptian Materials and Technology. Pp. 609–55. Cambridge,
UK: Cambridge University Press.
Murray, M.-A. 2000c. Viticulture and wine production. In:
Nicholson, P. T. and Shaw, I. (eds.) Ancient Egyptian
Materials and Technology. Pp. 577–608. Cambridge, UK:
Cambridge University Press.
Nalam, V., Vales, M., Watson, C., Kianian, S. and Riera-
Lizarazu, O. 2006. Map-based analysis of genes affect-
ing the brittle rachis character in tetraploid wheat
(Triticum turgidum L.). Theoretical and applied genetics,
112, 373–81.
Nasiri, J., Haghnazari, A. and Saba, J. 2009. Genetic diver-
sity among varieties and wild species accessions of pea
REFERENCES 223
(Pisum sativum L.) based on SSR markers. African Journal
of Biotechnology, 8, 3405–17.
Navot, N. and Zamir, D. 1987. Izozyme and seed protein
phylogeny of the genus Citrullus (Cucurbitaceae). Plant
Systematics and Evolution, 156, 61–7.
Navot, N., Sarfatti, M. and Zamir, D. 1990. Linkage rela-
tionships of genes affecting bitterness and flesh color in
watermelon. Journal of Heredity, 81, 162–5.
Nayar, N. M. 1995. Sesame, Sesamum indicum L. (Pedaliaceae).
In: Smartt, J. and Simmonds, N. W. (eds.) Evolution of crop
plants. 2 ed, Pp. 404–7. New York, Wiley.
Neef, R. 1989. Plants. In: van der Kooij, G. and Ibrahim, M.
M. (eds.) Picking up the threads: A continuing review of
excavations at Deir Alla, Jordan. Pp. 30–7. Leiden,
University of Leiden Archaeological Centre.
Neef, R. 1990. Introduction, development and environ-
mental implications of olive cultivation: The evidence
from Jordan. In: Bottema, S., Entjes-Nieborg, G. and
van Zeist, W. (eds.) Man’s role in the shaping of the east-
ern Mediterranean landscape. Pp. 295–306. Rotterdam,
Balkema.
Neef, R. 1998. Status and perspectives of archaeobotanical
research in Jordan. In: Gebel, H. J. K., Kafkafi, Z., Rollefson,
A. and Rollefson, G. O. (eds.) The prehistory of Jordan II:
Perspectives from 1997. Pp. 601–9. Berlin, ex oriente.
Negbi, M. 1999. Saffron cultivation: past, present and
future prospects. In: Negbi, M. (ed.) Saffron: Crocus
sativus L. Pp. 1–18. Australia, Harwood Academic
Publishers.
Nelson, D. E., Korteling, R. G. and Stott, W. R. 1977.
Carbon-14: direct detection at natural concentrations.
Science, 198, 507–8.
Nesbitt, M. 1993. Archaeobotanical remains. In: G.
Herrmann, V.M. Masson and K. Kurbansakhatov et al.
‘The International Merv Project. Preliminary Report on
the First Season (1992). Iran, 31, 56–8. Oxford, British
Institute of Persian Studies.
Nesbitt, M. 1994. Archaeobotanical research in the Merv
oasis. In: Hermann, G. and Kurbansakhatov, K. The
international Merv project: preliminary report on the
second season (1993). Iran, 32, 71–3. Oxford, British
Institute of Persian Studies.
Nesbitt, M. 1996. Chalcolithic crops from Kuruçay Höyük:
an interim report. In: Duru, P. (ed.) Results of the excava-
tions 1978–1988, the late Chalcolithic and early Bronze set-
tlements. Pp. 90; 134–7. Ankara, Türk Tarіh Kurumu
Basimevі.
Nesbitt, M. 2001. Wheat evolution: integrating archaeo-
logical and biological evidence. In: Caligari, P. D. S. and
Brandham, P. E. (eds.) Wheat taxonomy: the legacy of John
Percival, Vol. 3, Linnean Special Issue. Pp. 37–59. London,
Linnean Society.
224 REFERENCES
Nesbitt, M. 2002. When and where did domesticated cere-
als first occur in southwest Asia? In: Cappers, R. T. J.
and Bottema, S. (eds.) The dawn of farming in the Near
East. Pp. 113–32. Berlin, ex oriente.
Nesbitt, M. and Samuel, D. 1996a. Archaeobotany in Turkey:
a review of current research. Orient Express, 3, 91–6.
Nesbitt, M. and Samuel, D. 1996b. From staple crop to
extinction? The archaeology and history of hulled
wheats. In: Padulosi, S., Hammer, K. and Heller, J. (eds.)
Hulled wheats. Pp. 41–100. Rome, International Plant
Genetic Resources Institute.
Nesbitt, M. and Summers, G. D. 1988. Some recent discov-
eries of millet (Panicum miliaceum L. and Setaria italica
(L.) P. Beauv.) at excavations in Turkey and Iran.
Anatolian Studies, 38, 85–97.
Nesbitt, M., Simpson, S. J. and Svanberg, I. 2010. History
of rice in Western and Central Asia. In: Sharma, S. D.
(ed.) Rice: origin, antiquity and history. Pp. 312–44, 535–41.
Enfield, NH: Science Publishers.
Netolitzky, F. 1914. Die Hirse aus alten Funden. Sitzungsber
Kaiserl Akad Wissensch Math Naturwissensch Classe, 73,
725–59.
Neumann, F., Schölzel, C., Litt, T., Hense, A. and Stein, M.
2007. Holocene vegetation and climate history of the
northern Golan heights (Near East). Vegetation History
and Archaeobotany, 16, 329–46.
Neuweiler, E. 1935. Nachträge urgeschichtlicher pflanzen.
Vierteljahrsschrift der Naturforschenden Gesellschaft in
Zürich, 80, 98–122.
Neuweiler, E. 1946. Nachträge II urgeschichtlicher pflan-
zen. Vierteljahrsschrift der Naturforschenden Gesellschaft in
Zürich, 91, 122–36.
Nevo, E. 1992. Origin, evolution, population genetics and
resources for breeding of wild barley, Hordeum sponta-
neum, in the Fertile Crescent. In: Shewry, P. R. (ed.) Barley,
genetics, biochemistry, molecular biology and biotechnology.
Pp. 19–43. Oxon, UK: C.A.B. International, Wallingford.
Newton, C., Terral, J.-F. and Ivorra, S. 2006. The Egyptian
olive (Olea europaea subsp. europaea) in the later first mil-
lennium BC: origins and history using the morphomet-
ric analysis of olive stones. Antiquity, 80, 405–14.
Nguyen, T. T., Taylor, P. J. W., Redden, R. J. and Ford, R.
2004. Genetic diversity estimates in Cicer using AFLP
analysis. Plant Breeding, 123, 173–9.
Nixon, R. W. 1951. The date palm—‘tree of life’ in the sub-
tropical deserts. Economic Botany, 5, 274–301.
Noy, T., Legge, A. J. and Higgs, E. S. 1973. Recent excava-
tions at Nahal Oren, Israel. Proceedings of the Prehistory
Society, 39, 75–99.
Nürnberg-Krüger, U. 1960a. Cytogenetische untersuchun-
gen an der gattung Secale L. Zeitschrift für Pflanzen-
züchtung, 44, 63–72.
Nürnberg-Krüger, U. 1960b. Cytogenetische untersuchun-
gen an Secale silvestre Host. 1. Der Bastard mit Secale cere-
ale L. Züchter, 30, 147–50.
Olmo, H. P. 1995a. Grapes. In: Smartt, J. and Simmonds, N.
W. (eds.) Evolution of crop plants. 2 ed, Pp. 485–90.
London, Longman.
Olmo, H. P. 1995b. The origin and domestication of the
vinifera grape. In: McGovern, P. T., Fleming, S. J. and
Katz, S. H. (eds.) The origins and ancient history of wine.
Pp. 31–43. Luxembourg: Gordon and Breach.
Opravil, E. 1979. Rostlinnĕ zbytky z Mohelnice 1. a 2. Cǎsopis
Slezského muzea v Opave,̌ Série A, 28, 1–13, 97–109.
Opravil, E. 1981. Z histoire Inu b našich zemich a ve štrdní
Europě (Aus der geschichte des leines in unseren
ländern und in Mitteleuropa). Archeologicke rozhledy, 33,
299–305.
Osborn, D. J. 1968. Notes on medicinal and other uses of
plants in Egypt. Economic Botany, 22, 165–77.
Oybak-Dönmez, E. 2005. Early Bronze Age crop plants
from Yenibademli Höyük (Gökçeada), Western Turkey.
Environtal Archaeology, 10, 39–49.
Özkan, H., Brandolini, A., Schäfer-Pregl, R. and Salamini,
F. 2002. AFLP analysis of a collection of tetraploid
wheats indicates the origin of emmer and hard wheat
domestication in Southeast Turkey. Molecular biology and
evolution, 19, 1797–801.
Palmer, J. D., Jorgensen, R. A. and Thompson, W. F. 1985.
Chloroplast DNA variation and evolution of Pisum:
Patterns of change and phylogenetic analysis. Genetics,
109, 195–213.
Pals, J. P. 1984. Plant remains from Aartswoud, a Neolithic
settlement in the coastal area. In: van Zeist, W. and
Casparie, W. A. (eds.) Plants and ancient man. Pp. 267–76.
Rotterdam, Balkema.
Pals, J. P. and Voorrips, A. 1979. Seeds, fruits and charcoals
from two sites in northern Italy. Archaeo-Physika, 8, 217–35.
Pashkevich, G. 1999. New evidence for plant exploitation
by the Scythian tribes during the Early Iron Age in the
Ukraine. Acta Palaeobotanica, 2(Suppl), 597–601.
Pashkevich, G. 2003. Paleoethnobotanical evidence of
agriculture in the steppe and the forest-steppe of East
Europe in the Late Neolithic and Bronze Age. In: Levine,
M., Renfrew, C. and Boyle, K. (eds.) Prehistoric steppe
adaptation and the horse. Pp. 287–97. Cambridge,
McDonald Institute Monographs.
Pashkevich, G. 2005. Paleoethnobotanical evidence of the
Tripolye Culture. In: Dumitroaia, G., Chapman, J.,
Weller, O., Preoteasa, C., Munteanu, R., Nicola, D. and
Monah, D. (eds.) Cucuteni. 120 years of research. Pp. 231–
45. Piatra Neamt, Time to sum up.
Pasternak, R. 1998. Investigations of botanical remains
from Nevali Çori PPNB, Turkey: a short interim report.

In: Damania, A. B., Valkoun, J., Willcox, G. and Qualset,
C. O. (eds.) The origins of agriculture and crop domestica-
tion. Pp. 170–6. Aleppo, Syria: ICARDA.
Pazouki, L., Mardi, M., Shanjani, P., Hagidimitriou, M.,
Pirseyedi, S., Naghavi, M., Avanzato, D., Vendramin, E.,
Kafkas, S., Ghareyazie, B., Ghaffari, M. and Khayam
Nekoui, S. 2010. Genetic diversity and relationships
among Pistacia species and cultivars. Conservation
Genetics, 11, 311–18.
Pelling, R. 2008. Garamantian agriculture: the plant
remains from Jarma, Fazzan. Libyan Studies, 39, 41–71.
Peña-Chocarro, L. 1999. Prehistoric agriculture in Southern
Spain during the Neolithic and the Bronze Age, the applica-
tion of ethnographic models. Oxford, Archaeopress.
Peña-Chocarro, L. 2007. Early agriculture in central and
southern Spain. In: Colledge, S. and Conolly, J. (eds.)
The origins and spread of domestic plants in Southwest Asia
and Europe. Pp. 173–87. Walnut Creek, CA: Left Coast
Press.
Peña-Chocarro, L., Zapata, L., Iriarte, M. J., Gonzalez
Morales, M. and Straus, L. G. 2005. The oldest agricul-
ture in northern Atlantic Spain: new evidence from El
Miron Cave (Ramales de la Victoria, Cantabria). Journal
of Archaeological Science, 32, 579–87.
Percival, J. 1921. The wheat plant. London, Duckworth.
Pérez de la Vega, M. and Allard, R. W. 1984. Mating system
and genetic polymorphism in populations of Secale cere-
ale and S. vavilovii. Canadian Journal of Genetics and
Cytoloty, 26, 308–17.
Perrino, P., Laghetti, G., Antuono, L. F., Al Ajlouni, M. F.,
Kanbertay, M., Szabó, A. T. and Hammer, K. 1996.
Ecogeographical distribution of hulled wheat species.
In: Padulosi, S., Hammer, K. and Heller, J. (eds.) Hulled
wheats. Pp. 101–19. Rome, International Plant Genetic
Resource Institute.
Pessina, A. and Rottoli, M. 1996. New evidence on the ear-
liest farming cultures in northern Italy: Archaeological
and palaeobotanical data. Poročilo Oraziskovanju
Paleolitika, Neolitika i Eneolitica v Sloveniji, 23, 77–94.
Phillips, L. L. 1961. The cytogenetics of speciation in
Asiatic cotton. Genetics, 46, 77–83.
Piening, U. 1989. Pflanzenreste aus der Bandkeramischen
siedlung von Bietigheim-Bissingen, kreis Ludwigsburg.
Fundberichte aus Baden-Württemberg, 14, 119–40.
Pinto da Silva, A. R. 1976. Carbonized grains and plant
imprints in ceramics from the Castrum of Baiões (Beira
Alta, Portugal). Folia Quaternaria, 47, 3–18.
Pinto da Silva, A. R. 1988. A paleoetnobotanica na arqueo-
logica Portuguesa. Resultados desde 1931 a 1987. In:
Queiroga, F. M. V. R., Sousa, I. M. and Oliveira, C. M.
(eds.) Paleoecologia e Arqueologia. Pp. 5–36. Camara
Municipal de Vila Nova de Famalicao.
REFERENCES 225
Polignano, G. B., Bisignano, V., Tomaselli, V., Uggenti, P.,
Alba, V. and Della Gatta, C. 2009. Genotype × environ-
ment interaction in grass pea (Lathyrus sativus L.) lines,
Article ID 898396. International Journal of Agronomy
[Online], 2009. [Accessed 25/10/2010].
Popova, T. 1992. Analyse carpologique. In: Demoule, J.-P.
and Lichardusitten, M. (eds.) Kovačevo. Foullies
Néolithiques franco-bulgares dans la vallée du Strymon.
Rapport annuel. Pp. 21–3. Paris, Université de Paris I.
Popova, T. 1995. Palaeoethnobotanical remains from
Early Bronze Age settlement of Galabovo (South
Bulgaria). In: Kroll, H. and Pasternak, R. (eds.) Res
Archaeobotanicae, International Workgroup for
Palaeoethnobotany, Proceedings of the 9th Symposium.
Pp. 261–5. Kiel, Oetker-Voges.
Postgate, J. N. 1987. Notes on fruits in the cuneiform
sources. Bulletin on Sumerian Agriculture, 3, 115–44.
Potokina, E., Tomooka, N., Vaughan, D. A., Alexandrova,
T. and Xu, R.-Q. 1999. Phylogeny of Vicia subgenus Vicia
(Fabaceae) based on analysis of RAPDs and RFLP of
PCR-amplified chloroplast genes. Genetic Resources and
Crop Evolution, 46, 149–61.
Poyarkova, U. 1988. Morphology, geography and infra-
specific taxonomics of Triticum dicoccoides Körn., A retro-
spective of 80 years of research. Euphytica, 38, 11–23.
Poyarkova, U. and Gerechter-Amitai, Z. K. 1991. Two vari-
ants of wild emmer (Triticum dicoccoides) native to Israel:
morphology and distribution. Canadian Journal of Botany,
69, 2772–89.
Prasada Rao, K. E., de Wet, J. M. S., Brink, D. E. and
Megesha, M. H. 1987. Intraspecific variation and sys-
tematics of cultivated Setaria italica, foxtail millet
(Poaceae). Economic Botany, 41, 108–16.
Price, H. J., Dillon, S. L., Hodnett, G., Rooney, W. L., Ross,
L. and Johnston, J. S. 2005. Genome evolution in the
genus Sorghum (Poaceae). Annals of Botany, 95, 219–27.
Pruessner, A. H. 1920. Date culture in ancient Babylonia.
American Journal of Semitic Languages and Literature, 36,
213–32.
Quazzini, N., Lumaret, R., Villemur, P. and Di Giusto, F.
1993. Leaf allozyme variation in cultivated and wild
olive trees (Olea europaea). Journal of Heredity, 81, 34–42.
Raamsdonk, L. V. D., Wiestma, W.A. and de Vries, J. N.
1992. Crossing experiments in Allium L. section Cepa.
Botanical Journal of the Linnean Society, 109(2), 293–303.
Rafalski, J. A. 2002. Novel genetic mapping tools in plants:
SNPs and LD-based approaches. Plant Science, 162,
329–33.
Ramil Rego, P. and Aira Rodríguez, M. J. 1993. A palaeo-
carpological study of Neolithic and Bronze Age levels of
the Buraco da Pala rock-shelter (Bragança, Portugal).
Vegetation History and Archaebotany, 2, 163–72.
226 REFERENCES
Ramming, D. W. and Coicu, V. 1995. Plums (Prunus). In:
Moore, J. N. and Ballington, J. R. (eds.) Genetic resources
of temperate fruit and nut crops. Pp. 235–87. Wageningen,
International Society for Horticultural Science.
Ramón-Laca, L. 2003. The introduction of cultivated citrus
to Europe via Northern Africa and the Iberian Peninsula.
Economic Botany 57(4), 502–14.
Rao, P. S. and Smith, E. L. 1968. Studies with Israeli and
Turkish accessions of Triticum turgidum L. emend. var.
dicoccoides (Körn.) Bowden. Wheat Information Service,
26, 6–7.
Rashed-Mohassel, M. H. 2006. Saffron Botany. In: Kafi, M.,
Koocheki, A., Rashed, M. H. and Nassiri, M. (eds.)
Saffron (Crocus sativus L.): Production and Processing.
Pp. 13–38. USA, Science Publishers.
Rehder, A. 1940. Manual of cultivated trees and shrubs hardy
in North America. New York, Macmillan.
Reimer, P. J., Baillie, M. G. L., Bard, E., Bayliss, A., Beck,
J. W., Bertrand, C., Blackwell, P. G., Buck, C. E., Burr,
G., Cutler, K. B., Damon, P. E., Edwards, R. L.,
Fairbanks, R. G., Friedrich, M., Guilderson, T. P.,
Hughen, K. A., Kromer, B., McCormac, F. G., Manning,
S., Bronk Ramsey, C., Reimer, R. W., Remmele, S.,
Southon, J. R., Stuiver, M., Talamo, S., Taylor, F. W.,
van der Plicht, J. and Weyhenmeyer, C. E. 2004.
IntCal04: Calibration Issue. Radiocarbon, 46(3),
1029–58.
Reimer, P. J., Baillie, M. G. L., Bard, E., Bayliss, A., Beck, J.
W., Blackwell, P. G., Bronk Ramsey, C., Buck, C. E., Burr,
G., Edwards, R. L., Friedrich, M., Guilderson, T. P.,
Hajdas, I., Heaton, T. J., Hogg, A. G., Hughen, K. A.,
Kaiser, K. F., Kromer, B., McCormac, F. G., Manning, S.,
Reimer, R. W., Richards, D. A., Southon, J. R., Talamo, S.,
Turney, C. S. M., van der Plicht, J. and Weyhenmeyer, C.
E. 2009. IntCal09 and Marine09 radiocarbon age calibra-
tion curves, 0–50,000 years cal BP. Radiocarbon, 51,
1111–50.
Renfrew, J. M. 1968. A note on the Neolithic grain from
Can Hasan. Anatolian Studies, 18, 55–6.
Renfrew, J. M. 1972. The plant remains. In: Warren, P. (ed.)
Myrtos: an early Bronze Age settlement in Crete. Pp. 315–17.
London, British School at Athens.
Renfrew, J. M. 1973. Palaeoethnobotany: the prehistoric food
plants of the Near East and Europe. New York, Columbia
University Press.
Renfrew, J. M. 1974. Report on carbonized grains and
seeds from Obre I, Kakanj and Obre II. Wissenschaftliche
mitteilungen aus Bosnisch-Herzegowinischen landesmuse-
ums, 4A, 47–53.
Renfrew, J. M. 1976. Carbonized seed from Anza. In: Gimbutas,
M. (ed.) Neolithic Macedonia as reflected in the excavation of
Anza, south-east Yugoslavia. Pp. 300–12. Los Angeles,
Monumenta Archaeologica. University of California.
Renfrew, J. M. 1979. The first farmers in South East Europe.
Archaeo-Physika, 8, 243–65.
Reuveni, O. 1986. Fruit set and fruit development of the
date palm (Phoenix dactylifera L.). In: Halevi, A. (ed.)
Handbook of flowering. Pp. 119–44. Boca Raton, CRC
Press.
Rhouma, S., Dakhlaoui-Dkhil, S., Ould Mohamed Salem,
A., Zehdi-Azouzi, S., Rhouma, A., Marrakchi, M. and
Trifi, M. 2008. Genetic diversity and phylogenic relation-
ships in date-palms (Phoenix dactylifera L.) as assessed by
random amplified microsatellite polymorphism mark-
ers (RAMPOs). Scientia Horticulturae, 117, 53–7.
Richards, C., Volk, G., Reilley, A., Henk, A., Lockwood, D.,
Reeves, P. and Forsline, P. 2009. Genetic diversity and
population structure in Malus sieversii, a wild progeni-
tor species of domesticated apple. Tree Genetics and
Genomes, 5, 339–47.
Richardson, I. B. K. 1986. Chestnuts (genus Castanea). In:
Hora, B. (ed.) The Oxford encyclopaedia of trees in the world.
2 ed, Pp. 133–5. Oxford, Oxford University Press.
Riehl, S. 1997. Bronze Age environment and economy in the
Troad: the archaeobotany of Kumtepe and Troy. Ph.D.,
University of Tübingen.
Riehl, S. 2000. Erste ergebnisse der archäobotanischen unter-
suchungen in der zentralen oberstadt von Tall Mozan/
Urkeš im rahmen der DOG-IIMAS-Kooperation.
Mitteilungen der Deutschen Orient-Gesellschaft zu Berlin
132, 229–38.
Riggs, T. J. 1995. Umbelliferous minor crops. In: Smartt, J.
and Simmonds, N. W. (eds.) Evolution of crop plants. 2 ed,
Pp. 481–5. London, Longman.
Riley, R. 1965. Cytogenetics and evolution of wheat. In:
Hutchinson, J. B. (ed.) Crop plant evolution. Pp. 103–22.
Cambridge, UK; Cambridge University Press.
Rindos, D. 1984. The origins of agriculture: an evolutionary
perspective. New York, Academic Press.
Riviera Núñez, D. and Walker, J. 1989. A review of palaeo-
botanical findings of early Vitis in the Mediterranean
and of the origin of cultivated grape-vines, with special
reference to new pointers to prehistoric exploitation in
the Western Mediterranean. Review of Palaeobotany and
Palynology, 61, 205–17.
Robinson, D. E. 2000. Det slesvigske agerbrug i yngre ste-
nalder og bronzealder. Arkæobotanikkens udsagn
(Agriculture in Schleswig during the Neolithic and
Bronze Age—the archaeobotanical evidence). In:
Ethelberg, P., Jørgensen, E., Meier, D. and Robinson, D.
E. (eds.) Det Sønderjyske landbrugs historie. Pp. 281–98.
Haderslev, Sten- og Bronzealder.

Robinson, D. E. 2003. Neolithic and Bronze Age agricul-
ture in Southern Scandinavia – recent archaeobotanical
evidence from Denmark. Environmental Archaeology, 8,
145–65.
Rollefson, G. O., Simmons, A. H., Donaldson, M. L.,
Gillespie, W., Kafafi, Z., Köhler-Rollefson, I. U.,
McAdam, E., Ralston, S. L. and Tubb, M. K. 1985.
Excavation at the Pre-Pottery Neolithic B village of ‘Ain
Ghazal (Jordan), 1983. Mitteilungen der Deutschen Orient-
Gesellschaft zu Berlin, 117, 69–116.
Roose, M. L., Soost, R. K. and Cameron, J. W. 1995. Citrus.
In: Smartt, J. and Simmonds, N. W. (eds.) Evolution of
crop plants. 2 ed, Pp. 443–8. London, Longman, UK.
Rösch, M. 1998. The history of crops and crop weeds in
south-western Germany from the Neolithic period to
modern times, as shown by archaeobotanical evidence.
Vegetation History and Archaeobotany, 7, 109–25.
Rosenberg, M., Nesbitt, M., Redding, R. W. and Strasser, T.
F. 1995. Hallan Çemi Tepesi: some preliminary observa-
tions concerning Early Neolithic subsistance behaviors
in Eastern Anatolia. Anatolica, 21, 3–12.
Roshevitz, R. I. 1947. Monograph of the genus Secale L. Acta
instituti botanici academiae scientiarum USSR, 1, 105–63.
Rottoli, M. 1993. ‘La Marmotta’, Anguillara Sabazia (RM).
Scavi 1989. Analisi paletnobotaniche: prime risultanze.
Appendice 1. In: Fugazzola, M. A., D’eugenio, G. and
Pesina, A. (eds.) ‘La Marmotta’ (Anguillara Sabazia, RM).
Scavi 1989. Un abitato perilacustre di età Neolitica. Pp. 305–15.
Roma, Bullettino di Paletnologia Italiana.
Rottoli, M. 2002. Zafferanone selvatico (Carthamus lanatus) e
cardo della Madonna (Silybum marianum), piante raccolte
o coltivate nel Neolitico antico a ‘La Marmotta’? Bullettino
di Paletnologia Italiana (2000–2001), 91–92, 47–61.
Rottoli, M. 2005. Un nuovo frumento vestito nei siti neo-
litici del Friuli-Venezia Giulia (Italia nord orientale).
Gortania - Atti del Museo Friulano di Storia Naturale, 26,
67–78.
Rottoli, M. and Castiglioni, E. 2009. Prehistory of plant
growing and collecting in Northern Italy, based on seed
remains from the Early Neolithic to the Chalcolithic (c.
5600–2100 cal B.C.). Vegetation History and Archaeobotany,
18, 91–103.
Rottoli, M. and Pessina, A. 2007. Neolithic agriculture in
Italy: an update of archaeobotanical data with particu-
lar emphasis on northern settlements. In: Colledge, S.
and Conolly, J. (eds.) The origins and spread of domestic
plants in Southwest Asia and Europe. Pp. 141–54. Walnut
Creeck, CA: Left Coast Press.
Rowley-Conwy, P. 1979. Forkullet korn fra Lindebjerg. En
boplads fra aeldre bronzealder. KUML: Årbog for Jysk
Arkaeologisk Selskab, 1978, 159–71.
REFERENCES 227
Rowley-Conwy, P. 1983. Bronzealderkorn fra Voldtofte.
KUML: Årbog for Jysk Arkaeologisk Selskab, 1982–3, 139–52.
Rowley-Conwy, P. 1991. Sorghum from Qasr Ibrim,
Egyptian Nubia, c. 888 bc - ad 1811; a preliminary study.
In: Renfrew, J. (ed.) New light on early farming. Pp. 191–212.
Edinburgh, Edinburgh Univ Press.
Rowley-Conwy, P. 1994. Dung, dirt and deposits: site for-
mation under conditions of near-perfect preservation at
Qasr Ibrim, Egyptian Nubia. In: Luff, R. and Rowley-
Conwy, P. (eds.) Whither Environmental Archaeology.
Pp. 25–32. Oxford, UK: Oxbow.
Rowly-Conwy, P. A., Deakin, W. J. and Shaw, C. H. 1997.
Ancient DNA from archaeological sorghum (Sorghum
bicolor) from Qasr Ibrim, Nubia. Implications for domes-
tication and evolution and a review of the archaeologi-
cal evidence. Sahara, 9, 23–34.
Rowly-Conwy, P. A., Deakin, W. J. and Shaw, C. H. 1999.
Ancient DNA from sorghum: the evidence from Qasr
Ibrim, Egyptian Nubia. In: van der Veen, M. (ed.) The
exploitation of plant resources in ancient Africa. Pp. 23–34.
New York, NY: Kluwer Academic.
Rubtzov, G. A. 1944. Geographic distribution of the genus
Pyrus and trends and factors in its evolution. American
Naturalist, 78, 358–66.
Rudenko, S. I. 1970. Frozen tombs of Siberia, London, Dent.
Runesson, H. 2007. Den goda ordningen. In: Gustafsson, P.
and Spång, L. G. (eds.) Stenålderns stationer. Arkeologi i
Botniabanans spår. Pp. 71–116. Murberget, Länsmuseet
Västernorrland: Riksantikvarieämbetet.
Saisho, D. and Puruggana, M. D. 2007. Molecular phyloge-
ography of domesticated barley traces expansion of
agriculture in the Old World. Genetics, 177, 1765–76.
Samuel, D. 1996. Investigation of ancient Egyptian baking
and brewing methods by correlative microscopy. Science,
273, 488–90.
Samuel, D. 2001. Archaeobotanical evidence and analysis,
In: Berthier, S. (ed.) Peuplement rural et aménagements
hydroagricoles dans la moyenne vallée de l’Euphrate fin VIIe-
XIXe siècle. Pp. 343–481. Damascus, Institut Français
d’études arabes de Damas.
Sandvik, P. U. 2007. Plantemakrofossil og mikrofossil. In:
Bårdseth, G. A. (ed.) Hus og gard langs E6 i Råde kom-
mune. E6-prosjektet Østfold. Band 1. Varia 65. Oslo:
Kulturhistorisk museum, Fornminneseksjonen.
Sandvik, P. U. 2008. Naturvitskapleg syntese. In: Bårdseth,
G. A. (ed.) Evaluering— resultat. E6-prosjektet Østfold. Band
5. Varia 69. Pp. 61–77. Oslo, Kulturhistorisk museum,
Fornminneseksjonen.
Sarpaki, A. 1992a. A palaeobotanical study of the west
house, Akrotiri, Thera. The Annual of the British School of
Archaeology at Athens, 37, 219–30.
228 REFERENCES
Sarpaki, A. 1992b. The study of palaeoethnobotanical
material from Akrotiri, Thera and its aims. To archaeio-
botaniko hyliko: stochoi kai problimatiki. Akrotiri Thiras.
Eikosi chronica ereynas (1967–1987), Imerida. Athinai,
Anatypon ektos emporioy.
Sarpaki, A. 1995. Toumba Balomenou, Chaeronia: plant
remains from the Early and Middle Neolithic levels. In:
Kroll, H. and Pasternak, R. (eds.) Res Archaeobotanicae,
International Workgroup for Palaeoethnobotany, Proceedings
of the 9th Symposium. Pp. 5–15. Kiel, Oetker-Voges.
Sarpaki, A. 2009. Knossos, Crete: Invaders, ‘sea-goers’, or
previously ‘invisible’, the Neolithic plant economy
appears fully-fledged in 9,000 BP. In: Fairbairn, A. and
Weiss, E. (eds.) From foragers to farmers: papers in honour
of Gordon C. Hillman. Pp. 220–34. Oxford, UK: Oxbow
Books.
Sarpaki, A. and Jones, G. 1990. Ancient and modern culti-
vation of Lathyrus clymenum L. in the Greek islands. The
Annual of the British School at Athens, 85, 363–8.
Schäfer, H. I. 1973. Zur taxonomie der Vicia narbonensis-
gruppe. Kulturpflanze, 21, 211–73.
Schick, T. 1988. Nahal Hemar cave: cordage, basketry and
fabrics. Atiqot, 38, 31–43.
Schick, T. 1998. The Cave of the Warrior: a fourth millennium
burial in the Judean Desert. Jerusalem, Israel Antiquity
Authority.
Schick, T. 2003. A 6,000-year-old nobleman: finds from the
Cave of the Warrior. Jerusalem, The Israel Museum.
Schiel, V. 1913. De l’exploitation des dattiers dans
l’ancienne Babylonie. Revue d’Assyriologie et d’Archéologie
Orientale, 10, 1–9.
Schiemann, E. 1948. Weizen, roggen, gerste. Systematik, ges-
chichte, und verwendung. Jena, Fischer.
Schiemann, E. 1956. Fünfzig jahre Triticum dicoccoides.
Berichte der Deutschen Botanischen Gesellschaft, 69,
309–22.
Schlichtherle, H. 1977/1978. Vorläufiger bericht über die
archäobotanischen untersuchungen an Demircihüyük
(Nordwestanatolien). In: Korfmann, M. (ed.) Demircihüyük.
Pp. 45–53. Istambul, Istambuler Mitteilungen, Deutsches
Archäologisches Institut.
Schlichtherle, H. 1981. Cruciferen als nutzpflanzen in neo-
lithischen ufersiedlungen Südwestdeutschlands und
der Schweiz. Zeitschrift für Archäologie, 15, 113–24.
Schlumbaum, A., Blatter, R. and Neuhaus, J. M. 1998.
Coexistence of tetraploid and hexaploid naked wheats
in a Neolithic lake dwelling of central Europe: evidence
from morphology and ancient DNA. Journal of
Archaeological Science, 25, 1111–18.
Schneider, M. 1994. Verkohlte pflanzenreste aus einem
neolithischen Brunnen in Schletz, NÖ. Archäologie
Österreichs, 5, 18–22.
Schoch, W. H., Pawlik, H. and Schweingruber, F. H. 1988.
Botanische makroreste. Bern, Paul Haupt.
Scholz, H. 1983. Die unkraut-hirse (Panicum miliaceum
subsp. ruderale)-neue tatsachen und befunde. Plant sys-
tematics and evolution, 143, 233–44.
Schultze-Motel, J. (1968–94). Literatur über archäologische
Kulturpflanzenreste. [Literature on archaeological
remains of cultivated plants]. For 1965–67: Kulturpflanze
16, 215–30 (1968). For 1968: Jahresschrift für mitteldeutsche
Vorgeschichte 55, 55–63 (1971). For 1969: Kulturpflanze 19,
Kulturpflanze 21, 61–76 (1973). For 1972/3: Kulturpflanze
22, 61–76 (1974). For 1973/4: Kulturpflanze 23, 189–205
(1975). For 1974/5: Kulturpflanze 24, 159–78 (1976). For
1975/6: Kulturpflanze 25, 71–88 (1977). For 1976/7:
Kulturpflanze 26, 349–62 (1978). For 1977–8: Kulturpflanze
27, 229–45 (1979). For 1978/9: Kulturpflanze 28, 361–78
(1980). For 1979/80: Kulturpflanze 29, 447–63 (1981). For
1980/1: Kulturpflanze 30, 255–72 (1982). For 1981/2:
Kulturpflanze 31, 281–97 (1983). For 1982/83: Kulturpflanze
32, 229–43 (1984). For 1983/84: Kulturpflanze 33, 287–305
(1985). For 1984/85: Kulturpflanze 34, 317–33 (1986). for
1985/86: Kulturpflanze 35, 401–20 (1987). For 1986/87:
Kulturpflanze 36, 549–69 (1988). For 1987/88: Kultur-
pflanze 37, 427–51 (1989). For 1988/89: Kulturpflanze 38,
387–416 (1990). For 1989/90: Vegetation History and
Archaeology 1, 52–62 (1992). For 1990/91: Vegetation
History and Archaeology, 2, 47–57 (1993). For 1991/92:
Vegetation History and Archaeology, 3, 33–61 (1994). [The
reviews up to 1992 are in German; since then in
English.]
Schultze-Motel, J. 1979a. Die anbaugeschichte des leindot-
ters: Camelina sativa (L.) Crantz. Archaeo-Physika, 8, 267–81.
Schultze-Motel, J. 1979b. Die urgeschichtlichen reste des
schlafmohns (Papaver somniferum L.) und die enstehung
der art. Kulturpflanze, 27, 207–15.
Schultze-Motel, J. 1988a. Archäologische kulturpflanzen-
reste aus der Georgischen SSR (Teil 1). Kulturpflanze, 36,
421–35.
Schwartz, G. M., Curvers, H. H., Gerritson, F. A.,
MaCcormick, J. A., Miller, N. F. and Weber, J. A. 2000.
Excavation and survey in the Jabbul Plain, western
Syria: The Umm el-Marra project 1996–1997. American
Journal of Archaeology, 104, 419–62.
Schweinfurth, G. 1891. Aegyptens auswärtige beziehun-
gen hinsichtlich der kulturgewächse. Zeitschrift für
Ethnologie (Organ der Berliner Gesellschaft für Anthropologie,
Ethnologie, und Urgeschichte), 23, 649–69.
Schweingruber, F. H. 1979. Wildäpfel und prähistorische
Äpfel. Archaeo-Physika, 8, 283–94.
Scora, R. W. 1988. Biochemistry, taxonomy and evolution
of modern cultivated Citrus. Proceedings of the
International Society of Citriculture, 1, 277–89.

Sears, E. R. 1969. Wheat cytogenetics. Annual Review of
Genetics, 3, 451–68.
Sebastian, P., Schaefer, H., Telford, I. R. H. and Renner, S. S.
In press. Cucumber (Cucumis sativus) and melon (C.
melo) have numerous wild relatives in Asia and
Australia, and the sister species of melon is from
Australia. Proceedings of the National Academy of Sciences.
Sencer, H. A. and Hawkes, J. G. 1980. On the origin of cul-
tivated rye. Biological Journal of the Linnean Society, 13,
299–313.
Sethy, N. K., Choudary, S., Shokeen, B. and Bhatia, S. 2006.
Identification of microsatellite markers from Cicer retic-
ulatum: molecular variation and phylogenetic analysis.
Theoretical and Applied Genetics, 112, 347–57.
Shang, H.-Y., Wei, Y.-M., Wang, X.-R. and Zheng, Y.-L.
2006. Genetic diversity and phylogenetic relationships
in the rye genus Secale L. (rye) based on Secale cereale
microsatellite markers. Genetics and Molecular Biology,
29, 685–91.
Shanjani, P., Mardi, M., Pazouki, L., Hagidimitriou, M.,
Avanzato, D., Pirseyedi, S., Ghaffari, M. and Khayam
Nekoui, S. 2009. Analysis of the molecular variation
between and within cultivated and wild Pistacia species
using AFLPs. Tree Genetics and Genomes, 5, 447–58.
Sharma, H. G. and Waines, J. G. 1980. Inheritance of tough
rachis in crosses of Triticum monococcum and T. boeoti-
cum. Journal of Heredity, 71, 214–16.
Sharma, S. K., Knox, M. R. and Ellis, T. H. 1996. AFLP anal-
ysis of the diversity and phylogeny of Lens and its com-
parison with RAPD analysis. Theoretical Applied Genetics,
93, 751–8.
Sherratt, A. G. 1995. Alcohol and its alternatives: symbol
and substance in pre-industrial cultures. In: Goodman,
J., Lovejoy, P. E. and Sherratt, A. G. (eds.) Consuming hab-
its: drugs in history and anthropology. Pp. 11–46. London
and New York, Routledge.
Shewry, P. R., Napier, J. A., Sayanova, O., Smith, M., Cooke,
D. T., Stoker, G., Hill, J., Stobart, A. K. and Lapinskas, P.
1997. The use of biotechnology to develop new crops
and products. In: Smartt, J. and Haq, N. (eds.)
Domestication, Production and Utilisation of New Crops.
Pp. 76–87. Southampton, International Centre for
Under-Utilised Crops.
Shimotsuma, M. 1963. Cytogenetical and evolutionary
studies in the genus Citrullus. Seiken Zihô, 15, 24–34.
Simchoni, O. 1998. Reconstruction of the landscape and human
economy 19,000 BP in the Upper Jordan Valley by the botani-
cal remains found at Ohalo II. Ph.D. Thesis, Bar-Ilan
University. (Hebrew, English summary).
Simchoni, O. and Kislev, M. E. 2009. Relict plant remains
in the ‘caves of the spear’. Israel Exploration Journal, 59,
47–62.
REFERENCES 229
Simchoni, O., Kislev, M. E. and Melamed, Y. 2007. Bet-
Shean as a trade center of crops in the Bronze Age:
botanical and entomological evidence. In: Mazar,
A. and Mullins, R. A. (eds.) Excavations at Tel Bet-Shean
1989–1996. Vol. II, the Middle and Late Bronze Age strata
in Area R. Pp. 702–15. Jerusalem, The Israel Exploration
Society and The Institute of Archaeology, The Hebrew
University.
Singh, A. K. 1990. Cytogenetics and evolution in the
Cucurbitaceae. In: Bates, D. M., Robinson, R. W. and
Geffrey, C. (eds.) Biology and utilization of the Cucurbitaceae.
Pp. 10–26. Ithaca, NY: Cornell University Press.
Singleton, V. L. 1994. An enologist’s commentary on
ancient wines. In: McGovern, P. E., Fleming, S. J. and
Katz, S. H. (eds.) The origins and ancient history of wine.
Pp. 67–77. Philadelphia (PA), Gordon and Breach.
Širjaev, G. 1932. Generis Trigonella L. revisio critica V, sectio
Foenum graecum, Brno, Folia Facultatis Scientiarium
Naturalium Universitatis Masarykianae Brunensis.
Small, E. 1995. Hemp. In: Smartt, J. and Simmonds, N. W.
(eds.) Evolution of crop plants. 2 ed, Pp. 28–32. London,
Longman.
Small, E. 1997. Culinary Herbs. Ottawa, Ontario: National
Research Council.
Small, E. 2006. Culinary Herbs. 2 ed. Ottawa, Ontario: NRC
Research Press.
Small, E. and Cronquist, A. 1976. A practical and natural
taxonomy for Cannabis. Taxon, 25, 405–35.
Smartt, J. 1990. Grain legumes: evolution and genetic resources.
Cambridge, UK: Cambridge University Press.
Smartt, J. and Hymowitz, T. 1985. Domestication and evo-
lution of grain legumes. In: Summerfield, R. J. and
Roberts, E. H. (eds.) Grain legume crops. Pp. 37–72.
London, Collins.
Smith, B. D. 1998. The emergence of agriculture. New York,
Scientific American Library.
Smithson, J. B., Thompson, J. A. and Summerfield, R. G.
1985. Chickpea (Cicer arietinum L.). In: Summerfield, R. J.
and Roberts, E. H. (eds.) Grain legume crops. Pp. 312–90.
London, Collins.
Snogerup, S., Gustafsson, M. and von Bothmer, R. 1990.
Brasica sect. Brassica (Brassicaceae) I. Taxonomy and
variation. Willdenowia, 19, 271–365.
Solecki, R. S. 1975. Shanidar IV, a Neanderthal flower bur-
ial in Northern Iraq. Science, 190, 880–1.
Soltis, D. E. and Soltis, P. S. 1989. Isozymes in plant biology.
Portland, Dioscorides Press.
Soltvedt, E. C. 2000. Carbonised cereal from three Late
Neolithic and two Bronze Age sites in western Norway.
Environmental Archaeology, 5, 49–62.
Soltvedt, E. C., Løken, T., Prøsch, D., Børsheim, R. L. and
Oma, K. 2007. Bøndene på Kvålehodlene. Boplass-, jor-
230 REFERENCES
dbruks- og landskapsutvikling gjennom 6000 år på
Jæren, SV Norge. AmS-Varia 47, 215.
Song, K., Osborn, T. C. and Williams, P. H. 1990. Brassica
taxonomy based on nuclear restriction fragment length
polymorphism (RFLPs): 3. genome relationships in
Brassica and related genera and the origin of B. oleracea
and B. rapa (syn. B. campestris). Theoretical and Applied
Genetics, 79, 497–506.
Sonnante, G., Galasso, I. and Pignone, D. 2003. ITS
Sequence Analysis and Phylogenetic Inference in the
Genus Lens Mill. Annals of Botany, 91, 49–54.
Sonnante, G., Hammer, K. and Pignone, D. 2009. From the
cradle of agriculture a handful of lentils: History of
domestication. Rendiconti Lincei, 20, 21–37.
Spataro, G., Taviani, P. and Negri, V. 2007. Genetic varia-
tion and population structure in a Eurasian collection of
Isatis tinctoria L. Genetic Resources and Crop Evolution, 54,
573–84.
Spiegel-Roy, P. and Kochba, J. 1981. Inheritance of nut and
kernel traits in almond (Prunus amygdalus Batsch).
Euphytica, 30, 167–74.
Stager, L. E. 1985. First fruits of civilization. In: Tubb, J. N.
(ed.) Palestine in the Bronze and Iron Age: papers in honour
of Olga Tufnell. Pp. 172–87. London, Institute of
Archaeology.
Stearn, W. T. 1978. European species of Allium and allied
genera of Alliceae: a synonymic enumeration. Annales
Musei Goulandris, 4, 83–198.
Stewart, R. T. 1974. Paleobotanical investigation: 1972 sea-
son. In: Stager, L., Walker, A. and Wright, G. E. (eds.)
American expedition to Idalion. Pp. 123–9. Cyprus, Bulletin
of the American Schools of Oriental Research.
Stika, H.-P. 1991. Die paläoethnobotanische untersuchung
der Linearbandkeramischen siedlung Hilzingen, Kreis
Konstanz. Fundberichte aus Baden-Württemberg, 16,
63–104.
Stika, H.-P. 1996. Vorgeschichtliche pflanzenreste aus
Heilbronn-Klingenberg. archäobotanische untersuchungen
zum Michelsberger Erdwerk auf dem Schlossberg (Bandkeramik,
Michelsberger Kultur, Späthallstatt/Frühlatene). Stuttgart,
Konrad Theiss Verlag.
Stika, H.-P. 1999. Erste archäobotanische ergebnisse der
ausgrabungen 1997 in Ambrona (Prov. Soria). Madrider
Mitteilungen, 40, 61–5.
Stika, H.-P. 2005. Early Neolithic agriculture in Ambrona,
Provincia Soria, central Spain. Vegetation History and
Archaeobotany, 14, 189–97.
Stika, H.-P. and Frank, K. S. 1988. Die kirschpflaume: sys-
tematik, morphologie, verwendung, genetik und
archäologische funde. In: Küster, H. (ed.) Der prähistor-
ische mensch und seine umwelt: Festschrift U. Körber-
Grohne. Pp. 65–71. Stuttgart, Konrad Theiss.
Stol, M. 1987a. The Cucurbitaceae in the cuneiform texts.
Bulletin on Sumerian Agriculture, 3, 81–92.
Stol, M. 1987b. Garlic, onion, leek. Bulletin on Sumerian
Agriculture, 3, 57–80.
Storey, W. B. 1976. Fig. In: Simmonds, N. W. (ed.) Evolution
of crop plants. Pp. 205–8. London, Longman.
Storey, W. B. and Condit, I. J. 1969. Fig (Ficus carica). In:
Werda, F. P. and Wit, F. (eds.) Outlines of perennial crop
breeding in the tropics. Pp. 259–67. Wageningen, The
Netherlands.
Stuiver, M. and Reimer, P. J. 1993. Extended 14C data base
and revised CALIB radiocarbon calibration program.
Radiocarbon, 35, 215–30.
Stuiver, M., Kromer, B., Becker, B. and Ferguson, C. W.
1986. Radiocarbon age calibration back to 13300 years
bp and the 14C age matching of the German oak and
bristlecone pine chronologies. Radiocarbon, 28, 969–79.
Stutz, H. C. 1972. On the origin of cultivated rye. American
Journal of Botany, 59, 59–70.
Suess, H. E. 1970. The three causes of secular 14C fluctua-
tions, their amplitudes and time constants. In: Olsson, I.
U. (ed.) Radiocarbon variations and absolute chronology.
Pp. 595–605. New York, Wiley Interscience Division.
Sweeney, M. and McCouch, S. 2007. The complex history
of the domestication of rice. Annals of Botany, 100,
951–7.
Swidrak, I. 1999. A Celtic, La Tène trade centre in Ramsautal
in the Dürrnberg, Austria: macrofossil data towards
reconstruction of environment and food plants.
Vegetation History and Archaeobotany, 8, 113–16.
Täckholm, V. 1976. Faraos Blomster. Trelleborg, Sweden:
Generalstabens Litografiska.
Täckholm, V. and Drar, M. 1950. Cyperus esculentus. Phoenix
dactylifera. Flora of Egypt, Vol. II. Pp. 60–9, 165–273. Cairo,
Cairo University Press.
Täckholm, V. and Drar, M. 1954. Allium in ancient Egypt.
Flora of Egypt, Vol. III. Pp. 93–106. Cairo, Cairo University
Press.
Takahashi, R. 1955. The origin and evolution of cultivated
barley. Advances in Genetics, 7, 227–66.
Takahashi, R. 1964. Further studies on the phylogenetic
differentiation of cultivated barley. In: Broekhuizen, S.
(ed.) Barley genetics I, oroceedings first international barley
genetics symposium. Pp. 19–26. Wageningen, Centre for
Agricultural Publications and Documentation.
Takahashi, R. 1972. Non brittle rachis 1 and non brittle
rachis 2. Barley Genetics Newsletter, 2, 181–2.
Tanaka, M. and Ishii, H. 1973. Cytogenetic evidence on the
speciation of wild tetraploid wheats collected in Iraq,
Turkey and Iran. In: Sears, E.R. and Sears, L.M.S. (eds.)
Proceedings, 4th international wheat genetics symposium.
Columbia, MI: University of Missouri.

Tanaka, M., Kawahara, T. and Sano, J. 1979. The origin and
the differentiation of the B and G genomes of the tetra-
ploid wheats. Report of the Plant Germplasm. Institute,
Kyoto University, 4, 1–11.
Tanno, K.-i. and Willcox, G. 2006a. How fast was wild
wheat domesticated? Science, 311, 1886.
Tanno, K.-i. and Willcox, G. 2006b. The origins of cultiva-
tion of Cicer arietinum L. and Vicia faba L.: early finds
from Tell el-Kerkh, north-west Syria, late 10th millen-
nium. Vegetation History and Archaeobotany, 15, 197–204.
Tanno, K-i. and Willcox, G. (In press). Distinguishing wild
and domestic wheat and barley spikelets from early
Holocene sites in the Near East. Vegetation History and
Archaeobotany.
Tempír, Z. 1966. Výsledky paleoetnobotanického studia
pĕstování zemĕdĕlských rostlin na územi ČSSR (Results
of paleoethnobotanic studies on the cultivation of agri-
cultural plants in the ČSSR). Vĕdecké práce Československého
zemĕlského muzea (Praha), 1966, 27–144.
Tempír, Z. 1969. Archeologické nálezy zemĕdĕlských rost-
lin a plevelŭ na Slovensku (Archaeological finds of food
plants and weeds in Slovakia). Agrikultúra, 8, 7–66.
Tempír, Z. 1973. Nálezy prayekych a stredovékych zoyt-
sku pestovanych a uzitkovych rosdin a plevelu na nek-
terych lokalitách v Cechách a na Morave (Finds of
prehistoric and early historic remains of food plants and
weeds in some sites in Bohemia and Moravia). Vĕdecké
práce Československého zemĕlského muzea (Praha),1973,
19–47.
Tempír, Z. 1979. Kulturpflanzen im Neolithikum und
Äneolithikum auf dem Gebiet von Böhmen und Mähren.
Archaeo-Physika, 8, 302–8.
Tengberg, M. 1999. Crop husbandry at Miri Qalat: Makran,
SW Pakistan 4000–2000 BC. Vegetation History and
Archaeobotany, 8, 3–12.
Terral, J.-F., Tabard, E., Bouby, L., Ivorra, S., Pastor, T.,
Figueiral, I., Picq, S., Chevance, J.-B., Jung, C., Fabre, L.,
Tardy, C., Compan, M., Bacilieri, R., Lacombe, T. and
This, P. 2010. Evolution and history of grapevine (Vitis
vinifera) under domestication: new morphometric per-
spectives to understand seed domestication syndrome
and reveal origins of ancient European cultivars. Annals
of Botany, 105, 443–55.
Thanheiser, U. 1991. Untersuchungen zur Landwirtschaft
der vor- und Frühdynastischen Zeit in Tell-el-Fara‘in—
Buto. Ägypten Und Levante, 2, 39–45.
Thanheiser, U. 1992. Plant-food at Tell Ibrahim Awad: pre-
liminary report. In: van den Brink, E. C. M. (ed.) The Nile
delta in transition; 4th–3rd Millennium B.C. Pp. 117–21.
Jerusalem, Israel Exploration Society.
Thanheiser, U. 1997. Botanische funde. In: Hiller, S. and
Nikolov, V. (eds.) Karanovo, die ausgrabungen im Südsektor
REFERENCES 231
1984–1992. Pp. 429–54. Salzburg—Sofia, Ferdinand
Berger and Sohne.
Thanheiser, U. 2004. Die pflanzenreste. In: Hein, I. and
Jánosi, P. (eds.) Tell el-Daba XI. Areal A/V. Siedlungsrelikte
der späten 2. Zwischenzeit. Vienna, Österreichischen
Akademie der Wissenschaften.
Thanheiser, U., Walter, J. and Hope, C. A. 2002. Roman
agriculture and gardening in Egypt as seen from Kellis.
In: Hope, C. A. and Bowen, G. E. (eds.) Dakhleh Oasis
project: preliminary reports on the 1994–1995 to 1998–1999
seasons. Pp. 299–310. Oxford, U.K.: Oxbow Books.
Thiebault, S. 1989. A note on the ancient vegetation of
Baluchistan based on charcoal analysis of the latest peri-
ods from Mehrgarh, Pakistan. In: Frifelt, K. and
Sorensen, P. (eds.) South Asian srchaeology 1985. Papers
from the 8th international conference of the association of
South Asian Archaeologists in Western Europe. Pp. 186–8.
London, Curzon Press.
Thompson, R. C. 1949. A dictionary of Assyrian botany.
London, The British Academy.
Tosi, M. 1975. Hasanlu project 1974: paleobotanical survey.
Iran, 8, 185–6.
Townsend, C. C. 1974. Vicia ervilia. In: Townsend, C. C. and
Guest, E. (eds.) Flora of Iraq. Pp. 526. Baghdad, Iraq:
Ministry of Agriculture and Agrarian Reform.
Tutin, T. G. 1993. Ficus L. In: Tutin, T. G., Burges, N. A.,
Chater, A. O. and Edmondson, J. R. (eds.) Flora Europaea.
2 ed, Pp. 78. Cambridge, UK: Cambridge University
Press.
U, N. 1935. Genome analysis in Brassica. Japan Journal of
Botany, 7, 389–452.
Valamoti, S. 2011. Ground cereal food preparations from
Greece: the prehistory and modern survival of tradi-
tional Mediterranean ‘fast foods’. Archaeological and
Anthropological Sciences, 3, 19–39.
Valamoti, S. M. and Kotsakis, K. 2007. Transitions to agri-
culture in the Aegean: the archaeobotanical evidence.
In: Colledge, S. and Conolly, J. (eds.) The origins and
spread of domestic plants in Southwest Asia and Europe. Pp.
75–92. Walnut Creeck, CA: Left Coast Press.
Valdeyron, G. and Lloyd, D. G. 1979. Sex differences and
flowering phenology in the common fig Ficus carica.
Evolution, 33, 673–85.
Valkoun, J., Waines, J. G. and Konopka, J. 1998. Current
geographical distribution and habitat of wild wheats
and barley. In: Damania, A. B., Valkound, J., Willcox, G.
and Qualset, C. O. (eds.) Origins of agriculture and crop
domestication. Pp. 293–9. Aleppo, Syria: ICARDA.
van de Ven, W. T. G., Duncan, N., Ramsey, G., Phillips,
M., Powell, W. and Waugh, R. 1993. Taxonomic
relationships between Vicia faba and its relatives
based on nuclear and mitochondrial RFLPs and
232 REFERENCES
PCR analysis. Theoretical and Applied Genetics , 86 ,
71–80.
van der Maesen, L. J. G. 1972. Cicer L., a monograph on the
genus with special reference to the chickpea (Cicer arietinum),
its ecology and cultivation,. Wageningen, Agricultural
University.
van der Maesen, L. J. G. 1987. Origin, history, and taxon-
omy of chickpea. In: Saxena, M. C. and Singh, K. B.
(eds.) The Chickpea. Pp.11–34. Wallingford, UK: CAB
International Publications.
van der Veen, M. 1992a. Botanical evidence for Gara-
mantian agriculture in Fezzan, southern Libya. Review
of Palaeobotany and Palynology, 73, 315–27.
van der Veen, M. 1992b. Garamantian agriculture: the
plant remains from Zinchecra, Fezzan. Libyan Studies,
23, 7–39.
van der Veen, M. 1995. Ancient agriculture in Lybia: a
review of the evidence. Acta Palaeobotanica, 35, 85–98.
van der Veen, M. 1998. A life of luxury in the desert? The
food and fodder supply to Mons Claudianus. Roman
Archaeology, 11, 101–16.
van der Veen, M. 1999. The Exploitation of Plant Resources in
Ancient Africa. London, Kluwer Academic and Plenum
Publishers.
van der Veen, M. 2001a. The botanical evidence. In:
Maxfield, V. A. and Peacock, D. P. S. (eds.) Survey and
excavations at Mons Claudianus 1987–1993. volume 2: the
excavations: part 1. Pp. 174–247. Cairo, Institut Français
d’Archéologie Orientale du Caire.
van der Veen, M. 2001b. Survey and excavations at Mons
Claudianus 1987–1993. Vol. 2: The excavations: Part 1.
In: Maxfield, V. A. and Peacock, D. P. S. (eds.) Documents
de fouilles. Pp. 174–247. Cairo, Institut Français
d’Archéologie Orientale du Caire.
van der Veen, M. 2003. Trade and diet at Roman and medi-
eval Quseir al-Qadim, Egypt: a preliminary report. In:
Neumann, K., Butler, A. and Kahlheber, S. (eds.) Food, fuel
and fields, progress in African Archaeobotany. Pp. 207–12.
Cologne, Heinrich Barth Institut.
van der Veen, M. 2006. Food and farming in the Libyan
Sahara. In: Mattingly, D., McLaren, S., Savage, E.,
Fasatwi, Y. and Gadgood, K. (eds.) The Libyan Desert:
natural resources and cultural heritage. Pp. 171–8. London,
Society for Libyan Studies, Monograph.
van der Veen, M. and Tabinor, H. 2007. Food, fodder and
fuel at Mons Porphyrites: the botanical evidence. In:
Maxfield, V. A. and Peacock, D. P. S. (eds.) Survey and
excavation at Mons Porphyrites 1994–1998. volume 2: the
excavations. Pp. 83–142. London, Egypt Exploration
Society.
van der Veen, M., Grant, A. and Barker, G. 1996. Romano-
Libyan agriculture: crops and animals. In: Barker, G.,
Gilbertson, D., B., J. and Mattingly, D. (eds.) Farming the
desert: The UNESCO Libyan valleys archaeological survey.
Vol. 1: synthesis. Pp. 227–63 (bibliography 365–391).
Paris, UNESCO.
van der Veen, M., Morales, J. and Cox, A. 2009. Food and
culture: the plant foods from Roman and Islamic Quseir,
Egypt. In: Fairbairn, A. S. and Weiss, E. (eds.) From forag-
ers to farmers: papers in honour of Gordon C. Hillman.
Pp. 263–70. Oxford, Oxbow Books.
van Oss, H., Aron, Y. and Ladizinsky, G. 1997. Chloroplast
DNA variation and evolution in the genus Lens Mill.
Theoretical and Applied Genetics, 94, 452–7.
van Slageren, M. W. 1994. Wild wheats: a monograph of
Aegilops L. and Amblyopyrum (Jaub. and Spach) Eig
(Poaceae). Wageningen Agricultural University Papers,
The Netherlands.
van Zeist, W. 1994. Some notes on second millennium BC
plant cultivation in the Syrian Jazira. In: Gasche, H. and
Tanret, M. (eds.) Cinquante-deux réflections sur le Proche-
Orient ancien offertes en hommage à Léon de Meijer.
Pp. 541–53. Leuven, Peeters.
van Zeist, W. 2001. Third to first millennium B.C. plant
cultivation on the Khabur, north-eastern Syria.
Palaeohistoria, 41–42, 111–25.
van Zeist, W. 1972. Palaeobotanical results in the 1970 sea-
son at Çayönü, Turkey. Helinium, 12, 3–19.
van Zeist, W. 1975. Preliminary report on the botany of
Gomolava. Journal of Archaeological Science, 2, 315–25.
van Zeist, W. 1976. On macroscopic traces of food plants in
southwestern Asia (with some references to pollen
data). Philosophical transactions of the Royal Society of
London, Biological Sciences, 275, 27–41.
van Zeist, W. 1979–80. Plant remains from Girikihaciyan,
Turkey. Anatolica, 7, 75–89.
van Zeist, W. 1980. Aperçu sur la diffusion des végétaux
cultivés dans la région Méditerranéene. In: Emberger, F.
L. (ed.) Colloque sur la mise en place, l’évolution et la cara-
ctérisation de la flore et la végétation circumméditerranéene.
Pp. 129–45. Montpellier, Naturalia Monspeliensia,
Special volume 1980.
van Zeist, W. 1981. Plant remains from Cap Andreas-
Kastros (Cyprus). In: Le Brun, A. (ed.) Un site Néolithique
Précéramique en Chypre: Cap Andreas-Kastros. Pp. 95–100.
Paris, Recherche sur les Grandes Civilisations.
van Zeist, W. 1983. Fruits in foundation deposits of two
temples. Journal of Archaeological Science, 10, 351–4.
van Zeist, W. 1988. Some aspects of early Neolithic plant
husbandry in the Near East. Anatolica, 15, 49–67.
van Zeist, W. 1991. Economic aspects. In: van Zeist, W.,
Wasilikowa, K. and Behre, K.-E. (eds.) Progress in Old
World Palaeoethnobotany. Pp. 109–30. Rotterdam,
Balkema.

van Zeist, W. 2003. Plant husbandry and vegetation of tell
Gomolava, Vojvodina, Yugoslavia. Palaeohistoria, 43/44,
87–115.
van Zeist, W. and Bakker-Heeres, J. A. H. 1975. Evidence
of linseed cultivation before 6000 bc. Journal of
Archaeological Science, 2, 215–19.
van Zeist, W. and Bakker-Heeres, J. A. H. 1982 (1985).
Archaeological studies in the Levant 1. Neolithic sites in
the Damascus basin: Aswad, Ghoraifé, Ramad.
Palaeohistoria, 24, 165–256.
van Zeist, W. and Bakker-Heeres, J. A. H. 1986.
Archaeological studies in the Levant 2. Neolithic and
Halaf levels at Ras Shamra. Palaeohistoria, 26, 151–70.
van Zeist, W. and Bottema, S. 1971. Plant husbandry in
Early Neolithic Nea Nikomedeia, Greece. Acta botanica
neerlandica, 20, 524–38.
van Zeist, W. and Buitenhuis, H. 1983. A palaeobotanical
study of Neolithic Erbaba, Turkey. Anatolica, 10,
47–89.
van Zeist, W. and Casparie, W. A. 1968. Wild einkorn
wheat and barley from Tell Mureybit in northern Syria.
Acta botanica neerlandica, 17, 44–53.
van Zeist, W. and de Roller, G. J. 1991–2. The plant hus-
bandry of aceramic Çayönü, S.E. Turkey. Palaeohistoria,
33/34, 65–96.
van Zeist, W. and de Roller, G. J. 1993. Plant remains from
Maadi, a predynastic site in lower Egypt. Vegetation
History and Archaeobotany, 2, 1–14.
van Zeist, W. and de Roller, G. J. 1995. Plant remains from
Asikli Höyük, a pre-pottery Neolithic site in central
Anatolia. Vegetation History and Archaeobotany, 4,
179–85.
van Zeist, W. and de Roller, G. J. 2003. The Çayönü
Archaeobotanical Record. In: van Zeist, W. (ed.) Reports
on archaeobotanical studies in the Old World. Pp. 143–66.
Groningen, The Groningen Institute of Archaeology,
University of Groningen.
van Zeist, W. and Palfenier-Vegter, R. M. 1981. Seeds and
fruits from the Swifterbant S3 site. Palaeohistoria, 23,
105–68.
van Zeist, W. and Vynckler, J. 1984. Palaeobotanical inves-
tigations of Tell ed-Der. In: De Meyer, L. (ed.) Tell ed-Der
IV. Pp. 119–43. Leuven, Uitgeverij Peeters.
van Zeist, W. and Waterbolk-van Rooijen, W. 1985. The
palaeobotany of Tell Bouqras, eastern Syria. Paléorient,
11, 131–47.
van Zeist, W., Roller, G. and Fahmy, A. G. 2003. An archaeo-
botanical study of Ma’adi, a Predynastic site in Lower
Egypt. In: van Zeist, W. (ed.) Reports on archaeobotanical
studies in the Old World. Pp. 167–207. Groningen, The
Groningen Institute of Archaeology, University of
Groningen.
REFERENCES 233
Varshney, R. K., Graner, A. and Sorrells, M. E. 2005. Genic
microsatellite markers in plants: features and applica-
tions. Trends in Biotechnology, 23, 48–55.
Vaughan, D. A., Lu, B.-R. and Tomooka, N. 2008. The
evolving story of rice evolution. Plant Science, 174,
394–408.
Vaughan, J. and Geissler, C. 2009. The New Oxford book of
food plants. Oxford, UK: Oxford University Press.
Vavilov, N. I. 1949–50. Phytogeographic basis of plant
breeding. In: Vavilov, N. I. (ed.) The origin, variation,
immunity, and breeding of cultivated plants. Pp. 46–54.
Waltham, Mass: Chronica Botanica.
Vavilov, N. I. 1987. Origin and geography of cultivated plants.
Cambridge, UK: Cambridge University Press.
Velasco, R. et al. 2010. The genome of the domesticated
apple (Malus × domestica Borkh.). Nature Genetics,
42(10), 833–41.
Vences, F. J., Vaquero, F., Gracia, P. and Perez De La Vega,
M. 1987. Further studies on phylogenetic relationships
in Secale: on the origin of its species. Plant Breeding, 98,
281–91.
Vermeeren, C. and Cappers, R. T. J. 1997/2002. Ethno-
graphic and archaeobotanical evidence of local cultiva-
tion of plants in Roman Berenike and Shenshef (Red Sea
coast, Egypt). Pp. 1–14. BIAXiaal, 140, Zaandam,
Nederland.
Villaret-Von Rochow, M. 1958. Die pflanzenreste der
Bronzezeitlichen pfahlbauten von Valeggio am Mincio.
Berichte über das Geobotanische Forschungsinstitut Rübel in
Zürich für das Jahr 1957, 96–144.
Villaret-Von Rochow, M. 1967. Frucht- und samenreste aus
der Neolithischen station Seeberg, Burgäschisee-Süd.
Acta Bernensia, 2, 21–64.
Villaret-Von Rochow, M. 1971. Avena ludoviciana Dur. im
Schweizer Spätneolithikum, ein beitrag zur abstam-
mung des Saathafers. Berichte der Deutschen Botanischen
Gesellschaft, 84, 243–8.
Vishnu Mittre and Savithri, R. 1982. Food economy of the
Harappans. In: Possehl, G. L. (ed.) Harappan civilization:
a contemporary prospective. Pp. 205–21. New Delhi,
Oxford and IBH Publishing Co.
Vollesen, K. 1987. The native species of Gossypium
(Malvaceae) in Africa, Arabia and Pakistan. Kew Bulletin,
42, 337–49.
von Bothmer, R., Jacobsen, N., Jørgensen, R. B. and Linde-
Laursen, I. 1991. An ecogeographical study of the genus
Hordeum. Systematic and ecogeographic studies on crop
genepools 7. Rome, International Board for Plant
Genetic Resources.
Vuorela, I. and Lempiäinen, T. 1988. Archaeobotany of the
site of the oldest cereal grain find in Finland. Annales
Botanici Fennici, 25, 33–45.
234 REFERENCES
Wagner, I. and Weeden, N. F. 2000. Isozymes in Malus syl-
vestris, Malus domestica and in related Malus species.
Acta Horticulturae (ISHS), 538, 51–6.
Waines, J. G. 1975. The biosystematics and domestication
of pea (Pisum L.). Bulletin of the Torrey Botanical Club, 102,
385–95.
Waines, J. G. 1996. Molecular characterization of the
einkorn wheat. In: Padulosi, S., Hammer, K. and Heller,
J. (eds.) Hulled wheats. Pp. 193–7. Rome, International
Plant Genetic Resources Institute.
Waines, J. G. and Barnhart, D. 1992. Biosystematic research
in Aegilops and Triticum. Hereditas, 116, 207–12.
Waines, J. G. and Stanley Price, N. P. 1975–1977. Plant
remains from Khirokitia in Cyprus. Paléorient, 3, 281–4.
Wang, A., Yu, Z. and Ding, Y. 2009. Genetic diversity anal-
ysis of wild close relatives of barley from Tibet and the
Middle East by ISSR and SSR markers. Comptes Rendus
Biologies, 332, 393–403.
Warburg, O. 1905. Die gattung Ficus im nichttropischen
vorderasien. In: Ign, U. and Graebner, P. (eds.)
Festschrift Paul Ascherson. Pp. 364–70. Borntraeger,
Leipzig: Gebr.
Ward, C. 2003. Pomegranates in eastern Mediterranean
contexts during the Late Bronze Age. World Archaeology,
34, 529–41.
Wasylikowa, K. and Kubiak-Martens, L. 1995. Wild sor-
ghum from the Early Neolithic site at Nabta Playa,
south Egypt. In: Kroll, H. and Pasternak, R. (eds.) Res
archaeobotanicae, international workgroup for palaeoethno-
botany, proceedings of the 9th symposium. Pp. 345–58. Kiel,
Oetker-Voges.
Wasylikowa, K. and van der Veen, M. 2004. An archaeobo-
tanical contribution to the history of watermelon, Citrullus
lanatus (Thunb.) Matsum. and Nakai (syn. C. vulgaris
Schrad.). Vegetation History and Archaeobotany, 13, 213–17.
Wasylikowa, K., Barakat, H. N., Boulos, L., Butler, A.,
Dahlebrg, J. A., Hather, J. and Mitka, J. 2001. Vegetation
and subsistence of the Early Neolithic at Nabta Playa,
Egypt, reconstructed from charred plant remains. In:
Wendorf, F. and Schild, R. (eds.) Holocene settlement of the
Egyptian Sahara. Pp. 544–91. New York, Kluwer Academic/
Plenum Publishers.
Wasylikowa, K., Carciumaru, M., Hajnalová, E. P.,
Hartyányi, B., Pashkevich, G. A. and Yanushevich, Z. V.
1991. East-Central Europe. In: van Zeist, W., Wasylikowa,
K. and Behre, K.-E. (eds.) Progress in Old World
Palaeoethnobotany. Pp. 207–39. Rotterdam, Balkema.
Watanabe, N. Fujii, Y. Kato, N. Ban, T. and Martinek, P.
2006. Microsatellite mapping of the genes for brittle
rachis on homologous group 3 chromosomes in tetra-
ploid and hexaploid wheats. Journal of Applied Genetics,
47, 93–8.
Watanabe, N. Sugiyama, K. Yamagishi, Y. and Sakata, Y.
2002. Comparative telosomic mapping of homoeolo-
gous genes for brittle rachis in tetraploid and hexaploid
wheats. Hereditas, 137,180–5.
Watkins, R. 1986. Apples (genus Malus), pears (genus
Pyrus), and plums, apricots, almonds, peaches, cherries,
(genus Prunus). In: Hora, B. (ed.) The Oxford encyclopae-
dia of trees of the world. 2 ed, Pp. 197–201. Oxford, Oxford
University Press.
Watkins, R. 1995. Cherry, plum, peach, apricot and almond.
In: Smartt, J. and Simmonds, N. W. (eds.) Evolution of
crop plants. 2 ed, Pp. 423–8. London, Longman.
Watson, A. M. 1983. Agricultural innovation in the early
Islamic world: The diffusion of crops and farming techniques
700–1100. Cambridge, UK: Cambridge University
Press.
Way, R. D., Aldwinckle, H. S., Lamb, R. C., Rejman, A.,
Sansavini, S., Shen, T., Watkins, R., Westwood, M. N.
and Yoshida, Y. 1991. Apple (Malus). In: Moore, J. N. and
Ballington, J. R. (eds.) Genetic resources of temperate fruit
and nut crops. Pp. 3–62. Wageningen, International
Society for Horticultural Science.
Webb, D. A. 1968. Prunus , Vitis. In: Tutin, T. G.,
Heywood, V. H., Burges, N. A., Moore, D. M.,
Valentine, D. H., Walters, S. M. and Webb, D. A.
(eds.) Flora Europaea . 2 ed. Cambridge, UK:
Cambridge University Press.
Weinstein-Evron, M. 1983. The paleoecology of the early
Würm in the Hula basin, Israel. Paléorient, 9, 5–19.
Weiss, E. 2002. Issues in reconstruction the human economy and
society of the Epipalaeolithic site Ohalo II inhabitants by the
macrofossil botanical remains. Ph.D., Bar-Ilan University.
Weiss, E. 2009. Glimpsing into a hut: economy and society
of Ohalo II inhabitants. In: Fairbairn, A. and Weiss, E.
(eds.) From foragers to farmers: papers in honour of Gordon
C. Hillman. Pp. 153–60. Oxford, UK: Oxbow Books.
Weiss, E. and Kislev, M. E. 2004. Plant remains as indica-
tors for economic activity: a case study from Iron Age
Ashkelon. Journal of Archaeological Science, 31, 1–13.
Weiss, E., Kislev, M. E. and Hartmann, A. 2006. Autonomous
cultivation before domestication. Science, 312, 1608–10.
Weiss, E., Kislev, M. E., Simchoni, O., Nadel, D. and
Tschauner, H. 2008. Plant-food preparation area on an
Upper Paleolithic brush hut floor at Ohalo II, Israel.
Journal of Archaeological Science, 35, 2400–14.
Weiss, E., Wetterstrom, W., Nadel, D. and Bar-Yosef, O.
2004. The broad spectrum revisited: Evidence from
plant remains. Proceedings of the National Academy of
Sciences of the United States of America, 101, 9551–5.
Wendel, J. F. 1995. Cotton. In: Smartt, J. and Simmonds, N.
W. (eds.) Evolution of crop plants. 2 ed, Pp. 358–66.
London, Longman.

Wendel, J. F. and Cronn, R. C. 2003. Polyploidy and the
evolutionary history of cotton. Advances in Agronomy,
73, 139–86.
Wendel, J. F., Olson, P. D. and Stewart, J. M. 1989. Genetic
diversity, introgression, and independent domestication
of Old World cultivated cottons. American Journal of
Botany, 76, 1795–806.
Wendrich, W. Z. and Cappers, R. C. 2005. Egypt’s earliest
granaries, evidence from the Fayum. Egyptian
Archaeology, 27, 12–15.
Werker, E., Marbach, I. and Mayer, A. M. 1979. Relation
between the anatomy of the testa, water permeability
and the presence of phenolics in the genus Pisum. Annals
of Botany, 43, 765–71.
Werneck, H. L. 1949. Ur-und frühgeschichtliche kultur-
und nutzpflanzen in den Ostalpen und am rande des
Böhmerwaldes. Schriftenreihe der ober- Österreichischen
Landesbaudirektion, 6, 288.
Werneck, H. L. and Bertsch, K. 1959. Zur ur- und frühges-
chichte der pflaumen im oberen Rhein- und
Donauraume. Angewandte Botanik, 33, 19–33.
Western, C. 1971. The ecological interpretation of ancient
charcoals from Jericho. Levant, 3, 31–40.
Wetterstrom, W. 1993. Foraging and farming in Egypt:
the transition from hunting and gathering to horti-
culture in the Nile Valley. In: Shaw, T., Sinclair, P.,
Andah, B. and Okpoko, A. (eds.) The archaeology of
Africa: food, metals and towns. Pp. 165–226. London,
Routledge.
Wetterstrom, W. 1998. The origins of agriculture in Africa:
with particular reference to sorghum and pearl millet.
The Review of Archaeology, 19, 30–46.
White, K. D. 1970. Roman farming. Ithaca, Cornell
University Press.
Wild, J. P. 1997. Cotton in Roman Egypt: some problems of
origin. Al-Rafidan, 18, 287–98.
Willcox, G. 1991. Carbonized plant remains from
Shortughai, Afghanistan. In: Renfrew, J. (ed.) New light
on early farming. Pp. 191–212. Edinburgh, Edinburgh
University Press.
Willcox, G. 2000. Présence des céréales dans le Néolithique
précéramique de Shillourokambos à Chypre: résultats
de la campagne 1999. Paléorient, 26, 129–35.
Willcox, G. 2002. Charred plant remains from a 10th mil-
lennium B.P. kitchen at Jerf el Ahmar (Syria). Vegetation
History and Archaeobotany, 11, 55–60.
Willcox, G. 2005. The distribution, natural habitats and
availability of wild cereals in relation to their domesti-
cation in the Near East: multiple events, multiple cen-
tres. Vegetation History and Archaeobotany, 14, 534–41.
Willcox, G. 2007. The adoption of farming and the begin-
nings of the Neolithic in the Euphrates valley. In:
REFERENCES 235
Colledge, S. and Connoly, J. (eds.) Agriculture in south-
west Asia and Europe. Pp. 21–36. Walnut Creek, CA: Left
Coast Press.
Willcox, G. and Fornite, S. 1999. Impressions of wild cere-
als chaff in pisé from the 10th millennium uncal B.P. at
Jerf el Ahmar and Mureybet: northern Syria. Vegetation
History and Archaeobotany, 8, 21–4.
Willcox, G., Buxo, R. and Herveux, L. 2009. Late Pleistocene
and early Holocene climate and the beginnings of culti-
vation in northern Syria. The Holocene, 19, 151–8.
Willcox, G., Fornite, S. and Herveux, L. 2008. Early
Holocene cultivation before domestication in northern
Syria. Vegetation History and Archaeobotany, 17, 313–25.
Willerding, U. 1970. Vor- und frühgeschichtliche kulturp-
flanzen in Mitteleuropa. Neue Ausgrabungen und
Forschungen in Niedersachsen, 5, 287–375.
Willerding, U. 1980. Zum Ackerbau der Bandkeramiker.
In: Kruger, T. and Stephan, H.-G. (eds.) Beiträge zur
Archäologie Nordwestdeutschlands und Mitteleuropas.
Materialhefte zur Ur- und Frühgeschichte Niedersachsens.
Pp. 421–56. Hildesheim, A. Lax.
Willerding, U. 1996. Zur agrarproduktion von der jüngeren
Vorrömischen Eisenzeit bis ins frühe Mittellalter.
Historicum (Linz, Austria), Frühling, 96, 10–20.
Wilson, H. 1988. Egyptian food and drink. Aylesbury, Shire
Publications.
Winterhalter, P. and Straubinger, M. 2000. Saffron —
renewed interest in an ancient spice. Food Reviews
International, 16, 39–59.
Wright, T. W. 1986. Hazels (genus Corylus). In: Hora, B.
(ed.) The Oxford encyclopaedia of the trees of the world. 2 ed,
Pp. 143–4. Oxford, Oxford University Press.
Zaitschek, D. V. 1961. Remains of cultivated plants from
the cave of Nahal Mishmar: preliminary note. Israel
Exploration Journal, 11, 70–2.
Zaitschek, D. V. 1962. Remains of cultivated plants from
the Cave of the Pool. Israel Exploration Journal, 12 184–5.
Zapata, L. 2002. Origen de la agricultura en el Pa´ıs Vasco y
transformaciones en el paisaje: An´alisis de restos vegetales
arqueol´ ogicos. Bilbao, Diputacion Foral de Bizkaia.
Zaprjagaeva, V. I. 1964. Dikorastuščie plodovye Tadžikistana
(Wild growing fruits of Tadzhikistan). Moskva/Leningrad,
Nauka.
Zech-Matterne, V. and Leconte, L. 2010. New archaeobo-
tanical finds of Isatis tinctoria L. (woad) from Iron Age
Gaul and a discussion of the importance of woad in
ancient time. Vegetation History and Archaeobotany, 19,
137–42.
Zeder, M. A. 2011. The Origins of Agriculture in the Near
East. Current Anthropology, 52 (suppl. 4): S221–S235.
Zettler, R. I. and Miller, N. F. 1995. Searching for wine in
the archaeological record of ancient Mesopotamia of the
236 REFERENCES
third and second millennia B.C. In: McGovern, P. E.,
Fleming, S. J. and Katz, S. H. (eds.) The origins and ancient
history of wine. Pp. 123–31. Luxembourg, Gordon and
Breach.
Zhao, Z. 2011. New archaeobotanic data for the study of
the origins of agriculture in china. Current Anthropology,
52(suppl. 4), S295–S306.
Zhimin, A. 1989. Prehistoric agriculture in China. In:
Harris, D. R. and Hillman, G. C. (eds.) Foraging and farm-
ing: the evolution of plant exploitation. Pp. 643–9. London,
Unwin and Hyman.
Zhukovsky, P. M. 1964. Kulturnie rastenia i ich sorodiczy.
Leningrad, Kolos.
Zielinski, Q. B., Reimer, F. C. and Quackenbush, V. L. 1965.
Breeding behavior of fruit characteristics in pears, Pyrus
communis L. Proceedings of the American Society for
Horticultural Science, 86, 81–7.
Zohary, D. 1960. Studies on the origin of cultivated barley.
Bulletin of the Research Council of Israel, 9D, 21–42.
Zohary, D. 1965. Colonizer species in the wheat group. In:
Baker, H. G. and Stebbins, G. L. (eds.) The genetics of colo-
nizer species. Pp. 403–23. New York, Academic Press.
Zohary, D. 1969. The progenitors of wheat and barley in
relation to domestication and agriculture dispersal in
the Old World. In: Ucko, P. J. and Dimbleby, G. W. (eds.)
The domestication and exploitation of plants and animals.
Pp. 47–66. London, Duckworth.
Zohary, D. 1971. Origin of southwest Asiatic cereals:
wheats, barley, oats, and rye. In: Davis, P. H., Harper,
P. T. and Hedge, I. (eds.) Plant life of south-west
Asia. Pp. 235–63. Edinburgh, Botanical Society of
Edinburgh.
Zohary, D. 1984. Modes of evolution in plants under
domestication. In: Grant, W. F. (ed.) Plant biosystematics.
Pp. 579–86. Montreal, Academic Press Canada.
Zohary, D. 1989a. Domestication of southwest Asian
Neolithic crop assemblage of cereals, pulses, and flax:
the evidence from the living plants. In: Harris, D. R. and
Hillman, G. C. (eds.) Foraging and farming: the evolution of
plant exploitation. Pp. 358–73. London, Unwin and
Hyman.
Zohary, D. 1989b. Pulse domestication and cereal domesti-
cation: how different are they? Economic Botany, 43,
31–4.
Zohary, D. 1991. The wild genetic resources of lettuce
(Lactuca sativa L.). Euphytica, 53, 31–5.
Zohary, D. 1992. Is the European plum, Prunus domestica
L., a P. cerasifera Ehrh. x P. spinosa L. allopolyploid?
Euphytica, 60, 75–7.
Zohary, D. 1994. The wild genetic resources of the culti-
vated olive. Acta Horticulturae, 356, 62–5.
Zohary, D. 1995a. The domestication of the grapevine
Vitis vinifera L. in the Near East. In: McGovern, P. T.,
Fleming, S. J. and Katz, S. H. (eds.) The origins and
ancient history of wine. Pp. 45–56. Luxembourg, Gordon
and Breach.
Zohary, D. 1995b. Lentil, Fig, Olive. In: Smartt, J. and
Simmonds, N. W. (eds.) Evolution of crop plants 2nd ed,
Pp. 271–4, 366–70, 379–82. London, Longman.
Zohary, D. 1996. The mode of domestication of the founder
crops of Southwest Asian agriculture. In: Harris, D. R.
(ed.) The origins and spread of agriculture and pastoralism in
Eurasia. Pp. 142–58. London and New York, Routledge.
Zohary, D. 1998. The diffusion of south and east Asian and
of African crops into the belt of Mediterranean agricul-
ture. In: H. D. V. Prendergast, N. L. Etkin, D. R. Harris,
and P. J. Houghton (eds.) Plants for food and Medicine. Pp.
123–34. Kew, UK: Royal Botanic Gardens.
Zohary, D. 1999. Monophyletic vs. polyphyletic origin of
the crops on which agriculture was founded in the
Near East. Genetic Resources and Crop Evolution, 46,
133–42.
Zohary, D. 2002. Domestication of the carob (Ceratonia sili-
qua L.). Israel Journal of Plant Sciences, 50, S141–5.
Zohary, D. and Brick, Z. 1961. Triticum dicoccoides in Israel:
notes on its distribution, ecology and natural hybridiza-
tion. Wheat Information Service, 13, 6–8.
Zohary, D. and Hopf, M. 1973. Domestication of pulses in
the Old World. Science, 182, 887–94.
Zohary, D. and Plitman, U. 1979. Chromosome polymor-
phism, hybridization and colonization in the Vicia sativa
group (Fabaceae). Plant systematics and evolution, 131,
143–56.
Zohary, D. and Spiegel-Roy, P. 1975. Beginnings of fruit
growing in the Old World. Science, 187, 319–27.
Zohary, D., Harlan, J. R. and Vardi, A. 1969. The wild dip-
loid progenitors of wheat and their breeding value.
Euphytica, 18, 58–65.
Zohary, M. 1972. Flora Palaestina. Jerusalem, Israel
Academy of Sciences and Humanities.
Zohary, M. 1973. Geobotanical Foundations of the Middle
East. Stuttgart, G. Fischer.
 Index

Page numbers in italic refer to Figures and Tables, and those in bold to Maps

Aartswoud 189, 195 apple (Malus domestica) 5, 115, 135–8 Azmaška Mogila 180, 195. Colour
Aegilops archaeological remains 136–8 plate 6
cylindrica 50 wild ancestry 135–6. Colour
squarrosa 24, 46–49 plate 14 Bab edh-Dhra 174, 194
tauschii 24, 29, 30–33, 46, 46, 47–49, 50 distribution map 137 Bajč-Medzi kanálmi 186
’Ain Ghazal 2, 194. Colour plate 6 apricot (Armeniaca vulgaris) 7, 116, 144 Balbridie 191, 195. Colour plate 6
Aknashen 176, 194. Colour plate 6 Aratashen 176, 194. Colour plate 6 Balma Margineda 192, 196. Colour
Albertfalva 183 archaeological evidence 9–13, 10, 11 plate 6
Aldenhovener Platte 188, 195. Colour availability of 4–5 banana (Musa) 8
plate 6 charred (carbonized) remains barley (Hordeum vulgare) 21, 52–8, 54
Ali Kosh 2, 169–70, 194. Colour plate 6 10–11 archaeological remains 1, 56–8, 57
Allium chemical tests 13 classification 16–17, 57
ampeloprasum 156, 174 desiccated remains 12 molecular analysis 51, 55–56
cepa (onion) 6, 157 digested remains 12–13 seed dispersal 55
longicuspis 157 impressions 11–12 six-rowed cultivated barley 55
macrochaetum 157 mineralization 12 two-rowed cultivated barley 52–3
oschaninii 157 phytoliths 12 wild progenitor 3, 4, 53–6, 55
porrum (leek) 6, 7, 155–6 preservation by metal oxides 12 distribution 55–6, 57
sativum (garlic) 6, 156–7, 156 south-west Asian Neolithic sites 2 Battonya-Basarága 182, 195. Colour
truncatum 157 waterlogged preservation 12 plate 6
tuncelianum 157 Armeniaca vulgaris (apricot) 7, 144 beet (Beta vulgaris) 7, 159–60
vavilovii 157 Arukhlo 1 and 2 176, 194. Colour Belgium 189
almond (Amygdalus communis) 6, 116, plate 6 Bercy 191
147–9. Colour plate 17 Asikli Höyük 2, 171–2, 194. Colour Berettyóújfalu-Szilhalom 182, 195
archaeological remains 148–9, 149 plate 6 Beta
wild ancestry 147–8 asparagus (Asparagus officinalis) adanensis 159
Alvastra 190, 195 161–2 macrocarpa 159
amplified fragment length Atlit Yam 174 maritima 159
polymorphism (AFLP) 15, 35, Aubechies 189, 195, 196. Colour vulgaris (beet) 7, 159–60
41, 54, 66, 88, plate 6 Bietigheim-Bissingen 188, 195, 196.
Amygdalus aubergine (Solanum melongena) 8 Colour plate 6
bucharica 148 Austria 183–4 bitter orange (Citrus aurantium) 7, 146–7
communis see almond Avena bitter vetch (Vicia envilia) 1–2, 2, 61,
fenzliana 148 abyssinica 66 75–76, 79, 92–5
korschinskyi 147 byzantina 66, 67 archaeological remains 76, 92–5, 94
kuramica 148 fatua 66, 67 wild progenitor 4, 92
persica 144 ludoviciana 66, 67 distribution 93
webbii 148 macrocarpa 67 Bizkaia 192
Anau 176–7 nuda 66, 67 Bjästamon 190. Colour plate 6
Anethum graveolens (dill) 164 occidentalis 66 black cumin (Nigella sativa) 164–5
Anza 179, 195. Colour plate 6 sativa see oat Blatné 186, 195. Colour plate 6
Anzingerberg/Hundssteig 183 sterilis 41, 66, 67, 68. Colour plate 8 Bölcske-Vörösgyír 183
Apium graveolens (celery) 160–1 strigosa 66 Borge Vestre 190, 195. Colour plate 6

237
238 INDEX

bottle gourd (Lagenaria siceraria) 155 caprification 128 Citrus

Bovenkarspel 189 carob (Ceratonia siliqua) 145–6 aurantiifolia 7, 147
Brassica archaeological remains 145–6 aurantium (bitter orange) 7, 146–7
bourgeaui 158 reproductive biology 145 limon (lemon) 7, 146
campestris 112, 159 carrot (Daucus carota) 7, 160 maxima (pumello) 7, 146
cretica 158 Carthamus medica 7, 146
hilarionis 158 flavescens 168 paradisi 147
insularis 158 gypsicola 168 reproductive biology 146
juncea 112 oxyacanthus 168 reticulata 147
macrocarpa 67, 158–9 palaestinus 168 citrus fruits 146–7
napus (swede) 112, 159 persicus 168 Ćmielów 185, 195
nigra 112, 181 tinctorius (safflower) 168 Colocasia esculenta (taro) 8
oleracea (cabbage) 158–9 Castanea sativa (chestnut) 150–1 common millet see broomcorn millet
rapa (turnip/oil-seed rape) 7, 112, Çatalhöyük 171, 194 common oat see oat
159 Caucasia 176 common vetch (Vicia sativa) 95
rupestris 158 Çayönü 2, 171, 194. Colour plate 6 archaeological remains 95
villosa 158 Celei (Sucidava) 181 condiments 163–5
bread wheat (Triticum aestivum) 24, celery (Apium graveolens) 160–1 see also specific plants
29, 47–51 Ceratonia siliqua see carob coprolites 12–13, 108
archaeological remains 50–1 cereals 21–74 coriander (Coriandrum sativum)
club wheat 28, 29, 48 see also specific plants 163–4
cytogenetics 47 Cerro de la Virgen 192, 196 Corylus
free-threshing bread wheats 31, 33, Chateauneuf-les-Martigues 191, 196. avellana (hazelnut) 151
48, 51 Colour plate 6 maxima 151
hulled bread wheats 30, 31–2, 48, cherries (Prunus avium and cottons (Gossypium arboreum and
50–1 P. cerasus) 143–4 G. herbaceum) 7, 100, 107–9
molecular analysis 50 archaeological remains 144 archaeological remains 108–9
spelt wheat 29, 31–2, 48, 50–1 sour cherry 116, 143 genetic analysis 108
wild ancestry 48–50 sweet cherry 115, 143 Cova de Can Sadurní 192, 196.
see also wheats chestnut (Castanea sativa) 150–1 Colour plate 6
Britain 190–1 chickpea (Cicer arietinum) 75, 87–9 Cova de Cendres 192, 196. Colour
broad bean see faba bean archaeological remains 1–2, 2, 76, plate 6
broccoli 158 88–9 Coveta de l’Or 192, 196. Colour
Brødrene Gram 189–90 molecular analysis 88 plate 6
broomcorn (common) millet (Panicum seed dispersal 87 cow pea (Vigna unguiculata) 8, 73
miliaceum) 5, 7, 20, 69–71. wild ancestry 3–4, 87–8 crab apples (Malus sylvestris) 135–7
Colour plate 9 distribution map 88 distribution 137
archaeological remains 69–71, 70 Choga Mami 171, 194 Crasto de Palheiros 193
Brussels sprout (Brassica oleracea) 158 Chokh 176, 194. Colour plate 6 Crete 179
Brześć 185, 195. Colour plate 6 chronological chart 197–9 Crocus
Budapest 183 chufa (Cyperus esculentus) 5, 6, 158 cartwrightianus 165
Bulgaria 179–81 Cicer sativus (saffron) 165
Buraco da Pala 193, 196. Colour plate 6 arietinum see chick pea Csepel-Vízmü 183
Bylany 186, 195. Colour plate 6 bijugum 87 Cuconeştii Vechi 182
echinospermum 87 cucumber (Cucumis sativus) 7, 155
cabbage (Brassica oleracea) 158–9 judaicum 87, 89 Cucumis
Cafer Höyük 2, 194 pinnatifidum 87 callosus 155
Camelina reticulatum 87, 88, 89. Colour plate 10 chate 155
microcarpa 111 Circea (Cârcea) 181, 195. Colour melo see melon
sativa see gold of pleasure plate 6 sativus (cucumber) 7, 155
Can Hasan III 171, 194 citron (Citrus medica) 7, 146 Cueva de La Vaquera 192, 196
Cannabis Citrullus Cueva de los Murciélagos 192, 196.
indica 106 colocynthis 154 Colour plate 6
sativus see hemp ecirrhosus 154 Cueva del Toro 192, 196
Canton of Grisons 187 lanatus see watermelon Cueva de Nerja 192, 196
Canton of Valais 187 rehmii 154 cumin (Cuminum cyminum) 163, 164
Cap Andreas-Kastros 177, 194 vulgaris 153 black cumin 163, 164–5
 INDEX 239

Cydonia emmer and durum-type wheats molecular analysis 101–3

oblonga (quince) 144–5 (Triticum turgidum) 23, 24, 26, wild ancestry 3, 101–3, 102
vulgaris 144–5 27, 29, 40–7. Colour plate 4 distribution map 103
Cyperus esculentus (chufa) 5, 158 archaeological remains 1, 42–7, 43, 45 Forsandmoen 190
Cyprus 177 hulled emmer wheats 41–47, 43, foxtail (Italian) millet (Setaria italica)
cytogenetics 14 45. Colour plate 5 5, 7, 71–2. Colour plate 9
see also specific plants free-threshing wheats 46–7 archaeological remains 70, 71–2
Czech Republic 185–6 characterization of cultivated seed dispersal 71
forms 26, 27, 29, 40, wild progenitor 71
damson 140–141 cytogenetics 41 France 191–2
Dannau 188, 195 grain dispersal 31, 41. Colour Franchthi Cave 177–8, 195. Colour
date palm (Phoenix dactylifera) 5, 114, plate 2, 3 plate 6
116, 131–4 molecular analysis 41, 42 fruit crops 5–6, 114–16
archaeological remains 115, 133–4, wild progenitor 3, 26, 27, 34, 40–2. see also specific species
133 Colour plate 1, 2 Füzesabony-Gubakút 182, 195.
molecular analysis 133 distribution 16, 41–2, 42 Colour plate 6
reproductive biology 131 see also wheats
wild progenitor 131–3 Enkehøj 190 Galabovo 180–1
distribution map 132 enzyme polymorphism 15 garlic (Allium sativum) 6, 156–7, 156
Daucus carota (carrot) 7, 160 Erbaba 172, 194 Germany 188–9
dendrochronology 17–18 germination inhibition 22, 76, 82, 100
Denmark 189–90 faba (broad) bean (Vicia faba) 75, Ghassul see Tuleilat Ghassul
Dévaványa Réhelyi 182, 195 89–92. Colour plate 11 Gilgal 173
Dhali Agridhi 177 archaeological remains 76, 91–2, 91 Gilmosevej 190
dill (Anethum graveolens) 163, 164 characterization of cultivated Girikihaciyan 172, 194
Dimini 178, 195 forms 90 gluten proteins 23
disarticulation scars 1, 22, 26, 27, cytogenetics 90 Gniechowice 185, 195. Colour plate 6
31–2, 53, 54, 116, molecular analysis 90 gold of pleasure (Camelina sativa) 7,
Djade el Mughara 173 wild ancestry 90–1 16, 100, 111
DNA analysis 13–15, 32, 35, Farafra Oasis 174 Gomolava 179, 195
domestication syndrome 9, 13 Fayum 175, 194. Colour plate 6 Gossen 190
cereals 23 Federseeried 188, 195 Gossypium
pulses 75–7 fenugreek (Trigonella foenum-graecum) arboreum see cottons
Dresden-Nickern 188, 195 75, 97–8 barbadense 107
Dürrnberg/Hallein 183–4 archaeological remains 98 herbaceum see cottons
durum wheat see emmer and durum- fibre-producing crops 100–13 hirsutum 107
type wheats (Triticum turgidum) see also specific plants Graetheide 189, 195. Colour plate 6
dye crops 166–8 Ficus grafting 5, 6, 114–115
see also specific plants carica see fig see also specific plants
dyer’s rocket (Reseda luteola) 167 colchica 128 grain size 22
geraniifolia 128 grapefruit (Citrus paradisi) 146, 147
Eberdingen-Hochdorf 188 hyrcanica 128 grapevine (Vitis vinifera) 5, 6, 114,
Egolzwil 3 186, 195, 196. Colour plate 6 johannis 128 116, 121–6
Egypt 174–6 palmata 128 archaeological remains 124–6
einkorn wheat (Triticum pseudosycomorus 128 pip morphology 122–3, 123, 124–5
monococcum) 24, 25, 29, 34–40 sycomorus see sycamore fig reproductive biology 124
archaeological remains 1, 32, virgata 128 sex determination 124
36–40, 37, 38 fig (Ficus carica) 5, 6, 114, 116, 126–30, 127 wild progenitor 121–4, 122
beginnings of domestication 1 archaeological remains 115, 129–30 distribution 122, 123
grain dispersal 31, 35 reproductive biology 126–8 grass pea (Lathyrus sutivus) 2, 95–6
molecular analysis 35 wild ancestry 128–9 archaeological remains 96, 97
wild progenitor 4, 26, 34–6, 33 distribution map 129 cytogenetics 95
distribution 35, 36 Finland 190 Greece 177–9
see also wheats flax (Linum usitatissimum) 1, 2, 61, greengage 141
Eker 190, 195 100, 101–6. Colour plate 12 Grésine 191–2
El Argar 192–3, 196 archaeological remains 2, 103–6, Grotta dell’Uzzo 184, 195. Colour
El Mirón 192, 196. Colour plate 6 104, 105 plate 6
240 INDEX

Grotte G, Baudinard 191 Kastanas 178–9, 195 Linum

Gwoździec 185. Colour plate 6 Khirokitia 177, 194 angustifolium 101
Kissonerga-Mylouthkia 2, 177, 194. bienne 101, 102
Hacilar 171, 194 Colour plate 6 usitatissimum see flax
Hala Sultan Tekke 177 Knossos 179, 195. Colour plate 6 Liubcova 181, 195. Colour plate 6
Hambledon Hill 190, 195. Colour Kobaederra 192, 196. Colour plate 6 Luca Vrublevecaja 182
plate 6 kohlrabi 158 lupins (Lupinus) 98–9
hazelnut (Corylus avellana) 151 Kom el-Hisn 175 albus (white lupin) 98
Heilbronn 188, 195, 196. Colour Kovacevo 180, 195 wild progenitor 98–9, 98
plate 6 Krabbesholm 189, 195. Colour plate 6 angustifolius 98
hemp (Cannabis sativa) 7, 100, 106–7 Kückhoven 188, 195. Colour plate 6 graecus 99
Hienheim 188, 195. Colour plate 6 Kukoneshti 195 luteus 98
Hierakonpolis 175 Kumtepe 172 termis 98
Hilzingen 188, 195, 196. Colour Kvåle 190
plate 6 Maadi 174–5
Hjelle 190. Colour plate 6 La Baume Fontbrégoua 191 Maastricht-Randwijck 189
Hordeum Lablab purpureus (hyacinth bean) 73 madder (Rubia tinctorum) 166, 167–8
agriocrithon 53, 57 ’La Bosse de la Tombe’ à Givry 189 Majaki 182
distichum 52, 53, 55, 57 Lactuca Majdaneckoe 182
hexastichum 52, 57 sativa (lettuce) 6, 7, 157–8 Malus
spontaneum 41, 53–9, 54, 55, 56. serriola 157 domestica see apple
Colour Plate 157 Lagenaria siceraria (bottle gourd) 155 kirghizorum 136
tetrastichum 52 Lake Biel 187 orientalis 136. Colour plate 14a
vulgare see barley Lake Neuchatel 187 prunifolia 136
Hornstaad 188–9 La Lámpara 192, 196 pumila see apple
horse bean see faba bean La Marmotta 185, 195. Colour plate 6 sieversii 136. Colour plate 14b
horticulture 5–6, 114–17 Lathyrus sylvestris 136. Colour plate 14a
see also specific plants cicera 95–6 turkmenorum 136
Hungary 182–3 clymenum (Spanish vechling) 97 mandarin (Citrus reticulata) 146, 147
hyacinth bean (Lablab purpureus) 73 sativus see grass pea marrowstem kale 158
leek (Allium portum) 6, 7, 155–6 Melissa officinalis (lemon balm) 163
Imiris-Gora 176, 194 legumes 75–7 melon (Cucumis melo) 6, 154–5
Indigofera 166–168 see also specific plants archaeological remains 155
arrecta 168 lemon balm (Melissa officinalis) 163 Menneville and Aisne Valley
tinctoria (indigo) 7, 166, 168 lemon (Citrus limon) 7, 146–7 sites 191, 195, 196. Colour
Indigo (Indigofera tinctoria) 7, 166–168 Lens plate 6
Iran 169–70 culinaris see lentil Merimde 175, 194. Colour plate 6
Iraq 170–1 ervoides 77 microsatellites (SSR) 15, 84, 119
Ireland 190–1 lamottei 77 migrants
Isatis tinctoria (woad) 166–7 nigricans 77 temperate crops from central and/
Israel 173–4 odemensis 78 or east Asia 7
Italian millet see foxtail millet orientalis 77–9, 78, 80 warm-weather crops from south
Italy 184–5 tomentosus 77, 78 and/or east Asia 7–8
lentil (Lens culinaris) 75, 77–82 warm-weather crops from
Jarmo 2, 170, 194. Colour plate 6 archaeological remains 1–2, 76, sub-Saharan Africa 8
Jeitun 176. Colour plate 6 79–82, 80 millets see broomcorn millet; foxtail
Jerf el Ahmar 173, 194 cytogenetics 77–9 millet
Jericho 2, 174, 194. Colour plate 6 identification of cultivated forms 81 mineralization 12
Jerma 176 molecular analysis 78 Mohelnice 185–6, 195. Colour plate 6
Jordan 173–4 seed dispersal 79 Moldavia 181–2
Juglans regia (walnut) 149–50 wild progenitor 4, 77–9, 79 molecular biology tools 14–15
distribution 78 recessive mutations 61
Kaf Taht el-Ghar 176, 196. Colour Lerna 178, 195 see also specific plants
plate 6 lettuce (Lactuca sativa) 6, 7, 153, 157–8 Mondsee 183, 195
kale 158–9 Libya 176 Monte Còvolo 184–5, 195
Kapitan Dimitrievo 180, 195. Colour lime (Citrus aurantiifolia) 7, 146–7 Morocco 176
plate 6 Lindebjerg 190, 195 Musa (banana and plantain) 8
Karanovo 179–80, 195. Colour plate 6 linseed see flax mustard 112
 INDEX 241

Nabta Playa 174 parsley 163 Pont de Roque Haute 191, 196.
Nahal Zehora II 173 parsnip (Pastinaca sativa) 153, 161 Colour plate 6
Naqada 175 parthenocarpy 116, 127, 130 poppy (Papaver somniferum) 5,
Nea Nikomedeia 178, 195. Colour peach (Persica vulgaris) 7, 116, 144–5 109–11, 110
plate 6 pea (Pisum sativum) 75, 82–7, 84 archaeological remains 109–11
Neolithic south-west Asian crop archaeological remains 1, 2, 76, wild poppy distribution map 110
assemblage 1–3, 3 84–7, 85 Portugal 193
spread 3, 4. Colour plate 6 cytogenetics 82, 83–4 Pre-Pottery Neolithic B (PPNB) 1,
Netherlands 189 molecular analysis 84 2–3, 42, 43
Netiv Hagdud 173, 194 wild ancestry 4, 82–4 preservation of plant remains 9–13, 11
Nigella sativa (black cumin) 164–5 distribution map 83 carbonization 10–11
Nitriansky Hrádok 186, 195 morphological types 82–3 desiccation 12
nitrogen fixation 75, 97 pearl millet (Pennisetum glaucum) digested remains 12–13
Niuskala 190, 195. Colour plate 6 8, 72, 73 impressions 11–12
Norsminde 189, 195. Colour plate 6 pear (Pyrus communis) 5, 115, 138–40 metal oxides 12
Norway 190 archaeological remains 140 mineralization 12
Novye Rusešty 182 wild ancestry 138–40 waterlogging 12
Nowa Huta-Mogila 185, 195 distribution map 139 protein variation 15
Pennisetum glaucum see pearl millet Prunus
oat (Avena sativa) 7, 16, 21, 66–9 Persica vulgaris (peach) 7, 116, amygdalus 147
archaeological remains 69, 70 144–5 armeniaca 144
cytogenetics 66 Phoenix avium see cherries
molecular analysis 66 dactylifera see date palm brigantina 142
seed dispersal 67–8 reclinata 133 caspica 142
wild ancestry 66–7, 68 sylvestris 133 cerasifera 141–2
Obre 179, 195. Colour plate 6 theophrastii 133 cerasus see cherries
Ohalo II 173, 194 Pienza 184, 195. Colour plate 6 cocomilia 142
oil-producing crops 100 pistachio (Pistacia vera) 7, 115, 116, divaricata 141
oil-seed rape (Brassica rapa) 112, 159 151–2 domestica see plum
Olea Pistacia dulcis 147
europaea see olive atlantica 151 fruticosa 143
oleaster 116, 117–18. Colour plate 13 khinjuk 151 insititia 141
olive (Olea europaea) 5, 6, 114, 116–21. palaestina 151 persica 144
Colour plate 13 terebinthus 151 salicina 141
archaeological remains 115, 118, vera (pistachio) 7, 115, 116, 151–2 sogdiana 142
119–21 Pisum spinosa 141, 142
grafting 117–18 elatius 82–3, 84 ursina 142
molecular analysis 119 fulvum 82 pulses 75–7
vegetative propagation 117 humile 83, 84 see also specific plants
wild progenitor 117–19. Colour sativum see pea pummelo (Citrus maxima) 7, 146, 147
plate 13 syriacum 83 Punica granatum see pomegranate
distribution 117–19, 119 plantain (Musa) 8 Putineşti 182
onion (Allium cepa) 6, 157 plum (Prunus domesticus) 5, 115, 116, Pyrus
orange 146, 147 140–3 amygdaliformis 140
Seville or bitter orange (Citrus archaeological remains 142–3, bretschneideri 140
aurantium) 7, 146–7 143 caucasica 139, 140
Oryza cytogenetics 141, 142 communis see pear
nivara 74 wild ancestry 141–2 domestica 138
rufipogon 74 Poduri 181, 195. Colour plate 6 elaeagrifolia 140
sativa see rice Pokrovnik 179, 195 korshinskyi 140
Overhespen 189. Colour plate 6 Poland 185 pyraster 139, 140
Polish wheat 29, 39 pyrifolia 140
Panicum polymerase chain reaction (PCR) 15 salicifolia 140
milliaceum see broomcorn millet polyploidy 14 spinosa 139–40
spontaneum 69 see also specific plants syriaca 140. Colour plate 15, 16
Papaver pomegranate (Punica granatum) 6, ussuriensis 140
setigerum 109, 110, 111 115, 134–5
somniferum see poppy archaeological remains 134–5 quince (Cydonia vulgaris) 144, 145
242 INDEX

radiocarbon dating 17–19, 18, 19 vavilovii 59, 61, 62 archaeological remains 130–1
radish (Raphanus sativus) 112 secondary crops 7 reproductive biology 130
Rákoskeresztúr-Újmajor 182 seed dispersal Syria 172–3
random amplified polymorphic DNA cereals 22–3 Szeged-Gyálarét 182, 195. Colour
(RAPD) 15, 66, 101, 119 barley 53 plate 6
recessive mutations 60 foxtail millet 71 Szilhalom 195
Reseda luteola (dyer’s rocket) 167 oats 66–8
restriction fragment length rye 60 Tankardstown 191, 195. Colour plate 6
polymorphism (RFLP) 15, 41 wheats 30–1, 35, 40 taro (Colocasia esculenta) 8
Revilla del Campo 192, 196 pulses 76 Tell Abu Hureyra 2, 172, 194. Colour
rice (Oryza sativa) 5, 7, 21, 73–4 chickpea 87 plate 6
Rivne 182. Colour plate 6 lentil 79 Tell Aswad 2, 173, 194. Colour plate 6
Rosdorf 189, 195. Colour plate 6 flax 106 Tell es-Sa’idiyeh 174
Röszke-Lúdvár 182, 195. Colour self-pollination 20–1 Tell es-Sawwan 170, 194
plate 6 sesame (Sesamum indicum) 7, 112–13 Tell Hârşova 181
Rovno see Rivne archaeological records 113 Tell Mureybit 172–3, 194
Rubia tinctorum (madder) 167–8 Sesamum Tell Ramad 194
Rumania 181 indicum see sesame Tepe Hasanlu 170
Ruseştii Noi 182 orientale 112 Tepe Sabz 170, 194
rush nut see chufa Sesklo 178, 195. Colour plate 6 Tepe Yahya 170
rye (Secale cereale) 7, 59–65, 63 Setaria The Stumble 191, 195. Colour plate 6
archaeological remains 2, 37, italica see foxtail millet Timopheev’s wheat (Triticum
64–5, 64 viridis 71 timopheevi) 24, 29, 42, 51
molecular analysis 59 Seville or bitter orange (Citrus cultivated 24, 51
wild ancestry 59–62 aurantium) 7, 146–7 cytogenetics 51
distribution map 62 Shillourokambos 2, 177, 194 wild 41, 51
Shiqmim 174 see also wheats
Sacarovca 181–2, 195. Colour plate 6 Shortugai 177 Toos-Waldi 187
Saccharum officinarum (sugar Sion 187, 195. Colour plate 6 Toumba Balomenou 178, 195
cane) 7–8 simple sequence repeat (SSR) 15, 84, 119 Transcaucasia 176
safflower (Carthamus tinctorius) 166, Sinapis alba (white mustard) 112 Trigonella foenum-graecum
168 single nucleotide polymorphisms see fenugreek
saffron (Crocus sativus) 165 (SNPs) 15 Triticum
Sammardenchia 184, 195. Colour sloe (Prunus spinosa) 141, 142 aegilopoides 39, 33, 34, 35
plate 6 Slovakia 185–6 aestivum see bread wheat
San Marco 184 Solanum melongena (aubergine) 8 araraticum 29, 42, 51
Saqqara 175, 194 sorghum (Sorghum bicolor) 8, 20, 72–3 baeoticum see einkorn wheat
Šarišské Michalˇany-Fedelemka 186 south-west Asian crop assemblage 1–4 carthlicum 29, 40
Sarup 189, 195 spread of 3, 4. Colour plate 6 compactum 29, 48
Scamuso 184, 195. Colour plate 6 Spain 192–3 dicoccoides see emmer and
Scania 2, 190, 195. Colour plate 6 Spanish vechling (Lathyrus durum-type wheats
Schletz 183, 195. Colour plate 6 clymenum) 97 dicoccum see emmer and
Secale spelt wheat, see bread wheat durum-type wheats
afghanicum 59 Starčevo 179, 195 durum see emmer and durum-type
anatolicum 61 Starye Kukoneshti 182 wheats
ancestrale 59, 60, 63 Stary Zamek 185, 195 macha 29, 47
cereale see rye Station III de Clairvaux 191 monococcum see einkorn wheat
ciliatoglume 61 Stillfried 183 parvicoccum 29, 40
dalmaticum 61 Strachów 185, 195. Colour plate 6 polonicum 29, 40
dighoricum 59 Sucidava-Celei 181, 195 sativum 29, 48
iranicum 60, 61 sugar cane (Saccharum officinarum) 7 sinskajae 29, 34
kupriyanowii 61 swede (Brassica napus) 112, 159 spelta see bread wheat
montanum 61, 63–4, 66. Colour Sweden 190 sphaerococcum 29, 48
plate 7 Swifterbant S3 189, 195. Colour plate 6 thaoudar 29, 35
segetale 59 Switzerland 186–7 timopheevi see timopheev’s wheat
strictum 61 sycamore fig (Ficus sycomorus) 115, turgidum see emmer and
sylvestre 60, 62 116, 130–1 durum-type wheats

urartu 23–4, 29, 36, 41 narbonensis 90, 91, 92
vulgare 29, 48 sativa see common vetch
Troy 172 serratifolia 90
Tuleilat Ghassul 174, 194 Vigna unguiculata see cow pea
Turkey 171–2 Visborg 189, 195. Colour plate 6
turnip (Brassica rapa) 7, 112, 153, 159 viticulture see grapevine
Tutankhamun tomb 175–6 Vitis
Twisk 189, 195 sylvestris 121, 122, 123
vinifera see grape vine
Ukraine 181–2 Voldtofte 190
Usatovo 182 Vršnik 179, 195
vegetables 6–7, 153 walnut (Juglans regia) 149–50
see also specific plants Wange 189, 195. Colour plate 6
vegetative propagation 114–15 watermelon (Citrullus lanatus) 6,
carob 145 153–4
fig 126–7 archaeological remains 154
grapevine 124 weeds 7, 10, 16
olive 117 oil plants 100
vetch see bitter vetch; common vetch wheats (Triticum) 20, 23–32
Vicia beginnings of domestication 1
ervilia see bitter vetch classification 16, 23–4, 29
faba see faba bean cytogenetics 23–4
galilea 90, 91 free-threshing wheats 24–32, 27,
hyaenis-ciamus 90 28, 31, 32
johannis 90 tetraploid vs hexaploid
kalakhensis 90 free-threshing wheats 32, 32
INDEX 243
hulled wheats 24–30, 26, 30, 31
wild progenitors 3–4, 13–16, 33
see also bread wheat; einkorn wheat;
emmer and durum-type
wheats; Timopheev’s wheat
white lupin (Lupinus albus) 98
wild progenitor 98–9, 98
wild progenitors 3–4
geographic distribution 3–4,
15–16
identification 13–15
classical taxonomic methods 14
cytogenetic analysis 14
molecular analysis 14–15
see also specific plants
woad (Isatis tinctoria) 166–7
Yarym Tepe 170–1, 194
Yiftah’el 2. Colour plate 6
Yugoslavia (former) 179
Yunatzite 181
Zambujal 193, 196
Zánka-Vasútállomás 182, 195. Colour
plate 6
Zinchecra 176. Colour plate 6
Zürich 186–7, 195, 196. Colour plate 6
This page intentionally left blank
Plate 1 Stands of wild emmer wheat, Triticum dicoccoides (upper), and wild barley, Hordeum spontaneum (lower)—the progenitors of
domesticated emmer wheat and domesticated barley (photographs kindly provided by O. Fragman-Sapir, Jerusalem Botanical Garden).
Plate 2 A spikelet of wild tetraploid emmer wheat, T. turgidum Plate 3 A spikelet of domesticated, non-shattering, tetraploid emmer
subsp dicoccoides. Note the smooth disarticulation scars, both wheat, T. turgidum subsp. dicoccum. Note the rough disarticulation scars,
below and above the internode. These smooth scars are the both below and above the internode. These rough scars are the most
diagnostic elements for the identification of the wild forms. reliable diagnostic elements for the identification of the domesticated forms.
 Plate 4 An ear of domesticated, non-shattering, tetraploid emmer wheat, T. turgidum subsp.
dicoccum.

Plate 5 A–Remains of emmer wheat ears retrieved from Saqqara, Lower Egypt. B–Details of two individual ears (courtesy of Agropolis-Museum,
Montpellier, France).
 60 Legened
61
einkorn wheat
emmer wheat
58 barley
flax
71 62
lentil
pea
2,500-2,000 BP
63 3,000-2,500 BP
3,500-3,000 BP
72 59 4,000-3,500 BP
4,500-4,000 BP
5,000-4,500 BP
5,500-5,000 BP
70 66 6,000-5,500 BP
69 6,500-6,000 BP
65 40 7,000-6,500 BP
56 53
68 7,500-7,000 BP
67 57 38 8,000-7,500 BP
39 21 8,500-8,000 BP
52,55 9,000-8,500 BP
73 35 37 9,500-9,000 BP
41
54 36 10,000-9,500 BP
51 10,500-10,000 BP
48 49 47
33 22
50 34 1
44 23
81 32 4
82
64 74
75 25
78 6
31 24
77 42 29
28 5
43
27 7
79 30 26
45
76 18

80 19 8
3
46
17 9 2

20 14
10
13
11

12

16

Scale 1:16,000,000 15
0 125 250 375 500 miles

0 250 500 km
83

Plate 6 (Map 2) The spread of the south-west Asian Neolithic crop assemblage in Europe, west Asia, and North Africa. For details on the numbered sites, see Appendix C (p. 200). These are the
earliest sites in which domesticated grain crops were found, in each country.
Plate 7 Unripe ear of wild perennial rye, Secale montanum Plate 8 A seed dispersal unit of wild oat, Avena sativa subsp.
(photograph kindly provided by O. Fragman-Sapir, Jerusalem Botanical sterilis. Note the smooth disarticulation scar in the lower tip of the
Garden). spikelet, and the drill-like awans, which helps in the insertion of the
spikelet into the ground.

Plate 9 Millet crops of southwest Asia. A and B–Broomcorn millet [= common

millet], Panicum miliaceum; A–Fruiting panicle; B–Ripe spikelets. C and D–Foxtail
millet [= Italian millet], Setaria italica; C–Fruiting panicle; D–Ripe spikelets (after
Vaughan and Geissler 2009; with kind permission of Oxford University Press).
Plate 10 Fruiting branch of wild chickpea, Cicer arietinum subsp. Plate 11 Faba bean, Vicia faba. A–plant, B–flowers, C–opened pod,
reticulatum. D–seed (after Vaughan and Geissler 2009; with kind permission of
Oxford University Press).

Plate 12 Flax stems, heckled flax line, and spun linen thread, the work of Alverna Messersmith Learn, early 1900s (with kind permission of Linda
Learn, Class Act Fabrics).
Plate 13 Fruiting branches and stones of wild olive, Olea europaea subsp. oleaster (left), and domesticated olive, Olea europaea subsp.
europaea (right).

Plate 14A Fruiting branches of wild apple, Malus sylvestris subsp. Plate 14B Flowering branches of wild apple, Malus sieversii
orientalis (photograph kindly provided by O. Fragman-Sapir, Jerusalem (photograph kindly provided by O. Fragman-Sapir, Jerusalem
Botanical Garden). Botanical Garden).
Plate 15 Flowering branch of wild pear, Pyrus syriaca. Plate 16 Wild pear, Pyrus syriaca: cluster of maturing fruits
(photograph kindly provided by O. Fragman-Sapir, Jerusalem Botanical
Garden).

Plate 17 Almond, Amygdalus communis: cluster of flowers (photograph kindly provided by O. Fragman-Sapir, Jerusalem Botanical Garden).