Plant Domestication and the Origins of Agriculture in the Ancient Near East [New ed.] 1108493645, 9781108493642

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P LA N T DO M E S T I C A T I O N A N D T H E O R I G I N S O F A G R I C U LT U R E I N TH E A N CI E NT N E A R E A S T The Agricultural Revolution – including the domestication of plants and animals in the Near East – that occurred 10,500 years ago ended millions of years of human existence in small, mobile, egalitarian communities of hunter-gatherers. This Neolithic transformation led to the formation of sedentary communities that produced crops such as wheat, barley, peas, lentils, chickpeas and flax and domesticated a range of livestock, including goats, sheep, cattle and pigs. All of these plants and animals still play a major role in the contemporary global economy and nutrition. This agricultural revolution also stimulated the later development of the first urban centres. This volume examines the origins of plant domestication in the ancient Near East, along with various aspects of the new Human–Nature relationship that characterizes food-producing societies. It demonstrates how the rapid, geographically localized, knowledge-based domestication of plants was a human initiative that eventually gave rise to Western civilizations and the modern human condition. Shahal Abbo is an agronomist and plant geneticist at the Hebrew University of Jerusalem, Israel. Through comparative study of grain legumes and cereals, both domesticated and wild, across Mediterranean agro-eco-systems, he has developed several new practical and conceptual tools pertaining to plant domestication and crop evolution. Avi Gopher is an archaeologist at Tel Aviv University, Israel. He has conducted research on time-space systematics – seriation analyses reconstructing both the chronology and pace of the diffusion of Neolithic cultural elements in the interaction sphere of the early Neolithic in the Near East. Gopher is a member of a research group on plant domestication in the Near East and focuses on the archaeological aspects. Gila Kahila Bar-Gal is a molecular geneticist at the Hebrew University of Jerusalem, Israel. She studies host–pathogen interaction and human activities that affect animals with the aim of conserving future biodiversity.

Published online by Cambridge University Press

Published online by Cambridge University Press

PLANT DOMESTICATION AND THE ORIGINS OF AGRICULTURE IN THE A N C I E N T NE A R E A S T Shahal Abbo and Avi Gopher With a contribution by Gila Kahila Bar-Gal Translated by Halo (Hilla) Ben Asher

Published online by Cambridge University Press

University Printing House, Cambridge CB2 8BS, United Kingdom One Liberty Plaza, 20th Floor, New York, NY 10006, USA 477 Williamstown Road, Port Melbourne, VIC 3207, Australia 314–321, 3rd Floor, Plot 3, Splendor Forum, Jasola District Centre, New Delhi – 110025, India 103 Penang Road, #05–06/07, Visioncrest Commercial, Singapore 238467 Cambridge University Press is part of the University of Cambridge. It furthers the University’s mission by disseminating knowledge in the pursuit of education, learning, and research at the highest international levels of excellence. www.cambridge.org Information on this title: www.cambridge.org/9781108493642 DOI: 10.1017/9781108642491 © Cambridge University Press 2022 This publication is in copyright. Subject to statutory exception and to the provisions of relevant collective licensing agreements, no reproduction of any part may take place without the written permission of Cambridge University Press. First published 2022 Printed in the United Kingdom by TJ Books Limited, Padstow Cornwall A catalogue record for this publication is available from the British Library. Library of Congress Cataloging-in-Publication Data Names: Abbo, Shahal, author. | Gopher, Avi, author. | Bar-Gal, Gila Kahila, contributor. | Ben Asher, Halo, translator. Title: Plant Domestication and the Origins of Agriculture in the Ancient Near East / Shahal Abbo and Avi Gopher, with a contribution from Gila Kahila Bar-Gal ; English translation: Halo (Hilla) Ben Asher. Description: First edition. | New York, NY : Cambridge University Press, 2022. | Includes bibliographical references and index. Identifiers: LCCN 2021036430 (print) | LCCN 2021036431 (ebook) | ISBN 9781108493642 (hardback) | ISBN 9781108737708 (paperback) | ISBN 9781108642491 (epub) Subjects: LCSH: Ethnobotany–Middle East. | Agriculture, Prehistoric–Middle East. Classification: LCC GN476.73 .A22 2022 (print) | LCC GN476.73 (ebook) | DDC 630.9394–dc23 LC record available at https://lccn.loc.gov/2021036430 LC ebook record available at https://lccn.loc.gov/2021036431 ISBN 978-1-108-49364-2 Hardback Cambridge University Press has no responsibility for the persistence or accuracy of URLs for external or third-party internet websites referred to in this publication and does not guarantee that any content on such websites is, or will remain, accurate or appropriate.

Published online by Cambridge University Press

P LA N T DO M E S T I C A T I O N A N D T H E O R I G I N S O F A G R I C U LT U R E I N TH E A N CI E NT N E A R E A S T The Agricultural Revolution – including the domestication of plants and animals in the Near East – that occurred 10,500 years ago ended millions of years of human existence in small, mobile, egalitarian communities of hunter-gatherers. This Neolithic transformation led to the formation of sedentary communities that produced crops such as wheat, barley, peas, lentils, chickpeas and flax and domesticated a range of livestock, including goats, sheep, cattle and pigs. All of these plants and animals still play a major role in the contemporary global economy and nutrition. This agricultural revolution also stimulated the later development of the first urban centres. This volume examines the origins of plant domestication in the ancient Near East, along with various aspects of the new Human–Nature relationship that characterizes food-producing societies. It demonstrates how the rapid, geographically localized, knowledge-based domestication of plants was a human initiative that eventually gave rise to Western civilizations and the modern human condition. Shahal Abbo is an agronomist and plant geneticist at the Hebrew University of Jerusalem, Israel. Through comparative study of grain legumes and cereals, both domesticated and wild, across Mediterranean agro-eco-systems, he has developed several new practical and conceptual tools pertaining to plant domestication and crop evolution. Avi Gopher is an archaeologist at Tel Aviv University, Israel. He has conducted research on time-space systematics – seriation analyses reconstructing both the chronology and pace of the diffusion of Neolithic cultural elements in the interaction sphere of the early Neolithic in the Near East. Gopher is a member of a research group on plant domestication in the Near East and focuses on the archaeological aspects. Gila Kahila Bar-Gal is a molecular geneticist at the Hebrew University of Jerusalem, Israel. She studies host–pathogen interaction and human activities that affect animals with the aim of conserving future biodiversity.

Published online by Cambridge University Press

Published online by Cambridge University Press

PLANT DOMESTICATION AND THE ORIGINS OF AGRICULTURE IN THE A N C I E N T NE A R E A S T Shahal Abbo and Avi Gopher With a contribution by Gila Kahila Bar-Gal Translated by Halo (Hilla) Ben Asher

Published online by Cambridge University Press

University Printing House, Cambridge CB2 8BS, United Kingdom One Liberty Plaza, 20th Floor, New York, NY 10006, USA 477 Williamstown Road, Port Melbourne, VIC 3207, Australia 314–321, 3rd Floor, Plot 3, Splendor Forum, Jasola District Centre, New Delhi – 110025, India 103 Penang Road, #05–06/07, Visioncrest Commercial, Singapore 238467 Cambridge University Press is part of the University of Cambridge. It furthers the University’s mission by disseminating knowledge in the pursuit of education, learning, and research at the highest international levels of excellence. www.cambridge.org Information on this title: www.cambridge.org/9781108493642 DOI: 10.1017/9781108642491 © Cambridge University Press 2022 This publication is in copyright. Subject to statutory exception and to the provisions of relevant collective licensing agreements, no reproduction of any part may take place without the written permission of Cambridge University Press. First published 2022 Printed in the United Kingdom by TJ Books Limited, Padstow Cornwall A catalogue record for this publication is available from the British Library. Library of Congress Cataloging-in-Publication Data Names: Abbo, Shahal, author. | Gopher, Avi, author. | Bar-Gal, Gila Kahila, contributor. | Ben Asher, Halo, translator. Title: Plant Domestication and the Origins of Agriculture in the Ancient Near East / Shahal Abbo and Avi Gopher, with a contribution from Gila Kahila Bar-Gal ; English translation: Halo (Hilla) Ben Asher. Description: First edition. | New York, NY : Cambridge University Press, 2022. | Includes bibliographical references and index. Identifiers: LCCN 2021036430 (print) | LCCN 2021036431 (ebook) | ISBN 9781108493642 (hardback) | ISBN 9781108737708 (paperback) | ISBN 9781108642491 (epub) Subjects: LCSH: Ethnobotany–Middle East. | Agriculture, Prehistoric–Middle East. Classification: LCC GN476.73 .A22 2022 (print) | LCC GN476.73 (ebook) | DDC 630.9394–dc23 LC record available at https://lccn.loc.gov/2021036430 LC ebook record available at https://lccn.loc.gov/2021036431 ISBN 978-1-108-49364-2 Hardback Cambridge University Press has no responsibility for the persistence or accuracy of URLs for external or third-party internet websites referred to in this publication and does not guarantee that any content on such websites is, or will remain, accurate or appropriate.

Published online by Cambridge University Press

P LA N T DO M E S T I C A T I O N A N D T H E O R I G I N S O F A G R I C U LT U R E I N TH E A N CI E NT N E A R E A S T The Agricultural Revolution – including the domestication of plants and animals in the Near East – that occurred 10,500 years ago ended millions of years of human existence in small, mobile, egalitarian communities of hunter-gatherers. This Neolithic transformation led to the formation of sedentary communities that produced crops such as wheat, barley, peas, lentils, chickpeas and flax and domesticated a range of livestock, including goats, sheep, cattle and pigs. All of these plants and animals still play a major role in the contemporary global economy and nutrition. This agricultural revolution also stimulated the later development of the first urban centres. This volume examines the origins of plant domestication in the ancient Near East, along with various aspects of the new Human–Nature relationship that characterizes food-producing societies. It demonstrates how the rapid, geographically localized, knowledge-based domestication of plants was a human initiative that eventually gave rise to Western civilizations and the modern human condition. Shahal Abbo is an agronomist and plant geneticist at the Hebrew University of Jerusalem, Israel. Through comparative study of grain legumes and cereals, both domesticated and wild, across Mediterranean agro-eco-systems, he has developed several new practical and conceptual tools pertaining to plant domestication and crop evolution. Avi Gopher is an archaeologist at Tel Aviv University, Israel. He has conducted research on time-space systematics – seriation analyses reconstructing both the chronology and pace of the diffusion of Neolithic cultural elements in the interaction sphere of the early Neolithic in the Near East. Gopher is a member of a research group on plant domestication in the Near East and focuses on the archaeological aspects. Gila Kahila Bar-Gal is a molecular geneticist at the Hebrew University of Jerusalem, Israel. She studies host–pathogen interaction and human activities that affect animals with the aim of conserving future biodiversity.

Published online by Cambridge University Press

Published online by Cambridge University Press

PLANT DOMESTICATION AND THE ORIGINS OF AGRICULTURE IN THE A N C I E N T NE A R E A S T Shahal Abbo and Avi Gopher With a contribution by Gila Kahila Bar-Gal Translated by Halo (Hilla) Ben Asher

Published online by Cambridge University Press

University Printing House, Cambridge CB2 8BS, United Kingdom One Liberty Plaza, 20th Floor, New York, NY 10006, USA 477 Williamstown Road, Port Melbourne, VIC 3207, Australia 314–321, 3rd Floor, Plot 3, Splendor Forum, Jasola District Centre, New Delhi – 110025, India 103 Penang Road, #05–06/07, Visioncrest Commercial, Singapore 238467 Cambridge University Press is part of the University of Cambridge. It furthers the University’s mission by disseminating knowledge in the pursuit of education, learning, and research at the highest international levels of excellence. www.cambridge.org Information on this title: www.cambridge.org/9781108493642 DOI: 10.1017/9781108642491 © Cambridge University Press 2022 This publication is in copyright. Subject to statutory exception and to the provisions of relevant collective licensing agreements, no reproduction of any part may take place without the written permission of Cambridge University Press. First published 2022 Printed in the United Kingdom by TJ Books Limited, Padstow Cornwall A catalogue record for this publication is available from the British Library. Library of Congress Cataloging-in-Publication Data Names: Abbo, Shahal, author. | Gopher, Avi, author. | Bar-Gal, Gila Kahila, contributor. | Ben Asher, Halo, translator. Title: Plant Domestication and the Origins of Agriculture in the Ancient Near East / Shahal Abbo and Avi Gopher, with a contribution from Gila Kahila Bar-Gal ; English translation: Halo (Hilla) Ben Asher. Description: First edition. | New York, NY : Cambridge University Press, 2022. | Includes bibliographical references and index. Identifiers: LCCN 2021036430 (print) | LCCN 2021036431 (ebook) | ISBN 9781108493642 (hardback) | ISBN 9781108737708 (paperback) | ISBN 9781108642491 (epub) Subjects: LCSH: Ethnobotany–Middle East. | Agriculture, Prehistoric–Middle East. Classification: LCC GN476.73 .A22 2022 (print) | LCC GN476.73 (ebook) | DDC 630.9394–dc23 LC record available at https://lccn.loc.gov/2021036430 LC ebook record available at https://lccn.loc.gov/2021036431 ISBN 978-1-108-49364-2 Hardback Cambridge University Press has no responsibility for the persistence or accuracy of URLs for external or third-party internet websites referred to in this publication and does not guarantee that any content on such websites is, or will remain, accurate or appropriate.

Published online by Cambridge University Press

CONTENTS

List of Tables · vii Foreword · viii by Simcha Lev-Yadun Foreword · xi by Paul Gepts

Preface and Acknowledgements · xv IN T RO D U CT IO N

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1

W HAT I S T HE AGRIC ULTURAL REVO LUTIO N?

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F R O M HU N T E R - G A T H E R E R S T O F A R M E R S I N TH E N E A R E A S T : A R C H A E O L O G I C A L B A C K G R O U N D · 21

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M O DELS T HAT D ESC R IBE A ND EXP L AIN T HE AG RI CU LTU RAL R E VO L U T I O N , I N C L U D I N G P L A N T D O M E S T I C A T I O N · 80

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THE PLANT FORMATIONS OF THE FERTILE CRESCENT AND THE WILD P R O G E N I T O R S OF TH E D O M E S TI C A T E D F O U N D E R C R O P S · 100

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T H E D I F F E R EN C E B E T W E E N W I LD A N D D O M E S T I C A T E D P LA N T S

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T R A D I T I O N A L VE R S U S M O D E R N A G R I C U L T U R E – STAB ILITY V S M A X I M I Z A T I O N · 122

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T H E D I F F E R E N C E S BE T W E E N P L A N T DO M E S T I C A T I O N A N D C R O P EVOLU TI ON U NDER TRADITI ON AL AND M OD ERN F A R M I N G S Y S T E MS · 128

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T H E D I F F E R E N C E S BE T W E E N C E R E A L A N D L E G U M E CR O P S I N T H E N E A R E A S T · 135

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T H E C H O I C E O F P L A N T S P E C I E S F OR DO M E S T I C A T I O N : A G R O N O M I C AN D D IETARY C ON S IDERATIO NS · 147

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W H E R E , W H E N A N D HO W D I D N E A R E A S T E R N P L A N T D O M E S T I C A T I O N OC C U R ? · 156

CONTENTS

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D O M E S T I C A T I O N OF F R U I T T R E E S I N T H E N E A R E A S T

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P LA N T E V O L U T I O N U N D E R DO M E S T I C A T I O N

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A GL O B A L V I E W O F PL A N T D O ME S T I C A T I O N I N OT H E R W O R L D R E G I O N S : A S I A , A F R I C A A N D A M E R I C A · 188

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A N I M A L D O M E S T I C A T I O N I N TH E N E A R E A S T by Gila Kahila Bar-Gal

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P LA N T DO M E S T I C A T I O N A N D E A R L Y N E A R E A S T E R N A G R I C U L T U R E : S U M M A R Y A N D CO N C L U S I O N S · 229

Notes · 239 Further Reading · 242 References, Chapter 14 · 247 Glossary · 254 Index · 265

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CONTENTS

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166

· 179

· 199

TABLES

4.1 5.1 6.1 8.1 9.1 11.1 11.2 14.1 14.2 14.3

Founder crops of Near Eastern agriculture and their wild progenitors · 106 Summary of main fitness characteristics in wild and domesticated plants · 117 Differences between traditional farming and modern agriculture · 124 Summary of biological differences between Near Eastern legumes and cereals (wild and domesticated) · 136 Grain yield obtained from foraging wild cereals and legumes · 149 Sexual versus asexual plant reproduction · 168 Main differences between fruit trees and grain crops domesticated in the Near East · 173 Domesticated animal species in the Near East · 201 The ‘Big Four’ – characteristics considered favourable for animal domestication · 208 The feeding behaviour of the ‘Big Four’ Near Eastern livestock · 212

LIST OF TABLES

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FO REWOR D

It is with much pleasure that I take the opportunity to contribute a foreword to this work by Professors Shahal Abbo and Avi Gopher, my long-time colleagues in the study of this fascinating and hugely complicated question of the origin of agriculture in the Near East. Plant domestication and the origins of agriculture in the Near East were the most significant innovations of our species Homo sapiens, second only to the use of fire, which was initiated hundreds of thousands of years ago by an earlier member of the genus Homo. This book tells the plants’ part of the story of the Neolithic or Agricultural Revolution, a foundational change in human existence that took place less than 11,000 years ago. Since its beginnings, that revolutionary turn of events has resulted in an increase of more than 1,000-fold in the global human population and remains a cornerstone of modern human life. After several million years of hominin existence based on hunting and gathering of natural resources, the consequences of both food production and sedentism by most of humanity were enormous. Their technological and cultural outcomes stem from the ability of agriculture to support large human groups where people (at first only a few and later many multitudes) are not preoccupied with food production. Those early free people invented and adopted the usage of measuring, counting, writing, metallurgy, and in due time established urbanization, large social human organizations such as ancient Sumer, Egypt, Greece, Rome, China, and in more recent times modern states, professional armies, art and literature, musical instruments and ensembles, science and technology, modern medicine, people walking on the Moon and sending vehicles to Mars.

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The move towards tending the soil and growing crops was at least as much a cultural as a technological phenomenon, and perhaps even more so. It required a dramatic change, involving ideological and spiritual elements, in the perception of nature. Unfortunately, far too little is known about these aspects and their roles in the striking changes taking place in human behaviour and human relationships, both within society (human–human) and towards nature (human–world). This is because some of those aspects are archaeologically transparent, or at least do not lend themselves to an easy reading through the archaeological record. The actual data that are much more amenable to study are those from botanical and zoological finds recovered from archaeological sites, from genetics of archaeological and extant plants, animals and humans, and from archaeological material culture including site characteristics, architecture, stone industries, burial data, language and other features. To some readers, this book may be considered provocative. That is part of its value. Its fifteen chapters are focused on evidence-based aspects of the subjects pertinent to plant domestication in the relevant chronological (Neolithic) range and geographic (Near Eastern) expanse. To the study of plant domestication in the Neolithic Near East, Abbo and Gopher bring a balanced and complementary fund of knowledge. Shahal Abbo is a geneticist and agronomist who specializes in Mediterranean legume and cereal crops, including field biology and genetics of the wild progenitors of domesticated crops, and Avi Gopher is an expert in the archaeology of the Near Eastern Neolithic period. Their deep and consistent collaborations with botanists, population geneticists, experts in human nutrition and in plant diseases grants them a comprehensive view of this complex and multidisciplinary field of research. This book is a continuation of a long list of scholarly essays by leading scientists such as de Candolle, Vavilov, Braidwood, Harlan, Zohary, Ladizinsky and others, who, for over a century, have advanced our understanding of the origins of agriculture in the Neolithic Near East. Since plant domestication is essentially a cultural phenomenon, it must be reconstructed and understood in the context of its cultural background – that is to say, of its make-up and its time-space systematics. These must relate firstly to the time, and secondly to the place of plant domestication. The earliest evidence for the origin of any cultural element is likely to be found in the area where the phenomenon first appeared. The absolute and accurate dating of the cultural element (in the case of this book, plant domestication) becomes central, and when data are lacking, or are marred by inaccuracy or selective use or confusion, misunderstanding might result. Archaeological data are carefully collected and assigned to archaeological entities (such as archaeological cultures), following well-established systematics. When such cultural entities lose distinctiveness and their temporal borders become blurred, as was done and even prized by some scholars, the potential for further mistakes in reconstructing past phenomena and processes grows exponentially. For instance, some basic facts, too often overlooked, are that not a few Neolithic cultural phenomena and materials within the Near East spread from the northern to the southern Levant, and that such diffusion of

FOREWORD

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innovations from the north to the south could take a couple of centuries. Disregarding this possibility might interfere with the construction of models of the origin of agriculture and mislead scholars discussing Neolithic plant domestication and agriculture. It is imperative, therefore, that all archaeobotanical and genetic data be considered in light of the archaeological facts and cultural dynamics as revealed by the many excavations of the Pre-Pottery Neolithic period. This book, without delving into detailed discussions on the relevant periods, emphasizes the law and order that are essential in these respects. The same philosophy applies when contemplating the biology of the progenitors of the eight-species package constituting the founder crops of Near Eastern Neolithic agriculture, namely, emmer wheat (Triticum turgidum), einkorn wheat (T. monococcum), barley (Hordeum vulgare), pea (Pisum sativum), chickpea (Cicer arietinum), lentil (Lens culinaris), bitter vetch (Vicia ervilia) and flax (Linum usitatissimum). Abbo and Gopher systematically present the essence of the issues that relate to the origin of Near Eastern agriculture in the Pre-Pottery Neolithic B period some 10,500 years ago, as well as to the origin of fruit tree domestication that occurred several millennia later. Their book illuminates and reconstructs plant domestication in an accessible way to a readership of knowledge-seekers. It is also aimed at students, scholars, archaeologists, geneticists, archaeobotanists, botanists, plant breeders and anyone interested in human culture and this fascinating critical aspect of history. Still currently under debate are some of the partly unresolved questions about the geographic origin of Near Eastern plant domestication, its mechanisms and pace and the consciousness of the active prehistoric communities who established it. Rather than going into detailed polemic discussions on those issues, this book presents the coherent view of the authors (with which I agree) that plant domestication originated in a limited region in south-eastern Turkey/northern Syria, and that it was a knowledge-based initiative, conscious and episodic in time. As an experienced scientist, I expect that the views presented here will stimulate further relevant studies, and I am certain that this book will help others to deal carefully with the complicated story of the origin of agriculture in the Pre-Pottery Neolithic B of the Near East. Simcha Lev-Yadun

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F O R E WO R D

It is hard to overestimate the significance of events that took place some 10,000 years ago and more or less contemporaneously in different places across several continents. The Earth, coming out of the latest Ice Age episode, had started warming up by fits and starts. Humans inhabiting the Earth adapted gradually to the higher temperature, variable precipitation regimes, changing seasons and higher carbon dioxide concentrations in the atmosphere. Whether out of necessity to increase their food security or as an opportunity, hunter-gatherers made a momentous change in the way they procured food: they started planting some of the plants that were well-known to them but which earlier they merely harvested. This apparently simple planting gesture became seminal in the development of agriculture: at that time, an entirely new way of acquiring food and other plant resources (‘le geste auguste du semeur’ of Victor Hugo). The initiation of planting represented – depending on one’s viewpoint – either the apotheosis of the hunting-gathering era or the dawn of the agricultural era. It represented a major milestone in the evolution of the human lineage up to this day. The whence and whereto, when, how and, above all, why of agricultural origins in the Fertile Crescent of the Middle East are the topic of this most interesting book authored by Professors Shahal Abbo (Hebrew University) and Avi Gopher (Tel Aviv University). I am proud to count S. Abbo and A. Gopher as colleagues in the scientific area of crop evolution and agricultural origin studies. My own focus is on the origins of crops in Mesoamerica and the Andes, with emphasis on Phaseolus beans, but I have always followed the research of my two colleagues with great interest for their innovative approaches and thought-provoking writings on the topic. Plant genetics/breeding and

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archaeology are the two main, complementary sources of evidence on the origins of agriculture and domestication. The two authors have an extensive record of collaborative research that has led to this publication and speak with authority about the complex topic of the origins of agriculture and crop domestication. The transitions from hunting-gathering to agriculture took place independently and more or less simultaneously in multiple regions of the world. Not only were multiple new technologies introduced around that time such as domesticated plants and animals and pottery, giving rise to the term ‘Neolithic Revolution’, but the practice of agriculture was associated with major changes in human society around the time of the huntinggathering to agriculture transitions, including a more sedentary lifestyle, the development of city-states and ultimately the appearance of ancient and more recent civilizations. While these new societal structures and civilizations were not involved in the transitions to agriculture, they were one of the most significant outcomes. Agriculture was a necessary condition for the development of civilizations such as Sumer, Assur, Akkad and Babylon in Mesopotamia and the Olmecs, Mayas and Mexicans in Mesoamerica, further highlighting its crucial importance. Nevertheless, the specific causes and conditions of these transitions from huntinggathering to agriculture remain difficult to ascertain, in part because the biological, climatic, economic and social circumstances that surrounded the transitions differed in the various regions of agricultural origins. The plant (or animal) characteristics (such as its genome attributes, life history and reproductive system), the environment (including its climate and biological interactions with pollinators, herbivores and microbes) and human factors (for example, cultural advancement such as plant and animal knowledge and tools) varied markedly. This is illustrated by the contrasting crops that were the founder domesticates in their respective regions. These included annual herbaceous crops such as cereals (wheat, barley), grain legumes (chickpea, lentil, pea) and flax in the Fertile Crescent, as a consequence of the Mediterranean climate; perennial herbaceous plants such as bananas and sugarcane in south-east Asia; and perennial lignified plants such cacao, peach palm and cassava in the Amazon. It is, therefore, not a surprise that the transitions from hunting-gathering to agriculture followed different trajectories in different regions of the world. One of the best-studied regions in this respect is the Fertile Crescent. This region includes the Levant, southern Turkey and northern Syria, and eastern Iraq and western Iran, encompassing the mountainous regions surrounding Mesopotamia. The richness of the scientific record can be attributed to a relatively dry climate, conducive to the conservation of archaeological remains, and a long-standing interest by a multidisciplinary group of plant and animal scientists, including Abbo and Gopher. They address several issues that are the subject of current controversies in this scientific field, not only in the Fertile Crescent but in other regions of agricultural origins as well. For example, where were crops that originated in the Fertile Crescent domesticated? Did they all trace back to the same ‘core’ area or did they have a dispersed origin across

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this region? Abbo and Gopher propose a core area in south-eastern Turkey and northern Syria, based on the overlapping distribution of the wild progenitors of crops such einkorn wheat, barley and chickpea, and the oldest 14C isotope dates of archaeobotanical remains. The dispersed domestication origin across the Fertile Crescent proposed by others is part of the current paradigm of the origin of agriculture, which posits a slow progression from hunting-gathering to agriculture with limited human agency, in which crop domestication would have happened as – in the authors’ words – ‘an unguided, unintended and mostly unconscious development’, a sort of ‘immaculate domestication’ (in my own words). Instead, Abbo and Gopher argue that this paradigm ignores the significant knowledge accumulated by hunter-gatherers about their environment, including the life history, uses and adaptation of plants that surrounded them. Such knowledge was essential for their thrift and is still present among hunter-gatherers today and, in my experience, also subsistence farmers, who play an active role in the maintenance and development of their seed stocks. Abbo and Gopher argue that the transition to agriculture resulted from the huntergatherers’ awareness of plant (and animal) characteristics, which drove their intentionality and conscious selection of domesticated plants (and animals). They further posit a significant change in attitude towards the natural world on the part of hunter-gatherers, immediately preceding the transition to agriculture: from one that used natural resources to one that – through the acts of planting and domestication – created and exploited resources, which perdures to this day although in a markedly intensified way, with all the health and environmental consequences thereof. It is this integrated vision of plant (and animal) characteristics, environmental circumstances and human agency that makes this book so interesting. The combination of the multidisciplinary technical information gathered by scientists studying the origins of agriculture represents the foundational scientific research and information on which books such as Guns, Germs, and Steel by Jared Diamond and The Botany of Desire by Michael Pollan are largely based. I will end on a personal note. As an undergraduate in agricultural sciences in Belgium, I spent some time harvesting crops in Kibbutz Hamadyah, north of Beit Shean, in the Jordan Valley. The kibbutz grew wheat, pomegranates and dates, among other crops. Little did I know then that I would include my first-hand knowledge of these iconic crops of the Middle East in my course on crop evolution and agricultural origins at UC Davis. Paul Gepts

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PREFACE AND ACKNOWLEDGEMENTS

This book is about plant domestication that occurred in the Near East some 10,500 years ago. For a long time now, we have been investigating various aspects of plant domestication and the origins of agriculture in the region: Shahal Abbo studying aspects of agronomy, biology and genetics, and Avi Gopher studying archaeology. Shahal Abbo, as a dedicated student of Professor Gideon Ladizinsky, has been focusing on different aspects of legumes, particularly on the chickpea, while Avi Gopher, a disciple of the late Professor Ofer Bar – Yosef, has been concentrating on the origins of agriculture and investigating the Neolithic period – a prominent key research issue for over a century. Professor Simcha Lev-Yadun, a third partner (see the first Foreword), whose expertise lies in the botany, ecology and evolution of Near Eastern plants, chose not to partake in the writing of this book, yet his contribution is undisputed. Our joint work was publicly recognized when it was first published in the year 2000 in Science as an original paper conceived by Professor Lev-Yadun. There we claimed that plant domestication originated in a small, well-defined geographic core area spanning south-eastern Turkey and northern Syria, and that it occurred in a single, fairly rapid event (singular timing). Later, domesticated plants spread to other parts of the Near East and beyond – to Mediterranean islands (such as Cyprus), Europe, southeast Asia and Africa. This claim is based on (archaeo)botanical findings of various archaeological sites combined with geobotanical, genetic, agronomic and cultural considerations. Over the years we have published many papers in well-known international journals (see Further Reading at the end of this book) in which we explored diverse aspects of

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plant domestication. The research included fieldwork and controlled experiments and our findings support our original suggestion from 2000. Nevertheless, and despite other published works that supported our view, we found ourselves in a minority: most researchers of this issue did not and do not accept our arguments. In 2016, we presented our views and the full breadth of our research on the knowledge we have accumulated regarding plant domestication in Hebrew to a broad Israeli readership. That book (Plant Domestication and the Beginning of Agriculture in the Near East, Resling Publishing House, Tel Aviv) was written not only for an academic audience but for students and the broad readership of knowledge-seekers, including those interested in the origins of agriculture and plant domestication in the Near East. We believe that some of the topics discussed in this book are also relevant to modern plant breeders, agronomists and farmers. Feedback from Israeli readers was surprising and rewarding. It became clear to us through various professional and non-professional readers that a clear statement summarizing plant domestication in the Near East was necessary, and we decided to translate the Hebrew book into English before embarking on the original mission, namely, to write a detailed, fully referenced polemic discussion on plant domestication in the Near East. In the volume presented here, we refrain from reviewing the dynamic discussion and details of plant domestication that we have presented in professional publications. Rather, we state our opinion and ideas in full, yet concisely, acknowledging differences of opinion. We will attempt to convince the reader that our suggested reconstruction of plant domestication and the emergence of agriculture stands the test of both available Near Eastern data and results of recent professional analytical research work carried out in the Near East. We acknowledge that our reconstruction must be tied to the fact that we live in Israel, and it cannot be divorced from that or from our respective professional and personal backgrounds. We wish to thank the many good people who collaborated with us. First and foremost, we thank Professor Simcha Lev-Yadun, our partner, who read an early draft of the Hebrew text and the published Hebrew version of this book and made valuable remarks. We thank Professor Gila Kahila Bar-Gal, who kindly contributed a chapter on animal domestication in the Near East. We thank the Israeli Science Foundation for its Bikura Special Personal Track research grant No. 1406/05 as well as funding targeted at the publication of the Hebrew version of this book. We thank the research authorities of the Hebrew University of Jerusalem and Tel Aviv University for their support of the Hebrew publication. Shahal Abbo thanks the Jacob and Rachel Liss Chair of Agronomy at the Hebrew University of Jerusalem for its support. We thank our scientific publication partners, Professor Gideon Ladizinsky, Professor Baruch Rubin, Professor Tzion Fahima, Professor Avraham Korol, Professor Yehoshua Saranga, Professor Dani Shteinberg, Professor Ram Reifen, Professor Gideon Neeman, xvi

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Professor Zohar Kerem, Dr Amir Sherman, Dr David Bonfil, Professor Zvi Peleg, Dr Orit Shamir, Dr Moshe Fleishman, Professor Ofer Reany and Mrs Toni Friedman. We thank our students, who have broadened our minds, wrote theses and worked with us in the fields; the vision of some went beyond that of our own – Dr Ran Hovav, Dr Judith Lichtenzveig, Dr Roi Ben-David, Dr Omer Frenkel, Dr Ruth Pinhasi van-Oss, Ms Inbar Zezak, Ms Yael Zehavi, Ms Efrat Schwartz, Mr Erez Rachamim, Mr Itai Ofner and Dr Raanan Tzarfati. We also thank the Israeli farmers who hosted our experiments in their fields – agronomists Yaacov Yinon and Assaf Avneri, Mr Aharon Glazer, Tal and Eli Galili and others. We also thank Michael Ben-Dor for his assistance. We thank Itamar Ben-Ezra for preparing illustrations and for re-organizing the figures for publication. We thank Yael Saranga for preparing illustrations 5.1, Glossary 1 and 2 and for editing Figure 13.1. We thank Heeli Schechter, who assisted in the preparation of graphics and text boxes displayed in this book. We thank Adi Halpern for the preparation of Figure 2.3. We thank our partners from abroad: Professor Canan Can (Gaziantep University, Turkey), Professor Manfred Heun (Norwegian University of Life Sciences, Ås, Norway), Professor Francesco Salamini (Italy), Professor Tobin Peever (Washington State University, Pullman, USA), Professor Neil Turner, Dr Jens Berger and Dr Nick Galwey (the University of Western Australia, Australia). We thank those who allowed us to use their photos and illustrations: Professor Erella Hovers, Dr Francois Valla, Mrs Marjolain Barazani, Professor Juan José Ibáñez, the late Professor Ofer Bar-Yosef, Professor Nigel Goring-Morris, the late Dr Klaus Schmidt, Dr Hamoudi Khalaily, Dr Ianir Milevski, Professor Dany Nadel, Professor Gary Rollefson, Dr Andrew Moore, Professor Miquel Mollist, Professor Mehmet Özdoğan, Dr Ehud Galili, Professor Yosef Garfinkel, Dr Danielle Storduer, Mr Nimrod Getzov, Professor Simcha Lev-Yadun, Professor Jared Diamond, Dr Yuval Cohen, Professor Zvi Peleg, Dr Jeff Paull, Ms Talia Oron, Ms Rachel Gabrieli, Mr Alex Kantorovich, Mr Amir Balaban and Professor Yigal Elad. Our gratitude is especially extended to Professor Gideon Ladizinsky and Professor Ran Barkai, who read the Hebrew manuscript, and to Professor Amram Ashri, who read the English translation. All three readers have made many important contributions, assisted in eliminating some errors and offered significant advice and commentary. We thank the Resling Publishing House for accepting our Hebrew version of this book for publication and for the good work of their editors and staff. We thank Ms Halo (Hilla) Ben Asher, who translated this book into English, and Ms Myrna Pollak for copyediting the English translation. We also thank Beatrice Rehl and Edgar Mendez of Cambridge University Press and all who partook in the creation of this book and its print production process. Naturally, responsibility for this book’s content (including any mistakes or inaccuracies) is solely ours. PREFACE AND ACKNOWLEDGEMENTS

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INTRODUCTION

This book is about plant domestication and the origins of agriculture in the Near East (Figure Introduction 1), which were major components of the process known as the Agricultural Revolution or the Neolithic Revolution.1 The expansive discussion we offer in this book is restricted to plant domestication; animal domestication, a broad subject in its own right, is treated in a contribution by Professor Gila Kahila Bar-Gal (see Chapter 14). The term ‘domestication’ (or ‘plant domestication’) in the context of the current work carries both biological and cultural significance. From the biological perspective, it implies the acquisition of new traits that differ from the prevalent wild type plant while the cultural perspective denotes a change in worldviews and life-ways enabling the adoption of plants for food production. Throughout this book we distinguish between the Agricultural Revolution (see Glossary, General Terms, Agricultural Revolution) as a general socio-cultural transformation and the domestication (see Glossary, General Terms, Plant domestication) of plants (and animals) as a single aspect of this multifaceted human development. This book, then, aims mainly at addressing key questions concerning plant domestication in the Near East (some 10,500 years ago) while exploring the new and particular relationship that ensued between humans and plants as well as the general interaction that developed between human/culture and nature. We offer a discussion on some of the fundamental questions that relate to the broader cultural transformation (the Agricultural Revolution), including: What is the Agricultural Revolution? When and where did it occur? How did it occur? And, perhaps most important, why did it occur?

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The Blac k

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Int. 1 General map of the Near East; the core area we suggest as the home of plant domestication is denoted by an arrow.

Some of the answers may be simple and fairly straightforward, but others are difficult and complex, pertaining to the very foundations of our human existence. Plant domestication in the Near East occurred some 10,500 years ago, during the Neolithic period (the New Stone Age). The Near Eastern founder crops, that is, the group of plants that were to be domesticated and become the Neolithic plant package, were

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Int. 2 Wild progenitors of the Near Eastern domesticated plant package: (a) emmer or durum wheat, (b) barley, (c) bitter vetch, (d) einkorn wheat, (e) pea, (f ) flax or linseed, (g) lentil, (h) chickpea (refer to Table 4.1 for Latin names and progenitors). Colour versions of these images can be found at www.cambridge.org/abbo-gopher.

barley, two wheat species, pea, lentil, chickpea, bitter vetch and flax (Figure Introduction 2). These crops are still in use, providing a significant part of the agricultural produce used to feed both humans and livestock. Considering the animals that were domesticated at the same time – goats, sheep, cattle and pigs – it is easy to see that some of the prominent food products consumed worldwide to this very day were, in fact, singled out 10,500 years ago and adopted as part of the Agricultural Revolution. The transition to an agricultural way of life necessitated far-reaching changes in

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humankind’s perceptions, worldviews, social structure and institutions, all of which led humanity to its familiar, modern state. It is important to remember that agricultural revolutions also occurred independently in other world regions, where other crop plants were adopted, including cereals (e.g., rice, maize), legumes (e.g., soy bean, common bean) and species from other plant families (Solanaceae, Cucurbitaceae, Compositae, Malvaceae), according to the wild species repertoire of each region (e.g., potato, pumpkin, sunflower, cotton respectively). We are in awe of the creativity and vision of Neolithic societies and their striking capability worldwide to find useful plant and animal species suitable for domestication as well as their adeptness in applying the delicate and sophisticated decision making that was required for domestication. Despite typical challenges in the early days of agriculture, this was a highly successful system,2 as attested by its global sweep, and to date it underlies the socio-economic organization of most human societies.3 In fact, it is rather difficult to find an inhabited region anywhere in the world today that is devoid of agriculture. This is neither a botany nor archaeology textbook.4 It is rather a volume aimed at a wider intellectual readership of knowledge-seekers. To facilitate reading, we hope the text below describing how the book is structured, and the logic threaded in and between its chapters, will assist readers in following our arguments and suggestions regarding plant domestication in the Near East. Certain chapters are also laced with explanatory boxes and illustrations that clarify terms and introduce the archaeological sites or data upon which we base our claims. A Glossary with short definitions and explanations of professional terms appears at the end of the volume. Any discussion of plant domestication is multifaceted and complex. In the Near East, this discussion is characterized by a series of bipolar, dichotomous questions regarding the where, when (and at what pace) and how of plant domestication. The first question to ask, then, as it governs answers to all other questions, is whether or not there was a core area (within the Near East) in which plants were first domesticated and from which they spread, and if such an area did exist, where it was located. We respond positively to the first question, and determine, based on available data, that this area was located in south-eastern Turkey and northern Syria. Accordingly, we maintain: • that founder crops were domesticated in a single episode; • that this single episode was led by a specific set of considerations (e.g., cultural, economic, agronomic, nutritional); • and that these considerations, in turn, affected other aspects of the process, such as pace (was it rapid or slow?); or consciousness (were the actions of the domesticators deliberate?); or knowledge (was the process knowledge-based and pre-planned, or was it an accidental by-product of human behaviour?). We now map out for the reader the plan and scope of the book. The brief overview of each chapter stresses our main insights regarding the different aspects of

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plant domestication, and provides the basic knowledge required to understand and support our claims. In Chapter 1, we review the background and the time frame of plant domestication and the Agricultural Revolution in the Near East. We show the place of the Agricultural Revolution on a long continuum of revolutions (or transformations) from ancient to modern times, including ‘man the tool-maker’, the emergence of agriculture, the rise of city-states or urban centres, the Industrial Revolution and the Digital (computerized, virtual, information/communication technology) Revolution that is still unfolding around us. We introduce plant domestication in the broad context of domestications that build a picture of cultural change, through which humans from the very earliest times came to appropriate, dominate and regulate the world’s natural resources – a time, for example, when they came to control stone and fire, domesticate plants and animals (the Agricultural Revolution), manage water (dig wells and irrigate farm land with canals), manipulate clay (vessel making) and metal (copper and iron making) and possibly even domesticate and discipline themselves and their own species. Since the domesticators of plants (and animals) were hunter-gatherers living in small communities, we devote a short discussion to the characteristics of these societies while also comparing them to the subsequent food-producing societies. The dwelling sites of these two types of societies, their mobility patterns across the landscape, the economic and demographic aspects of the two systems, the social structure and organization of their respective lifestyles, and the differences in worldviews between them are briefly presented, especially as they pertain to the human–world or culture–nature relationships. In essence, in this chapter we sketch, in general, the socio-economic consequences of the Agricultural Revolution and seek to establish how this socio-cultural transformation led to the modern human condition and the birth of our current civilizations. Chapter 2 is devoted to the archaeology of the Fertile Crescent, especially its western wing, towards the end of the last Ice Age (Pleistocene) and the early Holocene, the present age.5 We offer a short historical review pertaining to human dwelling sites, technology and tools (mainly flint and stone) and life-ways during the long period known as the Epipaleolithic (ca. 23,000–12,000 years before present, or cal BP).6 We then introduce the Neolithic period (ca. 11,800/700–6,500 cal BP), during which Near Eastern plant domestication occurred and farming economies were established. Some relevant archaeological sites and findings are described in text boxes and illustrations. We offer only a brief and broad overview in the hope of stimulating our readers to seek further knowledge of these topics. Through archaeological findings, we delineate the transformation in the archaeological landscape before and after the Agricultural Revolution. The establishment of farming communities was accompanied by massive changes in human culture and worldviews, marking the end of some three million years of Paleolithic human existence and perceptions. The new world, the Neolithic world, was distinguished by its large sedentary populations dwelling in permanent sites, a new economy that was based on food production, and a new, more complex and less

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egalitarian (ranked and later stratified) social organization, to name but a few of the characteristics of this new era. In Chapter 3, we review the different explanations offered over the years for the questions we raised earlier: When did the Agricultural Revolution occur? Where? How? And why? Additional features of hunter-gatherer societies are discussed as background to some of the explanations and models of the Agricultural Revolution. The explanations presented here are divided into two categories: (1) the models that view the Agricultural Revolution as a consequence of external (e.g., environmental) influences and stress factors (resource depletion) that led to the adoption of new subsistence resources; (2) explanations that consider changing perceptions, worldviews and the socio-economic order as the transformational drive of social restructuring and new life-ways. The first set of explanations emphasize the reactive nature of humans and their culture when facing external forces. The second group of explanations – the one to which we subscribe – consider human agency and human initiative as the major drivers of social and cultural changes that were but one possible choice of action and not necessarily one related to stress. In our view, the Agricultural Revolution and plant domestication were the result of human social dynamics that emerged due to perceptual and ideological changes. Reviewing both types of models emphasizes their respective underlying viewpoints, thereby helping to elucidate our own stance that plant domestication was a proactive, knowledge-based skilful development. In Chapter 4, we describe the environment in which the Agricultural Revolution and plant domestication took place. We discuss the natural arena of these events, including climate, physical landscape and the ecology of the Near East. The founder crop species and their wild progenitors are presented as necessary background for understanding the bio-mechanisms of domestication.7 The geology and climate regime (mostly precipitation seasonality) of the eastern Mediterranean Basin create a rich and highly diverse but also highly vulnerable environment, the richness of which is not always easily detected. The potential of floral diversity, including the wild progenitors of the founder crops (as well as the animals to be domesticated), was in this case a hidden treasure awaiting use by local ‘owners’, to draw on Ecclesiastes (Chapter 5:13, KJB). Such areas, rich in the species of plants (and animals) that were later domesticated, coupled with suitable conditions for such a move, are rare in the world, as noted by Jared Diamond in his book, Guns, Germs, and Steel (1997). In Chapter 5, we describe the main differences between wild and domesticated plants, we explore the traits that allow wild plants to adapt to their habitat and show how these traits are relevant to domesticated plants. Some of these traits, known collectively as the domestication syndrome, are described with respect to the Near Eastern founder crops, including seed dispersal, seed dormancy, pest and disease resiliency, adaptation to climatic rhythm (seasonality), the extent of genetic diversity, nutritional value and the economic value of various plant parts, among others. We conclude that although general similarity renders most domesticated plants ‘alike’ (e.g., most Near

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Eastern annual crops are self-pollinating), researchers are not exempt from studying the unique peculiarities of each of the founder crops that required specific attention by the ancient domesticators. In Chapter 6, we probe the differences between modern and traditional (subsistence) agricultural systems, preparing the reader to understand the economic, nutritional and agronomic considerations underlying the adoption of species that finally made the Neolithic crop package. We present the differences between low-input traditional systems targeted at subsistence farming and modern systems that necessitate expansive infrastructures targeted at large-scale commercial industrial production. In Chapter 7, we explore a highly controversial issue in current plant domestication research: plant domestication (adoption of a wild plant for cultivation) versus crop evolution under domestication (all post-domestication improvements and changes, including modern breeding). Our main contention is that domestication occurred in a short, rapid episode, often requiring just a single (crucial) genetic change in the wild type to enable profitable cultivation. Accordingly, many of the other traits that differ between wild and domesticated plants evolved over millennia under domestication in farmers’ fields and were not necessarily associated with the pristine domestication episode. The domestication syndrome is discussed with respect to three species of the founders’ crop package – chickpea, pea and emmer wheat – emphasizing the distinction between adaptations that occurred as part of the domestication episode and subsequent evolutionary changes that took place under domestication. We thus attempt to single out those traits that were crucial for domestication vis-à-vis those that did not interfere with usage or cultivation. We believe that this distinction is critical for understanding domestication and our ability to determine whether it was a long, protracted process or a rapid, short event. Literature advocating the former is deficient in its discussion of this issue and therefore bases its argument on plant adaptations that are unrelated to domestication and which continue to evolve as part of the evolutionary trajectory of domesticated plants (such as in modern plant breeding). Chapter 8 is dedicated to the distinction between cereals and legumes – the two prominent plant groups within the Near Eastern domesticated plant package. Cereal and legume crops are the backbone of agro-eco-systems in several world regions. The biological features of these two crop groups suggest that their mode of domestication was entirely different, determined by plant stature, growth habit, reproductive biology, seed dormancy and other traits. We also present traits of plants that belong to neither group, such as flax and a domesticated plant whose wild progenitor is unknown – the broad (faba) bean. We conclude that domesticating cereals and legumes required different approaches and skills, and that legume domestication probably required more ingenuity and sophistication on the part of the early farmers. This is because wild and domesticated cereals share similar traits and growth habits, with both growing rather densely. In contrast, the sparse patches in which the progenitors of the domesticated legumes grow in the wild are quite different compared to their appearance in full

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canopy cultivated fields, thus necessitating a different approach (a conceptual leap) from the one that would be applied in the domestication of cereals. Another important conclusion regarding this crop combination is that it improves yield stability of the farming system in addition to providing balanced amino acid (whole protein) nutrition. We believe that this was likely the driver of similar domestication patterns in other world regions as well. A third conclusion emerging from these facts seems to have eroded in our busy modern world: that the main goal of subsistence farming systems is not yield maximization but rather the maintenance of stability of crops and produce, thereby contributing to sustainability. In Chapter 9, we discuss the choice of the Near Eastern wild candidates for domestication with respect to availability, nutritional value of each individual species, agronomic potential both in isolation and as components of the whole crop package, and their contribution to overall agronomic balance and yield stability. The natural productivity of the individual wild species, that is, their caloric and nutritional value, does not always provide a satisfactory explanation for their adoption, namely, it is not always a good predictor of the agronomic potential and often does not reflect the likely incentive for adoption in antiquity. We therefore turn to other traits, such as the unique nutritional contribution of each species, their taste, agricultural compensation potential and ease of manipulation. The three crop cases we discuss as examples – chickpea, lentil and pea – provide support for the notion of knowledge-based, fully conscious plant domestication in the Near East. In Chapter 10, we return to our central questions: When, where and how did plant domestication occur? We offer our model, by which domestication was a short-lived episode, occurring in a limited area, ca. 300 km in diameter, some 10,500 years ago. We show in many diverse ways that domestication was premeditated (conscious) and knowledge-based, and that it involved the full, harmonic plant package rather than each crop individually. We bring ample evidence from a broad range of disciplines to support our model, including archaeology, archaeobotany (the study of archaeological botanical findings), geobotany and genetics. In Chapter 11, we divert from the discussion of grain crops to discussing fruit tree domestication, which sheds additional light on the deliberations involved in producing the Near Eastern domesticated plant package. We focus on the emblematic Near Eastern fruit trees – olive, fig, grape, date, pomegranate – which were the first to be domesticated in this region and which form five of the seven biblical species of food plants (fruits and grains) grown in the Land of Israel (and with the remaining two – barley and wheat – make up a food package in their own right). We compare the domestication of annual grain crops (cereals and legumes) to the adoption of perennial woody species as crop plants, thus depicting a broader general picture of plant domestication. These fruit trees were probably assimilated into farming systems several millennia after the domestication of annual crops, as this necessitated greater experience stemming from fundamental biological differences between annual and perennial growth, the long juvenile

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period (see Glossary, Botany, Ecology and Agronomy, Juvenile period) and the understanding that the tree may change in appearance after its early years, in its reproductive biology (i.e., the flowering and fruiting patterns), and clonal (vegetative) propagation (see Glossary, Botany, Ecology and Agronomy, Clonal propagation). Additionally, tree growing involves a higher degree of delayed return, that is, it might take several years before trees yield produce and generate actual gain. Tree growing also involves a higher degree of long-term planning due to farmland allocation considerations and the prolonged lifetime of trees (e.g., several hundred years in the case of the olive). In Chapter 12, we present post-domestication processes, namely, evolutionary processes that characterize domesticated plants in their domesticated habitat. We address the naturally occurring genetic variability of crop plants and explain the evolutionary forces that promote variability as well as those that restrict genetic variation, such as selective sweeps like the domestication bottlenecks. An immediate – although not innovative – conclusion arising from this discussion is that safeguarding genetic variability of crops (and wild plants) is fundamental for modern plant breeding, especially in light of the ever-increasing demand for the supply of high-quality foods in a globalizing world. In Chapter 13, we extend our view to other world regions (domestication centres in America, Africa, Asia) and their respective crops. This review attests to independent, primary domestication centres of both annual crops and trees. Surveying the works of veteran bio-geographers such as de Candolle or pioneering geneticists like Vavilov raises questions concerning the ecological (dis)similarity of all domestication sites or the cultural independence (or lack thereof ) of domestication centres. We discuss contemporary genetic variability and its significance to the sustainability of modern farming systems. Animal domestication was part and parcel of the Near Eastern Agricultural Revolution and it is described separately in Chapter 14 by Professor Gila Kahila Bar-Gal, which unlike all other chapters includes a full academic apparatus (citations and a References section on p. 247). To make it reader-friendly, brief explanations of some of the terms and references to the Glossary are included. This chapter completes the picture of domestications in the Near East and provides a brief discussion and summary of the features of animal domestication that can be examined vis-à-vis plant domestication. Sheep, goats, cattle and pigs – the package of Near Eastern domesticated animals, or the ‘Big Four’ – were all domesticated within (or in the close vicinity of ) the proposed core area of plant domestication in south-eastern Turkey and northern Syria. The wild progenitors and their ecological affinities are introduced and the differences between the wild forms and the domesticated livestock are highlighted. The archaeozoological record shows that domesticated morphologies appear rather abruptly and within a similar time frame to plant domestication. It is stressed that the differential feeding habits of the four livestock species grant flexibility to the farmer since there is little competition between them for food resources. As with plants, this chapter tends to suggest a knowledge-based choice of the four animals for domestication. Discussing the

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possible incentives for animal domestication Professor Bar-Gal rejects climatic deterioration or resource depletion. She rather proposes that the motivation for animal domestication ought to be looked for in the cultural arena. In Chapter 15, our final chapter, we summarize our views on the central questions of this work: When, where and how did plant domestication occur? We discuss the time frame of domestication, the domestication of fruit trees and genetic variability. We briefly touch upon the geographic spread of Near Eastern domesticated crops in ancient Europe and Asia and the consolidation of the founder crops (annuals and perennials) into a coherent agro-economic system. We look into cultural (historical and perceptual) aspects of domestication and humbly attempt to answer the question that remains open: Why domesticate? Was an economic need, driven by external factors, the major motivating force? Or was it a change in perception and ideology that initiated and led this irreversible cultural transformation? Undoubtedly, plant (and animal) domestication involved a transformational conceptual and socio-cultural restructuring of human society, and this ultimately led to the formation of complex, stratified, urban and modern societies. Plant and animal domestication and the institutionalization of agriculture have had a far-reaching influence on humanity, and they continue to influence our lives today. First and foremost, they have reshaped the human–environment relationship; they have driven many technological innovations; brought about the emergence of new industries and professional specializations; generated social reorganization from time to time and the construction of social institutions including labour division, gender relations and social ranking; and they might even have propelled the advent of gods and religions. These are all elements that have led humanity to its modern reality; they are still prevalent in our contemporary societies, and they are likely to continue and influence humanity in the near future.

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1 WH AT IS TH E A GRICULTU RAL REVO LU TIO N?

The Agricultural Revolution (which made us all – humans – food-producers) is a major landmark in human history. It reflects a significant transformation in the general organization of human society and its components (see Glossary, General Terms, Agricultural Revolution). Since the advent of humans as tool-makers, some three million years ago, that is, since the moment we began producing tools and using them as a key vehicle in our daily activities, no transformation was as significant as the Agricultural Revolution. After three million years of living in small, mobile communities while subsisting by hunting animals and gathering plant foods, the Agricultural Revolution, which took place post-Pleistocene, during the Neolithic period, just over 10,000 years ago, brought about a prominent transformation in human life-ways. Mostly, it allowed humans to become food-producers, rendering them a unique and singular being on Earth. Small or large sedentary farming villages, characterizing the new way of life in the Near East, soon grew in size to very large villages, also known in Neolithic research as towns,1 which later became cities (Figure 1.1). Some claim that the development of cities in the Near East about 6,000–5,000 years ago, compared to the large villages that preceded them, was but a natural development of phenomena that had originated in the Agricultural Revolution. Others see a major significance in this transformation of urbanization. In their view, and we concur, this was another revolution that bore additional extensive consequences for the socioeconomic fabric. Among the changes of this revolution was the expansion of settlements, the concentration of large populations in newly emergent cities and the birth of new social institutions – kings and princes, states and taxes, temples and priests, armies and generals – all representing yet another step towards the formation of the modern

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Figure 1.1 (a) An illustration of work in the fields; (b) Tell Abu Hureyra reconstruction of the village – courtesy of Andrew Moore and Oxford University Press, New York.

world as we know it. The next significant transformation would not occur until thousands of years later. The Industrial Revolution that took place in the Western world some 200 years ago could also be perceived as an important transformation in the history of humankind, which further changed human perceptions as well as social structure and organization. We view the borderless Digital (information technology) Revolution currently taking place as yet another transformation, unfolding before us as it drives far-reaching changes in human societies and human behaviour. This short historical summary of human transformations clearly shows that changes in the way humans perceive their world and the structure and organization of their societies and economies are occurring at shorter time intervals and at an accelerating pace. With their technological and economical proficiencies combined with curiosity and diverse capabilities, humans have succeeded in densely populating the Earth and taking over its resources. The extent of human activities in the world is a cause of great concern for all due to the (possibly irreversible) disruption of life-sustaining ecological systems. A look at the arguments of those working to preserve the planet and put an end to the harm inflicted upon it or upon the flora and fauna that have existed on it since the dawn of time easily shows that such arguments are, more than anything, an attempt to change human awareness and perceptions. We presume that this was always the case and that each of the transformations or innovations we mentioned above was built on a 12

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revolutionary alteration of mind-sets, the worldviews, or ideologies, of the people who led and made these changes come about. Struggle is never easy in a society that is required to address conceptual or ideological changes. Therefore, the historical process presented here should be regarded as a development that involves inspection and thought (discovery and innovation), discussion, persuasion and social struggle before all of its components are accepted and realized. As human populations grew in size and diversity, widespread changes involving perceptions, society and economy increased in their complexity while becoming much more difficult to achieve, and yet such transformations had indeed occurred. In many respects, this history of humankind can be viewed as a timeline during which people had been distancing themselves from their natural state (i.e., as natural beings, the hunter-gatherers living in nature and subsisting only on natural resources). As time passed, they became manipulative creatures, of a unique state in nature, subsisting by virtue of capabilities that are prominently and multidimensionally expressed in the material and technological arenas. They became creatures of outstanding and extraordinary cognitive capabilities. Tying the above to the word ‘domestication’, as in the title of this book, the history we present could be perceived as a chain story of domestications. The emergence of stone tools and the control over their shaping millions of years ago can figuratively be seen as the domestication or taming of stone; the discovery, control and daily use of fire can also be figuratively perceived as the domestication or taming of fire. The Agricultural Revolution can, of course, be directly related to the domestication of plants and animals, and some propose to view the building of houses and large permanent settlements as a form of domesticating space or the environment. In this context, we may mention the emergence in the Near East of flint and stone axes and adzes during the Neolithic (as of ca. 11,500 years ago or somewhat earlier), which were used for tree-felling as well as for shaping wood for construction and therefore may be regarded as the domestication of the pristine, uninhabited environment. The emergence of pyrotechnology, the use of fire as energy required for the irreversible transformation of matter (such as limestone to lime-plaster or clay to pottery vessels and their firing), could be regarded as the domestication of energy and matter. The discovery and deployment of technologies for the collection of water (such as the digging of water holes and wells and the construction of irrigation systems) could be regarded as the domestication of water. The discovery of metalcontaining ores, their quarrying and/or mining, the extraction of the metals and the formation of metal vessels and tools are all part of the (figurative) domestication of metal. And finally, the complex social systems that developed as a result of all of the above and the ‘disciplining’ of the individual within these newly established systems are also a type of domestication – humans’ self-domestication. Turning to the word ‘domestication’, a derivative of the word domus, the Latin representative of home, we can clearly see the very essence of the phenomena we introduced: adopting innovation as part of our existential home, part of the system under our governance, and a component of the foundations that facilitate and lead our existence. 1 WHAT IS THE AGRICULTURAL REVOLUTION?

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H UN T E R- G A T HE R E R S A ND F O OD - P R ODU CI NG F A R M ER S

In a broad generalization, we can say that the Agricultural Revolution expresses the life-ways of the ‘New Human’, the farmer, as compared to those of the prerevolution hunter-gatherers. Below we illustrate some of the central aspects associated with the transition from hunter-gatherer to farming life-ways as understood from both ethnographic and archaeological evidence. For hunter-gatherers this must be viewed as a conventional generic, in a way simplistic, stereotypic and maybe even a naive way of presenting the complex story and the diverse cultural and socioeconomic landscapes of hunter-gatherers,2 yet we find it necessary, at least, for some of the readers.

Nature of occupation sites and mobility patterns. Hunter-gatherers typically live in small

communities in provisional, small settlements comprised of temporary, transient structures. They often move from one site to the next following available resources in their area, their distribution and extraction methods as well as input-output considerations. Mobility patterns of such groups in their habitats are diverse, and where these sites are intact (not disturbed by modernity), settlements also appear to be similar, exhibiting no clear spatial hierarchy but rather they are spread in space according to specific considerations related to the character of the area and its resources. In contrast to hunter-gatherers, food-producing farmers are usually sedentary, and their sites adorn the landscape according to the availability of those resources necessary for agriculture, namely, land and water. Settlements exhibit permanent dwellings and installations, such as those in which agricultural produce is processed and stored. Neolithic farmer settlements are of different sizes and include local and regional settlement centres that offer ‘services’ not available at other sites or villages and which host large-scale events (extending beyond the family or local community).

Social structure and organization. Hunter-gatherers live their daily lives in small commu-

nities, typically comprising only a few dozen individuals. The social formation is fluid, allowing individuals and families to move from one community to another fairly easily, and the autonomy of individuals (children included) is an important element. Huntergatherers do not usually exhibit a permanent structure of leadership or a hierarchy related to the individual’s position in the (kin) system. In other words, the status and prestige an individual might enjoy in their community is based on daily activities as well as on gender and age (e.g., an infant, a pregnant woman or an elderly man would not be expected to take part in large-game hunting and in providing meat for the community). Therefore, sharing and mutual responsibilities, on the one hand, and autonomy of group members, on the other, are fundamental principles that guide groups of hunter-gatherers.

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Figure 1.2 Modern hunter-gatherer life-ways. (a) Hunter-gatherers building a hut; (b) gathering women.

In contrast, food-producing farmers live in large and growing communities comprised of dozens, hundreds or even thousands of individuals. Social structure is much more rigid, and social mobility is difficult and sometimes impossible. Such societies typically promote social status based on kinship or on the individual’s success in their daily life that accords positions of power, which eventually become fixed within the community.

Economy. Hunter-gatherers consume wild resources available to them and do not grow

or produce foods – they hunt animals and gather plant foods that are found near their residential area to satisfy their needs (see Box 1 Paleolithic nutrition, p. 16). They engage in a variety of activities that are targeted at food procurement, build temporary dwelling sites and produce the tools they require to carry out all of their activities (Figure 1.2). Their intervention with nature is usually minimal and local. Ethnographic research attests to various methods by which these groups adapt to different environments from the coldest to the tropical-equatorial regions. Their consumption of subsistence resources is typically immediate, or shortly deferred, as hunter-gatherers do not ordinarily store foods in a systematic, orderly manner other than under exceptional circumstances. A central food consumption principle of hunter-gatherers is the sharing of hunted food among community members. We shall return to this issue later. In contrast, food-producing farmers are manipulators who have appropriated different types of plants and animals, and grow and nurture them according to their needs. Their aim is to increase production, accumulate resources and store these for future consumption – a practice termed delayed consumption by researchers of hunter-gatherers. The social organization in agricultural villages is based on the nuclear or extended family unit. As a rule, the burden of production is not shared by the entire community although it is likely that many activities are shared and that mutual responsibility is exercised at the community level in various ways.

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Box 1 Paleolithic nutrition: The Paleo (natural) diet Recent years have seen the flourishing of a rising number of people and groups that profess awareness of the harms of productivity and the optimization of production as well as the industrial character of our foods. These groups attempt to revert to a ‘sane’ diet, as they claim, known as Paleolithic nutrition, the Paleo diet or Stone Age, caveman and ancient diet. This pre-agriculture diet is based on our long evolutionary history as hunter-gatherers (inasmuch as this is currently possible). In other words, this is the hunter-gatherers diet based on gathering food from nature. Such a diet, for individuals living in modern cultural environments, in which most of the world’s population lives, means a decrease in the consumption of processed foods and increase in the intake of raw foods, which are – these groups claim – suitable for humans due to their evolutionary history as hunter-gatherers. These groups, in all of their variations, promote the Paleo diet mostly in the virtual arena. Each such group presents different perceptions and knowledge sources, and each emphasizes a choice of select issues. Their various websites offer practical advice and many recommendations. A parallel field of alternative healthcare has also emerged that is tied to the Paleo diet. Generally speaking, we might say that the Paleo approach to health is guided by knowledge concerning the biological evolution of Homo sapiens combined with modern scientific knowledge in order to make daily choices regarding nutrition, fitness and health. Following this logic, human evolution saw a significant increase in the volume of the human brain, which could only be supported by additional bodily adaptations in order to capitalize on the advantages of the large brain. The Expensive-Tissue Hypothesis proposes that one of these adaptations was a cutback in our digestive system over the millions of years of evolution during which our brains

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expanded. This process implies that humans can no longer consume and digest large quantities of plant foods (unlike, for example, our chimpanzee relatives) and so the human body has been adapted to the high consumption of animal protein and fat. In modern times, and given the conditions of our contemporary lives, the Paleo diet advocates abstinence from cereals (especially wheat), sugar and unsaturated industrial seed oils while focusing on an intake of animal protein and fat, eggs, fruits and vegetables. According to this approach, enjoying an animal-based diet is the most natural nutritional option, and it is, therefore, also the safest for most of us. This approach obviously stands in complete contradiction to the arguments of vegetarian groups. Paleo advocates also claim that our teeth and digestive system (as well as other bodily systems) are adapted to the consumption of large quantities of meat and fat. Moreover, consuming foods that are not aligned with our evolution, so the theory goes, is the cause of many diseases in general, and in particular heart and autoimmune diseases as well as cancer. Paleo proponents further claim that if we maintain a proper diet, one to which we are evolutionarily adapted, we could avoid these diseases altogether or at least considerably reduce their prevalence. The issue of the Paleo diet, in all of its diversity, has generated immense volume on the Internet and in different media outlets. It has achieved great success in the USA (in 2011 it was awarded first place by the magazine Sharp as the nutritional fad of the year, and in 2013 the Paleolithic diet was among the most popular diets on the Google search engine). In recent years, the application of Paleo diet principles has widened to include whole grain, seed oil and sugar-free Ketogenic and Low Carbohydrate diets. Together the popularity of these terms in the Google search engine continues to rise. Since (successful) scientific experiments with this diet have only just begun, this triumph has yet to be backed by research institutes and governmental

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agencies. At the moment, then, it appears to be based on the many success stories published over the Internet. These stories typically pertain to weight loss, although some relate to the healing or relief of incurable bowel and autoimmune diseases while a few emphasize an enhancement of alertness in waking hours during the transition to the Paleo diet. The Paleo diet has reached people across the globe (the website Paleo-Israel included, https://paleo.co.il). Those who advocate Paleolithic nutrition truly believe that it will benefit our health. Proponents are also typically environmentally aware and wish to promote sustainability in our world. Their suggestions, however, are difficult to realize in

large societies on a planet inhabited by some seven to eight billion people, many of whom traditionally rely on diets that are comprised of very little animal-related foods. Basing a diet mainly on animal protein and fat requires extensive resources (including energy) as many of these animals are secondary foodproducers that feed on plants to generate their proteins (e.g., cattle eating grass or maize, and chickens pecking grain). It would only be possible to assess retrospectively the influence of the multiple, diverse nutritional trends of the present industrial age. Here, as part of our brief discussion on the subsistence and diet of hunter-gatherers, we merely wished to bring attention to their existence.

Demography. Hunter-gatherer groups tend to grow very slowly. The number of births

per woman is low, and the small number of children surviving birth and early childhood eventually leads to a stable population size or very little growth. The presence of many infants is a hindrance to the mobility of both male and female hunter-gatherers in the field and in their execution of daily economic functions. These societies therefore adopted a set of rules and behaviours that could be regarded as limiting birth rates, such as rules concerning marriage (e.g., age restriction), codes governing accessibility to fertile women and rules regarding breastfeeding and general postpartum nutrition. In contrast, food-producing farmers are sedentary and therefore infants do not interfere with their daily economic activities. On the contrary, children, including very young children, facilitate the promotion of economic needs and production; therefore an increased birth rate is an economic advantage, and the demography of the foodproducers is ‘high’, leading to a perpetually growing society. Although child mortality is still high, the increased childbirths result in significant population growth. Therefore, farming societies adopted behaviours and rules that increase childbearing. Thus, as far as the anthropological literature reveals, hunter-gatherer societies worldwide maintain very low demographic growth and live with a very low density of people per square kilometre, while the food-producing farming populations have always proliferated at a growing rate, and still do. The intensified production of food-producing farmers was primarily ‘invested’, in a very Malthusian way (which some would call the Malthusian trap), in demographic growth. This has accompanied humans around the world until the very recent past, when modernity shifted to investing profits in the welfare and quality of life of individuals.

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Worldviews. The hunter-gatherer ethos can be summarized in two central themes:

human-to-world relations and human-to-human relations. Human–world relations include the former’s approach towards their environment and elements of nature, and particularly towards resources available for their subsistence as well as the place of humans in this world. Hunter-gatherers typically exhibit a positive attitude in their approach to the world. The work of Israeli anthropologist Nurit Bird-David shows that for huntergatherers, the world, or nature, is a giving environment; they consider it a first-degree relative committed to their subsistence and treat it as such. In this kind of world, there is no need to hoard or produce food as all human needs can be met through available natural resources. Moreover, this first-degree relative (the tree, bush, animal, river, spring, etc.) can always be counted on to provide needs tomorrow just as it did yesterday and today. This approach also results in a high degree of freedom in the mobility of individuals and groups over the terrain. Naturally, this understanding necessitates a mutual relationship, in which humankind is committed to both refraining from harming the world or the environment and to maintaining the equilibrium required for its further existence. This relationship is one of trust, which is reflected in stories of hunter-gatherers about animal hunting. The human-to-human relationship reflects the community’s self-perception and the underlying worldview that determines its behavioural code with respect to the sharing of hunted food among community members. In contrast, glorifying production, the food-producing farmer groups have adopted a confrontational approach towards the world and their environment. Their lives are a daily struggle as they toil on the land, much as described in the biblical story of man’s banishment from the Garden of Eden (see Box 2 From the fruit of Paradise. . ., p. 19). Food-producing farmers live in this world as though they are constantly battling for their existence, perpetually fearing a world that might harm their ability to secure, grow and expand production. Their worldview is therefore devoid of an inherent component of sustainability, preservation of natural resources and safeguarding of the delicate equilibrium between nature’s many components, humans included. This relationship lacks trust, and, in its essence, it expresses elements of domination (objectification). In fact, this is the guiding principle of the modern age – ever-increasing growth to the point of maximizing production and profits as well as agricultural (and later industrial) output. This relationship is not meant to achieve equality between different humans. Instead, it advocates power games and social ranking, which are increasingly taking hold within it. These are political moves meant to enforce the control of the strong over the weak. Eventually, this relationship culminated in the appearance of ranked, and later stratified, societies, governed and controlled by powerful elite groups. In the next chapter we will discuss the expression in archaeological findings of these significant differences between groups of hunter-gatherers and food-producing farmers. We will briefly present selected aspects of the vast knowledge accumulated within the sphere of archaeological research that is relevant to the transition from the life-ways of the hunter-gatherers to those of food-producing societies in the Near East – that is, we

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Box 2 From the fruit of Paradise to ‘by the sweat of your face you will eat your food’ The Bible describes Paradise, the Garden of Eden, as a place where ‘every tree that is pleasant to the sight, and good for food [grows] . . . And a river went out of Eden to water the garden’ (Genesis 2:9–10, King James Bible (KJB)). Because they ate from the forbidden tree of knowledge, Adam and Eve were banished from the garden and cursed: ‘cursed is the ground for your sake; in sorrow shall you eat of it all the days of your life. Thorns also and thistles shall it bring forth to you; and you shall eat the herb of the field. In the sweat of your face shall you eat bread, till you return to the ground’ (Genesis 3:17–19). Using the terminology that we introduced in this book, we might suggest that in Paradise, Adam and Eve were given all that was required

for their subsistence. Having sinned, however, they were forced to move to a world where the land yields thorns and thistles, and only by the sweat of their brow would they (and we, subsequently) eat food. The story of Adam and Eve goes beyond its biblical tradition and faithrelated message; it could easily be read as the transition from the life-ways of huntergatherers to a life based on farming. We believe that it is quite likely that the biblical story of Paradise echoes an ancient tradition relating to the ‘old’ world and its ‘giving environment’ in the days when humans were not required to grow their own foods. Water resources (rivers) and the diverse natural resources of our suggested core area in southeastern Turkey and northern Syria allow us to further build on this notion and picture the area that lies between the Euphrates and the Tigris as part of the biblical Garden of Eden.

will describe the last of the hunter-gatherers (see Chapter 1, note 2, p. 240) and the first of the farmers in this region. We reiterate that such knowledge was obtained in each of the domestication centres of the world but herein we focus on the Near East.

K E Y PO IN TS A ND BE Y ON D

• As we look at the multi-million-year history of humankind we see a series of transformations – significant changes in behaviour as well as in the structure and organization of human societies. The Agricultural Revolution is but one of these transformations, albeit a most prominent one. The Agricultural Revolution and the ensuing emergence of food production were the most significant changes that led to the deepening of the gap and further alienation between humans and nature. This gap has continued to grow over the last ten millennia and is probably climaxing now, in our own times. • The life-ways and daily activities of hunter-gatherers significantly differ from those of the food-producing farmers. These differences are reflected in many areas of life: the nature and character of dwelling sites, mobility patterns, social structure and organization, economy, demographics and worldviews (ideology). Worldviews of the huntergatherers are founded on elements of sustainability, equilibrium and a trusting relationship with nature, whereas worldviews of the food-producing farmers are confrontational, feeding off the human need to control and maximize the resources they need or produce. Archaeological evidence shows that Near Eastern hunter-

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•

•

•

•

gatherers at the end of the Ice Age had already sprouted buds of change in aspects related to the character and size of dwellings, mobility patterns, social order and probably demographics. At the turn of the Neolithic period some 11,800/700 years ago, in what is known as the Pre-Pottery Neolithic A period, we seem to see additional changes in scale and site characteristics. This included monumental (round) architecture and a rich symbolic sphere of Near Eastern communities. At the same time, the model of subsistence remained essentially unchanged, showing no sign of domestication. This growth represents a climax of sorts of hunter-gatherer societies and may have caused inevitable social tensions and instability that could, at least partly, have brought about further change in their worldviews. The economic change involving the domestication of plants and animals and food production emerged a short while later, some 10,500 years ago, with the beginning of what is known as the Pre-Pottery Neolithic B period. It was accompanied by yet additional changes, including the move from round to rectangular architecture, new lithic (flint) technologies and a new symbolism with a growing emphasis on human representations. Domestications – that is, the human ability to take control of natural species and adapt them to their new desires – brought about a transformation, a different and innovative relationship between culture and nature. This new relationship effected an irreversible distancing from the previous hunter-gatherer worldviews, leading to a growing rift between human and nature to the unprecedented point of alienation with which we are currently familiar. As a subsistence model, the hunter-gatherer socio-economic system is still a valid alternative, although it is practised in very few areas of the world, typically those that remain ‘unconquered’ by food-producing civilizations. Moreover, in vast areas, for example in Australia, the economy was based on hunting, gathering and traditional resource management (founded on trust while lacking an element of domination) until the encounter with Western civilizations. We draw our generalizations from the fields of ethnography and social anthropology on the one hand, and the archaeology and prehistory of hunter-gatherers and early farmers on the other. The mutual relationship between the two above-mentioned research disciplines requires the establishment of a theoretical and methodological framework, which we do not elaborate upon here. We merely note, then, that we use both these disciplines as sources for data analysis and various interpretations. Notably, the history of hunter-gatherers in the last few millennia throughout the world is complex and has included interrelationships with and has been influenced by foodproducers. Hunter-gatherers may have even experienced historical chapters when they became food-producers themselves.

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2 FROM HUNTER-GATHERERS TO FARMERS IN TH E N EAR E AST Archaeological Background

The area in which the domestication of plants – the Neolithic crop package with which we are concerned – and animals took place is known as the Fertile Crescent (Figure 2.1, and see Chapter 4), the Levant or the Near East. These terms differ in source and meaning according to the scientific or geographic context in which they were first introduced. For our purposes, however, we will not delve any further into this issue and will alternate between the terms based on which is most prevalent in the academic literature of each topic. The map of the area in Figure 2.1 details archaeological sites of the period located in the northern Levant (northern Syria and south-eastern Turkey) that are relevant for our understanding of plant domestication. For additional context, we also note sites from the central and southern Levant, an area that includes Israel, Jordan and the Palestinian Authority and small portions of Lebanon and southern Syria. We briefly discuss a selection of these and other sites in the boxes that accompany the chapter in order to provide some background information on the societies that were involved in plant domestication, their settlements and the general landscape of their daily lives. We begin our archaeological survey of the region with the Epipaleolithic period (the end of the Old Stone Age), which began some 23,000 years ago and ended around 11,800/700 years ago with the advent of the Neolithic period (the New Stone Age). The most prominent characteristic of the Epipaleolithic is its minuscule flint tools, known as microliths (Figure 2.2). Epipaleolithic sites are spread across Israel, including its Negev Desert area as well as Jordan, Syria and Lebanon. They have been assigned to different cultures by their material culture, mainly the various types of flint tools, and were given additional local names throughout the southern Levant (Figure 2.3). This is a

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Bla

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Figure 2.1 Map of the Fertile Crescent, illustrated by Itamar Ben Ezra. 1. Çayönü, 2. Mureybet, 3. Tell Abu Hureyra, 4. Tell Aswad, 5. Jericho. A colour version of this map can be found at www.cambridge .org/abbo-gopher.

well-known period in Israel comprising several local cultures, including the Kebaran (named after Kebara Cave on Mount Carmel in northern Israel), the Geometric Kebaran and the Natufian (named after Wadi an-Natuf, which descends from Samaria into the Mediterranean through the Yarqon River).1 The early Epipaleolithic climate was cold and dry, the culmination of the last Ice Age in this area (the Last Glacial Maximum); later in the period, the climate became milder. Climatic shifts characterized the end of the Ice Age, and with the advent of the current era, the Holocene (or Anthropocene; see Introduction, note 5, p. 239), in the midtwelfth millennium cal BP, the landscape and the climate regime stabilized to what is known today. Multiple and prominent changes in the landscape that occurred from that time onwards can be attributed to humans.

T H E E P I P A L E O L I T H I C PE R I O D

Epipaleolithic settlements in Israel and in neighbouring countries and the archaeological record of their cultures have been researched and analysed; the settlements all exemplify

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Figure 2.2 Kebaran and Geometric Kebaran minuscule flint tools (microlithics); note the scale the tools on the right – courtesy of Ofer Bar-Yosef.

the archaeological presence of groups of hunter-gatherers at the end of the Ice Age. These sites are typically open-air, small and shallow, indicating they were short-lived. A few sites are exceptionally large and deep, attesting to longer use: several hundred square metres or even a few thousand square metres like Kharane IV in Jordan, Ohalo II and Nahal Ḥadera V in Israel (see Box 3 for Ohalo II, p. 25, and Box 4 for Nahal Ḥadera _ _ V, p. 27). It is unclear, however, if the entire range of these sites was occupied concurrently. Most sites of the Epipaleolithic cultures were small open-air sites situated in open landscapes and from above they would look like small specks, short-lived and abandoned. Sites attesting to continuous, long-term residence are rare, although it was suggested for some sites that they were year-round sites, for example Ohalo II (see Box 3 Ohalo II, p. 25). These Epipaleolithic sites typically have no permanent stone-built structures; some boast temporary hut-like structures,2 and, in rare cases, structures had stone foundations. These structures were usually round or amorphous; with a few installations, mostly relating to fire, within and around them. Prominent among the findings at these sites are large assemblages of flint tools as well as pounding and grinding tools of basalt and limestone, accompanied by a few bone tools. Remains of hunted animals at these sites include deer and gazelle as well as smaller animals. The site of Ohalo II, on the shore of the Sea of Galilee, Israel, also contained large quantities of fish bones (Box 3 Ohalo II, p. 25).

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Information concerning vegetal foods at Epipaleolithic sites is rare. Fortunately, one site, that of Ohalo II (Box 3 Ohalo II, p. 25), was well-preserved due to the anaerobic conditions beneath the water level of the Sea of Galilee. Over 100 plant species were identified by Professor Ehud Weiss that were used as food plants (including cereals, legumes, grapes and various other fruits), to build huts and for diverse purposes. Kebaran and Geometric Kebaran sites have not yielded many additional floral remains. The small number of burials and human remains is too modest to offer any insight into the demography and health conditions of these populations. Burial offerings were sometimes found next to buried humans, such as stone pounding tools. Other findings include body ornaments made especially of processed molluscs and some rare cases of engraved stone plates (Figure 2.6). This and additional evidence attest to a generally homogeneous system of mobile communities that resided for short (or occasionally long) periods of time in small sites, subsisting on the hunting and gathering of local resources. No evidence was found of social ranking or any other kind of social differentiation among the inhabitants of these sites. Although it is difficult to trace the worldviews of these populations, it is clear that they were not food-producers. It is therefore reasonable to assume, based on anthropological data, that their culture, worldviews and ideological landscape resembled that which is known in recent small, mobile hunter-gatherer communities. Some 15,000 years ago, with the emergence of the Natufian culture in Israel, a significant change transpired in the archaeological landscape. The Natufian is a local

Figure 2.3 Chrono-cultural timeline of the end of the Pleistocene and early Holocene; the number axis represents thousands of years before our time; illustrated by Adi Halpern.

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Box 3 Ohalo II: An Early Epipaleolithic site on the western bed of the Sea of Galilee (Lake Kinneret, Israel) Ohalo II is an early Epipaleolithic site dated to ca. 23,000 years ago based on an extended

series of radiocarbon (14C) measurements. It is thus the earliest site assigned to the Epipaleolithic in the Near East and researchers currently consider it a site of the Mazraqan culture pre-dating the Kebaran culture. Its

Figure 2.4 The site of Ohalo II. (a) The banks of the Sea of Galilee and the site of Ohalo II, view towards the north-west (photographed by Arik Baltinaashter); (b), (c) and (d) the huts (dark spots) found at the site (the scale in (c) is of 1 m); (e) reconstruction of an Ohalo II hut; (f ) hut plan (note the hearth at its centre and the weed cover near its edges), with a section of the hut illustrated at the bottom of this image; (g) flint tools, particularly microliths; (h) shaped stone weight (possibly used in net-fishing); (i) points made of animal bones; (j) burial in supine position; (k) engraving on bone tools. Illustrations and photographs – courtesy of Dani Nadel.

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main contribution in our context is its yield of microlithic assemblages and rich, well-preserved archaeobotanical assemblages (plant remains found in archaeological sites). The site was discovered at a time when the low lake level allowed surveying previously submerged beach areas. The site was discovered in 1989 and thoroughly excavated until the year 2001 by Professor Dani Nadel. It is located on the south-western bank of the Sea of Galilee, near Beth Yerach. A short time after the site was abandoned by its inhabitants

it was covered by the waters of the Sea of Galilee (and clay deposited in the lake) and remained submerged until today. Occasionally, when the water level in the Sea of Galilee drops considerably, the site is exposed. The site spans some 2,000 sq m, boasting several round and oval-shaped brush huts, some of which are 3–4.5 m in length (Figure 2.4c–f). The floors of these huts, made of compacted mud, are slightly below the surface. The huts were built of plant materials only, comprising tree

Figure 2.4

(cont.)

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branches of oak, terebinth, tamarisk, shrubs and weeds; no stones were used. Sleeping areas could be detected (including ‘mattresses’ (bedding) made of plant materials), as were areas devoted to cooking and to food consumption. Installations were found near the huts, in particular, fireplaces. Burial of a single person was found in a shallow pit near the huts and additional remains of one other person were preserved less well. Burials at Epipaleolithic sites are rare and known only at a small number of sites in Israel and Jordan. The rich, well-preserved assemblage of botanical findings comprises over 100 plant species, including cereals, legumes, fruits, various grasses and additional plants. Remains of hunted game include bones of fallow deer,

pig, aurochs and, most prominently, gazelle, many fish (mainly carp and tilapia) and water fowl (loon, duck, goose, coot and others). Additional findings include stone tools for grinding and pounding as well as other activities (such as possible stone weights for net-fishing), a multitude of flint tools, the most prominent of which are the microliths (small flint tools made on narrow blades known as bladelets), bone and wood tools, and also ornaments, including molluscs, dentalium and rustic dove shells. This site appears to have served a group of huntergatherers for many months during the year, and judging by the variety of plants and animals, perhaps even throughout the year.

Box 4 Nahal Ḥadera V: A Kebaran site on the Israeli_ coastal plain

partially subterranean, built ca. 1 m into the ground (Figure 2.5j). Surrounding the hut were many animal bones, grinding and pounding stone tools, and many thousands of flint tools (and flint industry debris) that were used for hunting, butchering and food preparation. Flint tools include scrapers, burins, shaped flakes and blades and massive tools, but are dominated by microliths within which the most conspicuous are types of points and obliquely truncated bladelets (known as Kebara points) (see Figure 2.5i). Inhabitants of the site had been hunter-gatherers who subsisted mostly on hunted gazelle and fallow deer slightly supplemented by equids, aurochs, deer, turtles as well as predators such as fox, wolf, wild cat, lion and hyena. Plant-based nutrition is unknown as no such foods were preserved. Among other findings were molluscs and small masses of ochre, a red-brown earth pigment.

The site of Nahal Ḥadera V was first excavated _ by Ofer Bar-Yosef and Earl Saxon in the early 1970s and then later by Avi Gopher and Ran Barkai at the end of the 1990s. The site is located less than 1 km from the current shores of the Mediterranean Sea on the southern bank of the Ḥadera Stream (Nahal in Hebrew) _ and some 40 km north of Tel Aviv. The site spans around 500 sq m and is about 20,000–18,000 years old. This relatively large site was part of an array of extensive Kebaran settlements on the coastal plain of Israel. The site was assigned to the Kebaran culture based on its flint tool findings, including minuscule flint tools – microliths – which are typical of the period. In the highest area of the site, situated within red-brown soil layers that were covered by young sand, a nearly rectangular hut approximately 6 m long  4 m wide was uncovered. The hut was

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Figure 2.5 The site of Nahal Ḥadera V. (a) The site prior to excavation in the late 1990s, view _ facing north-west; (b) the site prior to excavation in the early 1970s, view facing north-northwest, towards the Mediterranean Sea; (c) the 1990s excavation, view facing north-east; (d, f) views towards levels rich in faunal remains outside the hut; (e) kurkar (solidified dunes) slabs at the base of the hut (dark layers are the interior of the structure); (g) a short and wide limestone pestle; (h) bones (vertebrae) of a large animal in articulation, in situ; (i) flint microliths, especially obliquely truncated bladelets; (j) plan of the hut (drawing: Karin Raneberg-White). Colour versions of these images can be found at www.cambridge.org/abbo-gopher.

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Figure 2.5

(cont.) culture whose ‘homeland’ is in the Mediterranean zones of Israel (Judea, Carmel, Galilee; although sites were also found in the northern Negev and the Negev Mountains). Natufian sites are spread across the Near East, from Israel and Jordan through Syria and the Zagros Mountains to the east. It is commonly assumed that this was the culture that planted the first seeds of the Agricultural Revolution. In areas where research on the Natufian culture is fairly intensive (mostly Israel and Jordan), this culture boasts considerably larger sites than those that preceded it and contains many densely placed round or oval stone structures. The presence of smaller Natufian sites resembling the preceding sites of Kebaran and Geometric Kebaran cultures is limited. The Natufian, then, is characterized by relatively large, home-base camps or sites, some of which appear to have been permanent, alongside short-lived temporary sites in which specific resource-procuring activities took place on a seasonal or task-specific basis. Thus, the Natufian model of residence included both permanent and ephemeral sites, and the movement between them is known as logistic mobility, that is, inhabitants would leave the permanent home-base in order to procure resources or conduct specific short-term activities, and would then return. This model diverges considerably from the classic hunter-gatherer model of residential mobility, in which the entire community moved from one place to the next based on seasonal and economic consideration of inputs and outputs. The presence of animals such as the house sparrow (Passer domesticus) or the

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Figure 2.6 Engraved stone plate from the site Urkan e-Rub, a Kebaran site in the Jordan Valley. Photographed by Zeev Radovan – courtesy of Erella Hovers.

house mouse (Mus musculus) in Natufian settlements attests to permanent and continuous residence at the sites throughout the year and over many years. This argument was recently extended to include sedentary settlements starting at the onset of the Natufian culture based on the analysis of bone assemblages of hunted animals at the Natufian site of el-Wad Cave, located in Wadi el-Mughara (Arabic) or Nahal Me’arot (Cave Creek, in _ Hebrew) on Mount Carmel. Thus, small communities settled in permanent residences as early as the emergence of the Natufian. This transition to sedentary settlements bears significant implications for our understanding of the steps that led to the Agricultural Revolution. The large (home-base) Natufian sites may span thousands of square metres. They are often open-air sites such as ‘Eynan (‘Ain Mallaha) in the Ḥula Valley (Box 5 _ ‘Eynan, p. 31) or the site of Wadi Ḥammeh 27, located to the east of the Jordan River. Other sites are located on cave fronts and within the caves, such as Ha-Yonim Terrace in the Western Galilee or the caves of el-Wad and Kebara on the Carmel Ridge. Some outstanding Natufian sites were limited to a single purpose, such as the Galilean Ḥilazon (Snail, in Hebrew) Cave that was used mostly for burial. Some Natufian sites were found in basal layers of large, multi-layered archaeological mounds (tell sites) that remained inhabited subsequent to the Natufian, mainly during the Neolithic period, such as the site of Jericho in the Jordan Valley or the Syrian sites of Tell Mureybet and Tell Abu Hureyra on the banks of the Middle Euphrates (Figure 2.8). These home-base Natufian sites are rich in findings that are absent from earlier sites. For example, round, stone-built structures were found at these sites that were likely to have been covered by a superstructure made of wood and other organic matter. Stonebuilt installations deck these sites within and between structures, some of which resemble stone-lined silos or granaries (Figure 2.9). Large quantities of flint and stone

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Box 5 ‘Eynan: A Natufian site in the Ḥula Valley, northern Israel The Natufian site of ‘Eynan (‘Ain Mallaha _ in Arabic) was discovered in 1954 and was first excavated by Jean Perrot in the 1950s and 1960s. In the 1970s, Monique Lechevallier and François Valla joined the excavating team. The site was excavated again by François Valla and Dr Hamoudi Khalaily in the 1990s and early 2000s. The site is located on a north-northeastern slope near the spring of ‘Eynan (Mallaha), _ which is one of the most affluent springs in the area, at the edge of Ḥula Lake. The site spans some 2 dunams (2,000 sq m) and is up to 3 m in depth. All of its layers have been dated to 15,000–11,800/ 700 years ago and assigned to the Natufian culture. All layers of this multi-layered site contain stone-built structures, a richly diverse assortment of findings, and overall about 100 human burials were found at the site. The structures are round or semicircular and are located on several terraces partially dug into the slope. In early phases, structures spanned up to about 9 m in diameter, but over time they reduced in size to 3 m. Structures appear to have had a supporting stone wall against the slope while the superstructure was built of wood and other plant materials. Some structures exhibited stonelined and stone-paved pits used as postholes, holding wooden poles that supported the superstructure (Figure 2.7c and see reconstruction in g). Various installations were found within and between structures, including hearths, ovens and stone-lined storage installations. In one of the early structures, a plaster-covered bench was found along the wall. The numerous burials found throughout site layers were located both within and between residential structures as well as in separate areas dedicated to burials. Group

burials were found among the burials. These included adults and children, men and women. Most of the buried were placed in flexed or semi-flexed positions. Some burials were adorned with marine shell and bone body ornaments, similar to burials at other Natufian sites, including that of el-Wad Cave on Mount Carmel, Israel. An exceptional burial was that of a woman and a dog placed side by side, presumably attesting to the deep integration of dogs in human societies; the dog was probably domesticated. A similar finding was uncovered in the front terrace of the Natufian site of Ha-Yonim (the Doves, in Hebrew) Terrace in the Western Galilee in Israel. The site of ‘Eynan is very rich in finds. Among the flint tools, microliths are prominent, especially lunates. It yielded many stone tools used for food (and other material) processing, including large, well-shaped basalt mortars and pestles as well as other grinding tools (Figure 2.7d). Many bone tools were found at the site such as points and spatulae as well as tools made of antler such as sickle hafts. Faunal assemblages include many gazelles in addition to deer, fallow deer, boars, aurochs and a great variety of hunted birds, some of which were migrating birds passing through on their way from Europe to Africa or back. Site residents subsisted on intensive hunting and gathering in their nearby, rich environment (plant foods as well as resources related to the lake and the swamp in the Ḥula Valley). Other than the single dog mentioned earlier, no evidence was found of domesticated animals or plants. Art and body ornament assemblages at ‘Eynan are large and diverse compared to earlier periods, and they include stone figures or figurines of animals and schematically shaped anthropomorphic (human) imagery. A large number of molluscs and bone beads were also uncovered at the site. Molluscs recovered at the site attest to ties to the Mediterranean area. Similarly, obsidian artefacts – artefacts made of volcanic glass

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Figure 2.7 The site of ‘Eynan. (a) View eastward of the site towards Highway 90 south of Kiriyat Shmona; (b) a round structure boasting a mortar and hearth on its floor; (c) the complex of structures 51–131 during excavation; (d) basalt mortars and a pestle (note, black and white scale is 30 cm); (e) large flint tool known as a pick; (f, h) round and semi-round structures exposed during the excavation works of François Valla and Hamoudi Khalaily in the 1990s; (g) suggested reconstruction of a semiround hut (following François Valla). Illustrations and photographs – courtesy of François Valla and Marjolain Barazani. originating in Turkey – attest to ties and exchange in a fairly large geographic area as early as the Late, or final, Natufian. It seems that the community residing at the Natufian site of ‘Eynan was larger than earlier communities, even if the site was not inhabited in its entirety at all points of time

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during its existence. The supporting evidence indicates that ‘Eynan appears to have been a sedentary community that spent most of the year, if not all of it, at the site. As a sedentary settlement, ‘Eynan is highly consequential in understanding subsequent steps that led to the Agricultural Revolution.

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Figure 2.7

(cont.) tools typically characterize these sites, including flint sickle blades hafted in bone, antler or wooden handles used to harvest cereals, pounding tools (mortars and pestles) and basalt or limestone grinding tools. Additional characteristic findings include animal bones, bone tools, burials, human skeletal remains (Figure 2.10), body ornaments made of sea shells, bone or stone and imagery items – mostly

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Figure 2.8 Natufian layer 1 at the Syrian site of Tell Abu Hureyra, on the banks of the Euphrates; two views of round pits (structures?) – courtesy of Andrew Moore and Oxford University Press, New York.

Figure 2.9 Stone-lined installation (70 cm in diameter, 40 cm in depth) from the Natufian site of HaYonim Terrace in the Western Galilee – courtesy of Francois Valla.

zoomorphic (Figure 2.11) but also some anthropomorphic. The change in the extent of artistic-symbolic findings and the prevalence of zoomorphic representations, which were typical of earlier times, are a testament to an essentially Paleolithic (hunter-gatherers) worldview. The emergence of anthropomorphic figures (mostly of unclear gender) attests to the beginning of a highly significant perceptual change (see below, Chapter 3).

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Figure 2.10 Natufian burials and human remains. (a) Flexed position burial next to a large limestone mortar; (b) burial nearby a stone installation and hearth enclosed by flat stones, both from the Natufian site of Nahal Oren on the Carmel Ridge; (c) burial boasting an adorned hip area at Ha-Yonim Cave, the_ Western Galilee – courtesy of Ofer Bar-Yosef; (d)–(e) skeletons adorned by marine shells from Natufian layers at el-Wad Cave on the Carmel Ridge.

The Natufian economy was based on hunted animals, typically gazelle, alongside deer, aurochs, boar and small animals such as turtles, and predators. Natufians appear to have domesticated the dog, as suggested by the dog bones found in two different human burials at Israeli sites, Ha-Yonim Terrace and ‘Eynan. The little knowledge accumulated regarding Natufian plant foods, particularly at early phases of this culture, seems to indicate that the Natufian people had gathered cereals and legumes. Recent publications regarding botanical remains at Syrian Natufian sites indeed verify this notion while also revealing the use of fruit such as almonds and figs. Towards the end of the Natufian, the climate turned relatively dry and cool for about a millennium. This is known as the Younger Dryas event, and it has been identified in climatic reconstructions conducted in Europe, mostly its central and

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Figure 2.11 Assortment of Natufian animal and human imagery items, decorated stone and bone tools, and body ornaments. (a) Reconstruction of a necklace made of polished, drilled bone items and shells, el-Wad Cave; (b) beads made of shells, stone and polished drilled teeth, el-Wad Cave; (c) turtle, ‘Eynan, Ḥula Valley; (d) a gazelle figure, Judean Desert; (e) representation of a human head, el-Wad Cave; (f ) assortment of shaped stone items, probably human representations, and decorated vessels and stones, ‘Eynan; (g) illustration and photograph of the end of an animal-shaped sickle haft from Kebara Cave (collection of the Israel Antiquities Authority, © Israel Museum, zoom photograph by David Harris, illustration by Florica Weiner; courtesy of the Israel Museum).

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Figure 2.11

(cont.)

northern areas. Although there is no clear evidence for significant modifications in the floral landscape or faunal populations of the Near East, many researchers suggest that the Younger Dryas caused some environmental stress, forcing the Natufian people to revert to ephemeral settlement and mobility patterns that allowed them to continue and subsist in the dwindling landscape. This proposition also gains support from the analysis of bone assemblages of hunted animal species and their age profiles. This environmental deterioration is portrayed by many researchers as a key factor in the cultural transformation of the Natufian culture and the emergence of Neolithic cultures, and, eventually, the Agricultural Revolution. Researchers such as Professor Gordon Hillman suggested that during the Younger Dryas, late Levantine (northern Syrian) Natufian communities began cultivating wild plants. Hillman argued that some plants were even domesticated in the late Natufian, but most researchers, ourselves included, disagree with this proposition. All in all, the size of the permanent home-base, the general layout of the Natufian settlement and recent findings attesting to designated burial sites collectively suggest that this culture was significantly different from entities preceding it. While this is still a predominantly hunter-gatherer society, it appears to have been organized in larger communities throughout long parts of the year (perhaps year round) in sedentary home-base settlements. Certainly, this innovation is highly significant when compared

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to earlier Epipaleolithic cultures, a possible precursor of the next phase. We reiterate that the Natufian seems to indicate sedentary settlements, a major point in the discussion of the Agricultural Revolution. T H E N EO L I T H I C P E R I O D

The Neolithic period began somewhat after 12,000 years ago, and it is subdivided in the southern Levant into the Pre-Pottery Neolithic (PPN, some 11,800/700–8,500 years ago) and the Pottery Neolithic (PN, some 8,500–6,500 years ago). The sequence is similar in the northern Levant, although pottery appeared slightly earlier, around 9,000 years ago. Each of these periods is further subdivided into subperiods (e.g., Pre-Pottery Neolithic A, Pre-Pottery Neolithic B, Pre-Pottery Neolithic C – see Figure 2.3). Periods and subperiods are characterized by cultural entities, each defined by uniquely recurring material culture assemblages found in well-defined territories and time ranges; these are known as archaeological cultures. Several main cultures coexisted during the first part of the PPN, known as the Pre-Pottery Neolithic A (PPNA). These include the Khiamian, the Sultanian and the Mureybetian cultures, typically named after the first site in which they were identified. Sites of these cultures, in both the southern and northern Levant, especially the Khiamian, are sparse; our knowledge is limited to the few sites, some of which are impressive in scope and findings, that have been excavated and whose findings have been published. Changes that distinguish sites of the early Pre-Pottery Neolithic A Khiamian (named after the el-Khiam Rockshelter in the Judean Desert; see Box 7 Mureybet, p. 41) from their Natufian predecessors are not highly conspicuous. Based on the paucity of evidence we have, it appears that the Khiamian sites are considerably larger than the Natufian ones. The few structures of which we are aware were round and built of stone with the innovative use of sun-dried bricks. PPNA flint tools boasted new manufacturing techniques as well as new tool types, the most prominent of which is the arrowhead (earlier periods used a variety of flint points for hunting); and new types of sickle blades also appeared, used in cereal harvesting. The Khiamian lifestyle, which attests to a huntinggathering subsistence economy, is reflected in only a few sites located in Israel, Jordan and Syria. As noted earlier, some researchers also argue that wild food plants were grown at this time in fields or in small plots and that animal domestication (other than the dog) had already begun at this early time. The next noticeable PPNA culture is the Sultanian culture, named after the nearby spring ‘Ain as-Sultan (early Jericho), in the Jordan Valley. Sultanian sites were found mostly in Israel, Jordan and Syria – in Mediterranean and slightly drier areas. Sultanian sites are considerably larger than their Natufian predecessors, spanning tens of thousands of square metres: 20, 30 or 40 dunams (2–4 hectares) as in Jericho (see Box 6) or Netiv Hagdud (see Box 8) in the Jordan Valley, or the Syrian Mureybetian culture of Tell Mureybet (we discuss sites of the Mureybetian culture along with Sultanian ones as they

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Box 6 Jericho: A Pre-Pottery Neolithic A ‘city’ in the Lower Jordan Valley Jericho is one of the oldest excavation sites in the region. It was excavated several times during the nineteenth and twentieth

a

centuries: first by a British team directed by Sir Charles Warren (1868), then by a German team directed by Ernest Sellin and Carl Watzinger (1907–1909), followed by another British team directed by John Garstang (1930–1936), and, finally, by the

b

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possible course of Proto-Neolithic wall and ditch

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Figure 2.12 The site of Jericho. (a) Tell Jericho today, view towards south-southwest; (b) the Tell’s plan, including excavation areas of Kathleen Kenyon; (c) view towards deep excavation sections in Area D at the west of the Tell; (d) the PPNA round tower and wall of Jericho; (e) a section of the PPNA tower and wall of Jericho.

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British team of Kathleen Kenyon (1952–1958). The site of Jericho is situated in the Jordan Valley, some 10 km north of the Dead Sea; it is located in a rich oasis in which ‘Ain as-Sultan, the most abundant spring of the Jordan Valley, flows. It is where Kenyon coined the terms Pre-Pottery Neolithic A and B, Pottery Neolithic A, etc. The multi-layered mound, or tell, is over 20 m high and very rich in finds throughout its layers, starting with a Natufian presence at its base, and followed by the PPNA, the PPNB and Pottery Neolithic A, and ending with the Pottery Neolithic B and the subsequent Early Bronze Age and Iron Age layers. We shall discuss only the PPNA layers of this site, which were assigned to the Sultanian culture (named after the nearby spring). The site’s PPNA layers were radiocarbon dated to ca. 11,800/700 through ca. 10,400 years ago and it spans some 30–40 dunams (3–4 hectares) in its PPNA occupations. The PPNA site is impressive. On its western end, a well-preserved wall, over 5 m in elevation, was exposed, as was a round tower some 8 m in diameter. Both were built of stone (Figure 2.12). A dry moat was also exposed – a trench dug in the rock some 10 m wide and 2 m deep.* Other structures uncovered in the same area of the site are suggested to be large storage facilities. The excavation also exposed round structures partially embedded in the ground and built of plano-convex sun-dried mudbrick in addition to many burials hosting hundreds of skeletons. For adult burials, skulls were removed from the body and buried separately, occasionally in groups. The multitude of findings include many types of flint tools, including sickle blades, arrowheads, flint and stone

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axes, in addition to basalt and limestone pounding and grinding tools and bone tools. The PPNA subsistence economy at the site comprised the gathering of food plants and the hunting of local animals, mostly gazelle. Excavation reports and various published papers suggest that some plants at Jericho may have been domesticated (such as barley and einkorn wheat), but this claim is not sufficiently supported by the available evidence. Among the interesting substances found at Jericho were bitumen, a glue substance extracted from areas near the Dead Sea and the wadis of the Judean Desert, as well as obsidian (volcanic glass) that originated in Turkey. Despite the fact that the PPNA layer at Jericho is not rich in art findings, it yielded imagery items, figurines, decorated items and many body ornaments (beads, amulets and the like) made of stone, bone and shell. The site of Jericho appears to have been very important during the PPNA, and it is part of an array of settlements located in the Jordan Valley and its periphery, including sites such as Gilgal I and III, Salibiya IX, Netiv Hagdud, Dhra’, Wadi Faynan 16, Gesher, ‘Iraq ed-dub Cave and others. * The architectural complex of the Jericho wall-tower-moat has been the target of a number of interpretations since K. Kenyon’s original suggestion that they served as defensive fortifications. For example, O. Bar-Yosef proposed it was a defensive system against the floods from the west; D. Naveh recommended that the structure was a political symbol of communal power and territorial claim; R. Barkai and R. Liran suggested it was a monument of social and perceptual significance, the location of which was derived from astronomic considerations; and A. Ronen and D. Adler suggested the wall functioned in the ideological and magical sphere, separating the built from the unbuilt landscape.

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Box 7 Mureybet: A Pre-Pottery Neolithic A tell site in the northern Levant The site of Tell Mureybet was excavated as part of a salvage project of sites around the Middle Euphrates prior to the planned flooding of large areas due to dam construction along the river. The site was first excavated by a Dutch team directed by Maurits van Loon (1964–1965) and later by a French team directed by Jacques and Marie-Claire Cauvin (1971–1974). The site is currently flooded and inaccessible. Mureybet is a large and tall archaeological tell that spans some 30 dunams (3 hectares); it stands 5 m tall. We provide here a glimpse into its PPNA layers only, which have

primarily been assigned to the Khiamian and Mureybetian cultures. These layers are radiocarbon dated to a period between 12,000/ 11,700 and 10,500 years ago. The architecture of the early layers (assigned to the Khiamian culture) is comprised of round wood and mud structures with floors made of mud and pebbles. These structures are 6 m in diameter and were slightly embedded in the ground. Various installations were found mostly outside the structures. In the later Mureybetian layers, structures were still round in outline, made of soft limestone and mud, with floors made of dirt and pebbles. Additionally, for the first time at Mureybet, round structures were occasionally internally subdivided by straight-angled walls that

Figure 2.13 The site of Mureybet. (a) A round structure internally divided – plan and reconstruction; (b) sickle blades and two awls (Nos. 11–12 bottom right); (c) arrowheads (scale in cm); (d) three flint tools known as herminettes; scale in cm; (e) bone awls; (f ) bone spatula; (g) bone tool of unknown use; (h, i) bone combs?; (j) drilled stone items including a miniature head carved in stone (bottom right); (k) body ornaments – stone beads; figures and photographs – courtesy of JuanJosé Ibáñez.

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Figure 2.13

(cont.) created rectangular spaces as well as a veranda of sorts at the front. In the late Mureybetian layers of Jerf el Ahmar in northern Syria, clear _ rectangular architecture was discovered. Sometimes the space was subdivided into

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rooms, and occasionally there were floors lined with pebbles and stone plates (see Figure 2.16a for these structures). Some of the rooms were used as residences while others are suggested to have been used

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Figure 2.13

(cont.)

for storage. Little is known of burials at this site. The findings at the site are rich. There were many flint tools, including arrowheads, sickle blades and axe-like tools known as herminettes (see Figure 2.13d). Stone tools included grinding and pounding tools. Other tool types, such as bone tools, were also found. Khiamian body ornaments at the site included mostly beads of a large variety of materials and colours (see Figure 2.13k). Imagery items were made of soft limestone and included both human and animal figurines.

The Mureybetian layer yielded various unusual artefacts such as decorated bowls made of colourful minerals and varied stone beads. Imagery items were few, including human figurines made of clay, mostly interpreted as representations of women. Both the Khiamian and the Mureybetian layers yielded engraved stone artefacts. The site’s excavator, Jacques Cauvin, suggested that the early Neolithic art world be considered as reflecting the beginning of a process of change in the worldviews of the inhabitants. He believed that this change eventually led to a socioeconomic transition (the Agricultural

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Revolution) and the establishment of ancient religions in the Levant which centred on supernatural characters: a male deity represented by the ox and a female deity represented by female figurines. Site subsistence was based on the hunting of local animals, including equids, deer, fallow deer, wild sheep and aurochs, as well as small animals and fowl that were prominent in the Khiamian layer but declined considerably in the later Mureybetian. Plant gathering was focused mainly on cereals, legumes and fruit. In our view, claims made concerning land cultivation – growing or domesticating

plants in PPNA sites, including Mureybet – do not withstand the evidence. We believe the so-called evidence is indirect and strewn with problems that require further research before this issue is resolved – although we acknowledge that the Middle Euphrates was part of the core area of plant domestication. There was an abundance of early Neolithic sites in the area, and it was clearly an important and highly significant region with respect to the transformations that were required to endorse and sustain the Agricultural Revolution, plant domestication included.

Figure 2.14 Stone pounding and grinding tools from the sites of Jericho (a) and Netiv Hagdud (b) in the Lower Jordan Valley; some tools are 40 cm long, weighing dozens of kilograms.

coexisted in the northern Levant). Smaller sites of these cultures, spanning only a few dunams each, were also found, such as Gilgal I in the Jordan Valley (see Box 8), the sites of Dhra’, Wadi Faynan 16, el-Hemmeh and Zaharat-adh-Dhra’ 2 south-east of the Dead Sea, as were truly small sites such as Nahal Oren on Mount Carmel in Israel, Ain Darat in _

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Box 8 Netiv Hagdud and Gilgal I: Two Lower Jordan Valley Pre-Pottery Neolithic A Sultanian sites Netiv Hagdud and Gilgal I are two Pre-Pottery Neolithic A Sultanian sites located in the Jordan Valley north of the Dead Sea, some 12 km north of Jericho. Gilgal I was excavated by Tamar Noy between 1974 and 1994. Netiv Hagdud was excavated by Ofer BarYosef and Avi Gopher between 1983 and 1986. Detailed monograph summaries of these excavations were published in 1997 and 2010, respectively.{ Both these tell sites are found in the fairly enclosed Salibiya Basin, 200 m below sea level near the intersection between the Jordan Valley and the mountains to its west. Gilgal I appears to span some 5 dunams (0.5 hectare), and is some 3 m high, while Netiv Hagdud is larger, spanning some 15 dunams (1.5 hectares) and is some 4.5 m high. Gilgal I was radiocarbon dated to have been active between ca. 11,500 and 11,000 years ago while Netiv Hagdud was active shortly thereafter. The sites are surrounded by many other prehistoric sites found in the Salibiya Basin and assigned to the earlier Natufian culture as well as the Khiamian culture, which occurred just slightly earlier than the Sultanian – which is the heart of the current description. The excavation at Gilgal I exposed oval mud structures constructed over stone foundations in which several installations and many artefacts were found. These structures were 3–7 m in diameter. Netiv Hagdud boasts slightly larger oval structures (up to 10 m on their long axis) as well as smaller round structures (up to 4 m in diameter), which were made of stone and sun-dried mudbrick. Installations meant for a variety of activities were uncovered, as were storage installations. While burials at Gilgal I were few and mostly in deeper layers, Netiv Hagdud yielded

numerous burials. As in other contemporary PPNA sites, adult skulls were placed separately from the skeletons, occasionally on residential floors designated for this purpose. Assemblages collected at both sites are comparable and associated with the Sultanian culture. Among others, these assemblages include sickle blades, arrowheads, flaked flint axes, polished stone axes and unique tools known as Hagdud truncations and Gilgal truncations, which seem to have been used in the preparation of arrows. Also uncovered were rich assemblages of stone (basalt or limestone) pounding and grinding tools (see Figure 2.14) and bone tools. Subsistence at both sites was based on hunting and gathering, and no evidence was found at either site of plant or animal domestication. Site residents hunted gazelle, pig, fallow deer, fox, hare, reptiles, fresh water crabs and many fowl that probably lived nearby at seasonal shallow bodies of water. The plant assemblage of Gilgal I is comprised of oat, barley, wheat, lentil and vetch as well as fruit such as figs. The assemblage of Netiv Hagdud is slightly richer, including almonds, acorns and grapes too. Both sites yielded obsidian, imported from Turkey, quite a distance away, as well as beads made of molluscs and green stone minerals. Taken together, these findings attest to a large trade network spanning a great geographic expanse. Figurines, decorated stone items, and body ornaments such as beads and pendants made of bone and molluscs were found at both sites. These include anthropomorphic figures and a few zoomorphic items. Decorated stone items display geometric engravings (see also Figure 2.15h).

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Figure 2.15 The site of Netiv Hagdud: (a) the site (marked by the arrow) and the nearby modern water reservoir, view from the west; (b) excavation plan of the western part of the site, including structures made of stone and mudbrick; (c) oval stone structure and round mudbrick structure; (d) el-Khiam arrowheads (1–2), Hagdud truncates (3–4), awl (5), flaked flint axe (6), large sickle blade (7); (e)* small clay female figurine; (f )* head of small clay female figurine; (g) stone beads and pendants; (h) engraved stone pallet.

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Figure 2.15

(cont.)

the western part of the Judean Desert, Gesher in the Jordan Valley and the site of Iraq eddub Cave in Wadi el-Yabis east of the Jordan River. To date, no Sultanian sites have been found in the desert areas of Israel or Jordan, nor have any been found in the Transjordanian mountain areas or most of the Israeli coastal plain. Sultanian sites typically show a dense pattern of round or oval structures built of stone and sun-dried mudbrick, some of which are very large. Some of these sites boast particularly impressive structures, such as the stone-built wall and tower of Jericho (Box 6 Jericho, p. 39), and the communal building at Wadi Faynan. Outstanding in size, architecture and rich findings are the Sultanian or contemporary sites in more northern areas along the Euphrates and the Tigris in northern Syria and south-eastern Turkey (e.g., see Box 7 Mureybet, p. 41). These findings attest to an unprecedented scale of activity (see Box 9 Göbekli Tepe, p. 53). In both Israel and Jordan, archaeological assemblages from Sultanian sites boast a large variety of flint tools, including arrowheads (mostly of the el-Khiam type; see

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Figure 2.15 (cont.) The site of Gilgal I**: (i) view from the site westwards, towards the Judean Mountains; (j) excavation plan of the central area (structures marked in light colour); (k) plan and section of the structure known as Locus 11; (l) plan and photograph of the structure known as Locus 3; (m) stone and polished bone beads; (n, o) bone tools, mainly points; (p)*** stone figurine, perhaps of an animal (bird?); (q)–(r)*** anthropomorphic stone figures. Colour versions of these images can be found at www.cambridge.org/abbo-gopher.

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Figure 2.15 (cont.) * Photographed by Zeev Radovan. ** All figures and photographs – courtesy of Ofer Bar-Yosef, Nigel Goring-Morris, Anna BelferCohen and Debby Hershman. *** The Antiquities Authority collection © Israel Museum. {

For monograph summaries of the projects at these two sites see:

Bar-Yosef, O. and Gopher, A. 1997. An Early Neolithic Village in the Jordan Valley, Part I: The Archaeology of Netiv Hagdud. American School of Prehistoric Research Bulletin 43, Peabody Museum of Archaeology and Ethnography, Harvard University; and

Bar-Yosef, O., Goring Morris, A. N. and Gopher, A. 2010. Gilgal: Early Neolithic Occupations in the Lower Jordan Valley: The Excavations of Tamar Noy. Oxbow Books, Oxford on behalf of the American School of Prehistoric Research.

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Figure 2.15d) and typical sickle blades. Two innovations in these assemblages, both of which became prominent tools in the Neolithic and the Chalcolithic tool kits, are the polished stone axe and the flaked flint axe, which were used for wood-felling, chopping and other wood-processing activities. These tools were typically required to prepare wood used as pillars and beams for roofs and walls, mostly by postSultanian cultures. Sultanian subsistence was comprised mostly of the gathering of food plants and fruit as well as the hunting of animals, mostly gazelles. At sites situated in the Jordan Valley, birds of different types that relied on the Jordan River and other seasonal or permanent bodies of water were also hunted (see Box 8 Netiv Hagdud and Gilgal I, p. 45). Hunting and gathering also characterized the economy of contemporaneous cultures in the northern Levant. The Sultanian sites yielded many human burials and skeletons. The dead were buried in various structures, both above and under floors, near thresholds or in between structures, in open areas. They were buried in shallow pits, typically in primary burials (i.e., the body was buried there as a whole, in articulation). Skulls of buried adults were removed from their skeletons as was customary in the Sultanian culture of the PPNA period. Some of the structures that hosted burials and skulls had exceptional properties: for example the Jericho tower or a sunken round structure at the Syrian site of Jerf elAhmar in the Middle Euphrates (Figure 2.16a, b). _ Sultanian imagery (art) items were made of stone or clay. Among others, these assemblages included: stone pallets engraved with varying patterns and human figurines, the gender of which was usually female or ambiguous (Figure 2.17). Some prominent body ornaments included beads and pendants made of stone, bone and sea shells. One of the most impressive sites occupied during the PPNA (but not quite Sultanian or Mureybetian) is the site of Göbekli Tepe in south-eastern Turkey. The site, unique in both size and content (see Box 9 Göbekli Tepe, p. 53), also boasted later, Pre-Pottery Neolithic B layers (Figure 2.18d). Despite the significant difference between the Sultanian (and the northern Mureybetian) culture and preceding cultures, and despite the major expansion of site size and construction intensity that reflect relatively large communities occupying each site (akin to small villages), subsistence economy seems unchanged. As their Late Natufian and Khiamian predecessors, early Sultanian and Mureybetian communities continued to subsist on hunting and gathering. Claims made regarding early Neolithic communities (and late Natufians before them) who raised crops near their sites of residence as yet remain unsupported by archaeological and archaeobotanical and faunal findings.

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Figure 2.16 (a) A general view of the area of the round structure including adjacent structures; (b) round structure subdivided internally and boasting a burial over its floor – the site of Jerf el-Ahmar, _ north Syria; (c) close-up over the skeleton – courtesy of Danielle Stordeur.

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Figure 2.17 (a)–(b) Figurines from the site of Gilgal I (see Box 8 Netiv Hagdud and Gilgal I, p. 45); (c) an incised stone from the site of Gilgal I – the Israel Antiquities Authority collection © Israel Museum – courtesy of Debby Hershman and Anna Belfer-Cohen; (d) anthropomorphic stone figurine from the site of Nahal Oren – courtesy of the _ Israel Museum.

After about one thousand years of PPNA existence in the Levant, prominent changes occurred at sites assigned to a new period known as the Pre-Pottery Neolithic B. PPNB cultures bear no designated names and sites of this period deck the entire Levant including the desert areas in the southern and eastern Levant. Among the characteristics noted at PPNB sites from the very beginning of the PPNB are: 1. The emergence of rectangular structures (Figure 2.19a, b, c; and see also Box 10 ‘Ain Ghazal, p. 61, Box 11 Çayönü, Nevalı Çori and Tell Ḥalula, p. 64), replacing the circular structures characteristic of earlier periods.

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Box 9 Göbekli Tepe: A Pre-Pottery Neolithic A sacred site in south-eastern Turkey, near Urfa The site of Göbekli Tepe was discovered during an archaeological survey conducted in the 1960s. Its significance, however, was not immediately recognized, and it was only excavated starting in 1995 by the late Dr Klaus Schmidt. For nearly two decades, as excavations were in full force, it yielded findings that are considered amazing on any archaeological scale. The site is located in south-eastern Turkey, on a hill near the city of

Şanlıurfa. The site spreads across over 100 dunams (10 hectares) and its occupation sediments are 10 m in height. Site layers were radiocarbon dated to the Pre-Pottery Neolithic: PPNA and early PPNB. Our discussion focuses on the PPNA, from about 11,500 years ago onwards. Round and oval structures were found in parts of the excavation areas, yet, the major architectural findings are comprised of a series of impressive circular-curvilinear stone structures or enclosures reaching a maximal diameter of almost 30 m (see Figure 2.18b, c),

Figure 2.18 The site of Göbekli Tepe. (a) General view northwards, towards the hill upon which the site is situated; (b) the complex of round temples with their stone statues; (c) one of the sacred circular enclosures with its stone statues, bird’s-eye view; (d) an early PPNB stone structure with small T-shaped statues; (e)–(i) stone statues boasting a variety of animal figures including snakes, foxes, lions, birds and cattle; (j) threedimensional statue representing a mammal (and perhaps also a human?); (k) the head of a bird of prey made of stone; (l) relief of a reptile; (m) animal sculpture – courtesy of Klaus Schmidt. Colour versions of these images can be found at www.cambridge .org/abbo-gopher.

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Figure 2.18

(cont.) which have been presented by the site excavators as sacred structures designed for (religious) communal ceremony and ritual. Most researchers of the period concur with this interpretation. Within each of these structures, there were massive statues weighing several tons each and standing up to 5.5 m tall. Most of these statues had been integrated

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into the circumferential wall of the structure in which they were found, while a couple were placed at its centre. These statues, dubbed T-shaped statues due to their general shape, were each made of a single stone mass that had been quarried from a nearby hill and transported to the site, where it was shaped and positioned. To date, dozens of such

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statues have been found in the four excavated curvilinear enclosures. The statues are decorated, exhibiting incisions and reliefs, with a variety of animal images: foxes, aurochs, pigs, snakes, reptiles, fowl, insects, scorpions and others, as well as with geometric patterns. The two statues in the centre of Enclosure D are unique and appear to have human-like arms engraved over their sides, and hands shaped on their front side, as well. Similar, yet small and less adorned, T-shaped statues have been found in the subsequent early PrePottery Neolithic B layers at both Göbekli Tepe and the nearby site of Nevalı Çori that was also excavated in the 1990s prior to the flooding involved in the building of a dam over the Euphrates (see Box 7 Mureybet, p. 41). Göbekli Tepe also yielded many zoomorphic three-dimensional stone statues. In addition to this impressive yield, thousands of flint and stone tools, including pounding and grinding tools, were recovered. A significant number of animal bones was also found at the site, including bones made into tools. In contrast, however, the botanical assemblage is rather meagre, including wheat, barley, lentil, vetch, pea, broad (faba) bean and possibly also chickpea. The subsistence economy at this huge site during the Pre-Pottery Neolithic A was based on hunting and gathering, with no evidence of plant or animal domestication. Based on the excavators’ estimate that this site did not serve for permanent residency, it appears to provide impressive, large-scale evidence of a

site designated for mass congregation to which PPNA pilgrims assembled for ceremonial and ritualistic purposes. The excavators claim that these sacred compounds, in which tremendous effort was invested, went out of use within a short time and were covered in dirt by the site users themselves – that is, they were ‘buried’. If we accept the excavators’ interpretation of this site and the significance of its highly adorned circular enclosures, the site attests to a complex, potent society of hunter-gatherers that had excellent engineering and management skills, which allowed not only for the construction of this site but also for its use. Whether or not one accepts the excavators’ interpretation, the early layers of this site represent an exceptional climax that attests to the high competence of the last huntergatherers in the area. Since this overwhelming growth was not accompanied by a change in economy (there is no evidence of domestication) we view it as a sort of anomalous condition for hunter-gatherers. This may have caused tension and may, at least in part, explain or reflect a transformation in worldview – a manifestation of change in the relationship between humans and their world that may soon have led to further socioeconomic adjustment. Indeed, plant domestication is evidenced in this very region, including the PPNB layers of sites such as Nevalı Çori, shortly after the PPNA site of Göbekli Tepe went out of use.

2. Conspicuous technological changes in the production of flint tools, and in particular the emergence of a new technology that facilitated the production of long (non-curved, non-twisted) flint blades from which new types of sickle blades and arrowheads were shaped (Figure 2.20a–d). 3. Flint axes produced by flaking and polishing that grew in size and weight compared to their PPNA predecessors and that were used in tree-felling and wood-working. 4. New types of stone grinding tools appear including stepped querns and there is a decrease in pounding tools.

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Figure 2.19 Rectangular PPNB structures. (a)–(b) Jericho; (c) plastered floors at Yiftahel (Lower _ Galilee) – courtesy of Hamoudi Khalaily.

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Figure 2.20 (a) Flint blades from Yiftahel – courtesy of Hamoudi Khalaily; (b) naviform (boat-like) flint _ of Yosef Garfinkel, and see also items k, l in (c); (c) illustrated core from Yiftahel – courtesy _ flint tools and arrowheads (a–e) from Ḥorvat Minha (Munhata) in the Jordan Valley; (d) a _ _ Yosef Garfinkel, and see also flint sickle blade, photographed by H. Salomon – courtesy of items i, j in (c).

5. Changes in burial customs in the northern Levant can be seen in the burial-designated ‘skull building’ of Çayönü or the ‘house of the dead’ at Dj’ade. They are conspicuous in the southern Levant somewhat later in the PPNB with flexed burials (Figure 2.21) including the treatment of detached skulls, separated from the rest of the skeleton and buried disjointedly or placed in designated locations at the site (Figure 2.22). 6. New, innovative types of art objects, including stone sculpture (mostly in the northern Levant), clay statues (in the south), a variety of stone and clay, including many zoomorphic and anthropomorphic (human figures) imagery items, a small presence of bone imagery items, stone masks in the southern Levant, decorated stone vessels and decorated (painted) plastered floors (Figure 2.23a–g). And, the most prominent change observed at PPNB sites as early as their onset (that is, from the very beginning of the PPNB over 10,500 years ago) or somewhat earlier is:

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Figure 2.21 PPNB burial in flexed body position from ‘Ain Ghazal nearby Amman, Jordan – courtesy of Gary Rollefson and the ‘Ain Ghazal delegation, photographed by Curt Blaire.

Figure 2.22 Decorated and treated PPNB skulls. (a) Plastered and painted skull from Kfar HaḤoresh – courtesy of Nigel Goring-Morris; (b)–(c) Glued bitumen net pattern from Nahal Ḥemar _ published Cave, nearby the Dead Sea – courtesy of the Israel Antiquities Authority, originally in Bar-Yosef, O. and Alon, D. 1988. Nahal Hemar Cave, Atiqot 18; (d) skull modelled with plaster from Jericho; (e) partially modelled skulls from ‘Ain Ghazal – courtesy of Gary Rollefson and the ‘Ain Ghazal delegation, photographed by John Tsantes; (f ) plastered and painted skulls from Tell Aswad, nearby Damascus – courtesy of Danielle Stordeur. Colour versions of these images can be found at www.cambridge.org/abbo-gopher.

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the emergence of clear evidence of plant domestication first in the northern Levant, at sites located in northern Syria and south-eastern Turkey, and later in the entire area. The same pattern emerges for animal domestication too. We reiterate the magnitude of this change, which, within a relatively short time frame, had brought about the end of the subsistence based on traditional huntergatherer life-ways that had lasted several million years and in its stead set the foundation for production-oriented societies. This is also a major reason for the long-standing research interest in this occurrence, our work included. In the southern Levant (Israel, Jordan, the Palestinian Auhority, Lebanon and southern Syria), the PPNB is characterized by the emergence of huge sites (known as mega sites) that span up to 150 dunams (15 hectares) and feature many densely built structures suitable for large populations. Such sites were found within and near the Jordan Valley (Wadi Shu’eib), at the edge of the Arabah (Basta), and the Transjordanian Plateau east of the Jordan River (‘Ain Ghazal; see Box 10 ‘Ain Ghazal, p. 61), and recently, for the first time west of the Jordan River in the Judean Hills (Motza). Pre-Pottery Neolithic B large and medium size sites were also found in the northern Levant, including Çayönü, Tell Abu Hureira, Göbekli Tepe and Nevalı Çori (see Box 11 Çayönü, Nevalı Çori and Tell Ḥalula, p. 64). While we have no direct, clear-cut evidence that the sites were fully occupied at any given point in time, it is nevertheless clear that their size significantly diverges from that of PPNA sites and certainly the size of Natufian sites. Pre-Pottery Neolithic B sites of a different nature are found in the desert area of Israel, the Negev, as well as in Sinai (Egypt), the east and south of Jordan, and Syria (see Box 12 Nahal ‘Issaron and Wadi Tbeik, p. 68). This testifies to a mild climate that _ facilitated habitation in the area. It is as yet unclear whether the desert hosted independent populations or whether the sites were merely seasonal (or even permanent), affiliated with groups that originated in the large, permanent sites of the Mediterranean vegetation zone. All in all, then, PPNB sites increased in size, and they traversed the entire area, attesting to a general population growth. Towards the end of the Pre-Pottery Neolithic, a period some researchers named PPNC, another change occurred in the archaeological map. While site numbers dwindled, mega sites such as the Jordanian ‘Ain Ghazal and Wadi Shu’eib were still in use. Some researchers suggest calling the cultural entity comprising the PPNC period the Ghazalian culture, after the site of ‘Ain Ghazal, where its presence was first discovered. Among medium-sized sites, some specialized in unique economic activities, for example ‘Atlit Yam, a site that is currently submerged off the coast of ‘Atlit, just south of Haifa, hosted a community of fishermen who left behind a large variety of well-preserved findings, including rectangular residential houses and a water well over 5 m deep (see Box 13 ‘Atlit Yam, p. 71). The PPNC period shows a relatively low density of sites spread throughout the Mediterranean zones of Israel, Jordan and the more northern parts of the Levant. For example, the site of Hagoshrim in the northern Israeli Ḥula Valley, named after the

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Figure 2.23 PPNB statues and figurines and body ornaments: (a)*–(b) clay imagery of cattle and a cattle figurine stabbed by flint blades from ‘Ain Ghazal, photographed by Peter Dorrel and Stuart Laidlaw; (c)** coloured stone mask from Nahal Ḥemar Cave; (d)–(e)** photograph and illustration of _ Ḥemar Cave; (f )* human figure about 1 m tall made of bone human face statuettes from Nahal _ clay over a cane frame from ‘Ain Ghazal, photographed by John Tsantes; (g) photograph of human statue with shells for eyes from Jericho; (h)*** stone beads from Nahal Ḥemar Cave. _ Colour versions of these images can be found at www.cambridge.org/abbo-gopher. * Courtesy of Gary Rollefson and the ‘Ain Ghazal delegation. ** Courtesy of The Israel Antiquities Authority, originally published in Bar-Yosef, O. and Alon, D. 1988. Nahal Hemar Cave, Atiqot 18. *** The Israel Antiquities Authority collection @ The Israel Museum. 60

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adjacent kibbutz, yielded oval stone structures and a broad variety of findings. A PPNC site in Ashkelon, near the sea, suggests that a community of hunter-gatherers resided there. It has been suggested that the large Pre-Pottery Neolithic C of ‘Ain Ghazal hosted a large community whose subsistence economy was derived from growing cereals and

Box 10 ‘Ain Ghazal: A Pre-Pottery Neolithic B mega site in Jordan The Jordanian Neolithic site of ‘Ain Ghazal was discovered in 1974 during the construction of a main highway to Amman, Jordan’s capital city. Since 1982 and until the end of the 1990s, the site was excavated by Gary Rollefson and Zeidan Kafafi. ‘Ain Ghazal is a very large site, one of a few mega sites, spanning, in some layers, up to 150 dunams (15 hectares) (see Figure 2.24b). The site is located on the Transjordanian Plateau, some 1,000 m above sea level at the edge of the Irano-Turanian vegetation zone. It consists of PPNB and Pre-Pottery Neolithic C layers, which have been extensively radiocarbon dated to the period of ca. 10,200–8,500 years ago. Above these layers is a Pottery Neolithic layer that had been assigned to the Yarmukian culture, dated ca. 8,500–8,000 years ago. Here we discuss the PPNB layers that show rectangular stone structures with plastered floors and additional stone installations. Slopes were levelled and retaining walls were built in order to erect numerous structures. The floors in many of the structures were made of lime plaster and some of the walls had a lime plaster cover as well, occasionally decorated with colour. Some unusual structures at this site have been interpreted as Neolithic shrines, and large erect stones found within them were interpreted as Massebot (standing stones) (Figure 2.24f). Many burials were found dispersed within and between houses under the plastered floors. As with other PPNB sites, here, too, skulls were separated from adult skeletons. Treated and decorated skulls, buried individually or in groups, were found too (see Figure 2.22e).

The rich assemblage of flint tools included arrowheads, sickle blades, tree-felling and wood-processing axes, as well as other types of flint tools. Findings included many grinding and some pounding stones as well as a few bone tools. The site excavators claimed that occupants intensively exploited their surroundings, thereby damaging the sensitive steppe ecosystem of the desert frontier and reducing the quantities of accessible resources near the site. This, in turn, would have necessitated that occupants travel far for food. Most Pre-Pottery Neolithic B layers at the site show subsistence based on plant gathering and animal hunting (deer, aurochs, pig and small mammals such as hare and predators). Evidence of domesticated plants was found quite early in the sequence, including wheat, barley, lentil, pea, chickpea and flax (as well as broad (faba) bean), while domesticated animals were found only in the later occupations of the site, including mainly goat. These PPN layers attest to the transition to production economy, growing cereals, legumes and livestock. The trading network at ‘Ain Ghazal was geographically dispersed. Basalt tools were brought from far away; asphalt from the Dead Sea; obsidian from Turkey; molluscs from the Mediterranean and the Red Sea; carnelian and turquoise from the southern Negev and Sinai, to name just a few. The artistic-symbolic element of the site is demonstrated in the large quantity of clay statues and figurines found. Especially notable are two large hoards of clay statues, both comprising over thirty human images up to 1 m tall, including men, women, children and figures of ambiguous gender. Smaller female figures of

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Figure 2.24 The site of ‘Ain Ghazal. (a) Site location; (b) general plan (grey line lunate/oval) marks the site area); (c) reconstruction of the (Middle) PPNB village; (d) two-storey stone structure; (e) general view of an area with rectangular structures and two circular structures interpreted by the excavator as shrines; (f ) rectangular stone structure interpreted as a shrine with standing stones; (g) elongated rectangular structure of the same period; (h) and (k) two rectangular structures showing plastered floors and installations; (i) apsidal structure (rectangular with one arched end) of the Yarmukian culture, interpreted by the excavator as a shrine; (j) close-up on one of the round structures; (l) flint sickle blades; (m) flint arrowheads; (n) stone figurine of a woman; (o) clay statue of ‘twins’ made over a cane frame (see Figure 2.23 for another statue of the same type) – photographs and drawings by Gary Rollefson, John Tsantes, Jonathan Mabri, Curt Blair, Yusef Zo’abi, Ali Omari and Muwafaq Bathaineh – courtesy of Gary Rollefson and the ‘Ain Ghazal delegation. stone are also notable (Figure 2.24n). Among smaller items were many animal figurines, including fox, goat, pig and cattle. Some of the cattle figurines were stabbed by a flint item with traces of tied rope – perhaps evidence of a hunting ritual or a symbol representing control.

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The PPNB population residing at ‘Ain Ghazal was large and characterized by agricultural life-ways. This notwithstanding, its members continued to hunt and gather, requiring a sophisticated seasonal work plan to subsist and provide for themselves and their livestock at the semi-arid desert frontier.

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Figure 2.24

(cont.)

legumes as well as domesticated herds, especially of goats and sheep. A reconstruction of environmental damage near the site led the researchers to suggest that this was the home-base of shepherds who travelled far with their herds, and for long periods of time, in search of pasture, yet they maintained some rights at their home-base and perhaps

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Box 11 Çayönü, Nevalı Çori and Tell Ḥalula: Pre-Pottery Neolithic B sites in Turkey and Syria

overview of some interesting aspects that are unique to these sites and provide a glimpse into the cultures of the period in question. All three sites yielded impressive findings in both scope and content. Their architecture revealed special structures while their material assemblages (stone and flint industries, beads and other body ornaments, figurines, etc.) are of high interest. In the context of this book, the most important finding shared by all three sites is the clear evidence of domesticated

Three sites were selected to represent the PPNB of the northern Levant: Çayönü near the Tigris as well as Nevalı Çori and Tell Ḥalula, in the area of the Middle Euphrates. The first two are located in Turkey while the third is situated in Syria. Refraining from particular detail of each site, we instead offer an

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Figure 2.25 The site of Çayönü. (a) General view; (b) round stone structure from an early, PPNA, layer; (c) and (g) rectangular PPNB structures with their underlying cell structure, also known as grill buildings; (d)–(e) the skull building and a close-up on human remains in its northern rooms; (f ) square PPNB structure with multiple rooms; (h) a unique stone item (divider?); (i) bone tool (needle?); (j) and (l) stone pendants and beads; (k) obsidian drills and sickle blades. All figures – courtesy of Mehmet Özdoğan.

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plants. At both Çayönü and Nevalı Çori, this is among the earliest evidence found attesting to plant domestication in the Near East, dated to the early PPNB, some 10,500 years ago. At

Çayönü, for example, barley, wheat, chickpea, lentil and pea were found (as well as almond, fig and grape). The small archaeobotanical assemblage of Nevalı Çori yielded

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Figure 2.26 The site of Tell Ḥalula. (a) General view; (b) rectangular structures made of brick; (c) pattern of floor decoration; all figures – courtesy of Miquel Molist. einkorn and emmer wheat, barley, lentil, pea, chickpea and broad bean (along with pistachio, almond and grape). Tell Ḥalula is a later PPNB agricultural site that eventually yielded domesticated wheat, barley, lentil, pea, vetch and broad bean. The architectural variation at all three sites is notable. There are both residential and other impressively designed and constructed buildings. Structures at Çayönü (Figure 2.25c, f) and Nevalı Çori were built on parallel foundation walls that slightly elevate structure floors above the ground (the so-called ‘grill buildings’). This has been interpreted by some experts as an attempt to

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maintain dry interiors in wet seasons and ventilate in hot seasons. Both sites yielded unusual structures. At Çayönü, a rectangular structure with one rounded end contained many burials and was dubbed the skull house (Figure 2.25d, e); at Nevalı Çori large Tshaped stone statues were found in some square structures with reliefs of human hands at their front, and one of the structures was surrounded by stone benches (Figure 2.27); they were interpreted as shrines. Tell Ḥalula showed rectangular structures made of mud and stone as well as some unique structures; the floors were painted in black and red and had representations of humans and animals

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Figure 2.27 The site of Nevalı Çori. (a)–(b) The site’s temples; (c) reconstruction of temples with stone pillars; courtesy of Harald Hauptmann.

(Figure 2.26c). Of note are the Tell Ḥalula ‘sitting burials’. Worldviews of the site residents and their world of symbols are expressed in a large variety and quantity of statues, figurines

and other artefacts that attest to their nature as agricultural, sedentary societies whose subsistence economy is far removed from that of the earlier huntergatherer societies.

even left property, storage rooms or even homes behind them (see Figure 2.31). All in all, then, this settlement seems to portray a complex, multifaceted picture. Taken together, Pre-Pottery Neolithic cultures represent a cultural interaction sphere (a koine à la J. Cauvin) that is among the most distinct and active in the history of the Levant. Intensive ties that persevered for some 3,000 years were strewn throughout the area, facilitating the distribution of matter, technology, knowledge and ideas, disseminating the many innovations in life-ways and socio-economy throughout the area. In our view, the northern Levant, and in particular those areas that we call the core area (northern Syria and south-eastern Turkey), were the stage upon which plant (and animal) domestication unfolded and from which domesticated plants proliferated. This claim is based on botanical findings (plant parts, especially seeds), genetic studies,

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Box 12 Nahal ‘Issaron and Wadi Tbeik: Desert sites_of the Pre-Pottery Neolithic B period

of only a few hundred sq m. Wadi Tbeik is located on the hill slope, and it, too, is small, spanning some 200 sq m. Both sites were radiocarbon dated to ca. 9,500 years ago and therefore represent the PPNB of the desert areas that did not undergo the socioeconomic developments of the Agricultural Revolution in the richer, northern areas of the Levant. These sites hosted small communities with just a few dozen members each, who occupied the sites on a seasonal basis, according to prevalent conditions and available resources. Both sites showed round stone structures that were densely organized in a configuration that may be called a beehive pattern

Nahal ‘Issaron is located in the south of the _ Arabah, in the ‘Uvda Valley in Israel, while Wadi Tbeik is located on the high mountain mass in the south of the Sinai Desert in Egypt. Nahal ‘Issaron was excavated in the early _ 1980s by Nigel Goring-Morris and Avi Gopher; the site of Wadi Tbeik was excavated in the late 1970s by Ofer Bar-Yosef, Nigel Goring-Morris and Avi Gopher. Located in the outlet of Nahal ‘Issaron _ towards the ‘Uvda Valley, the site of Nahal _ ‘Issaron is fairly small, extending over an area

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Figure 2.28 The site of Nahal ‘Issaron. (a) General view westwards; (b) close-up on structure _ No. 26; (c) (passive) grinding stone and two (active) grinding stones (scale is 20 cm long); (d) site plan (each unit on the frame represents a single metre); (e) flint drills (left) and flint arrowheads (right); (f ) body ornament (nose ring?) made of a cowry sea shell; (g) long flint drills; (h) pendants made of mother of pearl sea shell. Colour versions of these images can be found at www.cambridge.org/abbo-gopher.

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(Figures 2.28a, 2.29d). The structures, used for residence, are 3–5 m in diameter and some of them included stone-built installations. Burials were only rarely found at desert sites; none was discovered at Nahal ‘Issaron _ while Wadi Tbeik had skeletons of only two individuals who were buried in an abandoned structure.

Material culture findings at both sites included a wide, rich variety of flint tools as well as stone pounding and grinding tools. The sites also yielded bone tools. Prominent tools included different types of arrowheads, awls and borers (Figures 2.28e, 2.29f). Particularly notable was the absence of sickle blades, which were typical of all

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Figure 2.29 The site of Wadi Tbeik. (a) General view; (b) close-up on structures Nos. 101, 12 and 10; (c) close-up on structure No. 11; (d) site plan (scale at the bottom right); (e) stone slab with circular depressions (interpreted as a game board); (f ) arrowheads; all items – courtesy of Nigel Goring-Morris and Ofer Bar-Yosef. Colour versions of these images can be found at www.cambridge.org/abbo-gopher.

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Neolithic flint tool assemblages both in Mediterranean areas and in the desert frontier. Of note is the absence of flint axes and adzes too. Subsistence economy at both sites was based on gathering as well as hunting of local animals such as gazelle, ibex, hare, equids, ostrich, some

reptiles and birds. Nahal ‘Issaron also _ yielded a few fish vertebrae. Engraved and incised stone and bone artefacts were uncovered at both sites as were stone beads, pendants and bracelets and molluscs originating in the Mediterranean and the Red Sea.

Box 13 ‘Atlit Yam: A Pre-Pottery Neolithic C site at the bottom of the Bay of ‘Atlit

prolonged exposure to cold, deep waters. Bones of deep-water fish species were indeed uncovered at the site, thereby corroborating the notion of deep-water fishing. The diverse flint assemblages at the site included arrowheads, sickle blades, flint axes and adzes and a chipping floor rich in knapping debris. Additional findings included pounding and grinding tools made of limestone, a few wooden tools and some bone tools including spatulae and awls. Artistic-symbolic imagery items as well as body ornaments made of processed molluscs were also found. In addition, stone was used to make beads and a few decorated stone items. The findings suggest that site subsistence was based on hunted animals, the most prominent of which were cattle, goats and pigs. Gazelle and deer were found in small numbers. Subsistence was complemented with diverse fish, typically triggerfish. Information regarding domesticated animals is unclear because in the early phase of research, the cattle, goats and pigs were considered wild, while at a later phase domestication was suggested based on the composition of the faunal assemblage (e.g., the high percentage of goats) as well as the high numbers of animals slaughtered at a young age. Additionally, cereals and legumes were grown at the site, including wheat, barley, chickpea and lentil. The site also

The site of ‘Atlit Yam is located 12–18 m below sea level off the coast of ‘Atlit, Israel (Figure 2.30a). It was discovered in 1984, excavated for over a decade and investigated until the early 2000s by Dr Udi Galili. The site is located in a trough (marzeva in Hebrew) valley between two submerged kurkar (solidified dunes) ridges and near a shallow lagoon. The site was radiocarbon dated to ca. 9,000–8,500 years ago. It is estimated to have spanned 50–60 dunams (5–6 hectares), hosting a large community of fishermen who were also engaged in farming. It is among the earliest fishermen’s villages known in the Near East. The site has rectangular structures and structures dubbed corridor houses (comprised of two rows of rooms, one on each side of a corridor) as well as many installations located both within and without of the structures. In one part of the site, a well was dug into bedrock, while at another Massebot (or perhaps just large stone statues) were placed (Figure 2.30c, f). Numerous burials were uncovered, mainly concentrated in one area of the site. Most individuals were buried with their skulls, thereby indicating the cessation of the earlier PPNA and PPNB custom of skull separation. Some skulls attest to an ear disease that results from

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Figure 2.30 The site of ‘Atlit Yam. (a) General view, photographed by X. Clive; (b) (reconstructed) rectangular structure; (c) the large erect stones structure (a photograph of the large erect stone structure (note scale by comparing to the diver in the background)); (d) the well during excavation; (e) skeleton in flexed position; (f ) a circle of large erect stones interpreted as a ceremonial facility; (g) flint arrowheads; all items – courtesy of Udi Galili. Colour versions of these images can be found at www.cambridge.org/abbogopher.

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yielded flax seeds. Weeds typical of cultivated (tilled) fields were recently discovered in the botanical assemblage by Dr ‘Anat Hartmann-Shenkman and

her colleagues at Bar-Ilan University: Professor Mordechai Kislev, Dr Yoel Melamed and Professor Ehud Weiss.

Figure 2.31 A rectangular structure at PPNC ‘Ain Ghazal comprising a corridor with small cells – photographed by Yusef Zo’bi, courtesy of Gary Rollefson and the ‘Ain Ghazal delegation.

genomic DNA analyses and archaeological findings that offer a vast amount of information regarding the cultural and perceptual landscapes within which domestication transpired. Following the Pre-Pottery Neolithic came the PN (Pottery Neolithic period), which, as its name attests, is characterized by the presence of pottery at archaeological sites – some recognizable pottery vessels alongside a multitude of pottery sherds (see Figure 2.32). The PN began in the northern Levant some 9,000 years ago, while in the southern Levant it began around 8,500 years ago. It is characterized by a large variety of local, small-scale cultures. In the northern Levant, some extensive cultural entities had emerged, such as the Halafian, the influence of which can be observed in vast geographic areas across the entire Fertile Crescent. In the southern Levant, in contrast, several cultures emerged consecutively during the PN period in different parts of the

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Figure 2.32 PN pottery vessels from: (a)–(b) Nahal Qanah Cave in western Samaria; (c) Ein el-Jarba; (d, e, f ) Sha‘ar Hagolan – photographed_ by D. Kharis, courtesy of Yosef Garfinkel. Colour versions of these images can be found at www.cambridge.org/abbo-gopher.

area: the Yarmukian (named after the Yarmuk River, which feeds into the Jordan River just south of the Sea of Galilee where the site of Sha‘ar Hagolan is located and where it was first discovered); the Lodian (named after a site in the city of Lod despite being first uncovered at the Tell of Jericho); the Wadi Rabah (named after the wadi in which it was excavated – near the city of Petah Tikva, Israel – and one of the first sites to reveal this _ culture); the Qatifian (named after Tell Qatif on the southern coastal plain of the Gaza Strip); and the Besorian (named after sites situated on the banks of Wadi Besor in the northern Negev), to name a few. We will not elaborate on this period because the domestication of plants (and animals) occurred long before the PN. We merely note that significant institutionalization processes took place during the PN at both the economic and conceptual levels, which were the aftermath of domestication. The apex of domestication occurred with the full institutionalization of the agricultural society, which was based on the combination of crop plants (cereals and legumes) and livestock (e.g., goats, sheep, cattle and pigs). But by the end of the Neolithic period, the PN had fully divorced itself from and obliterated the last remaining vestiges of the hunter-gatherers’ ethos, which had still been present in the earlier Neolithic. By the end of the PN, hunting became a dying practice, no longer serving as a significant element of subsistence. It can thus be claimed unequivocally that the Agricultural Revolution fully materialized during the PN period (see Box 14 Sha‘ar Hagolan and Nahal Zehora II, below). _

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In light of the brief historical-archaeological review we have presented, the remainder of the book is devoted to the systematic analysis of plant domestication that eventually gave rise to the Neolithic ‘crop package’, or founder crops. Among the founder crops were barley, two species of wheat, pea, lentil, chickpea, bitter vetch and flax and their domestication is the very heart of the Neolithic Revolution and the institutionalization of agriculture as a sustainable system of subsistence lasting to this very day. In the next chapters we will explore various aspects – anthropological, genetic, botanical, ecological and agronomical – that pertain to plant domestication based on

Box 14 Sha‘ar Hagolan and Nahal Zehora _ II: Pottery Neolithic institutionalization of the Agricultural Revolution The Levantine PN is represented here by the Yarmukian culture, the earliest of the PN cultures in the southern Levant, as it is reflected in the site of Sha‘ar Hagolan; and the Wadi Rabah culture, the last of the PN cultures, as it is reflected in the site of Nahal _ Zehora II.* The site of Sha‘ar Hagolan is situated in the Jordan Valley near the Yarmuk River. It was discovered at the end of the 1940s and excavated in the early 1950s by Moshe Stekelis, who defined this culture and named it after the adjacent river. This Yarmukian site, found to cover an area of some 200 dunams (20 hectares), was later excavated more extensively by Professor Yosef Garfinkel. The site is rich in innovative architecture, notably different from its forerunners with courtyard houses, that is, rooms, arranged around a closed courtyard (Figure 2.33a–c). The site excavators ascribed this finding to a transformation in the social structure towards extended family units or households. The site, which had an impressive dug well, 5 m deep, yielded rich material assemblages. The large pottery vessel assemblage included bowls, pedestalled bowls, goblets, jars, small jars, holemouth jars and pithoi that are all typical of the Yarmukian; some were incised with herringbone patterns and painted red. The rich flint tool assemblage included arrowheads, sickle blades, axes, scrapers, burins and

grinding tools made of stone. The artisticsymbolic imagery assemblage is also highly diverse, including anthropomorphic (human) and zoomorphic (animal) clay figurines as well as a broad variety of engraved stone imagery items (Figure 2.33d, e). Compared to Sha‘ar Hagolan, Nahal _ Zehora II is a fairly small site, covering only 8 dunams (0.8 hectare). It was excavated by Avi Gopher from 1988 to 1998. While the bottom layer of this site was assigned to the Yarmukian culture, here we describe the Wadi Rabah culture of the top layers, which yielded typical rectangular structures and small plastered installations. The rich pottery assemblage includes typical forms, forming techniques and painted and incised decorations (surface treatments). The flint tool assemblage includes a small (and dwindling) number of arrowheads alongside a significant number of sickle blades, as well as diverse axes, adzes and chisels. The artistic-symbolic imagery assemblage includes zoomorphic clay figurines as well as anthropomorphous figures and incised stone items interpreted as representations of humans. Additional artefacts include stone items with incised patterns, beads and other body ornaments (Figure 2.34). Inhabitants of both sites subsisted on domesticates – cereals, legumes and flax among plants, and goat, sheep, cattle and pig among animals. Hunting of gazelle, deer and other animals continued but decreased in scope. Altogether, these were clearly agricultural sites. Both cultures yielded evidence of

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Figure 2.33 The site of Sha‘ar Hagolan. (a) General view of excavation Area E; (b) reconstruction of Structure I; (c) plan of Structure I, Area E; (d) clay figurines; (e) incised stone imagery items – courtesy of Yosef Garfinkel. Colour versions of these images can be found at www.cambridge.org/abbo-gopher. the Secondary Products Revolution, namely, the exploitation of animal secondary products (beyond meat) such as wool or milk. Spinning (and possibly weaving) was carried

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out at both sites, attested to in the form of round spindle whorls, while churning took place during the Wadi Rabah culture, attesting to secondary milk products such as

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Figure 2.34 The site of Nahal Zehora II. (a) A view of the site (marked by arrow) against the Menashe Hills;_ (b) Yarmukian stone-paved structure shaped as half a hexagon, opening towards the west; (c) Wadi Rabah jar, which, when opened, revealed a buried foetus; (d) the opened jar and foetus bones; (e) and (j) photographed and illustrated flint arrowheads; (f ) incised Yarmukian stone imagery item; (g) Yarmukian, Lodian and Wadi Rabah assortment of decorated clay sherds; (h) Wadi Rabah trapeze stone imagery item with an incised triangle; (i) PN clay spindle whorls; (k) flint sickle blades; (l) flint axe (top) and adze (bottom). Colour versions of these images can be found at www.cambridge.org/abbo-gopher.

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(cont.) butter and cheese. Taken together, this evidence implies the full institutionalization of the Agricultural Revolution, including its Secondary Products Revolution.

* Subsequent, post-Wadi Rabah cultural entities of the Pottery Neolithic period are not clear enough to be presented in this context.

which we will attempt to clarify the biological and cultural foundations of plant domestication in the Near East.

K E Y PO IN TS A ND BE Y O ND

• During the Paleolithic period, the archaeological landscape of the Fertile Crescent, as may be seen from a bird’s-eye view, remained stable for long periods of time, including few cave settlements and many open-air sites. The early and middle parts

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of the Epipaleolithic period (23,000–15,000 years ago) stereotypically represents, nearly to perfection, the common reconstruction and image of the world of huntergatherers. By the end of the Ice Age and especially with the emergence of the late Epipaleolithic Natufian culture (some 15,000 years ago), a change occurred in the archaeological landscape, as ‘spots’ in the view – that is, settlements – became more obvious and permanent. With the rise of the Neolithic period (some 11,800/700 years ago), the archaeological landscape was further boosted in the scope of its human ‘stains’, with some significantly larger sites and architectural features that are seen from afar, such as stone walls, a stone tower, monumental structures and even gigantic stone statues. Although subsistence was still based on hunting and gathering, these changes appear to be related to a transformation in worldviews and behavioural patterns of contemporary people and their social structure and organization. With the emergence of domestication, of both plants and animals, some 10,500 years ago, and the onset of agriculture, the archaeological landscape presents not only large settlement sites but also cultivated areas (fields) and herds of pasturing animals. In subsequent phases of the Neolithic period (PN, starting 9,000–8,500 years ago), the agricultural landscape changed beyond recognition, never to return to its previous form. Institutionalization increased in the agricultural landscape, which boasted a large spread of villages of varying sizes. From small sites (less than 1 hectare) to huge sites (tens of hectares), agricultural communities had erected residential and public structures, storage facilities and water wells, while establishing field and pasture lands both near sites and off at a distance. The Chalcolithic period, post-dating the PN period, began ca. 6,500 years ago and brought about additional change, culminating in the early Bronze Age urbanization that started just in advance of 5,000 years ago, where the bird’s-eye view would have shown true cities. The urban (sometimes fortified) landscape as combined with rural villages remains familiar to this day. The influence and ecological footprint of humankind in its residential surroundings and beyond increased and continues to increase. The domestication of plants (and animals) and the institutionalization of agriculture as a sustainable system of subsistence were key to this process.

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3 MODELS TH AT DESCRIBE AN D E XPLAIN TH E AGRICULTURAL REVOLUTION, INCLUDING PLANT DOMESTICATION

We introduce this chapter with a partial discussion of the conceptual framework that underlies the research on the Agricultural Revolution (Glossary, General Terms, Agricultural Revolution). We first explain how hunter-gatherers gained their knowledge and put it to use. We then present key ideas that have been offered throughout the years to explain the advent of the Agricultural Revolution. Many researchers have attempted to answer the question of why the Agricultural Revolution occurred. Some answers are broad, driven by a perception that human action is a reaction to some external force(s). Others emphasize the social dynamics that might have led to this revolution. Most of these explanations were formulated during the last half of the twentieth century. To facilitate a better understanding of these views, we will elaborate on prevalent research perceptions regarding plant domestication (Glossary, General Terms, Plant domestication), which was central to the Agricultural Revolution. In recent years, and mostly since the beginning of the twenty-first century, the complex discussion on plant domestication in the Near East has been characterized by a series of bipolar, dichotomous questions. The first question is: Where? Did plant domestication take place in a specific and well-defined (core) area within the Near East (highly localized) or did it occur independently (autonomously) in different places (diffused) in the Near East, and if it originated in a single area, where was that area? The second question is: When (at what pace)? Beyond dating, was plant domestication rapid, a revolutionary change achieved in a single major move, or was it a protracted, slow and gradual process, perhaps involving numerous stages –

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a recurrent event repeated multiple times? And the third question is: How? Did domestication occur in an exclusive, single episode for each species and for the package as a whole (singular timing) or were there multiple independent domestications per species and thus, by definition, for the package as a whole? This raises the question of how the Neolithic package coalesced. Was the process of domestication (including the choice of plant species and the selection of specific phenotypes) unconscious, incidental or circumstantial (a result of human–plant co-evolution or by natural selection occurring inadvertently in cultivated fields), or a designated human initiative, a conscious effort to select and manipulate various plant species that would eventually become transformed? It is difficult to separate the answers to these distinct yet interrelated questions, and we would argue that taking a stance concerning one of the ‘leading’ pairs (e.g., on the question of where) can invariably tip the balance concerning other undecided ‘pairs’, promoting potential conclusions in line with it and providing a coherent cultural and biological scenario of plant domestication in the Near East. For example, a researcher who rejects the notion of a core area in which the domestication of the founder crops has taken place and supports the claim of multiple, autonomous domestications in various parts of the Near East will find it difficult to accept that the process of domestication involved a selected package of plants based on complementary nutritional value and agronomic advantages of its components. Similarly, scholars who disagree with the contention that plant domestication was a conscious, knowledge-based process will fail to see that this was a very rapid process or virtually an episode.1 In other words, the various aspects and arguments pertaining to the discussion of plant domestication are interlaced, bound by the research position adopted. The above dichotomies as a matter of fact converge into two major models reconstructing plant domestication in the Near East, and the major research questions currently debated in plant domestication research are: Which model best reconstructs plant domestication? Or, which of the two models requires fewer assumptions? Of the two models, the protracted-autonomous model, which has thrived in Near Eastern Neolithic studies for more than a decade, emphasizes three major aspects of domestication: (1) a long, protracted process that was (2) geographically autonomous (noncentred, multi-focal) characterized by (3) an unconscious nature. In contrast, the alternative core area-one event model views Near Eastern Neolithic plant domestication as knowledge-based and conscious, occurring in a limited core area and during a short, single event. Although the second model is corroborated by multiple lines of evidence, including geobotanical, archaeological, archaeobotanical, agronomic and genetic, it represents a minority view. We critique the protracted-autonomous model while briefly discussing its key elements under the subsection titled ‘The Niche-Construction Model (or the Protracted Symbiosis and Domestication Model)’. At the end of this chapter, we offer a short, combined discussion of the pace of plant domestication and the role of the domesticators’ consciousness.

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O BT A I NIN G , O R G A NI ZI NG, EM P LO Y IN G A N D T R A N S M I T T I N G K N O WL E D G E AMONG HUNTER-GATHERERS

While anthropological research is a vibrant field with newly introduced frameworks of thought, the statements we make in this section are generalizations that should be read as a basic background (and see comments in Chapter 1). The first plant domesticators were hunter-gatherers who relied upon experience accumulated during a long history of knowledge-based and sustainable usage of natural resources. Resources were used by hunter-gatherers based on an intimate and broad acquaintance with the environment and its elements. Hunter-gatherers typically defined in detail (classification, organization and naming by way of a taxonomy) many hundreds of plants and animals, and they were generally acquainted with dozens, and sometimes hundreds, of species that served them in different purposes, starting with food and ending with tools, ornaments and other artefacts. This intimate familiarity of hunter-gatherers with different species and their specific characteristics included comprehensive knowledge of geographic availability and the seasonality of life cycles and different processes associated with plants and animals throughout the year. An important characteristic of hunter-gatherers is their use of environmental resources in a way that does not impair the resources’ future potential. Such societies are known for their ‘savage thinking’ (dubbed thus following the famous book of Claude Lévi-Strauss, The Savage Mind)2 and their preservation and transfer from one generation to the next of rich and detailed knowledge regarding the environment and the plants and animals it hosts. This is unsurprising, given that these societies subsisted on these environmental resources and were therefore committed to their preservation for practical, existential reasons; it was also in line with their worldviews. It is thus easy to understand why hunter-gatherers invested considerable time in order to become acquainted with their surroundings – including the various plants and animals, the significance they ascribe to this knowledge and their manner of utilization that facilitates an unthreatened, perpetual existence. While we cannot answer whether this approach emerged from the (conscious) perception of environmental sustainability as understood today in the Western world, some researchers would adamantly claim so, despite the difficulty in supporting this claim. In the absence of a writing system, hunter-gatherers preserved in memory all of their accumulated knowledge regarding the environment and its resources through traditional behaviours, a culture of discourse, and mechanisms, both oral and practical, for the transmission of knowledge to younger generations. The accumulated knowledge and the intimate understanding of plant and animal biology were the lifeblood of the hunter-gatherer society, and so the transmission of knowledge down through the generations was an inseparable component of their life-ways. The hunter-gatherers shown in recent years in different documentaries (irrespective of the limitations embodied in these films) present the ‘noble savage’, resourceful and highly knowledgeable individuals, skilfully navigating their environment thanks to their vast and intimate familiarity with it.

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Accumulated knowledge of hunter-gatherers often seems to extend beyond the requirements of daily subsistence. Their highly detailed general knowledge was accumulated because of both their innate curiosity and their lively discussions about the environment and its species, as well as their vast field experience and constant observations on processes and events that took place in their environment. This knowledge served hunter-gatherers daily not only at the economic level but also at ideological-social levels, as a foundational institution, a resource representing wealth that could be used in times of decision or crisis, a fundamental, existential element in the worldviews of these societies. In our view, this accumulated knowledge (and especially the knowledge that was amassed in the name of wisdom, akin to modern basic scientific research) was at the very heart of the process that – given adequate perceptual and cultural adaptation – eventually led to plant and animal domestication. M O D E L S T H A T D E SC R I B E A N D E X P L A I N T H E O N S E T O F A G RI C U LT U RE

Since their onset models explaining the Agricultural Revolution and plant domestication have included external influences that generated human distress as a driving force, which in turn drove humans to seek new subsistence resources. One response to this perceived reality was the intensification of food production by the growing of wild plants in cultivated fields, that is, taking action to improve the yield of various wild species that had already been established as food sources, or cultivating plants. These actions bear an implicit assumption that selection and some treatment (husbandry of sorts) of plants would result in greater yield (and predictability) than was available at the time in natural lands, which would facilitate overcoming the shortage caused by the deteriorating environment and its depleted resources. Although the root cause in these explanations is external, such as a climatic change, they embody an important element by attributing domestication and later improvement of domesticated species to activities initiated by humankind. For example, the epic model of Gordon Childe from the 1930s, known as the Oasis Theory (see more below), as well as the model suggested by Lewis Binford in the 1960s, known as the Marginal Zone Theory (also see below), are both characterized by this attribution. Other explanations of the Agricultural Revolution (and its component of plant and animal domestication), which did not focus on an external factor, attributed this transition to a change in humans’ perception, their implementation capabilities, resourcefulness and creativity in addition to social processes that took place in human communities. Early on in the research of plant domestication, different views were also voiced, suggesting that plant domestication was merely an unplanned ‘accident’ or the outcome of circumstances that led individuals to pay attention to random phenomena taking place in the environment. Consequently, benefits were noticed by them that could emerge from a new relationship with certain plants. An example of this view was the Dump-Heap Hypothesis of the German scholar T. H. Engelbrecht, suggested early in and further

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developed throughout the twentieth century (see below). The popularity of this approach grew in the twenty-first century, and, particularly through new, recent research, this view – that plant domestication was a prehistoric ‘accident’, a process to which humans were oblivious – became prevalent. According to this approach, plant domestication was the result of actions that humans had taken without any particular goal to domesticate plants; there was thus no intentionality. Proponents of this approach also posit that in the beginning of plant domestication, early farmers had not noticed the proliferation of plants with domesticated appearance (phenotypes; see Glossary, Genetics, Phenotype) in their fields. These researchers base their argument on the great similarity between domesticated plants and their wild progenitors. We believe this approach is patronizing and derogatory, especially as it originates with modern, Western individuals, separated by thousands of years from the lifestyle of the Neolithic domesticators, and who have long since lost almost all direct relationship and any interrelations with the environment and its natural resources. Both anthropological research and archaeological evidence show that humans’ actions in the past were both calculated and knowledge-based, guided by experience; these were systematic, deliberate actions in any field, but particularly so in the context of plant and animal domestication. The choice of plant species and plant packages that would become the founder crops in each primary centre of domestication worldwide, and certainly in the Near East, was a thoughtful craft that attests to high capabilities and extensive knowledge. To date, with very few exceptions (e.g., canola), modern scientists have not managed to harness new species that can make such a considerable contribution to the overall food basket beyond those staple crops that were domesticated in prehistoric times. This leaves the founder crops as the cornerstone of present-day global food production, and hence renders them highly significant from the economic perspective, even after thousands of years of agricultural activity, including the most modern agricultural research pursuits. This fact alone should teach us that the people who domesticated the founder crops over 10,000 years ago must have understood their choices and acted in an informed manner to achieve this domestication. In developing a model that could describe and explain the Agricultural Revolution and the emergence of a subsistence economy based on food production, it is difficult to distinguish between those elements that pertain to the biology of domesticated plants and those that pertain to the socio-economic nature of the prehistoric communities driving the Agricultural Revolution. Nevertheless, such a distinction can be made, and it would be useful in order to facilitate a better understanding of the processes involved in plant domestication. Three groups of models emerge through this prism: models emphasizing the biological dimension and the manner by which mutations (see Glossary, Genetics, Mutation) are exposed that give domesticated plants their unique expression in the form of desirable traits; models that emphasize the ecological aspects of plant domestication; and models focusing on a key theme in the research of the Agricultural Revolution, namely, the cultural and behavioural aspects of humans and

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the role of Neolithic community members in plant (and animal) domestication as well as in the Neolithization process (see Introduction, note 1, p. 240) as a whole. In addressing the ‘role’ of Neolithic people in plant domestication, we refer to the question of whether plant domestication was a result of knowledge-based initiative for the purpose of food production, or whether domesticated plants (and perhaps agriculture as a whole) developed randomly following a selection process of which Neolithic individuals were unaware despite partaking in it. To demonstrate this point let us assume that seeds of wild cereals are sown in order to intensify the naturally available local grain yield. Subjecting wild seed stocks to annual cycles of sowing and harvesting (with sickles) is likely to expose naturally occurring traits that confer a selective advantage under the man-made husbandry regime. One such important trait is spike non-shattering, the frequency of which is likely to increase under repeated cycles of sowing and reaping, and eventually dominate the managed (cultivated) population. This gave rise to the important notion of automatic selection, which expresses the view that the emergence of morphologically domesticated genotypes (see Glossary, Genetics, Genotype) was an inevitable, unconscious (thereby unintended) outcome of human activities in the nascent cultivated fields. One of the prominent advocates of this approach was the late Israeli geneticist Professor Daniel Zohary, who over several decades of original research made many important contributions to the study of Near Eastern plant domestication (see further reading list at the end of this volume).

Childe’s Oasis Theory

An early model explaining the Agricultural Revolution suggested in the 1930s was the Oasis Theory of Gordon Childe. Childe believed that a climatic change and a continuous decrease in rainfall in the Near East resulted in the depletion of vegetal resources in the area as well as shrinkage in herbivore populations. Valleys of large rivers such as the Euphrates, Tigris and Nile continued to pass water from high precipitation areas, standing out as green, fertile paths, against an arid background. Following Childe’s approach, these river valleys drew many animals while also sheltering grain-yielding plants that were used as a food source by gatherers. The forced coexistence in these valleys brought humans, plants and animals intimately close to each other. This in turn allowed humans to observe plant and animal biology from close up, and domestication became possible following this knowledge gain. The model is unacceptable to most researchers in the field since some of its key tenets – such as a climatic change or the geography of domestication – do not withstand the test of time and knowledge we have gained since then. Why, then, should we pay attention to this model? Because of the important underlying assumption that domestication was driven by an external factor (climatic change) and the intimate coexistence of the desertification refugees. This notion is repeated in other models, albeit involving

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different factors, such as the dwindling of natural resources following perpetual, massive hunting (e.g., gazelle, deer) and gathering. The Nuclear Zone Theory

Robert Braidwood suggested this theory in the 1960s following his archaeological work in Iraqi Kurdistan at the margins of the Zagros and Taurus mountain ridges. Together with geomorphologists and environmental researchers, climatologists, zoologists and botanists, Braidwood argued that this was the area in which the founder crops originated, since it is the natural range of the progenitors of those species that had been domesticated during the Agricultural Revolution in the Near East. Braidwood claimed that these wild species are still present in the area today, and that the current climate and natural resources are similar to those that had prevailed at the time of domestication. He advocated that technological developments and the emergence of sedentary settlements brought hunter-gatherers to specialize and become deeply familiarized with the plant and animal species that had domestication potential. In a gradual process, following many prolonged trials and errors, incipient agriculture had developed, following which agricultural sedentary villages were established, whose subsistence was based on domesticated plants and animals. To the question of why the Agricultural Revolution had occurred at that time rather than earlier (in concordance with similar climatic conditions), Braidwood offered a response that became an inalienable tenet of Agricultural Revolution literature: culture was not ready. While it is clear that Braidwood was referring to the technological, social and structural readiness of the prehistoric society, we might also add conceptual readiness (see below). In the middle of the twentieth century, several suggestions were made attributing the Agricultural Revolution to demographic pressures, that is, a population increase. Researchers such as Ester Boserup, Philip Smith or T. Cuyler-Young believed that the process was related to the end of the Ice Age and subsequent climatic improvement, which increased the availability of accessible resources and the possibility of sedentary settlement. This, in turn, prompted a demographic expansion, which necessitated the intensification of food production, which again drove demographic expansion and so forth. During this process, technologies and skills required for the intensification of production were honed and enhanced until plant and animal domestication were fully mastered as part of the intensified production. A key question remains with such explanations: Which came first – demographic expansion or the intensification of food production (see Introduction, note 3, p. 239)? The Marginal Zone Theory

One of the most instructive models suggested pertaining to demographic pressures is the Marginal Zone Theory. It offers a global explanation for the emergence of sedentary

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settlements and the onset of agriculture in the spirit of the New Archaeology (also known as Processual Archaeology) that materialized in the 1960s. According to Lewis Binford, who suggested this theory in the late 1960s, melting glaciers at the end of the last Ice Age and the subsequent rise of sea levels covered lands that had previously been accessible to humans while also generating new, rich environments in coastal areas and around the rivers flowing towards them. Richness was manifested in fish and fowl populations as well as additional animal and plant resources. Thus, at the end of the last Ice Age, hunter-gatherers were able to take advantage of a great variety of faunal and floral species in these food-rich areas, which drew them to establish sedentary settlements. The increased availability of these resources at the end of the last Ice Age and the subsequent climatic improvement inspired the demographic expansion of humans, too. This, however, increased competition over available resources and thereby also increased environmental stress, which led to food production. Suitable areas were taken over by the growing hunter-gatherer populations that opted for sedentary life. Some communities then developed social structures that would stabilize the population based on a hunting-gathering subsistence economy. However, in other communities, where social structures were flexible and open, social unrest tore communities apart to the point of relocation (budding off ) of certain groups to nearby areas. This process was repeated until some groups were pushed to the marginal areas of fertile lands, where natural floral resources were insufficient to meet subsistence demands. Having knowledge of plants and animals from the originally settled homelands, immigrants to these marginal zones brought with them seeds of the species they were familiar with and which they then grew in their new habitats. Binford believed that annual planting and harvesting aimed at supplying food sources to the inhabitants of the marginal zones were the actions that eventually led to the domestication of the select few species that were grown by Neolithic communities. We wish to emphasize that the underlying premise of the above theories is the need and urgency of supplying greater quantities of accessible foods (by intensifying production) as a result of the distress driven by environmental and/or demographic changes. These models assume that change was not the primary choice of the people who experienced it and who, so the theories stipulate, explicitly or implicitly would have preferred to continue living the hunter-gatherer way. The Dump-Heap Hypothesis

Another group of models explains the Agricultural Revolution and plant domestication through circumstantial reasoning as a process that is unplanned and unguided by humans. A classic example of these models is the Dump-Heap Hypothesis that was developed in the first half of the twentieth century in the works of researchers such as T. H. Engelbrecht, John G. Hawkes and Edgar Anderson. The underpinning assumption of the model is that

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human activity obstructs the natural growth of plants thereby favouring some species over others. Ecological niches (Glossary, Botany, Ecology and Agronomy, Ecological niche) that are impacted by human activity are invaded by plants that can leverage the advantages offered by these niches. Examples of such niches are sites used for butchering of hunted game or other areas in which waste was discarded, characterized by high concentrations of organic matter and minerals that fertilized the soil and encouraged the growth of certain plants. Such highly fertile sites, due, for example, to human or animal excretions, stimulated growth of randomly germinated plants, thereby increasing their yields. Some plants thrive around human dwellings (often called camp-followers), others along roadsides (ruderal plants), but their density is low in balanced ecological niches with no frequent disturbances. In Israel, bull mallow, wild barley and corn marigold are among many species that show a propensity to populate disturbed habitats. It is reasonable to believe that such propensities were also present in prehistoric times. According to this model, plants favouring these ecological niches would have been more competitive (reflected in opulence) than their counterparts, with a greater number of stems, leaves and yield (e.g., larger and more fertile barley spikes, more pea pods), and would therefore have caught the attention of site inhabitants and been adopted as candidates for husbandry.3 Randomness is a key in this model, too, as humans had no influence over the plants that favoured these man-made ecological niches. This model therefore does not explain the appearance or endurance of mutations leading to the domesticated morphology (phenotype) of the plants (the suite of phenotypic traits known as the domestication syndrome, a term we delve into below). The model thus explains only the reason for which certain species were identified and adopted at early stages of agriculture. An ecological affinity of plant species to disturbed niches is often referred to as weediness or ‘a weedy tendency’. However, we note here that the current agronomic usage of the term ‘weed’ refers to plant species that grow in arable land and cultivated fields and which interfere with the farming activities therein. Such species take advantage of the various human farm operations (e.g., ploughing, harrowing, removal of competitors). This ecological phenomenon is analogous to the occurrences described as part of the Dump-Heap Hypothesis along animal and human trails, around habitation sites and manmade refuse heaps (Box 15 Weeds, p. 89). The Dump-Heap Hypothesis was further supported by researchers who investigated plant domestication in the Americas, such as the domestication of sunflower or quinoa (also known as goosefoot). However, a deeper inspection of the ecological preferences of the wild progenitors of Near Eastern agriculture raises doubts concerning the applicability of the Dump-Heap Hypothesis to this area. Wild progenitors of emmer wheat, lentil, chickpea, pea and flax do not thrive in disturbed habitats, that is, they do not have weedy tendencies. Indeed, these species are not recognized in the area as weeds – not by modern agriculturalists nor by traditional farmers. Moreover, their presence is limited or nonexistent in areas that are disturbed by humans, along roadsides, or in areas that are intensively used as pasture. This fact poses a significant limitation on the explanatory

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power of the Dump-Heap Hypothesis regarding the origin of the Near Eastern crops and agriculture. In contrast, the wild progenitors of barley, einkorn wheat and bitter vetch may thrive in mildly disturbed niches (cultivated fields and roadsides) – wild barley (Hordeum spontaneum) is found in these habitats in the southern Levant while einkorn wheat is typical of such habitats in Turkey, Armenia and the Balkans. The wild form of bitter vetch is sometimes found as a weed in traditional pistachio plantations and farmed fields in the central area of its distribution in south-eastern Turkey. Although this fact initially seems to support the Dump-Heap Hypothesis, two additional criteria must be examined before we define these plants as weeds: (1) whether they are aggressive in their growth; and (2) their propagation coefficient, that is, the amount of seeds produced in each growth cycle. Given these two criteria, none of the founder crops in their original form, including barley, einkorn wheat and bitter vetch, can be considered a true weed.

Box 15 Weeds From an agricultural (weed science) perspective, weedy species are defined as follows: any plant that grows in an undesired location that interferes with or causes damage to human welfare. This definition includes both woody and non-woody plants, as well as annuals or perennials. Many researchers perceive weeds as a testimony to the presence of worked fields (or cultivation; Glossary, General Terms, Cultivation: plant husbandry) and agricultural activities. Identifying these plants among archaeobotanical remains and assigning them to specific species is highly challenging (see Box 13 ‘Atlit Yam for an example of successful identification, p. 71). The question arises: Where did these species reside prior to human preparation of plots for seeding and planting? Nature has its own share of unstable habitats in which available niches emerge and which pioneer plants (see Glossary, Botany, Ecology and Agronomy, Pioneer plants) invade. Some examples include outcrops that are exposed following landslides or river erosion, areas where young lava emerged near volcanoes, or areas characterized by shifting sands. It is likely that with the appearance of the first man-made agricultural niches, where early Neolithic farmers cleared the original

vegetation and introduced a sowing and planting regime, some plants established themselves in these niches and began to accompany the cultivated crops (later termed segetal plants). Weed population in cultivated fields is dynamic, and continuously changing according to the husbandry regime and diverse crops raised. For example, different weeds would characterize irrigated fields compared to dryland (rainfed) fields, while tree plantations would be characterized by different weeds than those typical of fields where different annual crops are grown in rotation. Modern dynamics of weeds is related to the fact that some weeds are invasive taxa (Figure 3.1). This often results from the transfer of millions of tons of produce (for both human and animal consumption) between diverse parts of the world. Although great efforts are extended to minimize contamination of grain shipments, it is impossible to guarantee the purity of freight and completely separate desired produce from undesired weeds. Thus, the Near East currently hosts weeds from the New World while California and Australia, for example, are now hosting many weedy species of Mediterranean origin.

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a

b

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d

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Figure 3.1 Cultivated fields disturbed by weeds. (a) Daisies in a vetch field; (b) thistle in a wheat field; (c) thistle and barley in a bread wheat field; (d) bindweed in a bread wheat field; (e) Diplotaxis in a pea field; (f ) oatmeal in a bread wheat field; photographs – courtesy of Baruch Rubin and Zvi Peleg. Colour versions of these images can be found at www.cambridge.org/abbo-gopher.

Therefore, the Dump-Heap mechanism (dubbed the commensal pathway), which relies solely on incidental developments such as the location of the plant and the extent of fertility of the land in its habitat, cannot explain the adoption of certain species by humans as food-producing crops in the Near East.

The Niche-Construction Model (or the Protracted Symbiosis and Domestication Model)

A modern model that has gained popularity in recent years describes plant domestication and the transition to agriculture as a gradual process that spanned thousands of years. Its roots are found in the routine food-collection activities of our ancestors that had nothing to do with plant domestication. This model describes the transition to a subsistence economy that was based on farmed plants as one stop along a long continuum of subsistence strategies that began with hunting and gathering and some simple actions aimed at increasing the effectiveness of gathering. Among these actions, which intervene with nature, were seasonal pruning and selective removal of plants and perhaps even sowing (of wild types) in suitable natural niches or plots prepared for the purpose. Removal of competing vegetation could be achieved by deliberately setting fires to burn undesired plants, for example, by Australian Aborigines. Additionally, selective (rather than exhaustive) harvest could leave enough seeds for natural dispersal and germination

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in the following season. Some experts in the field compare these human activities, which were carried out in order to increase the yield of wild plants, to animal actions (known as niche construction) that inadvertently influence the environment. For example, fruit eaten by bats and seeds distributed through their excretions result in intensive growth (a plantation of sorts) of the fruit-bearing species favoured by bats. Following the NicheConstruction approach, bats would be considered horticulturalists that encourage growth of certain fruit trees for the purpose of their subsistence. Similarly, the beaver also impacts its environment by building dams that slow the water flow in North American and European rivers while cutting down trees for this purpose. It therefore ecologically changes its environment to create its most favourable conditions. The Niche-Construction approach thus posits that organisms can, in fact, influence their own evolutionary trajectory. In contrast to Darwin’s epic model, where the environment influences, through different selection pressures, the fitness and survivability of different species, the Niche-Construction approach argues that organisms can change the selection pressures to which they are exposed, and thereby change (increase or reduce) the fitness of different individuals in the population. Organisms thus take part in the evolutionary process, a role that the classic Darwinian model has reserved to environmental (biotic and abiotic) factors imposing natural selection. A vivid debate takes place between the proponents and opponents of the Niche-Construction Theory (as summarized by Kevin Laland et al. and Gregory A. Wray et al. in two back-to-back comments published in Nature magazine in 2014). Its status and applicability to plant domestication notwithstanding, through its unique evolutionary perspective, the Niche-Construction Theory has certainly contributed to modern research on evolution. Applying the Niche-Construction approach to the origins of agriculture, no essential difference is considered to exist between those actions carried out by humans and those instinctively and routinely performed by different organisms. Increasing the yield of food plants (e.g., planting wheat or barley in some plots) is no different from the flower pollination of the hummingbirds, secretion of tree fruit seeds by bats, or fungi growing achieved by ants and termites. Therefore, proponents of this model emphasize the continuity spanning between different states, starting with the minute intervention in natural processes (as per hunter-gatherers) and culminating in different strategies to increase the yield of wild species that eventually lead to their domestication. Across this continuum of subsistence strategies, the interaction between humans and plants (domestication included) is viewed as a fluid and indistinct series of transitional situations in which the significance of wild foods slowly declined over thousands of years along with the rise in the significance of resources that were developed through environmental manipulation, while encouraging the proliferation of certain species. The process started with the encouragement of certain wild plants, passed through the deliberate cultivation of those wild species until, finally, domesticated plants were grown and consequently the agricultural system was established.

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The common denominator between the Dump-Heap Hypothesis, which attributes a great role to randomness, and the approach that views the transition to agriculture as a step in a process of instinctive actions (such as the bats’ secretion of tree fruit seeds), is their consideration of a long, protracted random and unintentional process. In neither model is there room – or need – for initiative on behalf of the grower, whose actions are perceived as automatic and instinctive, not the result of curiosity, expansion of knowledge and analysis, inference, strategizing or learning and application of knowledge from past experience. All this notwithstanding, the Dump-Heap Hypothesis allows for some observation and choice on the part of humans: they take notice of the exceptional growth in the unique niches and can choose to sample seed corn from relevant species for the purposes of sowing and grain production. The models described above focus on external drivers for change and the emergence of the Agricultural Revolution. In contrast, other theories assign plant and animal domestication as well as the emergence of agriculture to social-cultural or perceptual transformations4 that took place among hunter-gatherer communities living in the Near East towards the end of the Pleistocene and the beginning of the Holocene. Below we review a few such suggestions. The Competitive Feasting Theory

Some three decades ago, in 1990, the American anthropologist Brian Hayden suggested a model by which food production (plant and animal domestication as well as the onset of agriculture) was the result of intergroup (social) competition within local communities. The competition was expressed through expansive, festive feasting involving many participants and initiated by members of the community. These feasts were meant to enhance the status of the host and would have significantly impacted social status in the community or society. Requiring large quantities of food and thus the intensification of food production, these competitive feasts encouraged excessive food preparation, which eventually led to plant and animal domestication for the production of both foods and alcoholic drinks (such as beer), which were consumed during these feasts. The Ideological Model of Jacques Cauvin

Jacques Cauvin, a French archaeologist who worked for many years at Neolithic sites in Syria, suggested that a perceptual (ideological) change was required before the economic change, of which plant and animal domestication were but a part, could transpire. Cauvin argued that a changed ideology was a foundation necessary for the realization of the social and economic transformations that occurred during the

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Agricultural Revolution. Following this approach, a change in perception is a significant psycho-ideological adjustment in the relationship between humans and their world (culture–nature relations). Cauvin argued that human psychology during this period was preoccupied with supernatural elements, expressed symbolically in the figure of a woman and the figure of an ox, which was representative of the male. These expressions would later be institutionalized as the grounds upon which Near Eastern religions developed. According to Cauvin, this initial institutionalization of religion – one of the most significant and complex social structures in our lives – occurred alongside the institutionalization of the Agricultural Revolution. He based his arguments on archaeological findings reflecting different aspects of prehistoric life and especially noted the changes that had transpired in the symbolic arena, expressed mainly in artistic (symbolic) artefacts (imagery items) but also in architecture and other findings (such as hunting tools). Explanations such as this move further away from the approach that perceives the Agricultural Revolution as a random process; they emphasize decision making in face of a variety of choices and social and perceptual processes that led to change. Cauvin emphasized that ideology preceded economy; since food production was an antithesis of the hunter-gatherers’ ethos, without the ideological-perceptual developments that allowed hunter-gatherers to change their attitude towards nature and the world, the Agricultural Revolution could not have transpired. That is to say that change in human societies originates in new ideological-perceptual settings without which it would not have been activated. To clarify this point, readers may recall recent public debates in certain Western democracies regarding the role of the state in ensuring the welfare of its citizens, guiding the economic system and dividing public wealth, and the policy of deliberate departure from the welfare state model in some Western countries. Naturally, different parties to these debates hold fundamentally different worldviews (ideologies), which guide their actions. We believe that there is no reason to assume that prehistoric dynamics and social processes were very different in general than they are today. We have illustrated the dichotomy between models based on external factors (climatic, demographic or ecological fluctuations) driving the change in ancient subsistence strategies and those based on internal (cultural, social and perceptual) drivers of a process that was both informed and deliberate, accompanied by social-cultural transformations among local communities starting with the onset of the Neolithic period. A second dichotomy characterizing models explaining plant and animal domestication and the emergence of agriculture relates to time. Models that describe plant domestication as a fast, almost instant, transformation can be easily distinguished from those that perceive the process to have been long and protracted, occurring over thousands of years.

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O N T HE P A CE O F PL A N T D O M ES T I CA T IO N A N D TH E A W A R EN E S S O F T H E DO M E S T I C A T O R S

While those models advocating rapid domestication emphasize human agency and initiative (consciousness), others that describe a slow, protracted domestication process are based on the principles of the Niche-Construction Theory. Whether advocating a rapid process or a long one spanning thousands of years, both models base their arguments on the biology of the Neolithic founder crops. In our view, any discussion on the pace of plant domestication should involve the level of awareness that humans had while choosing the founder crops in nature and developing diverse ways to control their growth. Neolithic farmers’ awareness of processes that occurred in their fields is inseparable from this discussion. We thus posit that in answering the question of pace, relevant botanical, agronomic and genetic knowledge concerning these plants must also be considered. In many wild cereals, including wheat, barley, rice and sorghum propagules (seeds, or seed-bearing organs), dispersal occurs following the dehiscence (disarticulation) of the mature inflorescence (e.g., spike, panicle). Botanists and geneticists specializing in the domestication of cereals have suggested that domestication was triggered by a sowing and harvesting regime (annual repetition of harvest using a sickle followed by planting of seeds from the harvested plants) applied to a wild population of seeds (collected in its natural habitat from shattered spikes). This regime would lead, within several years, to an increase in the frequency of individuals that carry mutations in genes that influence the shattering of their spikes (panicles, etc.), favouring non-shattering types (see Figure 5.1). Simple consideration of population genetics shows that even a very small number of mutants among millions of seeds would eventually take over the entire population within a few years to a maximum of 200 years given an adequate harvesting and planting regime. This dynamic is simply explained, deriving from the fact that prior to or during harvest, some of the propagules produced by shattering spikes fall to the ground before ever reaching the bag from which the next cycle would be planted. If they are not ground into flour (or used in other ways), the grains of nonshattering spikes would end up in some storage installation at the end of harvest. In other words, grains from non-shattering spikes are less likely to be lost due to shattering and therefore have a higher chance to form part of the next season’s crop. Thus, the population faces a selection pressure that offers an advantage to non-shattering specimens over shattering ones (a larger portion of whose yield would always be lost). A similar selection process may be described for legumes with non-dehiscent pods, the yield of which would reach the barn in its entirety, compared to dehiscent pods, which are bound to suffer losses due to pod dehiscence and seed dispersal. This selection process, driven by the husbandry regime, is known as automatic selection (see above) as it requires no intent or attention on behalf of the farmer in order to reach the point where the great majority of the population comprises genotypes that lost their wild mechanism of seed dispersal, that is, that are characterized by the domesticated non-shattering traits.

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The literature on plant domestication also refers to this selection process as unconscious selection. This implies an underlying assumption about the lack of awareness of early farmers of processes and crop population dynamics that were taking place in their fields, that is, early farmers are assumed to have been unaware of annual fluctuations occurring in the frequency of certain spike or pod types as a result of their husbandry regime. Some researchers even claim that the Neolithic farmers were simply unable to distinguish between shattering (wild) and non-shattering (domesticated) spikes as the two are highly similar. This, in their view, justifies advocacy of a millennia-long, protracted domestication process. Since we have no direct evidence concerning the extent of awareness of Neolithic farmers regarding the different aspects of domestication and their agricultural practices where cereals are concerned, scholars and researchers must establish their own individual beliefs. Nevertheless, studying legume biology, and specifically seed dormancy of wild legumes, led Professor Gideon Ladizinsky of the Robert H. Smith Faculty of Agriculture, Food and Environment at the Hebrew University of Jerusalem to suggest in the late 1980s that legume domestication would have had to be a conscious and very rapid process. Seed dormancy rates in wild lentil amount to 90%, that is, 90% of the seeds would not germinate in the first year. Furthermore, wild lentil in nature yields an approximated average of ten seeds per plant. Therefore, following the planting of 100 seeds in the field, given 90% dormancy, only about ten plants will germinate, each yielding some ten seeds. In other words, by the end of the season (if all goes well) the farmer could expect a yield of 100 seeds, which is quite similar to the number of seeds planted to begin with. These conditions offer no incentive for planting since the work invested in preparing and maintaining the field or plot leads to no gain. Thus, Ladizinsky’s theory suggests that early Neolithic farmers had identified a natural lentil population characterized by a high rate of freely germinating individuals (mutant stock with relatively low seed dormancy), from which they took the initial seed pool that they would grow. This innovative suggestion was groundbreaking as it was the first time it was suggested that pulse domestication significantly differed from cereal domestication, a claim that was proved to require further attention. Ladizinsky named his provocative domestication model Pulse Domestication before Cultivation. His model, based on the biology of wild lentil species and agronomic consideration, was rejected by most experts in the field. However, our own students (Inbar Zezak, Yael Zehavi and Erez Rachamim) have conducted controlled experiments with three species of wild peas in Israel. During four years, experimental nurseries were sown in several locations in the country. Controls comprised the same pea species in which the seed coat was scarified (partial removal) as well as plots in which domesticated pea cultivars were sown. The scarified seed coat affects the germination rate, as seed dormancy in legumes (e.g., lentil, pea and chickpea) is caused by a water impermeable hard seed coat (seeds cannot absorb water from the soil). In contrast, after scarification wild seeds germinate at a rate of 95%, comparable to domesticated cultivars.

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Harvest was timed in each plot to a point where a few pods had already opened but most were intact and ripe, in order to minimize loss due to shattered pods and dispersed seeds. Findings were conclusive, showing that in the large majority of cases, seed yield from the wild types whose seed coats were intact was lower than the volume of seeds originally planted; that is, there was a deficit in seeds. Nurseries where wild types were sown whose seed coats were scarified showed high germination rates, and, by harvest, some pods had opened, allowing seeds to fall to the ground. Findings also showed that even after the loss of some yield (sometimes to a considerable extent) due to pod shattering, high germination rates guaranteed a significantly larger yield than the volume originally sown. In other words, in growing wild peas, free germination guarantees gain to the farmer. Naturally, non-shattering pods would increase the farmer’s gain, but this useful trait is not imperative to secure a productive crop. These experiments, then, support the claim that with respect to the grain pulses, free germination is the key trait without which profitable cultivation is impossible. They also lend support to the suggestion that an informed choice of a natural (free-germinating) source of seeds enables immediate domestication, rendering reliance on unconscious, long-term processes altogether redundant. These experiments also imply that it is unlikely that early Neolithic farmers would have invested years (tens, hundreds or thousands) to sow and treat fields of peas or lentils from which they would have consistently generated yields that were lower than their investment (i.e., consistently produced loss rather than gain) and yet persevere with the process until one field had unexpectedly produced a free-germinating seed yield, giving rise to our familiar domesticated pea. It is difficult to perceive that such a scenario would have lasted more than a year or two, and certainly not for numerous years. Any model that implies that these early Neolithic farmers had sown grain pulses for years without any gain also suggests that these farmers acted unintelligently and had no biological understanding of their environment or the ability to make simple loss and gain calculations. This, of course, stands in contrast to the knowledge that hunter-gatherers had concerning their environment, as discussed earlier. Alternatively, we may suggest that early farmers scarified their pulse seeds to ensure growth and gain as occurred in our control nurseries. This is technically possible, and it is also possible that the early farmers knew that seed scarification relieves dormancy, but then the question arises of how free germination is established in a population that is consistently sown following seed scarification. This would be puzzling since seed scarification results in germination regardless of the condition of the genes that control the water permeability of the seed coat and the dormancy trait. Therefore, even if a freegerminating mutant had spontaneously emerged in such a population, it would have been difficult to distinguish such individual(s) from other plants, all of which would have also germinated. It is very difficult to discern whether the initial choice of legume seeds to be sown had occurred as Ladizinsky suggested. Nevertheless, his model is significant as it offers a solid biological and perceptual alternative to those models that

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were based on notions of an unconscious and protracted process, and which lack support from the biological and agronomical perspectives.

K E Y PO IN TS A ND BE Y ON D

• Hunter-gatherers had extensive knowledge regarding plants and animals, and they meticulously used natural resources based on this knowledge in a manner that did not harm the future potential of these resources, unless some exceptional circumstances forced them to do so. • Hunter-gatherers maintained a continuous, detailed and rich discourse around natural resources. This discourse extended beyond that which is necessary for daily subsistence. Knowledge accumulation and sharing among community members (including young members) was part of the hunter-gatherers’ life-way; in addition to their economic significance, these practices were also embedded in the ideological and social levels of the hunter-gatherer culture. We argue that this accumulation of knowledge over millions of years of hunting and gathering was the foundation upon which Levantine Neolithic communities based their choices when opting for plant (and animal) domestication for the sake of food production. • The hunter-gatherers’ ethos as it generally emerges from anthropological studies of the last century would almost by definition oppose assuming control (or dominating) of plants and animals. Thus, there is no possibility for domestication if there is no change in perception. In this regard, we note that we are discussing here the very first domesticators/farmers, who had no model of anything comparable to guide them. • Attempts to understand and explain the Agricultural Revolution have been made for many years. On the one hand, descriptive models have attempted to reconstruct the unfolding of the revolution while, on the other hand, explanatory models have attempted to answer what drove the Agricultural Revolution in the first place. • Gordon Childe’s Oasis Theory is a somewhat naive narrative arguing that change was driven by an external force (dry climate), which led to a more intimate familiarity between humans, plants and animals living in rich geographic areas (the river valleys of the Nile, the Euphrates and the Tigris) and eventually also to domestication. Echoed in this narrative are ancient texts emphasizing the centrality of the large Near Eastern river valleys and the cultures that emerged on their banks. • The Dump-Heap Hypothesis is one of the earliest scenarios to have suggested the manner by which plant domestication was realized. Advocating an ecological mechanism, it thus attempts to answer the ‘how’ question of plant domestication. It offers a circumstantial justification for this process that is not concerned with human initiative but is rather related to plant traits and their ecological affinities. Consequently, according to this scenario, certain plants increased in frequency in locations where humans had disturbed the environment and its ecological balance and these species

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later became domesticated crop plants (dubbed ‘the commensal pathway to domestication’). Investigation of the plant biology of the wild progenitors of the Levantine founder crops refutes this hypothesis. Models by which agriculture emerged in core areas are based on the trivial assumption that it was the potential of plant species in these areas that allowed for their eventual domestication. In this sense, and based on these plants’ potential, it is obvious that the environment plays a significant role. Demographic explanations argue that plant (and animal) domestication and the intensification of production were the result of growth in the population, and were a response to the food crisis that these growing populations faced. While it remains unclear what came first, today it is commonly accepted that demographic growth was not the driving force for the Agricultural Revolution and domestication but rather their result. Voices advocating circumstantial reasoning for plant domestication (and the Agricultural Revolution as a whole) posit that the economic and social changes were the result of a prehistoric ‘accident’, an unguided, unintended and mostly unconscious development. In our view, this is a demeaning, condescending, unjustified approach. Nevertheless, the Dump-Heap Hypothesis, a representative of these approaches, has impacted subsequent scholarship, even among researchers that do not explicitly embrace it but have adopted some of its main tenets. Models explaining the emergence of agriculture based on the Niche-Construction Theory are yet another example of circumstantial explanations that regard plant domestication as a long, protracted process, on a (nearly) evolutionary scale. We view as groundless the analogy between plant-domesticating humans and activities of animals such as the beaver, the amoeba or fungi-‘domesticating’ insects. Social explanations for domestication and intensification of production are based on the competitive tendencies of individuals and groups in ancient human societies towards the end of the Paleolithic and in the early Neolithic periods. The process described as the drive for change is the emergence of competitive feasting, the materialization of which was dependent on intensified food production. In our view, this social drive is certainly a feasible and relevant component of the perceptual change that led to the domestication of plants and animals. Explanations tying plant domestication and the Agricultural Revolution as a whole with changed perceptions and ideologies may be considered poetic, or even naive, but they are not necessarily mistaken. History shows that many important large-scale processes (and revolutions) occurred, at least at face value, due to ideological reasons. We believe that a preceding ideological foundation and changed worldviews are an important infrastructure for any human activity, the Agricultural Revolution and domestications included. Why such a perceptual change occurs remains an open question and studies of the psychic or neurological perspectives of this change are only in their infancy.

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• Studying the biology and genetics of grain legumes in the Levant shows that legume domestication differed from cereal domestication because profitable legume cropping is dependent on the trait of free germination. Ladizinsky’s provocative domestication model posits that this trait was adopted by early farmers directly from wild types and was therefore the (conditional) starting point of pulse domestication. From an agronomic perspective, the increase in the frequency of free-germination types cannot develop through an evolutionary (long) process, rendering pulse domestication a rapid, revolutionary, episode. • In our view, the awareness of domesticating farmers was the starting point of domestication as is the assumption that their actions embodied intentionality. We perceive no other way for domestication to have occurred. It is likely that Neolithic people had not envisioned the many developments that followed plant and animal domestication and that they could not conceive the long-term, far-reaching consequences of their initiative. However, perceiving the Agricultural Revolution as a whole (and with it, plant and animal domestication) as the ‘biggest mistake in the history of mankind’, or in biblical terms, as the ‘expulsion from the hunter-gatherer heaven’ (see Box 2 From the fruit of Paradise. . ., p. 19), is an extreme negative approach, based on retrospective wisdom.

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4 T H E P L A N T F O R M A T I O N S O F T H E FE R T I L E C R E S C E NT AND T HE WI LD P R O G E N I T O R S OF T H E D O M E S T I C A T E D F O U N D E R CR O P S

The Fertile Crescent, named so after its lunate silhouette, spans from Khuzestan Province in Iran, across the Zagros Mountains in western Iran (Kurdistan), to the river valleys of the Euphrates and Tigris in Iraq, south-eastern Turkey and northern Syria, and then westwards towards Lebanon, the Mediterranean zone of Israel and Jordan and finally spanning southwards towards the Nile (see Figure Introduction 1). Geologically, most of the Fertile Crescent is covered by rocks that were formed at the bottom of the Tethys Ocean millions of years ago and soils that eroded from these rocks. Additional geological formations include extensive basalt flows characterized by the fertile soils they generate, outcrops of igneous rocks such as granites that are hundreds of millions of years old, and the sandstone deposits formed in the coastal areas of the ocean mostly during the Lower Cretaceous. Valleys of the region are characterized by deep alluvial soils and colluvial deposits from mountains, especially after humans cut down forests, thereby accelerating soil erosion. The coastal plains of the eastern Mediterranean are characterized by low ridges of beach-rocks (kurkar, solidified dunes), sandy loam soils (hamra) and dunes _ originating from quartz particles that eroded from the granite rocks of East Africa, and which were transported to the Mediterranean by the Nile and pushed further to the eastern coast of the Mediterranean by sea currents. The climate in the west and north of the Fertile Crescent is Mediterranean (Figure 4.1), characterized by two main seasons: a cool, rainy winter and a dry, hot summer; spring and autumn are relatively short. Fluctuating precipitation comprises mostly rain at an average of 400–900 mm annually and little snow, which is limited to high mountains or the northern part of the region. This climatic rhythm drives annual

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Figure 4.1 Vegetation belts (phytogeographic regions) in the Near East.

plants to grow mostly in the winter and flower mainly in the spring, whereas the dry season barely supports green annuals. Real forests are spread throughout the region; the southern, warmer area boasts evergreens, such as the oak, carob and olive, and the colder north hosts deciduous winter trees and various types of oaks and pines (Figure 4.2). East of the Mediterranean region are steppe lands known as the Irano-Turanian vegetation zone due to the Iranian and Turanian (central Asian) origins of major components of its plants. As in the Mediterranean zone, this area is characterized by a

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Figure 4.2 Mediterranean flora – courtesy of Zvi Peleg. Colour versions of these images can be found at www.cambridge.org/abbo-gopher.

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Figure 4.3 Irano-Turanian flora. A colour version of this image can be found at www.cambridge.org/ abbo-gopher.

cold winter and a hot summer, but here both are dry. The fluctuating precipitation averages 150–300 mm rainfall annually. Vegetation in this region is characterized by tall- and short-statured shrubs, deciduous winter trees and herbaceous annuals and perennials. There are no dense forests in the Irano-Turanian zone, and trees are distributed in fairly open park forest formations, that is, trees typically stand solitarily, separated by herbaceous flora, shrubs or dwarf shrubs. Typical deciduous species include Pistacia atlantica (terebinth), Pistacia vera (true pistachio) and diverse almond species (Figure 4.3). South and east of the Irano-Turanian zones, the Fertile Crescent borders desert lands, which are the northern extensions of the Sahara and deserts of the Arabian Peninsula. In this desert region, annual rainfall is less than 150 mm, and may even drop to as low as 50 mm. Vegetation in this hot, arid region is comprised mainly of herbaceous flora, shrubs or dwarf shrubs with few trees; the occasional trees are mostly found in wadis (Arabic, dry creeks with seasonal run-off ) that have collected run-off water and around springs. Species typical of this Saharo-Arabian region are various types of Tamarix and Atriplex that can secrete salt through their leaves and thereby survive in areas where the soil or waters are salty. Due to significant

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Figure 4.4 Saharo-Arabian flora. A colour version of this image can be found at www.cambridge.org/ abbo-gopher.

fluctuations in the little precipitation that characterizes the region, flora there is also subject to significant fluctuations (Figure 4.4). Mediterranean, Irano-Turanian and Saharo-Arabian vegetation elements often grow together in the Fertile Crescent. Following the establishment of farming societies across the Near East, the long tradition of hunter-gatherers’ minimal intervention in the natural vegetation was replaced by intensive and therefore destructive influence. This direct impact by cutting, burning, ploughing and uprooting and indirect damage through continuous pasturing of domesticated livestock (Figure 4.5) has changed, often to an unrecognizable degree, the natural vegetation and produced plant formations that are typical of disturbed habitats. Many of the plants boast biological mechanisms that confer resistance to grazing (thorns, toxicity, low growth or underground storage organs). Others have developed mechanisms that enable recovery following cutting and burning (e.g., renewal through branch buds, at stem bases or underground buds, seeds that are distributed or which germinate only after fire). And still other plants have developed diverse mechanisms that allow wild species to capitalize on cultivated lands (segetal plants) as well as fallow areas and field edges. The high degree of fertility of the river valleys in Syria and Iraq (Mesopotamia) at one end and Egypt at the other is not related to the meagre precipitation in these areas but to the fact that the water and minerals that reach these areas are from rainier regions of the

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Figure 4.5 Pasture in open lands. Colour versions of these images can be found at www.cambridge.org/ abbo-gopher.

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Euphrates and the Tigris or the Nile. In addition to the large rivers, the Fertile Crescent boasts many smaller rivers (such as the Orontes, Litani, Zaharani, Yarmuk and Jordan) and several lakes that are surrounded by fertile soils. In the past, however, flood plains adjacent to permanent bodies of water served as breeding grounds for mosquitoes that spread a malaria-causing parasite. In such areas, permanent settlements were rare and agricultural activity was restricted. The varied climatic conditions spanning from the Mediterranean through the semiarid (Irano-Turanian) steppes to desert lands result in a highly rich region in terms of floral diversity. For example, Israel, which spans a very small area in this region (little more than 20,000 sq km), boasts some 2,700 species of wild plants compared to, say, Scandinavia, which is fortyfold larger than Israel and in which only about 1,000 wild plant species have been recorded. Many plants in the Fertile Crescent are annuals and the proportion of annual plants in this region is considerably higher than it is in many areas in the world. In addition, in a disproportionate manner and due to unclear historical circumstances, the region boasts many of the wild annual cereals that have a relatively large seed compared to other regions of the world. Similarly, the Fertile Crescent hosts several wild legumes that have large, non-toxic seeds. In other words, the evolutionary history of the regional vegetation allowed for the botanical potential that facilitated the domestication of annual grain crops. A group of eight annual plants that originated in the Fertile Crescent have been collectively termed the founder crops of Near Eastern agriculture (see Table 4.1). The group is comprised of three cereals – einkorn wheat, emmer or durum wheat and barley; four legumes – pea, lentil, chickpea and bitter vetch; and flax or linseed, which is a source of fibre and oil. The group has gradually been recognized along with the development of Table 4.1 Founder crops of Near Eastern agriculture and their wild progenitors

Common name Einkorn wheat Emmer or durum wheat Barley Lentil Chickpea Pea Bitter vetch Flax or linseed Broad bean, horse bean, fava*

Scientific name of domesticated species

Scientific name of wild progenitor

Number of chromosomes

Triticum monococcum Triticum turgidum

Triticum boeoticum Triticum dicoccoides

14 28

Hordeum vulgare Lens culinaris Cicer arietinum Pisum sativum Vicia ervilia Linum usitatissimum Vicia faba

Hordeum spontaneum Lens orientalis Cicer reticulatum Pisum humile Vicia ervilia Linum bienne unknown

14 14 16 14 14 30 12

* We presume that Daniel Zohary and Maria Hopf did not include the broad bean among their list of Near Eastern founder crops probably due to the fact that its wild progenitor is not known to modern science.

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archaeobotanical research conducted at Neolithic sites in the region. Hans Helbaek was the first to note the cereals, pea and flax. Daniel Zohary and Maria Hopf later formulated the concept of the founder crops as it is known today, based on the frequency in the archaeobotanical record at the Neolithic sites of different species and their roles in traditional farming systems since then and to date in Turkey and the Near East at large. A brief botanical description of the wild progenitors of the founder crops follows. The wild progenitor of einkorn wheat is Triticum boeoticum, a typical element of park forests (woodlands) and grasslands in the northern Fertile Crescent. It is commonly distributed from the west of Anatolia through eastern Turkey towards Armenia and the Zagros Mountains in Iran and from Turkey in the north to the Lebanon mountain range in the south. This species flowers late and requires a cold period (see Box 17 Response to vernalization and control of flowering time, p. 130) to initiate spike development and bloom. It grows to a height of 1.5 m, with spikes 15–20 cm long; the grain protein content of this species amounts to 12%. Because of its developmental cold requirement for spike initiation, the domesticated forms derived from this species have spread mostly to Europe and other cool areas of the world. The wild progenitor of emmer (durum) wheat is Triticum dicoccoides, often called ‘the mother of wheat’. This species is common in grasslands in basalt-derived soils and rocky areas where soils formed from hard limestone (or dolomites). Its distribution spans both the eastern and western sides of the Mediterranean region of the Fertile Crescent – from Israel and Jordan, across to Lebanon, Syria and Turkey, to western Iran. Compared with the wild progenitor of einkorn wheat, not all populations of this species have developmental cold requirements, and often only to a limited degree. This has rendered durum wheat the prominent cereal throughout the Mediterranean to date. The species can exceed 1.5 m in height, and its impressive spikes, sometimes longer than 20 cm, bear grains averaging 40 mg in weight. The protein content of the grain is fairly high, amounting to 20% or even more. In Israel, large and dense populations of this wild wheat are known mostly in the north of the country – the Gilboa Mountains, Eastern Galilee, the Yahudia Forest Reserve in the Golan Heights and on Mount Hermon. Throughout its geographic distribution areas, this species is often accompanied by the wild progenitor of barley – Hordeum spontaneum, which spans a geographically wider region than either of the wheats discussed here. Wild barley spans from Cyrenaica Province in eastern Libya to central Asia. It mostly flowers earlier than wheat and can therefore avoid the early summer (terminal) drought in areas where the rainy season ends early. This trait has rendered barley an important cereal in areas with little precipitation, and especially at marginal desert areas at the rims of the Mediterranean zones of the Fertile Crescent. Wild barley also exceeds 1.5 m in height, and its spikes bear twenty-five to forty grains, averaging at 30 mg in weight and a protein content of 12–15%. The three founder crop cereals would have been used mainly as a source of carbohydrates and were most likely consumed during Neolithic times in some form of porridge or flat bread after being pounded and ground into flour.

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We do not know when baking, which would have required the ‘domestication’ of microorganisms, that is, yeast, began. The character of protein of the hard durum wheat grain (weak gluten that is unable to sustain gas produced by yeast fermentation) does not allow for the baking of the airy loaves favoured by Western present-day consumers. This is why many types of traditional breads throughout Asia and the Mediterranean region, where durum wheat was prominent, are flat (e.g., pitta breads, Turkish pide bread, pizza and so forth).1 The baking of airy breads became possible only after the appearance of bread wheat, which is a cultigen devoid of a wild progenitor. How is that possible? Cultivated fields usually host weeds (see Box 15 Weeds, p. 89) that originate from neighbouring plant formations that are region-specific. With the arrival of agriculture that was based on species found in the Near East towards the Caspian Sea north-east of Turkey, fields of domesticated emmer (durum) wheat were invaded by a wild relative of wheat, goat grass (Aegilops tauschii), which has seven pairs of chromosomes, similar to einkorn wheat. Spontaneous hybridization occurred (most probably more than once) in the Caspian Sea Basin between the ancient durum wheat and the goat grass. When such interspecific hybrids underwent chromosome doubling, the resulting progeny were in fact the species known today as bread wheat (Glossary, Genetics, Ploidy). This is a special case of the emergence of a domesticated species under human-made conditions rather than directly from a botanically and genetically similar wild progenitor. Since bread wheat emerged under domestication, it is not discussed in this book despite its enormous significance to humans both nutritionally and economically. The wild progenitor of the domesticated lentil (Lens culinaris) is the oriental lentil (Lens orientalis). This wild pulse grows in rocky and stony habitats at 300–1,700 m above sea level, characterized by shallow soils and found throughout the area between Israel and Tajikistan in central Asia. The Israeli populations of this species are small and sporadic whereas in eastern Turkey, the centre of the species’ distribution area, populations are dense and occur over larger areas. The species usually grows up to 20 cm in height in its natural habitat, yielding ten to twenty seeds averaging 10 mg in weight with a protein content of 25%. The wild progenitor of the chickpea is Cicer reticulatum, a species known only from stony habitats in south-eastern Turkey; it is found among deciduous oak and pistachio woodlands at an elevation of ca. 1,000 m above sea level. This species, too, does not grow taller than 20 cm in the wild and typically yields up to ten pods, each containing a single seed of 100 mg with a protein content of 25–35%. This Turkish species was unknown to the early students of Near Eastern plant domestication and was first described only in the early 1970s by Ladizinsky. The wild progenitor of the domesticated pea is Pisum humile. Compared to the three pulses discussed thus far, wild pea is taller and has the ability to climb trees and shrubs. As with the other pulses, this species, too, is prevalent in the Mediterranean zones of the Fertile Crescent, especially in the oak and pistachio woodland belt in Turkey, Syria, Lebanon, western Iran, Armenia and Turkmenistan at an elevation of 1,000 m above sea

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Figure 4.6 Wild legumes (pea, bitter vetch and chickpea) that had been domesticated in the Near East.

level as well as in limestone-derived soils, in basaltic areas and in granite rock formations. The large pods of this species each contain some eight seeds averaging 100 mg in weight and 20% protein content. In eastern Turkey, this species is often found alongside wild lentil and bitter vetch, all of which may easily be found growing together (Figure 4.6). The wild progenitor of bitter vetch is Vicia ervilia, a wild legume commonly found in stony habitats at elevations of 800–2,000 m above sea level in the Mediterranean vegetation zones between western Turkey and Armenia, western Iran and southern Turkmenistan and from Turkey to Mount Hermon in Israel and Jabel ed-Druze in Syria. In the wild, this species accompanies the oriental lentil, a plant which is slightly smaller than bitter vetch. Its pods contain some four seeds that typically resemble a pyramid in shape. In contrast to the oriental lentil and wild chickpea, this species is less susceptible to habitat disturbances and may flourish on field margins and sometimes even within fields, under conditions of traditional cultivation. An example of this can be seen in Turkish olive or pistachio groves (when only shallow cultivation is used) and as long as weed control by herbicides is not used in the system. The pea, lentil and chickpea are used as a source of vegetal protein, although seeds include a significant amount of carbohydrates and about 5% fat. Bitter vetch seeds are toxic to humans and are currently used to feed animals, typically in traditional agricultural systems in Turkey and across the Mediterranean Basin. The above notwithstanding, there exist some traditional Greek bread-making recipes in which bitter vetch flour is one of the ingredients. It is unclear whether the domestication of bitter vetch preceded the domestication of animals, or whether it was included in the package of founder crops simply because it is easily identifiable among archaeobotanical remains due to its unique seed shape (akin to a coin under the archaeobotanical lamp).

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The wild progenitor of flax (Linum bienne) is well known throughout the Mediterranean Basin as well as east of Turkey; its distribution, like that of barley, is thus considerably wider than most other species discussed here. In Turkey this is typically a tall plant, reaching about 1 m in height and boasting light-blue blossoms. Unlike the cereals, but similar to the legumes mentioned above, this plant grows in sparse populations, typical of rocky soils in woodland formations. Today, flax is used as a source of fibre and the high-quality oil produced from its seeds. It is likely that during the Neolithic, too, seeds were eaten in various forms and fibre was produced from the plant’s stem. As indicated in the note to Table 4.1 (see p. 106), no known species has been identified as the wild progenitor of the broad bean. This includes even several species among the Vicia genus with some morphological resemblance. None of the tested species have a chromosomal constitution similar to that of the broad bean. And likewise, none of the tested species were cross-compatible with the broad bean, which is the decisive and ultimate test for establishing such ancestry (see below). Identifying the wild progenitors of the founder crops involves botanical and genetic criteria. Botanists first have to identify a species botanically reminiscent of the domesticated form in terms of the shape of reproductive organs (flowers), fruit and seeds. This is sometimes very easy, as in the case of wild and domesticated barley, which are both well known to generations of botanists, as well as in the case of bitter vetch. In contrast, the wild progenitor of chickpea was only discovered some fifty years ago when the correct species was found and described. For some plants, such as faba bean (also known as broad bean or horse bean, which appears in archaeobotanical assemblages of Neolithic sites as early as the EPPNB; but see Chapter 8, note 1, p. 240), the absence of a sufficiently similar wild form led some researchers to conclude that the wild form is now extinct. It is nevertheless possible that the scarcity of the progenitor hindered identification and that botanists have yet to investigate its habitat in greater depth to uncover it. In addition to botanical resemblance, genetic evidence is also required for the identification of the wild progenitor. The most essential and important genetic index is the crossability potential (hybridization capabilities), that is, the candidate wild species must be able to produce fertile offspring when crossbred with the domesticated form. Following hybridization, the development of the hybrid offspring is examined for regular development and its reproductive potential, which is dependent on the chromosomal homology (similarity) of the two parent species. If the domesticated species has the same number of chromosome pairs and their chromosomal organization is similar, chromosome pairing would be full, leading to regular meiotic (reproductive) divisions of chromosomes, which leads to the formation of pollen grains and eggs in the floral organs of the hybrid plant. This enables the development of male and female reproductive cells, thereby allowing for the flower fertilization processes to occur, after which normal, viable seeds will develop. In contrast, differences in the chromosomal arrays of

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the two species will result in irregular divisions of the chromosomes in the meiotic cell divisions leading to pollen grain and egg formation, thereby impairing the fertility of both pollen and eggs, which usually leads to partial or total sterility of the hybrid plant. For example, five wild species of lentil are known, two of which are quite similar to the domesticated form based on common botanical criteria. However, in crossbreeding the different wild species with the domesticated lentil, only one species (the oriental lentil) results in fully fertile hybrids in which the chromosomal organization allows for the regular division of chromosomes and proper formation of sex cells. This is the reason why the oriental lentil was recognized as the wild progenitor of the domesticated lentil. Three known species of wild pea are another example. Two of these are morphologically similar to the domesticated form, but only one species results in a fully fertile hybrid offspring. This is also how the wild progenitors of domesticated emmer wheat (first recognized in the field by Aaron Aaronsohn in 1906) as well as barley, chickpea, bitter vetch and flax were discovered (see Figure Introduction 2). Current genomic technology allows for the examination and comparison of DNA sequences of the species under investigation. While DNA sequence comparisons provide important estimates of genetic relatedness based on statistical computation, examining chromosomal pairing during reproductive cell division offers conclusive inference on crossability relations that cannot be obtained from DNA data alone. Innovative DNAbased methods can nevertheless be used in the attempt to locate wild populations that are the closest relatives, genetically speaking, of any given domesticated species. This type of investigation was carried out in the three cereals in the founder crop package: einkorn and emmer or durum wheats as well as barley, and two legumes: pea and lentil. We further examine this issue in Chapter 10, where we discuss the question of where in the Fertile Crescent domestication first occurred.

K E Y PO IN TS A ND BE Y ON D

• The Near East is typified by limestone bedrock with bursts of volcanic matter (basalt) in different areas as well as ancient igneous outcrops that emerge in fairly small areas. The varied scenery comprises steep altitudinal and precipitation gradients, as well as extreme temperature differences. These are expressed in relatively small geographic regions, akin to a laboratory setting. • The Near Eastern climate is characterized by a short, cool rainy season, and a second dry, hot season. This climatic rhythm selects for a multitude of annual plant species. The gradient of precipitation and the diverse rocks and soils all encourage a mosaic of rich flora populations including diverse dense forests, woodlands (park forests), grasslands and steppe formations. • The wild progenitors of the founder crops of the Near East are all typical of the eastern Mediterranean flora, and they usually have no affinity with disturbed habitats (niches

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in which the ecological balance has been disturbed; see Glossary, Botany, Ecology and Agronomy, Ecological niche and Box 15 Weeds, p. 89). • The wild progenitors of the founder crops have been identified based on botanical similarities and genetic affinity by hybridization tests between the domesticated forms and their candidate wild progenitors as well by comparing DNA sequences.

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5 THE DIFFERENCE BETWEEN WILD AND DOMESTICATED PLANTS

In this chapter, we discuss the difference between wild and domesticated plants, focusing on those crops (wheat, barley, lentil, pea, chickpea and flax) that were domesticated in the Near East during the Neolithic period. We first review the traits that endow wild plants with selective advantage (increased fitness) in their natural habitats and then we look at domesticated plants through the prism of that list of traits. One fitness mechanism is seed dispersal. To ensure their survival over evolutionary time, wild plants require an effective mechanism of seed dispersal. By mechanism we mean a trait that is governed by one or more genes. An example of such a mechanism is the tuft of hairs (pappi) characteristic of the dandelion blowball that helps the seeds blow in the wind and spread. Devoid of an effective seed dispersal mechanism, ripe seeds fall at the foot of the parent plant so that during the coming rainy season, all seeds compete over the limited habitat resources. This considerably reduces chances of survival in semiarid regions such as the Near East. In contrast, an effective seed dispersal mechanism minimizes competition and allows the species to launch in a variety of habitats while broadening its distribution. Another fitness mechanism is seed dormancy, which is prevalent among the region’s legumes but is less important for cereals. In the bi-seasonal Mediterranean climate of a rainy winter and a dry summer, wild plants that exhibit a mechanism that delays germination are at an advantage. This mechanism facilitates coping with early rains during the autumn, after which precipitation stops for a long period of time – a typical climatic feature of the eastern basin of the Mediterranean. Under such conditions, seedlings that are not deeply rooted are likely to dry out and die. Therefore, a low

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germination rate allows dormant seeds to germinate and develop after the resumption of the rains later in the winter; this ensures plant establishment and later flowering and seed maturity at the end of spring. Early cessation of the rainy season is another frequent cause of severe droughts across the east Mediterranean Basin. Under such circumstances, many species wither immediately prior to (or upon) flowering and fail to produce any seeds. Thus, seed dormancy that persists for two to three years is a trait, or physiological mechanism, that encourages the accumulation of a ‘seed bank’ in the soil, thereby ensuring germination in years following extreme drought, in which seed production is often either low or nonexistent. Many wild plants have different mechanisms such as thorns or toxins to defend against pests and predators. These mechanisms reduce damage caused by grazing herbivores. Given that gazelle, fallow deer, aurochs and other herbivores grazed during the Neolithic, these mechanisms offered plants an advantage under such conditions. Wild plants exhibit many mechanisms that offer fitness with respect to the climatic rhythm prevalent in their habitat as well as genetic systems that ensure timely germination, adequate development and well-timed flowering. The development of cool-season cereals in the Near East (such as wheat and barley) is influenced by the length of day and temperature. Simply put, this influence dictates that following germination, at the beginning of the growth season, plants will develop leaves but their stems will not lengthen until days are long enough to facilitate differentiation (Glossary, Genetics, Cell differentiation) of the shoot apex to form a spike. Only after differentiation of the spike can stem elongation occur and the plant canopy develop, parallel with spike development to flowering in the early spring. This mechanism ensures that flowering will not occur too early in the season when exposure to sub-0 C temperatures might cause spike sterility. By ensuring that the flowering does not occur too late, this mechanism causes seeds to develop at the end of spring before the soil dries after the rainfall has ceased. Ecologists might say that the plants’ response to day length and temperature thus allows them to time their flowering to the optimal time slot (per season) in order to ensure the production of vital seeds. Typically, populations of wild plants boast genetic diversity that generates different plant types within each species, distinguished from each other by various traits (height, colour, seed size, resistance to disease and pests and so forth). This genetic diversity serves as a security (insurance of sorts) that allows the species to contend with fluctuations in its habitat. Since the Near East experiences many seasonal fluctuations (e.g., early or late end of rainy season, moderate or extra hot spring), wild populations are dynamic in terms of their genetic repertoire, which changes on a seasonal basis according to the prevalence of disease, pests, predators and especially the unique climatic profile of the season. Thus, in a season when rainfall ends early, the majority of seed yield will originate in early flowering plants, while in a season where rain still falls in April or May, considerable yield will also be produced by late-flowering individuals. Over time this leads to a fluctuation in ratio between early flowering and late-flowering

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types among the population. Genetic variation can also increase the ability to respond to mild climatic fluctuations. The significance of this genetic variation is appreciated when one understands that adaptability of wild and domesticated plants cannot be discussed at the level of the individual plant but only at the level of the population at large. To clarify, imagine a population of wild wheat in which an individual had randomly emerged (due to mutation or a new genetic combination) that is highly resistant to water shortages. At face value, one might think that the chances for this specimen to ripen its seeds in a drought year are significantly higher than for others that are less resistant to water shortage. However, this resistance set aside, there are other factors that will influence the chances for resistant or sensitive plants to survive and reproduce. Such factors include the presence of pathogens that cause leaf disease or the presence of a grazing gazelle herd that feeds upon spike-developing plants and is probably unaware of the difference between those plants that are resistant and those that are susceptible to water shortage. Thus, the survivability of the individual plant is dependent not only on its genetic constitution but also on random factors (such as the presence of a grazing herd). It is therefore clear that adaptation (in general and to the environment) can only be discussed in the context of incidence among the population of individuals that have a different genetic make-up (and therefore express different traits) and their relative productivity rates in response to the environmental challenges that they face. This is what we mean when we talk about genetic variability within populations that is found across the landscape over time (years). At the level of population, wild species usually occupy diverse habitats, each hosting plants of various botanical families, both annuals and perennials, with different seasonality. For this reason, the different plant groups maintain long-term reciprocal relationships among themselves as well as with animals and microorganisms that also subsist in these ecological niches. Many reciprocal influences can be found among natural plant communities, some beneficial to the parties (such as nitrogen-fixing bacteria in legume roots) while others are damaging (such as pathogens that harm the plants and their seed yield). In contrast to natural habitats, Near Eastern agriculture usually involves fields in which each crop is sown separately and mixtures are found only in special cases. When discussing domestication and domesticated plants, experts usually refer to the term domestication syndrome (see Box 16 The domestication syndrome, p. 116). When Near Eastern domesticated crops are examined vis-à-vis the above criteria, it is evident that with hardly any exception, domesticated species lack a seed dispersal mechanism (non-shattering cereal spikes and pod shattering resistant legumes) as well as a seed dormancy mechanism. Thus, given appropriate moisture, seed germination is made possible shortly after ripening and at a high rate, so no mechanism is set to guarantee long-term regulation of germination. While these two qualities are considered most important in Near Eastern domestication, the ease and convenience of cultivating and produce processing have also been emphasized, resulting in easy threshing of domesticated plants compared to their wild counterparts (i.e., the ability to separate

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Box 16 The domestication syndrome A syndrome is a collection of symptoms that occur together and can be used for the purpose of diagnosing a biological condition. With respect to plants, the term is generally employed to describe suites of traits that are visible (morphological) or sensed (smell, taste, aroma, chemical composition), as well as other traits (physiological, chromosomal) of domesticated plants by which they may be distinguished from their wild progenitors (e.g., the wild and domesticated wheat or pea). The domestication syndrome concept was developed by Dr Karl Hammer in the mid-1980s following the recognition that domesticated plants share many traits that set them apart from wild types. For example, all Near Eastern (domesticated) crops lack an efficient seed dispersal mechanism, so that a ripe plant still bears its seeds – legumes in closed pods, cereals in whole spikes and flax in closed capsules. In contrast, dispersal of ripe seeds in the wild occurs through shattering wheat spikes, dehiscent or fallen legume pods and open flax capsules. Other qualities that are often considered part of the domestication syndrome are seed dormancy (free germination in domesticated plants), the size of the organ that has economic significance (e.g., the seed; seeds of wild plants are usually smaller) and the

absence of thorns and toxicity (both of which may be present in wild forms). The term ‘domestication syndrome’ was originally developed to describe grain crops such as wheat, barley, beans or vegetables. The criteria developed by K. Hammer may still require further examination to suit other crops. For example, potato or sweet potato, leafy crops such as lettuce, kale or cabbage and spice crops such as basil, thyme or mint each necessitate a different set of characteristics, and it is sometimes difficult to find actual morphological differences between wild and domesticated forms. We emphasize that in contrast to the traditional use of the term ‘domestication syndrome’, we do not discuss traits that have changed throughout the plants’ evolution under domestication (that is, after domestication). In our view, domestication syndrome traits should be divided between those that were necessary for the purpose of successful cultivation and domestication (such as free germination of legumes) and those that are useful to the farmer but were not mandatory for domestication (such as non-shattering pods in legumes). This is not merely a semantic distinction but rather a distinction that embodies many advantages – from both the cultural and biological perspectives – to students of plant (and animal) domestication.

the grains from the chaff, or the grain from the straw and husk, which, in wild types, is a difficult task). The nutritional value of the seeds is also important. For example, protein content of domesticated cereals is lower than that of wild types whereas domesticated legumes vary in their differences compared to wild types. Of special note is chickpea in which the amino acid of tryptophan increased significantly following domestication. Other qualities of domesticated plants, which are not unique to the founder crops discussed throughout this book, are related to reduced toxicity (e.g., a wild almond may cause cyanide poisoning), the adaptation of the growth cycle to new habitat(s) or the work cycle of the farmer, the preservation potential of reproduction units (seeds, bulbs, tubers) from one season to the next, the competitiveness of the plants vis-à-vis

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Table 5.1 Summary of main fitness characteristics in wild and domesticated plants Wild plants

Domesticated plants

Effective seed dispersal

Prevention of spontaneous seed dispersal

Ensuring long-term germination potential through seed dormancy

Ensuring free and uniform germination

Defence against pests and predators by means of thorns and toxicity

Lessened extent of thorns and toxicity

Adaptation to climatic rhythm

Adaptation to local climate or farming induced environmental changes

Extensive genetic variability

Limited genetic variability

Co-evolution with sympatric species*

Competitiveness in dense plant communities (wheat and barley)

Not applicable**

Nutritional value (protein content, ease of digestion)

Not applicable**

Economic value of different plant organs

Not applicable

**

Ease of produce processing and use (threshing, storage, cooking)

* See Glossary, Botany, Ecology and Agronomy, Sympatric and allopatric distribution. ** In the long evolutionary history of wild plants, and certainly in the time phases that preceded plant domestication, these qualities were not relevant.

neighbouring plants and weeds, and the economic value of the plant (grains consumable by humans and animals or straw for construction and basketry). Table 5.1 shows that the qualities listed here, and especially the first two, mirror each other (e.g., seed dispersal and dormancy in wild types vs non-shattering and free germination in domesticated types). Another way to define the domestication syndrome is that it comprises a group of traits that are essential for the survival of plants in the wild that were genetically altered to ensure desired yield and convenience of processing under domestication. In a different version, these characteristics are absent from wild types and are therefore new, rendering the plant dependent on humans for its existence and distribution (especially given the impaired seed dispersal and dormancy mechanisms). These definitions of the domestication syndrome are not universal and cannot be applied to all domesticated plants. For example, it is customary to consider the prevention of inflorescence shattering in cereals (wheat, barley, rice) as the central characteristic that transformed with domestication and secondary to it the ease of threshing (the actions required to separate the grain from the chaff ). However, based on both theoretical considerations and following legume-growing experiments, it may be said that in pea, chickpea and lentil, the elimination of seed dormancy is considerably more important than the prevention of seed dispersal. Thus, a unique list of domestication traits should be compiled for each examined species based on its singular biology (perennial or annual), the quality of the organs in which the farmer is interested (dry seed, fresh fruit,

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ripe nut, edible leaves or stems, tubers for cooking, bulbs, etc.) or other criteria. Thus, the importance of the term ‘domestication syndrome’ lies in its setting a good starting point for the biological study of domesticated plants and their wild relatives and the comparison of each test case to known species. Many traits found under the conceptual framework of the domestication syndrome are recessive (see Glossary, Genetics, Recessive trait), where the wild type exhibits dominance (see Glossary, Genetics, Dominant trait) (as the pink flower allele is dominant over the white allele in Gregor Mendel’s peas and the brown eye allele in humans that conceals by its darker pigment product the expression of blue eyes). For example, free germination in domesticated lentil or chickpea is a recessive trait as is the nonshattering domesticated wheat and barley spike. As explained below, the floral biology of the Near Eastern wild progenitors facilitated the rather easy isolation of such agronomically desirable types out of wild stocks characterized mostly by dominant traits. The wild progenitors and the domesticated founder grain crops in the Near East are all self-pollinating. Thus, with rare exceptions of cross-fertilization, seeds in these plants are created following the merging of the male and female germ cells that share an identical genotype (see Glossary, Genetics, Genotype). The mechanism that ensures selffertilization in these species is based on the opening of the anthers and the release of pollen grains prior to flower opening, so that pollen grains begin to germinate on the stigma before any other pollen (from different plants) carried with the wind or by pollinators (e.g., insects, birds) has a chance to reach the flower. This mechanism ensures that in these species, other than in rare cases of mutation, the genetic pool of offspring is identical to that of the mother plants. Thus, self-pollinating plant populations are really a collection of pure (true breeding) lines (see Glossary, Genetics, Homozygous) that may differ genetically from each other but only rarely would they randomly mate. Over fifty years ago, Daniel Zohary noted in this respect that such a genetic structure is particularly suitable for plant domestication because the different types found among the population are mostly genetically isolated from each other. Moreover, given that self-pollination is the least common state among the plant kingdom (which evolved different mechanisms to ensure cross-fertilization), Zohary wondered about the chances of all early domesticated species in the Near East having been selected from self-pollinating populations. Naturally, the chances are low, and Zohary regarded the mode of reproduction a crucial consideration in the choice of species that would be domesticated (see more on this in Chapter 9). No evidence exists to support Zohary’s retrospective analysis that pollination biology guided the domesticators to have chosen the eight founder crops. Nevertheless, we accept Zohary’s reasoning. We believe those farmers knew that certain plant species produced offspring that were true to type while other (cross-fertilizing) species had more diverse offspring, and that they consciously chose to domesticate plants of the first group, even if they did not fully understand the genetic principles or the biology of pollination and their relation to offspring configuration.

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The genetic architecture of the domestication traits combined with the genetic population structure of the wild progenitors of the crops as well as the fact that these were self-pollinating species enabled early farmers to easily and quickly identify and isolate the desired plant types as domestication candidates. How so? As mentioned, the non-shattering of wheat and barley spikes is a recessive trait. In other words, the wild expression of this trait is governed by dominant alleles (see Glossary, Genetics, Dominant trait). Therefore, when the gene that controls spike-shattering is mutated in the reproductive cells of a wheat plant, it will not prevent the expression of genes (in cells of its spike) that are required for shattering. Moreover, in a plant that bears both the wild allele (see Glossary, Genetics, Allele) and the mutant allele (in all of its cells, somatic and reproductive; see Glossary, Botany, Ecology and Agronomy, Somatic tissues), the presence of that single wild allele is sufficient to induce shattering. If a pollen or egg cell carrying a mutant allele participates in the fertilization process resulting in a vital seed, the ensuing plant will be heterozygous, carrying a single domesticated allele and a single wild allele (see Glossary, Genetics, Heterozygous). This plant will perform as a wild type because the wild allele is dominant over the domestication allele. The expression of a non-shattering spike trait requires that the mutant allele be present in two copies, and, indeed, given a single generation (following self-pollination), about a quarter of the offspring of that heterozygous plant will perform as domesticated types (Figure 5.1) and half of the offspring will be heterozygous. The next generation of the heterozygous offspring will similarly produce 25% of domesticated-performing plants. Thus, in self-pollinating plants, once a gene that controls any recessive domestication trait has been mutated, the frequency of domesticated-performing individuals will rise incrementally within a few generations (years), rendering its identification (and selection for it) easy. In contrast, among plant populations characterized by a high rate of random cross-pollination, plants that are homozygous at one certain chromosomal site (e.g., host two copies of the recessive allele of non-shattering spikes) are exposed to massive quantities of pollen that is produced by wild types and therefore bear dominant alleles governing the shattering trait. Therefore, the majority of offspring of these plants, pollinated by random pollen, are expected to perform as the wild type (shattering spike). The fact that the rates of cross-fertilization events are marginal in self-pollinating plants decreases the extent to which the activity (performance, phenotypic expression) of the recessive allele is masked by the dominant allele in such populations. As noted, this offers the farmer a great advantage. Early fields were likely small in size compared to the natural habitats in which massive populations of the wild progenitors of the domesticated plants were growing. Thus, had the discussion involved cross-fertilization species, where most pollination events entail pollen that has originated in wild types (and so most offspring would also perform as wild types), it would have been difficult to isolate true breeding lineages of domesticated types desired by the farmer. Despite this, plants, such as maize in Central America and rye, which probably originated in

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BrBr A plant with a brittle spike phenotype (the wild type condition), with a BrBr (homozygote) genotype.

Brbr

A random mutation leads to a heterozygote condition with a Brbr genotype, but a wild (brittle spike) phenotype due to the dominance of the wild allele Br. The progeny of self-pollination are obtained from different combinations of pollen grains carrying a Br or br genotype and respective egg cells with a Br or br genotype. Therefore, non-brittle spike (domesticated phenotype) individuals are expected to make about 25% of the progeny. Following another generation of selfing, the proportion of non-brittle spike individuals will increase significantly. The rise in the proportion of the non-brittle plants is indicated in the table under the assumption that each of the parental genotypes yield a similar number of seeds. And likewise in the following generations.

pollen

Br 50%

br 50%

egg-cells

Br 50%

brittle spike

BrBr brittle 25% spike

Brbr 25%

br 50%

brittle spike

Brbr non-brittle brbr 25% spike 25%

BrBr 25%

Brbr 50%

brbr 25%

100% wild

25% 50% 25% BrBr Brbr brbr

100% domesticated

In each generation of selfing, the progeny of the heterozygotes' group will constitute 25% of wild-type (brittle spike) homozygotes, 25% non-brittle domesticated ones, and 50% heterozygotes. Therefore, the frequency of heterozygote individuals will decrease by 50% in each generation, with a respective increase in the proportion of homozygotes. A simple calculation will indicate that after ten generations of selfing about half of the population will show a domestic (non-brittle) spike phenotype, the other half will retain its wild type (brittle spike) phenotype, with a negligible number of remaining heterozygotes.

Figure 5.1 The inheritance of recessive traits in self-pollinating plants and the emergence of the domesticated cereal spike morphology. A colour version of this image can be found at www .cambridge.org/abbo-gopher.

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Turkey, were domesticated by farmers in ancient times. This fact further emphasizes the skill and competencies of early farmers while at the same time it slightly lessens the significance of the fact that all founder crops in the Near East are self-pollinating. K E Y PO IN TS A ND BE Y ON D

• Wild plants are dynamic entities that show diverse fitness and adaptation mechanisms that are genetically controlled. Among the most important of these adaptations are the mechanisms that ensure seed dispersal and mechanisms that delay seed germination. • Domesticated plants differ from their wild progenitors. In the case of Near Eastern grains, the most conspicuous differences involve the loss of the mechanisms that ensure dispersal and seed dormancy. The collective term describing the multitude of differences found between wild and domesticated plants is known as the domestication syndrome. This suite of differences may not be generalized, but rather specific definitions must be developed for different species based on their unique biology. • Other than broad bean and rye (that are usually not considered among the founder crops), all Near Eastern grain crops are self-pollinating plants, a fact that facilitates the identification and maintenance of desired mutants.

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6

T R A D I T I O N A L VE R S U S M O D E R N A G R I C U L T U R E – STABILITY VS MAXIMIZATION

To better understand the term ‘domestication syndrome’ (the group of traits that differentiate wild plants from domesticated crop plants; see Box 16 The domestication syndrome, p. 116), a deeper investigation into the differences between natural habitats, where wild species grow, and cultivated fields is required. In the agricultural context, actions taken are typically aimed at ensuring crop yields. These (husbandry) activities involve efforts that alter soil conditions, for example ploughing that aerates the soil, removes weeds and prepares it for sowing. To protect their fields and produce, farmers often create defensive systems (such as fencing) to minimize damages incurred by grazing wild animals (and later pasturing domesticated animals). Additionally, farming activities and crop growth may take place in a seasonal cycle that is different from that of wild populations. For example, under the Mediterranean climate, wild cereals begin to germinate immediately after the first autumn rains (October in the southern Levant, slightly earlier in the northern Levant). However, due to the absence of machinery with which to plough clayrich soils prior to wetting of the soil profile in early winter, farmers in this area, working the land in traditional ways, would not have been able to sow before the month of December. The farming package may thus even have included completely off-season operations, such as the sowing of chickpea in the spring (further discussed below). To reduce competition over water and other resources between crops and wild weeds, farmers often sowed crops quite densely. This density of genetically similar or identical plants facilitated the spread of epidemics of different harmful agents – a phenomenon that is atypical of wild populations due to the wide species

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diversity in any given habitat and the genetic heterogeneity that characterizes each species. Later agricultural developments of irrigation and fertilization deepened the distinction between cultivated and natural habitats. Since we have no direct evidence regarding the actions of the first Neolithic farmers, we are unable to characterize the farming environment that became the arena in which plant domestication took place. This is why experts in the field tend to examine traditional agricultural systems, which are considered to be the closest in nature to the Neolithic crop husbandry. Traditional agricultural systems were prevalent throughout the inhabited world until the early twentieth century. Accelerating modernization during that century in combination with the developing intercontinental transportation system and globalization of recent decades have caused traditional agricultural systems to degenerate to the degree that they no longer reflect past agricultural systems. Even in remote areas in Asia, South America or Africa, current agricultural systems only partially reflect the past. Despite this, traditional agricultural systems have been documented in the literature. In Israel, for example, Yitzhak Elazari-Volcani (Wilkanski), an early Israeli agronomist who was among the planners of modern agricultural settlements in the Land of Israel, documented in his book The Fallah’s Farm early twentieth-century agricultural practices and subsistence of the Arab fallaheen (peasants) before these were modified by subsequent British man_ date modernization and the intensive Jewish agricultural settlement efforts that continuously grew in scale. The current landscape observed in Judea and Samaria as well as other regions in Israel and throughout the Mediterranean Basin offers a glimpse into the impact of modernization on traditional agricultural practices in these regions. A major scenic marker of traditional farming – terrace farming – has recently been changing. Terraces, agricultural steps typical of mountain terrains throughout the world, are an important component of the physical infrastructure of traditional farming as they prevent soil erosion and facilitate the utilization of slopes for the purpose of agriculture. Terraces turn the slope into a continuum of small, levelled areas that are easy to work, sow or plant and in which movement is made convenient. Terraces are typically built as dry stone walls with no additive (cement) and therefore require ongoing maintenance to prevent collapse and consequential soil erosion. The decreasing profitability of agriculture compared to other vocations has driven many farmers to abandon their terraces, and, indeed, forsaken, crumbling terraces can be observed in the Galilee (northern Israel), Judea and Samaria. In some of these areas, new terraces are being constructed using mechanical tools that stabilize the land based on large boulder rocks, thereby creating units that may be worked with tractors instead of the traditional, narrow terraces that allow for only manual labour or perhaps animalassisted labour (e.g., with mules or oxen). The dynamics of these agricultural systems and related scenery in different areas of the world (e.g., the Balkans, the Maghreb, China to name but a few) is closely related to the impact of social and economic

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Table 6.1 Differences between traditional farming and modern agriculture Traditional farming

Modern agriculture

Goal: Provide local subsistence economy

Goal: Supply the food industry, local and far-flung markets

Strategy: Favouring stable yields

Strategy: Maximizing economic yield

Diverse crops grown in small plots

Specializes in few crops raised in large farming units

Heterogeneity of species, also known as polyculture (see Glossary, Botany, Ecology and Agronomy, Polyculture)

Genetically homogenous species, also known as monoculture (see Glossary, Botany, Ecology and Agronomy, Monoculture)

Manual or animal-assisted farming

As mechanized as possible

Necessitates minimal input

Necessitates intensive, and especially energyintensive, input

Necessitates minimal infrastructures

Necessitates intensive infrastructures, particularly for transportation

transformations, and their accelerating rate. Traditional agriculture was also practised over broad valleys or levelled planes; however, following the introduction of modern technologies in most areas, such landscapes have since changed. In Israel, a few remains of those past systems, marked by dry stone walls, watch-towers and various installations constructed from stones that were removed from worked fields, can still be seen in the Beit Netofa Valley in the Lower Galilee as well as in the Western Galilee. To better understand the traditional farming system, we compare it to modern agriculture as exercised today in industrial countries (Table 6.1). Modern agriculture emphasizes high yield (economic efficacy) to ensure profit, whereas traditional farming places higher emphasis on a stable (albeit relatively low) annual yield. This is an essential difference that merits further elaboration. In traditional farming, affluent years characterized by high yields affect many farming communities across large regions. In the absence of infrastructures that would facilitate effective, long-term storage or the transportation of produce, most of the excess yield is likely to be unwanted in the local community (while some is kept by the farmers for future needs) and would eventually benefit mice and other pests, the populations of which would thereby grow. Similarly, lean years and drought are regional phenomena impacting many neighbouring farming communities across large areas. With no neighbour upon which to rely, the outcome of dry years would often be food shortage and even hunger, which increase mortality among the elderly, the ill and the young. Thus, annual yield stability was crucial in such systems in semi-arid areas. As noted earlier, in Mediterranean climate, no two rainy seasons are alike, and throughout the years, affluent, average or lean seasons change at varying rates. In traditional farming systems, then, a moderate yield surplus collected during infrequent affluent years offsets impaired production, thereby allowing subsistence during drought.

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Because of its emphasis on maximal yield, in modern agriculture, large, uniform fields are advantageous as is the specialization in only few crop types. In contrast, in traditional farming, the large variety of crops, and even different types within each crop designated at different uses (such as wheat for flour versus wheat for bulgur groats), are favoured as they ensure a stable local supply. Crops with different seasonality also allow the farmer to decrease the workload during the (peak) seasons and effectively manage work in the fields. This explains the incentive to include both summer and winter crops. For example, in the Near East, wheat and barley are currently sown in the winter and reaped in May or June, while sorghum, which is a summer cereal, is sown in March, when the wheat and barley require little fieldwork, and is reaped after the winter cereals harvest. Most summer crops were not domesticated in the Near East but rather in other world regions, such as the African sorghum and cowpea or the Mexican common bean and maize. This will be elucidated in Chapter 9, where we discuss the choice of species that would have been candidates for domestication. Modern agriculture employs as much machinery as is economically possible in contrast to traditional farming, which was based on manual labour and later also on working (tract) animals. Modern agriculture thus necessitates intensive inputs and requires certain infrastructures. Being the product of recent globalization, modern agriculture generates scenarios that are on the verge of the absurd or which can develop only under heavy subsidy. For example, rapeseed is raised in Australia for the purpose of producing canola oil. However, its seeds are then transported to China on bulk carriers, and a minor portion of the packaged produce is returned to the Australian retail market. This scenario has evolved because Australians do not require most of their rapeseed yield (or wheat, barley, lentil and chickpea yield) and thus often have not built local facilities in which to process rapeseed produce. Similarly, in another field, Australia exports iron ore but does not enjoy the added value of steel manufacturing, which is carried out in countries where manpower is cheaper. In contrast, traditional farming does not necessitate heavy infrastructures and requires minimal input. Modern agriculture often results in the adaptation of the environment to the needs of grown crops (such as trellising, greenhouses or temperature and daylight control) whereas traditional farmers would match the crop to the environment. In spite of this, traditional farmers invest greatly in some environment-adapting infrastructure construction activities, such as terraces, dams, granaries and water aqueducts. While this infrastructure requires immense input and labour and necessitates subsequent maintenance activities, some of which could last for thousands of years, their goal was not to serve one specific crop but rather to allow for the existence of agriculture in the first place. Readers should note that we do not offer a ‘romantic’ view of traditional farming systems and that our comparison between traditional low-input systems and modern farming involves two time windows. Since our ability to reconstruct Neolithic husbandry is at best limited, we describe the Near Eastern systems of the late nineteenth to

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early twentieth centuries, before modern medicine and the advent of antibiotics led to sharp demographic growth worldwide; that is our first time window. Our second time window is that of post-World War II modern farming, for example in the industrial nations of Western Europe, North America and Australia. We also emphasize that the mere fact that traditional farming involves minimum input and is aimed at subsistence rather than the export of primary or processed produce to remote markets does not necessarily mean that it is sustainable. For example, due to massive population growth during the past 150 years, subsistence farmers in Ethiopia were forced to start ploughing steep mountainsides (because all prime land was long occupied), occasionally without adequate terracing. This, of course, resulted in soil erosion, and a quick decline in soil fertility and yields, which in turn may send the poor farmer to clear more marginal land for ploughing to feed his family. Similar processes occur not only in semi-arid regions like the Near East or East Africa, but also in more humid regions that sustain rainforests. For example, in certain tropical countries, subsistence farmers have practised swidden agriculture (often termed ‘slash and burn’) for centuries. In this system, the trees are felled, the slashed material dries out and, following burning, the ash contributes nutrients to the soil. The cleared land is subject to cultivation and after several years (and decline of fertility) is totally abandoned for eight to ten years. This allows the natural forest ecosystem to rehabilitate, thereby restoring soil fertility. However, population growth (and consequently the establishment of new villages and farming communities) across tropical Africa and south-east Asia increases the ecological pressure by dramatically shortening the gaps between cultivation cycles (e.g., two to three years), with the inevitable result of ecological damage to local forests and fauna, as well as a decline in soil fertility and yields. The above notwithstanding, we think that the features of pristine traditional farming (to the extent that we can grasp it these days) and its traditional cultivars (often termed heirloom crop varieties) provide the best reference for comparisons with the wild progenitors of our crop plants, rather than modern farming systems and modern high-yielding cultivars.

K E Y PO IN TS A ND BE Y O ND

• To understand the domestication syndrome, wild plants must be compared with crop plants that characterized traditional farming systems. We use diverse ethnographic sources for this purpose. The diversity of crops (number of genera, species and cultivars within each species) for different purposes is an important characteristic of these systems, determining seasonal work distribution throughout the year. • In traditional farming, attempting to stabilize yields year after year is more important than generating record high yields. This represents a deep perceptual-existential difference between traditional farming and modern agriculture, in which crop

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maximization is valued – a testimony to the distance we have crossed since the inception of subsistence farming. • Modern agriculture is guided by short-term profit considerations, requiring intensive inputs and costly infrastructures. Retrospectively, following short periods of time (decades at most), both short- and long-term damage are apparent which have been inflicted by modern agriculture upon the environment as well as on our health.

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7 T H E D I F F E R E N C E S BE T W E E N P L A N T D O ME S T I C AT I O N AN D C RO P E V O LU TI O N UN D E R T R A D I T I O N A L A N D M O D E R N F A R M I NG S Y S T E M S

Similar to wild populations, traditional landraces (varieties, cultivars) of many annual crops form dynamic populations, the genetic make-up of which changes over time. This point was clarified in our earlier discussion (Chapter 5, and see Chapters 8, 9) on the changing incidence of early and late bloomers responding to seasonal rainfall among both wild populations and cultivated plants. Usually, traditional farmers would preserve sowing materials from the last yield of traditional cultivars (see Glossary, Botany, Ecology and Agronomy, Traditional cultivar (landrace)), which comprise a sample of sorts of the entire yield. The population make that farmers thus own reflects the historical evolution that occurred in their fields. This includes natural selection originating in pressures caused by agents such as pest and disease epidemics, extreme climate, frequently occurring random mutations and the farmers’ own selection processes when choosing the plants or plots in their fields from which seeds will be cached for the next season. Since traditional landraces are dynamic, it would be unreasonable to assume that all of the differences found between the progenitors of the founder crops and the domesticated types are solely the result of the pristine domestication episode. In other words, some of the differences between these two plant groups are the results of selection processes that took place during the thousands of years that have elapsed since domestication and under (a cultivation regime) domestication. In light of the above, it may now be understood that our comparison of wild and domesticated plants based on the classical framework of the domestication syndrome (see Chapter 5) included, by necessity, not only crucial domestication traits such as the absence of bitterness (lack of cyanide-generating materials) in the domesticated almond

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or free germination in legumes but also traits that appeared and accumulated in domesticated landraces during the millennia following the domestication episode. Such traits are the size, colour and taste of fruits worldwide – for example, the high diversity in fruit characteristics among prune, peach or apple cultivars, relative to the uniformity among wild forms. Other examples of such features differentiating between wild and domesticated species include protein content of grain or the time (early or late) of flowering in ornamental, industrial (e.g., tobacco, cotton), food and feed crops alike. The question, then, is, how can we discern between those traits that were essential for domestication and those that accumulated in subsequent phases under domestication? In other words, which trait of the wild species under investigation is crucial to change and without which it is impossible or unprofitable (involving loss) to grow this crop? If such a trait can be identified, it is likely that most (or all) other traits became modified under (post-) domestication and were perhaps not even involved in the pristine domestication episode. For example, seed size of many cultivars of pea, chickpea and lentil are larger than those of their wild progenitors. It is important, then, to ask if large seed size is a necessity in domesticated pea, chickpea and lentil. The answer to this question would determine whether seed size was a domestication imperative. This question has additional value as some researchers base their arguments in favour of the protracted (millennia-long) model of domestication on the size of seeds found in the archaeobotanical record retrieved at Neolithic sites. Any attempt to identify the traits imperative for domestication among those traits collectively known as the domestication syndrome involves the study of the biology of wild species and their domesticated derivatives for each crop separately and the examination of genetic profiles among the diagnostic plant traits. Below we discuss chickpea, pea and wheat as test cases. To answer the question of whether seed size was a crucial element in domestication, we look at several commonalities and differences between domesticated and wild chickpea. All cultivars of domesticated chickpea are free-germinating and do not require a growth period under cold temperatures to time their flowering (a trait known as vernalization) (see Box 17 Response to vernalization and control of flowering time, p. 130). Chickpea cultivars vary greatly in several traits: seed weight at 120–650 mg/ seed, flower colour (white or pink) and growth habit – from low, bushy plants to tall plants with few branches. Chickpea cultivars also have non-shattering pods that do not shed with maturity. In other words, the seed distribution mechanism that is characteristic of wild species is inactive in domesticated chickpea. In a very few cultivars mature seeds fall from pods, but only under very hot and dry conditions and only to a minor extent. Compared to domesticated cultivars, all types of studied wild chickpea (Cicer reticulatum) exhibit hardseededness, hindered germination and response to vernalization, that is, they flower early following exposure to near-zero temperatures while in seedling stage. The range of seed weight, at 90–180 mg/seed, slightly overlaps that of smallseeded domesticated cultivars. Pods in wild chickpea shatter only partially, but ripe pods often fall off from the mother plant. Since pods in wild chickpea typically contain only a

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Box 17 Response to vernalization and control of flowering time Plant physiologists offer the following definition for vernalization: a plant’s growth period under temperatures that are considerably lower than those required for its optimal growth. The result is a considerably earlier time of flowering. This adaptive trait

(mechanism) is typical of both annuals and biannuals in temperate regions or regions in which below-zero temperatures may occur during the winter or even early spring. If the plant does not flower unless it undergoes a cold growth period, we say it requires vernalization. Vernalization ensures that an autumngerminating plant will not develop flowering

Figure 7.1 Vernalization in wild chickpea. Top: plants that underwent a vernalization treatment; bottom: control, unvernalized plants with retarded development and delayed flowering. A colour version of this image can be found at www.cambridge .org/abbo-gopher.

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buds until the end of the cold period in which temperatures drop below zero, thereby safeguarding flower and fruit development from potential frost damage. Vernalization corresponds with the mechanism responsible for the response to day length, which controls flowering in many annual wild plants that are typical of the Mediterranean region of the Fertile Crescent. The combination of these two mechanisms in local plants offers an adaptive edge (survivability) so that if the required doses of cold have already accumulated (during winter) but the day is still short (twelve hours in March), flowering would

not occur. In contrast, subsequent to spring germination, the plant is likely to experience long days but without the required doses of cold. Such conditions will delay (or prevent) flowering except in vernalization-insensitive plants. Such plants might flower and set pods in early summer and disperse their seeds before the rainy season. However, in the following autumn germination, and without the vernalization requirement, these plants are likely to flower too early. Such untimely flowering will be exposed to below-zero temperature damage, resulting in low fecundity (Figure 7.1).

single seed, for the purpose of seed distribution it is immaterial whether the dispersal unit is the naked seed or the seed-bearing pod. This minimizes the diagnostic value of pod shattering in chickpea, which drove Ladizinsky to suggest in 1979 that wild chickpea was pre-adapted to domestication since it required little deliberate selection efforts from early farmers to achieve cultivars with non-shattering pods. In contrast, as we have shown, free germination was crucial for crop profitability as was the elimination of response to vernalization (being sown in the spring, the plant would not receive the dose of cold required to advance flowering). Devoid of these two traits, chickpea growth could not be made profitable. This infers that all other traits that characterize certain domesticated cultivars of chickpea, such as large seeds, early flowering, white flowers and tall growth, have all developed under domestication, that is, later than the actual domestication episode. In other words, these traits emerged in cultivated fields only after farmers had possessed a free-germinating stock of chickpea. Said more directly and unequivocally, seed size cannot be used in chickpea for the purpose of describing or documenting domestication. As with chickpea, when examining the importance of seed size in pea, we look at the commonalities and differences between the domesticated stocks and wild species. Domesticated pea cultivars exhibit a great variety of seed sizes and colours (brown, green, grey, black and different mottling patterns of dark colour over a lighter background), white or pink flowers, early and late bloomers and types that require vernalization for timely flowering and those that do not. However, all domesticated pea cultivars, without exception, are characterized by free germination and non-shattering pods. Even some pea landraces that originated in the Ethiopian and Eritrean highlands, which germinate slowly (spanning a few days) compared to modern types, cannot be considered to be dormant in the sense of dormancy that characterizes seeds of wild peas.

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When paralleled with domesticated pea cultivars, all studied wild pea species exhibit strong seed dormancy, shattering pods, a variety of flowering times and different degrees of response to day length and temperatures as expressed in their flowering time. In the past, it was claimed that the most important trait in legume domestication was the elimination of the seed dispersal mechanism, that is, a non-shattering pod (analogous to the non-shattering cereal spikes). However, having learned that profitability in pea crops is dependent on free germination, and especially after learning that profitable yields can be obtained from wild pea stocks characterized by shattering pods as long as germination occurs at a rate of 30% or more, we may say with certainty that the crucial trait for pea domestication was free germination. Thus, no other trait – shattering of pods or size of seeds – can be used to describe or document pea domestication. This insinuates that variation in other traits developed after domestication, under a cultivation regime. This process is termed crop evolution or plant evolution under domestication. Our third test case is durum, or emmer, wheat. Studying domesticated cultivars shows a wide array of traits such as spike form, ease of threshing, seed colour and size, grain quality (including bread and other product features as determined by the flour quality), plant height, time of flowering, flowering response to vernalization and day length, awn length and response to pests and diseases. The wild progenitor (Triticum dicoccoides) also exhibits a broad range of traits such as time of flowering, spike pigmentation, seed weight, seed protein content, plant height and response to pests and diseases. All known types of wild wheat, however, are hard to thresh, that is, their seeds are covered by lignified chaff and exertion is required to separate it from the grain and to prepare the grain for pounding or grinding. In comparing the differences between wild types and domesticated cultivars, only spike shattering can be considered explicit. All wild wheats, without exception, are characterized by shattering spikes, while all domesticated cultivars are characterized by non-shattering spikes. A minor caveat to this statement concerns some traditional, relatively hard-to-thresh landraces that exhibit a limited degree of shattering that scarcely impacts grain yield and is therefore agronomically inconsequential. In contrast to this qualitative difference between shattering wild types and nonshattering domesticated cultivars, the overlap in seed weight between wild and domesticated wheats combined with the fact that some primitive landraces are hard to thresh and have non-shattering spikes seems to indicate that spike shattering was the most important trait for wheat domestication. Spike shattering is thus the sole diagnostic trait that can be used for the purpose of documenting domestication through botanical remains found at archaeological sites. The great variation recorded in all other plant traits, important and available to the professional archaeobotanist as they may be, are irrelevant for tracing wheat domestication. The variety seen in domesticated cultivars (as noted above for both the chickpea and the pea) is the result of plant evolution (diversification) under domestication, created and amassed in times subsequent to the domestication episode. For the three test cases reviewed here, understanding in detail the biology of each domesticated species and its wild progenitor allows us to both refine and expand the

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concept of the domestication syndrome. This in turn enables us to transform the concept into a valuable instrument that can be used in documenting the significant episode of domestication and to achieve an improved distinction between traits that changed or were accumulated under (after) domestication and those underpinning the domestication episode itself. Furthermore, it seems plausible to generalize and say quite confidently that the crucial domestication traits are typically qualitative, resulting in a uniform expression (monomorphic; see Glossary, Genetics, Polymorphism) in both the wild gene pool and among domesticated cultivars, albeit with wild phenotype in the former and with the alternative, domesticated alleles (and morphology) in the latter gene pool. In contrast, genetically variable traits (see Glossary, Genetics, Polymorphism) in both wild types and domesticated cultivars, and especially traits that show a phenotypic overlap between domesticated cultivars and wild populations (such as seed size), are likely traits that changed only after domestication and played no crucial role in the domestication episode. In closing this chapter, we would like to note that wild and domesticated cereals were, and still are, at the heart of scientific and agricultural research most likely because of their enormous significance to the human food (caloric) supply and economies on all inhabited continents. The main conclusion of all studies that have investigated domesticated crops in comparison to their wild progenitors is that biological changes pertinent to domestication and further evolution of these crops are similar in nature. For example, we reiterate that most domesticated grain crops do not self-distribute seeds, their seeds lack dormancy and in many domesticated crops seeds and fruit are significantly larger than in their wild progenitors, to name but a few traits that typify domesticated crop plants at large. The intensive study of cereals caused the biological characteristics of cereal domestication to dominate the collective awareness of researchers as an informal standard of domestication. This, combined with observations on what appears to be convergent or parallel evolution of many domesticated plants (in which the physiological mechanisms of seed dispersal and seed dormancy were inactivated upon domestication), is what bred the concept of domestication syndrome. Undoubtedly, shattering spikes (cereals) or pods (legumes) serve the same biological end, namely, seed dispersal, and thus non-shattering spikes and pods in domesticated cultivars are important traits producing high value to farmers because they minimize seed losses. From this perspective, cereal and legume domestication may seem similar. Over the years, however, legumes were studied only sporadically, resulting in little genetic and agronomic insight vis-à-vis domestication. Following Ladizinsky’s pioneering work (see discussion on seed dormancy in Chapter 5) and our own work in recent years, we have been able to advance research on wild and domesticated legumes in the Near East and show that crucial domestication traits of cereals and legumes are different. These works also show that it may be possible to arrive at additional insights regarding the differences between cereals and legumes through further studies rather than be satisfied with the traditional emphasis on

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similarities between domestication and crop evolutionary processes of the two plant groups. This comparison lies at the heart of the next chapter. K E Y PO IN TS A ND BE Y O ND

• Cultivars of different crops form dynamic populations, the genetic make-up of which changes over time. It is improbable that all the differences found between domesticated plants and their wild progenitors originated in the early domestication episode. Examining the biology of the different crops may direct us towards the traits crucial (imperative) for domestication, thereby helping us to determine which traits likely changed under (post)domestication. • Generally, crucial domestication traits have simple genetic control (a single gene or a small number of genes) whereas many of the traits that show a phenotypic continuum between wild progenitors and domesticated cultivars (such as seed size) are governed by multiple (poly)genes. • The distinction between those traits that were crucial for domestication and those that changed under domestication as part of crop evolution (and continue to change even today) is a fundamental element in understanding plant domestication, and it is imperative for any discussion concerning the pace of domestication.

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T H E D I F F E R E N C E S BE T W E E N C E R E A L A N D LE G U M E C R O P S I N TH E N E AR E AS T

As noted earlier, three cereals (barley, durum or emmer wheat and einkorn wheat), four species of legumes (pea, lentil, chickpea and bitter vetch) and flax, which is neither a cereal nor a legume but rather belongs to the Linaceae family, form the founder crops of the Near East. In this chapter, we compare the biological characteristics of the two main crop groups, the cereals and the legumes. Through this comparison, we shall attempt to trace the manner in which Neolithic farmers assembled their agricultural crop package. As in previous chapters, where we compared wild and domesticated plants and traditional and modern farming systems, in this chapter, too, we compare cereals and legumes based on a detailed study of the different wild species and their domesticated derivatives. Here, as in Chapter 7, we use traditional cultivars (landraces) (see Glossary, Botany, Ecology and Agronomy, Traditional cultivar (landrace)) as a reference rather than modern cultivars that were bred during the last century. The first trait that we discuss is plant height (Table 8.1). The three wild cereals – wild barley, wild emmer wheat and wild einkorn – can all reach or surpass the height of 1.5 m under optimal growing conditions. Similarly, traditional wheat cultivars and barley are tall, erect plants, and some may reach up to 2 m under particularly affluent conditions. Among the wild legumes, however, the pea alone is an erect plant that can exceed 1 m at maturity and even exceed 2 m when climbing an adjacent tree or shrub. Domesticated pea landraces also grow tall. In contrast, wild lentil, bitter vetch and chickpea exhibit compact, prostrate growth, rarely exceeding a height of 20 cm. Domesticated chickpea, lentil and bitter vetch are also short-statured and exhibit inferior yield potential compared to domesticated wheat and barley. The positive correlation

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Table 8.1 Summary of biological differences between Near Eastern legumes and cereals (wild and domesticated) Trait

Legumes

Cereals

Plant height

Short-statured, prostrate, or bushy plants (up to 30 cm)

Erect, exceeding 1.5 m

Dispersal unit

Camouflage-coloured seeds

Spikelets with long awns

Growth pattern

Indeterminate

Determinate growth: shoot apex ends with spike

Seed dormancy*

Very high (strong), approximately 90%

Low, approximately 50%

Pollination

Mostly self-pollinating

Mostly self-pollinating, 0.5–10% cross-pollination

Population

Sparse and patchy

Large and dense

Source of nitrogen

Symbiosis with soil bacteria

Nitrogen minerals in the soil

Competitiveness

Poor (uncompetitive)

Competitive, aggressive growth

* The seed dormancy figures refer to the wild types.

between plant height and biomass production, found in many studies of various plants (both wild and domesticated), is often expressed as a positive correlation between height and seed yield in grain crops. This positive correlation is easily explained as it reflects the correlation between the number of leaves (and other green plant organs) intercepting sun radiation and their total surface area and between the plant’s photosynthetic capacity. In other words, with greater photosynthetic capacity comes higher yield potential. This is why the grain yield (and straw mass) of domesticated wheat and barley, which typically exceed 1.5 m in their height, is considerably greater than the yield of lentil or bitter vetch, which typically do not grow beyond 0.5 m even as domesticated plants. Another difference between cereals and legumes is in the dispersal units of their wild progenitors. In wild legumes, the dispersal unit is the single seed that shoots off the shattered pod as it dries up. As legume seeds are typically camouflaged, matching the colour of the soil in their habitat, it is difficult to identify them against the cracked soil, among stubble or when they are hidden in rocky crevices. The cereals, in contrast, are distributed through their shattering spikes. These comprise spikelets that are arranged in alternating ‘levels’ along the spike’s axis. Upon maturity, a specialized tissue (abscission zone) develops in the joints that separate one spikelet from another so that when the plant dries up at the end of spring, the spike disarticulates into individual spikelets that fall to the ground. In barley, each spikelet produces a single flower, and it therefore carries a single seed covered in chaff. In wheat, however, each spikelet may produce two or more flowers (florets), resulting in a number of grains matching the number of fertile flowers (typically two). The bottom bract of the cereal floret ends with an awn (the ‘hair’ typical of cereal spikes), so that the dispersal unit of the barley carries a single awn while that of the wheat carries two awns. After spikelets have fallen to the ground,

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alternating cycles of (sun) drying and (dew) wetting of the serrated awns trigger their movement, which facilitates – due to the ratchet-like serration of the awns – the movement and thereby penetration of spikelets into crevices in the ground and under rocks. This is where the ripe pair of wheat seeds or the single barley seed will await the next autumn rains, covered in their chaff. Unlike the legume seeds, the size of which is measured in millimetres, cereal spikelets are approximately 1–2 cm long and may reach up to 10–15 cm when the awn is considered. For hunter-gatherers this translated into great effort, with little success, invested in the collection of legume seeds as opposed to the potential for the fairly easy collection of considerable amounts of wheat and barley spikelets even weeks after maturity was achieved and spikes shattered. This was proven more than a decade ago in a controlled experiment conducted in the Eastern Galilee by Professor Mordechai Kislev and his students at Bar Ilan University. The third trait concerns growth patterns, including plant structure, location and the manner by which their reproductive organs – cereal spikes and legume flowers – develop on the adult plants. The three cereals among the founder crops exhibit determinate growth habits, that is, following germination, the plants develop a certain number of leaves (at a series of compact internodes) at the base of the major growth axis. At each leaf axil, a lateral vegetative bud forms that may later elongate and develop another spike-bearing stalk (tiller). As the days grow longer at the end of winter, the shoot apices (apical meristems) in cereals transition from a vegetative state (growing leaves and lateral tillers) to a reproductive state (developing the spike; see Glossary, Genetics, Cell differentiation). Elongation of the upper nodes (and hence, the stem) occurs only after the embryonic spike has developed, pushing out the developing spike through leaf sheaths until emergence. A few days after the spike has emerged through the sheath of the uppermost leaf, the uppermost node ceases to elongate. At the same time, the anthers open and pollen grains land on the stigma within the florets. Wheat and barley spikes alike each carry approximately twenty spikelets. Following the fertilization of female egg cells in the ovaries by male nuclei from pollen grains, cells begin to rapidly divide and grains develop for the next thirty to forty days until physiological maturity. In this growth profile (determinate), most spikes in a given field emerge within just a few days, and therefore flowering and grain maturation in cereals occur rather simultaneously (synchronously) among the entire field population. In contrast, the legumes among the Near Eastern founder crops exhibit indeterminate growth. The lower stem segments (nodes) are compact, and elongate only slightly, dictating a prostrate growth habit. Lateral branches begin to develop at basal internodes and later also at internodes of higher stem segments. Additional (secondary, tertiary, etc.) branches develop from stem internodes on the first lateral branches, gradually creating a bushy, branched plant. This is in sharp contrast to the cereals in which lateral growth (tillering) occurs only at the basal stem internodes. In this respect, the pea differs slightly from its three counterparts (lentil, chickpea and bitter vetch) as its growth is somewhat more vigorous, its leaves are relatively larger, it has tendrils that assist in

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climbing over adjacent plants and it is often not highly branched. As with cereals, in legumes, too, reproductive organs (flowers) begin to emerge after the end of winter when days grow longer. Unlike cereals, however, in which the apex develops into the inflorescence (spike), flowering in legumes occurs in upper internodes (since low internodes carry lateral branches). The shoot apices of legumes thus continue to grow while budding leaves and flowers at their internodes. Most chickpea cultivars develop a single flower per node whereas pea, lentil and bitter vetch each develop one to three flowers per node. Since flowers develop in leaf axils, increased seed yield requires continued stem growth and leaf development. Unlike the cereal spikes, where twenty to forty grains develop rather simultaneously after flowering, in legumes seed pods may develop two to three days apart from one flower to the next, so that flower initiation, flowering, fruit (pod) set and maturity are each asynchronous, even within a single plant. Because of their indeterminate growth, at the end of spring, a single wild legume plant may bear pods at different ripening stages, from shattered, empty pods at lower stem internodes to green pods alongside continued flowering at higher positions. Compared to the number of grains in cereal spikes, legumes yield a limited number of seeds in each pod. Lentil and chickpea pods typically carry one to two seeds, bitter vetch might carry two to four seeds per pod, and the pea, in line with its other unique traits, develops four to eight seeds per pod. These differences in growth profiles between cereals and legumes are present in both wild and domesticated types and therefore bear enormous significance for farmers. The rather synchronous nature of ripening in cereal spikes exposes the crop to the hazards of environmental stresses; for example, an extremely dry event (Khamsin, in both Hebrew and Arabic) during the crucial stage of grain development (April or May) may cause shrivelling of a large majority of grains. In contrast, a similar event would only partially damage legumes since at any given point in time only a few of the pods or flowers are at their sensitive stage: mature pods will not be affected by such an event while damage to top flowers, even if they wither, would cause only partial damage to the final yield. Furthermore, legumes would respond better to late rains, as precipitation would encourage additional growth and podding and hence greater yield. Cereals, in which the number of grains per spike is determined at an early stage, long before the end of winter, would benefit only to a limited degree from late rains. The next trait in Table 8.1 is seed dormancy. Seed coats in wild Near Eastern legumes are hard and impermeable to water (a trait shared by some legumes such as the African cowpea and Asian soybean that were domesticated in other world regions), limiting germination to about 10% at the start of the rainy season. The remainder of the seeds will germinate in following seasons at similar rates. In domesticated cultivars, this mechanism was genetically altered (one or more mutated genes) so that seed coats allow seeds to swell (imbibe water) as they come in contact with soil moisture (allowing for about 90% germination rate). For the farmer, this is a highly important feature as it facilitates the uniform development of crop populations at similar

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developmental stages and at desired densities. Without this trait of free germination, no significant cultivation would be feasible for the legume species that were domesticated in the Near East. The wild founder cereals also exhibit a certain degree of seed dormancy. Whereas germination rates of wild cereals are approximately 50% (considerably higher than germination rates in wild legumes), domesticated cereals lack dormancy and mature seeds will germinate at a rate higher than 90% when they come in contact with moisture. The high germination rate of wild cereals (relative to wild legumes) serves as the first clue that the importance of this trait could not have been equal for cereals and legumes, and, therefore, the domestication episodes of the two groups, as well as the evolutionary processes that occurred under (i.e., after) domestication, could not have been identical, despite the widespread belief that they were. Most of the plants that were domesticated in the first domestication episode that occurred over 10,000 years ago in the Near East (excluding broad bean) are selfpollinating plants, that is, the pollen is released within the flower, upon the stigma (the female receptive organ), thus starting the fertilization process required for seed development. Despite the high rate of self-pollination, approximating 100% in cereals and most legumes, the two groups exhibit differences in this respect, too. In legumes, anthers release the pollen while the petals are still closed, so that the chances of pollen grains becoming airborne (or being transmitted by insects or birds) are nearly nonexistent as are the chances of alien pollen grains, carried by the wind or other agents, landing on the stigma and fertilizing the flower. In cereals, however, several mechanisms allow for pollination by neighbouring plants. One such mechanism allows the anther filaments to extend beyond the flower after releasing most of the pollen within the flower (an event that does not occur in legumes). Even the smallest amount of pollen released from the anthers and carried in the wind (thereby causing springtime hay fever) to the shortest distance (ca. 30–50 m) would enable cross-fertilization. This is made possible because another mechanism in cereals dictates that flowers will remain open for a short while after the pollen grains have been released within them. Wild cereal populations vary in the extent of cross-pollination they facilitate, from 0.5% or less to approximately 10%. Even a rate lower than 1% would still ensure gene flow between individual types as well as between populations and has the power to create new genetic combinations within a population over evolutionary time frames. The rate of cross-fertilization in the population is determined based on the rate of heterozygosity (see Glossary, Genetics, Heterozygous), a term that describes the condition in which the genetic profile of a certain gene facilitates the identification of a contribution that did not originate in the female (the seed parent) plant but rather from an adjacent (or remote) male (pollen donor) plant. In contrast, and given the low heterozygosity indices in Near Eastern wild legumes, we assume that individual members of wild legume populations are highly similar due to the high rate of selfpollination and the absence of mechanisms that allow for the technical potential of crossfertilization. Most genetic variability in such populations would therefore be found

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across sites in the geographic expanse, while individual plants that are in physical proximity are expected to be quite similar to each other and often identical. Wild barley and wheat populations in most areas of their natural distribution are fairly large and dense. Even today, after thousands of years of continuous damage incurred upon wild ecosystems and intensive modern impairment, Israel boasts large expanses such as the Gilboa, Golan Heights or Galilee (Eastern and Lower) where large populations of wild barley still grow. Based on what is known regarding the distribution of this species, it is highly likely that in prehistoric times near-continuous populations of wild barley would have occupied the area spanning between the Mediterranean through the Fertile Crescent to the border of Iran and Afghanistan, and perhaps even further eastwards towards central Asia. The habitat of wild emmer wheat is far more constrained, limited to the boundaries of the Fertile Crescent, but within its habitats, it too establishes dense populations in which thousands of individuals grow within a distance of just a few minutes’ walk (in the Gilboa and Golan Heights, in Israel). Similarly, wild einkorn wheat also establishes large, dense populations within its area of distribution in the northern Levant, eastern Turkey and the Balkans. Compared to the massive populations generated by wild cereals, small legume populations, each comprising but a few individual plants, are scattered in their environment. The patchy legume populations often comprise only two or three plants spanning just a few metres (e.g., of wild lentil or wild chickpea) whereas the next spot would only be found at a distance of some tens or hundreds of metres away with no other individual of the species found in between spots. Wild pea populations also comprise just a few individual plants, and sometimes only a single one. Next in our comparison is the source of nitrogen, a substance required by plants for their normal development and seed production. Chlorophyll – the molecule that grants plants their green colour and facilitates the conversion of solar energy into chemical energy required for growth and development of the plant, its fruit and seeds – is a nitrogen-rich compound. Nitrogen is also required for the production of proteins, without which life cannot be sustained. Nitrogen comprises 70% of our atmosphere, but only small concentrations of nitrogen minerals are found in the ground. Nitrogen penetrates the ground only following bacterial activity and the decomposition of organic plant matter or animal carcasses. Absorbed by plant roots, this soil mineral nitrogen is the main source of nitrogen for plants. Legumes, however, developed in their evolution an additional source of nitrogen, which is supplied to them through symbiosis with nitrogen-binding soil bacteria. These bacteria recognize legume roots, and through the secretion of different organic compounds trigger the development of designated organs within the root (known as nitrogen nodules), where they fix atmospheric nitrogen. In return, the host plants supply the bacteria (which settle in these root nodules) with sugars that are produced in their leaves as part of the photosynthesis process. This symbiotic system is active until flowering occurs, and seeds begin to develop in the pods. When the latter takes place, the plant slows the flow of energy-packed

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carbohydrates to its roots, and the symbiotic system collapses. At this point, however, the plant would have already amassed about two-thirds of the nitrogen it requires to generate its seed protein. No equivalent system has developed in cereals, leaving cereals dependent on available soil mineral nitrogen for their growth. A direct outcome of this difference in nitrogen availability is the difference in protein concentrations found in the seeds of legumes and cereals. In wild cereals, seeds comprise mostly carbohydrates, amounting to approximately 80% of their weight, whereas protein amounts to 12–19% of seed weight, and the remainder comprises fat and different minerals. In contrast, legume seeds comprise 65% carbohydrates, 19–31% protein and 4% fat. Additionally, the amino acids forming the protein in each crop group differ in composition, so that a diet that combines both cereals and legumes is considered balanced and healthy. Because legumes need not rely on soil fertility and have an independent source of nitrogen owing to the nodules that host nitrogenbinding bacteria, they are capable of sustaining high protein content in seeds even in unfertilized fields. In early cereal fields, as well as under modern cultivation, for various reasons (such as ploughing and straw removal that reduce natural organic matter in the soil or the limited variety of plants in the cultivated field), it is difficult to obtain a level of soil fertility that will ensure high protein content in crop yields. We therefore surmise that during the Neolithic period and at early stages of agriculture, wheat and barley landraces with relatively low seed protein content had a yield advantage. Indeed, many domesticated wheat and barley cultivars produce low seed protein content, averaging at 12%, much below the values documented in wild wheat and barley. We further infer that this decrease in seed protein content was acceptable to farmers as they relied mainly on grain cereals for their carbohydrate content whereas legume seeds, with their higher protein content, probably served as the main source of vegetal protein. The two plant groups, the cereals and legumes found among the Near Eastern founder crops, differ significantly in each and every one of the traits (parameters) detailed in Table 8.1. We therefore conclude that the domestication of legumes had to have been different from the domestication of cereals and that the successful domestication of each necessitated proper attention to the different biological traits that characterize the two plant groups. Similarly, their evolution after domestication would have been different due to the agronomic implications of the biological differences between the two groups. The combined effect of plant height and growth habits causes cereals to grow more aggressively and more competitively than legumes (also attested to by the large and continuous wild cereal populations as opposed to the patchy growth of wild chickpea, lentil and pea that thrive only in stony habitats in which the soil is shallow, and other locations in which cereals are less successful). This difference in the composition of wild populations emphasizes yet another aspect in which the two plant groups differ. We might say that cultivated cereal fields during the Neolithic period emulated their wild conditions, as wild cereals naturally grow in large, dense populations. In contrast, the notion of a legume field is a human invention, since this is not a typical

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a

b

Figure 8.1 Faba bean (Vicia faba) plants (a) with flowers and (b) with pods – courtesy of Dr Jeff Paul.

natural phenomenon. We would thus argue that the domestication of both cereals and legumes was part of a revolutionary human shift with respect to the human relation to natural vegetation. However, whereas the sowing of domesticated cereals in dense fields replicated a landscape that was natural and familiar to the domesticators, domestication of legumes resulted in an innovation that must have involved a significant conceptual shift (leap) in their minds, since a dense field of legumes was not a familiar scene prior to domestication.

C O M M E N T S O N B R O A D ( F A B A ) BE A N , A LE G U M E O F N O KN O W N WILD ANCESTRY

Botanists have classified broad bean (Figure 8.1) under the botanical genus of Vicia (vetch) although we have no knowledge of the wild progenitor of the species and despite the fact that it is genetically unrelated to any known Vicia species. The reason for this may have been its morphological resemblance to certain wild Vicia species, including Vicia narbonensis, Vicia johannis and Vicia galilea. These species have thick stems, their leaves are fairly large with wide leaflets and their large pods carry seeds that are significantly bigger than seeds of other Vicia species. Relying on their thick stems, these species grow tall, similar to the upright, tall growth of the broad bean. However, since the broad bean cannot be crossbred with any of the known Vicia species, these botanical criteria are insufficient to classify it under the genus of Vicia, let alone suggest that either one was the wild progenitor of the broad bean.1

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Assumptions about the possible primary growth area (outside the Near East) of the elusive wild broad bean have been raised (such as in the vicinity of Afghanistan) because the local types of broad beans bear small seeds and appear less advanced compared to other cultivars of broad bean documented from elsewhere in the world. While the wild progenitor of the domesticated broad bean is yet to be identified, the earliest remains of broad bean are charred seeds found in archaeological sites concentrated in northern Israel and north-western Syria. In Israel, they originated in the Lower Galilee in the 10,000 (or slightly later) year old PPNB sites of Yiftahel and Nahal Zippori, as well as the somewhat earlier site of _ _ Ahihud (ca. 10,200 years ago). Unfortunately, due to the lack of foresight of _ some of the Israeli planning authorities, these important sites are currently buried under new road interchanges. In Syria, faba bean remains were found in the area of Idlib, not very far from the Turkish border, in the early PPNB layers of Tell ‘Ain el-Kerkh, dated some 10,400–10,300 years ago. This finding is highly important as it may attest to the domestication of the faba bean within or at least near the same core area in which all of the Neolithic Near Eastern founder crops were domesticated, and probably also around the same time. We believe that this finding indicates that the progenitor of the faba bean is likely to be found in south-eastern Turkey or northern Syria, should botanists be allowed to systematically survey these areas, which in the past few decades have been suffering from political instability and civil wars. Other than being orphaned, devoid of a known wild progenitor, having thick stems and large pods compared to the other three edible legumes we have thus far discussed (lentil, chickpea and pea), faba bean is exceptional in several other traits, the most important of which is its cross-pollination nature. The plant has the ability to grow, develop pods and ripen seeds by self-pollination, but this inbreeding typically leads to a limited yield. Indeed, faba bean farmers often place beehives in their fields to ensure cross-fertilization by bees, thereby increasing the rate of seed set and hence seed yields. The faba bean is also distinct among the group of legumes discussed here in that it causes anaemia in susceptible humans following the destruction of red blood cells. This phenomenon leads to symptoms reminiscent of neonatal jaundice in individuals carrying specific variants of the G6PD enzyme encoding gene. This gene is carried on the human X chromosome, and therefore most individuals exhibiting these symptoms following the consumption of faba bean are males. Females bearing two healthy (homozygous) gene variants or only one healthy (heterozygous) allele will not develop anaemia following faba bean consumption, but females bearing two impaired alleles would, of course, be as susceptible as males. Impaired versions of this gene are common in Mediterranean, African and south Asian populations.

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F L A X : NE I T H E R C E R E A L , N O R LE G U M E

Flax biology has not been studied as extensively as the biology of cereals and legumes. Pale flax, the wild progenitor, resembles legumes in its sparse, patchy distribution in small populations comprising single members, and it resembles cereals in its determinate growth habit – that is, after leaves and a few stems have appeared, a few flowers will develop close to the apex of the stem from which seed-containing capsules emerge. For this reason, flax plants respond to rain and irrigation regimes in a similar manner to cereals, rather than legumes. Several wild flax species are known in the Mediterranean Basin. One of these – the hairy pink flax – is commonly found in Israel and Jordan, growing densely and forming typical expanses of pink flowers (Figure 8.2), often encountered by hikers in the Lower Galilee and the Judean Mountains. The stems of these wild plants can be used to produce fibre (Figure 8.3). Several methods are used to produce fibre from flax, the most convenient of which involves soaking the stems in water (retting) for approximately one week. Retting in still water (or dew in some world regions) allows for the decomposition of some stem tissues and cell-to-cell connections by natural microorganisms, further facilitating the easy separation of phloem fibre (elongated cells that are resistant to decomposition because of the chemical constitution) from other stem tissues. Indeed, as part of her graduate studies, a student of ours (Ms Inbar Zezak) successfully produced fibre from hairy pink flax, which was effectively used to spin thread by Ms Toni Friedman from Kibbutz Nir Etzion (Figure 8.3). Among wild flax species known in the Mediterranean Basin, the wild progenitor of domesticated flax (see Table 4.1) is not the most common. We regard this fact as further evidence that Near Eastern plant domestication was not a random event but rather a knowledge-based development. We further discuss the choice of Near Eastern domestication candidates in the next chapter.

K E Y PO IN TS A ND BE Y O ND

• Cereal and legume plants that were domesticated in the Near East exhibit several essential differences. These differences have important agronomic implications to both crop groups, and include plant height, growth habits and structure, seed dormancy rates, floral biology (self- or cross-pollination), nitrogen economy and the structure of wild populations. The cumulative outcome of these differences is aggressive growth among cereals and less competitive growth among legumes. • The difference between wild cereal and legume population structure indicates that the two groups must be differentiated and that the domestication of legumes necessitated a larger, perhaps bolder, leap in mind-set, supported by a cultural view that was

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a

b

Figure 8.2 Hairy pink flax in the wild (a) and flowers (b). Colour versions of these images can be found at www.cambridge.org/abbo-gopher.

KEY POINTS AND BEYOND

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a

b

d

c

f e

Figure 8.3 Flax fibre and threads. (a) Fibres produced following retting; (b) hairy pink flax plant picked for the purpose of fibre production; (c) naked stems from which fibre has been removed; (d)–(e) threads spun from flax fibre by Ms Toni Friedman; (f ) fibre of cultivated flax for the purpose of comparison – courtesy of Orit Shamir. Colour versions of these images can be found at www.cambridge.org/abbo-gopher.

contrary to nature, certainly when compared to that which was needed for the domestication of cereals. • Among the crops reviewed here, faba bean and flax are exceptional in several aspects. The wild progenitor of faba bean is as yet unknown, and the plant differs from the other legumes in the nature of its cross-pollination. Flax is neither cereal nor legume, and despite its determinate growth profile (the stem ends with flowers and then the ripened fruits), it does not grow aggressively.

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9 T HE CHO IC E O F P L AN T S P EC IE S F O R DO M ES T I C A T I O N Agronomic and Dietary Considerations

In this chapter, we trace the biological and agronomic logic underpinning the choice of plant species that were eventually domesticated in the Near East. Lacking direct evidence on this issue, we rely on conclusions drawn from the study of current wild and domesticated plants. In Chapter 3, we mentioned the application of the Dump-Heap Hypothesis for explaining plant domestication, stipulating that the chosen domestication candidates were plants that thrived in particularly fertile niches, or those created in proximity to human settlements that experienced frequent ecological disturbances, which encouraged their proliferation. However, when we looked at the ecological preferences of the wild progenitors of the Near Eastern founder crops, we saw that this theory does not hold true for most of them. An alternative suggestion advocated that the chosen founder crops were naturally highly productive and therefore served hunter-gatherers as (staple) food sources prior to domestication. Indeed, a few of the models that describe economic changes leading to the formulation of the agricultural system describe domestication as the end result of a long co-evolutionary process during which two genetically unrelated species (humans and plants) underwent changes over an evolutionary timeline due to biological reciprocity that benefited both parties. Following this approach (under NicheConstruction Theory explanations), humans’ reliance on a specific species of cereal or legume led them to take actions that could increase the grain yield of that desired species. Those actions (e.g., tillage, sowing and harvesting) would have increased the incidence of individuals carrying gene mutations that control spike and pod shattering, which would eventually have led to a plant population that was incapable of sustaining

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itself in the wild, rendering it dependent on humans for its existence, that is, the domesticated species. This approach thus implies the development of a symbiotic relationship between humans and plants: while humans became dependent on plants for their nutrition, domesticated plants became dependent on humans for their continued existence, since without purposeful sowing (having lost essential wild adaptations like effective seed dispersal mechanisms and the trait of seed dormancy), they would become extinct. According to this approach, then, it may be stated, perhaps cautiously, perhaps in jest, that it is unclear who domesticated whom: had the humans domesticated plants, or had plants domesticated humans? Given the collective world efforts invested in research, nutrition and protection from pests and disease in wheat, maize and rice fields, humans can ‘easily’ be seen as ‘serving’ plants rather than the other way around. In the Mediterranean vegetation zone within the Levant, most annual plants germinate in the autumn, once the rainy season has begun, and they then wither and distribute their seeds at the end of spring or early in the summer. Only a few plants exhibit a life cycle beginning with spring germination, which would require green foliage during the dry summer season. Our colleague Professor Simcha Lev-Yadun has conducted plant surveys throughout Israel and has shown that many plant species that maintain green summer foliage – including Scolymus sp. (spotted golden thistle), Alhagi sp. (camelthorn), Carthamus sp. (distaff thistle), Ammi sp. (bishop’s weed), Euphorbia sp. (spurge) and Ononis sp. (restharrow) – are toxic or thorny, or they boast other defences aimed at repelling grass-eating animals. Their thorny and toxic qualities offer these species an advantage in an environment shared by grazing animals (both wild and domesticated herds). LevYadun thus notes that it is unsurprising that most annual domesticated crops in the Near East begin their cycle after the onset of the autumn rains and develop during the cold and rainy season, given that the local summer flora is naturally sparse, difficult to treat or truly toxic. Studying the yield potential of Near Eastern wild cereals shows that large extents of natural populations may be harvested in a short time with simple instrumentation. Such controlled experiments were conducted in eastern Turkey and in Israel, demonstrating that enough spikes can be harvested within an hour to produce a clean (post-threshing and clean of chaff ) grain yield of between 250 g and up to 1 kg. It is therefore easily conceivable that wild wheat and barley served as an important food source even before domestication and the emergence of agriculture.1 It is thus clear why some scholars adopt an approach, as discussed earlier, describing plant domestication as a long-term reciprocal relationship between humans and their cereal food plants. In contrast, the natural grain yield of wild peas and lentils in Israel portrays quite a different picture (Table 9.1). Grain yield of wild legumes is considerably lower than that of cereals. Gathering capacity of wild lentil and pea in Israel was respectively typically under 10 and 50 g of seed per hour of gathering. Given the sparse nature of legume populations in the wild, sites that yield even over 500 g of seed per hour of gathering

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Table 9.1 Grain yield obtained from foraging wild cereals and legumes

Plant species Legumes:** Odem lentil Lens odemensis Oriental lentil Lens orientalis Oriental lentil Lens orientalis Wild pea Pisum humile Wild pea*** Pisum fulvum Wild pea*** Pisum elatius Cereals: Einkorn wheat Triticum boeoticum Wheat and barley Wheat Triticum dicoccoides Barley Hordeum spontaneum Oats Avena sterilis

Site,* year

Grain yield (gram per hour of gathering by single individual)

Shipon Mountain, 2006

8–25

Mount Hermon, 2006

5–15

Mount Hermon, 2007

3–45

Mount Avital, 2006

5–150

Judean, Carmel and Galilee mountains, 2006 Western and Eastern Galilee, 2006

1–250

Viranshahir, East Turkey, by J. R. Harlan, mid-1960s Northern Israel, by Kislev and students, 2000–2002 Upper Jordan Valley, by Ladizinsky and students, early 1970s Upper Jordan Valley, by Ladizinsky and students, early 1970s Upper Jordan Valley, by Ladizinsky and students, early 1970s

1,000

Cereals harvested by sickle and grain separated after threshing:** Oats Judean Mountains and Rosh Pina, 2006 Wheat and barley Judean Mountains and Rosh Pina, 2006

6–600

50–240 240 100 50

4–400 1,300–3,400

* All sites are in Israel unless otherwise noted. ** Experimental harvests conducted by Gopher, Abbo and students. *** Wild pea species that were not domesticated.

are restricted in their total yield due to the limited size of the population. It is thus difficult to consider the lentil as a potential symbiosis partner (prey–human predator relationship) prior to its domestication. The same is true for pea, chickpea and bitter vetch, the wild forms of which share the same characteristics of low grain yield and sparse, patchy populations. Moreover, in wild pea gathering experiments that we have conducted in Israel over the years, we noticed that two wild species (which were never domesticated; see Table 9.1) produced higher yields than the species that was eventually domesticated. In other words, with respect to wild pea, yield

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considerations and gathering capacity must not have been key factors in choosing the domestication candidate. A special case that we encountered in 2006 in the area of Mount Avital in Israel offers a glimpse into the spatial distribution and efficacy of wild legume yield gathering. A single patch comprising several Pisum humile plants produced a yield of approximately 5 g in two minutes of gathering from adjacent pod-bearing plants. However, in those two minutes, the gatherer had exhausted the yield potential of peas in that patch. Table 9.1 shows a computed grain legume yield of 150 g per hour of gathering, but we never encountered a population of Pisum humile or Pisum elatius that allowed for more than just a few minutes of gathering from the few individual members of the patch’s populations. This contrasts with the continued, large and dense wild cereal populations that sustain high and stable yield potential. We must thus seek a different motive for legume domestication. In jest we say that only a person who has not tasted hot lentil soup on a wintery day requires scholarly explanations regarding the importance of lentil domestication. Legumes are protein-rich compared to cereals, and the composition of both, consumed together, supplies a healthy, balanced diet. The nutritional logic is therefore clear for adopting a selection of crops from both plant groups when it is decided that the economy should be based on food production, so that ‘man doth not live by bread alone’. The importance of lentil and its place in the Mediterranean diet is emphasized in the biblical story of Esau, who sold his birthright to Jacob in exchange for a bowl of lentil stew. Having returned hungry and tired from the field, he petitioned to Jacob: ‘Feed me, I pray thee, with that same red pottage’ (Genesis 25:30, KJB). We do not know what caused the stew to turn red since tomato arrived in the region only after the discovery of the Americas. According to the Bible, Jacob supplied Esau with bread and lentil stew in exchange for his birthright; note the cereal and legume, which together supply whole nutrition with respect to protein composition. In addition to protein content, it is likely that the flavour and scent of the different grain domestication candidates would have influenced the choice of domesticators in their adoption of specific grains. We may assume that, like ourselves, our predecessors would have been picky eaters (see Glossary, Botany, Ecology and Agronomy, Selective nutrition). In our view, the choice of species to be domesticated is likely to have also been based on additional knowledge, the source of which is unknown to us, and perhaps Neolithic individuals had keener senses and were more attuned to subtleties than we are. A comparative biochemical study conducted by Professor Zohar Kerem in wild and domesticated chickpea may shed additional light on the nutritional aspects of domestication. Chickpea seeds are rich in tryptophan, an essential amino acid in human nutrition. Amino acids are the building blocks of protein in all living organisms, humans included. An essential amino acid denotes that the human body is unable to produce it, and we are therefore required to consume it through our foods (similar to vitamins). Tryptophan is found only infrequently in vegetarian foods, and its concentration in wild chickpea is also quite low. Nevertheless, most domesticated chickpea cultivars examined showed higher

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concentrations of this amino acid compared to wild chickpea. It thus appears that subsequent to domestication (having obtained a free-germinating stock), early farmers focused their attention on seed content to choose those types that boasted the higher nutritional value. With no biochemical education and matching equipment, how could early farmers distinguish between seeds based on their tryptophan concentration? The answer lies in our sense of taste: different nutrients, such as sugars, fat, aromatic compounds and apparently also amino acids such as tryptophan, influence our taste preferences in food. Research shows that animals are able to identify within seconds or minutes tryptophan-rich foods among a broad choice of foods, and that indeed it is favoured by them. Tryptophan plays another important role as it is the precursor for the molecule of serotonin, a neurotransmitter that is essential for healthy brain function in humans. A tryptophan-packed diet increases tryptophan blood concentrations, and hence increases the chances of tryptophan reaching the brain and being converted to serotonin. Furthermore, consumption of tryptophan-packed foods (such as hummus, chickpea _ spread) together with carbohydrates (such as pitta bread) increases the transferability of tryptophan to the brain due to the secretion of insulin by the pancreas, which occurs when eating carbohydrates. Hence, hummus with pitta, falafel with pitta or lentil and _ bulgur (known as mjadra) – all typical Mediterranean foods – are quite ‘high-tech’ dishes designed specifically to allow for optimal exploitation of the food and its ingredients. It is nevertheless puzzling how farmers, devoid of biochemical knowledge and equipment, were able to conjure these shrewd food combinations. It seems that tryptophan-packed nutrition had many outcomes. For example, satiety at the end of a meal. Visitors to the Near East may remember the feeling of comfort they experienced after eating a dish of hummus with pitta bread. A breastfeeding mother on a _ diet rich in tryptophan may increase the intervals between the meals of her breastfed baby. Tryptophan consumed directly by babies, when they are able to consume foods that contain it, leads to a similar result. Just as modern-day mothers can discern their infant’s comfort or discomfort following a meal, or as we are able to tie the feeling of drowsiness to some foods and alertness to others, it is likely that Neolithic people were also able to associate different bodily feelings with the composition of their foods. Studies have indicated that serotonin may affect impulsiveness, increase obedience and impact social functions under stress. Serotonin has also been found to influence the secretion of hormones that are related to monthly cycles (and hence frequency of ovulation) of mammalian females, humans included. It is therefore plausible that a diet rich in tryptophan would increase the frequency of ovulation, which would have certainly merited the important part that chickpea was awarded in Levantine and Mediterranean nutrition. Soaking or cooking chickpea, as tested under laboratory conditions, shows that tryptophan remains unharmed. No less important is the fact that daily recommended tryptophan values (by modern standards) are achieved through the consumption of fairly few chickpea seeds.

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Chickpea is not the sole source of tryptophan, which is found in animal protein as well as other legumes. Despite this, no information is available regarding levels of tryptophan in wild relatives of the domesticated pea and lentil. We therefore cannot establish whether chickpea represents a unique case or a general inclination. In any event, it is clear that the (agricultural and nutritional) value farmers assigned to the chickpea must have been considered important enough for them to overcome many agronomic challenges, especially a susceptibility to a destructive fungal disease (ascochyta blight, which spreads via rain splash), and the need to develop the spring sowing husbandry in order to avoid the disease. The complementary nutritional nature of cereals and legumes was probably only one reason for their joint adoption as domesticated crops. They are additionally complementary in the agricultural-agronomical sense. As discussed in Chapter 8, the cereal’s determinate growth, ending with a spike, differs significantly from the legume’s indeterminate growth, where seed-bearing pods develop at branch internodes. This growth pattern allows cereals to make better use of early winter rains and usually ensures higher grain yield compared to legumes. In contrast, the growth habit of legumes allows for more efficient use of late seasonal rains, which would increase grain yield of legumes while only affecting cereal grain to a limited extent. The differences in the responses of the two plant groups to the seasonal rain profile results in an interesting phenomenon with respect to the compensation potential of grain yield. An analysis of annual yields over time in traditional farms in early twentieth-century Israel as well as in Greek villages in the 1960s shows that in those years in which cereal yields were above average, legume yields were fairly small, and vice versa; in years characterized by low cereal yields, legume yields were higher than average. This is significant in the prevalent climate of the eastern Mediterranean Basin, where winter rainfall and temperature regimes vary considerably from year to year, between December and April. The different responses of cereals and legumes to these growth conditions result in seasonal variance, under traditional (and modern) farming, namely, an inverse pattern of record yields in the two groups over the years. Since subsistence farmers require stable yields (more than record yields), dividing fields into cereals and legumes serves as insurance of sorts for farmers, who are partially compensated by the high yield of one plant group when the other group produces an inferior yield. In the absence of transportation infrastructure for grain bulks and hence the inability to mobilize grains to remote locations in the world (as occurs in modern times), this agricultural compensation mechanism was of the highest, existential importance (see Chapter 6). In both traditional farming and modern agricultural systems, field areas dedicated to legumes make up approximately 15% of fields dedicated to cereals, which are the nutritional and agricultural backbone of the system. It should be noted here that due to their high protein content (20–25%) the contribution of grain legumes to human nutrition exceeds their humble role in the arable land area. Therefore, to maximize the yield compensation potential of the traditional systems, farmers rely on a different

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agronomic phenomenon, namely, a step-wise ripening pattern of the different crops in each group. Within cereals, barley is the earliest to mature, durum wheat matures next and einkorn wheat matures last. Farmers sowing plots with all three cereals will enjoy the same type of insurance noted above. If the rainy season ends earlier than expected, barley fields will supply most of the grain yield; in an average year, durum wheat fields will supplement the yield; and in an affluent year, the grain yield will be further supplemented by einkorn wheat. A similar gradient is observed among legumes: lentil is the earliest to mature and has fairly low growth potential, thus requiring little rain, whereas pea is a relatively late-maturing crop that requires considerable amounts of precipitation for its growth. Thus, in years of drought, most yields will originate in lentil and, in rainy years, in pea. In a moderate spring that follows a wet winter, the chickpea, sown in springtime, will bring in a high yield. Among fodder legumes domesticated as animal foods, bitter vetch is a small-statured crop and thus requires little precipitation whereas common vetch (Vicia sativa) is more vigorous and would yield better in an abundant year. Taken together, the considerations detailed above show that the choice of the Near Eastern Neolithic founder crops was truly perfect. On the one hand, it ensured a balanced diet, while, on the other, the composition of species produced an agronomic combination of the two (biologically differing yet complementary) plant groups. In other words, sowing fields of all three cereals and all three legumes, given the maturity gradient among the crops of each group as well as between the two groups, provided farmers with the best possible ‘insurance package’ they could have procured in ancient times against the uneven, often erratic, rainfall of the eastern Mediterranean Basin. Given the considerations we have portrayed thus far, we find it difficult to accept that the Near Eastern choice of grain species for domestication was the result of an ‘ecological mishap’, as proclaimed by the Dump-Heap Hypothesis. We also fail to see how this nutritional and agronomical combination could have developed randomly following continued gathering and manipulation, since, as we have shown, lentil or chickpea were not likely to have been used as a major food source prior to domestication. Thus, contrary to certain models portraying the transition to agriculture and the emergence of domesticated plant species as the result of a random process involving automatic and unconscious selection, we contend that domestication, including species selection, had to have been the result of a knowledge-based process. This process must have rested on the foundations of the hunter-gatherers’ long tradition of intellectual curiosity and knowledge accumulation for the sake of gaining knowledge, followed by knowledge classification, testing and transmitting to future generations. Without intimate knowledge of their surroundings, animate and still resources within it, the understanding of and ability to forecast seasonal phenomena, the ability to locate water resources, salt deposits, flint nodules and plants and animals that could all be used for different purposes, Paleolithic hunter-gatherers and early Neolithic societies would not have been able to successfully sustain themselves through millions of years prior to the emergence

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of agriculture or to settle all continents other than Antarctica, including locations characterized by extreme conditions such as desert lands and the Arctic Circle. This intimate knowledge and excellent familiarity with the environment, developed over a very long time, are the factors that enabled Neolithic societies to choose certain, desired plant species and types (within species) as prospective agricultural crops. We stress that our choice to name this chapter as we have, ‘The Choice of Plant Species for Domestication: Agronomic and Dietary Considerations’, is our way of stating that this was a knowledge-based move, undertaken in complete mindfulness by those involved in its initiation. Undoubtedly, choosing a source of seeds from wild plants, and subjecting it to a cultivation regime of sorts, amounts to a selective act; it is also quite clear that genetic processes taking place in cultivated fields are different from those occurring in wild populations. It is thus obvious that in any – natural or human-managed (including by cultivation) – population, processes take place that accompany selection. For example, selection of a stock characterized by non-shattering spikes within a large and variable population will immediately result in a lineage in which the genetic variability has been limited. We further elaborate on these and other processes in Chapter 12, where we discuss the evolution of crops under domestication. Even if unexpected genetic responses accompanied selection, there is no reason to assume that the process was essentially unguided or unconscious. In our view, the deep biological understanding and the intimate familiarity of our Neolithic forebears with plant species and their diverse populations, as well as the unique properties of each, would have certainly enabled farmers to notice and respond to developments taking place in their fields. For example, if a certain pool of seeds was found to be undesirable due to its taste or other qualities, or perhaps an undesired degree of susceptibility to a disease or a pest, we can safely assume that farmers could resample wild seeds in order to free their crops of the unwanted trait. It is otherwise difficult to decipher the success and economic significance that the Near Eastern founder crops currently enjoy in the Western world and beyond. The initial success of the founder crops gave birth to ancient Near Eastern cultures that eventually became the cradle of Western civilization. Over the years, wealth accumulation that was the result of agricultural progression allowed European superpowers to achieve global dominance, which increased enormously following the industrial revolution of the eighteenth and nineteenth centuries. As we understand it, the central economic role played in current world economics by grain crops that had been domesticated in the Near East during the Neolithic period over 10,000 years ago serves as testimony to the biology-related knowledge, acuteness, initiative and resourcefulness of Neolithic populations; and likewise with the African sorghum and pearl millet, East Asian rice and soybean, Meso-American maize and beans, South American potato, quinoa and groundnut, and so forth. In concluding this chapter, we would like to reiterate that when discussing the differences between wild and domesticated plants (Chapter 5), we noted that most domestication traits are recessive (see Glossary, Genetics, Recessive trait) and that the

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Near Eastern founder crops are self-pollinating. Under these conditions, the establishment of mutants that lead to domesticated morphology (phenotype) in these populations when managed (in cultivated fields) under a regime of sowing, harvesting, sowing from the same yield that was harvested in that field, and so forth, is a fairly rapid process in which self-pollination is an advantage (see Figure 5.1). The fact that eight of the founder crops of Near Eastern agriculture are self-pollinating is significant. We also remind readers of Zohary’s claim that the choice of self-pollinating species for domestication was not coincidental and that it is highly unlikely to have been a random occurrence.2 We embrace Zohary’s approach to this issue and regard it as robust support of our claim that plant domestication in the Near East was a cultural, knowledge-based process that did not rely on randomness or unconscious selection. K E Y PO IN TS A ND BE Y ON D

• Plant domestication candidates originated among local species and were therefore adapted to the local climatic rhythm. The yield potential of the founder crops indicates that plant productivity was not the sole criterion in the choice of candidates. In this respect, the choice of legumes cannot be interpreted as the result of an ongoing predator–prey relationship, that is, wild legumes are unlikely to have been staple food sources for Paleolithic hunter-gatherers. • The crop package selected for domestication comprised species that complemented each other in terms of nutritional value and agricultural-agronomic qualities due to their varying responses to the local precipitation and temperature regimes. In the absence of insurance against natural disasters or help from other world regions, only such a complementary crop package could facilitate stable annual yield. • Evidence has been presented insinuating that early farmers successfully selected among the standing natural genetic variation accessible to them in the wild (and later in their fields) those plant types that were characterized by unique nutritional qualities. We believe that the vast economic significance of the crops selected by early Neolithic farmers in the current global economic system attests to the fact that this was a very successful, knowledge-based process that was far removed from randomness. • In many senses, choosing the different plant (and animal) species to be domesticated placed Neolithic domesticators at the forefront of the Neolithic ‘science of the concrete’ à la Claude Lévi-Strauss, inspiring in us awe and a sense of modesty.

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W H E R E , W H E N A N D H O W D I D NE A R E A S T E R N PLANT DOMESTICATION OCCUR?

In this chapter, we discuss the questions of where and when Near Eastern plant domestication occurred: Did it transpire in a single, defined area, or perhaps it took place in different areas across the entire Fertile Crescent? Was each crop domesticated separately, or perhaps several crops were domesticated together as a harmonious agricultural package, in which crops complement each other nutritionally and agronomically? Was each crop domesticated on several occasions, each in a separate location, or did each crop undergo a single domestication episode? We look into evidence for each of these questions, based on several fields of knowledge, including geobotany, archaeology, archaeobotany and genetics – some of which we touched upon in earlier chapters. Finally, we examine whether the picture that emerges through these combined lines of evidence allows us to trace the progression of the new, agriculturally based economy throughout the Near East and from there to Europe, Asia and Africa. Readers will recall Childe’s theory, presented in Chapter 3, which describes the inception of agriculture as a response to the physical proximity of humans to plants and animals – all taking shelter from a climatic crisis in the fertile river valleys of the Nile, Euphrates and Tigris. The deep contrast between the Near Eastern landscape, a semi-arid region including desert lands characterized by a hot, dry summer and the green vegetation typical of these river valleys, as well as the constant water flow that sustains fresh vegetation even in the dry season, conjointly bred the name of ‘the Fertile Crescent’. This name is meant to denote the area that spans from the Zagros Mountains and valleys of north-western Iran, the valleys of the Euphrates and Tigris in Iraq, northwards alongside the Zagros and rivers to their point of origin in south-eastern Turkey, and from there south-west through the Mediterranean

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zones of Turkey, Syria, Lebanon, Jordan, the Palestinian Authority and Israel; some also include the Nile Valley of Egypt within the area of the Fertile Crescent. All in all, the area is shaped like a crescent, the head of which is located in south-eastern Turkey, while one cusp faces south-east towards the Persian Gulf and the other south-southwest along the eastern Mediterranean coast (Figure 2.1). The mountainous regions of this area host diverse flora, both annual and perennial. Vast regions – located between current Israel and western Iran – enjoy an average of over 350 mm of annual precipitation and host predominantly oak and terebinth trees while common annuals include wild barley, wild emmer wheat and wild oriental lentil. The incidence of the remainder of the crop progenitors that are discussed here is more limited; for example, the wild progenitors of pea, bitter vetch and flax are far less prevalent in the southern Levant compared to the cereals. Similarly, einkorn wheat cannot be found south of the Lebanon mountain range. The wild progenitor of chickpea grows only in a very limited geographic area in south-eastern Turkey, between the provinces of Adiyaman and Hakkari. Drawing the distribution areas of all species on a single map shows an interesting pattern, because all of the wild progenitors of the Near Eastern domesticated grain crops converge in the south-east region of Turkey (Figure 10.1). Assuming that availability was a necessary condition for domestication and that community members would have first domesticated species that were commonly found and available to them in their natural environment, we must conclude that this area of south-eastern Turkey played an important role in plant domestication and the inception of the Agricultural Revolution at large. Archaeological data, too, point to the head of the Fertile Crescent (south-eastern Turkey and northern Syria) as a key area in which major, broadly encompassing cultural developments have unfolded. Archaeological findings in the area (see Chapter 2) indicate that many technologies spread from this centre in south-eastern Turkey and northern Syria to the southern Levant, central and western Turkey and then to the Balkans, into Europe and the western Mediterranean. Charred plant remains originating in archaeological excavations also show that south-eastern Turkey and the adjacent region of the Middle Euphrates Basin in northern Syria served as an important area. Plant remains can be dated based on the measurement of the radioactive isotope 14C, which is assimilated by the photosynthetic processes into plant matter. Since archaeobotanists are able to distinguish remains of wild from domesticated cereals (based on the disarticulation scars on ripened spikelets or the broken axes of the threshed domesticated spikes), it is possible to determine the age of the layer in which domesticated cereals first appear at archaeological sites. The earliest recognized domesticated plants indeed originated in south-eastern Turkey and the adjacent region of northern Syria. Genetic experts and other scientists studying plant domestication are interested in knowing the number of times each domesticated species was domesticated. This is an important question as it pertains to the diversity of the genetic pool available to farmers and breeders for each of the domesticated species. The importance and influences of this genetic diversity are further discussed in Chapter 12, where we look at crop evolution

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Figure 10.1 Distribution of wild progenitors of Levantine founder grain crops (circle denotes the core area of Neolithic agriculture); The small maps show the distribution areas of each of the wild progenitors of the Near Eastern founder crop package: A) chickpea (shaded); B) einkorn wheat; C) emmer wheat; D) barley; E) lentil; F) pea; G) bitter vetch. Relevant archaeological sites are numbered: 1) Çayönü; 2) Cafer Höyük; 3) Nevali Çori; 4) Göbekli Tepe; 5) Dja’de; 6) Jerf el Ahmar; 7) Mureybet; 8) Tell Abu Hureyra; 9) Hallan Çemi Tepesi; 10) Qermez Dere; 11) M’lefaat; 12) Tell Aswad; 13) Yiftael; 14) Jericho, Netiv Hagdud, Gilgal I. The red triangle denotes Karacadag. Modified from Lev-Yadun, Gopher and Abbo 2000). A colour version of this map can be found at www.cambridge.org/abbo-gopher.

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under domestication, that is, after the biological domestication was achieved and the cultural transformation known as the Agricultural Revolution was underway. In this respect, we note here that a crop that has been domesticated more than once is in fact a ‘genetic reservoir’ of sorts, created through several independent samplings of wild seed stocks for the purpose of domestication by Neolithic farmers at different points or at different sites (foci) within the geographic range. Whether genetic diversity was amassed through geographically distant sites is of lesser biological importance although it is significant from the cultural perspective. This information is likely to facilitate answering the question of whether domestication was an independent initiative or an invention that was promoted in different areas or whether it involved a flow of ideas – or seeds, in this case – from a source area to nearby neighbours who had adopted them. In the case of a cultural-perceptual influence, it may be that the recognition of the potential that agriculture may bring with it caused Neolithic community members to sample adequate local plant stocks (of the same species) that were available to them in their area of residence in an attempt to domesticate on their own – effectively, a secondary domestication episode. In the case of direct contact, they may perhaps have sown seeds that had originated in the source region, passed on through nearby or more distant neighbours. A classic genetic approach by which to evaluate the number of domestication events (which seems somewhat simplistic nowadays) is based on the genetic study of hybrid offspring from crosses between domesticated and wild types. The aim of these experiments is to identify the number of genes that influence the main domestication traits. It is assumed that a single, identical gene governing a single trait, for example the nonshattering trait of spikes, in all domesticated types (of the respective crop species) indicates a single domestication occurrence. Alternatively, if several such genes can be identified, each conditioning a non-shattering spike in a distinct cultivar group, a possible explanation could be that each cultivar group represents an independent domestication lineage. Another way to assess the multiplicity or exclusivity of domestication events is the chromosomal behaviour of hybrids from crosses between wild and domesticated stocks. Evaluation of the degree of chromosome pairing can be obtained from microscopic observations of meiotic cell divisions that lead to the formation of pollen grains in the anthers. Based on the extent of chromosome pairing, it is possible to determine the extent of genetic affinity between the domesticated and wild stocks used in the hybridization experiment. In wild species, and particularly those of wide geographic range such as the oriental lentil, several chromosomal types were found that differed both from each other and from the domesticated species. Similarly, wild lentil types were found to exhibit different chloroplast DNA profiles (see Glossary, Genetics, Chloroplast). In contrast, all tested domesticated lentil cultivars exhibit a single chromosome type and chloroplast DNA pattern. Moreover, only wild types of lentil in eastern Turkey were found to exhibit chromosomal and chloroplastic similarity to domesticated stocks. This evidence leads to the conclusion that, in all likelihood, the lentil underwent only a single domestication event and that the seed stocks chosen for domestication originated in

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eastern Turkey. Hybridization experiments conducted in pea among a fairly small number of wild and domesticated stocks showed that pea samples that originated in Turkey and the Golan Heights (northern Israel) were closest to the studied domesticated cultivars. In other words, for the pea, crossability relations data are not as conclusive as for the lentil, although Turkey’s honorary status still holds. Indeed, French scientists (led by Dr Judith Burstin) have recently analysed the domesticated and wild pea genome sequences and their findings support a south-eastern Turkish origin of the domesticated pea stock, thereby corroborating our core area hypothesis. It is commonly assumed that several past biological domestication events should be reflected in the population structure of the domesticated species. In other words, it is assumed that following an evolutionary history in which several adoption (and domestication) events of wild seed sources were included, a similar number of cultivar groups would be identified that are distinguished from one another, as each would be genetically affiliated with a wild population of a different geographic origin. It is expected that a domestication history occurring in multiple sites would allow for the identification of approximated domestication location(s) based on the genetic profile of domesticated stocks and affiliated wild populations that can be found in the current native range of the wild species. As above, then, it is assumed that domesticated cultivars will exhibit genetic affinity with current wild populations in the area of their origin (domestication). In recent decades, modern biology and DNA-related technologies have opened new vistas by which to compare genetic profiles in a similar manner to tests conducted to identify criminal perpetrators or for the purpose of establishing paternity. In a pioneering study published in 1997, Professor Manfred Heun tested DNA profiles of wild and domesticated einkorn wheat and his results showed high similarity among domesticated cultivars and that the latter most resemble wild types collected in south-eastern Turkey, in the vicinity of the mountainous basalt range of Karacadağ near the city of Diyarbakır. This finding revived the discussion of the questions that are at the heart of the current chapter: Was a single domestication event involved? And, if so, where did it occur? This genetic study indicated that south-eastern Turkey was the likely place of origin of einkorn wheat, while archaeological data are highly suggestive that this wheat was indeed domesticated in a single cultural centre located in south-eastern Turkey. The genetic profiles of (domesticated) durum wheat and the wild types of emmer wheat also show that domesticated cultivars most resemble wild stocks that originate in the very same area in south-eastern Turkey. One of the arguments against a single domestication core area in south-eastern Turkey (formulated by Dr George Willcox) is based on the adaptation of different plant types to climatic and soil conditions. Following this argument, barley, which grows successfully in fairly arid areas, was domesticated in the southern Levant (Israel, Jordan, Lebanon and southern Syria) whereas einkorn wheat, which requires greater precipitation and a colder climate, was domesticated in the northern Levant (northern Syria and south-eastern Turkey). This model suggests that chickpea was domesticated in Turkey while lentil was

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domesticated in both the northern and southern areas of the Levant. Moreover, some experts claim that both wheat and barley were independently domesticated by different communities that cultivated wild plants and that the farming activities carried out by them (sickleassisted harvesting, threshing, land tilling and so forth) created selection pressures similar to those we have discussed in earlier chapters, until domesticated forms (non-shattering spikes) emerged in their fields and became prevalent. Other experts advocate that the entire Levant was a mosaic of wild plant cultivation centres, some of which became domestication centres following (husbandry-driven) automatic selection and unconscious human activity. These claims cannot be solved with the DNA patterns of either durum and emmer wheat or einkorn wheat, which attest to a single domestication event. Proponents of the decentralized, multi-site domestication process (termed autonomous domestication) further claim that following the initial domestication at different sites, many genetic lineages of domesticated wheats became lost over the years, leaving only the most successful lineage, which was then distributed through different communities by means of exchange trade. No evidence has been found to support this reconstruction of events. It is also difficult to fathom that out of a broad range of domesticated lineages (within each species) all but one would have been lost, and that the surviving lineage (in each of the respective species, as documented for both einkorn and emmer wheat, pea and lentil) would be genetically associated with wild stocks originating in, of all places, south-eastern Turkey. If we were to accept that domestication occurred at different sites and that most lineages were lost, what are the odds that surviving lineages (other than barley, perhaps) would point to the same area in south-eastern Turkey? Note that suggestions of several geographic origins of domesticated barley include the northern Levant. In our view, the likelihood of such a scenario is extremely low, and therefore, based on currently available scientific knowledge, claims advocating dispersed (autonomous, independent) domestications across the Near East are unacceptable. In this context we wish to mention the idea, or the concept, of ‘preemptive domestication’, coined and formulated by Professor Jared Diamond. This idea suggests that the successful domestication of one species precludes the need to domesticate other plant species with a similar potential. The preemptive concept does not suggest that the domestication of a cereal would prevent the adoption of a legume; rather, it relates to similar species. This would be applicable to species such as chickpea (three sympatric wild species with large seeds native to south-eastern Turkey), lentil (three similar species native to the northern Levant) or pea (three large-seeded species native to the region). Following this logic, the likelihood is minimal for a domestication event in a plant species to repeat itself (in the presence of similarly useful wild species), and in the case that it is repeated, it would be unlikely that it would always involve the same combination of specific taxa. Diamond (1997: Chapter 12) uses the evolution of writing as a cultural analogy for the preemptive domestication concept, explaining that once it came about it was unlikely to happen again within a short time frame in the same cultural interaction sphere. Applying the idea of preemptive domestication to the Neolithic Levant offers a solid conceptual

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alternative to the protracted-autonomous model and corroborates our core area-one event reconstruction on Near Eastern plant domestication. As noted, the non-centred, dispersed (autonomous) domestication model claims that early farmers at first cultivated just the one (wild) cereal that fitted their environment. However, as we have shown when discussing the significance of yield stability, relying on a single crop can be risky from the agricultural and economic perspectives while reliance on diverse crops (mixed farming) from both the cereal and the legume plant groups is highly advantageous as it offers nutritional complementation and the potential of yield stabilization. Moreover, we are unaware of archaeological sites that yielded plant remains of a single wild or domesticated cereal or a single wild or domesticated legume. The archaeobotanical record of the relevant time period is diverse and includes many of the wild species that became domesticated as well as many of the domesticated forms. This fact further weakens the notion of the sequential (progressive) model of domestication, where several species were adopted, one at a time. Such a model has yet to explain the emergence of the ‘winning coalition’ of founder crops in light of archaeological (and other) evidence that negate such a development. The development of the full founder crop package sits better with the model we are suggesting – in which all crops were simultaneously and rapidly domesticated following agronomic and nutritional considerations. The combined evidence detailed above regarding the naturally overlapping distribution of all wild progenitors of the founder crops, and especially the fact that the wild progenitor of the chickpea is found only in that area of south-eastern Turkey (i.e., the chickpea could not have been domesticated elsewhere), near the location of origin of einkorn wheat, led Lev-Yadun and us, back in the year 2000, to suggest that this region in south-eastern Turkey was the cradle of Near Eastern agriculture and thereby the cradle of Western civilization at large. We further suggested that all founder crops were domesticated in this area (see Figure 10.1) within the same time frame, and – at least for most crops – probably in a single domestication event. Back in 2000, we also suggested that the model should be re-evaluated based on a similar genetic investigation of the geographic origin of durum wheat. Such an investigation was indeed conducted a few years later, and, based on DNA profiles, it was concluded that the wild stocks most similar to domesticated emmer and durum wheat did, in fact, originate in the same geographic region near the Karacadağ Ridge in south-eastern Turkey. Despite this additional support, our suggested model is not widely accepted. Most scholars of Near Eastern domestication prefer the view that domestication recurred several times, taking place in different regions of the Near East. We shall not elaborate here on the cultural preferences of students of Near Eastern plant domestication, yet, it is notable that most scholars also advocate that the transition to agriculture progressed in the wake of a random process based on automatic, unconscious selection. This approach implicitly assumes a food crisis that affected the entire Near East and that numerous populations throughout the region responded to this crisis through their attempt to intensify resource availability by cultivating the species that had already served as food sources; hence the

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justification for a multi-site (non-centred), autonomous process of domestication. Even if we accept the claim that attempts to intensify grain availability were conducted at different sites throughout the Near East in response to some environmental change, it is difficult to accept the second claim, that effective actions in handling this environmental crisis were taken unconsciously. We thus find the logic of these claims flawed. While the genetic investigation of domesticated and wild einkorn, emmer, lentil, pea, chickpea and flax indicates a probable single domestication in Turkey, findings concerning barley are still inconclusive. Several research teams have produced contradictory reports – some advocating a single domestication event while others stipulating two or more allegedly independent domestication events and centres (including our suggested core area in south-eastern Turkey). We have no choice but to wait for future research that may clarify the challenging issue of barley.1

K E Y PO IN TS A ND BE Y O ND

• Radiocarbon dating of plant remains from archaeological sites has made it possible to determine that the emergence of domesticated crops occurred approximately 10,500 years ago. The sites that yielded the earliest remains of domesticated crops are located in south-eastern Turkey and northern Syria, an area that sprouted many cultural and material innovations that are documented in archaeological records. • Some form of interaction, albeit at varying degrees, was present throughout most of the Levantine expanse during the Pre-Pottery Neolithic period. This expanse was termed the Levantine koine by Jacques Cauvin, while others called it the Levantine interaction sphere, implying a well-connected expanse through which both information and materials flowed. • The region spanning south-eastern Turkey and northern Syria is the only area within the Near East in which the wild progenitors of all founder grain crops grow in proximity (sympatry). We therefore suggest that this is the location of origin of Near Eastern agriculture. We term this region the core area. • The study of genetic profiles of domesticated species and their wild progenitors shows that cultivars of einkorn wheat, emmer and durum wheat, pea and lentil each are genetically affiliated to wild stocks originating in the core area or closely nearby. The wild progenitor of chickpea is limited in distribution to the core area alone, implying that chickpea domestication must have occurred in this region. • Limited studies of wild and domesticated flax suggest a single eastern Turkish origin. Bitter vetch, in both its forms, wild and domesticated, still awaits thorough investigation. On the other hand, a wealth of genetic studies of wild and domesticated barley suggest multiple adoptions from the wild (including in south-eastern Turkey) but with no evidence to support cultural independence of those putative domestications. • Despite multiple opinions, no genetic or archaeological evidence has been presented to support the notion of multi-site (autonomous) domestication throughout the Levant.

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Nor is there any genetic or archaeological evidence of any domestication event earlier than the one that occurred in our suggested core area. It is thus difficult to assume that some areas within the Levant, especially in the northern regions, maintained a cultural autonomy that completely disconnected them from the rest of the region to the degree that enabled them to undergo autonomous (millennia-long) domestication processes. • Interestingly, although the domestication of animals is fundamentally different from the domestication of plants, the natural distribution areas of the wild forerunners of the domesticated goat, sheep, cattle and pig converge in south-eastern Turkey, the core area, where we believe plants were first domesticated (see Chapter 14). Linguistic studies suggesting that Indo-European languages originated in Turkey • (slightly west of the core area, where we believe plants were first domesticated) may be of particular value to this discussion considering that agriculture (domesticated flora and fauna) spread from the core area to central and western Turkey, the Mediterranean islands, the Balkans and into Europe. We note, however, that in recent years other reconstructions of this linguistic spread have also been suggested. • Recent studies of human aDNA from the Near East and various parts of Europe provide a growing amount of evidence indicating a flow of immigrants with an agricultural package from the Near East (Turkey) and into Europe reaching as far west as Spain and England.

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11 D O M E S T I C A T I O N OF F R U I T T R E E S I N T H E NE A R E A S T

Scientists generally agree that fruit trees were domesticated several millennia after the domestication of grain crops. And just as there is a founder group of grain crops, there is also a founder group of fruit trees. It is generally believed that the olive, fig, grape vine, date palm and pomegranate (Figure 11.1) were the first trees domesticated in the Near East. Before embarking on a detailed discussion of fruit trees, we briefly delineate the biological differences between the reproduction system of grain crops, discussed in earlier chapters, and that of fruit trees (Table 11.1). Other than the cross-pollinating faba bean, the grain crops domesticated in the Neolithic Near East are self-pollinating. This means that the fertilization process occurs within the flower after the anthers open and shed pollen grains on the stigma of the same flower. In this mode of pollination, even in the unlikely event of cross-pollination, the hybrid plant (created from pollen and eggs of two genetically distinct individuals) will yield, after about eight generations (of selfpollination), an array of distinct genetic types that breed true (each yielding offspring that are generally identical to their mother plants in traits (see Figure 5.1)). This is true for wild populations of many cereals and legumes: their members comprise different types, each of which yields progeny that are identical to the mother plant in nearly all aspects, with the exception of rare mutations. In contrast, reproduction in the fruit trees mentioned above is based on crosspollination. The date palm, for example, is a dioecious tree. Dioecious trees cannot self-pollinate; there are male trees that do not bear fruit and female trees that do and it is therefore technically impossible for female reproductive organs to be pollinated by

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Figure 11.1 Tree species domesticated in the Near East. (a) Date palm – courtesy of Yigal Elad; (b) grape vine; (c)–(d) olive; (e)–(f ) pomegranate; (g)–(h) fig. Colour versions of these images can be found at www.cambridge.org/abbo-gopher.

genetically identical male pollen. Wild grapes are also dioecious whereas domesticated cultivars bear a mutation that enables the development of both male and female organs within a single flower. Figs exhibit a more complex system, in which the fruit that we eat is not a true fruit in botanical terms but rather a complex inflorescence (termed syconium; plural, syconia) carrying hundreds of flowers on the internal wall of their inflorescence cavity. The common fig is a dioecious species with two tree types: polleniferous male tree (caprifig-goat fig, in which inedible ‘fruits’ are developed) bearing both male and female flowers; and female trees in which male flowers are degenerated so that they are completely dependent on cross-pollination from male trees. Fig pollination is carried out by a type of wasp that is about 2 mm long and which lays its eggs in the syconia of the male (caprifig) trees. Once wasps emerge from their eggs

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Table 11.1 Sexual versus asexual plant reproduction Sexual reproduction from seeds (cross-pollination)

Asexual reproduction (vegetative propagation)

Tissue of origin

Reproductive cells (pollen and egg cells)

Somatic cells (buds, bulbs, tubers, branches)

Number of parents

Two (male and female)

One, source of vegetative tissue

Offspring composition

Diverse, each offspring represents a unique genetic combination

Uniform, identical to original source of vegetative tissue*

Likelihood of inherited genetic changes

Relatively high, according to allele composition of the two parents

Slim, dependent mostly on frequency of somatic mutations

Lifespan of individual

Throughout lifetime of individual specimen

As long as clonal propagation of that stock is carried on

Characteristic

* With the exception of somatic mutations that occur during regular cell division and which may give rise to plant organs that bear new qualities (such as nectarine-bearing branches on a peach tree). Such cases notwithstanding, all offspring originating in somatic tissues of the same branch would be identical.

they exit the male syconium through its small opening (ostiole) carrying pollen from male flowers. With this pollen load, the wasp enters the inflorescences of the female trees, thereby pollinating the female flowers and enabling seed development. The flower structure in female syconia prevents wasps from laying eggs and therefore the wasps cannot reproduce therein. Due to their sweetness and succulence, the ‘fruits’ that develop on the female trees are edible, unlike the male (caprifig), which is inedible (because the ovaries of its female flowers each develop into a seedless gall containing the wasp larvae). Thus, there exists a mechanism in figs that ensures cross-pollination, similar to the date palm. The olive releases, during its flowering period, great quantities of pollen, some of which is carried in the wind to flowers of neighbouring trees in the population and beyond. Finally, pomegranate flowers, too, are structured for cross-pollination. What, then, is the significance of these differences between sexual reproduction mechanisms of annuals and perennials? With the grain crops, once a desired type of wheat (e.g., characterized by a non-shattering spike) or pea (e.g., free-germinating) is identified as a self-pollinating plant, its offspring would typically be genetically identical to the mother plant and hence exhibit the same desirable traits. In self-pollinating annual plants, then, once the desired plant has been identified and seeds have been multiplied over a small number of growing seasons, it is possible to then generate a large population and, in fact, create a landrace (in effect, a domesticated cultivar). In contrast, once a suitable fruit tree is found that yields pomegranates to our liking or particularly fatty olives, a seedling growing from its seeds is expected to produce a tree that is quite unlike the mother tree from which the seeds were collected, and it is also probable that

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its fruit yield will be low (and of different quality). This is because in most fertilization events, seeds are created from female eggs, which bear the alleles of the desired traits, and male pollen, which is of an unknown source and will typically not carry the desired traits. In other words, farmers domesticating trees required a completely different approach to reproducing the trees that they desired. Indeed, a quick review of the tree species mentioned earlier that formed the foundation of domesticated orchards and plantations shows that they may all be propagated asexually (vegetative clonal reproduction; see Glossary, Botany, Ecology and Agronomy, Clonal propagation) fairly easily. The term ‘vegetative reproduction’ denotes reproduction that does not involve the creation of sex cells and seeds. For example, the shoots grown by the date palm in its early years are, in fact, side branches developed from buds found in leaf (palm) axils. Since these shoots are developed from somatic tissues (Glossary, Botany, Ecology and Agronomy, Somatic tissues) rather than from seeds, they are entirely identical to the tissues of the parent tree (other than in cases of rare mutations). The basal part of the side shoot can easily be wrapped together with moist soil (or potting mixture), which induces root growth. Once the offshoot has developed roots, it may be disconnected from the parent tree and planted in a desired location (Figure 11.2).

Figure 11.2 Reproduction of date palm and removal of offshoots – courtesy of Yuval Cohen.

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In this manner, during the first years of growth of the individual date palm, about a dozen offshoots may be obtained before the tree grows out of its juvenile phase and its axillary buds are no longer vegetative. At this point, the tree begins to develop axillary inflorescences and bear fruit, hence new offshoots are no longer available for reproduction. Mature olive trees grow shoots near the base of the tree; these are branches that may be removed from the parent tree along with a portion of its root system and be planted independently. Unlike the date palm, where the equivalent act does not affect the root system of the parent plant, the parent olive tree sustains some damage when its basal shoots are removed, thereby limiting the number of shoots that farmers can remove from each tree. Reproduction of grape vines as well as reproduction of the fig and pomegranate trees is a considerably easier process. All three species are deciduous and are dormant during the winter. It is thus possible to reproduce these species by cutting off a branch towards the end of winter and planting it in moist soil. Typically, roots will form at the base of the branch above the pruned point, growth and foliage development will be renewed in the spring, and within a few years the plant will have matured and begin to yield fruit. The vegetative reproduction methods applied to each of the tree species that founded orchard and plantation farming thus produce offspring that are identical in their qualities to the parent tree. It seems that the grafting technique – conjoining a bud or branch of a desired tree (scion) onto a seedling (sprouted from a seed) or a mature rootstock – was developed only several millennia later. Archaeobotanical remains of fruits such as dates and grapes as well as charred wood remains have been uncovered at a variety of archaeological sites in the southern Levant. Based on 14C dates, we are able to determine whether these remains originated in layers that accumulated before or after plant domestication. Interestingly, almond stone fragments with their typical pitted surface were unearthed in several Near Eastern archaeological sites that were assigned rather deep Paleolithic dates. Why then is the iconic almond1 not mentioned among the founder fruit tree species of the Near East? The answer is probably related to the above discussion on the ease of clonal reproduction of the other fruit tree species. Unlike grape vine, date palm or the pomegranate, almond trees cannot be propagated from branch cuttings since these do not root easily. The wild almond kernels contain the highly toxic, cyanogenic compound amygdalin, and crushing the kernel releases cyanide. To grow almonds as fruit trees Neolithic people had to identify non-toxic almond types and overcome the propagation issue. Note that domestication syndrome traits may appear as spontaneous mutants among wild populations. This may be true to free germination in legumes as well as for sweet (non-toxic) almond kernels. We assume that (Paleolithic and) Neolithic people encountered naturally occurring sweet-kernel almonds while foraging. However, with no clonal propagation technique and prior to the development of grafting, how could they recruit such sweet-kernel specimens to their early orchards? The answer we offer is related to the mode of inheritance of the toxicity trait of almond. It appears that the

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sweet-kernel trait is dominant over the wild type (bitter, toxic) phenotype. Hence, it follows that a naturally occurring individual tree that carries a mutation in the toxicity genes (which confers sweet kernels) will yield at least 50% non-toxic progeny. That is, at least 50% of trees grown from the seeds that mature on a sweet-kernel individual are expected to bear sweet kernels, regardless of pollination by adjacent bitter almond trees. This is in sharp contrast to the observation that most of the sexual progeny of a good olive, date palm, fig or grape vine are likely to have undesirable fruit traits. Hence, the Neolithic fruit tree domesticators did not have to wait for the (supposedly Roman) invention of grafting to maintain sweet-kernel almond cultivars. While the differences between wild and domesticated trees are usually expressed in the qualities of the fruit, the soft fruit tissues are typically not preserved in the archaeological record. How, then, do we determine whether remains originated in domesticated cultivars or wild trees? The answer to this question is complex and leads to occasional controversy. For example, it has been claimed that the remains of seedless (parthenocarpic) PPNA figs identified at Gilgal I (see Box 8 Netiv Hagdud and Gilgal I, p. 45) and dated to ca. 11,400–11,200 calibrated years ago attest to fig domestication at a date that precedes the arrival of domesticated wheat and barley into the region. The claim is based on the argument that trees that do not produce fertile seeds (due to some mutation) cannot survive independently in nature. This brought Professor M. Kislev and his students to conclude that the site’s inhabitants had identified and reproduced such trees in plantations for the purpose of fig harvesting. However, some fig trees produce succulent, edible figs even without pollination. These are the early ripening fruit that mature in spring prior to the main fruit season or those that grow in the autumn following the main fruit season. This fact offers a simple alternative explanation to the seedless fig findings of Gilgal I that is not necessarily related to fig domestication or to systematic, institutionalized plantation farming at the site. We can track the changes in frequency of certain plant species that produce large quantities of pollen for the purpose of pollination by wind; the olive is a good example of this. The airborne pollen released by these species leaves an imprint in layered sediments at the bottom of lakes such as the Sea of Galilee and Ḥula Lake in Israel or Lake Van (Van Gölü) in eastern Turkey. Microscopic inspection of the layered matter in lake sediments allows for the study of local plant composition based on submerged pollen types from different periods. The Pottery Neolithic period (dated roughly 8,500–6,500 years ago) in general shows a significant rise in the incidence of olive pollen in the region. An equivalent finding of the same period is evidence from submerged Pottery Neolithic sites at ‘Atlit Bay (on the coastal plain of Israel, south of the city of Haifa) of installations used to produce oil from olives as well as olive waste (pomace) produced through oil extraction (see Box 13 ‘Atlit Yam, p. 71).2 In addition, the Mediterranean Basin boasts wild populations of olive, grape vine, fig, pomegranate and date palm (the latter restricted to the southern desert region). It is plausible that cross-pollination has been taking place between wild and domesticated

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trees since the domestication of these trees by farmers millennia ago. It is likely that during the period close to domestication, when orchards and plantations were still few and wild populations were massive and opulent, pollination, and accordingly the direction of ‘gene flow’, was primarily from the wild to the domesticated trees. In contrast, in the modern age, with wild populations being small and isolated, and commercial domesticated olive, fig and date palm plantations being expansive, the gene flow is primarily from the domesticated gene pools to the wild. Therefore, probably many of the wild olive populations currently identified, for example, on Israel’s Carmel Ridge or in the mountainous Moroccan region of Riff, based on their small, low-fat fruit or their inferior fertility, and the fact that they grow in rocky areas that bear no evidence of early cultivation, carry a genetic pattern that was formed, to some extent, through pollen that originated in domesticated olives. In other words, in the olive populations that we currently consider to be genuinely wild, we might also find gene variants that originated in domesticated olive cultivars that grow in adjacent olive groves such as those near Druse villages on Israel’s Carmel Ridge. Similarly, wild olives that grow in Morocco, Spain, France and the Balkans may carry the genetic signature of local olive cultivars. It is therefore not surprising that the DNA profile of ‘wild’ and domesticated Mediterranean olive cultivars drove the claim that the olive was independently domesticated in different regions within the Mediterranean Basin. It is of course possible that following the spread of the Near Eastern grain crops and farming societies, farmers from Morocco, Spain or Greece have adopted local wild olive types with good fruit qualities for cultivation. Such local strains had the advantage of being adapted to the ecological conditions in these regions. As such, local olive types might have had better (or more stable) yields compared with introduced east Mediterranean trees. In effect, the above scenario depicts secondary domestication centres, as discussed in Chapter 10. However, the possibility of genetic contamination (via cross-pollination gene flow) cannot be ignored. This genetic contamination would have occurred when wild trees pollinated the early domesticated olive cultivars brought to the western parts of the Mediterranean from the east, where the earliest evidence of oil production installations and extensive use of olives were found. This potential contamination among current-day wild populations, bearing genes that originated in domesticated olive cultivars, hinders the possibility of identifying the geographic origin of olive domestication employing technologies aimed at tracing ‘genetic fingerprints’ as successfully implemented with self-pollinating crops (such as einkorn and emmer wheat). This fact highlights the importance of any archaeological evidence and its dating. Discounting our reservations about DNA profile analyses in trees due to contamination, we note a recent research investigation of genetic relations based on DNA sequences of domesticated and wild olives from the entire Mediterranean Basin combined with molecular dating of the time during which these sequences changed. Based on the findings, it has been suggested that olive was first domesticated in the northern Levant, that is, in the area in which the founder grain crops discussed in this

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Table 11.2 Main differences between fruit trees and grain crops domesticated in the Near East Characteristic

Fruit trees

Grain crops

Time of domestication

End of Neolithic (PN) and during Chalcolithic period

Pre-Pottery Neolithic B

Life cycle and longevity

Perennial, up to hundreds of years

Annual, single growth season

Reproductive system

Cross-pollination

Self-pollination

Number of selection cycles since domestication

Relatively few

Many thousands

Agro-ecological range of domesticated cultivars

Similar to distribution area of wild progenitors (Mediterranean climate or similar)

Quite extensive, from tropical to temperate zones

book were first domesticated. Similarly, recent genetic analyses suggest a north Levantine origin of the domesticated almond. Two additional important aspects of tree domestication concern the length of tree life cycles and the fact that fruit tree cultivars represent rare genetic combinations (Table 11.2). The perennial nature of trees combined with the fact that fruit tree cultivars represent rare genetic combinations that were expressed in seedlings (or as mutations in plantations) led farmers to adopt these individuals due to their exceptional fruit traits, and allowed ample time for vegetative reproduction over many human generations. At the same time, because of the long life of many fruit trees (e.g., several hundreds of years in the case of olive trees), only very few sexual reproduction and selection cycles had been completed to date, since the domestication of these trees. It must also be remembered that the asexually propagated olives or date palms from shoots and offshoots, as well as of figs, grape vines or pomegranates reproduced from shoot cuttings, represent genetic lineages (clones of a domesticated cultivar) that could persist (nearly) unchanged for hundreds or even thousands of years, with the exception of rare somatic mutations (see Glossary, Botany, Ecology and Agronomy, Somatic tissues; Genetics, Mutation). This is, of course, in full contrast to grain crops that are sown anew annually, which have therefore undergone thousands of reproductive cycles since their domestication in the Near East. During each of these cycles, mutations and random mating (cross-pollination, if even on a very limited scale) could have occurred, creating new genetic combinations that would have offered farmers a wide variation among the annual crops from which to select new stocks for cultivation. We emphasize that even if mutations occurred only rarely (one in a million or even less), or even if cross-pollination occurred in percentage fractions, over the evolutionary time frame of several millennia, significant genetic variation would have been accumulated. Among self-pollinating annuals, this genetic variation is rapidly exposed, and

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therefore the incidence of individuals exhibiting advantageous traits would rise rather quickly within the population. As noted by Daniel Zohary and Pinhas Spiegel-Roy, this fundamental difference between the limited number of selection cycles completed in fruit trees compared to the many thousands of selection cycles completed in annuals since their domestication is also reflected in the ecological fitness profile of these fruit trees. Fruit trees originating in the Near East grow in commercial plantations only in climatic zones that are characterized by a climate similar to that of the Levant. In other words, even after their import to other continents, such as the Americas or Australia, date palms, olives, figs and pomegranates are grown mostly in eco-geographic regions that resemble the Mediterranean conditions (such as south and south-west Australia or California in America). The grape vine seems to have originated in a colder climate (the border of Turkey, Armenia and Georgia), and it is more resistant to low temperatures compared to the date palm; it is therefore an exception to the rule and may be successfully grown in higher latitudes. In contrast, all annual grain crops originating in the Near East grow successfully in many areas, from near-tropical regions (Ethiopian highlands) to Scandinavia and the Great Plains in the USA and Canada. Since the sowing rate of annual crops is high (field stand density of 50–500 plants per sq m), and since their seeds result from sexual reproduction cycles, the chances of exposing desired traits are infinitely greater than the chances of exposing such mutations in fruit trees, which are planted sparsely at one specimen or less per 10 sq m and in which only a few sexual reproduction cycles have been completed because they, for the most part, underwent vegetative reproduction since domestication. In the context of the ecological fitness of fruit trees, an exceptional and important example is the successful endeavour of Abba Stein, a member of kibbutz ‘Ein Shemer in Israel, who developed sub-tropical apple cultivars (see Box 18 The sub-tropical apple, p. 175). In the next chapter, we discuss the relationship between ecological amplitude (the range of habitats in which a plant is found) of domesticated crops and the way in which the rarest genetic combinations that occur among domesticated crop populations may confer a selective advantage that will therefore become established. This occurs when farmers bring crops into new regions under growing conditions that differ from those found at the crops’ place of origin. While the apple originates in central Asia rather than the Near East, we opt to discuss the work of Abba Stein due to its great significance, both agronomically and economically, and the lesson that may be derived from it. Apart from the genetic and archaeological aspects reviewed here, the domestication of fruit trees involves additional interesting aspects. One concerns the issue of agricultural land designation. Unlike plots that are sown for the purpose of annual crops, such as barley or lentil, the designation for fruit trees is a priori made for the long-term. Ancient olive orchards in Israel located in the Judean Mountains, Samaria, Galilee and the Carmel are well-known in the local scenery. Fig orchards and grape vineyards in the Hebron Mountains and Galilee are also maintained over long periods of time, and likewise all around the Mediterranean Basin. The phenomenon of cultivar replacement

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Box 18 The sub-tropical apple Different fruit trees have different ecological requirements, and in this way they are no different from annual crops. Because of the genetic composition of the population of origin of the fruit trees, because of the frequency of cross-pollination in these species and because of the limited number of sexual reproductive cycles that were completed since fruit trees were first domesticated, the ecological range of fruit trees is quite similar to that of their wild progenitors. For example, sub-tropical species such as mango, banana and avocado are commonly grown across the Mediterranean Basin. However, due to occasional extreme winter temperatures, their plantations may be severely damaged and even destroyed. The apple is a deciduous tree and, like the apricot, pistachio, cherry, plum and peach, originates in central Asia. In that region, it sheds its leaves towards the cold winter and renews growth after snows melt in the spring. The apple blossoms with the emergence of its renewed growth, and the fruit ripens during the summer. Flower buds of the next season are formed during the long summer days, but they remain dormant, protected by scales. As autumn temperatures begin to drop, leaves are shed, and the tree enters a dormant period. With the progression of the cold season, a complex biochemical process unfolds within cells of the flower buds of the apple (as in all deciduous trees), which effectively ‘counts’ the accumulated ‘cold units’ (agronomists usually denote this as the average degrees Celsius  number of days) to which the tree has been exposed since dormancy began. In nature, during the evolution of the apple (and other deciduous species), genetic stocks were selected in which dormancy breaks at a time when temperatures are unlikely to drop, that is, during the spring. When attempting to grow fruit trees that have high cold requirements in regions where

winter is not sufficiently cold, their blossom is less abundant and asynchronous (between buds, branches and individual trees in the orchard). Under such circumstances, only a few flowers will blossom in the optimal season, hence creating inferior fruiting potential. For this reason, it is difficult in Israel to produce a profitable yield of pistachio or apple in trees grown on the coastal plain. How, then, did Abba Stein successfully develop an apple cultivar well adapted to regions characterized by moderate winters? In his work on his kibbutz’s plantations, in the years before the establishment of the State of Israel (in the 1940s), Stein noticed that European deciduous fruit tree cultivars brought to the country had not blossomed to a desirable extent, rendering their yield unprofitable. Realizing the solution must be genetic, he searched for a source of the desirable trait. Stein scanned abandoned orchards across the country in search of early flowering apple trees. Under traditional farming, seedlings can be found at the edges of fields or orchards that are offspring of sexual rather than vegetative, asexual reproduction. Most of these seedlings never reach maturity, and if they do happen to produce a yield, their fruits are often small or of poor quality, and their overall yield is marginal. As such, it is highly likely that these trees end their lives as raw material used for construction or combustion. However, if such a seedling bears an advantageous genetic combination, it may very well be protected from felling or burning. A mutation that produces early apple blossoms in a region where the winter is very cold, such as the Caucasus or central Asia, prevents the seedling from producing offspring, causing farmers to discard such mutants. In contrast, the same trait uncovered in a country characterized by a mild winter, such as Israel, could be advantageous as it allows farmers to cultivate apple orchards that will blossom profusely even

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with the accumulation of only a few cold units. Stein understood this and had identified several such trees near Jerusalem that blossomed early; he sampled some propagation material from those trees and planted them at his kibbutz. Later he crossbred those early flowering stocks with commercial apple cultivars. From among the offspring, he was able to isolate the cultivar that he named after his daughter, ‘Anna. The ‘Anna apple (Figure 11.3) was successfully embraced

worldwide, opening agricultural and economic opportunities to farmers in tropical countries such as Venezuela, Ecuador and Florida, where it was previously impossible to grow apples. Stein won many awards and much recognition for his work from farmers and scientists worldwide. Interested readers will find further details in the book, ‘Anna’s Father (Abba): The Work of Abba Stein in Breeding of Deciduous Fruit Varieties in Israel (in Hebrew), published by the Fruit Board of Israel, 1995.

Figure 11.3 ‘Anna apples – courtesy of Y. Doron. Colour versions of these images can be found at www.cambridge.org/abbo-gopher.

(by re-grafting) or of uprooting and replanting of fruit trees is a modern phenomenon that is rarely observed under traditional farming. Designating an area for any specific tree species throughout the lifespan of the farmer and perhaps future generations is no trivial issue. Furthermore, under traditional farming, which typically does not rely on intensive irrigation and fertilization, large planting intervals must be maintained between trees to allow each individual tree to grow a broad root system without competing with its neighbours. As a small number of individuals can be grown in each area unit under such conditions, securing a profitable yield requires additional land. An interesting feature of fruit trees we have yet to mention is their long juvenile period (a period during which the plant does not develop reproductive organs (flowers) and during which its branches may be thorny; see Glossary, Botany, Ecology and Agronomy, Juvenile period). In other words, during the growth period immediately following planting, the tree does not yield produce but merely grows in diameter and deepens its roots and thus may not require watering between winters. Due to this

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extensive growth, during this stage the tree develops very slowly. Thus, even without strict observance of the biblical prohibition of ‘orlah-fruit (uncircumcised fruit; Leviticus 19:23, KJB) for the first three years after planting and the provision that the fourth year’s yield be put aside as a donation for the Temple priests, it is unlikely that trees under a traditional farming regime would start yielding fruit in their first four years. Juvenility is most pronounced in fruit tree seedlings, but also in date palm offshoots and trees that develop from olive shoots. Juvenility is shorter and less of an issue in the grape vine, fig and pomegranate, which are propagated by branch cuttings, since these are typically removed from mature tree parts – a fact that further emphasizes the agronomic advantages of the first fruit trees that had been domesticated in the Near East. Given land designation considerations and the long waiting period until fruit is produced, we may deduce that the domestication of fruit trees necessitated a different approach to farm management as well as a different worldview from those involved in the management of grain crops. In contrast to the annual grains scenario, where the farmer sows in winter with the expectation of harvesting the yield in early summer, with trees, three to five years may elapse between planting and fruiting; in some cases, plantations might even yield only to the farmer’s next generation, as recounted in the Talmud. We thus introduce the borrowed term of delayed return to the current discussion – in contrast to the hunter-gatherers’ life-ways where accumulation was minimal and resources were immediately consumed, the agriculture-based economy relies on delayed returns. Work, time and resources must be invested before the grain yield can be harvested after the growing season and seeds may be consumed, whereas tree cultivation represents a different degree of delayed returns spanning several years, and perhaps even generations. Clearly, such subsistence requires an adequate cultural infrastructure alongside a suitable social organization and an economic regime that, together, facilitate the handling of such long delays in yield returns. This brings us to another important point, which we raised when discussing the way in which domestication candidates were chosen and whether plants were domesticated randomly or purposefully following educated decisions based on accumulated knowledge. During the long, fruitless juvenile period of trees, their appearance somewhat diverges from that of mature trees. Typically, leaves of juvenile trees are shaped differently from those of mature trees. In some species, juvenile trees bear thorns, probably in protection from plant-eating animals until they are established. This implies that growing fruit trees requires biological understanding and familiarity with the different growth phases of relevant species as well as the understanding that the juvenile manifestation (seemingly undesirable with its thorns, lack of fruit and later paucity of fruit) will eventually die, to be replaced with the desired qualities of the mature tree. As noted earlier, such an understanding negates arguments in favour of a random domestication process that advocates that domesticated cultivars were adopted from plants that happened to grow in man-made niches, along trails, or on riverbanks. Furthermore, the fruit of most of the trees discussed in this chapter can be eaten immediately upon

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ripening, including the pomegranate, the fig, the grape and the date; they can also be preserved in the form of raisins, wine, dried dates and dried figs. Ripe olives are bitter, and are consumed following processing, as preserved olives or olive oil, further emphasizing that Neolithic plant domesticators were not simpleton consumers of nature. We thus conclude this chapter by reiterating our main argument through the prism of tree domestication. To the best of our understanding, fruit tree domestication required extensive knowledge of tree biology, familiarity with the environment, longterm planning and manipulation skills. The knowledge and skills involved in fruit tree domestication thus further support our view that plant domestication could not have occurred randomly. K E Y PO IN TS A ND BE Y O ND

• The olive, grape, date, fig and pomegranate are the first fruits to have been domesticated in the Near East; these make up the founder group. • These species reproduce in the wild through cross-pollination, but they may be easily propagated vegetatively (asexually). The ease of asexual reproduction in these species allowed farmers to preserve desirable mutants as chosen cultivars. • Cross-pollinating mechanisms hinder the identification of the wild populations from which the domesticated stocks originated. This notwithstanding, for the almond and olive, a location of origin near the core area of grain crops has been suggested. Genetic studies suggest that the date palm most probably originated in remote desert oases in Oman, south-eastern Arabia. • Plantation (and orchard) farming necessitates long-term planning, land designation for several generations and investment in installations by which to process produce; it thus required great patience and the ability to delay returns. • Prehistoric farmers developed technologies by which fresh fruit could be processed and preserved; some of these successfully serve us to date (drying, olive pressing and wine production). • Similar to annual species, it is difficult to conceive fruit tree domestication as the outcome of an unconscious dynamic or a random process as it necessitated the correct choice of tree species and a thorough understanding of their biology, including vegetative propagation mechanisms, reproductive processes that determine fruit set, growth patterns, husbandry methods (such as pruning) and their establishment processes. • As such, tree domestication represents a complex and well-perfected process, seemingly easily executed in terms of actual reproduction, but highly complex in terms of perception. It is therefore unsurprising that the process occurred subsequent to the domestication of annuals and after years during which agricultural experience and agronomic knowledge were accumulated.

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P L A N T E V O L U T I O N U N D E R DO M E S T I C A T I O N

In this chapter, we discuss the evolution of domesticated plants in the periods subsequent to the episode of domestication. We unfold the influences of domestication on the genetic variability of different crops and trace the evolutionary forces that promote genetic diversification and its preservation over the years. At the same time, we also identify forces that restrict genetic diversity among the different domesticated crops. We end this chapter by reviewing the impact of these processes on current and future plant breeding and endeavours for crop improvement. As discussed earlier, wild plant populations are dynamic entities in which different genetic types (genotypes) coexist. The incidence of these genotypes varies over the years according to the environmental conditions that characterize each growing season. When seeds are sampled from wild populations for the purpose of domestication, as expected, only a small portion of the genetic diversity found within wild populations (over the entire distribution area of the relevant species) is represented in this small domestication candidate sample. This phenomenon is termed a genetic bottleneck or genetic drift (also known as the founder effect; see Glossary, Genetics, Genetic drift). Randomly sampling individual specimens from a large population for the purpose of domestication drives two phenomena that have importance for the topic discussed here. The first is the reduction of genetic diversity among domesticated species compared to that which is found among the populations of the wild progenitor. The second is the outcome of this ancient process of sampling for the purpose of domestication: in domesticated populations (landraces) that are established based on such sampling, the prevalence of certain alleles is considerably higher compared to the population of origin, so that the genetic profile

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of the new population is ‘biased’ or ‘drifts’ towards an evolutionary trajectory that may be different from the one that unfolds within the original population. An excellent example is the prevalence of genes that facilitate free germination in legumes or the prevalence of non-shattering spikes in cereals. It must nevertheless be remembered that in addition to those genes that govern desired traits – which probably guided domesticators in their educated choice – the prevalence of other alleles is also increased or decreased as the result of the selective sampling process. Thus, domestication involves the choice of desired traits while also fostering random changes in the genetic make-up of the new (landrace) population in chromosomal regions that bear genes controlling traits that are unrelated to the original reason for selection. If domestication restricts the gene pool and diversity of domesticated plants, how can we account for the vast range of variation found among many crop plants? We list here but a few of the numerous available examples: the broad range of seed weights in legumes such as faba bean, lentil or chickpea; the variability in size and colour of dry bean seeds or fresh bean pods; the diversity of size and colour of fruits such as peaches, plums, apples, pears, melons, watermelons, zucchinis, pumpkins, cucumbers or aubergines; the broad range of grape cultivars, known by any wine drinker; and perhaps the most extreme example is that of the cabbage group (Brassica crops). Many readers are probably unaware that cabbage, cauliflower, broccoli, Brussel sprouts, kohlrabi and kale are botanical cultivars of the same plant species, which may be easily crossbred to produce fertile offspring. How then can we settle this diversity of taste, shapes and colours with the phenomenon described earlier, which has been documented in many different domesticated species, regarding the narrow genetic variability found among crop plants compared to their wild progenitors? Well, among domesticated plant populations, several processes take place that increase genetic diversity while other processes restrict it. The final observable outcome reflects the balance between these two negating processes. It is important to remember that farmers continue to make selections among the domesticated species to promote desired traits even after their initial selection. For example, they are likely to remove undesirable types that may have an unpleasant taste or whose seed size or colour are undesirable, or which are otherwise unwelcomed. Additionally, the growing environment in the farmer’s field acts as a natural filter, affecting selection within the domesticated population. For example, seeds that germinate slowly might not develop into yielding plants because of the shade cast upon them by neighbours that successfully germinated earlier in the season. If this scenario were to recur, late germinators would slowly be removed from the population. Similarly, early blossoming individuals that are injured during cold episodes at the end of winter, or late bloomers whose seeds are unable to ripen before the soil dries in the summer, will be unable to produce offspring, or their yield will be so marginal that their prevalence among the population will be significantly reduced after only a few growing seasons. Pests or disease-causing agents (fungi, bacteria or viruses) will damage the yield in susceptible individuals, leaving

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mostly resistant or partially resistant plants in the field. This is another dynamic process, as pest populations and pathogens are dynamic entities in themselves, occasionally producing new types of virulence while older virulence variants may disappear. The reason for the occurrence of new pathogens and pest strains is rather simple. If all susceptible host individuals will be destroyed by the pathogen leaving no seeds for the next season, how could the pathogen be able to sustain itself among the remaining resistant types? This in itself grants a selective advantage to the emergence of new virulence genes by pests or pathogens. Otherwise, following total destruction of the host population, the pathogen may also become extinct. Thus, in nature, pests and disease agents do not completely destroy their host populations, but rather cause only partial damage, thereby allowing the hosts the opportunity to produce at least some offspring. Additionally, the high propagation coefficient of pests and disease agents (i.e., the number of progeny per generation and number of generations per growth season) ensures that in any generation some random mutation will occur in genes that determines their adaptation and/or virulence profile. Some of these mutants would be able to attack plants that showed resistance in an earlier generation, while others may cause the pathogen to be less virulent. Likewise, mutations occur in hosts’ genes that may confer them resistance (or susceptibility) to pest or pathogen strains. In fact, such selection processes, which consistently transpire over multiple generations, shape domesticated populations and adapt them to the needs of farmers and the husbandry regime of agronomic operations that take place on the farm. This is the manner by which all traditional cultivars (landraces; see Glossary, Botany, Ecology and Agronomy, Traditional cultivar (landrace)) have been established over millennia in different farming societies worldwide. Indeed, investigating crop cultivars in regions where traditional farming communities still operate (such as rice growers in Thailand’s highlands or bean growers in Mexico) shows that genetic differences can be identified between rice cultivars from different villages. Furthermore, genetic differences have also been recorded within the rice populations that neighbouring farmers from a single village grow in their fields. In this manner, by selectively choosing sowing resources for the next season during the harvest, local adaptations are shaped and desirable genotypes are preserved from one generation to the next. Wheat was domesticated in south-eastern Turkey from where it spread to many different world regions. In all likelihood, a seed sample from Turkey will successfully grow in the Balkans, Italy, the Iberian Peninsula and most other regions of the Mediterranean Basin, due to the similarity in latitude, which dictates a similar climate and hence a similar seasonal profile. This, however, is not the case in the Ethiopian highlands, which is closer to the equator and, while exhibiting a different day length and rain regime during the growing season, is nevertheless successfully able to host the Near Eastern founder crops. Due to its proximity to the equator, seasonal variation in day length is minor, and, during most of the year, days are nearly twelve hours long. In Chapter 8, we mentioned that the length of day during the growth season

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determines the developmental pattern of domesticated Near Eastern cool-season crops, emphasizing the differences between cereals and legumes. The short winter days dictate vegetative development of cereals, and reproduction can begin only when days grow longer at the end of winter and early in the spring and induce spike and flower development. The near-equal length of day in Ethiopia throughout the year does not provide the appropriate signal to induce timely flowering in plants originating in higher latitudes. How, then, did early farmers successfully adapt wheat, barley, lentil, faba bean and chickpea, originating in the northern, colder area of south-eastern Turkey, to the climate of the Ethiopian highlands? To answer this question, we must review the factors that promote genetic variability under domestication. The first factor involves mutations (Glossary, Genetics, Mutation). The above notwithstanding, some mutations renew the functionality of genes that were debilitated sometime earlier in the evolutionary past of the specific individual (also known as atavism). It is commonly assumed that mutations occur at a rate of approximately one in every million or more replications of the genetic material (DNA). Most mutations are harmful or even lethal so that if they occur, they are not passed on to future generations. However, a minor fraction of mutations could be beneficial, such as a mutation that confers protection against a new type of virulence exhibited by pests or disease agents, high temperatures or water shortages. These mutations regularly appear within traditional crop populations but are quickly lost merely because most of the seed yield is consumed by farmers, leaving only a small portion of the yield to be preserved for the next sowing season. However, should these mutants emerge during a plague or an attack of some harmful agent, their presence is exposed due to the resistance they confer, and they would be preserved by the farmer. Similarly, a mutation that enables flowering with no regard to the length of day, while being meaningless to a Mediterranean population, would be highly advantageous (adaptive) in regions where days are relatively short. For example, if such a day length insensitive genotype was present in a sack of seeds transported to Ethiopia, it is likely that plants of this stock would be the only ones that would yield seeds in the Ethiopian highlands. Thus, acclimating Mediterranean crops in Ethiopia relies on the fact that random mutations do occur (albeit at a low rate) and that relevant genotypes must have been present in seeds exported to Ethiopia in antiquity. Had growth failed, it could be assumed that Ethiopian farmers would have relinquished any further endeavours after just a few futile attempts. Similarly, there is no reason to raise tropical plants such as mango, coffee or banana in the British Isles or north-eastern United States where they are likely to fail year after year due to low winter temperatures. Can a rare genetic event indeed influence the evolution of a domesticated species? Considering a sowing rate of fifty to 100 wheat or barley seeds per square metre under traditional farming, and given fifty mature seeds yielded by each plant by the end of the season, a simple calculation shows that a single hectare of cereal crops would yield over twenty-five million seeds. Given one in a million random mutants, we would expect to

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see more than twenty mutants in any chromosomal region (including genes) in each hectare of wheat or barley. Even if most mutations are harmful and seeds bearing them would not germinate, over the course of several years, with cultivated fields growing in volume, an increasing number of mutants would be exposed to farmers. This genetic diversity (created spontaneously and at a low rate) is typically latent and exposed under special circumstances such as the transfer in ancient times of Near Eastern crops to Ethiopia or during epidemics, which disclose resistant individuals within a susceptible population. Mutations are also the force that creates genetic variability in fruit trees, but with one fundamental difference: the location of the mutant gene. A mutation occurring in leaf cells of barley or wheat will be lost once the plant matures and then withers. In contrast, a mutation occurring in sex cells of wheat or barley and passed on to the DNA of the egg or pollen cell has a chance to be passed on to offspring if the mutant seed is not consumed. Its survivability increases if the mutation is harmless, and certainly if it is beneficial. However, as noted in Chapter 11, propagation in fruit trees does not rely on seeds but rather on vegetative tissues such as buds or branch segments. What then creates and sustains the genetic diversity that we encounter in various familiar fruit species? Unlike annual plants, in trees, only a mutation that occurs in somatic cells (see Glossary, Botany, Ecology and Agronomy, Somatic tissues) may catch the farmer’s attention. Thus, contrary to common belief, the smooth-skinned nectarine with its peach pit was not selected from a cross between a peach and a plum but is rather a hairless peach. Nectarine-bearing branches appearing spontaneously in peach orchards are described by Charles Darwin in his book On the Origin of Species, along with other examples presented to support his arguments regarding the evolution of plants and animals under domestication. These rare phenotypes (traits) require special attention and understanding exercised by the farmer lest the mutation is lost by pruning. Should a mutant fruit-bearing branch exhibiting some unique trait such as colour, size, flavour or scent be inadvertently pruned, the mutation would immediately become obsolete. However, should it be noticed, preserved and grafted onto other trees, this branch will then become the outlet of a new cultivar or variety of the same species, as transpired with the Shamouti orange cultivar (also known as the Jaffa orange), which appeared a few hundred years ago as a mutation on a tree branch in an orange grove near the old Jaffa seaport in Israel. Typically, diversity in form and colour found in domesticated species (such as the cabbage group) is very limited in wild species, in other words, wild cabbage does not possess the great variety of leaf shapes, inflorescences or stems as do the abovementioned domesticated forms. Similarly, the fruits of wild plum or pear trees are not as varied in form and colour as those of domesticated trees. Why, then, do we claim that the genetic diversity of domesticated species is lower than that found among wild populations? Domesticated species indeed exhibit diverse forms, colours, flavours, scents or other traits regarded as useful by humans, and these are therefore a target in

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selection applied by growers and breeders. However, in fact, most of these traits are usually governed by several dozen genes. In contrast, in all the thousands of genes, which were not subjected to selection by growers since domestication, diversity is lower among domesticated fruit cultivars compared to the wild forms because of the founder effect mentioned earlier. And so, immigration to world regions that require some adaptation (such as the change in sensitivity to day length in the transition from high to low latitudes and vice versa) may expose concealed mutations, which enable the crop to become established in the new region. Imagine a sack, or perhaps several sacks, of seeds containing several millions of wheat or lentil seeds brought to Ethiopia from the Near East, or the other way around – imagine a sack of sorghum seeds making its way from sub-Saharan Africa towards Israel. When farmers sow the new seeds, only those individuals that bear the traits that will allow them to develop normally and blossom in the right season will produce any kind of seed yield. Given that mutations are rare events and that beneficial mutations are even rarer, it may be easily understood why the number of individuals fitting the new regions constitutes only a minuscule portion of the seed shipment brought to the new region (as the prevalence of those mutants is, by definition, infinitesimal). In fact, this is the description of a genetic bottleneck event as the entire population with which the growers are left has been derived from the mutant individuals that are used to establish the crop in the new region (and hence the origin of the genetic term, the ‘founder effect’). As with the adaptations that were required in the introduction of Near Eastern crops to Ethiopia, similar adaptations were required to bring lentils to the lower latitudes of India. Indeed, despite the fact that India is one of the largest lentil producers worldwide and although it hosts thousands of traditional lentil cultivars, the genetic diversity among Indian lentils is extremely limited. This is probably the result of the founder effect, which occurred when lentils were first brought to the Deccan Plateau in India several millennia ago. Since plant domestication took place in areas where wild populations grow naturally, cross-hybridization must have occurred subsequent to domestication between wild and domesticated types. Cross-pollination between wild types and plants growing within domesticated fields, even at a low incidence, would have driven the flow of different traits between the two populations, the wild, and the domesticated. Within this process, known as introgression, gene variants flow according to the rate of airborne pollen. Naturally, in the first few millennia following domestication, when wild populations were still massive and larger than domesticated populations, wild plants would have been the ones to enrich the ancient landraces (cultivars) with different alleles and genes (see Chapter 11). In fact, in this manner, wild populations increased the genetic diversity found in ancient cultivars, and, indeed, this is why we perceive traditional landraces to be more diverse. In contrast, currently, with the decline of natural ecosystems and the significant increase in arable land, it is generally assumed that genes flow primarily from domesticated fields to wild populations. Conservationists consider the

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current trend as ‘genetic contamination‘ since the domesticated alleles interfere with the integrity of the genuine wild gene pool. A new growing region has its own climatic profile, soils, weeds and local pests and disease agents as well as common husbandry actions of local farmers, the manner of seed selection for different purposes and more. Thus, following the process described above, genetic diversity that confers adaptation to local conditions and to the needs of farmers and their preferences as well as to regional conditions will be selected and preserved. In particularly remote geographical areas, distance and isolation from the population of origin found in the older growing regions will foster diversity – through mutation and cross-pollination – that is unique to the area and will not flow back to the original gene pool. Indeed, in the Ethiopian highlands a vast variability of wheat, barley, lentil and chickpea have evolved, inspiring the suggestion that Ethiopia was a primary centre of domestication, independent and separate from the Near East (see Chapter 13). This suggestion, however, is implausible as no wild progenitors of these crops grow in Ethiopia, so it would have been impossible to domesticate these species in that location. Similarly, it would have been impossible to domesticate wheat, chickpea and pea in mountainous regions of central Asia, where a broad variety of these crops was recorded, including types that are unique to that area. For the same reason, West Africa is considered a secondary centre of diversity (i.e., the original domestication occurred elsewhere) for groundnuts that were imported by the Portuguese and Spaniards after the discovery of the Americas (the relationship between genetic diversity and estimated location of domestication is discussed in the next chapter). We now turn to the implications of the phenomena described above for modern agriculture and plant breeding today and in the future. Genetically speaking, modern crop varieties are highly uniform compared to traditional cultivars, as growers and seed producers meticulously maintain product uniformity. It is impossible today to register a cultivar (similar to registering a patent) in industrialized countries without proof that the new cultivar is a uniform genetic stock and exhibits traits that distinguish it from other cultivars. The diverse populations of traditional wheat or lentil cultivars would not withstand these criteria. The modern requirement of uniformity among crop cultivars since the early 1900s has considerably restricted the number of crop cultivars grown today in Europe, North and South America, Australia, India and China compared to those grown in the past. Vast expanses spanning millions of hectares in which a single variety of wheat or rice is grown make fertile grounds for the spread of pests and disease agents. This has brought about the intensive use of pesticides, which not only negatively affect the environment and our health, but also heavily tax farmers. The decrease of genetic diversity through domestication-related bottlenecks hinders breeders’ endeavours to increase the yield of current cultivars or confer better quality and durability. The reason is simple: in order to select types that exhibit new traits, genetic variability is required as it is the raw material of selective breeding. Increased yields are currently the result of interactions between agro-technological improvement

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and new genetic combinations that allow the plant to exhaust its potential and overcome its yield-limiting factors. How can crop breeders make progress within the genetic boundaries of the domesticated gene pool? First, even in modern cultivars, and certainly in traditional ones, mutations occur within the extant genetic matter. Second, each year, breeders and researchers create thousands of new hybrid combinations and examine tens of thousands of their offspring; this is akin to buying as many tickets as possible for the lottery. Since this is a numbers game, the more combinations investigated, the better the chances of finding the right combination. Nevertheless, despite modern genetic knowledge and the vast selection endeavours, it is clearly difficult to maintain a desirable rate of yield increase in crop plants (compared with the post-World War II decades) or develop resistances to new pests and disease agents that are transferred cross-continentally by humans. This is the place to reiterate the importance of the wild progenitor populations of domesticated crops. One of the first biologists to understand this was the agronomist Aaron Aaronsohn, who discovered wild emmer wheat near a vineyard in Rosh Pina in Israel in 1906. Aaronsohn was the first to see wild emmer in its natural habitat, mostly in niches characterized by poor soil. With his keen senses, Aaronsohn understood that this wild wheat had become genetically adapted for rough environmental conditions – otherwise it would not have survived as a wild species in the semi-arid environment of the eastern Mediterranean Basin. In a review, published by the US Department of Agriculture in 1910, Aaronson predicted that wild emmer wheat could be used to develop wheat cultivars with the purpose of increasing its yield under stress. Indeed, in recent decades, scientists have successfully transferred genes from wild emmer wheat to modern wheat cultivars that confer resistance to various diseases. In the early 2000s, another research group, led by Professor Tzion Fahima, at the University of Haifa successfully isolated one of the genes responsible for the high protein content found in wild emmer grains. This is a first step in our ability to increase the protein content of domesticated wheat grains. These findings are an important reminder to the public regarding the importance, for future food production worldwide, of wild plant populations that thrive in natural ecosystems and the imperative to protect such areas from development and construction endeavours that are typically aimed at producing shortterm gains for real-estate entrepreneurs.

K E Y PO IN TS A ND BE Y O ND

• Plant domestication involves the adoption of a small segment out of a broad range of genetic diversity found among wild progenitors, creating a genetic bottleneck. • Despite these genetic bottlenecks, a broad range of traits is found among many domesticated plant species, converged at target organs (such as different lettuce or cauliflower heads, or the colours, flavours and scents of fruit and tuber crops).

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• Random mutations are the primary raw material from which new traits emerge in both wild and domesticated populations. Introducing plants to new growing regions or exposing them to extreme events such as pest or disease outbreaks may expose useful traits, which can then be preserved by farmers. • Modern cultivars are more uniform than traditional cultivars and hence more susceptible to damage incurred by climate, disease or pests. The uniformity of modern cultivars hinders continued breeding endeavours. This fact renders wild populations related to domesticated species an invaluable and irreplaceable genetic reservoir for future crop improvement and for future human generations.

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13

A G LO BAL V IEW O F P LAN T DOMESTIC ATI ON I N O T H E R WO R L D R E G I O N S Asia, Africa and America

Near Eastern plant domestication, which took place over 10,000 years ago, has become an important research topic worldwide, drawing many researchers from diverse disciplines. For us, who live in the area, it is only natural that we focus our attention on it. Plants domesticated in other world regions also contribute important elements to key food packages. In this chapter, we discuss plant domestication centres found in Africa, East Asia and America. The first researcher to notice a global geographic pattern in plant domestication was the Swiss botanist Alphonse de Candolle, who was active at the end of the nineteenth century. He developed a method by which to identify the location of origin of domesticated plants based on archaeological, botanical, paleobotanical (plant fossils), historical and philological information. Underpinning his method was the logical assumption that the joint use of these information sources would point to the area in which the plant was first domesticated. Botanical and paleobotanical evidence attests to the availability of the plant in ancient times, while the earliest dated archaeological evidence of the use of plants and of domesticated remains in conjunction with written testimonies from early historical periods and linguistic evidence in ancient languages point to the cultural origin of traditions involving the use of the plants under investigation. De Candolle’s general conclusion was that domestication origins in the world were unequally distributed; some regions, such as the Near East, India and China, provided many domesticated crops, while others, such as Europe or Australia, offered only a few. A highly important contribution to the understanding of the geography of plant domestication worldwide was made by the Russian botanist and geneticist Nikolai

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Vavilov. From 1916 to 1940, Vavilov travelled extensively to all five inhabited continents, collecting seeds of as many traditional crop cultivars as he could. His interest in these seed samples derived from his recognition that plant breeding and improvement of crop yields in his country required the broadest possible genetic diversity. Vavilov understood that cultivars originating in different world regions that share a climate similar to that found in northern Europe or North America would be successful crops in different areas of Russia with almost no further breeding efforts, while others might contribute desired traits through crosses and selection, as done regularly in plant breeding programmes. Vavilov’s seed collection is to this day considered one of the most important (if not the most important) crop germplasm (seed bank) collections worldwide; it is kept in an institute named for him in St Petersburg. It is said that employees at the institute were so cognizant of the immense significance of this seed collection that they protected the seed storage units with their bodies against hungry citizens during World War II when the city was under Nazi siege. Like de Candolle, Vavilov also noticed an unequal geographic pattern of crops in his journeys. Namely, that some world regions boast a highly diverse range of crop plants while in other areas the genetic diversity found among crops is rather limited. In the study of evolution, a fundamental assumption is that the more time that elapses from the emergence of a new species (or a lineage), the more likely it is that the species will have gained greater genetic diversity, which would be expressed in different traits such as colour, forms or fitness to different habitats, relative to younger lineages or species. Vavilov thus assumed that the area in which the greatest variability would be found was likely the oldest centre in which the crop originated. This implied that it would also be the area where the wild progenitor had been domesticated by early farmers. Based on the geographic distribution of genetic variation in the different crops that he collected on his journeys and which were analysed by staff at the research institute he founded, Vavilov defined eight independent centres of origin worldwide (often termed ‘domestication centres’; see Figure 13.1), listed below. Centres of origin were found in the following world areas (numbers correspond to those in Figure 13.1): 1. China, where different types of millet, buckwheat, soy, adzuki beans, different types of bamboo, aubergine, cucumber, citrus and litchi were developed; 2. India, where rice, sorghum, chickpea, pigeon peas, black-eyed peas, aubergine, cucumber, mango, some citrus, sugarcane and Indian mustard were developed; 2a. Malaya (Indochina), where coix (a grain cereal named Job’s tears in English), different types of bamboo, different types of yam (tuber plants), ginger, banana, durian, coconut and sugarcane were developed; 3. Central Asia, where wheat, rye, chickpea, pea, lentil, faba bean, flax, melon, carrot, radish, onion, garlic, basil, peach, pear, almond, grapes and walnuts were developed;

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Figure 13.1 Map of diversity centres of domesticated crops by Vavilov, adapted by both Jack Harlan and ourselves: (1) China; (2) India; (2a) Malaya (Indochina); (3) Central Asia; (4) Near East; (5) Mediterranean; (6) Ethiopia; (7) Central America; (8) South America; (8a) Chile; (8b) Bolivia-Paraguay.

4. Near East, where einkorn wheat, durum wheat, barley, oat, chickpea, lentil, bitter vetch, fenugreek, alfalfa, clover, sesame, poppy seeds, melon, pumpkin, carrot, cabbage, fig, pomegranate and pistachio were developed (Vavilov did not list here the pea and fruit trees we discussed earlier); 5. Mediterranean, where durum wheat, lentil, chickpea, pea, faba bean, grass-pea (Lathyrus), clover, flax, beet, parsley, celery, dill, lavender and mint were developed; 6. Ethiopia, where durum wheat, teff (Williams lovegrass), millet, chickpea, noog, lentil, pea, faba bean, sesame, Ethiopian mustard and flax were developed; 7. Central America, where maize, beans, cocoa, different types of pumpkin, sweet potato, sweet and hot peppers, cotton, agave, sugar apple (anona), avocado and tobacco were developed; 8. South America, where potato, quinoa, tomato, pumpkin, cotton, Aztec marigold, cocaine plants and tobacco were developed; 8a. Chile, where potato and strawberry were developed; 8b. Bolivia-Paraguay, where cassava, groundnuts (peanuts), cocoa, rubber tree, pineapple and Brazil nuts were developed. A brief analysis of this list exposes several challenges. First, that Vavilov found the need to divide some centres into secondary ones (as in South America) shows that defining a centre of crop origin (a domestication centre) is sometimes very difficult. The second and more essential challenge is the fact that some crops, such as wheat, chickpea and lentil, appear in several centres, with each appearing in over three centres of origin. The 190

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principle is not flawed, and we have mentioned the possibility of a multi-site domestication history as a suggestion that was made following one interpretation of the genetic findings related to olive and barley. However, one of our underpinning assumptions is that any species can be domesticated only in the area in which it grows wild or in close proximity to its natural area of distribution, whereas no evidence was found, for example, in Ethiopia of wild wheat, wild pea or the wild progenitor of chickpea. It is therefore difficult to accept the notion that any one of these crops was domesticated in Ethiopia. Similarly, no evidence is found in the Near East of wild sesame or pumpkin, rendering it unlikely that these crops were domesticated in that area. Likewise, the wild progenitor of chickpea is unknown in India, so that despite the extensive variability and antiquity of the crop in the Indian subcontinent, it is unlikely that chickpea domestication occurred in India. Thus, a few years after the death of Vavilov, who was a prominent authoritative figure in the scientific arena during his lifetime, doubts began to be raised regarding the equivalence he determined between crop variability centres and crop origin centres (see Figure 13.2 for the differences between domestication centres and prominent current grain production areas worldwide). Indeed, currently, variability centres are considered distinct from domestication centres, a fact that does not detract from the importance of Vavilov’s extensive pioneering work. It is also indisputable that the Ethiopian highlands were an important, independent domestication centre, as some crops are typical only of that area, such as teff (Figure 13.3), which is used to make injera (traditional Ethiopian flat bread) and noog, an oil plant of the Compositae (Asteraceae) family. Equally, Central America was an important, independent domestication centre from which maize and beans spread to many areas worldwide. The north-eastern region of the United States is considered the area of origin for sunflower, pumpkin and an additional grain crop (marsh elder or sumpweed) that was used by Native Americans but has since grown extinct, whereas it is widely accepted that the potato, amaranth, quinoa and groundnut originated in South America. Of course, in our view, as detailed throughout this book, there is no doubt regarding the origins of wheat (both einkorn and emmer), barley, lentil, pea, chickpea, bitter vetch and flax, which were found in the Near East, and, more specifically – in south-eastern Turkey and northern Syria. We emphasize that opinions differ regarding the exact number of domestication centres around the world, and some claim that over twenty such centres can be defined. Understandably, proponents of multi-centred domestication will identify more domestication centres than those who advocate centralized domestication for most crops. For example, some experts suggest that barley was domesticated several times, once in the Fertile Crescent, and once thousands of kilometres to the east of it; others suggest that barley was independently domesticated in two centres within the Fertile Crescent, one in the northern Levant and one in the southern Levant. Likewise, there is a claim for independent barley domestication in Morocco. In our view (as discussed in Chapter 9), such claims should not be endorsed unless archaeological findings show evidence of earlier domestication in one of the alternative locations (centres) and the independence – that is, the absence – of cultural contact between the areas. 13 A GLOBAL VIEW OF PLANT DOMESTICATION

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A R

NORTH AMERICA

EU

Eastern USA A T

P

L

Shael

A

A

N

C

A

T

I F

Meso America

I C

I C

Amazonia O

O

West A

C

C A

E A

E N

N

SOUTH AMERICA

The Andes

Ancient plant domestication centres Areas of important currently grown grain crops

Figure 13.2 Map of domestication centres worldwide, following Jared Diamond (2002), redesigned by Itamar Ben-Ezra. A colour version of this map can be found at www.cambridge.org/abbogopher.

Another characteristic of Vavilov’s list of crops assigned to each of his centres of origin is important for our argument on domestication as a conscious, deliberate process. All ancient domestication centres gave rise to diverse crops that together supply carbohydrates, protein and fat in addition to typical vegetables, spice herbs and fibre plants. In Chapter 8, we discussed the importance of the complementary nutritional potential of Near Eastern grain crops and that a diet that relies on wheat and barley (cereals), on the one hand, and lentil, pea and chickpea (legumes), on the other, is balanced. According to Vavilov’s lists, this phenomenon appears to be universal, and it is not unique to the Near East: East Asia is where soy and rice originated, sub-Saharan Africa is the homeland of sorghum and cowpea (black-eyed pea), maize and beans 192

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C T I C

O C E A N

OPE UR

ASIA

The Fertile Crescent China P A C I F I C O C E A N

AFR IC A

Africa

Ethiopia

New Guinea I N D I A N O C E A N

AUSTRALIA

originated in Mesoamerica (Mexico), and potatoes, quinoa and groundnut were adopted as crops in South America – all of which are complementary nutritional combinations. Vegetables and spices were grown in each region for flavour in addition to health, as plants are a source of vitamins and minerals, and some plants also have medicinal qualities. Nutritionists suggest that flavour is our signal by which we distinguish nutrients that are beneficial and healthy to our bodies, and this might have been a physiological mechanism determining the choice of founder crops. This pattern of domestication of nutritionally complementary crops, we believe, further supports the view that domestication was the result of thought and learning, rather than a random process; otherwise, what are the chances that a crop package that was balanced in both agricultural-agronomical aspects as well as nutritionally would have evolved via random incidental occurrences in each culturally independent world region? 13 A GLOBAL VIEW OF PLANT DOMESTICATION

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Figure 13.3 Teff in an Ethiopian field. Notice the safflower (oil crop) – plants that emerged from seeds scattered by the previous crop grown in this field. A colour version of this image can be found at www.cambridge.org/abbo-gopher.

Can we define a common ecological denominator of ancient domestication centres? Apparently, some regions share certain characteristics. The Ethiopian highlands, the Near East and the Oaxaca province in Mexico – the domestication area of maize – are characterized by semi-arid climates that comprise a defined rainy and dry season. Only a few plant types were domesticated in temperate regions such as Europe or North America, whereas in Australia, which has a broad climatic range, including tropical, semi-arid, desert and temperate regions, no plants were domesticated in antiquity (the macadamia nut, which originates in Queensland, was domesticated and developed as a crop in Hawaii after World War I by American agronomists). We may thus conclude that domestication was the outcome of a cultural process rather than climatic conditions or some environmental change. This, of course, further supports our claim regarding deliberate, knowledge-based domestication. What is the significance of defining domestication centres as independent? Vavilov’s list of criteria for defining domestication centres as independent included biological, botanical and agricultural contexts. We propose adding another criterion – a cultural one. Given the notion that domestication was primarily a cultural transformation, it is prudent, in our view, to ascertain that domestications in the locations under

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investigation (within the Near East) were indeed culturally independent. We emphasize that adding the criterion of cultural independence goes a long way beyond a technical definition for a domestication centre. The Levant is a good example of a culturally independent entity. Materials (e.g., obsidian from Turkey and marine shells from the Red Sea) and technological innovations traversed the Levantine interaction sphere prior to plant domestication. Likewise, ideas, worldviews and novel perceptions were exchanged across the entire Fertile Crescent, furnishing the new perceptual landscape that brought about domestication. It is in the context of these intensive relationships that the innovative idea of domestication spread across the region. To demonstrate this point, imagine a Neolithic community in eastern Turkey domesticating wheat, chickpea, lentil and pea. Naturally, this community interacts, to some extent, with adjacent communities with which it trades, and perhaps there are also some blood relations among these communities. It is likely, then, that during the period in which a crop-based economic system was being developed in the area, news about actions taken to grow plants for the purpose of food production spread. If, following the migration of the idea to remote locations, people in those locations began to examine local wild species and adopt them as agricultural crops, then from a plant genetic perspective, this domestication episode was independent of parallel domestications in the source area (south-eastern Turkey) or other parts of the region. But can we say that this development was culturally independent? Certainly not. Archaeological criteria determine the flow of cultural influence, and if this is the case with domestication, then the dating of archaeobotanical remains, which attest to the first domesticated plants in the affected area, should be later than dates of such remains at the driving centre (which served as the source of cultural innovation). To date, no evidence supports the cultural independence of the many hypothetical domestication centres around the world mentioned earlier in this chapter, whereas only little botanical and genetic evidence supports such a large number of domestication centres. We of course accept a global convergent (independent) model for plant domestication centres, yet we do not accept the ‘inflation’ in the numbers of such centres. It is important to clarify at this point that the Near East in general, and the Levant in particular, are among the most studied regions in these aspects of the archaeology, botany, ecology and genetics of wild progenitors of domesticated crops, thereby allowing us to probe deeply questions pertaining to plant domestication in this region. Lacking such an extensive array of knowledge in other world regions and due to the limitations of our own knowledge regarding other regions, we have for the most part refrained from discussing domestication centres on other continents (and their independence thereof ). Vavilov’s work is important not only for the study of the history of domestication research but also with respect to modern plant breeding. As shown in Chapter 12, where we discussed the evolution of plants under domestication, the genetic diversity of domesticated species is limited compared to that of wild types. Furthermore, since the end of World War II, the massive industrialization of many world regions and the

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Figure 13.4 Mixed field crop: genetic variability under traditional farming, Nagorno-Karabakh, Summer 2012. Note the variation in shape and colour of wheat spikes as well as differences in maturity levels of spikes. A colour version of this image can be found at www.cambridge .org/abbo-gopher.

opening of new transportation routes in remote areas, traditional cultivars have been replaced with high-yielding modern cultivars that were bred in research institutes and seed companies. In some remote areas, this process is still unfolding; in Armenia, for example, traditional farming is still prevalent in certain regions but cereal crop fields host mainly modern cultivars. At the same time, in other regions in Armenia for example, cereal fields are non-uniform, usually hosting a blend of traditional landraces (tall) and semi-dwarf cultivars (some lacking awns), which probably arrived in the region with international aid following the destructive earthquake of 1988 (Figure 13.4). These fields also typically host wild rye as a weed, akin to the wild barley that infests cereal fields in Israel. The traditional husbandry (such as the limited use of herbicides), therefore, often creates a confusing situation with respect to which is the genuine crop and which is the weed – the rye or the wheat? In many world regions, the transition from traditional to modern cultivars climaxed at the end of the 1960s and early 1970s and was termed the green revolution. High-yielding wheat cultivars developed by the Rockefeller Foundation in Mexico and rice cultivars

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developed by the International Rice Research Institute in the Philippines transformed many countries that were traditionally grain importers to exporters. Within just a few years, India, Mexico and Pakistan, which had been dependent on grain import to feed their populations, became not only self-reliant, but exporters as well. The traditional, tall wheat and rice landraces were susceptible to lodging upon high nitrogen fertilization rates, a fact that severely restricted crop yields. However, the new wheat and rice cultivars were shorter, thereby permitting farmers to apply high nitrogen fertilizer levels to their fields. The higher fertilization levels, and the fact that the plants were bred to yield more grain instead of tall culms (i.e., the cereals’ stem), increased grain yields considerably. Naturally, where the new seeds were available for sale, farmers neglected their traditional crops and started raising the new ones. At first, seed inventories of the new cultivars were few and demand was high, to the point where the Indian government placed armed guards in seed production fields of the new cultivars in an attempt to deter thieves. However, the continued population growth in Asia and Africa in the intervening decades raised opinions that the process has exhausted itself and that we would require yet another scientific and agricultural breakthrough to ensure food for all. The extreme food price shifts since 2006, which affect us all, should be considered in this context. The green revolution was a dramatic event that echoed throughout the world and won the Nobel Prize for Peace for the American wheat scientist Dr Norman Borlaug for his contribution to preventing world hunger. The process was also dramatic from a genetic perspective, as tens of thousands of unique cultivars that had been preserved from one generation to the next under traditional farming throughout Asia, Africa, Europe and America were lost forever as they were abandoned in favour of modern high-yielding cultivars. As argued in the previous chapter and the opening of the current chapter, genetic variability is the vital raw material from which new cultivars can be developed. Without genetic variability, no breeding programme can be successful. Thus, the loss of tens of thousands of domesticated cultivars drew the attention of agronomists and geneticists, who began to collect seed material in an attempt to conserve it. The outcome of this activity is an international network of gene banks, established in many countries, as well as the Norwegian seed storage facility in Spitsbergen, where seed samples from across the globe are stored in permafrost conditions, as a modern Noah’s Ark aimed at saving domesticated plant seeds and preserving them. Preserving such quantities of viable seed material involves many challenges, as it is a costly process that requires periodical propagation to renew seed viability. Another challenge is to preserve species that rely on asexual reproduction such as sweet potato or fruit trees that are vegetatively reproduced. We emphasize that projections concerning global climatic change and the frequent droughts in the American corn belt, the Australian wheat belt and across Asia are a constant reminder of the fragility of modern agricultural systems that lack the balancing

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and compensation potential that traditional farming had. The fact that modern systems are unsatisfied by low, stable yields but rather constantly strive to achieve higher yields is another reason for the price volatility mentioned earlier. Thus, from the perspective of crop breeders, there is no substitute for the gene pools of crop wild relatives that bear diverse genetic variability, among others in traits such as resistance to extreme temperatures or water shortages. We can only hope that the recognition of the important role wild species play in meeting future agricultural needs and guaranteeing continuity in food supplies to populations across the globe will trickle down to circles of decision makers in the Near East and across the world, and find an applicable expression in the form of effective protection against development initiatives targeted at regions in which these precious wild populations still grow. K E Y PO IN TS A ND BE Y O ND

• The significant questions raised in this chapter pertain to key issues of modern global economy and social order, and they are therefore highly relevant to our current realities. It is important to note that the fundamental questions remain unchanged, and that they accompany public discourse to this day. • Other than in the Near East, crop packages that are agriculturally and nutritionally balanced were independently developed in antiquity in several world regions (e.g., China, Central and South America and Ethiopia). The antiquity and location of domestication centres may be identified based on combined botanical, archaeological (cultural), philological and genetic evidence. • In isolated world regions and under traditional farming, extensive genetic variability is accumulated in important agricultural traits (as well as quality traits) of crop species, yet (genetic) variability centres of domesticated crops may not necessarily overlap their domestication centres. • Post-World War II modernization and industrialization encouraged the replacement of local landraces with modern cultivars in many world regions. Many traditional cultivars that represented local adaptability and qualities, which were selected by many farming generations, have been lost without a trace. As a result, a network of gene banks was founded in order to collect, document and preserve the beneficial genetic variability of traditional cultivars and wild progenitors, which can be used in future breeding of high-yielding modern cultivars. • Ironically, this pattern of too late (often futile) conservation attempts is repeating itself in many world regions and concerns landscape, wildlife, natural ecosystems and ancient historic sites that are constantly sacrificed for the sake of modernization, often in the name of ‘economic growth’. Dedicated individuals who foresee the results of the incurred damage initiate some conservation efforts. They establish their own collections, of seeds, for example, and raise public attention and awareness, at times followed by late governmental actions.

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14 A N I M A L D O M E S T I C A T I O N I N TH E N E A R E A S T Gila Kahila Bar-Gal

As indicated in the preceding chapters, the Agricultural Revolution in the Levant involved both plant and animal domestication. In this chapter, I discuss several basic aspects of Near Eastern animal domestication, including when, where and how it took place. I also discuss the genetic and biological differences between domesticated livestock and their wild progenitors, the value and role of domesticated animals in traditional farming systems, and the effect domesticated animals had on humans. Sheep, goat, cattle and pig were the four major animal species domesticated in the Near East in the Neolithic period. The common and scientific names of these species, known as the ‘Big Four’, as well as those of their wild progenitors are listed in Table 14.1 (and see also Figure 14.1). In addition, the dog was domesticated some time in the late Pleistocene and the cat, possibly parallel to the ‘Big Four’, but the two species became pets rather than food resources.1 T H E BI O G E O G R A P H Y OF T H E R E G I O N

The Levant is a continental corridor between Africa and Eurasia. Significant climate changes during the Pleistocene era resulted in major shifts in biogeography and faunal turnovers in the Near East, with cyclic climatic swings on the continents triggering animal and plant migrations. Such shifts in climate occurred during and after the Last Glacial Maximum (the Epipaleolithic period (23,000–ca. 12,000 cal BP); see Chapter 2). At the end of the Pleistocene and the beginning of the Holocene, the climate became warmer and vegetation increased, as shown in the cyclical upturns in humidity and precipitation on the Arabian

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a

b

c

d

e

f

g

h

Figure 14.1 The wild (right) and domesticated (left) package animals. (a, b) Goat; (c, d) sheep; (e, f ) cattle; (g, h) pig. Photo credits: domestic goat, Ms T. Oron; wild goat, Mr A. Kantorovich; domestic cattle, Ms Rachel Gabrieli; excavated skeleton of wild cattle, Yiftahel - courtesy of Dr M Khalaily; wild pig, Mr Amir Balaban. Colour versions of these images can be found at www.cambridge.org/abbo-gopher.

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Table 14.1 Domesticated animal species in the Near East Common name

Latin name

The wild progenitor

Chromosome number*

Approximate time of domestication (cal BP)^

Cattle, Zebu

Bos taurus Bos indicus

Aurochs (Bos primigenius)

60 (29+1)#

10,500–10,300 8,000 (zebu)

Sheep

Ovis aries

Asiatic mouflon (Ovis orientalis)

54** (27+1)

11,000/10,500

Goat

Capra hircus

Bezoar goat (Capra aegagrus)

60 (29+1)

10,500

Pig

Sus scrofa

Wild pig (Sus scrofa)

38*** (18+1)

10,500–10,300

Dog

Canis lupus familiaries

Grey wolf (Canis lupus)

78 (38+1)

ca. 15,000–14,000****

Cat

Felis catus

Wild cat (Felis silvestris)

38 (18+1)

9,500

* In brackets/parentheses the number of autosome pairs and one pair of sex chromosome (see Glossary, Genetics, Allosomes). ^ Approximate time frame of domestication (calibrated years before present) is based on MacHugh et al. 2017; Larson et al. 2014; and Larson and Fuller 2014. # Bos taurus and Bos indicus can be distinguished by their sex chromosomes (Jorge 1974; Jantarat et al. 2009). ** Variation was identified, range 48–58, in wild sheep 54–58; in domestic sheep 52–53 with majority of 54. *** In exotic pigs a fundamental number of sixty-four chromosomes was identified (Vishnu et al. 2015). **** In the Near East Natufian (Davis and Valla 1978; Tchernov and Valla 1997).

Peninsula. It was suggested that this climatic amelioration in the Near East (Levant, Arabian Peninsula and surrounding regions) may have been the driving force of faunal dispersals. For example, Arvicanthis ectos and Mastomys batei (African rodents), which required dense vegetation and considerable water supply, migrated at this time from Africa into the southern Levant. Consequently, the Levant’s large mammal community as it is known today comprises a mixture of species of various origins: Palearctic (deer (Cervus elaphus, Dama mesopotamica and Capreolus capreolus), wild boar (Sus scrofa), Bezoar goat (Capra aegagrus) and brown bear (Ursus arctus)), Holoarctic (wolf (Canis lupus), red fox (Vulpes vulpes) and wild cat (Felis silvestris)), African (gazelle (Gazella gazella), striped hyena (Hyaena hyaena)) and Asian (onager (Equus hemiones)). Archaeozoological (see Glossary, Zoology, Archaeozoology) evidence (animal bone remains in archaeological sites) further indicates that during the Epipaleolithic period the human population of the Levant consumed the common wild fauna that flourished throughout the oak–pistachio woodland surrounding their settlements. The assemblages comprise high frequencies of gazelles together with the wild progenitors of domesticated animals (i.e., Bos primigenius, Capra sp., Capra aegagrus and Sus scrofa, Felis sp. size of F. silvestris and Canis lupus). The rich floral and faunal environment of the Levant exposed human hunter-gatherer populations to a variety of edible plant and wildlife resources.

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Figure 14.2 The distributions of Bos primigenius and Sus scrofa extend from western Europe to east Asia, including the Near East, while the distributions of Ovis orientalis and Capra aegagrus are smaller. All four distributions cover much of the plant domestication core area in south-eastern Turkey and north Syria.

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E C O L O G Y , D I S T R I B U T I O N A N D D E S C R I P T I O N OF TH E F O U R W I L D PROGENITORS OF LIVESTOCK

Bezoar wild goat (Capra aegagrus): The Bezoar wild goat dispersed from central Afghanistan

and southern Pakistan, west through Iran, western Turkmenistan, northern Iraq, the Caucasus region (Armenia, Azerbaijan, north-eastern Georgia and southern Russia), as far as south-western Turkey (Figure 14.2). Based on archaeological evidence, it became extinct from the southern Levant (Israel, Jordan, Lebanon, the Palestinian Authority and southern Syria) some 10,500 years ago (Dayan et al. 1986; Grubb 2005; Alberto et al. 2018). In this wide range of distribution four subspecies of wild goat exist: C. a. aegagrus (Afghanistan, Armenia, Azerbaijan, Georgia, Russian East Caucasus, Turkey, Iran), C. a. blythi (Pakistan, Iran, Iraq, Turkmenistan), C. a. chialtanensis (Pakistan) and C. a. cretica (Greece). The status of the latter subspecies is questioned as it may be a form of an early domestic goat that became feral on the Greek islands. Whole mitochondria genome analyses of ancient samples from the sites of Hovk1 Cave, Armenia, Direkli Cave, Turkey, ‘Ain Ghazal and Abu-Ghosh, southern Levant, found that the Bezoar goat was probably the progenitor of the domestic goat (Daly et al. 2018). The Bezoar wild goat has distinctive colouration (brownish, with typical black stripes through the shoulders and side along the abdomen) and the males are characterized by a long black beard and large, arc-curved horns. The males usually live alone and the females gather with their young in small flocks (up to fifteen animals). The Bezoar wild goat is herbivorous, feeding on herbaceous plants and shrubs (see Box 19 Feeding behaviour and diet selection of the ‘Big Four’ livestock, p. 214). Their diet consists of a variety of plants, including selected specific species that are not necessarily the most abundant ones (Aldezabal and Garin 2000). The wild goat inhabits mountainous areas with rocky outcrops (including scree slopes) and shrubby thickets (maquis) or coniferous forests. Its diet includes plants with high tannin content as they are able to tolerate the intoxication (Aldezabal and Garin 2000). Despite anti-nutritional (see Glossary, General Terms, Anti-nutritional factors) and toxic effects of tannin-rich plants wild goats tend to feed on them as they have beneficial roles in animal nutrition (dietary protein protection from ruminal microflora attack) and health (Lamy et al. 2011). Hence, goats are less sensitive to tannins than sheep and cattle. Wild sheep (mouflon; Ovis orientalis orientalis): The distribution of the wild sheep ranges from Urial or Arkar in Afghanistan, north-western India (Kashmir), Iran, south-western Kazakhstan, Pakistan, Tajikistan, Turkmenistan, Uzbekistan, Armenia, southern Azerbaijan, northern Iraq, eastern Turkey, with an isolated population in south-central Turkey (Figure 14.2) and Oman (where it was possibly introduced). Along this region there are three subspecies of the wild sheep (Ovis orientalis laristanica (Laristan), Ovis orientalis orientalis (mouflon) and Ovis orientalis vignei (the Urial)). The contribution of the Urial (Ovis orientalis vignei) to the domestic sheep’s wild ancestor is debated. The

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International Union for Conservation of Nature (IUCN) considers it as a separate species (Ovis orientalis), while the Integrated Taxonomic Information System (ITIS) classifies it as a subspecies of Ovis aries. The wild sheep, Urial, is distributed in Asia and the Middle East on grassy terrains at high elevations. In several of the regions, an intermediate Laristan sheep (Ovis orientalis laristanica) occurs together with the mouflon (Ovis orientalis orientalis). The presence of wild sheep bones in the assemblages of the Late Pleistocene–Early Holocene sites in Anatolia (Hallan Çemi Tepesi), Iraq (Palegawra Cave and Zawi Chemi Shanidar), Iran (Warwasi Cave) and the southern Levant (Shubayqa 1 and 6, Nacharini Cave) indicates its ancient geographical distribution across the Fertile Crescent. The wide distribution of the wild sheep as found at the very end of the Epipaleolithic period in Middle Euphrates sites of the late Natufian culture (see Chapter 2) such as Mureybet and Tell Abu Hureyra (dated to ca. 13,000–12,000 cal BP) indicate that this species could inhabit relatively open environments beyond the foothills of the Taurus and Zagros Mountain ranges. The findings of wild sheep bone remains in the Negev, Israel, and southern Jordan (at pre-Neolithic sites such as Rosh Horesha, Abu Salem, Ramat Harif, Wadi Judayid and Wadi Mataha), in desert areas, stresses its adaptation, or reflects probable refugia populations adapted to arid and semi-arid habitats. Although the wild sheep, as herbivores, feed on grasses and shrubs, as well as grains and tree leaves, variation in body size is found throughout their wide distribution range associated with the environmental conditions and availability of food. For example, in Iran the mouflon occurs together with the Laristan, known as the smallest wild sheep in the world. It is of note that wild sheep bones have been found in the region in much earlier, Prehistoric sites such as Douara Cave in Syria, assigned to the Middle Paleolithic period (> 100,000 cal BP). Sheep are less selective than goats in their diet and they use pasture more effectively, but in harsh environments their productivity decreases greatly (Lamy et al. 2011). Most of the referred differences on diet selection between sheep and goats can be attributed to genetics, which account for differences in innate sensory ability and other morphophysiological characteristics (Lamy et al. 2011). The overlap of the wild sheep with the range of Bezoar goat and Nubian ibex (Capra nubiana) makes direct identification of archaeological Caprinea bones to the species level quite challenging. Aurochs (Bos primigenius): The aurochs is extinct but once had three subspecies: Bos primigenius primigenius from Europe and the Middle East; Bos primigenius namadicus from India; and Bos primigenius mauretanicus from North Africa. The Bos primigenius subspecies survived until recent times. The last recorded live aurochs, a female, died in 1627 in the Jaktorów (Jaktorowka) Forest, Masovia, Poland (Grubb 2005). The original distribution of the aurochs was from the British Isles and southern Scandinavia, through most of Europe to northern Africa, the Near East, central Asia and India (Figure 14.2). By the

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thirteenth century CE the aurochs’ range was restricted to Poland, Lithuania, Moldova, Transylvania and East Prussia.2 The aurochs were among the largest herbivores in postglacial Europe, and their size varied by region; in Europe, northern animals were larger on average than those from the south (bulls, height at shoulders 155–180 cm; cows, 135–155 cm). Like many bovids, aurochs lived in herds (up to thirty individuals) for at least part of the year. As with other wild cattle, ungulates that form unisexual herds, considerable sexual dimorphism was expressed. The wide distribution of aurochs questions its habitat selection. It is assumed that it inhabited open grasslands and was a grazer (rather than a browser; see Box 19 Feeding behaviour and diet selection of the ‘Big Four’ livestock, p. 214) with food selection similar to domestic cattle. One of the ecological consequences of such grazing activity was the maintenance of open woodland (the socalled park forest) areas. Isotope analyses of Mesolithic aurochs and present-day domestic cattle bones show that aurochs probably inhabited wetter areas than domestic cattle. After the beginning of the Common Era, the habitats of aurochs became more fragmented (see Glossary, Botany, Ecology and Agronomy, Habitat fragmentation) because of the steadily growing human population. During the last centuries of its existence, the aurochs was limited to remote regions, such as flood plain forests or marshes. Wild boar (Sus scrofa): Boars, pigs and hogs are dispersed throughout the world, except

for Australia, Antarctica, northern Africa and far northern Eurasia. The Eurasian wild pig has a wide geographic distribution from Europe, through the Fertile Crescent to Asia and India (Figure 14.2). Its distribution range has been greatly expanded by human agency. The wild boar is opportunistic, utilizing an extremely wide variety of habitats, typically living in grasslands, wetlands, rainforests, savannas, scrublands and temperate forests. It is omnivorous with a preference for vegetal food with high nutritional value. Its diet is largely determined by availability, which means that it is seasonal. In addition, environmental conditions (temperature, winter harshness, etc.), which affect vegetation productivity (i.e., food resources), are strongly correlated to wild boar food search and movement. Hence, the home range is influenced mainly by food availability and density of wild boar populations. Increasing food availability decreases movement and home range size to reduce risk and energy expenditure (Massei et al. 1997). Wild boars choose different habitats in their home ranges for day and for night: they choose open areas where they can be active during the night and the forest for resting during the day. Wild pigs vary greatly in size and weight (7–320 kg) and can take advantage of any forage resources, including roots, fruit, rodents and small reptiles. D E S C R I P T I O N OF TH E ‘ BIG F O UR ’ D O M E S T I C A T E D L I V E S T O C K

Goat (Capra hircus): Domestic goats are herd animals with a lifespan of fifteen to eighteen years. Breeding is on a seasonal basis with a gestation period of five months

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and an average birth rate of 2.2 kids per year. Goats are known as versatile animals; they can survive in diverse climate conditions, including harsh environments, and eat a wide variety of plants. Goats are browsers; they naturally prefer to eat and clear up weeds and brush before they eat grass, including plants rich in tannins. In their natural habitat, they roam mountaintops and climb as high as possible to pick out choice bits of forage. As mountain animals, they are very good at scaling whatever is before them; they have been known to climb to the tops of trees, or even dams. Studies have shown that there are differences in plants’ nutrient digestibility through the course of the year, higher in spring and summer and significantly lower during the autumn and winter (Sun et al. 2014) or during rainy periods (Sorathiya et al. 2016). On a rich diet, a dairy goat can deliver a tenth of its body weight (or more) in milk daily. A goat’s mass varies from 20 to 40 kg and is an excellent source of lean meat with high protein content.

Sheep (Ovis aries): Domestic sheep are extremely versatile and persist in a wide variety

of habitats worldwide, ranging from temperate mountain forests to desert conditions (MacDonald 1984; Grzimek 1990). They are extremely hardy animals with a lifespan of ten to twelve years. Domestic sheep vary distinctly from wild sheep. In domestic breeds, for example, the eye socket and brain case in the skull are reduced and the tail is larger and used as a fat reserve. There is also great variation in physical characteristics among domestic sheep breeds, most of the differences are associated with economically important traits such as body mass (20–200 kg), wool and horn features, colouration, etc. Rams (male sheep) are fertile year-round while ewes (female sheep) are fertile from early autumn to mid-winter. Breeding, therefore, is on a seasonal basis, with a gestation period of about five months (148 days) resulting with one or two lambs mostly born in mid-spring (Ensminger 1964). Sheep are grazers, feed mainly on grasses and can survive on a diet consisting of only cellulose, starch or sugars as an energy source. They have a large and complex stomach, which is able to digest highly fibrous foods that cannot be digested by many other animals (Ensminger 1964; Hecker 1983). Sheep are more sensitive to tannins than goats. The perceived sensitivity to tannins could be related to the different ability of species to tolerate and orally detect these compounds (Lamy et al. 2011). Sheep are one of the most economically important livestock, and they can be maintained in many environments at relatively low cost. In some cultures, sheep are highly prized as a sacrificial animal.

Cattle (Bos taurus and Bos indicus): Domestic cattle have a herd mentality; being together in a herd reduces stress levels and increases the safety level for each individual. Their lifespan is about twenty-five years. They have a gestation period of nine months, which usually

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Table 14.2 The ‘Big Four’ – characteristics considered favourable for animal domestication* Trait

Sheep

Goat

Cattle

Pig

Social structure

Dominance hierarchy Large gregarious groups Persistent groups Non-territorial groups

✓ ✓

✓ ✓

✓ ✓

✓ ✓

✓

✓

✓

✓

Food preference

Generalist herbivorous Omnivore

✓

✓

✓

Reproduction

Males dominant over females Males initiate

✓ ✓

✓ ✓

Behaviour (characteristics)

Non-aggressive Tamable Solicits attention Readily controlled

✓* ✓

✓* ✓

Plasticity and adaptation

Wide environmental tolerance Low sensitivity to environmental changes Small home range Exploits anthropogenic environments

Partial Partial

✓ ✓

✓

✓

✓

✓**

* Based on relevant literature, mainly on Driscoll et al. 2009. ** Except males in heat, those that are ready to propagate and are fertile.

results in one offspring. Cattle are commonly raised as livestock for meat, milk and hides/leather and are used as draught and riding animals. The weight of an adult cow varies enormously (272–1,362 kg) depending on the breed, with cattle raised for meat and draught being the biggest. Cattle are ruminants; they are grazers with a digestive system that is highly specialized to allow the use of poorly digestible plants as food. During grazing they vary in several aspects of their bite, that is, tongue and jaw movements, depending on characteristics of the plant they are eating. Cattle also adjust other aspects of their grazing behaviour in relation to the available food; foraging velocity decreases and intake rate increases in areas of abundant palatable forage (Bailey et al. 2016). As domestic livestock it is versatile and can exist in a wide variety of habitats worldwide. Hence, it was found that environmental factors, geographical location, climate conditions and seasonality have a direct influence on body mass (Bradford et al. 2016) – such as in beef cows in pasture-based production systems that show strong seasonal fluctuations in body weight, with substantial variation between animals (Meyer and Colditz 2015). For example, in Australia the extreme seasonal variation results in an annual pattern of weight gains and losses, depending on feed availability (Bradford et al. 2016). Pig (Sus scrofa): Pigs are considered very intelligent social animals and they have a

lifespan of about eight years. Female pigs, called cows or sows, give birth to

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offspring twice a year with an average normal litter of eight to twelve piglets. The gestation period is 115 days and the sows are ready to breed again about five to seven days after they wean their piglets. In the wild, they live in sounders (a closeknit group) that consist of one male, many females and their young. Males without a sounder remain solitary. In captivity domesticated pigs can have sounders with up to 300 members. Since they are omnivorous and can consume a wide range of foodstuffs, they can live in virtually any productive habitat that can provide enough water to sustain their large bodies. Under domestication, pig breeds were selected by farmers for size and weight and like wild pigs they vary greatly from 50 to 350 kg. Domestic pigs are raised as livestock material that generate meat, leather and bristly hair, which is used for brush making. Their omnivorous diet, aggressive behaviour and their feeding method of rooting in the ground all combine to severely damage ecosystems, hence they are known as major drivers of extinctions and ecosystem disturbances. I have presented the biogeography and a history of the ‘Big Four’ progenitors, their ecology and geographical distribution, and then a survey of the four domesticates: their life cycle, breeding, ecology, diet and behaviour. As a sort of a summary, Table 14.2 highlights the features favourable for domestication vis-à-vis the ‘Big Four’.

T H E W H E N A N D W H E R E O F A N I M A L DO M E S T I C A T I O N IN T H E N E A R E A S T

Goat (Capra hircus): Goats were first domesticated from the wild Bezoar goat some time

between 11,000 and 10,000 cal BP in the northern parts of the Levant. The progenitor of the goat, the Bezoar goat, was distributed throughout Anatolia and the Fertile Crescent during the Epipaleolithic and early Neolithic periods prior to the domestication that occurred in the Fertile Crescent (Daly et al. 2018). The unique ability of goats to draw nutrients from woody vegetation not accessible to other ruminants like cattle and sheep enabled them to adapt and thrive in marginal environments and provide a reliable source of meat, milk and hide. Surveys of mitochondrial DNA (mtDNA; see Glossary, Genetics, Mitochondrial DNA) indicate that goats underwent rapid demographic expansion following domestication (Naderi et al. 2008). The high frequency of haplogroup A (90%) among goats (see Glossary, Genetics, Haplotype), independent of their geographic origin, suggests that their distribution was assisted by human migration and the exchange of animals (Naderi et al. 2007). In addition, less frequent haplogroups also dispersed across large geographic areas. These two points together indicate a population history characterized by high levels of animal movement and genetic admixture. Genome Wide Association Studies (GWAS) further support high rates of DNA diversity, which strongly suggests that goats are probably more polymorphic (see Glossary, Genetics, Polymorphism) than cattle, sheep and pig (Zhang and Plastow 2011). The prevalence of different haplogroups (based on mtDNA analysis) together with the GWAS

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variation indicate that domestication took place in a few foci in the Fertile Crescent within an active cultural interaction sphere, or, in other words, within communities that were in cultural contact (i.e., not independently). Genomic (see Glossary, Genetics, Genomics) analysis detected early selection for pigmentation and offers interpretation of allelic variants affecting stature, calving interval and mammary gland development, which can be associated with domestication traits such as reduction in size, milking and response to dietary change (Bailey et al. 2016). The genetic conclusions are supported by the archaeozoological data (Legge 1996; Peters et al. 1999, 2005; Naderi et al. 2008) showing morphological changes associated with domestication. Studies based on genomic analysis of modern specimens show differences between the wild progenitor, the Bezoar goat, and the domestic goat but at the same time imply low DNA polymorphism indicative of high inbreeding in the Bezoar compared to goats (Alberto et al. 2018). A recent genomic study, based on eighty-three ancient specimens, concluded that the Bezoar was the progenitor of the domestic goat and that goat domestication was a mosaic (diffused) rather than a singular (centred) process, with capture from the wild in different regions of the Fertile Crescent impacting genetic diversity. A similar picture was indirectly suggested based on the analysis of human genomes of Neolithic populations from North-western Anatolia and Iran (GallegoLlorente et al. 2016; Lazaridis et al. 2016; Broushaki et al. 2017). Yet viewing Near Eastern animal domestication as dispersed in space and time is not in accordance with archaeozoological data that can be viewed as indicating a centred, one-time domestication that has later spread throughout the region and beyond. Moreover, it is hard to claim and accept cultural isolation and independent domestications in the communities discussed since they were part of the same active Neolithic cultural interaction sphere.

Sheep (Ovis aries): Sheep were domesticated from the wild sheep (mouflon; Ovis orientalis) around 10,500 years ago in the Fertile Crescent region, south-eastern Anatolia and the Iranian Zagros Mountains (Alberto et al. 2018). Furthermore, the wild sheep overlapped with the range of the abundant Bezoar goat in the area prior to domestication (Figure 14.2). Mitochondrial genome analyses imply that the five sheep maternal lineages evolved at different times before domestication. Thus, the wild ancestors of domestic sheep underwent a major population expansion starting before the Last Glacial Maximum (~26,500–19,000 cal BP). The ancestor of domesticated sheep originated from one geographical location in the Fertile Crescent, and this is where it was domesticated. Sheep migrated post-domestication in two general directions, westwards into Europe as part of the Neolithization of Europe, and eastwards into Asia followed in two migratory waves at approximately 6,800–4,500 cal BP – one through the Caucasus, eastwards to Central Asia, the Mongolian Plateau region and to China, and the second from the Fertile Crescent and Iraq to Iran and India. The migration of domestic sheep with humans created the original maternal genetic make-up of modern sheep across

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Eurasia and China (Lv et al. 2015). Recent genomic studies suggest biological processes influenced by domestication and artificial selection including: (1) dorsal/ventral patterning as reduction in body size was quick and early; (2) regulation of lipid metabolism, which may reflect human-mediated alteration of muscularity and fatness as sheep were actively managed as a food source; (3) sexual maturation: as humans increasingly exerted control over breeding, pressure to maintain beneficial sexual fitness traits in the wild were removed, reducing sexual dimorphism and altering the timing of reproduction.

Cattle (Bos taurus and Bos indicus): Cattle were domesticated in the Near East (Vigne and

Helmer 2007; Bollongino et al. 2012; Larson and Fuller 2014; Scheu et al. 2015). The earliest archaeological evidence of domestic cattle (Bos taurus) in the Fertile Crescent is from the sites of Dja’de and Çayönü3 and dates to the second half of the eleventh millennium cal BP (possibly ca. 10,300 cal BP or somewhat earlier (Helmer et al. 2005; MacHugh et al. 2017)) and about 1,500–2,000 years later in the Indus Valley (Bos indicus). In south-west Europe domestic cattle can be traced back to the ninth millennium cal BP (9,000–8,000 cal BP) and in the next millennium in western, central and northern Europe. All of these findings are within the former geographic range of the aurochs, which can be differentiated from domestic cattle remains by the size of their bones. Recent genetic data further support the archaeological finding, indicating that taurine cattle originated in the Fertile Crescent and the zebu cattle originate from the Indus Valley. An interesting statement based on a continent-wide evaluation of the early spatio-temporal demography of Bos taurus by Scheu et al. (2015) concluded that: ‘domestic cattle indeed have a discrete and rather localized origin, very likely in Southeastern Anatolia and the Near East, a view that is consistent with a huge body of archaeozoological evidence from the 9th millennium BCE’ (11,000–10,000 cal BP) (Helmer et al. 2005; Peters et al. 2005; Achilli et al. 2008; Hongo et al. 2009). The domestic cattle dispersed while they accompanied human migrations and this resulted in adaptation to different environments ranging from green pastures to deserts and they developed considerable variation in appearance and performance. Neolithic cattle remains were found to be smaller than aurochs and continued to decrease in size until the Middle Ages. The decrease in body size was associated with the reduction of horn size and shape. Pig (Sus scrofa): Pigs were first domesticated in the Near East (Anatolia (Ervynck et al.

2001)) around 10,500 cal BP (8,500 cal BC (Ervynck et al. 2001)) and subsequently brought into Europe by agriculturalists (Larson et al. 2007); others would claim a somewhat later domestication in the Near East (10,300 cal BP; see MacHugh et al. 2017). Pigs were also (later) domesticated independently in the Mekong Valley in Vietnam about 9,000 cal BP (Cucchi et al. 2011). Analyses of ancient mitochondrial DNA (mtDNA) indicate that the first domestic pigs in Europe were transported by early

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Table 14.3 The feeding behaviour of the Big Four Near Eastern livestock

Feeding behaviour Herbivore

Omnivore

Livestock

Selective feeder*

Grazers

Cattle

Less***

Browsers

Goat

Most***

Intermediate feeders Foragers

Sheep

Medium***

Pig

Less***

* Selective feeder: food procurement in which the animal exercises choice over the type of food ** Secondary plant component: plants produce secondary metabolites, divided into three chemically *** Less/Most (High)/Medium: description of the animals’ reaction to (1) the choice of food –

farmers from the Near East, from Anatolia into Europe around 7,500 years cal BP, concordant with archaeozoological evidence for a single domestication origin of west Eurasian domestic pigs. However, a few thousand years after their introduction (2,500 cal BP), domestic pigs in Europe had completely lost their Near Eastern mtDNA signatures and instead acquired mtDNA haplotypes found in local European wild boar. These findings suggest post-domestication gene flow from wild boar populations to early domestic populations (Frantz et al. 2015). These exchanges of haplotypes between pigs and wild boars suggest extensive mobile swine herding throughout Europe and Anatolia, consistent with both archaeological and historical evidence. Although a common phenomenon, it has no clear advantage to pig production and this gene flow is thought to have occurred spontaneously. In spite of the random wild–domesticated gene flow there is a clear morphological and behavioural dichotomy between wild boars and domestic pigs that is evident in modern animals as well as in the archaeozoological record.

CO MPARI NG T HE TH REE RU M IN ANT S P EC IES

Although sheep, goat and cattle are all ruminants they differ in the way they grasp and ingest forage. With their wide mouths and inflexible upper lips cattle can take large clumps of forage into their mouths. This eating method causes a lack of selectivity and results in ingesting more dead material than sheep and goats, who have narrower mouths and more flexible lips (van Dyne and Heady 2018). Furthermore, goats as browsers differ from sheep and cattle in their preference for eating and clearing up

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Diet selection/preference Tolerance to secondary plant components** Less*** Most*** Medium*** Medium***

Grasses (%)

Forbs (%)

Shrubs/trees (%)

High*** (66–75) Medium*** (20–30) High*** (45–55) Availability

Medium*** (20–30) Low*** (10–30) Medium*** (30–40) Availability

Low*** (5–10) High*** (40–60) Low*** (10–20) Availability

being taken, preference to specific type of food. distinct groups – terpenes, phenolics and nitrogen-containing compounds. quantifying their selective response; (2) the secondary metabolites within their food.

weeds and brush before they eat grass (see Box 19 Feeding behaviour and diet selection of the ‘Big Four’ livestock, p. 214, and Table 14.3). This preference is an advantage in raising goats together with sheep or cattle as there is little competition for food resources; goats leave the grass for others. The amount of energy gained from grazing dry residues is affected by the plant species, duration of stubble availability and amount of grain available within stubble. Among the common stubble in the Mediterranean area are wheat, barley and oat straws. Wheat straw diets are low in protein but easier to process than barley and oat. The cereal stubbles contain starch that can cause acidosis if rapidly consumed and will not provide sufficient protein for maturing lambs. Differences in dietary chemical composition were detected among grazing animals, sheep and cattle (van Dyne and Heady 2018). During dry seasons or harsh conditions sheep and goats can feed on stubble because they are able to select the more nutritious parts of it. The content of the stubble, especially the protein, has an effect on weight change and, as a result, on pregnancy too. The increase of genetic and genomic information, including ancient DNA data, indicates patterns of genomic diversity in domestic populations, demonstrating that domestic animals have not evolved in complete isolation, but rather went through significant admixture with their wild progenitors. For example, in the case of cattle, which were domesticated from a local limited stock in the Near East (South-eastern Anatolia), alleles introgression (alleles exchange between distinct gene pools via mating) from wild aurochs into domestic herds during their expansion has been frequently observed (Bollongino et al. 2012; Scheu et al. 2015; MacHugh et al. 2017). The case of the domestic pig shows a similar pattern. Genetic and archaeological evidence reveals

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Box 19 Feeding behaviour and diet selection of the ‘Big Four’ livestock Each of the wild progenitors of the ‘Big Four’ livestock has its own ecological affinities as

indicated in the geographic range it has occupied prior to domestication (Figure 14.2). Plant life, in terms of species diversity, richness and potential production, is determined by the local conditions

Figure 14.3 A goat climbing a tree – courtesy of Ms Talia Oron. A colour version of this image can be found at www.cambridge.org/abbo-gopher.

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(e.g., yearly precipitation, temperature profiles and soil characteristics). The major plant formations of the Near East were described in Chapter 4 (see map therein). As indicated in the distribution maps of the ‘Big Four’ wild progenitors (Figure 14.2) two, namely, wild pig and wild cattle, span a huge natural range (extending from temperate and moist across to semi-arid regions) with no apparent adherence to any specific vegetation zone; while the wild progenitors of the domesticated goat and sheep had a significantly narrower distribution, mostly confined to the montane areas that sustain east Mediterranean and Irano-Turanian woodlands (Figure 4.1). Evolutionary adaptation to the available vegetal materials across their native range, as well as their inherent capacity to grasp (lips and tongue flexibility) and digest plant material (different complexity of the digestive systems), are expressed in the feeding habits and preferences of the ‘Big Four’ livestock. Cattle are considered grazers – namely, an animal that feeds on tender plants (low vegetation), grasses, clovers and the like, weeds and on other multicellular autotrophs (such as algae). Cattle are not considered highly selective in their feeding habits. Goats are considered browsers – animals that feed on leaves, twigs, vines, soft shoots or fruits of high-growing, generally woody,

plants such as shrubs. This feeding behaviour is of course assisted by their excellent climbing ability, and they may often be seen in treetops in Mediterranean rangelands (Figure 14.3). The goat’s high tolerance for secondary plant compounds (e.g., tannins) enables it to feed on sclerophyllous foliage (hard leaves), which is typical of many Mediterranean and IranoTuranian trees and shrubs. Sheep have intermediate behaviour, with an intermediate level of foodstuff selectivity and likewise intermediate tolerance of tannins, and lower climbing ability compared to goats. Wild pigs are foragers – animals that search for and feed on a wide range of wild food resources. These include (but are not limited to) weeds, grasses, browse and brush plant materials. Being omnivorous, pigs do not restrict themselves to a vegetal diet and may feed upon human refuse heaps and even small animals (e.g., reptiles). Considering the differential food preferences of the ‘Big Four’, it could be argued that there is rather limited competition for resources between these animals when grazing together around a certain territory. It is possible that this was one of the considerations of the Neolithic domesticators in their choice of wild animal species for domestication.

that the origin of European pigs in the Near East was followed by a complex history of trade and multiple importations that masked the genetic signature of the early domestic pigs from South-eastern Anatolia by local introgression from wild boars as the domestic pig was expanding to Western Anatolia and into Europe (Ottoni et al. 2013; Frantz et al. 2015; MacHugh et al. 2017). Genomic comparison between sheep and goats indicated opposite global patterns of genomic diversity. In Capra, the low DNA sequence diversity (i.e., genetic polymorphism; see Glossary, Genetics, Polymorphism) and high inbreeding in the Bezoar goat compared to domestic goats can be the result of several declines in the wild

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populations due to extensive poaching and habitat fragmentation (i.e., discontinuities in an organism’s preferred environment) (Weinberg et al. 2008). In Ovis, on the other hand, the wild populations exhibited higher diversity than their domestic counterparts. The increased presence of favourable alleles in the genes of sheep populations may represent a domestication signature, where demographic bottlenecks reduced the efficacy of negative selection in purging harmful mutations from the domestic gene pool (Alberto et al. 2018). The patterns of gene selection vary significantly between the genomes of very similar species, such as sheep and goats, suggesting that different ‘genetic solutions’ have given rise to similar phenotypic traits (Alberto et al. 2018). T H E BI O L O G I C A L DI F F E R E N C E S B E T W E E N W I L D A N D DOMESTICATED ANIMALS

The study of the ‘animal domestication’ phenomenon is based on archaeological faunal remains analysed together with genetic/genomic information obtained from modern forms of wild candidate progenitors and from traditional breeds of domesticated animals. The commonly used identification of a ‘domestic animal’ is based on morphological diversity (disparity) of the domesticated forms resulting from conscious human selection for favourable traits (strong artificial selection) including behavioural traits (Sánchez-Villagra et al. 2017). Therefore, in general, domesticated mammals exhibit a suite of behavioural, physiological and morphological traits not observed in their wild forebears. Today, these characteristics are known to include: increased docility and tameness, coat colour changes, reductions in tooth size, changes in craniofacial morphology, alterations in ear and tail form (e.g., floppy ears), more frequent and nonseasonal oestrus cycles, alterations in adrenocorticotropic hormone levels (see Glossary, Zoology, Adrenocorticotropic hormone (ACTH)), changed concentrations of several neurotransmitters, prolongation of juvenile behaviour and reduction in both total brain size and of particular brain regions (Wilkins, Wrangham and Fitch 2014; Wilkins 2017). The general combination of traits in domesticated mammals is an ensemble referred to as the ‘domestication syndrome’ (DS) (Brown et al. 2009) akin to the DS in plants (Hammer 1984 and see comments in Chapter 5). Unfortunately, except for skeletal variations all of the other DS characteristics such as behavioural changes and most of the morphological characteristics, such as coat colour, fur texture, etc., are archaeologically translucent. Therefore, the widely accepted morphological marker of domestication used by archaeozoologists, anthropologists and archaeologists is the sharp and rapid reduction in overall body size (Vigne et al. 2005; Zeder 2008; Wright 2015), with the skull being the most extensively used marker of morphological diversity, given its complexity in form (shape and size) and embryological origin. Although domestication has influenced postnatal growth trajectories, no DS trait for ontogeny (see Glossary, Zoology, Ontogeny) has been identified, because there is no

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single, universal pattern in skull morphology that accompanies domestication. The patterns that have been discovered exemplify the complex nature of evolutionary changes in the skull upon domestication, and indicate that there is no single, universal and global pattern of retention of juvenile characteristics by the adult (paedomorphosis; see Glossary, Zoology) or any other kind of development leading to changes in size and shape (heterochrony; see Glossary, Zoology) behind the morphological diversification between individuals (Sánchez-Villagra et al. 2017); this stresses the need to study each species separately. Consequently, using metric data to distinguish wild versus domestic forms is problematic, thus questioning past interpretations of archaeozoological results based on similarities or differences between single measurement sets (Vigne et al. 2005). Although traits such as tameness and juvenile behaviour listed in the DS cannot be studied from the archaeozoological records it is assumed that the founder ‘package’ of domesticated animals (sheep, goat, cattle and pig) bore characteristics considered favourable for domestication (dominance hierarchy, generalist herbivorous feeder or omnivore, non-aggressive and moderate sensitivity to environmental change, etc.; see Table 14.2). In other words, they were all behaviourally pre-adapted to domestication. These animals, unlike other wild types abundant in the Near East, such as gazelles and fallow deer, were herd animals with temperaments geared to adaptation to confinement, living on a flexible diet with fast growth rates and breeding regardless of the environmental conditions (Table 14.2), features that enabled them to subsist under human control without expending farmer resources. As the wild progenitors of the ‘Big Four’ livestock species are known to avoid anthropogenic (human) environments4 it is evident that their domestication was a conscious process on the part of humans, who deliberately selected them based on the knowledge and awareness they had about their biological and behavioural characteristics. Furthermore, animals that bred well in controlled environments went through a mixture of artificial selection for favourable traits, and natural selection for adaptation to captivity, with artificial selection being the prime mover. An interesting view from an economist (Svizzero 2016) suggests that animal domestication, specifically of sheep and goats, was initiated not only via selective hunting but also by baiting, and, more specifically, through the cultivation of food plots (whether wild or domesticated) for attracting wild game, either to kill them or to capture them alive. Recent genomic analyses comparing wild progenitors and domestic forms (including local breeds at the respective geographical regions suggested as the domestication areas (centres), traditional breeds and exotic breeds) indicate that the main force driving domestication was strong selection for morphological and behavioural traits required by humans such as pigmentation (KIT and KITLG genes; Kijas et al. 2012; Alberto et al. 2018), reduction of body size (SIRT1; Li 2013) and milk production (the STAT1 gene; Cobanoglu et al. 2006). The genes associated with the domesticated condition are related to multiple developmental processes of bones, teeth and the nervous system associated with morphological and morphometric (DS) changes and behaviour

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characteristics such as aggressiveness and feeding. Furthermore, genomic analyses among the four species (goat, sheep, pig, cattle) stress the existence of allele and gene exchange with the wild progenitors during and post domestication. The interbreeding of wild and domestic forms was possible as both shared the same habitats and it is an open question whether this was managed by humans or it occurred spontaneously with humans merely taking advantage of the useful natural progeny. In pigs, despite the undesired gene flow from wild boar, as it has no production advantage, the genomes of domestic pigs have strong signatures of selection at loci that affect behaviour and morphology. Recurrent selection for domestic traits likely counteracted the homogenizing effect of gene flow from wild boars resulting in genomic ‘domestication islands’. That is, chromosomal regions that maintain the desired domesticated allelic variants despite gene flow between wild and domesticated populations (Frantz et al. 2015). Among sheep and goats, genome-wide analyses have identified a number of variants that differentiate domesticates from their wild counterparts. Nevertheless, a genomic study of eighty-three archaeozoological specimens from various locations in the Near East found that the pigmentation loci, KIT and KITLG, are the only shared signals in Neolithic populations and are common signals in modern livestock (Daly et al. 2018). Selection for pigmentation may serve to distinguish individuals and maintain ownership5 or, as proposed by Trut et al. (2009), have a pleiotropic effect of selection for tameness. The finding of pigmentation as the only shared signal is an example of how evidence for trans-specific signatures of domestication remain largely unexplored and mostly elusive across domestic animals. This might reflect a scenario in which selection acted on species-specific traits during domestication. The imperative of addressing species-specific traits (referred to above as ‘different genetic solutions’ to obtaining a domestic phenotype) points to the important role of conscious knowledge-based selection rather than automatic or unconscious domestication dynamics. In addition, it is likely that several domestication traits are predominantly polygenic (a group of nonallelic genes that together control a quantitative character in an organism) and/or pleiotropic (having multiple phenotypic expressions) in nature, allowing selection to target different genes while resulting in similar phenotypes, for example polledness (the state of being hornless) is driven by different genes in sheep and goat (Kijas et al. 2012, 2013). The ancestor of domestic cattle, the wild aurochs (Bos primigenius), is now extinct; therefore, the study of cattle domestication is based solely on archaeological evidence and targeted DNA analyses using ancient DNA recovered from prehistoric specimens. Advances in genomic technology have enabled the sequencing of a whole genome from a specimen of a British aurochs’ humerus bone radiocarbon dated to 6,738  68 cal BP – the beginning of the Neolithic period in Britain. Comparison of the aurochs’ genome with domestic cattle’s genome flagged a number of genes associated with neurobiology, growth and metabolism, and immunobiology that exhibit evidence for positive selection within the time frame since cattle domestication ~10,500 years ago (Park et al. 2015).

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T H E DI F F E R E N T A P P R O A C H E S T O E VO L U T I O N A R Y M E C H A N I S M S O F ANIMAL DOMESTICATION

The need to address animal domestication within a wider evolutionary framework has inspired several scholarly theories and explanations of domestication. These include commensalism, habituations and proto-domestication; niche construction; and punctuated equilibrium theories. Below I will briefly survey the way archaeozoologists extract data from faunal remains at archaeological sites of the relevant time to reconstruct domestication (see Box 20 Animal domestication: How does it show in the archaeological faunal record and what mechanisms are suggested?, p. 222). Commensalism, habituations and proto-domestication: The proto-domestication theory is

founded on animal initiatives to exploit anthropogenic environments. It is one of the common theories used to explain domestication. Commensalism and habituations of animals require that species will have the ability to learn and alter their behaviour in response to human-sourced stimuli (Shettleworth 1998; Stankowich 2008). The behavioural-based boundaries between habituated wild individuals to those in natural habitats might lead to genetic differentiation between them due to segregation and mating preferences, causing pre-mating isolation among populations over short distances. It is commonly suggested that dog domestication had a phase of proto-domestication/commensalism when wolves in search of food were attracted by human camps where they could find some leftovers and they thus became progressively accustomed to the human presence (Clutton-Brock 1995). Stronger evidence of domestication was suggested at a burial site in Germany called Bonn-Oberkassel, dated to 14,000 years BP, indicating partnership between hunters and canines that might have helped track and retrieve wounded animals. The remains found at the site of ‘Eynan (‘Ain Mallaha) dated _ to ~15,000–14,000 years cal BP (Davis and Valla 1978; Bar-Oz et al. 2004a) of a human skeleton lying on its right side in a flexed position with its left hand on the thorax of a puppy stress the strong human–dog relationship (see Box 5 ‘Eynan, p. 31). Genomic studies estimated that dogs and wolves diverged genetically between 41,500 and 36,900 years ago, and that the split between eastern and western dogs occurred 23,900–17,500 years ago. By calibrating the mutation rate using the oldest dog, the timing of dog domestication was delimited to 40,000–20,000 years ago (Botigué et al. 2017). Unlike the dog, most of the species domesticated in prehistoric time, namely, the ‘Big Four’ livestock species, are known to have avoided anthropogenic environments (Driscoll et al. 2009). The commensal house mouse (Mus musculus domesticus) outcompeted the wild mouse (Mus macedonicus) and established durable populations that expanded with human societies starting at Natufian sites ca. 15,000 years ago (Weissbrod et al. 2017). The ‘domestication’ of the house mouse (becoming a laboratory mouse) occurred much later, in the twentieth century. In the Levant, Nubian ibex (Capra nubiana), a wild goat, is well known from the archaeological records in prehistoric

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and historic periods (Zohary et al. 1998; Kahila Bar-Gal et al. 2002; Ureña et al. 2018). Nevertheless, the Nubian ibex never became domesticated despite its close contact with humans, including interaction with nomadic sheep/goat flocks. Recent studies of the Nubian ibex show different behavioural patterns between individuals habituated to human townships (e.g., Mitzpe Ramon, Israel) and those in rural areas (e.g., ‘Ein Gedi Nature Reserve, Israel) but none of these animals are considered domesticated (Iribarren and Kotler 2012). Hence, it is important to stress that the terms ‘habituation/ commensalism’ indicate the ability of animals to live as free individuals in close contact with humans and this should not be mistaken for a conscious act of domestication.

Niche construction: The Niche-Construction Theory, used in ecological studies, refers to

the modification of selective environments by organisms, and emphasizes the capacity of organisms to modify natural selection in their environment and thereby act as codirectors of their own, and other species’, evolution (see Chapter 3; Laland and O’Brien 2010; Laland et al. 2016). It stresses that human niche-altering activities such as ecosystem engineering, relocation of plants and animals, and niche alteration served as selective factors in the environments shared by humans and the would-be domesticated species, to their mutual benefit (Zeder 2016). It was suggested that this was a protracted process that evolved in relatively stable environments, especially ones with abundant, diverse resources (Smith 2016; Zeder 2017). This theory implies a coevolutionary (mutualistic in nature) scenario for domestication and it relies (mostly) on unconscious prey–predator relations, thereby rather minimizing the role of human consciousness, initiative and agency (Zeder 2016). Punctuated equilibrium: The punctuated equilibrium theory was proposed in 1972 by

Eldredge and Gould. The theory explains the evolutionary process based on patterns of first appearances and subsequent histories of species in the fossil record. The abrupt origination of species may result from a rapid genetic switch from one species to its successor, following the law of natural selection (Weninger 2017). Applying this theory to animal domestication can explain the emergence of ‘new species’ using historical, generations and/or nanoseconds scales (for molecular reactions of DNA molecules) rather than a geological time scale.

T H E A R C H A E O Z O O L O G I C A L R E C O RD I N T H E NE A R E A S T , I T S I N T E R P R E TA T I O N A N D A N S W E R IN G T HE QU E S TI ON S O F W HE R E A N D WHEN WERE ANIMALS DOMESTICATED

The archaeozoological and archaeobotanical records from Neolithic sites in the Near East (especially in the Levant) indicate changes in morphology and genetic/genomic

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profiles associated with domestication lacking intermediate forms. If domestic animals were gradually pre-formed (i.e., domesticated) through long-term mutualistic biological interactions of wild animal and human, as suggested by the NicheConstruction Theory, we would expect to find specimens representing intermediate forms. Yet, the archaeozoological record lacks such intermediate types. The record shows abrupt appearance of domestic animal remains, mainly the ‘Big Four’, harbouring the morphological and genomic characteristics associated with domestication. In spite of the appearance of domestic animal remains, wild animal remains are represented in the assemblages of sites dated to the Pre-Pottery Neolithic (ca. 11,600–8,500 cal BP) and the Pottery Neolithic (8,500–6,500 cal BP) periods. In south Levantine Epipaleolithic and Pre-Pottery Neolithic A sites (23,000–10,500 cal BP) the main dominant prey species was the mountain gazelle (Gazella gazella) (Davis 1982; Tchernov 1993; Bar-Oz et al. 2004; Munro and Bar-Oz 2005). During the Early, Middle, Late PPNB and the PPNC (10,500–8,500 cal BP), the dominance of gazelle in archaeozoological assemblages ended and they were virtually replaced by remains of domestic goats and sheep (Peters et al. 1999; Bar-Yosef 2002), such as in the case of Motza (Sapir-Hen et al. 2009), Abu-Ghosh (Kahila Bar-Gal et al. 2002), ‘Ain Ghazal, Basta and Baja. This abrupt appearance of the ‘Big Four’ domestic animals in the archaeological record follows the pattern evident from analyses of the archaeological data from Near Eastern sites concerning the founder crop plants and human culture (as expressed in aspects such as architecture, lithic technology, burial customs and imagery (art) items), together with 14C dates and climatic data (Kolodny et al. 2015; Arbuckle et al. 2016; Weninger 2017). These analyses indicate a punctuated transition from wild to domesticated animals that occurred at 10.2  0.2 ka cal BP (Arbuckle et al. 2016), supporting previous data (for details see Box 20 Animal domestication: How does it show in the archaeological faunal record and what mechanisms are suggested?, p. 222). T H E E F F E C T O F A N I M A L DO M E S T I CA T I O N ON H U M A N C O M M U N I T I E S

One of the forces affecting animal adaptation and survival to anthropogenic changes is the exposure to pathogens. Recently, research into parasite–host relationships exposed the potential of parasites to influence animal hosts (Krasnov and Poulin 2015). In prokaryotes (see Glossary, Zoology, Prokaryotes), lateral gene transfers (LGTs) (see Glossary, Genetics, Lateral gene transfer) play a major role in providing novel protein coding genes and contribute adaptive traits. In eukaryotic genomes (see Glossary, Zoology, Eukaryotes), LGTs were found to influence the evolution of the coding capacity, predominantly affecting enzymes from metabolic pathways and other genes for putative proteins of unknown function (Huang 2013; Schönknecht et al. 2013; Hirt et al. 2015). The close proximity of domestic animals with the contained surroundings constantly exposed them to various pathogens that challenge the immune system, and,

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Box 20 Animal domestication: How does it show in the archaeological faunal record and what mechanisms are suggested? Animal domestication research is based on comparative studies of archaeozoological assemblages from the late Epipaleolithic Natufian culture sites (15,000–ca. 12,000 cal BP) and sites of the early Neolithic period (ca. 11,600–8,500 cal BP) and it focuses on: 1. How species were chosen for domestication: the abundance of species in the ecosystem was not reflected in the chosen species that were eventually domesticated; for example, gazelle spp. played a major role in human diet in the Epipaleolithic and early Neolithic periods and dominated the animal bone (faunal, archaeozoological) assemblages of the sites in the region

(Tchernov 1993, 1998; Bar-Oz et al. 2004; Munro and Bar-Oz 2005; Zeder 2012, 2017; Munro et al. 2018). 2. Phenotype changes: Most of the morphological and behavioural traits that are considered as representing domesticates cannot be detected in archaeozoological remains. Nevertheless, the detectable traits show similar alterations in the different domestic species (Wright 2015). Detectable changing traits including animal size and biometric data (e.g., in teeth, horns) do show that small-sized animals (sheep, goat and cattle) appear suddenly, with no transitional forms, in assemblages of archaeological sites from the Fertile Crescent (e.g., the Zagros Mountains and the Middle Euphrates) dated to the Neolithic period

Figure 14.4 Graphic summary of the decisive human steps that led to the formation of domestic livestock. A colour version of this map can be found at www.cambridge .org/abbo-gopher.

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(Zohary et al. 1998; Kahila Bar-Gal et al. 2002; Zeder 2008, 2012, 2016; Arbuckle 2015; Arbuckle et al. 2016). 3. Herd composition: The rationale leading the expectation of a change in herd composition between the wild and domesticated herds is that under domestication only a few males but large numbers of breeding females are required to ensure herd continuity. Consequently, an increase in culling young males drastically changes the herd’s sex and age composition resulting in reproductive biology changes. This phenomenon is evident from the

archaeozoological record (e.g., Zohary et al. 1998). 4. Genetic change: Principal changes in human–animal relations triggered the transformation of wild animal to domesticated livestock. The genetic changes that accompanied the shift from wild to domestic goats, for example, occurred within a relatively short period of time (Kahila Bar-Gal et al. 2002). A graphic summary of the decisive human steps that led to the formation of domestic livestock is depicted in Figure 14.4.

ultimately, in a short time, shape it (Perry et al. 2007, 2015; Axelsson et al. 2013; Thornhill and Fincher 2014). Furthermore, the exposure of animals to human refuse may have increased starch consumption, influencing amylase production (Perry et al. 2007, 2015; Axelsson et al. 2013) with a direct impact on the gut microbiota. Specific intestinal bacteria populations might have triggered the development of diseases through their collective metabolic activities and host interactions especially among susceptible hosts (Prakash et al. 2011; Lozupone et al. 2012). Another outcome of this post-domestication co-evolutionary process between hosts (human and animals) and pathogens (viral and bacterial) is the emergence of infectious diseases (EIDs) (Daszak et al. 2000). Domestic animals’ exposure to human waste has been associated with their introduction to new pathogens (Flint et al. 2016; Murray et al. 2016). The genetic diversity in wild populations enabled them to survive these pathogens and, in several cases, even to become their natural reservoir, for example Psammomys obesus and Procavia capensis reservoirs of Leishmania (Jaffe et al. 2004). For the ‘Big Four’, climate and anthropogenic changes are altering the distribution and incidence of infectious diseases, such as in the case of the bluetongue virus (BTV), foot and mouth disease (FMD) virus, African Swine Fever (ASF) and more. Recent outbreaks and the change in disease distribution have provided a reminder that livestock can quickly become exposed to the ravages of a new disease that takes hold because of changing conditions. A good example for a zoonotic disease with adverse effects on humans is Brucellosis (see Box 21 Brucellosis – a zoonotic disease, above).

BREED DEVELOPMENT

Once an animal was domesticated, human (artificial) selection for required traits continued, eventually establishing a range of breeds in each species. The process of animal evolution

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Box 21 Brucellosis – a zoonotic disease Brucellosis is one of the world’s major zoonotic pathogens, and is responsible for human morbidity and enormous economic losses in endemic areas (Boschiroli et al. 2001). Common zoonotic hosts for the Brucella bacteria are mainly Bovidae species such as goats (Brucella melitensis) and cows (Brucella abortus), but other forms are known such as Brucella suis from pigs, and Brucella canis from dogs. The close association between animals and humans provided an easy path for disease transmission. The source of human infection resides in domestic or wild animal reservoirs as humans do not maintain the pathogen. Routes of infection are multiple: foodborne, occupational or recreational and travel. New Brucella strains or species may emerge or existing Brucella species may adapt to changing social, cultural, travel and agricultural environments (Boschiroli et al. 2001; Godfroid et al. 2005). Therefore, at least from the Neolithic period to the present, an

evolutionary race between different strains of Brucella and the host immune systems has taken place. At least one case of brucellosis DNA has been found in the human remains from the offshore site of ‘Atlit Yam south of Haifa, Israel (see Box 13 ‘Atlit Yam, p. 71) dated to the Pre-Pottery Neolithic C period (ca. 9,000–8,500 cal BP). It stands out as one of the first cases where domesticated cattle remains found on-site may represent the agent of this disease (Kahila Bar Gal, unpublished). In addition, Brucella aDNA has been demonstrated in an adult human female with vertebral lytic lesions from Iron Age Siberia dated to 2,310–2,120 cal BP (360–170 BC; Bendrey 2008). Brucella spp. was also identified on skeletal remains of two adolescent males from the ancient Albanian city of Butrint, dated to the thirteenth to eleventh centuries CE. Anthropologists identified severe circular lytic lesions on the thoracic vertebrae, as well as porosity of the ribs. Molecular analysis based on the IS6501 insertion element and Bcsp31 gene confirmed that it was caused by Brucella (Mutolo 2006).

under domestication to accommodate farmers’ needs, consumers’ preferences and all biotic and abiotic factors affecting livestock survival and productivity is (by definition) still ongoing. Therefore, it may (and should) be seen as the evolutionary equivalent of postdomestication crop evolution processes in domesticated plants (Abbo et al. 2014). There is no strict definition, scientific or otherwise, of an animal breed. In 1999, the Food and Agriculture Organization of the United Nations adopted a broad definition of breed: (1) a subspecific group of domestic livestock with definable and identifiable external characteristics that enable it to be separated by visual appraisal from other similarly defined groups within the same species or (2) a group for which geographical and/or cultural separation from phenotypically similar groups has led to acceptance of its separate identity (FAO 2007). As of today, forty species and over 8,000 breeds of domestic animals are known (Yaro et al. 2016). Animal breeds were developed through selective pressures according to their functions and only those that adapted best to the environment, husbandry conditions and the demands of their holders have survived (Marsoner et al. 2018). The development of breeds, until the Industrial Revolution, was based on breeding animals targeting for valuable traits required by the farmers such as

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the ability to survive and produce under low food availability (Marsoner et al. 2018). Traditional breed development was a resource-driven activity bound to local cultural and environmental conditions – often focusing on multipurpose breeds. Today, livestock breeding targets highly specialized traits driven by demand for high-yielding breeds (Marsoner et al. 2018). Worldwide this ongoing process has led to rapid decline in the populations of indigenous breeds associated with geographical locations, respective cultures and environments. In the European Alps, especially Switzerland, over 80% of the indigenous farm animal breeds have become extinct (excluding poultry and dogs). For example, at first cattle breed development was through selection for aesthetic considerations, such as colour or horns, for various cultural and religious purposes and later selection for particular production purposes, including milk, meat and draught. During the eighteenth century the development of the ‘pedigree breeds’ raised the importance of the breed and combination of breeds, to create a new breed better suited to prevailing conditions. For example, during the twentieth century the goals in cattle farming were to increase production and efficiency to satisfy consumer markets that demanded an abundance of produce at low cost. In dairy cattle breeding the main aim of the last fifty years has been to improve production efficiency, with genetic selection focused on increasing milk yield. Indeed, milk production per cow has increased tremendously but it was accompanied by a declining ability to reproduce, increasing incidence of health problems and declining longevity of modern dairy cows – the consequence of negative genetic correlations between production traits, fertility traits and health in modern dairy cows (Oltenacu and Algers 2005). Hence, it is important to the dairy industry that animal welfare problems should be addressed to avoid widespread condemnation of breeding and management practices. W H A T M I G H T HA V E B E E N T H E M O T I V A T I O N F O R ANIMAL DOMESTICATION?

As there is no evidence of food shortage in the archaeozoological assemblages, what was the trigger that drove humans to domesticate animals? Various hypotheses have been raised to explain the will or need of humans to domesticate animals, climate changes being the most common explanation. The hypothesis suggests that during the Younger Dryas event (12,900–11,600 cal BP or somewhat earlier) the drier and colder climate caused resource depletion and foraging societies in the Fertile Crescent failed to adjust to the climate changes and looked for alternative food resources. This hypothesis was questioned because it was shown that humans were able to alter their behaviour and adapt to cyclic changes in their animal and plant resources (Rosen and Rivera-Collazo 2012; Roberts et al. 2018). The human responses to environmental change are thought to be based on embedded memory of situations of environmental stress and can last from years to generations. Human adjustment included flexible procurement strategies with changing resource availability such as altering between nuts and fruits to grass seed in times of environmental hardship (Rosen and Rivera-Collazo 2012). Unlike interpretations based on floral archaeobotanical

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remains, animal consumption was stable during the Younger Dryas event at many Near Eastern sites. Several animals (such as gazelle (Gazella gazella), deer (Dama mesopotamica), tortoise (Testudo graeca), hare (Lepus capensis) and partridge (Alectoris chukar)) adapted themselves to the changing environment and remained available to humans (for hunting) throughout the early and middle Epipaleolithic period (23,000–15,000 cal BP), throughout the early and middle Natufian culture of ‘Eynan in the northern Ḥula Valley (between 15,000 and 13,000 years cal BP; see Rosen and Rivera-Collazo, 2012, and Box 5 ‘Eynan, p. 31), and in late Natufian sites (mainly 13,000–12,000 cal BP and maybe somewhat later too) such as Mureybet (see Box 7 Mureybet, p. 41) and Tell Abu Hureyra (see Chapter 2) on the Middle Euphrates and in the late Natufian site of Nahal ‘Ein Gev II by the Sea of Galilee (Grosman et al. 2016). Furthermore, the utilization of the wildlife resources hypothesis – which assumes that continuous and accelerating hunting and trapping of wildlife reduced the number of available species, especially the ungulates, thus enforcing humans to find alternatives – was not supported either. Gazelles, fallow deer, hare and partridge have been hunted and are known in faunal assemblages throughout the PPN period (ca. 11,600–8,500 cal BP) and in decreasing numbers in the PN period (ca. 8,500–6,500 cal BP; see Chapter 2 and Box 14 Sha‘ar Hagolan and Nahal Zehora II, p. 75) in parallel to the _ presence of domesticated animals. Moreover, larger animals like hippopotamus, hartebeest, wild cattle and pigs also appear occasionally in the faunal records of Neolithic (and Chalcolithic and even Bronze Age) sites in varying (decreasing) quantities. Archaeozoological data from various habitats and landscapes in the Near East, especially the Levant, indicated that the first major ungulate extinction occurred during the local Iron Age (1,200–586 BCE; 3,150–2,536 cal BP), a period characterized by significant human population growth, rather than earlier (Tsahar et al. 2009). During that time the last of the largest wild ungulates, the hartebeest (Alcelaphus buselaphus), aurochs (Bos primigenius) and the hippopotamus (Hippopotamus amphibius), became extinct, followed by a shrinking distribution of forest-dwelling cervids (Tsahar et al. 2009). A second major wave of extinction of ungulates in the Near East occurred only during the nineteenth and twentieth centuries CE (Tsahar et al. 2009). Hence, the abundance of wildlife throughout the Neolithic period rules out this theory, too. Therefore, it seems that ‘push models’ as presented here (and see above in Chapter 3) do not provide a satisfactory explanation that may account for animal domestication. The motivation for animal domestication ought to be looked for in a cultural arena (and see Chapter 15, below, for a general view on ‘why domesticate, why agriculture?’).

K E Y PO IN TS A ND BE Y O ND

• All progenitors of the domesticated animals were present in the Near East (Fertile Crescent, Levant) and known to be hunted in Paleolithic times as observed in archaeozoological assemblages of the Epipaleolithic period (23,000–15,000 cal BP), and in

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•

• •

•

•

•

• •

• •

Natufian archaeozoological assemblages (15,000–ca. 11,800/700 cal BP) as well as in the early Neolithic period (prior to domestication). The geographical distributions of all livestock progenitors show a habitat overlap in parts of the Near East and west Asia, covering an area from central Turkey to the western parts of Afghanistan and from the Caspian coast to the Persian Gulf and the Indian Ocean – namely, an area of roughly 3,000 by 400–700 km. This area includes the northern part of the core area suggested for plant domestication. The sheep, goat and pig progenitors survived to modern times while those of cattle disappeared in the Neolithic period or later in time. Animal domestication as an integral part of the Agricultural Revolution in the Neolithic period is known only from the Near East and not in any other primary plant domestication centre in the world. Only at a later time were animals domesticated in other world regions, as for example in the case of the horse, domesticated in the central Asian steppe 5,500–4,000 cal BP (Gaunitz et al. 2018). Genetic and genomic databases tend to indicate a Near Eastern (Fertile Crescent, Levant) origin for the ‘Big Four’ (package) livestock. The ‘centre’ indicated by these studies encompasses the smaller in size Near Eastern plant domestication core area (Lev-Yadun et al. 2000). The ‘when’ question of animal domestication is difficult to answer. However, a date between 11,000 and 10,000 years cal BP would be acceptable to many of the involved researchers. In more accurate terms, this range could be limited to the 500 years between 10,500 and 10,000 cal BP. A thorough study of the archaeozoological evidence and the archaeological contexts from which it originates would certainly refine the time frame estimates. The archaeozoology of the Near East (Levant) provides data on the spread (out of the above-mentioned centre including south-east Turkey and northern Syria) of the ‘Big Four’ domesticated livestock, and likewise for the package of domesticated plants, and various cultural elements throughout the Near Eastern PPN interaction sphere. It is difficult to determine the pace of animal domestication as many of the DS characteristics are behavioural and are therefore elusive in the archaeozoological record. Animal domestication was an educated, knowledge-based process as far as the choice of species for domestication goes (from the naturally occurring fauna) and later selection of favourable (DS) traits. Enhancement of domesticated animal processes of improvement (breeding) continued over time and continue to this very day. Animal domestication influenced the health of both human and animal populations both because of the exposure to new pathogens and/or changes in pathogen abundance. Animals are not just meat (a dietary protein source): they provide secondary products that have become part and parcel of the human diet and economy including milk (with a genetic change in humans that enables digestion), dairy, hair/wool, hides, horn and, later, work (traction by bovids and transportation) and more. Milking started rather early and it is a significant element concerning Neolithic communities and demographics.

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• Livestock husbandry necessitated the domestication of many plant species utilized as feed. Some of these plants were grown mostly for freshly mown materials, or dry hay (e.g., alfalfa, clover), while others primarily served as a source of dense feed, that is, grain (e.g., bitter vetch, grass-pea). The species list includes (but is not limited to) clovers, vetches, alfalfa species, grass-pea and different grass species. • In many agro-eco-systems livestock rely (partly or exclusively) on grazing in rangelands. These pastoral systems are of different patterns including nomadic, sedentary and intermediate forms, each showing characteristic cultural and social adaptations. • In traditional societies where animal traction and transportation had a major role in the economy, a considerable percentage of the arable land had to be devoted to growing animal feed of different types including fibrous plant parts as well as starch and protein-rich feed materials, for example grains. • Post-World War II agricultural production in the industrialized nations was modernized by replacing (many millions of ) burden animals with motorized machinery. This enabled an increase in the areas devoted to grain production for human food into areas that had earlier been relegated to feed production. For example, the huge Canadian rapeseed oil (canola) industry mostly replaced oat production that had been required as animal feed. • Overall, livestock systems including the ‘Big Four’ species together with chicken, horse, donkey and camel play very important economic and socio-cultural roles for the wellbeing of human society. These systems occupy about 30% of the planet’s icefree terrestrial surface and are a significant global asset with a value of at least $1.4 trillion annually. The twentieth century has seen a new focus on a range of aquatic species (Duarte et al. 2007; Teletchea and Fontaine 2014; Teletchea 2015). Livestock production is likely to be increasingly affected by carbon constraints and environmental and animal welfare legislation. The demand for livestock products in the future could be heavily moderated by socio-economic factors such as human health concerns and changing socio-cultural values (Thornton 2010).

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P L A N T DO M E S T I C A T I O N AN D E A R L Y N E A R EASTERN AGRICULTURE Summary and Conclusions

In this chapter, we bring together most of the aspects discussed earlier, including biological, agricultural-agronomical and cultural facets related to plant domestication and the roots of Near Eastern agriculture. We briefly describe the spread of domesticated plants and the institutionalization of the agricultural system while discussing the historical and conceptual component underlying the Agricultural Revolution. Finally, we return to our lead question of why the Agricultural Revolution transpired. Let us revisit the issue of methodology and context of Near Eastern plant domestication research and researchers, to which we alluded in the introduction of this book. The data available to investigators of plant and animal domestication and the results of their analyses necessitate a professional look. Yet this is no easy task, as general explanations of plant and animal domestication are based on diverse schools of thought and research branches; scholars must thus remain open while being critical of the evidence before them. In our view, some of the reconstructions suggested for Near Eastern plant domestication would not stand a critical review; they rely on foreign and un-necessary assumptions far removed from the issue or on worldviews that hinder one’s ability to remain fully attentive to the data in its entirety while avoiding bias. It is important to probe deeper and understand these assumptions and worldviews and recognize that the communities that led and promoted plant (and animal) domestication and the Agricultural Revolution at large were groundbreaking pioneers, and it was these communities that first established the conceptual and socio-economic foundations of Western civilization as we know it today.

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P L A N T DO M E S T I C A T I O N : W H E R E A N D W H E N ?

Only one geographic area, rather limited in its range, is home to all wild progenitors of the grain crops domesticated in the Near East. This area is in south-eastern Turkey and northern Syria. • This area is characterized by a rich diversity of cereals and legumes as well as perennial vegetation comprised of deciduous oak trees, different types of pistachio, hawthorn, almond, pines and other trees. We therefore suggest that this (core) area is the cradle of plant domestication and the birthplace of Near Eastern agriculture. • This area also yielded the earliest evidence of domesticated cereals excavated at archaeological sites of the Neolithic period. • The core area hosts the wild stocks of einkorn and emmer wheat as well as the wild stocks of lentil and pea that exhibit a genetic profile that is closest to currently grown domesticated species. • The wild progenitors of flax and bitter vetch are also native to the core area. However, no wide-scale comparisons (of genetic affinity) of wild and domesticated germplasm of pale flax and wild bitter vetch were conducted. Therefore, at present it is impossible to infer the geographic origin of the wild stocks of these two crops. • The core area is also the homeland of the progenitor of chickpea, a wild species with a very limited natural distribution area, indicating that the chickpea could not have been domesticated anywhere but in this area. • Likewise, the natural ranges of the major livestock progenitors (goat, sheep, cattle and pig) domesticated in the Near East overlap in the proposed core area of southeastern Turkey. Following archaeological evidence, series of 14C dates and the archaeobotanical findings available from Near Eastern sites, it appears that plant domestication occurred approximately 10,500 years ago and this applies also to the approximate time of livestock domestication. Older, more ancient sites in this area yielded archaeological testimony of the gathering of the wild progenitors of domesticated plants. This is true, for example, at sites inhabited earlier than the Neolithic period such as Tell Abu Hureyra 1 and Tell Mureybet I (both boast late Natufian layers dated 13,000–12,000 years ago; see Box 7 Mureybet, p. 41), located on the banks of the Middle Euphrates. It is also true for early Neolithic sites in the area such as Jerf el-Ahmar, Tell Mureybet Layer III (the PPNA layer) and Dja’de in Syria or Çayönü in _ south-eastern Turkey (see Box 7 Mureybet, p. 41, and Box 11 Çayönü, Nevalı Çori and Tell Ḥalula, p. 64). Sites in the region inhabited starting ca. 10,500 years ago (the beginning of the early PPNB, e.g., Layer IV at Mureybet, Dja’de, Çayönü and Nevalı Çori; see Box 11 Çayönü, Nevalı Çori and Tell Ḥalula, p. 64) produced evidence of domestication, particularly manifested in cereals’ spike remains. Younger (later) sites in the Levant, and particularly in the areas west and south of the domestication core area, show that domesticated plants subsequently spread to other parts of the region (see Figure 15.1).

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Figure 15.1 Spatio-temporal spread of domesticated plants showing a ripples pattern from south-eastern Turkey and northern Syria. A colour version of this map can be found at www.cambridge.org/ abbo-gopher. T H E BO T A N I C A L - B I O LO G I C A L A N D A G R O N O M I C A S P E C T S

The domestication syndrome (see Box 16 Domestication syndrome, p. 116) comprises several botanical criteria (traits) by which we may distinguish wild from domesticated plants. The most important traits of the domestication syndrome in grain crops are seed dispersal and seed dormancy. Wild plants spontaneously disperse their seeds following maturation – cereals through shattered spikes and legumes through shattered pods from which seeds are scattered. In contrast, domesticated cereals exhibit non-shattering spikes while domesticated legumes have non-shattering pods. A thorough investigation into the nature of wild and domesticated cereals vis-à-vis wild and domesticated legumes shows that despite a superficial similarity between the two plant groups (stemming from the fact that in both domesticated groups the seed dispersal mechanism has been disabled), their biology is considerably different, pointing to free germination (non-dormant seeds) as the crucial trait for legume domestication. Whereas wild cereals are tall, aggressive plants originating in large, lush, wild populations and exhibiting approximately 50% seed dormancy, the wild Near Eastern legumes are compact, sprawling plants (other than the pea), growing in small, sporadic patchy populations and exhibiting seed dormancy rates of approximately 90%.

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We assume that due to these obvious biological differences between cereals and legumes as well as due to the uniqueness of each adopted species, domesticators were required to address each species individually. Here we would like to reiterate one of our basic assumptions – that domestication was based on knowledge with respect to seasonality of species, their nutritional value and their yield potential in the field. The Near Eastern crop package comprised three species of cereals (barley and two species of wheat), four species of legumes (pea, lentil, chickpea and bitter vetch) and flax, which was used as a source of fibre and perhaps also of oil. The three cereals facilitate flexibility within the farming system in terms of their response to the seasonal rain regime because barley matures early, einkorn wheat matures late and emmer wheat matures somewhere in between. Similarly, lentil and bitter vetch are small-statured plants so their seeds can mature even with limited precipitation whereas chickpea is sown in the spring and matures during the early summer. The Near Eastern crop package thus confers some protection upon farmers against drought in a region where the quantity and distribution patterns of rains varies annually. Additionally, a diet based on the combination of cereals and legumes provides all essential amino acids humans require. We stress the fact that both cereals and legumes were adopted for domestication in the biologically and culturally independent domestication centres of the Americas, Africa and East Asia. Following this understanding that farmers in all domestication centres had the sense to produce crop packages that were balanced both agronomically and nutritionally, it is clear why we believe that domestication was a knowledge-based initiative rather than circumstantial or the outcome of some ‘prehistoric ecological accident’, as some researchers suggest. The differential feeding behaviour of the Near Eastern domesticated livestock suggests that husbandry considerations (i.e., minimal competition for food resources, reproduction cycle and productivity) may have guided species selection as domestication candidates. T H E T I M E FR A M E A N D PA C E OF D O M E S T I C A T I O N

Some experts assert that domestication unfolded slowly (over thousands of years) in the early cultivated fields and was mostly an unconscious process in which domesticated types (such as non-shattering barley) gradually grew in prevalence due to selection pressures applied through harvest and seed handling activities. However, we believe that Neolithic farmers initiated a goal-oriented process guided by knowledge in which they selected the types they found desirable. They would have made their choice directly from among the great variability of wild populations in which they occasionally appear as mutants, as well as during the first years in their cultivated fields. In this respect, we believe that the approach by which domestication is described as a very long process that took place despite the farmers’ lack of awareness significantly undermines the extent of the achievement of farmers who, over 10,000 years ago and without the means available

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to us today, were able to domesticate the very same plants that today feed billions of people across the globe. We find it difficult to explain how it can be claimed that a protracted, random, automatic and unconscious process of plant domestication involved various ‘domestication experiments’ – some of which were successful while others failed – and perhaps also ‘candidate auditioning’ among various species, as researchers such as Bruce Smith, Dorian Fuller or Ehud Weiss advocate. By their very definition, experiments in specific wild species and their evaluation as potential crops would have necessitated premeditation, a careful selection of experimental species candidates, particular attention to results, and a process of inference. Thus, to the best of our understanding, the notion of unconscious experimentation is an oxymoron, and therefore any model that incorporates such ideas is logically flawed. In terms of the traceable (morphological – size and shape) differences between the wild and the domesticated livestock as evident from the archaeozoological record, the appearance of domesticated animals is abrupt. Contrary to the protracted domestication model, no transitional forms were identified in archaeozoological assemblages from the relevant strata in Neolithic sites. It is important to distinguish between the episode of domestication, which we regard as having been rapid, and other (later) changes that occurred in the plants and animals while under domestication. It is equally important, we claim, to distinguish between these two and the process of the institutionalization of agriculture as a whole socioeconomic system (see below), which indeed evidence shows had been much longer. FRUIT TREES

The domestication of fruit trees – grape vine, fig, olive, date palm and pomegranate – occurred after the domestication of the first annual crops. Here, too, the decisive factor was the biology of the wild species and the agronomic consequences involved in the unique traits of these species. The fruit trees we have discussed in this book are crosspollinating plants, that is, the pollen grains of one tree are carried on to the flowers of neighbour trees by the wind or insects where they fertilize their flowers. This is unlike the annual cereal and legume species, which are typically self-pollinating. In selfpollinating species, the isolation of desirable types (such as those exhibiting nonshattering spikes or free germination) is quite easily achieved because desired domestication traits are typically recessive, so offspring exhibit identical traits to the parent plants. In contrast, the cross-pollination mechanisms in fruit trees prevent the preservation of desired genetic combinations (found in parent trees) among their seedpropagated progeny. In other words, it is highly unlikely that a tree similar to the parent tree, which perhaps bears fruits that are sweet or high in fat content, will be found among its progeny. It is thus unsurprising that the first trees to have been domesticated in the Near East were those that lent themselves fairly easily to vegetative (clonal; see Glossary, Botany, Ecology and Agronomy, Clonal propagation)

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reproduction. In other words, once tree stocks were identified that bore desirable fruit, they could be reproduced by planting branch cuttings (in pomegranate, fig and grape vines), by removing shoots or suckers (in olive) or through side offshoots (in date palm). We emphasize the length of time required for fruit trees to reach maturity and begin to produce yield – five years or more without fertilization or irrigation – thereby requiring a greater degree of delayed returns compared to annual crops. It is therefore clear why we believe it is likely that tree domestication transpired within a more institutionalized agricultural environment and could not have occurred earlier than the domestication of annual crops. G E N E T I C DI VE R S I T Y I N D O M E S T I C A T E D P L A N T S A N D TH E F U T U R E O F FO OD PROD UCTI ON WORLDWIDE

Many domesticated species are characterized by a broad range of colours, shapes, flavours and qualities harnessed for different uses. These traits represent but a small fraction of all the traits essential for the functioning of plants in agro-eco-systems and for securing their yield. Yet we have shown that from the genetic perspective, the selection process that was the cornerstone of plant domestication involved a significant loss of genetic variability. This was due to the founder effect (see Glossary, Genetics, Genetic drift), in which only a portion of the genetic diversity that was available to the domesticators as they chose their desired stocks from among wild populations was sampled. This further intensified due to a continued selection process that occurred within farmers’ fields, and in the framework of public and commercial breeding programmes during the last century. This loss of genetic diversity led to the preservation of a fairly narrow segment of genetic diversity found in domesticated species compared to the wild populations of each species. Most scholars (ourselves among them) thus believe that it is highly important to effectively protect wild populations in large, sustainable nature reserves (and not only in deeply frozen seed collections) as a source of desirable traits that could serve the continued cultivation of these crops that are essential to the nurturing of humanity both now and in the future. During the twentieth century many traditional breeds of domesticated animals have been lost in many world regions as a direct consequence of the post-World War II rapid modernization and replacement of animal traction by modern machinery. Animals reproduce by cross-mating and therefore are highly heterozygous, contrary to most Near Eastern founder crop plants that are self-pollinators. Conservation of animal genetic diversity thus necessitates different measures. SPREAD AND INSTITUTIONALIZATION

As domestication transpired, and probably before agriculture was institutionalized, the knowledge, and perhaps also biological materials (seeds), began to spread throughout

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the Near East. The domesticated founder crops swept from their domestication centre (core area) in south-eastern Turkey and northern Syria westwards towards the Mediterranean coastal plains, Cyprus (with evidence of early domesticated plants and animals), as well as towards central Turkey (see Figure 2.1), the Balkans, the Danube Basin and further on into central and western Europe. Southwards the founder crops spread through central and southern Syria towards Israel and Jordan, while eastwards, they made their way towards Iran, Pakistan-Afghanistan to the western margins of the Indian subcontinent. This expansive development unfolded over a long period of time, and in several cases resulted in local adaptation and the domestication and adoption of local stocks. The exchange of plant species between Africa and the Near East did not occur until much later historical periods. Thus, after the rapid domestication that took place approximately 10,500 years ago in a well-defined region in the north of the Fertile Crescent, that is, in south-eastern Turkey and northern Syria, the idea took wing and the domesticated founder crops spread over a vast expanse, where domestication was institutionalized with its diverse expressions in the Near East, parts of western Asia, north Africa and the east African highlands as well as large portions of the European continent. Societies showing new life-ways and a new agriculture-based economy relying on domesticated plants and animals had taken hold in communities of Paleolithic (and Mesolithic) hunter-gatherers, who had previously lived in these areas, and became the heart and centre of what would later be known as Western civilization. Being an integral element of the Near Eastern farming package, the ‘Big Four’ livestock species have accompanied the grain crops in their spread across the Mediterranean Basin, into Europe and eastwards to central Asia, India and China.

H I S T O R I C A L A N D PE R C EP T U A L A S PE C T S O F P L A N T DO M E S T I C A T I O N AND THE AGRICULTURAL REVOLUTION

Plant (and animal) domestication and the transition from hunter-gatherer subsistence to a production economy were involved in revolutionizing socio-economic arenas, and, in fact, all walks of life. Following suggestions such as those offered by J. Cauvin (see Chapter 3), we regard the Agricultural Revolution, since its inception, as the manifestation of a new ideology that took root among human societies towards the end of the last Ice Age. At the heart of this transformation lay a new relationship between humans (culture) and the world (nature) that had divorced from the hunter-gatherer worldview, leading to a growing rift between humans and nature to the point of the alienation with which we are currently familiar. Underlying this change was the human ability to take control of natural species and adapt them to their new desires. Humans may have remained dependent on nature, as without natural resources the new subsistence economy could not exist, but their relationship with nature became irreversibly altered.

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High levels of environmental awareness involve, among other things, intelligent responses to changing circumstances. We thus do not claim that environmental or demographic changes did not to some degree influence the processes that led to the institutionalization of agriculture – surely they did. We do, however, argue that humans had a range of alternative choices, from which this one was made. Since we consider domestication as an educated, knowledge-led move of humans that reflected their extensive knowledge and competencies; and since domestication was a derivative of a cultural-perceptual transformation; and given that embracing this great new innovation was a complex, multifaceted process influenced by the conditions and circumstances prevalent at the time in the core area as well as any other area to which this innovation had spread – we find it difficult to accept that this transition had transpired as a result of external pressures or randomly. We have no unequivocal answers to the question of why this particular alternative was chosen. For example, what were the reasons that led to sedentary settlements? Did they play a significant role in the systems that later developed to become agriculture? And what led to the perceptual change in the relationship between humans and nature towards the end of the Pleistocene? What drove this ideological change? How did the social structure and organization change to the point that encouraged the institutionalization of agriculture? We are also unable to propose clear answers to questions such as why this particular manipulative direction was chosen, in which plants and animals were subjugated and production intensified? Why was another alternative, which did not necessitate the intensification of production, not chosen? Why had humans not developed other types of resources as a central subsistence resource? Although we have no answers to these challenging questions, there is no return from the choice that was made; the modern world in which we are currently living is the ultimate testimony to the apparent (evolutionary) success of this choice. Of course, some may claim that it might still be too early to determine whether this is indeed a successful achievement or whether we are headed towards an evolutionary failure due to the fact that we have yet to learn how to guarantee balances that can facilitate the sustainable existence of humanity. As to sedentary settlements, a fundamental element in the new relationship between humans and nature, we may say it has had a significant influence on the life-ways and landscape perceptions of humans. Clearly, the transition to sedentary settlements was a significant, highly influential process, for example, in the demographic arena and the infringement (or preservation) of equilibrium between humans and surrounding nature’s eco-systems comprising plants, animals and other natural resources. Undoubtedly, changes of this nature in settlement patterns and mobility of humans had impacted their social structure and organization, effectively being reflected in humans’ material world and expressed in a large range of symbolic manifestations that had attested to this change. About a quarter of a century ago, a discourse emerged involving the question of whether the perceptual and ideological changes, reflected among other aspects in

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changed symbolism, had occurred before sedentary settlements and plant and animal domestication or whether they were the result of these processes. In his book The Birth of the Gods and the Origins of Agriculture, Cauvin argued that the perceptual change (reflected in symbolism such as statues, imagery items or decorated vessels discovered among archaeological findings) preceded the economic change, and had, in fact, instigated change in other areas of life, and especially the growing divorce from nature. Cauvin also proposed that profound changes had taken place with regard to the supernatural and its perception as a driving force in the world or in nature. Therefore, the Agricultural Revolution rendered humans a unique species in nature – on the one hand, harmful and divorced from balance, and increasingly alienated from nature; while, on the other hand, humans had opened a window into expansive and thoughtful spiritual worlds, beyond the animistic perceptions (in their original meaning) that had characterized hunter-gatherers’ societies over many generations. All this has not transpired without social upheaval and required intensive social discourse and struggle that brought with it a whole new array of concerns and anxieties, part of which we carry to this very day. To conclude, plant and animal domestication and the institutionalization of the agricultural system brought about a new organization of all areas of human life. This was notable in gender relations, in which the equilibrium was reshaped between the reproductive (childbearing) element and productive (economic production) element of women; in the new division of labour among community members, which included the labour of youth and children and seasonal work plans; in the employment of many technological innovations as well as the emergence of processes that promoted and enhanced professional specialization; in numerous new activities that derived from grain crop production and later the growing of fruit trees and vegetables; in new industries that rested upon the novel worldviews such as the new lime plaster and clay industries that then evolved in the pottery industry – all industries that involve a pyrotechnical process in which energy (heat, fire) is used to irreversibly transform natural materials into new ones; and so forth (glass and metal working included). Plant and animal domestication had first and foremost served the need to exploit their primary potential as food sources for humans, replacing foods that had been gathered or hunted prior to the Agricultural Revolution. However, once domestication was achieved and agriculture institutionalized as a sustainable system, the system did not rest. Other than many ongoing processes aimed at improving domesticated species, humans also began to utilize new and additional plants (e.g., trees and vegetables) as well as secondary animal products such as milk, wool or labour (cattle, donkeys). In the early 1980s, the British researcher Andrew Sherratt designated these innovations as the Secondary Products Revolution. He assigned this revolution to the end of the Neolithic period, but, mainly, to the Chalcolithic and Early Bronze Age. In recent years, evidence has been accumulated – as well as corresponding arguments – that secondary products, at least of animal milk, were in use in early phases of the Agricultural Revolution while the domestication of some fruit trees and vegetables (including spices) had begun at different stages of the

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Neolithic period. Consequently, Gopher recently proposed calling the late institutionalization of Near Eastern agriculture the Second (Agricultural) Neolithic Revolution, which transpired during the Pottery Neolithic period and comprised the full institutionalization not only of the traditional Near Eastern agricultural farm and village but also of social change and the full divorce from the remains of the hunter-gatherer ethos towards a new era – an era of socially ranked societies, and, later, socially stratified societies in the Urban Era. WH Y DO MESTIC ATIO N, WH Y A GRIC ULTURE?

It is difficult to provide a decisive answer to the ‘why’ question of the Agricultural Revolution. That notwithstanding, our opinion in the matter is clear: we assign the revolution and its primary moves to the initiative and decision of humans with respect to social and cultural dynamics taking place towards the end of the last Ice Age and the early Holocene. It is unlikely that our view would be accepted by all researchers and thus end a long debate that has spanned many decades. The answer to the why question is inherently tied to the fundamental perceptions of human societies and the manner in which they operate and change over time. To a great extent, the perceptions of different researchers are derived from ideologies and schools of thought that are based on research and profound discussions on the history and sociology of human societies. It is thus improbable that this dialogue will come to an end any time soon, or even some time in the future. Nevertheless, in this book, we have presented readers with a broad range of evidence that is related to plant domestication, supporting the notion that plant (and likewise animal) domestication was a premeditated, knowledge-based and rapid move. This development went through the ‘sift’ of social debate, negotiations and struggles that eventually led to the institutionalization of a sustainable agriculturaleconomic system that thus lent full materialization to the Agricultural Revolution. Alongside the economic change, a ranked, complex social system crystallized, which shortly led to the emergence of cities and social elites (kings, princes, noblemen, rich men, priests, moral guardians and warriors) who effectively, despite upheavals, manage our lives to this very day.

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NOTES

INTRODUCTION 1 The terms ‘Agricultural Revolution’ or ‘Neolithic Revolution’ were coined by Gordon Childe in the early twentieth century. The term ‘revolution’ is indeed appropriate since, if we visualize the three million years of human and cultural existence as a clock, the Agricultural Revolution would be only a few minutes to twelve. Yet, focusing on these ‘few minutes’ at a higher resolution will make it possible to differentiate between the Agricultural-Neolithic Revolution as a long-term cultural and socio-economic transformation (also known as Neolithization) and short-term, specific events such as plant domestication. 2 From a long-term evolutionary point of view, caution should be exercised because it is as yet unknown where these developments – the Agricultural Revolution and our modern existence with its diverse agricultural and industrial organization – might lead. Many are of the opinion that it could lead to failure rather than success, especially with respect to the disruption of the Earth’s ecological equilibrium and the harm inflicted upon Earth as well as many plant and animal species, drawing us away from long-term sustainability. 3 In this context, we might add that the Agricultural Revolution is associated with demographic growth. Some claim that this growth instigated the revolution while the majority claim that it was the result of the revolution. Analysis of collected data seems to show that the most significant population growth started a short while (a few centuries) after plant domestication, reached a climax and then stabilized. Nevertheless, this is neither a direct nor a linear relationship, as this demographic growth occurred at an uneven rate, accelerating and decelerating at different times. Some demographic gain achieved through the domestication of plants (and animals) was overridden by the emergence and spread of disease that attacked sedentary and dense communities living in unsanitary conditions as well as diseases originating in nearby livestock (see some comments in Chapter 14). 4 Although it necessitates at least some prior knowledge of plants and their biology, as well as familiarity with the archaeology of relevant periods. 5 A growing trend in recent years is the assigned centrality of the human impact on nature, thereby replacing the term ‘Holocene’ with the term ‘Anthropocene’, which emphasizes humans’ role during this period. Farming and its consequences are listed as an important component of this impact. 6 All dates in this volume are presented in calibrated years before present (BP) based on the available radiocarbon record of relevant Near Eastern sites. 7 Key concepts are explained in text boxes 16 to 18 as well as in the Glossary at the end of this book. C HA P T E R 1 1 In Near Eastern Neolithic research, sites over 50 or 100 dunams (5 or 10 hectares) in size are called mega sites or towns despite being largely rural villages.

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2 Although it has become difficult to study hunter-gatherer populations as few, if any, such groups have maintained their original state, the current multifaceted field of hunter-gatherer research offers a broad range of study approaches that have long since broken through the barrier of stereotypes. Yet, they generally portray an unchanged picture of the hunter-gatherer ethos, which underlies our discussion.

C HA P TE R 2 1 Other Epipaleolithic cultures include the Mazraqan, Hamran, Mushabian and Ramonian and these are not presented in detail as this is beyond the scope of our discussion. 2 Such as the site of Kharane IV in Jordan, excavated by Professor Lisa Maher and her team, where the remains of a few amorphous-shaped semi-subterranean huts were found.

C HA P TE R 3 1 This relates to the selection of species and the biology of domestication as well as to the available archaeological resolution based on 14C dates. 2 Originally published in French under the title La Pensée Sauvage. 3 The commensal pathway as defined by Zeder (2012) or Larson and Fuller (2014). 4 It is of note that proponents of the Niche-Construction explanation to plant (and animal) domestication seem to assign human culture and action an important role in affecting the environment and thus the evolution of organisms (domesticates included) as well as their own evolution. Under this framework, the long domestication process is viewed as a co-evolving relationship (or partnership) between humans and plants or animals in which humans benefit due to their high (and unique) capabilities of learning and transmitting knowledge and information rapidly and changing their behaviour accordingly – used for enhancing fitness. This makes them successful players in the domestications arena. Yet, it seems that the economic cost-benefit (biological, evolutionary) element is governing the relationship for both humans and plants or animals. Thus, the term ‘Cultural Niche Construction’ accounts for this unique setting. Niche construction as well as cultural niche construction have never ‘sailed’ beyond the cost-benefit relationship between humans and plants (or animals) and statements of unconsciousness and no intentionality on the part of humans are repeatedly made. Thus, there is, in our view, a discrepancy between the highly praised human potential and the almost fully random co-evolutionary nature of domestication. Dynamics in perceptual/ideological or socio-cultural arenas that result in a mental and behavioural change (and action) are hardly discussed.

C HA P TE R 4 1 In 2018, charred remains of bread-like products, or ‘flat bread’, from the Natufian site of Shubayqa 1 in the Black Desert, northern Jordan, were reported by archaeobotanist Amaia Arranz Otaegui, the excavator Tobias Richter, and colleagues. This flat bread was made of wild cereals and some tubers and dated to some 14,000 years ago, thus pre-dating Neolithic plant domestication.

C HA P TE R 8 1 Recently, a few seeds, one of which shows the morphology of faba bean, were identified in a sample of legume seeds extracted from the Natufian site of el Wad Terrace, located on Mount Carmel in

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Israel. Carbon dating of one of these seeds to ca. 14,000 years ago prompted the authors to suggest they might have identified the elusive wild progenitor of faba bean. However, without a living plant species with genetic affinity to the domesticated faba bean, such a claim based on a single seed is rather doubtful.

C HA P T E R 9 1 Indeed, cereal grains were found in Paleolithic sites such as the 60,000–50,000 year old Kebara Cave and they were found in particularly large quantities in the 23,000 year old site of Ohalo II (see Box 3 Ohalo II, p. 25), suggesting they played a significant role in the diet of site inhabitants. 2 As noted in Chapter 5, Neolithic domesticators may not have defined their select domestication candidates as self-pollinating, and we cannot be sure that they could distinguish between the different pollination profiles. However, it is highly likely that their acquaintance with flora in general had taught them that some plants yield offspring that are similar to their parents while in other plants offspring are diverse with some very much resembling the parent plants and others less so.

C HA P T E R 10 1 Recent studies of barley genetics by the group of Terence Brown have eventually come to the conclusion that the population genetics of wild and domesticated barley indicate a ‘descent from a single founding population, which emerged in the western Fertile Crescent’ (see Civáň et al. 2021 in Further Reading).

C HA P T E R 11 1 See Jeremiah 1:11 (KJB): ‘Moreover the word of the LORD came unto me, saying, Jeremiah, what seest thou? And I said, I see a rod of an almond tree.’ 2 Residues of olive oil have recently been identified by D. Namdar and colleagues in a complete pottery vessel of the late Pottery Neolithic period (seventh millennium cal BP) at the site of ‘Ein Zippori (Lower Galilee, Israel), possibly the earliest olive oil found in the region.

C HA P T E R 14 1 See comments on the dog below; the cat is beyond the scope of this chapter, which focuses on the ‘Big Four’ livestock. 2 The IUCN Red List of Threatened Species, www.iucnredlist.org. 3 Dja’de in the Middle Euphrates Valley, in the the Early Pre-Pottery Neolithic B layer (EPPNB; 10,800–10,300 cal BP; Helmer et al. 2005); Çayönü in the High Tigris Valley, between the Early and Middle PPNB (around 10,200 cal BP; Hongo et al. 2009). 4 Akin to the ecological preferences of the Near Eastern plant, wild progenitors that are not campfollowers (see Chapter 3). 5 And see the negotiations between Jacob and Laban: ‘The speckled shall be thy wages; then all the cattle bare: and if he said thus, the ringstraked shall be thy hire; then bare all the cattle ringstraked’ (Genesis 31:8, KJB).

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F U R T H E R RE A D I N G

The list below comprises two groups of publications that relate to different aspects of plant domestication and the discussion presented in this book: • joint publications by the authors with many good partners, the most prominent of whom is Professor Simcha Lev-Yadun; • general publications and seminal works in the field of plant domestication, in particular in the Near East but also in other world regions, including key books and articles.

P UB LI C A T I O N S B Y S H A H A L A B B O , SI M C H A L E V - Y A D U N , A V I G O P H E R A N D O T H E R PA R T N E R S Abbo, S., Lev-Yadun, S. and Ladizinsky, G. 2001. Tracing the Wild Genetic Stocks of Crop Plants. Genome 44: 309–310. Abbo, S., Lev-Yadun, S. and Galwey, N. 2002. Vernalization Response of Wild Chickpea. New Phytologist 154: 695–701. Abbo, S., Shtienberg, D., Lichtenzveig, J., Lev-Yadun, S. and Gopher, A. 2003. Chickpea, Summer Cropping, and a New Model for Pulse Domestication in the Ancient Near East. The Quarterly Review of Biology 78: 435–448. Abbo, S., Berger, J. and Turner, N. C. 2003. Evolution of Cultivated Chickpea: Four Bottlenecks Limit Diversity and Constrain Adaptation. Functional Plant Biology 30: 1081–1087. Abbo, S., Lev-Yadun, S., Rubin, B. and Gopher, A. 2005. On the Origin of Near Eastern Founder Crops and the ‘Dump-Heap Hypothesis’. Genetic Resources and Crop Evolution 52: 491–495. Abbo, S., Gopher, A., Peleg, Z., Saranga, Y., Fahima, T., Salamini, F. and Lev-Yadun, S. 2006. The Ripples of ‘The Big (Agricultural) Bang’: The Spread of Early Wheat Cultivation. Genome 49: 861–863. Abbo, S., Frenkel, O., Sherman, A. and Shtienberg, D. 2007. The Sympatric Ascochyta Pathosystems of Near Eastern Legumes, a Key for Better Understanding of Pathogen Biology. European Journal of Plant Pathology 119: 111–118. Abbo, S., Zezak, I., Schwartz, E., Lev-Yadun, S. and Gopher, A. 2008. Experimental Harvest of Wild Pea in Israel: Implications for the Origin of Near Eastern Farming. Journal of Archaeological Science 35: 922–929. Abbo, S., Zezak, I., Schwartz, E., Lev-Yadun, S., Kerem, Z. and Gopher, A. 2008. Wild Lentil and Chickpea Harvest in Israel: Bearing on the Origins of Near Eastern Farming. Journal of Archaeological Science 35: 3172–3177. Abbo, S., Saranga, Y., Peleg, Z., Lev-Yadun, S., Kerem, Z. and Gopher, A. 2009. Reconsidering Domestication of Legumes versus Cereals in the Ancient Near East. The Quarterly Review of Biology 84: 29–50.

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Abbo, S., Lev-Yadun, S. and Gopher, A. 2010. Yield Stability: An Agronomic Perspective on the Origin of Near Eastern Agriculture. Vegetation History and Archaeobotany 19: 143–150. Abbo, S., Lev-Yadun, S. and Gopher, A. 2010. Agricultural Origins: Centers and Noncenters; a Near Eastern Reappraisal. Critical Reviews in Plant Sciences 29: 317–328. Abbo, S., Lev-Yadun, S. and Gopher, A. 2011. The Origin of Near Eastern Plant Domestication: Homage to Claude Levi-Strauss and ‘La Pensée Sauvage’. Genetic Resources and Crop Evolution 58: 175–179. Abbo, S., Rachamim, E., Zehavi, Y., Zezak, I., Lev-Yadun, S. and Gopher, A. 2011. Experimental Growing of Wild Pea in Israel and Its Bearing on Near Eastern Plant Domestication. Annals of Botany 107: 1399–1404. Abbo, S., Lev-Yadun, S. and Gopher, A. 2012. Plant Domestication and Crop Evolution in the Near East: On Events and Processes. Critical Reviews in Plant Sciences 31: 241–257. Abbo, S., Lev-Yadun, S., Heun, M. and Gopher, A. 2013. On the ‘Lost’ Crops of the Neolithic Near East. Journal of Experimental Botany 64: 815–822. Abbo, S., Zezal, I., Zehavi, Y., Schwartz, E., Lev-Yadun, S. and Gopher, A. 2013. Six Seasons of Wild Pea Harvest in Israel: Bearing on Near Eastern Plant Domestication. Journal of Archaeological Science 40: 2095–2100. Abbo, S., Pinhasi van-Oss, R., Gopher, A., Saranga, Y., Ofner, I. and Peleg, Z. 2014. Plant Domestication versus Crop Evolution: A Conceptual Framework for Cereals and Grain Legumes. Trends in Plant Science 19: 351–360. Abbo, S., Lev-Yadun, S. and Gopher, A. 2014. The ‘Human Mind’ as a Common Denominator in Plant Domestication. Journal of Experimental Botany 65: 1917–1920. Abbo, S., Zezak, I., Lev-Yadun, S., Shamir, O., Friedman, T. and Gopher, A. 2015. Harvesting Wild Flax in the Galilee, Israel and Extracting Fibers: Bearing on Near Eastern Plant Domestication. Israel Journal of Plant Sciences 62: 52–64. Abbo, S., Gopher, A. and Lev-Yadun, S. 2015. Fruit Domestication in the Near East. Plant Breeding Reviews 39: 325–377. Abbo, S. and Gopher, A. 2017. Near Eastern Plant Domestication: A History of Thought. Trends in Plant Science 22: 491–511. Abbo, S., Gopher, A. and Lev-Yadun, S. 2017. The Domestication of Crop Plants. In: B. Thomas, B. G. Murray and D. J. Murphy (Editors in Chief ), Encyclopedia of Applied Plant Sciences, Vol 3. Waltham, MA, pp. 50–54. Abbo, S. and Gopher, A. 2020. Plant Domestication in the Neolithic Near East: The Humans-Plants Liaison. Quaternary Science Reviews 240: 106412. Abbo, S., Peleg, Z., Lev-Yadun, S. and Gopher, A. 2021. Does the Proportion of Shattering vs. Nonshattering Cereal Remains in Archeobotanical Assemblages Reflect Near Eastern Neolithic Arable Fields? Review of Palaeobotany and Palynology 284: 104339. Ben-David, R., Can, C., Lev-Yadun, S. and Abbo, S. 2006. Ecogeography and Demography of Cicer judaicum, a Wild Annual Relative of Domesticated Chickpea. Crop Science 46: 1360–1370. Berger, J., Abbo, S. and Turner, N. C. 2003. Ecogeography of Annual Wild Cicer Species: The Poor State of the World Collection. Crop Science 43: 1076–1090. Frenkel, O., Shtienberg, D., Abbo, S. and Sherman, A. 2007. The Sympatric Ascochyta Complex of Wild Cicer judaicum and Domesticated Chickpea. Plant Pathology 56: 464–471. Frenkel, O., Sherman, A., Abbo, S. and Shtienberg, D. 2008. Differential Aggressiveness among Didymella rabiei Isolates Originated from Domesticated Chickpea and Sympatric Wild Cicer judaicum. Phytopathology 98: 600–608.

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Frenkel, O., Peever, T. L., Chilvers, M. I., Özkilinc, H., Can, C., Abbo, S., Shtienberg, D. and Sherman, A. 2010. Ecological Genetic Divergence of the Fungal Pathogen Didymella rabiei on Sympatric Wild and Domesticated Cicer spp. (Chickpea). Applied and Environmental Microbiology 76: 30–39. Gopher, A. 2012. The Pottery Neolithic in the Southern Levant – A Second Neolithic Revolution. In: Village Communities of the Pottery Neolithic Period in the Menashe Hills, Israel. Archaeological Investigations at the Sites of Nahal Zehora (Monograph Series of the Institute of Archaeology of Tel Aviv University 29). Tel Aviv, pp. 1525–1611. Gopher, A. 2020. The Neolithic Levant: Reflections on a Revolution and Deliberations on Then and Now. In: H. Khalaily, A. Re’em, J. Vardi and I. Milevski, eds. The Mega Project at Motza (Moza): The Neolithic and Later Occupations up to the 20th Century. Israel Antiquities Authority, Jerusalem, pp. 37–68. Gopher, A., Abbo, S. and Lev-Yadun, S. 2001. The ‘When’, the ‘Where’ and the ‘Why’ of the Neolithic Revolution in the Levant. Documenta Praehistorica XXVIII: 49–62. Gopher, A., Lev-Yadun, S. and Abbo, S. 2013. A Response to ‘Arguments against the Core Area Hypothesis’, for Plant Domestication. Tel Aviv 40: 187–196. Gopher, A., Lev-Yadun, S. and Abbo, S. 2017. Domesticating Plants in the Near East. In: Y. Enzel and O. Bar-Yosef, eds. Quarternary of the Levant. Cambridge, pp. 737–742. Gopher, A., Lev-Yadun, S. and Abbo, S. 2021. Breaking Ground, Plant Domestication in the Neolithic Levant: The Core-Area One-Event Model, the Sonia and Marco Nadler Institute of Archaeology, Occasional Publication No. 5, the Emery and Clair Yass Publications in Archaeology, the Institute of Archeology, Tel Aviv University, Tel Aviv. Heun, M., Abbo, S., Lev-Yadun, S. and Gopher, A. 2012. A Critical Review of the Protracted Domestication Model for Near-Eastern Founder Crops: Linear Regression, Long Distance Gene Flow, Archaeological and Archaeobotanical Evidence. Journal of Experimental Botany 63: 4333–4341. Kerem, Z., Gopher, A., Lev-Yadun, S., Weinberg, P. and Abbo, S. 2007. Chickpea Domestication in the Neolithic Levant through the Nutritional Perspective. Journal of Archaeological Science 34: 1289–1293. Lev-Yadun, S., Gopher, A. and Abbo, S. 2000. The Cradle of Agriculture. Science 288: 1602–1603. Lev-Yadun, S., Gopher, A. and Abbo, S. 2006. How and When Was Wild Wheat Domesticated? Science 313: 296. Lev-Yadun, S., Ne’eman, G., Abbo, S. and Flaishman, M. A. 2006. Comment on ‘Early Domesticated Fig in the Jordan Valley’. Science 314: 1681. Peleg, Z., Fahima, T., Krugman, T., Abbo, S., Yakir, D., Korol, A. and Saranga, Y. 2009. Genomic Dissection of Drought Resistance in Durum Wheat x Wild Emmer Wheat RIL Population. Plant Cell and Environment 32: 758–779. Tzarfati, R., Saranga, Y., Barak, V., Gopher, A., Korol, A. B. and Abbo, S. 2013. Threshing Efficiency as an Incentive for Rapid Domestication of Emmer Wheat. Annals of Botany 112: 829–837. van Oss, R., Abbo, S., Eshed, R., Sherman, A., Coyne, C. J., Vandemark, G. J., Zhang, H.-B. and Peleg, Z. 2015. Genetic Relationship in Cicer Sp. Expose Evidence for Geneflow between the Cultigen and its Wild Progenitor. PLoS ONE. DOI:10.1371/journal.pone.0139789. van Oss, R., Gopher, A., Kerem, Z., Peleg, Z., Lev-Yadun, S., Sherman, A., Zhang, H.-B. et al. 2018. Independent Selection for Seed Free Tryptophan Content and Vernalization Response in Chickpea Domestication. Plant Breeding 137: 290–300.

FU RTH E R R EAD IN G B Y OTHER A UTHO RS Aaronsohn, A. 1910. Agricultural and Botanical Exploration in Palestine. Bulletin Plant Industry 180: 1–63. US Department of Agriculture, Washington, DC, USA.

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G L O S S A R Y : BA S I C C O N C E P T S I N G E N E T I C S , B O T A N Y , E C O L O G Y , A G R O N O M Y AN D Z O O L O G Y G E NE T IC S

Allele. One of the alternative forms of the DNA sequence in any gene, which determines the expression of the controlled trait considering the dominant–recessive relations between the different alleles in the given locus (see below). Allosomes. The sex chromosomes of an organism, so called because often this pair of chromosomes do not have an identical morphology. For example, the human X and Y chromosomes vary in their size and respective gene content. The genes located on the sex chromosomes have a sex-linked inheritance (e.g., haemophilia in humans); see autosomes. Autosomes. The chromosomes that are not the sex chromosomes and appear in homologous pairs of similar morphology (size and centromere position) in somatic cells of eukaryotic organisms. The homologous chromosomes undergo a process of pairing during the meiotic division that leads to the formation of gametes (sex cells). The genes located on the autosomes follow a Mendelian inheritance, according to the chromosome segregation to the progeny (see allosomes). Cell differentiation. Plants boast special tissues, known as meristems, that maintain cell differentiation abilities throughout the plant’s lifetime (akin to animal stem-cells). These tissues produce cells that may become designated cells such as leaf, root, flowers, bark or other types of cells. Apical meristems are responsible for above-ground elongation and growth, while those found at root tips are in charge of the development of the root system. Proper development of the plant also necessitates the development of flowers at the adequate seasonal timing and in the right location in the plant. To this end, plants have numerous mechanisms, some of which are mentioned in this book, that determine the adequate timing of inflorescence development. Typically, the first step in developing flowers is the reception of an environmental signal (temperature or length of day), which is then followed by the instigation of a chain of control and expression of genes, ending with non-differentiated meristem cells transforming (differentiating) into a complex structure of the flower or inflorescence.

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Chloroplast. The cellular organelle containing the photosynthetic mechanism in green plants, granting plants their green colour due to the colour of the chlorophyll molecule. These organelles also contain circular DNA molecules that carry some of the information required to synthesize proteins that participate in the photosynthesis process. The chloroplast DNA sequence can serve to determine genetic proximity. With few exceptions, chloroplasts transfer from one generation to the next through female sex organs (maternal inheritance) whereas male sex cells (pollen grains) mostly do not carry any chloroplasts. Co-dominance. A condition in which the two alleles of a given locus are different but have an equal influence over the phenotype (see below). Dominant trait. An expression that is determined by a dominant allele compared to the alternative allele(s) in a given locus. For example, the allele that determines that the pea flower would be pink is dominant compared to that which produces a white flower; the allele that determines the shattering trait of the pea pod is dominant compared to that which produces a non-shattering pod. The expression of a dominant trait will also materialize when the organism bears one dominant and one recessive allele – a state known as heterozygosity (see homozygous below). Gene. The basic unit of heredity. A DNA sequence coding for a protein that has a certain function in the plant or a controlling sequence that governs the expression of other gene(s). A random change in the DNA sequence that changes the gene’s manner of operation or disables it altogether is a mutation (see mutation below). Genetic drift (genetic bottleneck, or founder effect). A phenomenon that occurs when a small group of individuals is removed from a large, diverse population, giving rise to a new population that is distinguished from the population of origin. Typically, the group of isolated individuals reflects only a small portion of the genetic diversity of the population of origin (see Glo. 1). Therefore, the genetic profile of the offspring population is expected to be different, and, in particular, less variable. The small number of individuals in the isolated population and the reduction in allelic variability may expose genetic irregularities due to the increased frequency of recessive allele combinations that may cause diseases. This is a known phenomenon among immigrants, for example among descendants of the Royal Navy vessel HMS Bounty rebels who settled on Pitcairn Island. This phenomenon is also known as a genetic bottleneck, and in many aspects we may regard the early seed sampling for the purpose of domestication as a private case of genetic drift. Genomics. A genome is an organism’s complete set of DNA, including all of its genes. Genomics is a scientific field focusing on the study of structure, function, evolution, mapping and sequence content of genomes.

GENETICS

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A wild population of a certain species that contains a wide range of genetic variation; a rich genepool.

Sampling of certain stocks for domestication results in a lesser degree of genetic variability in the population managed by the farmers.

Plant breeders are using a limited set of elite genotypes in their programmes, thereby leaving the modern cultivars genepool heavily eroded.

Glo. 1 Genetic drift. A colour version of this figure can be found at www.cambridge.org/abbogopher.

Genotype. The entire genetic information of the organism carried on its chromosomes with respect to the states of alleles in different loci (see locus below), gene sequences, number of chromosomes and so forth. In effect, the genotype is the overall genetic formula that determines the developmental potential of the individual. The actual expression of this potential is dependent on physical and biological environmental conditions that impact, in practice, its gene expression and hence the development of the individual. Haplotype. A haplotype is a group of DNA sequences (chromosomal segments) that for various reasons are transmitted together from parents to offspring. A haplogroup is a group of several such haplotypes, all of which have a common ancestor, that are inherited together. That is, they all share the same allelic variants at the respective DNA segments carried by different chromosomal regions.

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Heterozygous. A condition in which the two alleles in the locus are different, for example one is dominant while the other is recessive or the two are different but with a similar mode of action (see co-dominance above). Homozygous. A condition in which both alleles in the locus are identical (both are dominant or recessive). Lateral gene transfer. Regular mode of inheritance is by (vertical) transfer of genetic information from parents to offspring. Lateral gene transfer is the exchange of DNA by movement of DNA molecules from one cell to the other between unicellular organisms that are not necessarily closely related. Locus (plural: loci). The physical location of the gene along a chromosome. Mitochondrial DNA. The mitochondria are sub-cellular double-membrane organelles present within most eukaryotic organisms. Mitochondria convert chemical energy, via respiration, from food (carbohydrates) into a form that cells can use, thereby regulating the organisms’ metabolism. Like chloroplasts (see above) mitochondria carry circular DNA molecules that code for part of the proteins that function within the organelle. Mutation. A random change in the DNA sequence that alters the gene’s manner of operation or disables it altogether is a mutation. For example, a change that blocks pigment production and accumulation in the pea flower petals may result in white rather than pink flower colour (one of Gregor Mendel’s pea traits). Phenotype. The actual (external as well as biochemical and other intrinsic traits) appearance of the organism, which is determined by the genotype and the interaction effects between the genotype (all genes and any interactions among them) and environmental conditions that affect gene expression and the development of the organism. Ploidy. Describes the number of sets of chromosomes in the cells of the organism. A diploid plant bears two sets of chromosomes, one contributed by the maternal (egg) parent and another from the paternal (pollen donor) parent. The number of chromosomes in sex cells (gametes) is denoted by the letter n; in somatic tissues (see Somatic tissues below), regular cells will hold 2n. A range of levels of ploidy can be found in the plant kingdom, and some plant genera exhibit a ploidy series. The most familiar example is that of the wheat group, which includes diploid species of seven chromosomal pairs (2n=14), tetraploid species of fourteen chromosomal pairs (2n=28) and the hexaploid bread wheat of twenty-one chromosomal pairs (2n=42), and likewise in oat (Avena sp.). Among Near Eastern domesticated crops, only wheat exhibits this range of ploidy levels while other species are diploid. For plants that were domesticated in other world regions, polyploidy series are known among potato, beetroot, cotton and other species. Groundnuts also exhibit a ploidy range where wild species are diploid and the domesticated species is tetraploid. (see Glo. 2)

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A spontaneous crosspollination between the two species.

(see page 108).

Glo. 2 Polyploidy: definition and formation. A colour version of this figure can be found at www.cambridge.org/abbo-gopher.

Polymorphism. A condition in which variability is found among alleles of a certain trait, as opposed to monomorphism, where a single allelic version is found in a given locus within the populations of the organism. Qualitative trait. A trait that is determined by the activity of a gene that has strong influence over the phenotype, in most cases a single or only few genes. Due to the strong influence over the phenotype, individuals in diverse (polymorphic) populations can

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usually be classified into categories (discrete groups) with relative ease. For example, peas with white versus pink flowers or peas with yellow versus green seeds. Quantitative (metric) trait. A trait exhibiting a continuous phenotypic range that cannot be categorically classified into discrete groups. Classifying individuals by traits that are phenotypically continuous requires repeated measurements of the phenotypic values and then the calculation of averages and statistical analyses using adequate statistical tools. Due to the requirement of measurement (unlike the simple classification of qualitative traits; see above), these traits are known as metric. For example, seed weight in most cereal and legume crops is a quantitative, continuous trait: in a diverse population, individuals will exhibit different weights, the frequency of which is distributed in a typical bell-shaped (Gaussian) curve. Quantitative traits are governed by more than one or two genes, sometimes up to several dozen genes. Most of the traits that are important from the agricultural perspective (e.g., seed yield, quality and weight) are, by definition, quantitative because they express the developmental outcome of the function of many genes throughout the plant’s growth period. Genes that influence a quantitative trait are collectively called polygenes due to the multiplicity of genes involved in determining the phenotype; accordingly, a quantitative trait is also known as a polygenic trait. Recessive trait. An expression determined by a recessive allele that functions recessively compared to alternative allele(s) in a given locus. For example, the allele that produces a non-shattering pod in peas or lentils is recessive whereas the allele that produces shattering pods in these species is dominant. Thus, to produce the phenotype of a recessive trait, a homozygous state (see homozygous above) is required in the given locus; that is, a recessive trait will be expressed only in organisms bearing two recessive alleles.

B OT A NY , E CO LO G Y A N D A G RO NO M Y

Clonal propagation. Vegetative reproduction that does not involve sexual reproduction. The offspring of vegetative propagation are identical to the parent plant. The reproduction of many fruit tree species involves this type of reproduction based on branch segments or shoots, whereas in many ornamental plants, vegetative reproduction is usually achieved through such cuttings and other organs such as tubers and bulbs. Ecological niche. A term that describes the overall physical and biological factors that influence organisms (flora, fauna and microorganisms) in their habitat. In the context of the current volume, the term is mostly used to describe the bedrock and soil environment and climatic regime found in the habitat of the plants discussed. Typically, plants growing in lush habitats (deep, fertile soil, excessive precipitation, temperate temperatures) will be absent from poor habitats, in which the soil is shallow or precipitation is limited. Plants that favour poor habitats will be absent from lush habitats, as they typically cannot withstand competition with species typical of lush and fertile

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habitats. Species that grow side by side are in constant competition in natural habitats, yet they also share positive interaction relationships, for example shade-loving species will grow in the shadow of taller plants; or more complex relationships such as symbiosis, for example between legume root cells and nitrogen-binding bacteria. Over evolutionary time periods, in a process known as succession, each ecological niche develops vegetation (as well as faunal populations) typical of the conditions and carrying capacity of the terrain (in the Mediterranean Basin the most important factor determining carrying capacity is water availability). Normally, any intervention in such vegetation – by chopping, burning, partial or full removal, or grazing animals – will lead to changes in the composition of species in that habitat. For example, an intensive grazing regime will increase the prevalence of thorny plants, which animals refrain from eating as they focus on non-thorny species. Similarly, moderate, selective removal of the yield of certain plant species will generate change due the reduction of seed reservoirs of these species, opening the arena to other species. A cultivated field is also an ecological niche, albeit man-made, and will exhibit deliberately sown crop species and what we call weeds alongside their pests, symbionts, disease agents and so forth. Similarly, other areas suffering from human activity (grazing lands, roadsides, dumps, quarry or mine waste areas) are populated by species that can handle such conditions. Typically, species’ composition in habitats disturbed by humans differs from that of more balanced, or less disturbed, habitats. In the Mediterranean region of Israel, mallow, wild mustard and chrysanthemum show an ecological affinity to disturbed habitats. Field plant stand. The number of individual plants growing on a single unit of land. While the term usually refers to cultivated fields, it may be applied to wild populations as well. For all practical matters, the term represents the density of the target plant in a cultivated field (for the farmer) or the wild population (for the hunter-gatherer). Habitat fragmentation. A process by which an ecosystem with its native biota (plants, animals, microorganisms) is broken up into smaller, isolated patches of habitats. Consequently, many of the native organisms can no longer roam their former (larger and contiguous) home range. As a result, some populations survive as islands and can no longer participate in gene exchange with neighbouring population patches (thereby experiencing genetic drift; see above). In modern times, the major driver of habitat fragmentation is human development: for example, building, road construction, canal digging, mining, deforestation and the like. Juvenile period. The period in the plant’s life during which it does not yet develop reproductive (sex) organs but rather somatic tissues (see below), which are asexual. During this time, the plant builds its canopy, accumulating its reservoir of nutrients and photosynthetic products that are required for the reproductive process. In annuals, this

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period lasts a few months; however, in many perennials this period may last several years, contingent on growth conditions, where abundant settings will shorten this period and resource limitation may prolong this period. Monoculture. An agricultural system in which few plant species are grown, where each plot hosts a few or even a single cultivar over a relatively large acreage. This condition is typical of industrialized systems characteristic of modern times, in which uniformity is required from the perspectives of sowing and harvest timing, and the quality of produce designated for industrial processing that demands a stable and predetermined quality as much as possible. Pioneer plants. Plant species capable of colonizing newly cleared (recently disturbed) habitats. Such species usually form the first stage in the secondary succession of natural plant communities in ecosystems disturbed by landslides, fire, volcanic eruptions or human activity. If left undisturbed, the pioneer taxa are usually replaced by later succession species forming a more diverse plant formation. Identity of pioneer species will depend on the disturbed ecosystem and environmental conditions. For example, pioneer taxa colonizing abandoned agricultural fields will differ from those following a landslide in rocky terrain. Polyculture. The traditional alternative to monoculture (see above). It is commonly assumed that polyculture was the prevalent condition since the inception of agriculture and until the modernization of agricultural systems starting at the end of the nineteenth century. In this system, growers cultivate a broad range of crops and various cultivars of each to satisfy most of their needs, while relying only to a limited degree on trade and import from distanced areas. Under these conditions, disease and pests spread more slowly because among traditional landrace (cultivar) populations, some variants will show resistance. Variability in the developmental rate of heterogeneous traditional cultivars in such systems confers adaptability to climatic threats such as drought or cold events. This is because in such a system – unlike the system in which all plants are uniform – some plants are early while others are late to develop, allowing the population to sustain only partial injuries in the face of climatic calamity (or disease or pests’ epidemics). Selective nutrition. Individuals exhibiting unique preferences, in this case with respect to nutritional choices, based on flavour, scent or cultural conditioning (such as kosher, or vegetarian-vegan practices). Somatic tissues. In plants, typically those tissues that do not develop reproductive (sex) organs. Sympatric and allopatric distribution. These terms describe the physical proximity of plant (or animal) species in natural habitats. The term ‘sympatric distribution’ describes the condition where two or more species are found in great physical proximity, whereas

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the term ‘allopatric distribution’ describes the condition where said species do not inhabit the same habitat, or even the same geographic area. Traditional cultivar (landrace). A stock of a crop plant species that was maintained by farmers and passed from generation to generation, without any involvement of modern breeding or seed companies. Nearly all domesticated crop cultivars that persisted throughout the world prior to the advent of modern plant breeding by the second half of the nineteenth century were traditional cultivars.

Z OO L OGY

Adrenocorticotropic hormone (ACTH). A polypeptide (protein) hormone produced and secreted by the anterior pituitary gland (the hypophysis). It is important in the animal hormonal system and is often produced as a response to stressful conditions. Archaeozoology. The study of archaeological remains of animal origins (faunal remains). Eukaryotes. Organisms whose cells have a membrane-enclosed nucleus that contains most of the cell’s genetic material. Eukaryotes also possess other membrane-bound organelles such as mitochondria in their cells (see above). Heterochrony. A developmental change leading to modifications in size and shape of organs. This is different from structural (form) innovation created by change in the position of certain tissues. Ontogeny. The sequence of developmental events that occur during the lifetime of a living organism (from the time of fertilization of the egg to the organism’s mature form). Paedomorphosis. The retention of juvenile traits by the adult animal. Prokaryotes. Unicellular organism that lacks membrane-bound organelles such as a nucleus and/or mitochondria. Only the single-celled organisms of the domains Bacteria and Archaea are classified as prokaryotes.

GENERAL TERMS

The Agricultural Revolution. A comprehensive cultural, socio-economic process (or transformation) that brought about overarching changes in human perception and that was a cultural turning point in human history. It took place in Near Eastern Neolithic societies during the early Holocene period and instigated the emergence of rural, sedentary societies of food-producers. Plant and animal domestication is a key

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component of this revolution as were many innovations found in diverse fields: architecture, technology, burial customs, art and symbolism, and structural and organizational changes in society and subsistence. Archaeologists tend to view this revolution as a multifaceted process known as Neolithization, which climaxed with the institutionalization of a sustainable agricultural system and a specializing, ranked and later stratified society that soon became urban. Anti-nutritional factors. Compounds present in various plant foods (e.g., Trypsin inhibitors) that have adverse effects on proteolytic enzymes (which break down proteins into their animo acid building blocks) or interfere with the solubility of certain minerals (e.g., phytic acid). Thereby, those anti-nutritional factors reduce the efficiency of the digestive processes as well as the absorption of nutrients. Cultivation: plant husbandry. Scientific literature on plant domestication often refers to the term ‘cultivation’ as a behavioural corridor leading to domestication. In the current volume, we did not discuss this term or archaeological evidence that attests to such behaviour, and neither did we mention the debate on the use of this concept. Briefly, we can say that according to our understanding, this term defines the agricultural niche, differentiating it from the natural wild state – that is, overall activities carried out by growers in their fields and farms that might influence yield or crops’ population composition. These activities include working the soil, the elimination of weeds, unique selection of seeds for growing, sowing, pest and disease control (especially in modern times), fertilizing, pruning, harvesting, seed cleaning, and storing of sowing matter for the next season. Due to the ongoing need to decide which actions must be carried out and when, and the continued assessment of effectivity as the basis for further farm operations, in our view cultivation cannot be considered unconscious in any respect. Similarly, we cannot advocate cultivation as a prelude to domestication lasting several millennia during which farmers were unable to distinguish shattering from nonshattering wheat spikes as argued by researchers such as George Willcox and Dorian Fuller. Plant domestication. Plant domestication was only one element of the Agricultural Revolution: introduction of plants into human systems and their control. The term ‘plant domestication’ harnesses both biological and cultural aspects. In botanical or genetic terms, a plant is considered domesticated if it exhibits morphology and physiology that differ from those commonly found among wild populations and which were selected by humans to facilitate the application of an orderly growing regime (cultivation: plant husbandry; see above) to plants for the purpose of producing food, feed, fibre or other materials. For example, domesticated wheat and barley exhibit a spike that does not shatter upon maturity – a trait that is rarely found among wild cereals. This quality hinders the plant’s survival in nature but offers great advantage to the farmer. Similarly, domesticated chickpea, pea and lentil exhibit free germination (lack of seed

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dormancy), a trait that hinders the plant’s survival in nature, yet without which it would be agronomically impossible to produce profitable yield from these species. From a cultural perspective, plant domestication is an event in which humans select seeds (reproduction matter) of wild plants based on the knowledge and recognition that those seeds bear desirable traits (desirable flavour, nourishing nutritional composition, free germination and so forth) in order to subject them to a growth regime targeted at different, predefined purposes (producing foods or raw materials such as fibre, dye or oils).

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INDEX

Aaron Aaronsohn, 111, 186 abscission zone, 136 adaptation, 115, 161 bodily, 16 change, 184 cultural, 83 local, 181, 185 wild, 148 agricultural compensation, 152 agricultural package, 156 Agricultural Revolution, 11, 29, 43, 68, 74, 78, 80, 83–84, 86–87, 229, 235, 237 Ideological model, 92 why?, 238 Ain el-Kerkh, 143 Ain Ghazal, 59, 61, 204, 221 almond, 170 domesticated, 128, 173 wild, 116, 170 animal breed, 224 traditional, 225 animal domestication, 38, 59, 199, 210, 216, 219, 229 motivation, 226 species-specific traits, 218 animal evolution under domestication, 224 apple, 174 ‘Anna, 176 architecture, 64, 221 circular, 52–53 rectangular, 42, 52, 59, 61 Ascochyta blight, 152 ‘Atlit Yam, 71, 171, 224 aurochs, 205 automatic selection, 85, 94, 162 autonomous protracted model, 163 barley, 106–107, 213 domestication, 191 wild, 140, 157

Bar-Yosef Ofer, 27, 45, 68 Bezoar, 204 goat, 204–205, 209, 215 bitter vetch, 106, 109 boar, 206 wild, 206 Bolivia, 190 Bos indicus, 207, 211 primigenius, 205, 218 taurus, 207, 211 Braidwood, Robert, 86 bread flat, 107 wheat, 108 broad bean, 139 cultivars, 143 broccoli, 180 Brucellosis, 223 burials, 27, 33, 40, 50, 57, 61, 66, 69, 71 group, 31 cabbage, 180 wild, 183 Capra aegagrus, 204 hircus, 206 cattle, 199 dairy, 225 domestic, 207 domestication, 211, 218 introgression, 213 cauliflower, 180 Cauvin Jacques, 41, 67, 92, 235, 237 Çayönü, 64 Central America, 190 Central Asia, 189

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centre of domestication, 84 centres of origin, 189, 192 cereal, 135 cool-season, 114 cultivated fields, 141 domesticated, 231 domestication, 133, 141 yield, 152 Charles Darwin, 183 chickpea, 106, 108, 116 cultivars, 129 domesticated, 129, 150 wild, 129 Chile, 190 China, 189 Cicer reticulatum, 108 climate change, 85, 197, 199 clonal reproduction, 169 commensalism animal, 219 Competitive Feasting Theory, 92 conscious selection, 216 core area, 44, 67, 81, 143, 161, 230, 235 core area-one event, 163 core-area-single-event model, 81 crop evolution, 132, 224 under domestication, 160 crop improvement, 179 crop package, 75, 135, 232 crop variability centres, 191 crop wild relatives, 198 cross-pollinating, 233 cross-pollination, 139, 143, 166, 171 cultivation, 44, 89, 91, 154 centres, 162 cultural interaction sphere, 67 date palm, 166, 170, 174 delayed return, 177, 234 demographic expansion, 86–87 dispersal unit, 136 division of labour, 237 dog, 31 domestication, 219 domestic animal, 216 domesticated animals, 61, 71 abrupt appearance, 221 founder package, 217 wild progenitors, 201 domesticated herds, 63 domesticated plants, 61, 91

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earliest, 157 evidence, 65 origin, 188 populations, 180 spread, 229 domestication, 13 auditioning, 233 candidates, 119, 147, 150, 177 centres, 191, 194 cereal, 94 conscious, 217 episode, 128–129, 132, 139, 233 Ethiopia, 185 immediate, 96 independent, 160 initial, 162 legume, 95 mutualistic, 220 origins, 188 preadaptation, 217 protracted, 94 random process, 177 rapid, 94 secondary, 160, 172 single episode, 156 single event, 160, 163 standard, 133 traits, 117, 119 unconscious, 218 domestication centre, 235 independent, 191, 232 domestication islands, 218 domestication syndrome, 115–117, 122, 128–129, 133, 216, 231 domestication traits, 154 crucial, 128, 133 Dump-Heap Hypothesis, 83, 87, 92, 147, 153 durum wheat, 106, 108 einkorn wheat, 106–107 origin, 161, 163 emmer wheat, 132 domesticated, 107–108 wild, 140, 186 environmental awareness, 236 Epipaleolithic period, 21 settlements, 22 Ethiopia, 190–191 Eynan, 31, 219, 226

faba bean, 142 farm management, 177 farming package, 235 Fertile Crescent, 21, 73, 100, 156–157, 195, 205–206, 209, 225, 235 fig, 166, 174 domestication, 171 pollination, 167 seedless, 171 fishermen, 71 flax, 106, 110, 144 fibre, 144 food-producing farmers, 14 founder crops, 75, 107, 135, 141, 153, 163, 172, 181, 221, 235 choice, 193 founder effect, 179, 184, 234 free germination, 139 fruit trees, 175 clones, 173 cultivars, 173 domestication, 174, 177–178, 233 founders, 166 genetic variability, 183 juvenile period, 176–177 selection cycles, 174 wild populations, 171 fungi growing ants, 91 G6PD enzyme, 143 Garden of Eden, 18–19 gender relations, 237 gene banks, 197 gene flow, 172, 184, 218 post domestication, 213 genetic bottleneck, 179, 184 genetic contamination, 172, 185 genetic diversity, 114, 160, 179, 183–184, 189, 210, 223, 234 animal, 234 decrease, 185 loss, 234 reduction, 179 genetic variability, 197 loss, 234 genetic variation, 173 genomic diversity, 213 Gilgal I, 45, 171 giving environment, 18–19 global food production, 84

goat, 199 domestic, 206 domestication, 209 wild, 204 goat grass, 108 Göbekli Tepe, 53 grafting, 170 grain protein, 141 grape vine, 166, 174, 177 green revolution, 196 groundnut, 154, 185 growth habit, 141, 152 determinate, 137 indeterminate, 137 habituation animal, 219 ibex, 220 Hallan Çemi Tepesi, 205 Harlan J. R., 149 Helbaek Hans, 107 Hillman Gordon, 37 hog, 206 Holocene, 22 Hordeum spontaneum, 107 human agency, 94 hummus, 151 hunter-gatherers, 14, 80, 82, 153, 177, 201, 235 ethos, 18, 74, 93 knowledge, 83, 96 last, 55 society, 82 imagery items, 33, 40, 43, 71, 75, 93, 221 art, 50 incipient agriculture, 86 India, 189 infectious diseases, 223 institutionalization of religion, 93 introgression, 184, 213 Irano-Turanian vegetation, 101 Jericho, 38–39, 45 tower, 47, 50 Jordan Valley, 50, 59, 75 Karacadağ, 161, 163 Kebaran culture, 27

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Khiamian culture, 38, 41, 44, 50 knowledge transmission, 82 knowledge-based selection, 218 kohlrabi, 180 Kurdistan, 100 Iraqi, 86 Ladizinsky Gideon, 95, 108 land designation, 174, 177 landraces, 128, 132, 135, 179, 181, 184 traditional, 196 legume, 135 domesticated, 231 domestication, 141, 150 field, 141 populations, 140, 148 wild, 148 yield, 152 Lens culinaris, 108 lentil, 106, 160 cultivars, 184 domesticated, 107 soup, 150 wild, 95, 111, 157 Levant, 21, 52, 195, 199, 226 interaction sphere, 195 northern, 64, 67, 161, 172 southern, 73, 161 Lévi-Strauss Claude, 82 linseed, 106 Linum bienne, 110 livestock domesticated, 199 domestication, 230 progenitors, 230 spread, 235 logistic mobility, 29 macadamia nut, 194 maize, 119 Marginal Zone Theory, 86 marsh elder, 191 Mediterranean climate, 100 diet, 150 vegetation zone, 148 zone, 100, 107 Mesolithic, 206, 235 mixed farming, 163

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modern agriculture, 124 fragility, 197 mouflon, 204 Mureybet, 41 Mureybetian culture, 38, 41, 50 mutation, 84, 94, 118, 147, 166, 171, 175, 182–183, 186 beneficial, 182 Nahal Ḥadera V, 27 _ Issaron, 68 Nahal _ Nahal Zehora II, 75 _ Natufian, 230 economy, 35 plant foods, 35 Natufian culture, 24, 37, 45, 205 Near East, 21, 195 Near Eastern agriculture, 229 nectarine, 183 Neolithic farmers, 232 Neolithic period, 38 Netiv Hagdud, 45 Nevalı Çori, 64 Niche-Construction approach, 91 model, 90 Niche-Construction Theory, 91, 147, 220 nitrogen fixation, 140 non-shattering, 115 types, 94 Oasis Theory, 85 oat, 213 obsidian, 31, 40, 45, 61, 195 Ohalo II, 23, 25 olive, 8, 166, 168, 170–171, 174, 233 domestication, 172 independent domestication, 172 oil, 178, 241 orchards, 174 pollen, 171 wild, 172 orange Jaffa, 183 Ovis aries, 207 orientalis, 204 Paleo diet, 16 Paleolithic nutrition, 16

Paradise, 19 Paraguay, 190 park forest, 107 pea, 106 domesticated, 108, 131 domestication, 132 wild, 95, 111, 132 pig, 199, 208 domestication, 211 independent domestication, 211 introgression, 213 wild, 206 pistachio, 103 groves, 109 plantations, 89 Pisum humile, 108 plant breeding, 179, 185, 189 modern, 195 plant domestication, 65, 67, 80, 90, 184, 229–230, 238 evidence, 59 geography, 156 in America, 88 models, 81, 83 research, 229 research perception, 80 plant evolution under domestication, 132 pod shattering, 115, 131 pomegranate, 166, 174 flowers, 168 population growth, 59 post-domestication, 223 Pottery Neolithic, 73 PPNC, 59 Pre-Pottery Neolithic A, 38 Pre-Pottery Neolithic B, 40 sites, 59 preemptive domestication, 162 proto-domestication, 219 protracted domestication, 129 process, 95 protracted-autonomous model, 81 Pulse Domestication before Cultivation, 95 punctuated equilibrium, 220 residential mobility, 29 ruminant species, 208 rye, 119

secondary animal products, 237 Secondary Products Revolution, 76 sedentary settlement, 32 emergence, 87 sedentary settlements, 37 sedentary villages agricultural, 86 seed dispersal, 113, 116, 231 mechanism, 132 seed dormancy, 95, 113, 116–117, 138, 231 legume, 95 seed-bank, 189 self-pollinating, 118, 155, 233 self-pollination, 139, 166 serotonin, 151 Sha’ar Hagolan, 75 sharing, 14, 18, 97 sheep, 199 domestic, 207 domestic breeds, 207 domestication, 210 wild, 204 shepherds, 63 sickle blades, 33, 38, 40, 43, 50, 55, 61, 71, 75 absence, 69 Sinai, 59 slash and burn, 126 South America, 190 sowing and harvesting regime, 94 spike shattering, 132 subsistence strategies, 90 continuum, 91 Sultanian culture, 38, 45, 47, 50 Sus scrofa, 206, 208 sustainability, 18, 82 symbiotic relationship, 148 Tell Abu Hureyra 1, 230 Tell Ḥalula, 64 Tell Mureybet I, 230 The Nuclear Zone Theory, 86 traditional agricultural systems, 123 traditional breeds, 234 traditional cultivars, 128, 181, 185 traditional farming, 152, 175–176, 182, 197 communities, 181 subsistence, 126 traditional farming systems, 125 tree dioecious, 166 domesticated, 171

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Triticum boeoticum, 107 Triticum dicoccoides, 107, 132 tryptophan, 116, 150 unconscious selection, 95, 153, 163 ungulate extinction, 226 urbanization, 11, 79

Yarmukian culture, 75 yield compensation, 152 stability, 124, 163 Younger Dryas, 35, 225

Vavilov, Nikolai, 189 vernalization, 130 Vicia ervilia, 109 Wadi Tbeik, 68 weed, 73, 88 invasive, 89 species, 89

270

Western civilization, 154, 163, 229, 235 wheat, 213 domestication, 132 wild, 132, 148 wild progenitors, 107 populations, 186 woodland, 107, 110, 201, 206

Zohary Daniel, 85, 107, 118, 174 zoonotic disease, 223

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