handbook of plant palaeoecology · the late glacial and holocene. subfossil plant remains offer us...

Handbook ofPLANT PALAEOECOLOGY

GRONINGEN ARCHAEOLOGICAL STUDIES

VOLUME 19

EDITORIAL BOARDProf. dr. P.A.J. A� emaProf. dr. R.T.J. Cappers

Prof. dr. L. HacquebordDr. W. Prummel

Prof. dr. D.C.M. RaemaekersProf. dr. H.R. Reinders

Prof. dr. S. Voutsaki

GRONINGEN INSTITUTE OF ARCHAEOLOGY

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Handbook ofPLANT PALAEOECOLOGY

R.T.J. CappersGroningen Institute of Archaeology

University of Groningen

R. NeefDeutsches Archäologisches Institut

Berlin

BarkhuisGroningen University Library

Groningen 2012

Photomicrography: Judith Jans & René T.J. Cappers

Photomacrography: René T.J. Cappers & Dirk Fennema, unless otherwise noted in the captions

Book cover photographs: René T.J. Cappers

Book cover design: Nynke Tiekstra, ColfsfootMedia – Noordwolde

Book interior design and typese� ing: Nynke Tiekstra, ColfsfootMedia – Noordwolde

Copy editor: Suzanne Needs-Howarth

Cover photographs:

Top: Harvesting Bread wheat (Triticum aestivum) with a sickle (Ezbet Basili, Egypt; April 2002)

Bo� om le� : Goat browsing a dump area next to a roadhouse with concentrations of discarded

vegetables and burned cans (Ras Banas, Egypt; March 1998)

Bo� om right: Culms of Sorghum (Sorghum bicolor) used as kiln fuel (El Nazla, Egypt; November 2009)Sorghum bicolor) used as kiln fuel (El Nazla, Egypt; November 2009)Sorghum bicolor

Printed by: HooibergHaasbeek te Meppel

ISBN 9789491431074

Copyright © 2012 Groningen Institute of Archaeology (University of Groningen) and the Deutsches

Archäologisches Institut (Berlin).

All rights reserved. No part of this publication or the information herein may be reproduced, stored in a

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or otherwise, without prior permission from the Groningen Institute of Archaeology (University of Groningen)

or the Deutsches Archäologisches Institut (Berlin).

Although all care is taken to ensure the integrity and quality of this publication and the information herein, no

responsibility is assumed by the publishers nor the authors for any damage to property or persons as a result

of operation or use of this publication and/or the information herein.

Contents

Preface 91 General Introduction 13 1.1 Plant taxonomy 13 1.1.1 Taxonomic ranks 13 1.1.2 Abbreviations 14 1.1.3 Synonyms and type identifications 15 1.1.4 Naming of cultivated plants 15 1.1.5 Plant names in written sources 16 1.1.6 Vernacular names of common crops 20 1.1.7 Genetic research 22 1.2 Plant ecology 26 1.2.1 Seed production and seed predation 26 1.2.2 Seed dispersal 28 1.2.3 Environmental conditions 34 1.2.4 Water stress 37 1.2.5 Agricultural practices 47 1.3 Flora and vegetation 93 1.3.1 Landscape, flora, and vegetation 93 1.3.2 Flora 94 1.3.3 Vegetation 97 1.4 Subfossil plant remains 128 1.4.1 Biomolecules 128 1.4.2 Phytoliths 130 1.4.3 Spores and pollen 131 1.4.4 Seeds and fruits 135

2 Palynology 143 2.1 The archaeobotanical archive 143 2.1.1 Dispersal of spores and pollen 143 2.1.2 Pollen precipitation 144 2.1.3 Peat Formation 146 2.1.4 Sampling and microscopic analysis 151 2.2 Vegetation reconstruction 153 2.2.1 Pollen diagram 153 2.2.2 Anthropogenic pollen indicators 157 2.2.3 Pollen precipitation and vegetation 159 2.3 Pollen morphology 159 2.3.1 Anatomy and morphology of pollen and spores 159 2.3.2 Glossary of terms 163 2.3.3 Atlas of spores, pollen, and algae 165

3 Non-woody macro-remains 173 3.1 The archaeobotanical archive 173 3.1.1 Origin and taphonomy 173 3.1.2 Sampling 199 3.2 Morphology of fruits and seeds 239 3.2.1 Morphology of fruits 239 3.2.2 Fruit types 244 3.2.3 Morphology of seeds 246 3.2.4 Subfossil seeds, fruits, and threshing remains 247 3.2.5 Cereals 248 3.2.6 Pulses 322 3.2.7 Oil and fibre crops 333 3.2.8 Seed atlas 339

4 Vegetation history of the Netherlands 351 4.1 The Late Glacial 351 4.1.1 General overview 351 4.1.2 The Older Dryas 353 4.1.3 The Allerød 356 4.1.4 The Younger Dryas 356 4.2 The Holocene 357 4.2.1 General overview 357 4.2.2 The Preboreal (10 300–8800 BP) 358 4.2.3 The Boreal (8800–7500 BP) 359 4.2.4 The Atlanticum (7500–5000 BP) 361 4.2.5 The Subboreal (5000–2800 BP) 364 4.2.6 The Subatlanticum (2800 BP to present) 366 4.2.7 Case study 370

5 Food economy 375 5.1 Transition to farming 375 5.1.1 Modelling the dawn of farming 375 5.1.2 Domestication 380 5.2 Reconstruction of the diet 387 5.2.1 The food spectrum 387 5.2.2 Cereals 388 5.2.3 Pulses 397 5.2.4 Oil crops 402 5.2.5 Vegetables and fruits 403 5.2.6 Case studies 405

6 Fuel 423 6.1 Woody plants 423 6.2 Non-woody plants 425

7 Appendixes 435 7.1 Chronology of the Near East 435 7.2 Chronology of ancient Egypt 436

8 Literature 439 8. 1 Taxonomy 439 8. 2 Ecology 439 8. 3 Flora and vegetation 440 8. 4 Identification 443 8. 5 Spores and pollen 445 8. 6 Fruits, seeds, and mosses 446 8. 7 Vegetation history 447 8. 8 Food economy 449 8. 9 Fuel 451 8. 10 Additional references cited 451

9 Indices 457 9.1 Taxonomic and syntaxonomic index 457 9.2 Subject index 467

Burning of kitchen trash outside the camp area (Berenike excavation, Egypt; 2000–2001 season).

Preface

Plant palaeoecologists use data from plant fossils and plant subfossils to reconstruct ecosystems of the past. This book deals with the study of subfossil plant material retrieved from archaeological excavations and cores dated to the Late Glacial and Holocene. Subfossil plant remains offer us great opportunities for the reconstruction of former landscapes and the ways in which humans exploited and even transformed vegetations. However, to improve our knowledge of the past, we need to employ sampling procedures that will provide us with samples that contain the relevant kinds of plant material. This, in turn, means we need to have solid knowledge of all processes that act upon plant material. By modelling the transport of plant material from outside the settlement towards the settlement and its final destination in specific archaeological contexts, we can design an optimal sampling strategy to be used during the excavation. Most archaeological contexts contain plant remains that originate from more than one source and that entered that context by more than one pathway. Recognizing these pathways — which is a real challenge — may facilitate the identification of the different depositional origins within the archaeological context. It also may help to improve the typology of archaeological contexts. Fortunately, nowadays archaeobotanical research focuses much more than it did in the past on a clear link between archaeology, taphonomy, and biology. One of the main objectives of this book is to describe the processes that underlie the formation of the archaeobotanical archive and the ultimate composition of the archaeobotanical records, being the data that are sampled and identified from this immense archive. Our understanding of these processes benefits from a knowledge of plant ecology and traditional agricultural practices and food processing. This handbook summarizes the basic ecological principles that relate to the reconstruction of former vegetations and of agricultural practices in particular. Ethnoarchaeobotanical research offers the opportunity to document processes that assist us in interpreting subfossil records. We were fortunate to have many opportunities to observe traditional agricultural practices during our research stays in the Near East, the cradle of agriculture—especially in Turkey, Syria, and Egypt. Although there, too, globalization is resulting in the modernization of agriculture, some small-scale, traditional agriculture still exists. This offered us the possibility to research methods of crop growing, crop processing and storage, and food processing that in other parts of the world have long since been lost because of the transformation to large-scale modern agriculture. Although we recognize that, as with any ethnographic analogy, agricultural practices and food processing will have differed in the past because they were adapted to local conditions and traditions, our observations are nevertheless a valuable source of information that can be used to interpret

subfossil records. We grew up and were educated in the Netherlands, so because of this background, we have provided information on wild and crop plant ecology and history — and on vegetation ecology and history in general — from the Netherlands as well, as a means of verifying of our work in the Near East. Of course in the Netherlands, traditional agriculture has disappeared, and the definition of what is waste and garbage has changed enormously in an era of combine harvesters and waste disposal services. Botanical evidence relating to the past is extensive and includes both written sources and subfossil records. Our ability to interpret and use earlier written sources and identifications is, however, hampered by the ongoing changes in taxonomy, which recently have become tightly linked with genetic research. Because of these changes, the same plant can be presented under different names — which is especially true for crop plants. To facilitate the consultation of these sources, this handbook presents the valid scientific plant names together with their most important synonyms. In addition, we link the most commonly used plant names in classical Latin and Greek texts with modern plant taxonomy. A substantial part of this book gives a more detailed view of plant taxonomy and morphology, especially of the main crop plants in Europe, western Asia, and northern Africa. We hope this book will help palaeobotanists, environmental archaeologists, and colleagues from related disciplines optimize inferences based on what we could term “old-style” archaeobotany. And we hope that our observations will serve as an eye-opener and improve future research, not only as it is practised in our laboratories, but also as it is practised in the field. We would like to thank all those who supported our research and studies. In the first place, we want to express our gratitude to the people (too many to name) of the countries where we documented traditional agriculture — especially in Turkey, Syria, and Egypt — for their hospitality and sincerity. We also extend many thanks to the archaeologists (again, too many to name individually) with whom we collaborated, and of course to the institutions that have supported our work, the Groningen Institute of Archaeology (GIA) of the Rijksuniversiteit Groningen and the Deutsches Archäologisches Institut (DAI) in Berlin. We thank Suzanne Needs-Howarth (copy editor), Nynke Tiekstra (designer and typesetter), and Roelf Barkhuis (publisher) for giving a “face” to the book. We are grateful to Renée Bekker, Susanne Jahns, Benjamin Kilian, and Jan Frits Veldkamp for their critical remarks on some of the chapters. We thank our former professor Willem van Zeist, who introduced us to the field of palaeobotany. And, finally, we would like to express our gratitude to the late professor Sytze Bottema (1937–2005), who influenced both of us a lot by the way he incorporated field observations into his research.

René Cappers and Reinder Neef Groningen and Berlin, February 2012

22 Handbook of plant palaeoecology | Cappers & Neef

1.1.7 Genetic research

Until recently, the classification of plants was mainly based on morphological and anatomical features. Over the past 20 years, much research has been done on the genetics of plants, resulting in a better understanding of phylogenetic trees and in major shifts of plants between genera and even families. However, this work is still in progress and current floras only present a status quo of our increasing insight into phylogenetic relationships. For example, based on available genetic research, the traditional classification of the Brassicaceae family turns out to be rather artificial, and a revision may be expected. Genetic research is applied on two levels: the size of the genome and the nucleotide sequences of genes. The genome is the entire number of genes present in each cell of a particular plant taxon. A gene is a functional unit bearing the code for a specific feature of the organism. Genes are linked together in long molecules, called deoxyribonucleic acid (DNA). In organism with a large number of genes, the DNA is split up into several parts, each of which is called a chromosome. In comparison with mammals, the amount of DNA in the flowering plants varies tremendously. This is partly related to the presence of multiple copies of the chromosome set (see below). For example, the total genome of Arabidopsis thaliana contains 157,000,000 base pairs (157 megabases; Mb) representing 25,498 genes, which are subdivided over five chromosomes. Because of its relatively small genome size, Arabidopsis thaliana is one of the model plants for genetic research (fig. 2). Genes are concentrated in ‘gene islands’, and more than 80 percent of the genome consists of repetitive elements (e.g. retrotransposons).

Figure 2: Its small plant

size, short life cycle, and

small genome make

Arabidopsis thaliana one of

the model organisms for

genetic research.

1.1 Plant taxonomy 23

Most flowering plants are classified as diploid, which means that the genome is present as a double set of chromosomes in the nucleus of each cell. Only the gametes (pollen and ovules) of the plants have a single copy in their nucleus, being the result of a special cell division called meiosis. The division of the genome into a unique set of the genome makes it possible that each fusion of a pollen grain and an ovule results in a diploid cell. In this way, the size of the genome remains the same for a particular plant species. The number of the unique (haploid) chromosome number is designated with the letter n (1 x n = n). Diploid organisms have two homologous chromosomes in each cell nucleus, which is designated ‘2n’. A homologous set of chromosomes contains the same genes in a fixed sequence, although the expression of both genes may differ. Arabidopsis thaliana has five different chromosomes. The genome of this diploid plant species can thus be designated as n = 5 or 2n = 10. The genome is designated with a capital letter. This is of special interest for illustrating the composition and origin of the genomes of plant species in which changes of chromosome numbers occur (viz. polyploids; see below). Aberrant numbers of chromosomes may be present in organisms. Humans have 23 different chromosomes (n = 23), but abnormalities occur due to the absence of, for example, one of the sex chromosomes (genome is 2n − 1) or the presence of an extra copy of chromosome number 21 (genome is 2n + 1). In both cases, the aberrant number of chromosomes results in an abnormal human phenotype. The former leads to Turner syndrome and the latter to Down syndrome. Because the aberrant chromosome number results in a combination of specific, clinically recognizable features, the term syndrome is used. In plant species aberrant numbers of chromosomes also occur. This phenomenon is of special interest with respect to the process of plant domestication. The term domestication syndrome has been introduced to describe the spectrum of traits that characterize domesticated plants. A disadvantage of this term is that it is based on a clinical perspective rather than on an ecological framework. See section 5.1.2 for a further discussion on the use of this term. Variations in chromosome numbers can be achieved by reducing or increasing the number of chromosomes. The reduction of the number of chromosomes is largely restricted to diploid plant species, resulting in double haploid plants (monoploids) in which every nucleus has just a single copy of each chromosome. In fact, the genome of a monoploid, which in essentially is a diploid organism, is similar to that of a haploid gamete (pollen and ovules). Monoploids can be artificially produced by special tissue cultures in which, for example, plants are grown from pollen or unfertilized ovules. The use of pollen is preferred because they are more easily obtained in large quantities. Because genes can still be represented by several copies in the haploid genome, the phenotypic expression is only predictable when dealing with major genes. This kind of crop manipulation has been successfully applied to, for example, Soybean (Glycine max) and Common tobacco (Nicotiana tabacum). Once such monoploids have been cultivated, the chromosome number is doubled again by treating the cells with chemicals. In this way, double haploids are created having similar (homozygotous) chromosomes.


We are dealing with polyploids if more than two copies of the chromosome set are present. The genome of a triploid plant consists of three copies of the unique chromosome set (3n), that of a tetraploid has four copies (4n), that of a pentaploid has five copies (5n), that of a hexaploid has six copies (6n), and so forth. Each pair of chromosomes is designated with a capital letter. For example, the genome of the hexaploid Bread wheat (Triticum aestivum ssp. aestivum) is BBAADD, and can be written as 6n. This wheat species has 21 different chromosomes (7B, 7A and 7D), the total number of chromosomes being 6 x 7 = 42. Preferably, the genome is written as 2n = 6x = 42, in which n is the sum of a single set of chromosomes. This means that it behaves as a diploid plant (viz. the number of chromosomes of BAD = 3 x 7) and x is the number of one unique set of chromosomes (= 7). This description can be reduced to 2n = 42, in which the total number of chromosomes is mentioned but the polyploidy level remains obscure. The sequence of the genome designation in genomic formulas is determined by the taxon providing the cytoplasm (ovule); thus female parent x male parent. The BA genome of Triticum turgidum indicates that the B genome has been donated from a female Aegilops plant (present in the ovule) and the A genome stems from a male Triticum plant (present in the pollen grain). The hybrid offspring receives the organelles, residing in the cytoplasm, from the female parent. In plant cells, organelle genes are present in mitochondria (producing energy-rich ATP molecules) and chloroplasts (conducting photosynthesis). Recombination of mitochondrial DNA (mtDNA) and chloroplast DNA (cpDNA) does not occur during sexual reproduction, but changes in the DNA structure (viz. the sequence of the bases) result from rapid mutations. For this reason, studying the organelle genes is of much interest for understanding the evolutionary relationships of plants. The presence of more than two copies of the chromosome set is rather common in the plant kingdom. About one-third of all flowering plants are polyploids, and the proportion is even higher in economic plants. Because the number of chromosomes is large in polyploids, the nucleus and cell are often larger as well, resulting in plants that are more flexible and can adapt to different conditions and habitats. Polyploidy can be the result of chromosome duplication within a species, which is called autopolyploidy, but it can also be the result of a crossing between two different species (hybridization), which is called allopolyploidy. Autopolyploids have been cultivated because of the increase in flowers and fruits in particular. Fruits of Grape (Vitis vinifera) may serve as an example. The fruits of the diploid species (genome: 2n = 38) are normal-sized, whereas those of the tetraploid cultivars (genome: 4n = 76) have fruits that are almost twice as large. Allopolyploidy has played an important role in the cultivation history of several crops, including Wheat (Triticum), Banana (Musa), Banana (Musa), Banana ( ), Sugarcane (Saccharum), Potato (Solanum), and Sweet potato (Ipomoea). Triticum urartuand T. monococcum are both diploid, having two sets of seven chromosomes, but their genome is distinct, and the related chromosomes show little affinity, as a result of which hybridization is rare (Table 7). A superscript letter designates the different character of both genomes: Audesignates the different character of both genomes: Audesignates the different character of both genomes: A and Am. These letters are not fixed, and different combinations are used in publications. The two tetraploid wheat species, T. timopheevii and T. turgidum, both have the A genome, which is donated by Triticum urartu. The domestication of wheat also

1.1 Plant taxonomy 25

includes hybridization with Aegilops species. Recent research shows that the B and G genomes were probably donated by Ae. speltoides and the D genome by Ae. tauschii, although the origin of the B genome is still a matter of debate. The domestication history of wheat most probably included two independent hybridizations of Ae. speltoides with T. urartu. The first hybridization could have taken place some 300,000 years ago, by which the B genome was donated. During a second, more recent, hybridization between the species about 90,000 years ago, the G genome was donated. Due to the long time span since these hybridizations, the genome of the outcrossing Ae. speltoides has evolved, and identical alleles are only found in the G genome; hence it gets its own designation with the letter S. Most probably, specimens of Ae. speltoides having a genome that is almost identical with the B and G genomes cannot be found anymore (Kilian et al., 2009; Kilian et al., 2011; Wang et al., 2011).

Table 7: Genome designation of Aegilops and Triticum species. The two Aegilops species have donated their genome by hybridization.

Plant species Genome

S B G A D

Aegilops speltoides S

2n = 2x = 14Aegilops tauschii D

Triticum urartu Au

Triticum monococcum Am

Triticum timopheevii G Au

2n = 4x = 28Triticum turgidum B Au

Triticum aestivum B Au

2n = 6x = 42Triticum zhukovskyi G Au Am

Recently, artificial hybridization has resulted in new crops, such as Triticale (x Triticosecale) and several Cabbages (Brassica). Triticale is a hybrid of Wheat (Triticum) and Rye (Secale cereale), combining the high yield and protein content of wheat with rye’s the adaptation to more harsh environments and its high lysine content, one of the essential amino acids. Several hybrids exist in which genomes of different wheat species have been used. No valid names on a species level are available yet. Although the first crossings were made in the late 1890s, it only became a commercial crop in the 1970s. This commercially produced hexaploid Triticale (genome AABBRR) is a hybrid of the tetraploid Hard wheat (Triticum turgidum ssp. durum; genome BBAA) and the diploid Rye (genome RR). The octaploid Triticale (genome BBAADDRR), resulting from a cross between the hexaploid Bread wheat (Triticum aestivum; genome BBAADD) and Rye, has less favourable traits and is therefore not cultivated on a commercial basis. Within the genus Brassica, three diploid species are considered to be basic species: B. nigra (2n = 2x = 16), B. oleracea (2n = 2x =18), and B. rapa (2n = 2x = 20). Hybridization between B. nigra and B. oleracea resulted in the tetraploid B. carinata (2n = 4x = 34), hybridization between B. nigra and B. rapa in the tetraploid B. juncea (2n = 4x = 36), and hybridization of B. oleracea and B. rapain the tetraploid B. napus (2n = 4x = 38). Each of these tetraploids is represented by several varieties, demonstrating the large phenotypic potential within this genus.


The classification of plants has recently been improved by the analysis of DNA sequences. This kind of research focuses on the specific gene sequences, such as that of the gene rbcL (ribulose biphosphate carboxylase, large subunit). This gene is present in the chloroplast DNA and has the genetic code for the production of an enzyme that triggers an essential process in photosynthesis. Because of this essential trait, this gene is present in all angiosperms and is suitable for this kind of research. The phylogenetic relationship between plants is inferred by specific dissimilarities in the base sequences of the gene. The conserved regions of this gene are of special interest when comparing the same gene in different species, because they accumulate mutations over time. Assuming a rather constant rate of mutations, the dissimilarities in base sequences can be interpreted on a time scale and used for the reconstruction of phylogenetic trees. This kind of reconstruction is, however, hampered by specific events that can lead to a higher rate of mutations. The Angiosperm Phylogeny Group (AGP), which has published three overviews so far, governs this research (viz. APG 1 in 1998, APG 2 in 2003, and APG 3 in 2009).

1.2 Plant ecology

1.2.1 Seed production and seed predation

Subfossil plant remains can be used to reconstruct the former vegetation and the past food economy. Ideally, archaeobotanical data records should consist of the scientific plant name; the plant part; the number of plant remains; the preservation condition; and the presence of specific features dealing with fragmentation, predation, and processing. The number of seeds or fruits, being part of most archaeobotanical records, can be indicative of the relative contribution of each plant to the former vegetation or food diet. This does not imply, however, that the observed differences in numbers coincide with the original representation in the vegetation or diet. Several variables have to be taken into account for a more reliable interpretation. For the reconstruction of the former vegetation in a particular landscape, these include seed production, seed dispersal, and the taphonomic processes that act on seeds after deposition over time. A complicating factor for the reconstruction of vegetation is that descriptions of vegetation are based on the surface area coverage of the individual species, whereas archaeobotanical records predominantly consist of seed counts. A possible pathway to link the archaeobotanical data record to the syntaxonomy will be discussed in section 1.3.3. Variables that determine the presence in the archaeobotanical data record of edible plants deal with crop processing and food preparation in particular. A meaningful conversion of seed counts to energy levels will be discussed in section 5.2.5. Seed production is the primary variable that determines the number of seeds recovered from an archaeological sample. Differences in seed production are, however, not mirrored in the archaeobotanical archive. Seed dispersal and all kinds of taphonomic processes are responsible for a shift in the proportions between plants. The potential seed production is primarily determined by the number of ovules present in the lower part of the pistil. Actual seed production is primarily determined by the degree of fertilization. This, in turn, depends

1.2 Plant ecology 71

During the entire growing season, and close to harvesting time in particular, crops are vulnerable to predation. Agricultural experiments in salt marshes have, for example, shown that crops can be damaged by insects, cattle, and birds (figs. 63 and 64). The fencing of fields proved to be necessary to prevent such a loss of yield. Birds need trees in the vicinity of the fields to use as shelter; in the absence of trees, depredation by birds is no longer a serious threat.

1.2.5.6 Threshing, winnowing, and sievingThreshing is the first process after harvesting. It is aimed at separating the edible parts from the non-edible parts. Threshing can be followed by winnowing and sieving, during which the edible parts are cleaned to a higher degree. While threshing, if practised, takes place only once, winnowing and sieving can be applied several times before the grains are used for food preparation. Crop processing following harvesting can be rather complicated and is determined by the kind of crop, its use, the harvesting method, and the cultural traditions, which may be partly related to environmental conditions. Hulled and naked (or free-threshing and hull-less) cereals are threshed using different methods. In hulled cereals, the grain kernels are tightly enclosed by the chaff, whereas naked cereals have loose chaff. The threshing of hulled cereals is aimed at separating the spikelets from the rachis, whereas the threshing of naked cereals is aimed at separating the grain kernels from the ear. The threshing of naked cereals can be completed in one go. The grains are easily separated from the chaff on the threshing floor—hence its alternative adjective, free-threshing. The threshing remains of naked cereals consist of culm fragments; chaff (namely, glumes, lemmas, and paleae); and diaspores of field weeds (fig. 65). The lighter fragments can be removed by winnowing. The difference in husk tightness determines both the use that the threshing remains can be put to and the archaeological contexts that should be sampled.

Figure 64: Flower-heads of

Sunflower (Helianthus

annuus) with ripening fruits

are protected from bird

predation with plastic bags

(Çiflik, central Anatolia,

Turkey; August 2010).


Hulled cereals need two stages of threshing. The first threshing results in the fragmentation of the culms and spikes. Subsequent winnowing and sieving separate the spikelets from the threshing remains. Threshing remains from hulled cereals consist of culm fragments (the number depends on the reaping height) and diaspores of field weeds (fig. 66). A second threshing is necessary for dehusking the grain kernels. This can be done by pounding and grinding. To facilitate the removal of the chaff, spikelets can be slightly roasted.

Figure 65 : Threshing

remains from Bread wheat

(Triticum aestivum)

produced by a modern

threshing machine. These

remains are fragmented to

a high degree and are

relatively poor in

diaspores from associated

field weeds (Tunis, Egypt;

September 2002).

Figure 66: Threshing

remains from hulled 6-row

Barley (Hordeum vulgare

ssp. vulgare) produced by a

modern threshing

machine. These remains

contain a large number of

diaspores from associated

field weeds. Barley is only

represented by some

smaller sterile spikelets

(Calabria, Italy; July 2007).


The degree of fragmentation of the threshing remains depends on the implements used. The use of a flail or roller will bruise the crop. The same is true if animals, such as horses or oxen, are used for trampling the grain out of the crop (fig. 67). More advanced implements, such as the threshing sledge, having stones or metal on its underside, combine separating with fragmenting. Flailing and eventually rolling can be done inside, whereas trampling and sledging need more space and are done on a threshing floor that is located outside. Threshing may be delayed because of the pressure of work during the summer period. In that case, the unthreshed harvest is stored and threshing is done during the winter period at regular intervals.

When hulled cereals are ear-harvested, threshing is not necessary if the grain is to be used as fodder or for making beer, especially in the case of 2-row Barley. If used as fodder, the whole yield can be fed to the animals. There are several advantages to this. The spikes can be fragmented into their individual spikelets, and they can also be separated from the culm fragments. Fragmentation of the harvested crop also reduces the amount of space that is needed for storage. The spikelets have a higher nutritional value. And finally, spikelets are larger and are more easily chewed than the grain kernels themselves. For beer making, spikelets are isolated from the yield, but separating the grain kernels from the chaff (dehusking) is not necessary. Grain kernels are allowed to germinate, and this occurs naturally within the chaff. Only when grain is to be used for human consumption should an effort be made eventually to separate the grain kernels from the chaff. Threshing can be done off-site or on-site. In the latter case, the yield can be stored and threshing can be done in periods when less labour is required on the fields. Threshing is preferably done on a threshing floor that is clean, flat, and solid (fig. 68). The size and quality requirements for the floor also depend on the implements used for threshing. Cato (On Agriculture, 91) and Varro (On Agriculture, 11), for example, describe the requirements for a threshing floor. To facilitate the drainage of rainwater, the central part should be somewhat elevated, and the solid and well-packed floor may be coated with amurca, being the last liquid that is produced during olive pressing. This black water has antiseptic properties and protects against weedy plants, insects, and moles.

Figure 67: Wall painting

from the tomb of Mena at

Thebes (eighteenth

dynasty), showing

threshing of ears of

Emmer using longhorn

cattle. This is a

reproduction on Papyrus,

painted by Nadia Kotb.


Some crops, such as Sesame, can be threshed without special tools. When the uprooted plants have become sufficiently dry, they can be turned upside down, and the seeds can then be shaken from the fruits (fig. 69). The threshing of many crops is, however, done by animals or with special tools. By walking over the harvested crop, animals can tread the spikelets from the ears and the seeds from the fruits with their feet. Traditional tools used for threshing are flails, threshing sledges, and rollers (fig. 70). A flail is made from two rods of uneven length, connected to each other with a short chain or other strong, flexible material, such as the dried skin of an eel, which is extremely durable. The threshing sledge was used until recently around the Mediterranean and in the Near East. The most common type consists of a

Figure 68: Overview and

detail of a village

threshing floor. The floor

consists of large, flat

stones. The metal pole in

the centre was used for

fastening the rope used to

restrain the animals that

were used for threshing.

Having fallen out of use,

the threshing floor has

become a storage place

for fuel wood (Geraki,

Greece; July 2010).


wooden board with many small holes on the bottom, in which sharp stones or metal blades have been inserted (fig. 71). The use of sledge flints illustrates that even in modern times implements could be still partly made of stone. In Egypt, a different type of threshing sledge evolved: the nurag. This sledge consists of a frame that is furnished with metal or wooden rollers. Each roller carries several large metal disks (fig. 72).

Figure 69: Seeds of

Sesame (Sesamum

indicum) are shaken from

the fruits (Tunis, Egypt;

October 2003).

Figure 70: Threshing fruits

of Chickpea (Cicer

arietinum) using a roller.

Picking the fruits from the

plants and crushing the

fruits is done on the roof

(Murtazaköy, Turkey;

August 2010).


3.2.5.4.4 Subfossil remainsCharred grain kernels of Broomcorn millet are mostly preserved without chaff. The scutellum is large and the embryo may be exposed or broken off. In the latter case, the shape of the grain kernel is characterized by a hollow base (fig. 81).

3.2.5.5 Setaria (Bristle grasses)3.2.5.5.1 Classifi cation

Wild Domesticated

Setaria italica (L.) P. Beauv. Setaria italica (L.) P. Beauv.

ssp. viridis (L.) Thell. ssp. italica

Note: Panicum italica L. is a synonym of Setaria italica ssp. italica.

The progenitor of Foxtail millet (Setaria italica) is Green bristle grass (S. italica ssp. viridis). Although the wild ancestry is well defined, many floras still use the name Setaria viridis (L.) P. Beauv. for the wild progenitor. One of the synonyms of domesticated Foxtail millet is Panicum italica L.

3.2.5.5.2 EcologySetaria italica ssp. viridisGreen foxtail millet is native to Europe, the temperate part of Asia, the Indian subcontinent, and northern Africa. It is a weed that grows on both cultivated and waste ground and is thus a potential seed contaminant. Green foxtail millet propagates by means of shattering spikelets.

Setaria italica ssp. italicaFoxtail millet is propagated by sowing florets and by tillering. It grows in areas with a temperate climate and is adapted to poor soils, being less exacting than Panicum miliaceum. Soils should not be too dry or waterlogged. Foxtail millet can be harvested 2.5–3 months after sowing. If soils are not too poor, Foxtail millet produces higher yields than P. miliaceum. Mixed cropping with P. miliaceum is also practised.

Figure 81: Charred grain

kernels of Panicum

miliaceum from Tanais

(Russia), dating to the

Roman period.

3.2 Morphology of fruits and seeds 255

3.2.5.5.3 MorphologyThe infructescence is a dense, spike-like panicle. Spikelets of Setaria differ from those of Panicum by the presence of a large number of long scabrid bristles at the base of each spikelet, giving the infructescence a bristly appearance (fig. 82). The dispersal unit of the wild subspecies (ssp. viridis) is a spikelet (consisting of glumes and sterile and fertile floret), whereas that of the domesticated subspecies (ssp. italica) is the fertile floret (grain kernel enclosed by the lemma and palea) (figs. 83 and 84). In both subspecies, the bristles remain attached to the axis. The spikelet of Foxtail millet has a similar structure to that of Broomcorn millet (Panicum miliaceum). One of the main differences is the presence of small papillae on both lemma and palea, which are more or less arranged in short wavy lines on the lemma (thus being transversely rugose). The papillae on the palea are limited to the central area and are sharply bordered by smooth fringes (fig. 85). As in Broomcorn millet, the colour of the hardened lemma and palea of the fertile floret varies.

Figure 82: Infructescences

(spikes) of Setaria italica

ssp. viridis (left) and

S. italica ssp. italica (right).

In the wild subspecies

(viridis), part of the florets

has been shattered,

whereas all florets are still

present in the spike of

the domesticated

subspecies (italica).

Figure 83: Spikelets and

florets, the latter with

dark flecks, of Setaria

italica ssp. viridis.

1 mm 1 cm

1 mm


Figure 84: Spikelets

and florets of Setaria

italica ssp. italica.

Figure 85: Ventral view of

fertile florets of Setaria

italica ssp. viridis (left),

Setaria italica ssp. italica

(middle), and Panicum

miliaceum (right). Fringes

of the lemma cover those

of the palea and are thus

visible in a ventral view.

Papillae are present on the

lemma and the central

part of the palea of both

subspecies of Setaria

italica. The palea of

Panicum miliaceum

is characterized

by three nerves.

Figure 86: Charred grain

kernels of Setaria italica

ssp. viridis from Lebehn

(Germany), dating to the

Late Bronze Age.

1 mm

8. 1 Taxonomy 439

Literature

Most of the literature is organized in sections and also includes relevant references that are not cited in the text. Although there is some overlap among the sections because publications may cover several themes, we feel that this presentation facilitates the search for relevant literature on a certain topic. The last section concerns references from the text that are not directly related to any of the named sections.

8. 1 Taxonomy

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8


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8. 3 Flora and vegetation

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Stripping Date palm (Phoenix dactylifera) leaves (Marazig, Egypt; February 2010).

9.1 Taxonomic and syntaxonomic index 457

Abelmoschus esculentus, 425Abies, 166Acacia, 61 A. nilotica, 180 A. tortilis, 30-31, 42, 46, 89, 103, 423Acer, 161, 361, 366 A. pseudoplatanus, 164, 168, 243Ador, 19Adoreum, 19Adoxaceae, 348Aegilops, 24, 91, 252, 320, 322, 396 Ae. bicornis, 286, 321 Ae. biuncialis, 286 Ae. caudata, 286, 321 Ae. columnaris, 286 Ae. comosa var. somosa, 286 Ae. comosa var. subventricosa, 286 Ae. crassa, 286 Ae. cylindrica, 285-288, 290, 321 Ae. geniculata, 286 Ae. juvenalis, 286 Ae. kotschyi, 286 Ae. longissima, 286, 321 Ae. neglecta, 286 Ae. peregrina var.

brachyathera, 286 Ae. peregrina var. peregrina, 286 Ae. searsii, 286, 321 Ae. sharonensis, 286, 321 Ae. speltoides, 25, 285-289, 321 Ae. speltoides var. ligustica,

286-288, 321 Ae. speltoides var. speltoides,

286-289, 321 Ae. squarrosa, 285-286 Ae. tauschii, 25, 285-287, 289-291 Ae. triuncialis, 286

Ae. triuncialis var. persica, 286 Ae. triuncialis var. triuncialis,

286 Ae. umbellulata, 286 Ae. uniaristata, 286 Ae. vavilovii, 286 Ae. ventricosa, 286Aegopodium podagraria, 35Aerva javanica, 46, 103Agrostemma githago, 88, 92, 115, 246,

341Aizoon canariense, 45-46, 102Alhagi graecorum, 90, 116, 429,

432-433Alica, 19Alisma, 217-218 A. plantago-aquatica, 339Alismataceae, 339Allium cepa, 50, 114, 184Alnetea glutinosae, 99Alnus, 160, 359-360, 364 A. glutinosa, 164, 168, 345, 361Alopecurus aequalis, 110 A. myosuroides, 115 A. pratensis, 138, 340Alstroemeria, 133Amaranthus graecizans, 103-104Ammi majus, 211Ammophila arenaria, 340Ammophiletea, 99Anagallis, 245, 347 A. arvensis, 207-208, 210, 346 A. arvensis ssp. arvensis, 346Ananas comosus, 39Anchusa arvensis, 14 A. officinalis, 14Andromeda polifolia, 187

99 Indices

9.1 Taxonomic and syntaxonomic index

Accepted names are in roman face. Synonyms, invalid names and names mentioned in classical texts are in italics. Bold face signifies syntaxon names.


Angelica sylvestris, 106-107Angiosperms, 26, 96, 132, 135, 166, 240Anisantha sterilis, 95 A. tectorum, 95Anthemis, 126Antirrhinum, 96, 245Apera spica-venti, 115Aperion spicae-venti, 114Aphaka, 19Apiaceae, 14, 27, 159, 170, 208, 243-244,

347-348Apium, 14Aquifoliaceae, 169-170Arabidopsis thaliana, 22-23, 115Arabis, 20Arachis hypogaea, 187, 246Araliaceae, 170Areca, 14Arecaceae, 14Argania spinosa, 125Arinca, 19Aristida adscensionis, 89, 102 A. funiculata, 45, 89, 102, 104 A. mutabilis, 102, 104Armenia, 269 A. maritima, 341Armoracia rusticana, 246Arnebia hispidissima, 102-103Arrhenatherum elatius, 138Artedia squamata, 287Artemisia, 356, 365, 376 A. vulgaris, 111Artemisietea herbae-albae, 123Artemisietea vulgaris, 99, 107, 111Asplenietea trichomanis, 99,

107-108Aster, 14Asteraceae, 14, 27, 29, 123, 170-171, 208,

349Asteretea tripolii, 99, 123Astragaletea, 123Astragalus, 429-430 A. eremophilus, 89, 102 A. vogelii, 89, 102-103Atriplex littoralis, 164 A. patula, 14, 29, 115, 342 A. patula var. bracteata, 14 A. portulacoides, 36 A. prostrata, 29 A., 92, 117, 205, 217, 251-252, 319Avena, 92, 117, 205, 217, 251-252, 319 A. abyssinica, 21, 257-258, 264

A. barbata, 257-260 A. brevis, 21, 257-258, 264 A. byzantina, 257 A. diffusa var. segetalis, 257 A. diffusa var. vogelensis, 257 A. fatua, 118, 208, 210, 250, 257-258,

260-261, 267, 320 A. fatua var. glabrata, 260 A. ludoviciana, 257 A. nuda, 21, 257-259, 262-263, 267 A. nudibrevis, 257 A. orientalis, 257 A. sativa, 18, 21, 114, 209, 257-259,

264-268, 320, 383 A. sterilis, 209, 250, 257-258,

261-262, 267, 320 A. strigosa, 21, 257-258, 263-264, 320Balanites aegyptiacus, 31, 194, 196, 423Bellis perennis, 108Beta vulgaris, 53-54, 80, 114, 117-118,

209-210, 217, 225 B. vulgaris ssp. vulgaris, 133Betula, 164, 244, 356, 358, 365-366, 372 B. nana, 353-354 B. pendula, 29, 168, 353, 359 B. pubescens, 164, 168, 353, 363Betulaceae, 168, 345Bidens tripartita, 110, 349Bidentetea tripartitae, 99, 107, 110Bolboschoenus maritimus, 339Boraginaceae, 244, 346Borago officinalis, 346Brassica, 14, 191, 225 B. carinata, 25 B. juncea, 25 B. napus, 25, 114, 425 B. nigra, 25, 217, 345 B. oleracea, 25 B. rapa, 25, 106, 114, 168Brassicaceae, 14, 20, 22, 135, 164, 168,

209, 239, 245, 338, 345Bromopsis erecta, 138Bromus secalinus, 95, 383Bryophyta, 131Cakiletea maritimae, 99Calluna, 160, 245 C. vulgaris, 147, 149, 164, 169, 356, 357Calluno-Ulicetea, 98, 123Caltha, 244Camelina, 20-21, 338, 402 C. alyssum, 122, 337 C. sativa, 20-21, 337-338, 402


C. sativa ssp. microcarpa, 337 C. sativa ssp. pilosa, 337 C. sativa ssp. sativa, 337-338Campanula, 245 C. rapunculus, 348Campanulaceae, 348Cannabaceae, 345Cannabis sativa, 21, 334-335, 345 C. sativa ssp. sativa, 334-335 C. sativa ssp. spontanea, 334Capparis spinosa, 116, 191Capsella bursa-pastoris, 345Carduus crispus, 111Carex, 244 C. acuta, 15 C. elata, 15 C. flava, 15 C. hostiana, 15 C. inflata, 15 C. lepidocarpa, 15 C. nigra, 14-15 C. oederi, 15 C. otrubae, 15 C. riparia, 339 C. rostrata, 15 C. serotina, 15 C. trinervis, 15 C. vesicaria, 15 C. vulpina, 15, 340Carpinus, 366 C. betulus, 168, 364Carthamus tinctorius, 217, 402Carum carvi, 114Caryophyllaceae, 123, 209, 341-342Cassia, 17 C. fistula, 20Castanea, 244Caucalidion platycarpi, 114Caylusea hexagyna, 102, 104Cenchrus ciliarus, 103Centaurea cyanus, 115, 123, 157, 170,

238, 349, 356Centropodia forskalii, 103Cerastium, 245 C. fontanum ssp. vulgare, 341Ceratophyllaceae, 167Ceratophyllum, 167Charetea fragilis, 98Chelidonium majus, 108Chenopodiaceae, 123, 160, 167, 342,

356, 376

Chenopodium album, 29, 33, 115, 167, 342

C. ficifolium, 164 C. murale, 104, 209, 211, 217-218 C. polyspermum, 115 C. rubrum, 110Cicer arietinum, 18, 21, 64, 75, 77,

326-328, 399 C. arietinum ssp. arietinum,

326-328 C. reticulatum, 326Cicercula, 18Cichorium endivia, 114Cirsium, 29 C. vulgare, 111, 349Citrullus colocynthis, 103-104 C. lanatus, 187, 218, 404Citrus, 245-246Cladium mariscus, 187Clusia, 14Clusiaceae, 14Cocos nucifera, 245Coelachyrum brevifolium, 45, 102Coelastrum, 165Coix lacryma-jobi, 20Colchicum, 245Comarum palustris, 187Compositae, 14Coniochaeta lignaria, 165Coniochaetaceae, 165Convolvulaceae, 346Convolvulo-Filipenduletea, 99Convolvulus, 157, 205 C. arvensis, 115-116, 209-211, 217, 346Conyza canadensis, 96Coriandrum sativum, 217, 384Cornus, 245Coronilla, 245Corylus, 159-160, 359, 361, 364-366 C. avellana, 21, 30, 164, 168, 225Crambe maritima, 28Crassulaceae, 39Crotalaria aegyptiaca, 45, 103-104 C. microphylla, 102, 104Cruciferae, 14Cucumis melo, 240, 404 C. sativus, 30, 217, 404Cucurbitaceae, 245Cupressaceae, 132, 166Cuscuta epilinum, 122Cymbalaria muralis, 108, 110


Cynosurus cristatus, 138Cyperaceae, 164, 167, 217-218, 244,

339-340, 355Cyperus, 217-218 C. conglomeratus, 103-104 C. esculentus, 95, 123Dactylis glomerata, 138Danthonia decumbens, 87Datura, 245 D. stramonium, 115Daucus carota, 54, 114, 225, 347Dennstaedtiaceae, 165-166Dichanthium foveolatum, 89, 103-104Digitalis, 96Digitaria, 249Digitario-Setarion, 114Dipsacaceae, 348Dipsacus fullonum, 348Ecballium elaterium, 111-112Echinochloa, 249 E. colona, 209 E. crus-galli, 108Eleocharis, 217-218 E. palustris, 340Elytrigia repens, 111, 115Emex spinosa, 209-210Empetraceae, 356Empetrum, 245 E. nigrum, 169, 187, 356, 357Emys orbicularis, 363Epilobietea angustifolii, 99Epilobium hirsutum, 343Epipactis palustris, 29Equisetum arvense, 131Eragrostis cilianensis, 102 E. ciliaris, 45, 102-105 E. ciliaris var. laxa, 105Erica tetralix, 147, 149, 169, 346Ericaceae, 123, 143, 169, 346, 355-356Eriophorum, 147, 159 E. vaginatum, 167Erodium, 244 E. cicutarium, 343Erophila verna, 115Ervum, 19Eucalyptus, 154Euphorbia, 244 E. granulata, 102, 207 E. helioscopia, 209, 343 E. prostrata, 207Euphorbiaceae, 343Euphrasia, 96

Faba, 13-14, 19Faba vulgaris, 13Fabaceae, 13-14, 27, 48, 116, 123, 135,

239, 244, 247, 343-344, 399, 432Fabales, 13Fagaceae, 167-168Fagonia, 103Fagopyrum, 244 F. esculentum, 21, 114, 148, 369Fagus, 244, 364, 366 F. sylvatica, 164, 167Fallopia convolvulus, 115-116, 157Far, 19 F. adoreum, 19Farsetia ramosissima, 89, 103Festuca gigantea, 167 F. rubra, 106, 138Festuco-Brometea, 98Ficaria verna, 137, 246Ficus, 42 F. carica, 218, 225, 243, 345, 399,

404-405 F. sycomorus, 61, 405Filicales, 131Foeniculum vulgare, 348Formica rufa, 32Forsskålea tanacissima, 89, 103Fragaria vesca, 344Franguletea, 99Fraxinus, 244, 360-361, 365-366 F. excelsior, 169, 362Fumaria officinalis, 116, 340-341Fumariaceae, 340-341Fumario-Euphorbion, 114Fungi, 27, 131, 143, 161, 189Galanthus, 161Galeopsis, 115 G. bifida, 15 G. speciosa, 15, 347 G. tetrahit, 15, 157Galinsoga, 157Galio-Urticetea, 99Galium, 161, 205, 217, 244 G. aparine, 115, 346 G. spurium, 95 G. verum, 169Gentiana, 245Geraniaceae, 343Geranium, 244 G. phaeum, 343Gisekia pharnaceoides, 104Glycine max, 23


Gossypium, 425Gramineae, 14, 138Gratiola, 96Grossulariaceae, 342Guttiferae, 14Gymnosperms, 132, 135, 166, 239Halopeplis perfoliata, 45Hedera, 365 H. helix, 37, 134, 164, 170, 363Helianthemum, 356Helianthus annuus, 31, 69, 71, 81, 187,

240, 384, 407Helicoon dubium, 165 H. fuscosporum, 165Hieracium pilosella, 164Hippophae, 353 H. rhamnoides, 355Hippuridaceae, 97Hippuris vulgaris, 96Holosteum umbellatum, 115Homo sapiens, 380Hordeum, 133, 251-252, 365, 396 H. bogdanii, 275 H. bulbosum, 126 H. cantherinum, 18 H. distichum, 16, 18 H. galaticum, 18 H. hexastichum, 18, 273 H. murinum, 275 H. spontaneum, 16 H. tetrastichum, 273 H. vulgare, 16, 18, 21, 32, 39, 65, 72,

92, 114, 119, 167, 209, 217-218, 223, 226, 246, 250, 273-274, 277-285, 287, 291, 320-321, 375, 388, 392, 399, 405, 407, 414

H. vulgare ssp. distichon, 16, 18, 21, 92, 119, 250, 273, 277-280, 283, 287, 291, 320-321, 388, 392, 405

H. vulgare ssp. spontaneum, 16, 32, 273-274, 277-278, 320-321

H. vulgare ssp. vulgare, 16, 18, 21, 65, 72, 209, 217-218, 246, 250, 273, 277, 280-282, 284, 287, 291, 320-321, 388, 405, 407, 414

Humulus lupulus, 345Hydrodictyaceae, 165Hyoscyamus, 245 H. niger, 111, 346Hypericaceae, 343Hypericum perforatum, 343Hyphaene thebaica, 17

Ilex, 38, 245, 361, 365 I. aquifolium, 37, 164, 169, 170Indigofera, 40 I. articulata, 103-104Ipomoea, 24Iris, 245 I. pseudacorus, 30Isatis tinctoria, 21Isoetaceae, 131Isoeto-Nanojuncetea, 99Jatropha curcas, 425Juglans, 245 J. regia, 12, 225, 242Juncaceae, 245, 339Juncaginaceae, 339Juncus gerardii, 106 J. maritimus, 220 J. squarrosus, 339Juniperus, 353 J. communis, 166, 354Kickxia, 207Koelerio-Corynephoretea, 98, 123Labiatae, 14Lactuca, 114, 225 L. sativa, 114Lamiaceae, 14, 217, 244, 347Lamium, 14 L. purpureum, 347Lathyrus, 205 L. cicera, 331 L. hirsutus, 343 L. sativus, 18, 21, 116, 246, 330-332,

397-399 L. tuberosus, 123Launaea capitata, 102Leguminosae, 14Lemnetea minoris, 98Lens, 152, 162-163 L. culinaris, 13, 18, 21, 116, 217,

322-324, 397-399 L. culinaris ssp. culinaris, 322-324 L. culinaris ssp. orientalis, 322-323 L. esculenta, 322Leontodon autumnalis, 164 L. saxatilis, 33-34, 349Lepidium coronopus, 108, 207 L. sativum, 20Liguliflorae, 164Liliaceae, 166Lilium bulbiferum ssp. croceum, 123 L. martagon, 166Limonium vulgare, 36


Limosella, 96Linaceae, 343Linaria, 96Linum bienne, 333 L. usitatissimum, 20-21, 64, 79, 114,

217, 238, 333-334, 343, 399, 402 L. usitatissimum ssp.

angustifolium, 333 L. usitatissimum ssp.

usitatissimum, 333-334Lithospermum arvense, 431Littorella uniflora, 97Littorelletea, 98Lolium, 209-210, 217 L. perenne, 33, 138 L. remotum, 122Lonicero-Rubetea plicati, 99Lotononis platycarpa, 45, 102-103Lotus hebranicus, 103Lupinum, 18Lupinus albus, 18, 21, 217, 401Lycopodiaceae, 131, 165Lycopodiella inundata, 157Lycopodium, 154 L. clavatum, 165Lycopus europaeus, 347Lysimachia nummularia, 246Malus sylvestris, 21, 190Malva, 244 M. moschata, 164, 168 M. parviflora, 217, 345-346Malvaceae, 168, 345, 346, 425Matricaria discoidea, 108Medicago, 217, 237 M. intertexta, 209 M. lupulina, 343-344Melampyro-Holcetea mollis, 98Melampyrum, 96Melandrium album, 15 M. rubrum, 15Melilotion indici, 210Melilotus indicus, 209-210 M. messanensis, 80, 209Menyanthaceae, 170Menyanthes trifoliata, 164, 170, 187Mercurialis, 244Milium, 18Molinio-Arrhenatheretea, 98, 123Monsonia nivea, 103-104 M. senegalensis, 102Montio-Cardaminetea, 98Moraceae, 345

Morettia, 89Moringa peregrina, 45Musa, 24, 246Myosotis arvensis, 115 M. scorpioides, 346Myriophyllum, 220Myrsinaceae, 346Myrtus communis, 154Najas, 218-220Nardetea, 98Narthecium ossifragum, 147Nasturtium, 20Neurada procumbens, 31Nicotiana tabacum, 23Ocimum basilicum, 113Odontites vernus ssp. serotinus, 14Olea, 245, 404 O. europaea, 21, 198, 218, 236-237,

241Oleaceae, 169Onagraceae, 343Ophiostoma ulmi, 365Orbiliaceae, 165Orchidaceae, 29, 245Orobanchaceae, 29, 96Oryza glaberrima, 13 O. sativa, 13, 18, 21, 39, 407Oxalis, 245 O. acetosella, 28Oxycocco-Sphagnetea, 98Paliurus spina-christi, 126Palmae, 14Panicum, 19, 249, 319 P. italica, 254 P. miliaceum, 18, 21, 252-257, 320,

414 P. turgidum, 103Papaver, 123, 136, 245 P. rhoeas, 94, 115, 157 P. somniferum, 21, 238, 340 P. somniferum ssp. somniferum,

340Papaveraceae, 340Papaveretalia rhoeadis, 114Papilionoidae, 325Parietaria judaica, 108Parvocaricetea, 98Pastinaca sativa, 114, 348 P. sativa ssp. sativa, 348Pediastrum simplex, 165Pedicularis, 96Peganum harmala, 202


Pennisetum, 39Persicaria hydropiper, 110 P. lapathifolia, 341 P. maculosa, 115Petroselinum crispum, 191Phalaris canariensis, 209 P. minor, 209-211 P. paradoxa, 209-210, 217Phaseolus vulgaris, 397Phoenix dactylifera, 27, 57, 143-144,

180, 182, 187, 198, 218, 402, 407Phragmitetea, 98Phymatosorus scolopendria, 131Pinaceae, 37, 132, 166Pinguicula, 147Pinus, 144, 160, 287, 355- 356, 358-359,

361, 364, 378 P. contorta, 132 P. pinea, 236 P. sylvestris, 164, 166, 353, 360, 372Piper nigrum, 174-175Pistacia, 94, 245Pisum, 326 P. humile, 322 P. sativum, 19, 21, 114, 116, 247, 322,

324-325, 397, 399 P. sativum ssp. elatius var. elatius,

324 P. sativum ssp. elatius var. pumilio,

322, 324-325 P. sativum ssp. sativum, 324-325Plantaginaceae, 96-97, 169, 347Plantaginetea majoris, 98, 107Plantago, 209, 212, 245, 339, 356, 366 P. coronopus, 97, 108 P. lanceolata, 97, 164, 169, 365 P. major, 108, 158, 347 P. major ssp. major, 347Plumbaginaceae, 123, 341Poa, 14Poaceae, 13-14, 123, 160, 164, 167, 209, 217,

239, 244, 247-248, 340, 355, 365Poaceae tribe Triticeae, 248Polycarpaea repens, 103 P. robbairea, 103Polygala irregularis, 102Polygonaceae, 148, 341Polygonum aviculare, 108-109, 115, 341 P. monspeliensis, 209-211Populus, 363 P. tremula, 167Portulaca oleracea, 62-63

Potametea, 98Potamogeton, 166, 220 P. crispus, 212 P. lucens, 106 P. natans, 339 P. perfoliatus, 218-219 P. vaginatus, 187Potamogetonaceae, 166, 339Potentilla anserina, 108, 344 P. indica, 243Prunella vulgaris, 347Prunus domestica ssp. domestica, 344 P. dulcis, 245, 399 P. mahaleb, 245 P. persica, 20, 245 P. persica var. compressa, 20 P. persica var. persica, 20Pseudofumaria lutea, 108-109Pteridium aquilinum, 165-166Puccinellia distans ssp. distans, 340Pulicaria undulata, 103, 126Punica granatum, 404Pyrus communis, 344Quercetea robori-petraeae, 99Quercetum-mixtum, 365Querco-Fagetea, 99Quercus, 161, 244, 287, 359, 361, 364-366 Q. brantii, 378 Q. calliprinos, 38 Q. cerris, 38, 378 Q. cerris-type, 378 Q. coccifera, 38 Q. ilex, 38 Q. infectoria, 378 Q. ithaburensis, 38, 378 Q. libani, 378 Q. petraea, 378 Q. pubescens, 378 Q. robur, 38, 164, 168, 362, 378Ranunculaceae, 340Ranunculus, 244 R. aquatilis, 340 R. repens, 340 R. sceleratus, 110 R. subg. Batrachium, 14, 217-218Raphanus, 217, 245Reichardia tingitana, 102, 104Rhamno-Prunetea, 99Rhamnus, 245 R. frangula, 159Rhinanthus angustifolius, 96 R. minor, 96


Rhodanthe humboldtiana, 29Rhynchospora, 149Ribes rubrum, 342Ricinus, 244 R. communis, 113, 247Rorippa palustris, 110Rosaceae, 344Rostraria pumila, 102-103Rubiaceae, 169, 243, 346Rubus, 245 R. fruticosus, 32, 225, 344 R. idaeus, 32, 225Rumex, 209, 217, 356, 365 R. crispus, 126, 341 R. maritimus, 110 R. palustris, 110 R. vesicarius, 45, 102, 104Ruppieta, 98Ruta graveolens, 191Saccharum officinarum, 39, 133Sagina nodosa, 341 S. procumbens, 108Saginetea maritimae, 99Salicaceae, 167Salicetea albae, 99Salicornia europaea, 15, 36, 106, 342Salix, 143, 353-354 S. alba, 167 S. caprea, 134Saltera, 20Sambucus, 245 S. nigra, 242, 348Sanguisorba minor, 356Sapindaceae, 168, 243Scandix pecten-veneris, 115Scandulae sive speltae, 19Scenedesmaceae, 165Scheuchzerietea, 98Schoenoplectus lacustris, 353Scirpus supinus, 217-218Scleranthus annuus ssp. annuus, 341Scolytus, 365Scrophularia canina, 347Scrophulariaceae, 14, 96, 347Secale, 251-252, 396 S. cereale, 19, 21, 25, 114, 133, 157, 167,

248, 250, 269, 271-272, 317, 320-321, 383, 388, 414, 431

S. cereale f. clausopaleatum, 269 S. cereale ssp. cereale, 321, 388 S. cereale ssp. vavilovii, 250 S. iranicum, 269

S. montanum, 269 S. sylvestre, 269 S. vavilovii, 269-271Sedo-Scleranthetea, 98Senna alexandrina, 20 S. italica, 103-104Sequoiadendron giganteum, 239Sesamum indicum, 64, 75, 336-337,

384, 402-403, 425-426 S. indicum ssp. indicum, 336 S. indicum ssp. malabaricum, 336Setaria, 249, 252, 319 S. italica, 19, 21, 254-256, 320 S. italica ssp. italica, 254-256 S. italica ssp. viridis, 254-256 S. viridis, 254, 320Silene dioica, 15, 342 S. latifolia ssp. alba, 14-15Siligo, 19Similago, 19Sinapis arvensis, 94, 117, 119-120,

209-210, 217Sisymbrium officinale, 108, 111Sitellus suslyka, 182Solanaceae, 169, 346Solanum, 24, 53, 245 S. dulcamara, 169, 353 S. lycopersicum, 53, 407 S. melongena, 407 S. nigrum, 346Sonchus, 29, 211 S. arvensis, 96, 205 S. asper, 205, 349 S. oleraceus, 210Sordaria, 165Sordariaceae, 165Sorghum, 248, 408, 410 S. bicolor, 39-40, 69, 80, 84, 89, 117,

406, 425, 432-433Spalax, 182Sparganiaceae, 339Sparganium erectum, 30, 339Spartinetea, 99Speltae mundae, 19Spergula arvensis, 342Spergularia rubra, 108 S. salina, 342Sperguletalia arvensis, 114Sphagnaceae, 165Sphagnum, 146-148, 164, 230, 354, 363 S. cuspidatum, 165, 368Spinacia oleracea, 114


Stachys palustris, 347Stellaria media, 342Stellarietea mediae, 99, 112-114Stipagrostis hirtigluma, 102 S. plumosa, 103 S. uniplumis, 102, 104Suaeda monoica, 45Tamarix, 180 T. aphylla, 45 T. nilotica, 43, 44Taraxacum officinale, 170Taxaceae, 132, 166Taxus, 361 T. baccata, 166, 239Thalictrum, 356Thelypteridaceae, 166Thelypteris palustris, 164, 166Thero-Salicornietea, 99Thlaspi arvense, 345Tilia, 143-144, 153, 244, 360-361,

364-366, 368 T. x vulgaris, 168, 362Tiphe, 19Tragum, 19Tribulus terrestris, 89, 102Trichodesma africanum, 103 T. ehrenbergii, 89, 102, 104Trichophorum cespitosum ssp.

germanicum, 147Trifolio-Geranietea sanguinei, 98Trifolium, 48, 126, 217 T. alexandrinum, 53, 62, 127, 237 T. dubium, 344 T. repens, 87, 106, 108 T. resupinatum, 210Triglochin maritima, 339Tripleurospermum maritimum, 164,

171, 349Triraphis pumilio, 45, 102-104Trisetum flavescens, 138Triticeae, 248, 396Triticosecale, 25Triticum, 39, 80, 133, 241, 248, 251-252,

286, 365, 393, 395 T. aegilopoides, 293 T. aestivo-compactum, 292 T. aestivum, 19, 21, 24-25, 68, 70, 72,

78, 82, 114, 118, 136, 157, 184-185, 193, 207-208, 210, 224, 247, 249, 291-294, 296-297, 302-305, 308, 314-318, 320-321, 381, 388, 392, 394, 396, 406, 407, 431

T. aestivum ssp. aestivum, 19, 21, 24, 136, 157, 184-185, 193, 224, 247, 291-294, 296-297, 302-305, 308, 315-317, 321, 388, 392, 394, 396, 406

T. aestivum ssp. compactum, 19, 21, 291-294, 304, 317-318, 388, 396

T. aestivum ssp. macha, 291, 294, 304 T. aestivum ssp. spelta, 19, 21,

291-294, 296, 302, 304-305, 314-315, 321, 388, 396, 406

T. aestivum ssp. sphaerococcum, 291-292, 294, 304

T. aestivum ssp. vavilovii, 291 T. aestivum/durum, 316, 318, 320 T. araraticum, 292 T. baeoticum, 292-293 T. carthlicum, 292 T. compactum, 292 T. dicoccoides, 292 T. dicoccon, 292 T. dicoccum, 292 T. monococcum, 19, 21, 24-25, 183,

241, 291-295, 297-298, 303, 305, 310-312, 320-321, 375, 388-389, 392, 396, 406

T. monococcum ssp. aegilopoides, 291-295, 297, 303, 320-321

T. monococcum ssp. monococcum, 291, 293-295, 298, 303, 305, 310-312, 320-321, 388-389, 392, 396

T. oriëntale, 292 T. palaeocolchicum, 292 T. parvicoccum, 293 T. polonicum, 292 T. pyramidale, 292 T. ramosum, 19 T. robus, 19 T. sinskajae, 298 T. spelta, 292 T. thaoudar, 293 T. timopheevii, 21, 24-25, 291-295,

299, 303-304, 311-312, 321, 388-389

T. timopheevii ssp. armeniacum, 291-292, 299, 303

T. timopheevii ssp. timopheevii, 291, 293-295, 299, 304, 311-312, 321, 388-389

T. turanicum, 292 T. turgidum, 16, 18-19, 21, 24-25, 53,

86, 144, 180, 183, 185, 207, 217-218, 237, 247, 250, 275,


291-294, 296, 299-301, 303-309, 312-318, 320-321, 375, 381, 388, 390-392, 394, 396, 399, 405-406, 408, 414

T. turgidum ssp. carthlicum, 292, 294, 304

T. turgidum ssp. dicoccoides, 275, 291-292, 294, 296, 299-300, 303, 320-321, 381

T. turgidum ssp. dicoccon, 16, 18-19, 21, 53, 86, 183, 250, 291-294, 296, 299-301, 304-307, 312-314, 318, 320-321, 375, 381, 388, 390, 392, 396, 399, 405, 414

T. turgidum ssp. durum, 18-19, 21, 25, 144, 180, 185, 207, 217-218, 237, 247, 291-294, 296, 301, 304-305, 308-309, 314-317, 321, 381, 388, 392, 394, 396, 406, 408

T. turgidum ssp. paleocolchicum, 291-292, 294, 304

T. turgidum ssp. polonicum, 291-292, 294, 304

T. turgidum ssp. turanicum, 291-292, 294, 304

T. turgidum ssp. turgidum, 18-19, 21, 294, 296, 301, 304, 388, 390-391, 394

T. turgidum var. mirabile, 390 T. turgidum var. pseudocervinum,

390 T. urartu, 24-25, 291, 297, 303 T. vulgare, 292 T. zhukovskyi, 25, 291, 293-294Tuberculatisporites mamillarius,

186-187Tulipa sylvestris, 95Typha, 230 T. latifolia, 353Ulmaceae, 167Ulmus, 359, 364-366 U. minor, 164, 167, 361Umbelliferae, 14Urtica urens, 111Ustilago hordei, 285Utricularia, 147Vaccinio-Betuletea pubescentis, 99Vaccinio-Piceetea, 99Vaccinium, 246 V. myrtillus, 346Valeriana officinalis, 348

Valerianaceae, 348Veronica, 96 V. anagallis-aquatica, 347 V. arvensis, 115 V. hederifolia, 97, 115 V. officinalis, 97 V. triphyllos, 115Vicia, 205 V. ervilia, 13, 19, 21, 328-330, 332,

398 V. faba, 13, 19, 21, 80, 113, 217, 246,

328-331, 397 V. faba var. equina, 13, 19, 21, 80,

328-330 V. faba var. faba, 13, 19, 21, 328-330 V. faba var. major, 21, 328 V. faba var. minor, 21, 328 V. faba var. minuta, 13, 19, 21,

328-329 V. faba var. paucijuga, 328-329 V. hirsuta, 16, 95, 115, 117, 120-122,

405, 414, 418 V. sativa, 13, 19, 21 V. sativa ssp. nigra, 13 V. sativa ssp. sativa, 13Vigna unguiculata ssp. unguiculata, 80Viola, 247 V. tricolor, 343Violaceae, 343Viscum, 365 V. album, 363Vitaceae, 342, 343Vitis, 245-246 V. vinifera, 21, 24, 190, 218, 225,

342-343, 404Xanthium strumarium, 28, 96Zannichellia palustris, 217-218Zea, 13 Z. mays, 39, 69, 117, 184, 407Zilla spinosa, 31, 89, 103-104, 126Ziziphus spina-christi, 218, 236Zosteretea, 98Zygophyllum coccineum, 45, 103-104 Z. simplex, 45-46, 102-103

9.2 Subject index 467

Abaxial, 250-251, 278-279, 281, 303, 306-307, 312

Abiotic, 34, 93, 97, 135Achene, 244Achenium, 244Acidity, 34, 106, 113, 141, 147, 188-189Acute, 304Adaptation, 25, 29-30, 33, 37-38, 40, 45,

108, 147, 204, 385-386, 393, 396Adaxial, 250-251, 278-279, 281,

289-290, 298, 300, 306-307aDNA, 128After-grazing, 90-91After-ripening, 64, 68, 81, 120, 376, 382Agadir, 86A-genome, 291Agriculture, 18, 33, 35, 39, 43, 47-48,

51-52, 54, 58-59, 68-69, 71, 73, 78-79, 83, 91, 101, 105, 112, 115, 123, 127, 130, 157-158, 189, 202, 204-205, 207, 214, 216, 218-221, 228-229, 248, 273, 365-366, 369, 377-378, 380, 386-387, 392, 405-407, 416, 425, 428, 432

Agriophyte, 95Algae, 143, 165Alkaline, 269, 296Alkalinity, 323Allergy, 396Allerød, 351-353, 356, 358-359Alliance, 97, 113-114Allopolyploidy, 24, 291Amurca, 73, 86Anaerobic, 145Angiosperm Phylogeny Group, 26, 96Annual, 39-41, 44-45, 58, 113-114, 122, 148,

210, 213, 269, 273, 287, 333, 336, 352, 371, 376, 378-380, 385, 428

Annulus, 163, 365Anther, 132-133Anthropogenic, 104-105, 157-158, 366Antibiotic, 189AP, 154, 352, 364Arable, 29, 33, 35, 39, 58, 86-88, 112-113,

115-117, 122-123, 127, 136, 139, 144, 149-150, 157, 186, 200, 205, 207, 212, 216, 222-223, 274, 287, 325-326, 333, 337, 366, 369-370, 383, 405, 414-415, 428

Arboreal, 154, 356, 364Archaeophyte, 95Ard, 54-57, 87Arid, 45, 182, 193-194, 378Aril, 32, 239Ascospore, 165Ash, 63, 90, 91, 125, 148, 174, 204, 211,

214-216, 220, 222, 229, 337, 360-363, 365-366, 402, 413-414, 417, 423-424, 431-432

Assimilation, 35Association, 48, 97-98, 105, 138, 294Atlanticum, 351, 358, 360-361, 363-365,

368ATP, 24, 47Autopolyploidy, 24Awn, 117, 249-252, 257-259, 261-264,

267, 270, 274-275, 277, 279-281, 284, 288-290, 297, 300, 302-304, 308, 320, 385-386

Bacteria, 48, 129, 189Baculate, 163Baking, 377, 392, 396, 409, 423-424Berry, 225, 245-246Biennial, 113, 333Bin, 83-85, 180, 418Biomolecule, 128Biotic, 34, 93, 97, 106, 136, 141Biotope, 211, 414Bisexual, 27, 132-133Bog, 98, 139, 146-148, 150, 187, 368-369Bølling, 352-353, 358Boreal, 351, 357-361, 364, 370, 372Brackish, 98Bract, 34, 117, 240, 244, 249-251, 349Brewing, 188-189Brittle-rachised, 20, 67, 250, 295, 303,

305, 376, 381, 383-386Broadcast-sown, 52-54, 112, 114-115, 415Buoyancy, 225Burning, 8, 47, 90-92, 140, 183, 193-195,

201, 203, 221, 247, 337, 365, 402C3-plant, 38-39, 129, 410C4-plant, 38-39, 103, 252, 410Calcareous, 85, 101, 108, 273Calcifuge, 269, 294, 296Calcium, 47, 269, 294CAM-plant, 38-39Capsula, 245

9.2 Subject index


Capsule, 131, 244-245, 336-337Carbohydrate, 128, 133, 371, 396Carnivorous, 147Carpel, 27, 132, 244-245Carpellate, 27, 133Caryopsis, 244, 250, 275Caulking, 230Cerealia, 157-158, 248, 365, 379Cesspit, 193, 203-204, 222, 224-225, 387Chaff, 71-73, 78, 180, 186, 202, 205, 207,

210-212, 214, 217, 220-221, 225, 237, 240-241, 247-250, 254, 267, 276, 283, 293, 303-304, 308, 310, 313, 318-320, 381-383, 385-386, 391, 406, 413, 416, 428

Chalaza, 247Charcoal, 221, 229, 417, 423-424Charring, 91-92, 100, 138-140, 174, 183,

186-187, 190, 192-195, 200-201, 205, 217-218, 220-221, 223, 231-234, 250, 254, 256-257, 267-268, 272, 276, 283-285, 290, 304-307, 310-319, 324, 326-328, 330-332, 334, 337-338, 388, 399, 402, 410, 413-414, 418-419, 424, 431-432

Chemosynthesis, 35Chromosome, 22-24, 291Classification, 13, 16, 20, 22, 26, 94,

96-97, 106-107, 130, 239, 244, 248, 250, 252, 254, 257, 269, 273, 285-286, 291-293, 322, 324, 326, 328, 331, 333-334, 336-337

Clavate, 163-164Climate, 29, 34, 36-39, 45, 93, 123-124,

129, 145-146, 148, 154, 193, 229, 252, 254, 258-259, 296, 328, 333, 335-336, 351-353, 356-358, 363-365, 367, 376, 380, 385, 390, 392-393, 397, 428

Clubmoss, 157, 165CML-classification, 106CO2, 35, 38, 39, 47Coeliac disease, 396, 410Colporate, 160Colpus, 160-161, 163Columella, 18-19, 68, 162-163, 189, 191,

212Competition, 35-36, 53-54, 62, 86, 115,

122, 372, 386Cone, 83, 131-132, 239, 294Connate, 240

Contamination, 69, 80, 87, 91, 95, 105, 128-130, 135, 137-138, 152, 178, 182, 184, 186-187, 192, 215, 221, 227, 229, 232, 406, 419

Cooking, 377, 382, 392, 398, 409Coring, 144, 151, 204, 367Cotyledon, 247, 251, 322-323, 327, 330,

338Coverage, 26, 92-93, 100-104, 124, 144,

152-153, 192, 256, 296, 363, 367, 371-372

Crossbreeding, 15Cross-pollination, 27, 133-134, 252, 327,

388Cultivation, 24, 47-48, 63-64, 122,

148-149, 204, 212, 229, 291, 295-296, 328, 366, 369, 378, 380-381, 393, 397, 403, 405, 407-408, 410, 416, 421

Decay, 29, 86, 108, 129-130, 135-138, 145, 147, 191, 193, 224, 331, 413

Deciduous, 38, 101, 325, 366, 378Decumbent, 108-109, 332Deflation, 182, 220Deforestation, 139, 366Dehiscence, 120, 122, 244-245, 333, 336,

383-384, 417Dehusking, 72-73, 86, 226, 377, 383, 391Depletion, 47-48, 50, 368, 403Desiccation, 187, 205, 216, 304, 331Dew, 43-45, 58, 68, 103, 189, 257, 382D-genome, 291, 396Diaspore, 28-30, 32-33, 53-54, 63, 69,

71-72, 76, 79-80, 83, 92, 95-96, 100, 115, 117, 128, 131, 135, 146, 174-175, 183-186, 202, 205-207, 209-212, 215-216, 218-220, 231-232, 246, 250, 257-258, 260-261, 267, 287, 380-381, 418, 432

Dicotyledon, 247Diet, 20, 26, 129-130, 173, 175, 201, 203,

225-226, 371, 387, 391, 396-397, 403, 428, 431

Dioecious, 27, 143, 335Diploid, 23-25, 257, 291, 297, 305-307,

404Disarticulation, 249, 258, 260-261, 264,

266-267, 277, 286-288, 290, 303-304, 312, 314, 320, 386

Disarticulation scar, 249-250, 260-261, 264, 267, 303-304, 320, 386


Disease, 54, 192, 295, 328, 365, 377, 396-397, 410

Dissemination, 17, 250Disturbance, 35, 91, 104, 107, 115, 123,

125, 175, 365DNA, 22, 24, 26, 47, 96, 128-129, 189Domestication, 15-16, 21, 23-25, 27, 39,

56, 63-64, 67, 83, 85, 107, 116, 129-130, 212, 214, 220-221, 246-250, 252, 254-255, 257-260, 267, 269, 273-276, 286, 291-293, 295-296, 301, 303-304, 310, 319, 322-329, 331-334, 336-337, 365, 375-377, 379-381, 383-388, 390, 393, 405-407, 409

Dormancy, 29, 33, 87, 128, 138, 377, 385-386

Drainage, 73, 149, 369Draining, 40, 148, 232, 269, 296, 325,

368-369Drought, 39, 45, 46, 103, 269, 273, 287,

294, 296, 328, 353, 380Drought-resistant, 296, 328Drought-tolerant, 287, 380Dryas, 352, 355-356Dry-sieved, 216Dung, 47-51, 54, 63, 92, 106, 111, 143, 151,

194-195, 197, 211-213, 216, 220-221, 236, 369, 378, 382, 393, 409, 413-414, 422-432

Dung cake, 92, 194, 211-213, 216, 382, 393, 409, 413-414, 424-430, 432

Echinate, 163-164Ecotope, 97, 106Ectexine, 162Edaphic, 34, 123, 246Elaiosome, 32Eldest Dryas, 352, 358Embryo, 247, 249, 251, 254, 267, 322,

327, 338Endexine, 162Endocarp, 32, 194-196, 198, 212, 225,

236-237, 241, 245, 339, 344, 348, 402, 404

Endosperm, 225, 245, 247, 249, 304, 392, 395-396, 406, 409-410

Environment, 25, 38, 43, 45-46, 91-92, 101, 105, 107, 129, 135-140, 143, 145-147, 154, 182, 188-189, 191, 194, 204, 273, 286, 296, 333, 375-376, 392

Enzyme, 26, 38, 47, 189Ephemeral, 44-46, 101-104, 376Epicarp, 30, 239, 241, 245, 339Epidermis, 35, 37, 171Epipalaeolithic, 379Epithet, 14, 16Erosion, 47, 58, 182, 186, 215Ethnoarchaeobotanical, 86, 204Ethnographic, 207Eutrophic, 98, 147Evaporation, 37-39, 41, 43, 63, 111, 190,

245, 352Evergreen, 37, 38Excrement, 30, 32, 111, 224-225Exine, 159Exocarp, 239, 241-242Exotic, 154Exploitation, 51, 150, 211, 366, 368,

378-380, 425Faeces, 175, 203, 224-225, 403, 405Fallow, 48, 104, 111, 113, 116, 122, 212,

216, 325, 433Famine, 56, 331Farmer, 48, 53-54, 57-58, 61-62, 65, 67,

80, 86, 117, 122, 127, 212-213, 365-366, 377, 381-382, 385-387, 392-393, 395, 416

Farming, 18, 219, 352, 375-378, 380, 386, 428

Fenestrate, 163-164Fermentation, 188-189, 191Fern, 160, 164-165Fertile, 15, 205, 252, 255-256, 258-259,

263, 273, 275-276, 278-279, 281-282, 287, 291, 298, 316-317, 375, 378-380, 388-390

Fertility, 51, 100, 157, 248, 257, 259, 274, 325, 390

Fertilizer, 48, 50-52, 54, 431Fertilizing, 26, 27, 43, 48, 51, 133, 139, 143,

148, 188, 239, 332, 370, 377, 388Fibre, 64, 230, 333-335, 371, 403, 418, 425Firewood, 393, 432Flail, 73-74Floatation, 229, 231-234, 399-400Flora, 16, 22, 40, 93-96, 100, 107, 137,

139, 140, 205, 207, 212, 215-216, 254, 304, 353

Floret, 249-268, 275-276, 279-282, 284-285, 288-290, 297, 302, 304, 316-317, 319-320, 388-390, 415


Fodder, 56, 73, 127, 194, 212, 220, 273, 323, 365-366, 382-383, 392, 396, 406-407, 409, 413, 428, 432

Follicle, 244Folliculus, 244Food economy, 26, 174, 223, 377,

387-388, 405, 418, 421Foragers, 370, 372, 377-378Foraging, 375, 378, 380Fossil, 140Fossilization, 138Fractionation, 129Fragmentation, 26, 72-73, 92, 98, 128,

195, 250, 285, 305, 413Freatophyte, 46Free-threshing, 67-68, 71, 204,

207-208, 211-212, 249, 269, 271, 298, 305, 318, 381-383, 385, 392, 396, 408-409, 411, 415-416, 428

Fructose, 189Fruitcoat, 225, 335Fuel, 47, 63, 74, 84, 92, 125, 148, 150, 173,

180, 194, 212, 220, 229, 337, 369, 378, 382-383, 392-393, 395-396, 402-403, 409, 422-426, 428, 430, 432, 433

Fungus, 27, 165, 285, 355, 365Funiculus, 32, 121-122, 246-247, 249,

331, 397Garum, 191Gatherer, 377Gathering, 67, 258, 377, 379, 406Genome, 22-25, 129, 257, 286, 291-292,

396Germ, 138, 144, 204, 247, 249, 256, 258,

293, 311, 318, 353, 382, 416Germination, 27, 29, 31-35, 39, 46,

52-54, 87, 135-138, 285, 323, 325, 377, 385-386, 415

Glacial, 146, 148, 351-353, 356-358Glucose, 35, 38, 146, 189, 363Glume, 71, 247-253, 255, 262, 270-271,

274-275, 277, 280-282, 284, 288-290, 297-299, 302-308, 311-312, 317, 321, 381, 388, 410, 413, 431

Gluten, 395-396, 409-410Granary, 33, 83, 86, 179, 223, 231, 388, 405Grassland, 98, 101, 123, 127, 139, 212, 287,

322, 333, 356, 366, 425, 428, 432Grazing, 47, 91-92, 123, 125-130, 206, 211,

216, 269, 369, 386, 414, 428-429

Grinding, 72, 225-226, 377, 392, 395-396, 406-407, 409

Groundwater, 34, 36-37, 40-42, 46, 50, 98, 123, 145, 147, 154, 188, 352, 363, 368

Growth-forms, 100Gyttja, 146, 155Habitat, 97, 106, 269, 273Halophytic, 45Haploid, 23Harrowing, 47, 52, 54, 57Harvest, 48, 58, 73, 91-92, 113, 117, 121,

173-174, 205, 207, 211, 223, 246, 314, 331, 380, 387, 395, 414-416

Harvesting, 47-48, 63-64, 66,-69, 71, 90, 115-117, 120, 122, 189, 205-206, 211-212, 220, 246, 249, 252, 258, 271, 308, 376-377, 380, 382, 384-387, 392, 395-396, 414-418

Health, 392, 395-396, 404Hearth, 230, 413Heating, 138, 188, 378, 393, 396, 413,

423, 431Hemiparasite, 363Herbaceous, 41, 46, 62, 94, 154, 393,

422-423, 425, 432Herbivore, 129, 220Hexaploid, 15, 19, 24-25, 257, 291, 302,

305, 316, 381, 394Hilum, 32, 122, 247, 249, 251, 259, 271,

274, 276, 320, 322-323, 325-326, 329, 331

Hoe, 54-57, 87, 89, 90Hoeing, 54, 87, 112Holocene, 139-140, 146, 148, 178, 351,

357-359, 370, 376, 378Holoparasite, 363Homozygotous, 23Hulled, 18-19, 65, 67, 71-73, 86, 92, 212,

218, 225-226, 249-250, 252, 257, 259, 267, 269-270, 273, 275-276, 280-284, 291, 293, 296-297, 299, 302-303, 305, 310, 319-320, 381-384, 391-392, 395-396, 399, 405-409, 411-415, 418-419, 421, 428

Humidity, 36, 40, 43, 117, 147-148, 259, 325, 397

Hunter-gatherer, 189, 360, 370-372, 377

Hybrid, 25, 257


Hybridization, 15, 24-25, 291-292, 381Hypothesis, 200ICBN, 13, 15, 20ICPN, 130Ideotype, 386Imprint, 43, 192-193, 213, 216, 248, 267,

310, 318-320, 425, 427Indicator, 34-35, 62-63, 104, 106, 138,

140, 157-158, 205, 353, 366Infertile, 279, 388Inflorescence, 205, 209, 242, 248-249,

251-252, 287, 297, 334-335Infructescence, 27-28, 39, 64, 69, 117,

237, 239, 242, 246, 251-253, 255, 265, 270, 274, 276, 333, 386, 390, 392

Intectate, 163Interglacial, 351Internode, 208, 248, 251, 273-275, 277,

285, 289-290, 297-298, 304-307, 310, 313-314, 317, 319-321, 381, 410-411, 413, 416

Interstadial, 351Intine, 159Invader, 95-96Inventory, 101, 107, 113, 410, 415Irrigation, 47, 50, 53, 57-63, 90, 117, 127,

130, 177, 198-199, 204, 207, 210, 212, 215-219, 258, 327, 352, 392, 410

Isotherm, 353Isotope, 128-130, 130, 351Judgemental sampling, 200, 202-204,

219, 391Kiln, 193, 402, 433Kilocalorie, 370-371, 404Kilogram, 415-416Kilojoule, 370-371, 404Labour, 73, 392, 395, 409, 421Lactic fermentation, 188-189Landnam, 365-366Landscape, 26, 93-95, 97, 100, 104, 107,

127, 137, 140, 143-145, 148, 150, 174, 222, 352, 354, 356, 365-366, 369, 407, 424-428

Lax-eared, 208, 276, 416Legume, 48, 135, 244, 380Legumen, 244Lemma, 71, 180, 249-253, 255-257,

259-264, 267, 270-271, 274-277, 280-281, 283-284, 288-289, 297,

302-304, 307-308, 319-320, 381, 389-390, 431

Limestone, 325-326Lithology, 154Longevity, 33, 44, 116, 138, 159Macronutrient, 47-48Manuring, 47-48, 50, 130, 323, 369,

392, 431Megaspore, 186-187Meiosis, 23, 160Mericarpium, 244Mesocarp, 241, 245Micronutrient, 47Microorganism, 35, 80, 145, 188-189,

191-192, 224Micropyle, 247Micro-remain, 204, 227Midden, 176, 178, 220, 387Mimicry, 122Mineral, 34-35, 47-48, 50-51, 54, 63, 86,

91, 107, 113, 115, 123, 146-148, 175, 193, 213, 296, 352, 363, 365-366, 371-372, 392, 403

Mixing, 54, 135, 137, 178, 180, 182-183, 231-232, 234

Moisture, 36, 39, 43, 64, 80, 106-108, 115, 145-146, 149, 159, 193, 212, 244, 257, 287, 356, 382, 410, 426

Monocolpate, 161, 163Monocotyledon, 247Monolete, 160-161, 163-164Monoploid, 23Monoporate, 160, 163-164Mud brick, 52, 56, 60, 144, 180, 184-185,

207-208, 213-218, 220-222, 228-229, 232, 234, 414

Mutation, 24, 26, 381, 386Naked, 18, 21, 68, 71, 86, 130, 135, 180,

238, 249-250, 257-259, 267, 273, 275-276, 282-284, 291, 293, 299, 302, 304, 310, 316, 319-320, 381-383, 386, 391, 395, 408, 418-419, 421, 428

NAP, 154, 352, 364Nectar, 133, 143Neolithisation, 95, 393Neophyte, 95Nitrogen, 47, 48, 111, 129, 386Nomenclature, 13, 16, 20, 97, 130Non-arboreal, 154, 364


Non-brittle, 67, 249-250, 305, 376, 381, 385-386

Non-disarticulating, 263-264Non-shattering, 258, 269, 381, 384-385Nurag, 75Nutrient, 35, 47-48, 54, 98, 106-107,

113-114, 139-141, 246-247, 249, 337, 363, 371, 377, 390, 392

Nutrient-poor, 98, 139, 363Nutrient-rich, 98, 114Nux, 244Octaploid, 25Off-site, 29, 73, 204, 211Older Dryas, 351-353, 355-356, 358Oligotrophe, 147On-site, 73, 204Ovary, 132-133, 135, 239, 242, 251, 267Oven, 125, 229, 396, 402, 409, 413-414,

425, 430-432Overexploitation, 124, 369Overgrazing, 94, 124-126, 229, 369, 379,

393, 429, 431Ovule, 23-24, 26, 132-133, 135, 239-240,

247, 251, 332Palaeobiocoenose, 105-106Palea, 71, 180, 249-253, 255-257, 262, 271,

274-276, 280-281, 283-284, 288-289, 307-308, 319-320, 381, 431

Panicle, 117, 249, 251-252, 255, 259-260, 265, 267

Pantoporate, 163-164Pappus, 29, 34, 210, 230, 349Parasite, 96, 143, 363Parching, 377, 383PCR, 128-129Peat, 144, 146-148, 150, 155, 164, 186,

230, 363, 368-369, 379, 423Pedicel, 119, 208-209, 217-218, 251, 261,

266, 275Pentaploid, 24Perennial, 46, 62, 87, 96, 103-104, 138,

205, 269, 274, 333, 376, 428Perianth, 117, 209, 217, 249, 335, 339,

340-348Pericarp, 225, 244, 247, 249, 251, 319Pericolpate, 161, 163Pericolporate, 163Periporate, 160, 163-164pH, 188-189Phosphorus, 47, 111Photoperiodism, 385

Photosynthesis, 24, 26, 35, 37-39, 129, 146, 269, 363, 385-386, 423

Phytolith, 128, 130Pickling, 188Pingo, 145, 148, 352, 358Pioneer, 48, 99, 101, 110, 139, 175, 287Pistil, 26, 132-133, 143, 239, 242, 249Plant community, 39, 97, 100-101,

104-108, 110, 112, 114-115, 122-125, 140, 178, 186, 215, 220, 286, 372

Pleistocene, 351, 376, 378Pleniglacial, 351, 376Ploidy, 291, 305Plough, 54-57, 87, 123Ploughing, 47-48, 52, 54-56, 63, 87,

90-91, 116, 183, 194, 204, 207, 365Pollen diagram, 130, 153-159, 351, 356,

360, 364-367, 378-379Pollination, 27, 34, 133, 143, 159, 243,

377, 387-388, 404Polyploidy, 23-24Pore, 160-161, 163, 245Postglacial, 357-358Potassium, 47Pounding, 72, 310, 383Preboreal, 351, 358-359, 364Precipitation, 34, 36-37, 39-40, 43, 58,

93, 144-145, 147-148, 153-154, 159, 326, 352, 363, 376, 379, 392

Predation, 26-27, 52, 70-71, 135, 137-138, 195, 271, 371, 377

Preservation, 26, 56, 105, 128, 135, 137-140, 145-146, 152-154, 159, 173-174, 187-189, 191-195, 201, 205, 216, 221, 244, 248, 267, 282, 284, 304, 310, 319, 353, 392, 397, 402

Pressing, 73, 188-189Procumbent, 108-109Progenitor, 15-16, 116, 254, 257, 269,

291, 299, 310, 324, 328, 333, 336-337, 375-376, 380

Propagating, 28, 34-35, 67, 95-96, 129, 136-137, 254, 258, 354, 380

Propagule, 89Prostrate, 46, 63, 92, 108, 117-118, 122,

353, 417Protein, 25, 35, 47, 128, 130, 133, 189,

371, 396, 409Psilate, 163Rachilla, 249, 251, 261, 282, 290, 307, 390


Rachis, 19, 67, 71, 91, 180, 184-185, 193, 195, 208, 210, 237, 246, 248-252, 270, 272-279, 281-282, 284-285, 287, 289-290, 295, 297, 302-308, 310-314, 317, 319-321, 377, 381-386, 388-391, 396, 410-413, 415-416, 431

Rain-fed, 58, 352Random sampling, 78, 200-202, 219,

221, 391Raphe, 247Reallocation, 64, 68, 382Reaping, 63-65, 67-68, 72, 115, 204,

211-212, 214, 220, 376, 382Reduction, 188Refugium, 126-127, 356Regma, 244Reine Proben, 106, 178Relevé, 39, 100-104, 415Reproduction, 24, 28, 34, 36, 68, 73, 87,

95, 108, 125, 133, 246Reticulate, 162-164, 336Riparian, 97, 186, 215, 217-218, 220RNA, 47, 128, 189Ruderal, 99, 111-112, 139, 202, 286Saccate, 163Salinity, 34, 45, 63, 106-107, 258, 323Salsamentha, 191Salting, 188, 191Sampling, 135, 138, 151, 199-200,

202-205, 215, 219, 222, 227, 230-231, 353, 370, 387, 391-392, 413

Sarcocolla, 20Scabrate, 163Scar, 145, 148, 161, 163, 190, 247,

249-251, 258, 260-261, 264, 267, 280-281, 289-290, 303-304, 312, 319-321, 352, 358, 377, 381, 384, 386

Schizocarp, 243, 384Schizocarpium, 244Scutellum, 249-251, 253-254, 257, 298,

319-320Season, 252Seasonality, 112-115Sedentism, 370, 376-377Seed bank, 28-29, 33, 44, 62-63, 86-87,

90, 92, 104, 115-117, 122, 135-138, 140, 207, 246, 295

Seed dispersal, 26, 28-29, 32-34, 67-68, 87, 95, 97, 104-105, 108, 116, 121,

135-136, 212, 222, 246, 249, 377, 380-381, 384, 396

Seed flora, 135, 137, 206Seedcoat, 27, 29-30, 244, 246-247, 251,

322-323, 325-328, 330-331, 336, 401

Seedling, 46, 52-54, 113, 212, 386Self-pollination, 27, 143-144, 158, 252,

365, 388Self-fertilizing, 327Self-pollination, 133Semi-brittle, 250, 303, 305-306, 381,

386Semi-shattering, 269, 383Sepal, 132Septum, 119-120, 245, 333-334, 345Sexuality, 27Shade-tolerant, 36Shaduf, 61Shattering, 68, 254, 258, 269-270, 336,

381Short-lived, 123Sibakh baladi, 51Sibakh kufri, 51Sickle, 64-68, 117, 127, 205, 212, 214,

308, 376, 382, 387, 410Sieve, 76, 78-80, 129, 138, 180-181, 193,

195, 201-202, 208, 210, 214, 216, 218, 225, 231-232, 235-238, 309, 323, 331, 397, 399

Sieving, 71-72, 78-79, 92, 117, 193, 214, 231-232, 235-236, 238, 246, 323, 392, 397

Siliqua, 245Silique, 245Silo, 224, 405Sledge, 73-76, 425Smoking, 188Sociability, 100, 415Soil, 27-29, 33-36, 41-48, 50-54, 56-60,

62-65, 68, 80, 87, 93, 95-96, 100, 106-108, 110-117, 119, 123-124, 128, 135-140, 145, 148, 150-151, 157-158, 176, 182-183, 205, 207, 211-212, 215-216, 221, 231-232, 234, 238, 252, 254, 257-259, 269, 273-274, 287, 294-296, 309, 322-323, 325, 327-328, 331, 333, 336-337, 356, 361, 363, 366-369, 372, 376-377, 383, 386, 390, 392-393, 399, 407, 413, 414, 416, 419


Sowing, 20, 29, 33, 52-54, 57, 87-90, 95, 113, 115-117, 119-210, 246, 249, 254, 366, 376-377, 386-387, 390, 392-393, 405, 415, 418, 421

Spectrum, 18, 23, 30, 124, 153-154, 387, 392, 399, 418

Spike, 67, 72-73, 92, 131, 158, 162, 218, 249-251, 253, 255, 270-271, 274-276, 278-282, 286-290, 297-298, 300, 303-304, 306, 309, 313, 386, 389-391, 393-394, 415-416

Spikelet, 32, 53, 67-68, 71-74, 85-86, 92, 104, 117, 208-211, 217, 225-226, 246, 249-265, 267, 270-271, 273-279, 281-282, 285-290, 293, 295, 297-300, 302-307, 309-317, 320-321, 376, 381-383, 386, 388-391, 396, 405-406, 410-411, 413, 415

Spiny, 54, 92, 126, 393Split fruit, 243-244Sporangiophore, 131Sporangium, 131, 146Spore, 108, 128, 131, 133, 143, 145-146,

148, 152-154, 159-161, 163-166Spring-sown, 259Stamen, 27, 132-133, 249, 267Stefanocolpate, 161Stefanoporate, 160Stephanocolpate, 163Stephanocolporate, 163Stephanoporate, 163, 164Steppe, 101, 123, 229, 273, 294, 296, 325,

379-380, 427, 429Steppe-forest, 379-380Sterile, 72, 129, 191, 246, 252-253, 255,

275, 279, 282, 288-289, 297, 411Sterility, 274Stigma, 132-134Stomata, 35, 37-38Stone fruit, 241, 245, 402Storage, 27, 29, 45, 73-74, 80, 82-86, 130,

174, 191, 211, 214, 222-224, 232, 238, 285, 310, 314, 370-371, 376-377, 382-383, 385-386, 388, 391-392, 395-397, 399, 409-412, 414-415, 417-421, 425-426, 428, 431

Strategy, 33, 48, 108, 116, 124, 138, 175, 200, 204-205, 387, 391

Stratigraphy, 200

Stress, 35, 37, 40, 125, 258, 352Striate, 163-164Subassociation, 106Subatlanticum, 351, 358, 366, 369Subboreal, 351, 358, 364-365Subfossil, 26, 28-29, 91, 96, 105, 128,

130, 135-141, 143, 162, 182, 184, 190, 204-205, 220-221, 232, 235, 247-249, 254, 257, 267, 272, 282, 286, 290, 293, 304, 306, 316, 319, 323, 326-327, 330, 332, 334-335, 337-338, 375, 385-386, 388, 391, 401, 410, 415, 417-418, 423

Subsampling, 83, 154, 223, 230-231, 310, 392, 418

Subsistence, 375, 380, 409Succession, 107, 137, 365-366Succulent, 45, 62Sweetening, 188, 191Symbiosis, 34, 48, 387Syndrome, 23, 387, 396Synecology, 97Syntaxon, 97-98, 107, 123, 210Syntaxonomy, 26, 93, 97, 113, 138Tafla, 50-51Taphonomy, 26, 130, 137, 173, 205,

414-415, 417Taxon, 13-16, 18, 20, 22, 24, 94, 97, 154,

273, 291-293, 306, 399, 418Taxonomy, 13, 16, 93, 96, 269, 293, 334Tectate, 163Tectum, 162-163Temperature, 34-38, 40, 43, 93, 111, 138,

154, 188-189, 191-193, 269, 325-326, 337, 352-353, 356-357, 364-365, 385, 392, 402, 410, 423, 431

Territory, 371-372, 380Testa, 246, 251Tetrad, 160, 164Tetrahedral, 160Tetraploid, 15, 19, 24-25, 257, 291, 293,

299, 303, 305-306, 310, 316, 381, 390, 394

Thanatocoenose, 105-106Therophyte, 113Threshing, 64, 71-76, 79-80, 82, 84, 86,

92, 117-118, 174, 178, 180, 184, 189, 193-194, 201, 204-205, 207-208, 210-212, 214-216, 218, 220-221, 225, 228-229, 232, 247-250,


257-258, 260-261, 264, 267, 275-276, 280-282, 284-286, 288, 290, 304-309, 316, 318, 320, 323-324, 337, 377, 381-384, 386-388, 391-393, 395-397, 402-406, 409, 413-414, 425-426, 428, 432

Tillage, 115, 125, 158Tillering, 208, 254, 270, 297, 377,

385-386, 390, 392, 415Tough-rachised, 305, 376Toxic, 16, 246, 396Transient, 33, 138Trash, 8, 78, 85, 111, 173-174, 176-180,

182, 184, 186-187, 194-195, 197-198, 203, 217, 230-231, 308, 331

Tricolpate, 161, 163-164Tricolporate, 163-164Trilete, 160-161, 163-164Triploid, 24Triporate, 163-164Unisexual, 27, 132, 143Unleavened, 396Uprooting, 63-65, 68, 74, 88, 90, 104,

207, 212, 214, 220, 308, 376, 382, 390

Utricle, 244, 339-340Valve, 119-120, 122, 209, 245Varve, 379Vascular, 34, 38, 108, 147, 363, 365Vegetable, 189, 191, 197, 403Vegetation, 26, 33, 35-36, 41, 44, 47-48,

52, 56, 62-63, 87, 91-101, 104-108, 110-112, 115-117, 123-128, 130, 135-139, 143-147, 153-154, 157, 159, 174, 178, 183, 202, 204-205, 207, 211, 215, 221-222, 229, 231, 247, 273, 319, 351-352, 354-357, 359-360, 364-369, 372, 376, 379, 393, 415, 423, 428-429, 433

Vernalization, 297, 385-386Verrucate, 163Verruculose, 325Vesiculate, 163-164Vitamin, 35, 371Waste, 29, 35, 48, 50-51, 76, 78-80, 86,

104, 107, 112, 175, 177-178, 180, 186-187, 194, 199, 203-204, 212, 225, 248, 254, 258, 331, 387, 397-398, 401, 428

Waterlogging, 100, 138-140, 186, 192-193, 200, 205, 235, 244, 254, 267, 319, 326, 331, 335, 397

Weed, 29, 33, 35, 39, 47, 52, 54, 62-63, 69, 71-72, 78-80, 83, 86-92, 95, 97, 105, 108, 112-117, 120-123, 136, 139, 157-158, 186, 200-203, 205-208, 210-212, 214-216, 218-223, 246, 254, 258, 274, 294-295, 325, 333, 337, 383, 392, 405, 414-415, 417-418, 421, 432

Weeding, 52, 89, 204, 337Wet-sieved, 399-400Wind-dispersed, 29, 91, 96, 180, 211Winnowing, 71-72, 76-77, 79, 92, 117,

202, 208, 214, 246, 392Winter-grown, 328Winter-sown, 259Xeromorphic, 45Xerophyte, 45, 378Yield, 25, 27, 47-48, 53, 62, 67-68, 71, 73,

78, 80, 82-83, 85-86, 123, 143, 174, 189, 221, 252, 254, 259, 269-270, 273, 296-297, 327-328, 336-337, 366, 376-379, 382, 385-386, 390-392, 395-396, 410, 414-418, 421

Younger Dryas, 351-353, 356, 358, 380

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