A Manual for the Identification of PLANT SEEDS AND FRUITS

A Manual for the Identification of PLANT SEEDS AND FRUITS

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Groningen Archaeological StudiesVolume 23

Editorial boardProf. dr. D.C.M. Raemaekers

Prof. dr. P.A.J. A�emaDr. C. Çakirlar

Prof. dr. R.T.J. CappersProf. dr. P.D. Jordan Prof. dr. S. Voutsaki

Groningen Institute of ArchaeologyPoststraat 6

NL-9712 ER Groningenthe Netherlands

[email protected]

Websitewww.gas.ub.rug.nl

Publishers’ address

Barkhuis

Zuurstukken 37 9761 KP Eelde the Netherlands

Tel. 0031 50 3080936 Fax 0031 87 7844285

[email protected] www.barkhuis.nl

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A Manual for the Identification of

PLANT SEEDSAND FRUITS

R.T.J. Cappers & R.M. Bekker

Barkhuis &

University of Groningen Library

Groningen 2013

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Book cover design: Nynke Tiekstra, ColtsfootMedia - NoordwoldeBook interior design and typese�ing: Nynke Tiekstra, ColtsfootMedia - NoordwoldePhoto's: F. Bo�ema, R.T.J. Cappers, D. Fennema & J. JansCopy editor: Suzanne NeedsProofreader: Heather Sangster

Cover: Infructesence of Black medick (Medicago lupulina; le�) and fruit with seeds of Common whitlowgrass (Erophila verna; right).

ISBN: 9789491431265

This volume is part of the Digital Plant Atlas, a joint project of the Groningen Institute of Archaeology (GIA) of the University of Groningen (the Netherlands) and the German Archaeological Institute (DAI, Berlin, Germany)

Copyright © 2013 Groningen Institute of Archaeology (University of Groningen)

All rights reserved. No part of this publication or the information herein may be reproduced, stored in a retrieval system, or transmi�ed in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, without prior permission from the Groningen Institute of Archaeology (University of Groningen).

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.

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5

Cont

ents Contents

Introduction Organization of the manual 7

Family, genus, and species 9

Types of seeds and fruits 10

References 12

Plant family Amaranthaceae 16

Apiaceae 28

Asteraceae 44

Boraginaceae 66

Brassicaceae 76

Caryophyllaceae 94

Convolvulaceae 102

Cyperaceae 108

Fabaceae 122

Geraniaceae 146

Juncaceae 152

Lamiaceae 156

Malvaceae 164

Plantaginaceae 174

Poaceae 186

Polygonaceae 216

Ranunculaceae 228

Rosaceae 240

Solanaceae 254

Indices Index 1: Common names 262

Index 2: Scientific names 265

Index 3: Glossary of botanical terms 272

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Inho

ud Inhoud

Inleiding Opzet handboek 7

Familie, geslacht en soort 9

Zaad- en vruchttypen 10

Literatuur 12

Plantenfamilie Amaranthaceae 18

Apiaceae 30

Asteraceae 46

Boraginaceae 68

Brassicaceae 78

Caryophyllaceae 96

Convolvulaceae 104

Cyperaceae 110

Fabaceae 124

Geraniaceae 148

Juncaceae 154

Lamiaceae 158

Malvaceae 166

Plantaginaceae 176

Poaceae 188

Polygonaceae 218

Ranunculaceae 230

Rosaceae 242

Solanaceae 256

Indexen Index 1 Nederlandse plantennamen 264

Index 2 Wetenschappelijke plantennamen 267

Index 3 Botanische termen 274

Seeds and fruits of weed species found in a sample of flax from Argentina.

6

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Organization of the manual 7

Intr

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tion Introduction

Organization of the manual

To identify plant seeds and fruits, it is important to know to which family of plants they belong. To further identify them to species, one is able to narrow down the possibilities using both the scientific literature and reference collections. Especially in families with many genera, the morphology of fruits and seeds can vary considerably. This variation is a result of the fact that the grouping of genera into families is not only based on characteristics dealing with generative plant parts. In addition, specific adaptations to seed dispersal mechanisms have occurred within more than one family.

To narrow down the identification of fruits and seeds to a particular plant family, it is helpful to have some insight into the way in which the flowers, and in a later stage the fruits, are attached to the plant. It is also important to determine the dispersal unit: a seed, a fruit, or a fruit that has merged with parts of the flower.

This manual illuminates the variation in seeds and fruits belonging to 19 plant families (figure 1). These particular plant families have been included because the fruits and seeds of the wild plant species of these families often occur both in seed mixtures and in archaeological contexts. The emphasis is on the morphology of the wild species, as expressed in the choice of illustrative examples.

AsteraceaeFabaceaePoaceae

LamiaceaeMalvaceae

CyperaceaeApiaceae

BrassicaceaeRosaceae

CaryophyllaceaeSolanaceae

BoraginaceaeRanunculaceaeAmaranthaceaePlantaginaceae

ConvolvulaceaePolygonaceae

GeraniaceaeJuncaceae

0

0 5000 10000 15000 20000 25000

200 400 600 800 1000 1200 1400 1600 1800

Figure 1 Number of

genera (brown, numbers

on top axis) and species

(green, numbers on bottom

axis) per plant family that

have been described

worldwide (adapted from

Mabberley [2008]). �

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8 A Manual for the Identification of Plant Seeds and Fruits | R.T.J. Cappers & R.M. Bekker

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The introduction to each plant family briefly explains issues of taxonomy. Where applicable, attention is paid to nomenclature, subdivisions within the family, and defining characteristics that distinguish the family from other families. Recent DNA research has provided many new insights into the relationship between genera. Based on this research, some families have recently been split, while others have been combined. For this manual, scientific family names based on the type genus are used. The delineation of the families is based on Mabberley (2008).

This short introduction is followed by a paragraph describing the primary types of inflorescence and infructescence within each family. These terms refer to the way in which the flowers, and in a later stage the fruits, are attached to the stem (figure 2). A solid understanding of this distinction can make it much easier to place fruits and seeds in their correct family. Unfortunately, the way in which plants are grouped in terms of inflorescence and infructescence can be quite complicated and, in addition, can be difficult to determine when the pedicels of the flowers or fruits themselves are very short. Another major distinction, which will be elaborated on in the section ‘Fruit and seed’, is between species with multiple fruits and those with compound fruits. For more detail about the varies types of inflorescence, the reader is referred to botanical dictionaries and floras.

A paragraph titled ‘Fruit and seed’ describes the different types of fruits and the variation in seed morphology within the plant family. This manual follows the fruit typology of Spjut (1994). The accompanying photos show fruits and seeds native to various parts of the world whose appearance is representative for the entire family.

The diagnostic features of the fruits and seeds sometimes provide a clue to the plant’s dispersal methods, as noted in the paragraph titled ‘Diaspore’. This paragraph indicates which dispersal units (that is, diaspores) are most commonly found within that particular family. A diaspore is the seed-containing part of the plant, which eventually detaches to ensure the

� Figure 2 Inflores-

cence (left; 21 June 2009)

and infructescence (right;

11 July 2009) of

Buckwheat (Fagopyrum

esculentum). The flowers

and fruits are united in

panicles.

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Family, genus, and species 9

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creation of offspring in a new location. This dispersal unit can be loose seeds, but it can also be one or more seeds within a fruit or even entire fruits fused to parts of the flower. People often use the term seed dispersal, but what they actually mean is diaspore dispersal, which includes the dispersal of fruits and of fruits fused to parts of the flower.

For plants that are planted and harvested by humans, the dispersal unit can differ from that found in the wild, as determined by the method of harvest and the timing of harvest with respect to the ripening process. This explains why in seed samples we sometimes find dispersal units that deviate from the natural dispersal units (e.g. Cappers & Neef 2012, p. 286). It also explains why we sometimes encounter fragments of the infructescences or vegetative parts such as roots.

The paragraph titled ‘Heterocarpy and seed dimorphism’ discusses a phenomenon encountered in some families whereby a single plant can form more than one type of fruit (heterocarpy) or seed (seed dimorphism). This variation in fruit and seed morphology often relates to differences in dispersal mechanism and even in germination. For example, some species in the family Asteraceae form, within a single flowering head, fruits with a pappus and fruits without a pappus. The fruits with a pappus are adapted to effectively disperse over longer distances, for example by means of wind and/or water. The fruits without a pappus are often smaller, which allows them to more easily penetrate into the ground and thus contribute to a supply of seeds in a spot that has already been proven to be favourable by the mother plant.

Providing a comprehensive overview of the variation in fruit and seed morphology within the selected plant families is beyond the scope of this manual. Instead, a selection of genera that are representative of the respective families is presented. The selection of fruits and seeds in the concise seed atlas that concludes each chapter is mostly based on the Dutch flora (Van der Meijden, 2005). This concise seed atlas depicts fruits and seeds within a frame, accompanied by a scale. In addition, each image includes a silhouette that is depicted at the same scale for each of the images relating to that family (figure 3). For more extensive images, please refer to the publications of the Digital Plant Atlas project at www.plantatlas.eu.

Family, genus, and species

Plants are categorized in a hierarchical system based on relatedness. In the context of this manual, the most important levels are family, genus, and species. Family names and genus names consist of a single word. Species names consist of two words, namely, the genus name and the species name. Family names often end in -eae or -ae. In contrast, the endings of genus and species names vary.

New insights into relatedness have resulted in changes in the nomenclature and even reorganizations within plant families. The recent

� Figure 3 Photo of

Coriander (Coriandrum

sativum) from the seed

atlas. The 1 mm scale

pertains to the colour

image. The small

silhouette represents

the actual size of

the fruit depicted.

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large-scale research into certain loci of plant DNA, in particular, has led to species being reassigned to different families. When a species is reassigned to a different family, this need not necessitate changes to the species name. The species name does change, however, if new insights have led to the reorganization of species within a genus or to a species being assigned to a different genus. In order for researchers to be able to make the link between the new, so-called valid name and the older, so-called invalid name, the author’s name is added after the species name.

Some species are subdivided into subspecies or varieties. In that case, a third name is added to the species name, preceded by the abbreviation ssp. (subspecies) or var. (variety). Scientific names for genus, species, subspecies, and variety are set in italics; family names are not. The common names are written with a capital letter on the first word to avoid any confusion between adjectives and words belonging to the proper name of the plant (for example, creeping Cockspur versus Creeping Buttercup). If a plant has no common name, the binomial is given without parentheses. Nomenclature of common English names of plant follows Stace (2010).

Types of seeds and fruits

A seed develops from an ovule and becomes enclosed by a fruit or aril (figure 4). Plants whose seeds are not enclosed by a fruit mostly produce their seeds in cones and are often called gymnosperms (figure 5).

In angiosperms the ovules and thus the developing seeds are enclosed by a fruit. The distinction between seed and fruit presents no problem if the seed does not connate with the fruit (figure 6). Often the seed is an independent dispersal unit.

There are also plants in which the seed and fruit connate. One-seeded fruits in which the fruit is often not much larger than the seed are frequently wrongly called ‘seed’ (figure 7).

When the fruit is the dispersal unit, other plant parts may also become part of the dispersal unit. This may result in the fruit itself not being visible anymore. An example of plant parts that may partially or completely enclose the fruit are bracts. In cereals, such bracts can be distinguished as separate pieces of chaff (figure 8). In some species, these types of bracts are fused, as is the case with the green husk surrounding the fruit of the Walnut (figure 9). In the latter case, the tissue surrounding the fruit is sometimes referred to as the exocarp.

Some fruits are composed of visibly distinct layers. This is the case for stone fruits. In such fruits the inner layer is firm and is called endocarp. The endocarp is often enclosed by a middle layer, called mesocarp, and an outer layer, called epicarp. An example of a stone fruit with a clear differentiation in layers is the Olive (figure 10). Sometimes the term exocarp is used instead of epicarp. In accordance with Spjut (1994), in this manual we use the term exocarp only for those plant parts that are not part of the fruit, e.g. the husk surrounding the fruit of a Walnut (figure 9).

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Types of seeds and fruits 11

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Figure 4Seed of the Yew (Taxus baccata) surrounded by an aril (longitudinal section).

Figure 5Cone and seed of the Giant sequoia (Sequoiadendron giganteum).

Figure 6Longitudinal section of the Melon (Cucumis melo), a fruit with seeds.

Figure 7Complete fruit of the Sunflower (Helianthus annuus) and a longitudinal section of the fruit containing the seed.

Figure 8 Fruit of the Einkorn (Triticum monococcum) covered by chaff (left) and without chaff (right). Cereals have 1-seeded fruit.

Figure 9 Fruit of the Walnut (Juglans regia) with part of the exocarp (left), fruit without exocarp (middle), and a longitudinal section of a fruit containing the seed (right).

Figure 10 Complete fruit of the Olive (Olea europaea) (left), fruit whose mesocarp and epicarp (shell) were half removed (middle), and a cross-section through the fruit and seed (right). The epicarp and mesocarp are purple in colour, the endocarp is brown, and the seed is white.

Figure 11 Infructescence of the Elderberry (Sambucus nigra).

Figure 12 Multiple fruit of the Yellow-flowered strawberry (Potentilla indica) (top). A highly magnified fleshy axis contains a large number of small, 1-seeded fruit (bottom).

Figure 13 Exterior (top) and longitudinal section (middle) of the compound fruit of the Fig (Ficus carica). The fleshy axis is closed but for a small opening at the bottom. At the receptacle there are several hundred flowers. After pollination by a Fig wasp, a 1-seeded fruit may develop in each of these flowers (bottom).

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tion

Flowers are often arranged in a characteristic pattern. This pattern is called inflorescence, and the various types are indicated with specific terms. The term infructescence is used for a characteristic arrangement of fruit (figure 11).

In this manual no specific terms are used for the different ways fruits can be arranged. However, in accordance with Spjut (1994), we distinguish between simple fruits, multiple fruit, and compound fruit. A simple fruit develops from a single flower with a more or less isolated position in the infructescence. A multiple fruit develops from one flower with multiple pistils. In the ovary of each pistil, one or more seeds ripen (figure 12). A compound fruit develops from several flowers, each with its own pistil (figure 13). The transition from compound fruit to infructescence may sometimes be arbitrary.

References

Sources cited in the text:Brückner, C. (2000): Clarification of the carpel number in Papaverales,

Capparales, and Berberidaceae. The Botanical Review 66 (2), pp. 155-307.

Cappers, R.T.J. & R. Neef (2012): Handbook of plant palaeoecology. Groningen Archaeological Studies no. 19. Groningen: Barkhuis & Groningen University Library.

Imbert, E. (2002): Ecological consequences and ontogeny of seed heteromorphism. Perspectives in Plant Ecology, Evolution and Systematics 5 (1), pp. 13-36.

Mabberley, D.J. (2008; 3rd ed.): Mabberley’s plant-book. A portable dictionary of plants, their classification and uses. Cambridge: Cambridge University Press.

Meijden, R. van der (2005; 23rd ed.): Heukels’ flora van Nederland. Groningen: Wolters-Noordhoff.

Spjut, R.W. (1994): A systematic treatment of fruit types. Memoirs of the New York Botanical Garden 70, pp. 1-182.

Stace, C. (2010, 3rd ed.): New flora of the British Isles. Cambridge: Cambridge University Press.

Publications of the Digital Plant Atlas project:Cappers, R.T.J., R.M. Bekker, & J.E.A. Jans (2012; 2nd ed.): Digital seed atlas of

the Netherlands. Groningen Archaeological Studies no. 3. Groningen: Barkhuis & Groningen University Library [with text in Dutch, English, and German; http://www.plantatlas.eu].

Cappers, R.T.J., R. Neef, & R.M. Bekker (2009): Digital atlas of economic plants (3 vols.). Groningen Archaeological Studies no. 9. Groningen: Barkhuis & Groningen University Library [http://www.plantatlas.eu].

Neef, R., Cappers, R.T.J., & R.M. Bekker (2012): Digital atlas of economic plants in archaeology. Groningen Archaeological Studies no. 17. Groningen: Barkhuis & Groningen University Library [http://www.plantatlas.eu].

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References 13

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tion

Further readings on the typology of seeds and fruits:Anderberg, A.-L. (1994): Atlas of seeds Part 4: Resedaceae-Umbelliferae.

Stockholm: Swedish Museum of Natural History.Berggren, G. (1969): Atlas of seeds Part 2: Cyperaceae. Stockholm:

Swedish Natural Science Research Council.Berggren, G. (1981): Atlas of seeds Part 3: Salicaceae-Cruciferae.

Stockholm: Swedish Museum of Natural History.Black, M., J.D. Bewley, & P. Halmer, eds. (2006): The encyclopedia of seeds:

Science, technology and uses. Trowbridge: Cromwell Press.Bojňanský, V., & A. Fargašová (2007): Atlas of seeds and fruits of Central and

East-European flora: The Carpathian Mountains region. Dordrecht: Springer.

Brouwer, W., & A. Stählin (1975): Handbuch der Samenkunde. Frankfurt-am-Main: DLG-Verlag-GmbH.

Corner, E.J.H. (1976): The seeds of dicotyledons (2 vols.): Cambridge: Cambridge University Press.

Kesseler, R., & W. Stuppy (2006): Seeds: Time capsules of life. London: Papadakis Publishers & Royal Botanical Gardens Kew.

Roth, I. (1977): Fruits of angiosperms. Berlin: Gebrüder Borntraeger.Stuppy, W., & R. Kesseler (2008): Fruits: edible, inedible, incredible.

London: Papadakis Publishers & Royal Botanical Gardens Kew.

Further readings on the dispersal mechanisms of seeds and fruits:Bresinsky, A. (1963): Bau, Entwicklungsgeschichte und Inhaltsstoffe der

Elaiosomen: Studien zur myrmekochoren Verbreitung von Samen und Früchten. In: Bibliotheca Botanica, Heft 126, pp. 1-54.

Genderen, H. van, L.M. Schoonhoven, & A. Fuchs (1996): Chemisch-ecologische flora van Nederland en België. Utrecht: KNNV Uitgeverij.

Heywood, V.H., R.K. Brummitt, A. Culham, & O. Seberg (2007): Flowering plant families of the world. Kew: Royal Botanical Gardens.

Kleyer, M., R.M. Bekker, I.C. Knevel, J.P. Bakker, K. Thompson, M. Sonnenschein, P. Poschlod, J.M. van Groenendael, L. Klimeš, J. Klimešová, S. Klotz, G.M. Rusch, M. Hermy, D. Adriaens, G. Boedeltje, B. Bossuyt, P. Endels, L. Götzenberger, J. Hodgson, A-K. Jackel, I. Kühn, D. Kunzmann, W.A. Ozinga, C. Römermann, M. Stadler, J. Schlegelmilch, H.J. Steendam, O. Tackenberg, B. Wilmann, J.H.C. Cornelissen, O. Eriksson, E. Garnier, A. Fitter, & B. Peco (2008): The LEDA Traitbase: A database of plant life history traits. Journal of Ecology 96 (6), pp. 1266-1274.

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PLANT FAMILY

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16

Amaranthaceae Love-lies-bleeding (Amaranthus caudatus)

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Introduction 17

Am

aran

thac

eae Amaranthaceae

Introduction

The Amaranthaceae family is named after the genus Amaranth (Amaranthus). Plants in the Goosefoot family have recently been subsumed under the Amaranth family. Genetic research has proven that the Goosefoot family and the Amaranth family are closely related; distinguishing between them at the taxonomic level of family is therefore no longer desirable. The Amaranth family now contains about 2000 species in 175 genera.

Inflorescence and infructescence

The flowers are small. Because of their green or brown colour, they are not very noticeable. The flowers and fruits are united in tight clusters distributed over the length of the stem or at the end of the stem (figure 1).

The fruit is sometimes surrounded by a perianth (the collective name for the sepals and petals) and bracteoles. During fruiting, the perianth and bracteoles can become enlarged and partly or completely surround the fruit. In Oraches (Atriplex), the fruit is surrounded by two bracteoles that have fused together distally. These bracteoles vary in morphology and size within Oraches and can be toothed and nubbly. Sometimes the perianth is noticeably swollen, as in the Strawberry goosefoot (Chenopodium foliosum) (figure 2).

Fruit and seed

In some genera, such as Amaranth (Amaranthus), the fruit and seed are not fused together. The fruit of Amaranths is membranous and contains a single seed. This seed is disc-shaped and usually black and shiny. Because the seeds of the different species in this genus are similar in shape, they are usually not identified below the genus level. A species determination is, however, possible based on the characteristics of the fruit and the perianth. In some (sub)species, such as Indehiscent amaranth (Amaranthus

Figure 1 Terminal

inflorescence of the

Purple Amaranth

(Amaranthus cruentus). �

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Am

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Figure 2 Strawberry

goosefoot (Chenopodium

foliosum) with fruits

surrounded by red,

swollen perianths. �

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Fruit and seed 19

Am

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eae

hybridus ssp. bouchonii), the fruit does not open, whereas in other (sub)species the fruit opens transversely. In the latter case, the surface of the break can be irregular, as in Guernsey pigweed (A. blitum), or regular, as in Short-tepalled pigweed (A. graecizans) (figure 3).

In many other genera, the fruit and seed are fused together. In some of these genera the fruit wall is thin, fragile, and easily separated from the seed. This kind of fragile fruit wall occurs in Summer-cypress (Bassia), Perennial glasswort (Salicornia), Spinach (Spinacia), Saltworts (Salsola), Goosefoots (Chenopodium), and Oraches (Atriplex). Of those Goosefoots and Oraches that occur in cultivated fields, it is often the seeds rather than

Figure 3 Infructescence of

Short-tepalled pigweed

(Amaranthus graecizans).

The tops of some of

the fruits have torn

or fallen off. �

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Am

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intact fruits that are found, because the fragile fruit is removed during threshing (Berggren, 1981). In the Goosefoots, the outermost layer of the fruit is also referred to as ‘exocarp’ (Berggren 1981; Cappers et al., 2012). Here, both layers are considered part of the fruit (figure 4).

Winged fruits occur in Bugseed (Corispermum) and Saltworts (Salsola). In Bugseed, the wings are formed by the margins of the fruit, whereas in Saltworts the wing is formed by the perianth that surrounds the fruit. The seed of Glassworts (Salicornia) is partly covered with hooked hairs (figure 5).

The orientation of the fruit in the perianth can be horizontal or vertical (figure 6). The fruits of Oraches are all oriented vertically, while within the genus Goosefoots both orientations occur. In the case of horizontal orientation, the fruit is attached to the receptacle along the flattened side. These kinds of fruits have a scar on just one side, in the middle. This scar is often still visible under oblique light on the seed itself, as in Fat-hen (Chenopodium album) (figure 4, bottom centre). Fruits oriented vertically on the receptacle lack this scar, as in Clammy goosefoot (C. pumilio) (figure 6).

� Figure 4 Fruit with

perianth (left), fruit

without perianth (centre),

and seed (right) of

Fat-hen (Chenopodium

album). The perianth

consists of five keeled

tepals. The top row shows

the top, the bottom row

shows the underside. The

scar on the perianth in the

bottom left image marks

the location where the

flower was once attached

to the stem. The scar on

the fruit in the bottom

centre image marks the

location where the fruit

was once attached to the

receptacle. In the bottom

right photo the scar is

only faintly visible.

Figure 5 The seed of

Common glasswort

(Salicornia europaea) is

partly covered with

hooked hairs. �

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Diaspore 21

Am

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Diaspore

The dispersal unit can be the seed; the 1-seeded fruit; or the 1-seeded fruit with perianths, bracteoles, or a swollen receptacle (table 1).

Table 1 Composition of the dispersal unit in several genera of Amaranthaceae.

Seed 1-Seeded Fruit

1-Seeded Fruit with Perianth

1-Seeded Fruit with Bracteoles

1-Seeded Fruit with

Receptacle

Salicornia ○Amaranthus ○ ○Corispermum ○Suaeda ○Chenopodium ○ ○Salsola ○Bassia ○Spinacia ○Atriplex ○Beta ○

Figure 6 Fruits of

Many-seeded goosefoot

(Chenopodium

polyspermum) (left) are

oriented horizontally,

whereas those of Clammy

goosefoot (C. pumilio)

(right) are oriented

vertically. �

� Figure 7 Ripe fruits of

Saltmarsh goosefoot

(Chenopodium

chenopodioides)

surrounded by

the perianth.

Top: A fruit from a

terminal cluster with

four-five perianths

that are joined only

at their bases.

Bottom: A fruit from

a lateral cluster with

mostly three completely

fused perianths.

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Am

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Heterocarpy and seed dimorphism

Heterocarpy is common in this family. For example, the type of perianth surrounding the fruits of Saltmarsh goosefoot (Chenopodium chenopodioides) varies depending on the fruit’s location along the stem (figure 7).

The variation in the bracteoles in Oraches (Atriplex) relates to the position of the fruit on the plant. In some species the fruit also varies in size and shape. Therefore it is not always possible to obtain a secure species-level identification. The variation is especially great in such species as Common orache (Atriplex patula), Spear-leaved orache (A. prostrata), and Grass-leaved orache (A. littoralis). The fruits of the first two of these species are usually not differentiated; only large fruits (with a diameter of at least 3.2 mm) are identified as Grass-leaved orache. The fruits of Halogeton Halogeton glomeratus and Fat-hen (Chenopodium album) and the seeds of Common glasswort (Salicornia europaea) are also heterocarpic (Imbert, 2002).

The number of fruits in fruit clusters of the Beet (Beta vulgaris) varies from (1-)3-5(-7). In the Beet from the eastern Mediterranean, the number of fruits is small.

Variation in the shape of the fruit can also be the result of the selection of seed stock. For example, the bracteoles of Spinach (Spinacia oleracea) possess a spine. This spine is absent in fruits used today as seed stock (figure 8).

� Figure 8 Fruits of

the Spinach (Spinacia

oleracea) encapsulated by

bracteoles, with three

long spines (left) and with

only the bases of the

spines (right).

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Seed atlas Amaranthaceae 23

Am

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Seed atlas Amaranthaceae

Amaranthus standleyanus Amaranthus standleyanus

Atriplex prostrata Atriplex prostrata

Axyris amaranthoides

250 μm

250 μm1 mm

250 μm

1 mm

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Am

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Bassia scoparia Bassia scoparia

Beta vulgaris Beta vulgaris

Chenopodium rubrum Chenopodium rubrum

250 μm 250 μm

250 μm1 mm

250 μm 250 μm

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Seed atlas Amaranthaceae 25

Am

aran

thac

eae

Corispermum intermedium

Polycnemum arvense Polycnemum arvense

Salicornia europaea

1 mm

250 μm

250 μm

250 μm

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Am

aran

thac

eae

Salsola kali Salsola kali

Spinacia oleracea Spinacia oleracea

Suaeda maritima

250 μm 250 μm

250 μm1 mm

250 μm

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INDICES

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262 Handboek voor het determineren van zaden en vruchten | R.T.J. Cappers & R.M. Bekker

Ajowan, 31Almond, 247Alpine cat’s-tail, 194Amaranth, 17Amphibious bistort, 218Anemone, 231-232Annual knawel, 97Annual marguerite, 52, 53Apple, 248-249Apple-of-Peru, 255, 257Arabian grass, 198Asafoetida, 34-35Avens, 245Bamboo, 187Baneberry, 230, 234Barley, 187, 190, 192, 194-195,

198-199, 201, 2052-row Barley, 201, 2056-row Barley, 201, 205Beak-sedges, 114Beet, 22Berry catchfly, 97Bindweed family, 103Bittersweet, 254, 256Black mustard, 79-80Black nightshade, 257Black-bindweed, 220, 222-223Bladder campion, 94, 96Bladder ketmia, 166, 170Bladder-sedge, 116Bladder-senna, 127-128Bladder-sennas, 127Bloody crane’s-bill, 149Bluestem, 194Borage, 67, 71Borage family, 66, 157Bramble, 248Brambles, 241, 248Branched plantain, 176Bread wheat, 196-197, 201, 205Bristle club-rush, 109-110Bristle oat, 205Broad bean, 123

Broadleaf sermountain, 33Broad-leaved dock, 220Broad-leaved everlasting-pea, 126Broomcorn, 201, 204-205Broomrape family, 175Brown beak-sedge, 113Brown sedge, 111-112Buckwheat, 8, 217, 220-221Bugles, 159Bugseed, 20Bulbous barley, 199Burdocks, 53Buttercup family, 229Buttercups, 229, 232Cabbage family, 77Cabbages, 77Canary-grasses, 194Cape-marigold, 52Carrot family, 29, 35, 166Celery-leaved buttercup, 232-233Charlock, 79-80, 83-85Cherries, 241Cherry plum, 247Chinese mallow, 166Chinese wisteria, 123Cinquefoils, 245-246Clammy goosefoot, 20-21Claries, 159Cloud grass, 188Clove pink, 95Clovers, 133Clustered clover, 134Clustered dock, 221-222Coast button grass, 198Cockleburs, 48, 53Coffee senna, 134Common bistort, 219Common club-rush, 113Common comfrey, 66-67, 69, 71Common cottongrass, 113Common couch, 188-192Common dandelion, 44Common fleabane, 229

Common glasswort, 20, 22Common hemp-nettle, 158Common millet, 203, 205Common mouse-ear, 96Common orache, 22Common reed, 194-195Common sorrel, 217, 220Common spike-rush, 114Common stork’s-bill, 149Common vetch, 122Common whitlowgrass, 77, 81Common yellow-sedge, 111-112Copse-bindweed, 222-223Coral-necklace, 97Coriander, 9, 32-33Corn buttercup, 232-233Corn toadflax, 181Cornflower, 50Cow parsley, 33Cow-itch, 133Crown vetch, 129-130Cultivated carrot, 34-35Cut-grass, 188-189Daisy family, 29, 45Darnel, 198-199Dead-nettle family, 157Dead-nettles, 157, 159Deergrass, 109-110Docks, 217, 220-222Dodders, 103Dog-rose, 240Dutch rose, 249Dwarf cherry, 247Dwarf mallow, 165-166Early forget-me-not, 68Egyptian plum, 72Einkorn, 11, 195, 197, 201, 205Elderberry, 11Emmer, 195, 197, 200, 205False fox-sedge, 114False indigo, 127Fat-hen, 20, 22Fennel, 33

Index 1 Common names

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Index 1 Common names 263

Fenugreek, 125Field bindweed, 103-104Field penny-cress, 78, 83Field wood-rush, 154Fig, 11Fine-leaved water-dropwort, 34-35Flamboyant, 124Flat-sedge, 113Floating club-rush, 109Fluellens, 176, 181Forget-me-nots, 67, 71Foxglove, 180-181Foxtail barley, 195Foxtails, 194French bean, 131, 134-135Garden rocket, 81Germanders, 159Giant hogweed, 28Giant sequoia, 11Giant stock-bean, 131, 133Glaucous sedge, 114, 218Globe-thistles, 48Goldenseal, 233Goosefoot family, 17Goosefoots, 19-20Grass family, 109, 187Grass pea, 138Grass-leaved orache, 22Great fen-sedge, 114-116Greater honeywort, 71Greater knapweed, 45, 47Greater plantain, 174, 176, 181Greater sea-spurry, 97Greater spearwort, 228-229Grey hair-grass, 192-193Guernsey pigweed, 19Hairy buttercup, 232-233Hairy finger-grass, 197Hairy sedge, 115Hairy tare, 131, 133, 137Hard wheat, 194, 197, 201Hawkweed oxtongue, 47Heath rush, 154Hedge bindweed, 102Hedge hyssop, 181Heliotrope, 69

Hellebores, 233Henbane, 255, 257Herb-Robert, 147Himalayan sorbaria, 242Hollyhock, 164-165, 170Holy-grass, 188, 192Honesty, 76, 81-84Honeywort, 67, 71Hybrid chokeberry, 243Hyssop, 159Indehiscent amaranth, 17Ivy family, 29Juneberry, 243Kelch-grass, 198Kidney vetch, 124Knawels, 97Knotgrass, 216, 218, 220Knotgrasses, 217, 221Knotted hedge-parsley, 36-37Knotweed family, 217Knotweeds, 217-218, 221-222Laburnum, 141Large-flowered hemp-nettle, 156Larkspurs, 230, 233Least mallow, 166Lentil, 132Lesser canary-grass, 197Lesser celandine, 232Lesser hawkbit, 53, 55-56Lesser sea-spurry, 97Lime, 168-170Limes, 166Long-bracted sedge, 113Love-in-a-mist, 232-233Love-lies-bleeding, 16Lucerne, 103Lungwort, 71Lupins, 133Lyme-grass, 190Maize, 187, 201, 204-205Malling toadflax, 181Mallow family, 165Mallows, 165Many-seeded goosefoot, 21Marsh cinquefoil, 246Marsh dock, 221-222

Marsh pennywort, 29Marsh yellow-cress, 77Marsh-mallow, 165, 170Marsh-marigold, 232-233Marshworts, 29Mat-grass, 188Meadow clary, 159Meadow crane’s-bill, 148Meadow fescue, 195Meadow-grasses, 187Meadowsweets, 245Medicks, 129Medlar, 242Melilots, 129, 133Melon, 11Milk-vetches, 129Millets, 187, 201, 205Morning glories, 105Mouse-ears, 96Mousetail, 230, 233Mudwort, 181Musk thistle, 45-46Musk-mallow, 166-167Narrow-leaf water-dropwort, 35-36Nightshade family, 255Nightshades, 255Nile acacia, 124Nipplewort, 48-49Oat, 198, 201, 205Oats, 187, 192, 198, 201, 205Olive, 10-11Oraches, 17, 19-20, 22Oxeye daisy, 48Pale persicaria, 219Parsley-piert, 241Parsley-pierts, 241Parsnip, 35Pasqueflower, 232Pea, 136Pea family, 123Peach, 247Peanut, 137Peas, 133Perennial glasswort, 19Perennial rye-grass, 189Phacelia, 67, 71, 75

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Pink family, 95Plantains, 175-176, 179, 181Pod mahogany, 133Prickly lettuce, 51Purple amaranth, 17Quaking-grass, 195Red dead-nettle, 159Redshank, 220, 222, 224Restharrows, 133Rhubarb, 217Rhubarbs, 221-222Ribwort plantain, 178-179, 181Rice, 187, 189Rose family, 229, 241Rose of Jericho, 85Roselle, 167Rosemary, 159Rough cocklebur, 52Round-fruited rush, 154Ruptureworts, 97Rush family, 109, 153Rushes, 153, 187Rye, 187, 201-202, 205Rye-grasses, 188Sainfoin, 131Saltmarsh flat-sedge, 113Saltmarsh goosefoot, 21-22Saltworts, 19-20Sand sedge, 115Sandworts, 96Sanicle, 31Savory, 159Scottish dock, 221-222Sea club-rush, 109, 113Sea mayweed, 52Sea plantain, 176Sea rocket, 79-80, 85Sea rush, 154Sea-hollies, 31, 33Sea-kale, 78-80Sea-spurreys, 97Sedge family, 109Sedges, 110-112, 114, 116, 187, 218,

220Shoreweed, 175, 179Short-tepalled pigweed, 19

Sickle medick, 134Silverweed, 137Skullcap, 157-158Skullcaps, 159Slender rush, 153Slender tufted-sedge, 114Small mallow, 166-167Small melilot, 134Small toadflax, 176Small-flowered crane’s-bill, 148Smooth tare, 124Snapdragon, 181Soft-rush, 152-153Soybean, 133Spanish broom, 126Spanish-needles, 50Spear-leaved orache, 22Speedwells, 175-176Spelt, 201, 205Spignel, 35Spike-rushes, 114Spinach, 19, 22Spotted dead-nettle, 157-158Spurreys, 97St Lucie cherry, 247Starry lady’s-mantle, 241Steeple-bush, 244Stocks, 83Strapwort, 97Strawberry goosefoot, 17-18Subterranean clover, 137, 138Sugar cane, 187Summer pheasant’s-eye, 233Summer-cypress, 19Sunflower, 11, 49-50Sweet cicely, 35-36Sweet vernal-grass, 188Swine-cress, 77, 79Tall melilot, 125Tansy, 47Teak, 157Thistles, 45, 51, 53Thorn-apple, 256Thorn-apples, 255Three-nerved sandwort, 97Tidy-tips, 52

Toadflaxes, 181Tobaccos, 255Tomato, 257Tomatoes, 255Toothed medick, 129Touch-me-not, 129-130True fox-sedge, 115Tufted sedge, 115Twisted acacia, 134Velvetleaf, 165, 170Vervain family, 157Vetches, 123-133Wall barley, 193Wall lettuce, 48Wall speedwell, 176Walnut, 10-12Waterleaf family, 67Water-pepper, 219, 221Water-starworts, 181Wheat, 187, 195, 197, 201, 205White beak-sedge, 110, 113-114White campion, 95White henbane, 256White lupin, 131White sedge, 113Wild basil, 159Wild carrot, 29, 34Wild cherry, 247Wild emmer, 200Wild plum, 247Wild radish, 82-85Wild strawberry, 246Wild-oat, 193, 198, 200Willow dodder, 104Winter wild-oat, 198, 200Wood avens, 245-246Wood dock, 221Wood speedwell, 177Wood-rushes, 153Woundworts, 159Yellow chamomile, 45, 50Yellow-flowered strawberry, 11, 245Yew, 11Yorkshire-fog, 186

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Index 2 Scientific names 265

Abutilon, 165, 170Abutilon theophrasti, 170-171Acacia nilotica, 124Acacia tortilis, 134Acanthospermum hispidum, 57Aconitum, 231, 233Aconitum vulparia, 231, 236Actaea, 230-231, 233-234, 236Actaea pachypoda, 234Actaea spicata, 236Adonis, 231-232Adonis aestivalis, 233Adonis flammea, 236Aethionema, 85Aethusa cynapium, 38Ageratina altissima, 57Ageratum conyzoides, 57Ageratum houstonianum, 57Agrimonia, 245Agrimonia eupatoria, 250Agrostemma githago, 98Agrostis nebulosa, 188Ajuga, 157, 159Ajuga reptans, 160Alcea, 164-166, 170Alcea rosea, 170-171Alchemilla, 241, 245Alchemilla glabra, 250Alhagi graecorum, 139Alliaria petiolata, 86Alopecurus, 194Alopecurus myosuroides, 206Alopecurus pratensis, 206Althaea, 165-166, 170Althaea hirsuta, 171Alysicarpus monilifer, 138Alyssum alyssoides, 86Amaranthaceae, 7, 16, 21Amaranthus, 17, 21Amaranthus blitum, 19Amaranthus caudatus, 16Amaranthus cruentus, 17

Amaranthus graecizans, 19Amaranthus hybridus ssp.

bouchonii, 17Amaranthus standleyanus, 23Ambrosia artemisiifolia, 57Ambrosia psilostachya, 58Ambrosia trifida, 58Amelanchier, 245Amelanchier lamarckii, 243, 250Ammi majus, 38Amphicarpaea bracteata, 138Amsinckia micrantha, 73Anastatica hierochuntica, 85Anchusa arvensis, 73Anemone, 231-232Anemone nemorosa, 236Anethum graveolens, 38Anoda, 165Anoda cristata, 172Anthemis cotula, 52-53, 58Anthemis tinctoria, 45, 50Anthoxanthum, 192Anthoxanthum odoratum, 188, 206Anthriscus caucalis, 39Anthriscus cerefolium, 39Anthriscus sylvestris, 33Anthyllis vulneraria, 124, 139Antirrhinum, 175-176, 178, 181Antirrhinum majus, 183Aphanes, 241Apiaceae, 7, 28, 166Apium, 29Apium graveolens, 39-40Aquilegia, 231Aquilegia vulgaris, 236Arabidopsis thaliana, 86Arabis hirsuta ssp. hirsuta, 86Arachis hypogaea, 137Araliaceae, 29Arctium, 53Arctium minus, 58Arenaria, 96

Arenaria serpyllifolia, 98Aronia, 245Aronia x prunifolia, 243, 250Artedia squamata, 30Artemisia verlotiorum, 58Aruncus, 244-245, 250Aruncus dioicusAsperugo procumbens, 73Aster, 45Asteraceae, 7, 9, 29, 44, 123, 229,

245Astragalus, 128-129Astragalus eremophilus, 128Astragalus glycyphyllos, 139Atriplex, 17, 19, 21-22Atriplex littoralis, 22Atriplex patula, 22Atriplex prostrata, 22-23Atropa, 255Atropa bella-donna, 258Aubrieta deltoidea, 86Avena, 187, 192, 198, 201, 205Avena fatua, 193, 198, 200, 207Avena sativa, 198, 201, 205, 207Avena sterilis, 198, 200, 207Avena strigosa, 205Axyris amaranthoides, 23Baccharis halimifolia, 59Ballota nigra ssp. meridionalis, 160Bambusa, 187Baptisia australis, 127Barbarea vulgaris, 86Bassia, 19, 21Bassia scoparia, 24Berteroa incana, 87Beta, 21Beta vulgaris, 22, 24Bidens bipinnata, 50Blysmus, 109, 112Blysmus compressus, 113Blysmus rufus, 113, 117Bolboschoenus, 109, 112

Index 2 Scientific names

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Bolboschoenus maritimus, 109, 113, 117

Bombacaceae, 165Boraginaceae, 7, 66, 157Borago, 67Borago officinalis, 67, 71, 73Brassica, 77Brassica juncea, 77Brassica napus, 77Brassica nigra, 79-80, 87Brassica oleracea, 77Brassica rapa, 77Brassicaceae, 7, 76Briza media, 195Bromopsis inermis ssp. inermis,

207Brunnera macrophylla, 73Bunias orientalis, 87Caesalpinioideae, 123-125, 127,

133-134Cakile maritima, 79-80, 85, 87Calamintha, 157Calendula officinalis, 53-54, 59Calepina irregularis, 87Callistephus chinensis, 59Callitrichaceae, 175, 178Callitriche, 175-176, 178, 181-182Callitriche cophocarpa, 181Callitriche hermaphroditica, 183Caltha, 231Caltha palustris, 232-233, 236Caltha palustris ssp. palustris, 236Camelina sativa ssp. sativa, 87Canavalia ensiformis, 131, 133Capsella bursa-pastoris, 81-82, 88Cardamine chenopodifolia, 85Cardamine pratensis, 88Carduus, 51Carduus acanthoides, 59Carduus crispus, 59Carduus nutans, 45-46, 59Carex, 112, 114, 116, 220Carex acuta, 114Carex arenaria, 115Carex crawfordii, 115

Carex curta, 113Carex disticha, 111-112Carex elata, 115Carex extensa, 113Carex flacca, 114, 218Carex hirta, 115Carex lepidocarpa, 117Carex oederi ssp. oedocarpa, 111-112Carex otrubae, 114Carex spicata, 117Carex vesicaria, 116Carex vulpina, 115Carthamus tinctorius, 60Caryophyllaceae, 7, 94Caryophyllus, 95Centaurea benedicta, 60Centaurea cyanus, 50Centaurea scabiosa, 45, 47Centaurea solstitialis, 60Cerastium, 96, 98Cerastium fontanum, 96, 98Cerastium fontanum ssp. vulgare,

96, 98Cerinthe, 67, 71Cerinthe major, 71Chaenorhinum, 175-176, 178, 181Chaenorhinum minus, 176, 183Chaenorhinum origanifolium, 181Chaerophyllum temulum, 40Chenopodium, 19, 21Chenopodium album, 20, 22Chenopodium chenopodioides,

21-22Chenopodium foliosum, 17-18Chenopodium polyspermum, 21Chenopodium pumilio, 20-21Chenopodium rubrum, 24Chondrilla juncea, 60Cicer arietinum, 139Cichorium endivia, 60Cichorium intybus, 60Cirsium, 53Cirsium acaule, 61Cirsium arvense, 61Cladium, 112

Cladium mariscus, 114-116, 118Clematis, 231Clematis vitalba, 232, 237Clinopodium acinos, 160Clinopodium vulgare, 159Cochlearia danica, 88Coincya monensis, 88Colutea, 127Colutea arborescens, 128, 139Comarum, 245Comarum palustre, 246, 250Compositae, 45Conium maculatum, 40-41Conringia orientalis, 88Consolida, 230-231, 233Consolida regalis, 237Convolvulaceae, 7, 102Convolvulus, 103Convolvulus arvensis, 104, 106Convolvulus sepium, 102Cordia, 71-72Cordia myxa, 72Coriandrum sativum, 9, 32-33, 41Corispermum, 20-21Corispermum intermedium, 25Corrigiola, 97Corrigiola litoralis, 98Corynephorus canescens, 192-193Cotoneaster, 245Cotoneaster integerrimus, 251Crambe maritima, 78-80, 88Crataegus, 245Crataegus monogyna, 251Crotalaria juncea, 139Cruciferae, 77Crypsis aculeata, 189Cucubalus baccifer, 97Cucumis melo, 11Cuminum cyminum, 41Cuscuta, 103Cuscuta europaea, 106Cuscuta pedicellata, 103Cuscutaceae, 103Cymbalaria, 175-176, 178, 181Cymbalaria muralis, 183

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Cynodon dactylon, 208Cynoglossum officinale, 71, 73Cyperaceae, 7, 108, 187, 218Cyperus, 112Cyperus conglomeratus, 110-111Cyperus esculentus, 118Cytisus scoparius, 140Dactyloctenium, 198Dactyloctenium aegypticum, 198Datura, 255Datura stramonium, 256, 258Daucus carota ssp. carota, 29, 34Daucus carota ssp. sativus, 34-35Delonix regia, 124Descurainia sophia, 89Dianthus caryophyllus, 95Dianthus deltoides, 98Dichantium, 194Digitalis, 175-176, 178Digitalis purpurea, 180-181, 183Digitaria sanguinalis, 197, 208Dimorphotheca pluvialis, 52Diplotaxis muralis, 89Draba muralis, 89Echinacea angustifolia, 61Echinochloa crus-galli, 208Echinops, 48Echium, 67Echium vulgare, 70, 74Eclipta prostrata, 61Eleocharis, 112, 114Eleocharis multicaulis, 118Eleocharis palustris, 114Eleogiton, 109, 112Eleogiton fluitans, 109-110, 118Elytrigia repens, 188-192Emex spinosa, 222, 224-225Entada, 129Eranthis, 231Eranthis hyemalis, 237Eriophorum, 112Eriophorum angustifolium, 113Eriophorum latifolium, 118Eriophorum vaginatum, 108Erodium, 147, 149

Erodium cicutarium, 149-150Erophila verna, 77, 81, 89Eruca vesicaria, 81, 89Erucastrum gallicum, 89Eryngium, 31, 33Eryngium alpinum, 33Erysimum cheiranthoides, 90Faba, 123Faba vulgaris, 123Fabaceae, 7, 122Fagopyrum, 221Fagopyrum esculentum, 8, 217, 220,

226Fallopia, 217, 221-222Fallopia convolvulus, 220, 222-223,

226Fallopia dumetorum, 222-223Farsetia, 83Farsetia ramosissima, 84Ferula foetida, 34-35Festuca arundinacea, 209Festuca pratensis, 195Ficaria, 231Ficaria verna, 232, 237Ficus carica, 11Filipendula, 245Filipendula ulmaria, 251Foeniculum vulgare, 33, 42Fragaria, 245Fragaria vesca, 246, 251Galega officinalis, 140Galeopsis speciosa, 156, 160Galeopsis tetrahit, 158Galinsoga parviflora, 61Genista pilosa, 140Geraniaceae, 7, 146Geranium, 147-149Geranium dissectum, 150Geranium molle, 146, 148Geranium pratense, 148Geranium pusillum, 148Geranium robertianum, 150Geranium robertsianum, 147Geranium sanguineum, 149Geum, 245

Geum urbanum, 245-246, 251Glebionis segetum, 61Glechoma, 157Glechoma hederacea, 160Glycine, 133Glycine max, 140Gramineae, 187Gratiola, 175-176, 178Gratiola officinalis, 181, 183Gypsophila muralis, 98Halogeton glomeratus, 22Helianthus annuus, 11, 49-50, 62Heliotropium, 69Helleborus, 231, 233Helleborus foetidus, 237Heracleum mantegazzianum, 28Herniaria, 97Herniaria glabra, 99Hesperis matronalis, 90Hibiscus, 166, 170Hibiscus sabdariffa, 167Hibiscus trionum, 172Hieracium pilosella, 62Hieracium vulgatum, 62Hierochloe odorata, 188Hippocrepis, 129Hippocrepis comosa, 130, 140Hippuridaceae, 175, 178Hippuris, 176, 178-179, 182Hippuris vulgaris, 175, 177, 179, 184Hirschfeldia incana, 90Holcus lanatus, 186, 209Holosteum umbellatum, 99Honckenya peploides, 99Hordeum, 187, 190, 192, 194, 198,

201Hordeum bulbosum, 199Hordeum jubatum, 195Hordeum murinum, 193Hordeum vulgare, 205, 209Hordeum vulgare ssp. distichon,

205Hordeum vulgare ssp. vulgare, 205Hydrastis canadensis, 233Hydrocotylaceae, 29

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Hydrocotyle, 29Hydrophyllaceae, 67Hyoscyamus, 255Hyoscyamus albus, 256Hyoscyamus niger, 257-258Hyssopus, 157, 159Hyssopus officinalis, 160Iberis umbellata, 90Illecebrum, 97Illecebrum verticillatum, 99Inula conyzae, 62Ipomoea, 105Ipomoea carnea, 105Isatis tinctoria, 90Isolepis, 109, 112Isolepis setacea, 109-110, 119Iva xanthifolia, 62Jacobaea vulgaris, 62Juglans regia, 11Juncaceae, 7, 109, 152, 187Juncus, 153-154Juncus acutiflorus, 155Juncus compressus, 154Juncus effusus, 152-153Juncus maritimus, 154Juncus squarrosus, 154Juncus tenuis, 153Kickxia, 175-176, 178, 181Kickxia elatine, 184Labiatae, 157Lablab purpureus, 140Laburnum anagyroides, 141Lactuca sativa, 63Lactuca serriola, 51Lagurus ovatus, 194Lamiaceae, 7, 70, 156Lamiastrum galeobdolon ssp.

galeobdolon, 161Lamium, 157, 159Lamium maculatum, 157-158Lamium purpureum, 159, 161Lappula squarrosa, 74Lapsana communis, 48, 63Laserpitium latifolium, 33Lathyrus, 133Lathyrus ciliolatus, 138

Lathyrus latifolius, 126Lathyrus sativus, 138, 141Lathyrus sylvestris, 141Layia platyglossa, 52Leersia oryzoides, 188-189Leguminosae, 123Lens culinaris, 132, 141Leontodon saxatilis, 53, 55-56Leonurus cardiaca, 161Lepidium coronopus, 77, 79Lepidium didymum, 90Lepidium sativum, 91Leucaena leucocephala, 141Leucanthemum paludosum, 52-53Leucanthemum vulgare, 48, 63Leymus arenarius, 190Limosella, 175-176, 178Limosella aquatica, 181, 184Linaria, 175-176, 178, 181Linaria arvensis, 181Linaria vulgaris, 184Lithospermum arvense, 74Littorella, 175-176, 178-179, 182Littorella uniflora, 175, 179, 184Lobularia maritima, 91Lolium, 188Lolium multiflorum, 210Lolium perenne, 189, 210Lolium remotum, 210Lolium temulentum, 198-199Lotus pedunculatus, 141Lunaria annua, 76, 81-84, 91Lupinus, 133Lupinus albus, 131Lupinus angustifolius, 142Luzula, 153-154Luzula campestris, 154Luzula multiflora, 155Lycium, 255Lycium barbarum, 258Lycopersicon, 255Lycopus europaeus, 161Malcolmia maritima, 91Malus, 245Malus sylvestris, 248-249, 251Malva, 165-166, 170

Malva alcea, 172Malva moschata, 166-167Malva neglecta, 165-166Malva parviflora, 166Malva pusilla, 166-167Malva verticillata, 166Malvaceae, 7, 164Marrubium vulgare, 161Matricaria chamomilla, 63Matthiola, 83Matthiola tricuspidata, 83Medicago, 129Medicago falcata, 134Medicago lupulina, 4Medicago polymorpha, 129Medicago sativa, 103, 142Melilotus, 129, 133Melilotus altissimus, 125, 142Melilotus indicus, 134Melissa, 157Melissa officinalis, 161Mentha arvensis, 162Mespilus, 245Mespilus germanica, 242, 252Meum athamanticum, 35Mimosa pudica, 129-130Mimosoideae, 123-125, 127, 133-134Minuartia hybrida, 99Misopates, 175-176, 178, 181Misopates orontium, 180-181, 184Moehringia trinervia, 97, 99Moenchia erecta, 100Mucuna, 133Mucuna pruriens, 133Mycelis muralis, 48Myosotis, 67, 71Myosotis arvensis, 74Myosotis ramosissima, 68Myosoton aquaticum, 100Myosurus, 231Myosurus minimus, 230, 233, 237Myrrhis odorata, 35-36Nardus, 188Nasturtium microphyllum, 91Nepeta, 157Nepeta cataria, 162

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Neslia paniculata, 91Nicandra, 255Nicandra physalodes, 255, 257-258Nicotiana, 255Nicotiana tabacum, 258Nigella, 231-233Nigella arvensis, 238Nonea lutea, 74Ocimum basilicum, 162Oenanthe aquatica, 34-35Oenanthe silaifolia, 35-36Olea europaea, 11Omphalodes verna, 74Onobrychis, 131Onobrychis viciifolia, 142Ononis, 133Ononis repens ssp. spinosa, 142Origanum vulgare, 162Ornithopus, 129Ornithopus sativus, 142Orobanchaceae, 175Oryza, 187Oryza sativa, 189, 211Panicum, 187, 201Panicum capillare, 211Panicum miliaceum, 203, 205Papilionaceae, 123Papilionoidae, 123-125, 127, 133-135Paspalum dilatatum, 211Pastinaca sativa ssp. sativa, 35, 42Pelargonium, 147Pennisetum, 187Pennisetum glaucum, 212Pentaglottis sempervirens, 75Persicaria, 217, 221Persicaria amphibia, 218, 226Persicaria bistorta, 219Persicaria hydropiper, 219, 221Persicaria lapathifolia, 219, 226Persicaria maculosa, 220, 222, 224Persicaria minor, 226Petasites hybridus, 63Petrorhagia prolifera, 100Petroselinum crispum, 43Phacelia tanacetifolia, 67, 71, 75Phalaris, 194

Phalaris canariensis, 212Phalaris minor, 197, 212Phaseolus coccineus, 143Phaseolus vulgaris, 135, 143Phleum, 194Phleum alpinum, 194Phragmites australis, 194-195Physalis, 255Physalis alkekengi, 259Picris echioides, 63Picris hieracioides, 47Pisum fulvum, 138Pisum sativum, 136, 143Plantaginaceae, 7, 174Plantago, 175-176, 178-179, 181Plantago arenaria, 176Plantago coronopus, 182Plantago lanceolata, 178-179, 181,

185Plantago major, 174, 176, 181Plantago maritima, 176Poa, 187Poa pratensis, 213Poaceae, 7, 109, 123, 186Polycarpon tetraphyllum, 100Polycnemum arvense, 25Polygonaceae, 7, 216Polygonum, 217, 221Polygonum aviculare, 216, 218, 220,

226Potentilla, 245-246Potentilla anserina, 137Potentilla indica, 11, 245Potentilla tabernaemontani, 252Prosopis farcta, 143Prunella vulgaris, 162Prunus, 241, 245-246Prunus avium, 247Prunus ceracifera, 247Prunus cerasus, 247Prunus domestica, 247, 252Prunus domestica ssp. domestica,

252Prunus dulcis, 247Prunus mahaleb, 247Prunus persica, 247

Pulicaria dysenterica, 229Pulmonaria officinalis, 71, 75Pulsatilla, 231Pulsatilla vulgaris, 232, 238Pyrus, 245Pyrus communis, 252Ranunculaceae, 7, 228Ranunculus, 229, 231-232Ranunculus arvensis, 232-233Ranunculus bulbosus, 238Ranunculus lingua, 228-229Ranunculus peltatus, 233Ranunculus sardous, 232-233Ranunculus sceleratus, 232-233Ranunculus subgen. Batrachium,

232Ranunculus subgen. Ranunculus,

232Raphanus raphanistrum, 82-85, 92Rapistrum rugosum, 92Rheum, 221-222Rheum x hybridum, 217Rhynchospora, 112, 114Rhynchospora alba, 110, 113-114Rhynchospora fusca, 113, 119Robinia pseudoacacia, 143Rorippa palustris, 77, 92Rosa, 245Rosa canina, 240Rosa villosa, 252Rosaceae, 7, 229, 240Rosmarinus, 157, 159Rosmarinus officinalis, 162Rubus, 241, 245, 248Rubus fruticosus, 248Rumex, 217, 220-222Rumex acetosa, 217, 220Rumex acetosella, 217, 227Rumex aquaticus, 221-222Rumex conglomeratus, 221-222Rumex crispus, 227Rumex obtusifolius, 220, 227Rumex palustris, 221-222Rumex sanguineus, 221Saccharum, 187Sagina nodosa, 100

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Salicornia, 19-21Salicornia europaea, 20, 22, 25Salsola, 19-21Salsola kali, 26Salvia, 157, 159Salvia pratensis, 159Salvia verticillata, 163Sambucus nigra, 11Sanguisorba, 245Sanguisorba minor, 253Sanicula, 31Saponaria officinalis, 100Satureja, 157Satureja hortensis, 159, 163Scandix pecten-veneris, 35-36Schismus, 198Schismus arabicus, 198Schoenoplectus, 109, 112Schoenoplectus lacustris, 113Schoenoplectus tabernaemontani,

119Schoenus, 112Schoenus nigricans, 119Scirpoides, 109, 112Scirpoides holoschoenus, 119Scirpus, 109, 112Scirpus sylvaticus, 119Scleranthus, 97Scleranthus annuus, 97, 101Scleranthus annuus ssp. annuus,

97, 101Scolymus hispanicus, 64Scrophulariaceae, 175, 178Scutellaria columnae, 163Scuttelaria, 159Scuttelaria galericulata, 157-158Secale, 187, 201Secale cereale, 202, 205, 213Securigera, 129-130Securigera varia, 143Senna occidentalis, 134Sequoiadendron giganteum, 11Sesbania sesban, 144Setaria pumila, 213Setaria viridis, 214

Silene, 96Silene baccifera, 97Silene dioica, 101Silene latifolia, 95Silene vulgaris, 94, 96Silybum marianum, 64Sinapis arvensis, 79-80, 83-85, 92Sisymbrium altissimum, 92Solanaceae, 7, 254Solanum, 255Solanum dulcamara, 254, 256Solanum esculentum, 257Solanum nigrum, 257, 259Solanum nigrum ssp. nigrum, 259Solanum nigrum ssp. schultesii,

257Sonchus arvensis, 64Sonchus asper, 64Sonchus oleraceus, 64Sorbaria, 244-245Sorbaria sorbifolia, 253Sorbaria tomentosa, 242Sorbus, 245Sorbus aucuparia, 253Sorghum, 187, 201Sorghum bicolor, 204-205, 214Sorghum halepense, 214Spartium junceum, 126Spergula, 97Spergula arvensis, 101Spergularia, 97Spergularia media, 97, 101Spergularia salina, 97Spinacia, 19, 21Spinacia oleracea, 22, 26Spiraea, 244-245Spiraea douglasii, 244Spiraea salicifolia, 253Stachys, 159Stachys arvensis, 163Stellaria media, 101Sterculiaceae, 165Suaeda, 21Suaeda maritima, 26Subularia aquatica, 92

Symphytum officinale, 66-67, 69, 71, 75

Tagetes minuta, 64Tamarindus indica, 144Tanacetum vulgare, 47Taraxacum officinale, 44, 65Taxus baccata, 11Tectona grandis, 157Teesdalia nudicaulis, 93Tetragonolobus maritimus, 144Teucrium, 157, 159Teucrium botrys, 163Teucrium montanum, 159Thalictrum, 231Thalictrum flavum, 238Thlaspi arvense, 78, 83, 93Thymus, 157Thymus vulgaris, 163Tilia, 166, 170Tilia platyphyllos, 173Tilia x vulgaris, 168-169Tiliaceae, 165Tordylium apalum, 30Torilis nodosa, 36-37, 43Trachyspermum ammi, 31Trichophorum, 109, 112Trichophorum cespitosum ssp.

germanicum, 109-110, 120Trifolium, 133Trifolium dubium, 144Trifolium glomeratum, 134Trifolium subterraneum, 137-138Trigonella foenum-graecum, 125, 144Tripleurospermum maritimum, 52, 65Triticum, 187, 195, 201Triticum aestivum, 196-197, 201,

205, 215Triticum aestivum ssp. aestivum,

196-197, 201, 205Triticum aestivum ssp. spelta, 201,

205Triticum monococcum ssp.

monococcum, 195, 201, 205Triticum turgidum, 194-195, 197,

200-201, 205, 215

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Triticum turgidum ssp. dicoccoides, 200

Triticum turgidum ssp. dicoccon, 195, 200, 205

Triticum turgidum ssp. durum, 194, 197, 201, 205, 215

Ulex europaeus, 144Umbelliferae, 29Vaccaria hispanica, 101Verbenaceae, 157Verbesina encelioides, 65Veronica, 175-176, 178Veronica arvensis, 176Veronica montana, 177Veronica praecox, 185Veronicaceae, 175, 178Vicia, 133Vicia ervilia, 145Vicia faba, 123Vicia hirsuta, 131, 133, 137Vicia sativa, 122, 138, 145Vicia sativa ssp. amphicarpa, 138Vicia sativa ssp. sativa, 145Vicia tetrasperma, 124Vigna radiata, 145Vigna umbellata, 145Vigna unguiculata ssp. unguiculata,

145Wisteria sinensis, 123Xanthium, 48, 53Xanthium spinosum, 65Xanthium strumarium, 52, 65Zea, 187, 201Zea mays, 204-205, 215Zizania palustris, 215

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Achene, 49, 112, 178-179, 182, 230-233, 235, 241, 245-246, 249

Achenetum, 245Amphicarpy, 85, 138Angiosperms, 10Anthocarp, 244Aril, 10-11, 133Awn, 192-193, 196, 200Beak, 35, 51, 79-80, 82-85, 110,

113-114, 148-149Berry, 97, 230-231, 233-235, 255, 257Bilomentum, 82Bract, 110-111, 114-115, 124, 153, 170,

187-188Bristle, 109-110, 112-114, 116, 205Calyx, 33, 50, 68-72, 96-97, 129, 131,

157-159, 166-167, 177, 180, 244, 255-257

Calyx-tooth, 33Camara, 127, 129, 136Capitulum, 45-46, 48-50, 54, 109,

124-125, 153, 229Capsule, 71-72, 96, 103, 166-167,

178-182, 232, 255-257Carpel, 70, 79, 81-82, 96, 112-114,

116, 125, 127, 129, 136, 148-149, 154, 166, 179, 217, 222, 230-232, 244, 248-249, 255-256

Carpophore, 31, 33Caryopsis, 194Chaff, 10-11, 189, 198, 201Columella, 148-149Compound fruit, 8, 11-12, 229, 245Cone, 10-11Core, 248-249Corona, 134Corymbose inflorescence, 77-78,

125, 241Corymbose infructescence, 243Cotyledons, 131-132Craspedium, 127, 129, 136Cyme, 67, 69-70, 95, 103, 217,

255-256

Cypsela, 113Dichasial cyme, 241Dicotyledons, 45Drupe, 71-72, 245-248Drupetum, 245Elaiosome, 71, 97, 154, 159, 232-233Embryo, 131-132, 194Endocarp, 10-11, 71-72, 246-248Epicalyx, 166Epicarp, 10-11, 71, 247Exocarp, 10-12, 20, 244, 247-249False fruit, 244False septum, 70-71, 81, 128-129,

256Follicetum, 245Follicle, 230-233, 235, 244, 249Fruit wall, 19, 33, 35, 71Funicle, 79, 83-84, 125, 128, 131,

133-136Funicular residue, 131, 133Glandetum, 245Glandular hair, 148, 157Glandular scale, 157Glume, 188-194, 196, 199, 201-202,

204Gymnosperms, 10Heterocarpy, 9Hilar groove, 133-135Hilar region, 134-135Hilar rim, 133-135Hilum, 50, 70-71, 103, 131, 133-135,

159, 194-195, 197-198Involucral leaf, 46, 53, 55-56Legume, 127, 129, 131, 136Lemma, 188-189, 192-194, 196, 199,

201-204Lens, 134-135Locule, 70-71, 81, 84, 103, 154, 166,

249, 256Lodicule, 188Lomentum, 127, 129, 136Mericarp, 31-37, 70-72, 148-149, 157-

159, 166-167, 170, 181-182

Mesocarp, 10-11, 71, 247-248Micropyle, 133-135Monochasium, 67Multiple fruit, 8, 11-12, 229,

244-246, 248-249Ovary, 12, 110, 154, 230, 244,

246-249Ovule, 10, 70, 82, 126, 131, 248Palea, 188-189, 193-194, 199, 201,

204Panicle, 8, 109, 153, 187-188, 230,

241-242Pappus, 9, 50-51, 53, 55-56Pedicel, 8, 69, 77, 79, 81-82, 84, 125,

137, 166, 168, 187, 199, 217Perianth, 17, 20-22, 110, 125,

153-154, 177-178, 187-188, 217, 220-225, 246

Pericarp, 31, 33, 103-104, 127, 136, 159, 166, 194, 232

Pericarpium, 244Pistil, 12, 33, 50, 177, 188, 229-230,

232, 244, 246-249Placenta, 79, 81-82, 84, 96, 170,

179-180, 255Pleurogram, 133-134Pome, 245, 248-249Pometum, 245, 249Raceme, 77, 78, 109, 125, 165,

176-177, 187, 217, 230-231, 234, 241, 243, 255

Rachilla, 111, 190-192, 194, 199Rachis, 114, 187, 190, 192-194, 198,

200, 202Radicle, 133-134Receptacle, 11, 20, 21, 45-47, 50, 53,

56, 70, 82, 229-230, 233, 241, 244-246, 248-249

Replum, 129, 136Schizocarp, 31, 37, 70, 72, 157, 165,

178, 181Scorpioid cyme, 67, 69-70Scutellum, 194, 197-198

Index 3 Glossary of botanical terms

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Index 3 Glossary of botanical terms 273

Seedcoat, 83, 105, 131-135, 148, 154, 257

Septum, 70-71, 81, 83-84, 128-129Silicula, 77Silique, 77Simple fruit, 12, 242, 244-248Spiciform panicle, 187-188Spike, 109-112, 114, 125, 131, 153, 176,

178, 187-190, 195-198, 201, 217Spikelet, 187-193, 195-198, 200-202,

204Split seed, 131-132Stigma, 33, 50, 81, 83, 178Style, 33, 35, 50, 81, 111-112, 114, 116,

148, 165-166, 180-181, 218, 230-232, 244, 246, 249

Umbel, 29-31, 36-37, 147-148Utricle, 112, 114-116Valve, 79-85, 96, 127, 136-137, 181,

255-257Wing, 20, 51-52, 83, 97, 125-126,

129, 131, 133, 181, 222

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Documents

A Manual for the Identification of PLANT SEEDS AND FRUITS