A Manual for the Identiication of PLANT SEEDS AND FRUITS Groningen Archaeological Studies Volume 23 Editorial board Prof. dr. D.C.M. Raemaekers Prof. dr. P.A.J. Atema Dr. C. Çakirlar Prof. dr. R.T.J. Cappers Prof. dr. P.D. Jordan Prof. dr. S. Voutsaki Groningen Institute of Archaeology Poststraat 6 NL-9712 ER Groningen the Netherlands gia@rug.nl Website www.gas.ub.rug.nl Publishers’ address Barkhuis Zuurstukken 37 9761 KP Eelde the Netherlands Tel. 0031 50 3080936 Fax 0031 87 7844285 info@barkhuis.nl www.barkhuis.nl A Manual for the Identiication of PLANT SEEDS AND FRUITS R.T.J. Cappers & R.M. Bekker Barkhuis & University of Groningen Library Groningen 2013 Book cover design: Nynke Tiekstra, ColtsfootMedia - Noordwolde Book interior design and typeseting: Nynke Tiekstra, ColtsfootMedia - Noordwolde Photo's: F. Botema, R.T.J. Cappers, D. Fennema & J. Jans Copy editor: Suzanne Needs Proofreader: Heather Sangster Cover: Infructesence of Black medick (Medicago lupulina; let) 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 transmited 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. Contents 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 5 Inhoud 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 Index 3 Botanische termen 274 Seeds and fruits of weed species found in a sample of flax from Argentina. 6 267 Introduction 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. 0 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]). d 200 400 600 800 1000 1200 1400 1600 1800 Asteraceae Fabaceae Poaceae Lamiaceae Malvaceae Cyperaceae Apiaceae Brassicaceae Rosaceae Caryophyllaceae Solanaceae Boraginaceae Ranunculaceae Amaranthaceae Plantaginaceae Convolvulaceae Polygonaceae Geraniaceae Juncaceae 0 5000 10000 15000 20000 25000 Organization of the manual 7 Introduction a Figure 2 Inflorescence (left; 21 June 2009) and infructescence (right; 11 July 2009) of Buckwheat (Fagopyrum esculentum). The flowers and fruits are united in panicles. 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 8 A Manual for the Identiication of Plant Seeds and Fruits | R.T.J. Cappers & R.M. Bekker a 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. 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 Family, genus, and species 9 Introduction 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. Introduction 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). 10 A Manual for the Identiication of Plant Seeds and Fruits | R.T.J. Cappers & R.M. Bekker Introduction Figure 4 Seed of the Yew (Taxus baccata) surrounded by an aril (longitudinal section). 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 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 5 Cone and seed of the Giant sequoia (Sequoiadendron giganteum). 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 6 Longitudinal section of the Melon (Cucumis melo), a fruit with seeds. Figure 7 Complete fruit of the Sunflower (Helianthus annuus) and a longitudinal section of the fruit containing the seed. Figure 11 Infructescence of the Elderberry (Sambucus nigra). 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). Types of seeds and fruits 11 Introduction 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]. 12 A Manual for the Identiication of Plant Seeds and Fruits | R.T.J. Cappers & R.M. Bekker 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): Chemischecologische 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. References 13 Introduction Further readings on the typology of seeds and fruits: PLANT FAMILY Rosaceae Dog-rose (Rosa canina) 240 Rosaceae Rosaceae Introduction The Rose family (Rosaceae) comprises approximately 85 genera and 3000 species. It encompasses a richness of forms as a result of, among other things, the great variation in inflorescence and infructescence and the different forms of fruits. Some seeds develop without being fertilized, resulting in species that can be distinguished on only minimal differences, as in the genus Lady’s-mantles (Alchemilla). Recently the genus Parsley-pierts (Aphanes) has been combined with that of Lady’s-mantles. This merger is supported by the similarities in fruit type (figure 1). It can be difficult to assign a scientific name to taxa in various other genera, because they are capable of hybridizing with both wild relatives and feral domesticates – a phenomenon known in both Cherries (Prunus) and Brambles (Rubus). Inflorescence and infructescence The flowers and fruits are either located exclusively in the axils of the bracts or united into dense racemes, dichasial cymes, panicles, or corymbose inflorescences, terminal or otherwise (figures 2-5). These different inflorescences and infructescences can occur within the same genus, as in Prunus, which displays flowers and fruits that are simple as well as those that are united into panicles or corymbose inflorescences. Figure 1 An achene of Starry lady’s-mantle (Alchemilla acutiloba) (left) and Parsley-piert (Alchemilla arvensis, previously Aphanes arvensis) (right). There are few differences in the shape and size of the fruits. The base of the fruit does vary, depending on its position on the receptacle. d Introduction 241 Figure 2 Simple fruit of Rosaceae the Medlar (Mespilus germanica). d Figure 3 Terminal flowering panicles of Himalayan sorbaria (Sorbaria tomentosa). d 242 A Manual for the Identiication of Plant Seeds and Fruits | R.T.J. Cappers & R.M. Bekker Figure 4 Terminal, corymbose infructescence Rosaceae of Hybrid chokeberry (Aronia x prunifolia). d Figure 5 The terminal inflorescence (left) and infructescence (right) of the Juneberry (Amelanchier lamarckii) is a raceme. b Inlorescence and infructescence 243 Fruit and seed Rosaceae The variation in fruit forms in the Rosaceae is considerable. Because the fruits are part multiples and can also contain vegetative plant parts, classification is problematic and far from unanimous. The classification in this overview is restricted to genera that occur in the Netherlands, based on species that occur in the wild. It is based on the approach advocated by Spjut (1994). In this family, it is important to distinguish between, on the one hand, simple fruits and multiple fruits and, on the other hand, fruits without parts that developed from the flower and fruits united with such vegetative parts. The genera discussed below represent each of the four possible combinations (table 1). A simple fruit develops from a single pistil. In this situation, it is possible that, if the pistil consists of multiple carpels, the ovaries form individual units during flowering. The carpels do, however, form a communal style (Spjut, 1994). If a flower contains multiple pistils, it develops a multiple fruit. Fruits can fuse with certain parts of the flower, for example, the receptacle. A fruit with its seed or seeds, but without external parts originating from the flower, is termed a pericarpium by Spjut (1994). If such external vegetative parts are fused with the fruit, the fruit and external parts together are termed anthocarp. The vegetative parts are termed exocarp. The term false fruit is, in fact, synonymous with anthocarp, but its use could give the incorrect impression that no fruit is present. A multiple fruit consisting of several follicles occurs in Aruncus, Sorbaria, and Spiraea. The seeds that are formed in these types of fruits are spoolshaped and very small (figure 6). Figure 6 The Steeplebush (Spiraea douglasii) has a multiple fruit consisting of five follicles. Left: The fruit with the receptacle, which is surrounded by folded back calyx-lobes. Right: A seed. d 244 A Manual for the Identiication of Plant Seeds and Fruits | R.T.J. Cappers & R.M. Bekker g g g g g g g g g ○ ○ ○ ○ ○ ○ ○ ○ ○ ○ ○ ○ ○ ○ ○ ○ ○ ○ ○ ○ ○ ○ ○ A second type of fruit is formed by a number of achenes that are united on a non-enlarged communal receptacle, as in Wood avens (Geum urbanum) (figure 7). Because this is in fact a multiple fruit, Spjut refers to it as an achenetum (to distinguish it from an achene, which pertains to a single achene and that can thus be part of a multiple fruit, as in the Rosaceae, or of a compound fruit, as in the Asteraceae). These achenes are individually dispersed. Most species of Cinquefoils (Potentilla) have multiple fruit that, just like those of the Meadowsweets (Filipendula) and Avens (Geum), consists of a large number of achenes. A deviation from this pattern is the Yellow-flowered strawberry (Potentilla indica), whose receptacle is greatly enlarged. This specific type of fruit is termed glandetum by Spjut (1994). This type of fruit, Fruit and seed 245 Rosaceae Pometum g Pome g g Drupetum Glandetum g g Drupe Achenetum Simple fruit Multiple fruit Fruit without external parts Fruit with external parts Aruncus Sorbaria Spiraea Filipendula Geum Potentilla Comarum Fragaria Prunus Rubus Amelanchier Aronia Crataegus Malus Mespilus Pyrus Sorbus Agrimonia Alchemilla Cotoneaster Rosa Sanguisorba Follicetum Table 1 Classification of genera based on the number of fruits that forms in a flower and the presence or absence of external vegetative parts. d Figure 7 Flower (left) Rosaceae and multiple fruit (right) of Wood avens (Geum urbanum). In the flower, both the many pistils and the stamens grouped around the pistils are visible. The ovary of each pistil develops into an achene. Each achene contains a single seed. The end of the long style is hooked, allowing for animal dispersal. with a swollen receptacle, also occurs in Marsh cinquefoil (Comarum palustre) and Wild strawberry (Fragaria vesca) (figures 8 and 9). Because the difference is a subtle one, it has been suggested by some researchers (Weeda et al., 1987) to move these genera into the genus Cinquefoils (Potentilla). In the genus Prunus, the flower possesses a single pistil, forming a simple fruit without any parts of the perianth (figure 10). Characteristic of these fruits is the hard inner layer, known as the stone, or endocarp (figure 11). The entire fruit is called a drupe. The seed sits inside the endocarp. The inner, hardened fruit layer (endocarp) is surrounded by an intermediate layer known Figure 8 Flower (left) and multiple fruit (right) of Wild strawberry (Fragaria vesca). The dark red achenes are visible at the surface of the swollen receptacle. d d Figure 9 Individual achene of Wild strawberry (Fragaria vesca). d 246 A Manual for the Identiication of Plant Seeds and Fruits | R.T.J. Cappers & R.M. Bekker d Figure 10 Flowers (left) and simple fruit (right) of Rosaceae the Cherry plum (Prunus ceracifera). Each flower possesses approximately 20 stamens and one pistil. The ovary of the pistil develops into a drupe. d Figure 11 Endocarps of the Cherry plum (Prunus ceracifera). Each endocarp contains a seed. d as the mesocarp and an outer layer known as the epicarp. The outer layer is also known as an exocarp, but, following Spjut (1994), we reserve this name for vegetative plant parts that have fused with the fruit. The mesocarp is often well developed and edible, as in the Wild cherry (Prunus avium), Dwarf cherry (P. cerasus), Peach (P. persica), Wild plum (P. domestica), and Cherry plum (P. ceracifera). The seeds of the Peach are edible, but it takes some effort to extricate them from the hard stone; they are therefore sold as a delicacy. Of the fruits of the St Lucie cherry (P. mahaleb) and Almond (P. dulcis), only the seed is edible. Fruit and seed 247 Figure 12 Flower (left) Rosaceae and multiple fruit (right) of the Bramble (Rubus fruticosus). The multiple fruit consists of tens of small drupes. d a Figure 13 Endocarp of In the Brambles (Rubus), the ovary of the pistil develops into a drupe. Because the flower of a Bramble often possesses a large number of pistils, it is considered a multiple fruit with many individual drupes on a communal receptacle (figure 12). The mesocarps are juicy, and the endocarps, with their characteristic netting pattern, are hard and each contain a seed (figure 13). the Bramble (Rubus fruticosus). The stone contains a seed. Figure 14 A flower in bloom and in bud (left) and two simple fruits with In several genera within the Rosaceae, the fruit fuses with parts of the flower. Following Spjut (1994), a distinction is made between simple and multiple fruits. An example of a simple fruit that is fused with part of the flower is the Apple (Malus sylvestris). This type of fruit is called a pome (figure 14). The flower of the Apple possesses five styles, each consisting of a single carpel. Above the ovary the styles are fused. Each carpel contains two ovules that can grow into a seed. The ovary grows into a leathery fruit that we call the core. The receptacle becomes greatly swollen, growing around the fruit, and is edible (figure 15). The seeds are easily removed from all of the genera listed in table 1 that have such a fruit. exocarp (right) of the Apple (Malus sylvestris). b 248 A Manual for the Identiication of Plant Seeds and Fruits | R.T.J. Cappers & R.M. Bekker d Figure 15 Fruit with seeds and exocarp (left; Rosaceae horizontal cross-section) and seed (right) of the Apple (Malus sylvestris). The fruits are formed by the core, which is composed of five carpels. Two seeds can develop in each of the five locules. The core is surrounded by edible tissue (exocarp) that has developed out of the receptacle. In some genera in the Rosaceae, the flower possesses more than one pistil, each with its own style and ovary. These pistils are surrounded by a swollen receptacle. Spjut (1994) uses the term pometum for this type of multiple fruit surrounded by a receptacle. An example of this type of fruit is the Dutch rose (Rosa ‘Hollandica’). The individual fruits are achenes (figure 16). Diaspore In the Rosaceae either seeds or fruits are dispersed, depending on whether the fruit is a follicle or one of the other types discussed above. Seeds are freed from the follicles when these burst open. In the other types the fruit or fruit with fused external vegetative parts is the dispersal unit. This includes the pome, in which the seeds are clearly recognizable within the fruit. Heterocarpy and seed dimorphism Variation in fruit and seed form does not occur in this family. Figure 16 Multiple fruit of the Dutch rose (Rosa ‘Hollandica’). Left: Intact fruit with its five sepals. Right: The fruit cut open, showing the red, enlarged receptacle surrounding tens of white achenes. d Diaspore 249 Rosaceae Seed atlas Rosaceae 1 mm Agrimonia eupatoria 1 mm Amelanchier lamarckii 250 μm Aruncus dioicus 250 μm Alchemilla glabra 250 μm Aronia x prunifolia 250 μm Comarum palustre 250 A Manual for the Identiication of Plant Seeds and Fruits | R.T.J. Cappers & R.M. Bekker Rosaceae 250 μm Cotoneaster integerrimus 250 μm Filipendula ulmaria 1 mm Geum urbanum 1 mm Crataegus monogyna 250 μm Fragaria vesca 1 mm Malus sylvestris Seed atlas Rosaceae 251 Rosaceae 1 mm Mespilus germanica 1 mm Prunus domestica ssp. domestica 250 μm Rosa villosa 250 μm Potentilla tabernaemontani 1 mm Pyrus communis 250 μm Rubus idaeus 252 A Manual for the Identiication of Plant Seeds and Fruits | R.T.J. Cappers & R.M. Bekker Rosaceae 250 μm Sanguisorba minor 250 μm Sorbus aucuparia 250 μm Sorbaria sorbifolia 250 μm Spiraea salicifolia Seed atlas Rosaceae 253 INDICES Index 1 Index 1 Common names Ajowan, 31 Almond, 247 Alpine cat’s-tail, 194 Amaranth, 17 Amphibious bistort, 218 Anemone, 231-232 Annual knawel, 97 Annual marguerite, 52, 53 Apple, 248-249 Apple-of-Peru, 255, 257 Arabian grass, 198 Asafoetida, 34-35 Avens, 245 Bamboo, 187 Baneberry, 230, 234 Barley, 187, 190, 192, 194-195, 198-199, 201, 205 2-row Barley, 201, 205 6-row Barley, 201, 205 Beak-sedges, 114 Beet, 22 Berry catchfly, 97 Bindweed family, 103 Bittersweet, 254, 256 Black mustard, 79-80 Black nightshade, 257 Black-bindweed, 220, 222-223 Bladder campion, 94, 96 Bladder ketmia, 166, 170 Bladder-sedge, 116 Bladder-senna, 127-128 Bladder-sennas, 127 Bloody crane’s-bill, 149 Bluestem, 194 Borage, 67, 71 Borage family, 66, 157 Bramble, 248 Brambles, 241, 248 Branched plantain, 176 Bread wheat, 196-197, 201, 205 Bristle club-rush, 109-110 Bristle oat, 205 Broad bean, 123 Broadleaf sermountain, 33 Broad-leaved dock, 220 Broad-leaved everlasting-pea, 126 Broomcorn, 201, 204-205 Broomrape family, 175 Brown beak-sedge, 113 Brown sedge, 111-112 Buckwheat, 8, 217, 220-221 Bugles, 159 Bugseed, 20 Bulbous barley, 199 Burdocks, 53 Buttercup family, 229 Buttercups, 229, 232 Cabbage family, 77 Cabbages, 77 Canary-grasses, 194 Cape-marigold, 52 Carrot family, 29, 35, 166 Celery-leaved buttercup, 232-233 Charlock, 79-80, 83-85 Cherries, 241 Cherry plum, 247 Chinese mallow, 166 Chinese wisteria, 123 Cinquefoils, 245-246 Clammy goosefoot, 20-21 Claries, 159 Cloud grass, 188 Clove pink, 95 Clovers, 133 Clustered clover, 134 Clustered dock, 221-222 Coast button grass, 198 Cockleburs, 48, 53 Coffee senna, 134 Common bistort, 219 Common club-rush, 113 Common comfrey, 66-67, 69, 71 Common cottongrass, 113 Common couch, 188-192 Common dandelion, 44 Common fleabane, 229 262 Handboek voor het determineren van zaden en vruchten | R.T.J. Cappers & R.M. Bekker Common glasswort, 20, 22 Common hemp-nettle, 158 Common millet, 203, 205 Common mouse-ear, 96 Common orache, 22 Common reed, 194-195 Common sorrel, 217, 220 Common spike-rush, 114 Common stork’s-bill, 149 Common vetch, 122 Common whitlowgrass, 77, 81 Common yellow-sedge, 111-112 Copse-bindweed, 222-223 Coral-necklace, 97 Coriander, 9, 32-33 Corn buttercup, 232-233 Corn toadflax, 181 Cornflower, 50 Cow parsley, 33 Cow-itch, 133 Crown vetch, 129-130 Cultivated carrot, 34-35 Cut-grass, 188-189 Daisy family, 29, 45 Darnel, 198-199 Dead-nettle family, 157 Dead-nettles, 157, 159 Deergrass, 109-110 Docks, 217, 220-222 Dodders, 103 Dog-rose, 240 Dutch rose, 249 Dwarf cherry, 247 Dwarf mallow, 165-166 Early forget-me-not, 68 Egyptian plum, 72 Einkorn, 11, 195, 197, 201, 205 Elderberry, 11 Emmer, 195, 197, 200, 205 False fox-sedge, 114 False indigo, 127 Fat-hen, 20, 22 Fennel, 33 Hellebores, 233 Henbane, 255, 257 Herb-Robert, 147 Himalayan sorbaria, 242 Hollyhock, 164-165, 170 Holy-grass, 188, 192 Honesty, 76, 81-84 Honeywort, 67, 71 Hybrid chokeberry, 243 Hyssop, 159 Indehiscent amaranth, 17 Ivy family, 29 Juneberry, 243 Kelch-grass, 198 Kidney vetch, 124 Knawels, 97 Knotgrass, 216, 218, 220 Knotgrasses, 217, 221 Knotted hedge-parsley, 36-37 Knotweed family, 217 Knotweeds, 217-218, 221-222 Laburnum, 141 Large-flowered hemp-nettle, 156 Larkspurs, 230, 233 Least mallow, 166 Lentil, 132 Lesser canary-grass, 197 Lesser celandine, 232 Lesser hawkbit, 53, 55-56 Lesser sea-spurry, 97 Lime, 168-170 Limes, 166 Long-bracted sedge, 113 Love-in-a-mist, 232-233 Love-lies-bleeding, 16 Lucerne, 103 Lungwort, 71 Lupins, 133 Lyme-grass, 190 Maize, 187, 201, 204-205 Malling toadflax, 181 Mallow family, 165 Mallows, 165 Many-seeded goosefoot, 21 Marsh cinquefoil, 246 Marsh dock, 221-222 Marsh pennywort, 29 Marsh yellow-cress, 77 Marsh-mallow, 165, 170 Marsh-marigold, 232-233 Marshworts, 29 Mat-grass, 188 Meadow clary, 159 Meadow crane’s-bill, 148 Meadow fescue, 195 Meadow-grasses, 187 Meadowsweets, 245 Medicks, 129 Medlar, 242 Melilots, 129, 133 Melon, 11 Milk-vetches, 129 Millets, 187, 201, 205 Morning glories, 105 Mouse-ears, 96 Mousetail, 230, 233 Mudwort, 181 Musk thistle, 45-46 Musk-mallow, 166-167 Narrow-leaf water-dropwort, 35-36 Nightshade family, 255 Nightshades, 255 Nile acacia, 124 Nipplewort, 48-49 Oat, 198, 201, 205 Oats, 187, 192, 198, 201, 205 Olive, 10-11 Oraches, 17, 19-20, 22 Oxeye daisy, 48 Pale persicaria, 219 Parsley-piert, 241 Parsley-pierts, 241 Parsnip, 35 Pasqueflower, 232 Pea, 136 Pea family, 123 Peach, 247 Peanut, 137 Peas, 133 Perennial glasswort, 19 Perennial rye-grass, 189 Phacelia, 67, 71, 75 Index 1 Common names 263 Index 1 Fenugreek, 125 Field bindweed, 103-104 Field penny-cress, 78, 83 Field wood-rush, 154 Fig, 11 Fine-leaved water-dropwort, 34-35 Flamboyant, 124 Flat-sedge, 113 Floating club-rush, 109 Fluellens, 176, 181 Forget-me-nots, 67, 71 Foxglove, 180-181 Foxtail barley, 195 Foxtails, 194 French bean, 131, 134-135 Garden rocket, 81 Germanders, 159 Giant hogweed, 28 Giant sequoia, 11 Giant stock-bean, 131, 133 Glaucous sedge, 114, 218 Globe-thistles, 48 Goldenseal, 233 Goosefoot family, 17 Goosefoots, 19-20 Grass family, 109, 187 Grass pea, 138 Grass-leaved orache, 22 Great fen-sedge, 114-116 Greater honeywort, 71 Greater knapweed, 45, 47 Greater plantain, 174, 176, 181 Greater sea-spurry, 97 Greater spearwort, 228-229 Grey hair-grass, 192-193 Guernsey pigweed, 19 Hairy buttercup, 232-233 Hairy finger-grass, 197 Hairy sedge, 115 Hairy tare, 131, 133, 137 Hard wheat, 194, 197, 201 Hawkweed oxtongue, 47 Heath rush, 154 Hedge bindweed, 102 Hedge hyssop, 181 Heliotrope, 69 Index 1 Pink family, 95 Plantains, 175-176, 179, 181 Pod mahogany, 133 Prickly lettuce, 51 Purple amaranth, 17 Quaking-grass, 195 Red dead-nettle, 159 Redshank, 220, 222, 224 Restharrows, 133 Rhubarb, 217 Rhubarbs, 221-222 Ribwort plantain, 178-179, 181 Rice, 187, 189 Rose family, 229, 241 Rose of Jericho, 85 Roselle, 167 Rosemary, 159 Rough cocklebur, 52 Round-fruited rush, 154 Ruptureworts, 97 Rush family, 109, 153 Rushes, 153, 187 Rye, 187, 201-202, 205 Rye-grasses, 188 Sainfoin, 131 Saltmarsh flat-sedge, 113 Saltmarsh goosefoot, 21-22 Saltworts, 19-20 Sand sedge, 115 Sandworts, 96 Sanicle, 31 Savory, 159 Scottish dock, 221-222 Sea club-rush, 109, 113 Sea mayweed, 52 Sea plantain, 176 Sea rocket, 79-80, 85 Sea rush, 154 Sea-hollies, 31, 33 Sea-kale, 78-80 Sea-spurreys, 97 Sedge family, 109 Sedges, 110-112, 114, 116, 187, 218, 220 Shoreweed, 175, 179 Short-tepalled pigweed, 19 Sickle medick, 134 Silverweed, 137 Skullcap, 157-158 Skullcaps, 159 Slender rush, 153 Slender tufted-sedge, 114 Small mallow, 166-167 Small melilot, 134 Small toadflax, 176 Small-flowered crane’s-bill, 148 Smooth tare, 124 Snapdragon, 181 Soft-rush, 152-153 Soybean, 133 Spanish broom, 126 Spanish-needles, 50 Spear-leaved orache, 22 Speedwells, 175-176 Spelt, 201, 205 Spignel, 35 Spike-rushes, 114 Spinach, 19, 22 Spotted dead-nettle, 157-158 Spurreys, 97 St Lucie cherry, 247 Starry lady’s-mantle, 241 Steeple-bush, 244 Stocks, 83 Strapwort, 97 Strawberry goosefoot, 17-18 Subterranean clover, 137, 138 Sugar cane, 187 Summer pheasant’s-eye, 233 Summer-cypress, 19 Sunflower, 11, 49-50 Sweet cicely, 35-36 Sweet vernal-grass, 188 Swine-cress, 77, 79 Tall melilot, 125 Tansy, 47 Teak, 157 Thistles, 45, 51, 53 Thorn-apple, 256 Thorn-apples, 255 Three-nerved sandwort, 97 Tidy-tips, 52 264 Handboek voor het determineren van zaden en vruchten | R.T.J. Cappers & R.M. Bekker Toadflaxes, 181 Tobaccos, 255 Tomato, 257 Tomatoes, 255 Toothed medick, 129 Touch-me-not, 129-130 True fox-sedge, 115 Tufted sedge, 115 Twisted acacia, 134 Velvetleaf, 165, 170 Vervain family, 157 Vetches, 123-133 Wall barley, 193 Wall lettuce, 48 Wall speedwell, 176 Walnut, 10-12 Waterleaf family, 67 Water-pepper, 219, 221 Water-starworts, 181 Wheat, 187, 195, 197, 201, 205 White beak-sedge, 110, 113-114 White campion, 95 White henbane, 256 White lupin, 131 White sedge, 113 Wild basil, 159 Wild carrot, 29, 34 Wild cherry, 247 Wild emmer, 200 Wild plum, 247 Wild radish, 82-85 Wild strawberry, 246 Wild-oat, 193, 198, 200 Willow dodder, 104 Winter wild-oat, 198, 200 Wood avens, 245-246 Wood dock, 221 Wood speedwell, 177 Wood-rushes, 153 Woundworts, 159 Yellow chamomile, 45, 50 Yellow-flowered strawberry, 11, 245 Yew, 11 Yorkshire-fog, 186 Abutilon, 165, 170 Abutilon theophrasti, 170-171 Acacia nilotica, 124 Acacia tortilis, 134 Acanthospermum hispidum, 57 Aconitum, 231, 233 Aconitum vulparia, 231, 236 Actaea, 230-231, 233-234, 236 Actaea pachypoda, 234 Actaea spicata, 236 Adonis, 231-232 Adonis aestivalis, 233 Adonis flammea, 236 Aethionema, 85 Aethusa cynapium, 38 Ageratina altissima, 57 Ageratum conyzoides, 57 Ageratum houstonianum, 57 Agrimonia, 245 Agrimonia eupatoria, 250 Agrostemma githago, 98 Agrostis nebulosa, 188 Ajuga, 157, 159 Ajuga reptans, 160 Alcea, 164-166, 170 Alcea rosea, 170-171 Alchemilla, 241, 245 Alchemilla glabra, 250 Alhagi graecorum, 139 Alliaria petiolata, 86 Alopecurus, 194 Alopecurus myosuroides, 206 Alopecurus pratensis, 206 Althaea, 165-166, 170 Althaea hirsuta, 171 Alysicarpus monilifer, 138 Alyssum alyssoides, 86 Amaranthaceae, 7, 16, 21 Amaranthus, 17, 21 Amaranthus blitum, 19 Amaranthus caudatus, 16 Amaranthus cruentus, 17 Amaranthus graecizans, 19 Amaranthus hybridus ssp. bouchonii, 17 Amaranthus standleyanus, 23 Ambrosia artemisiifolia, 57 Ambrosia psilostachya, 58 Ambrosia trifida, 58 Amelanchier, 245 Amelanchier lamarckii, 243, 250 Ammi majus, 38 Amphicarpaea bracteata, 138 Amsinckia micrantha, 73 Anastatica hierochuntica, 85 Anchusa arvensis, 73 Anemone, 231-232 Anemone nemorosa, 236 Anethum graveolens, 38 Anoda, 165 Anoda cristata, 172 Anthemis cotula, 52-53, 58 Anthemis tinctoria, 45, 50 Anthoxanthum, 192 Anthoxanthum odoratum, 188, 206 Anthriscus caucalis, 39 Anthriscus cerefolium, 39 Anthriscus sylvestris, 33 Anthyllis vulneraria, 124, 139 Antirrhinum, 175-176, 178, 181 Antirrhinum majus, 183 Aphanes, 241 Apiaceae, 7, 28, 166 Apium, 29 Apium graveolens, 39-40 Aquilegia, 231 Aquilegia vulgaris, 236 Arabidopsis thaliana, 86 Arabis hirsuta ssp. hirsuta, 86 Arachis hypogaea, 137 Araliaceae, 29 Arctium, 53 Arctium minus, 58 Arenaria, 96 Arenaria serpyllifolia, 98 Aronia, 245 Aronia x prunifolia, 243, 250 Artedia squamata, 30 Artemisia verlotiorum, 58 Aruncus, 244-245, 250 Aruncus dioicus Asperugo procumbens, 73 Aster, 45 Asteraceae, 7, 9, 29, 44, 123, 229, 245 Astragalus, 128-129 Astragalus eremophilus, 128 Astragalus glycyphyllos, 139 Atriplex, 17, 19, 21-22 Atriplex littoralis, 22 Atriplex patula, 22 Atriplex prostrata, 22-23 Atropa, 255 Atropa bella-donna, 258 Aubrieta deltoidea, 86 Avena, 187, 192, 198, 201, 205 Avena fatua, 193, 198, 200, 207 Avena sativa, 198, 201, 205, 207 Avena sterilis, 198, 200, 207 Avena strigosa, 205 Axyris amaranthoides, 23 Baccharis halimifolia, 59 Ballota nigra ssp. meridionalis, 160 Bambusa, 187 Baptisia australis, 127 Barbarea vulgaris, 86 Bassia, 19, 21 Bassia scoparia, 24 Berteroa incana, 87 Beta, 21 Beta vulgaris, 22, 24 Bidens bipinnata, 50 Blysmus, 109, 112 Blysmus compressus, 113 Blysmus rufus, 113, 117 Bolboschoenus, 109, 112 Index 2 Scientiic names 265 Index 2 Index 2 Scientific names Index 2 Bolboschoenus maritimus, 109, 113, 117 Bombacaceae, 165 Boraginaceae, 7, 66, 157 Borago, 67 Borago officinalis, 67, 71, 73 Brassica, 77 Brassica juncea, 77 Brassica napus, 77 Brassica nigra, 79-80, 87 Brassica oleracea, 77 Brassica rapa, 77 Brassicaceae, 7, 76 Briza media, 195 Bromopsis inermis ssp. inermis, 207 Brunnera macrophylla, 73 Bunias orientalis, 87 Caesalpinioideae, 123-125, 127, 133-134 Cakile maritima, 79-80, 85, 87 Calamintha, 157 Calendula officinalis, 53-54, 59 Calepina irregularis, 87 Callistephus chinensis, 59 Callitrichaceae, 175, 178 Callitriche, 175-176, 178, 181-182 Callitriche cophocarpa, 181 Callitriche hermaphroditica, 183 Caltha, 231 Caltha palustris, 232-233, 236 Caltha palustris ssp. palustris, 236 Camelina sativa ssp. sativa, 87 Canavalia ensiformis, 131, 133 Capsella bursa-pastoris, 81-82, 88 Cardamine chenopodifolia, 85 Cardamine pratensis, 88 Carduus, 51 Carduus acanthoides, 59 Carduus crispus, 59 Carduus nutans, 45-46, 59 Carex, 112, 114, 116, 220 Carex acuta, 114 Carex arenaria, 115 Carex crawfordii, 115 Carex curta, 113 Carex disticha, 111-112 Carex elata, 115 Carex extensa, 113 Carex flacca, 114, 218 Carex hirta, 115 Carex lepidocarpa, 117 Carex oederi ssp. oedocarpa, 111-112 Carex otrubae, 114 Carex spicata, 117 Carex vesicaria, 116 Carex vulpina, 115 Carthamus tinctorius, 60 Caryophyllaceae, 7, 94 Caryophyllus, 95 Centaurea benedicta, 60 Centaurea cyanus, 50 Centaurea scabiosa, 45, 47 Centaurea solstitialis, 60 Cerastium, 96, 98 Cerastium fontanum, 96, 98 Cerastium fontanum ssp. vulgare, 96, 98 Cerinthe, 67, 71 Cerinthe major, 71 Chaenorhinum, 175-176, 178, 181 Chaenorhinum minus, 176, 183 Chaenorhinum origanifolium, 181 Chaerophyllum temulum, 40 Chenopodium, 19, 21 Chenopodium album, 20, 22 Chenopodium chenopodioides, 21-22 Chenopodium foliosum, 17-18 Chenopodium polyspermum, 21 Chenopodium pumilio, 20-21 Chenopodium rubrum, 24 Chondrilla juncea, 60 Cicer arietinum, 139 Cichorium endivia, 60 Cichorium intybus, 60 Cirsium, 53 Cirsium acaule, 61 Cirsium arvense, 61 Cladium, 112 266 Handboek voor het determineren van zaden en vruchten | R.T.J. Cappers & R.M. Bekker Cladium mariscus, 114-116, 118 Clematis, 231 Clematis vitalba, 232, 237 Clinopodium acinos, 160 Clinopodium vulgare, 159 Cochlearia danica, 88 Coincya monensis, 88 Colutea, 127 Colutea arborescens, 128, 139 Comarum, 245 Comarum palustre, 246, 250 Compositae, 45 Conium maculatum, 40-41 Conringia orientalis, 88 Consolida, 230-231, 233 Consolida regalis, 237 Convolvulaceae, 7, 102 Convolvulus, 103 Convolvulus arvensis, 104, 106 Convolvulus sepium, 102 Cordia, 71-72 Cordia myxa, 72 Coriandrum sativum, 9, 32-33, 41 Corispermum, 20-21 Corispermum intermedium, 25 Corrigiola, 97 Corrigiola litoralis, 98 Corynephorus canescens, 192-193 Cotoneaster, 245 Cotoneaster integerrimus, 251 Crambe maritima, 78-80, 88 Crataegus, 245 Crataegus monogyna, 251 Crotalaria juncea, 139 Cruciferae, 77 Crypsis aculeata, 189 Cucubalus baccifer, 97 Cucumis melo, 11 Cuminum cyminum, 41 Cuscuta, 103 Cuscuta europaea, 106 Cuscuta pedicellata, 103 Cuscutaceae, 103 Cymbalaria, 175-176, 178, 181 Cymbalaria muralis, 183 Erodium cicutarium, 149-150 Erophila verna, 77, 81, 89 Eruca vesicaria, 81, 89 Erucastrum gallicum, 89 Eryngium, 31, 33 Eryngium alpinum, 33 Erysimum cheiranthoides, 90 Faba, 123 Faba vulgaris, 123 Fabaceae, 7, 122 Fagopyrum, 221 Fagopyrum esculentum, 8, 217, 220, 226 Fallopia, 217, 221-222 Fallopia convolvulus, 220, 222-223, 226 Fallopia dumetorum, 222-223 Farsetia, 83 Farsetia ramosissima, 84 Ferula foetida, 34-35 Festuca arundinacea, 209 Festuca pratensis, 195 Ficaria, 231 Ficaria verna, 232, 237 Ficus carica, 11 Filipendula, 245 Filipendula ulmaria, 251 Foeniculum vulgare, 33, 42 Fragaria, 245 Fragaria vesca, 246, 251 Galega officinalis, 140 Galeopsis speciosa, 156, 160 Galeopsis tetrahit, 158 Galinsoga parviflora, 61 Genista pilosa, 140 Geraniaceae, 7, 146 Geranium, 147-149 Geranium dissectum, 150 Geranium molle, 146, 148 Geranium pratense, 148 Geranium pusillum, 148 Geranium robertianum, 150 Geranium robertsianum, 147 Geranium sanguineum, 149 Geum, 245 Geum urbanum, 245-246, 251 Glebionis segetum, 61 Glechoma, 157 Glechoma hederacea, 160 Glycine, 133 Glycine max, 140 Gramineae, 187 Gratiola, 175-176, 178 Gratiola officinalis, 181, 183 Gypsophila muralis, 98 Halogeton glomeratus, 22 Helianthus annuus, 11, 49-50, 62 Heliotropium, 69 Helleborus, 231, 233 Helleborus foetidus, 237 Heracleum mantegazzianum, 28 Herniaria, 97 Herniaria glabra, 99 Hesperis matronalis, 90 Hibiscus, 166, 170 Hibiscus sabdariffa, 167 Hibiscus trionum, 172 Hieracium pilosella, 62 Hieracium vulgatum, 62 Hierochloe odorata, 188 Hippocrepis, 129 Hippocrepis comosa, 130, 140 Hippuridaceae, 175, 178 Hippuris, 176, 178-179, 182 Hippuris vulgaris, 175, 177, 179, 184 Hirschfeldia incana, 90 Holcus lanatus, 186, 209 Holosteum umbellatum, 99 Honckenya peploides, 99 Hordeum, 187, 190, 192, 194, 198, 201 Hordeum bulbosum, 199 Hordeum jubatum, 195 Hordeum murinum, 193 Hordeum vulgare, 205, 209 Hordeum vulgare ssp. distichon, 205 Hordeum vulgare ssp. vulgare, 205 Hydrastis canadensis, 233 Hydrocotylaceae, 29 Index 2 Scientiic names 267 Index 2 Cynodon dactylon, 208 Cynoglossum officinale, 71, 73 Cyperaceae, 7, 108, 187, 218 Cyperus, 112 Cyperus conglomeratus, 110-111 Cyperus esculentus, 118 Cytisus scoparius, 140 Dactyloctenium, 198 Dactyloctenium aegypticum, 198 Datura, 255 Datura stramonium, 256, 258 Daucus carota ssp. carota, 29, 34 Daucus carota ssp. sativus, 34-35 Delonix regia, 124 Descurainia sophia, 89 Dianthus caryophyllus, 95 Dianthus deltoides, 98 Dichantium, 194 Digitalis, 175-176, 178 Digitalis purpurea, 180-181, 183 Digitaria sanguinalis, 197, 208 Dimorphotheca pluvialis, 52 Diplotaxis muralis, 89 Draba muralis, 89 Echinacea angustifolia, 61 Echinochloa crus-galli, 208 Echinops, 48 Echium, 67 Echium vulgare, 70, 74 Eclipta prostrata, 61 Eleocharis, 112, 114 Eleocharis multicaulis, 118 Eleocharis palustris, 114 Eleogiton, 109, 112 Eleogiton fluitans, 109-110, 118 Elytrigia repens, 188-192 Emex spinosa, 222, 224-225 Entada, 129 Eranthis, 231 Eranthis hyemalis, 237 Eriophorum, 112 Eriophorum angustifolium, 113 Eriophorum latifolium, 118 Eriophorum vaginatum, 108 Erodium, 147, 149 Index 2 Hydrocotyle, 29 Hydrophyllaceae, 67 Hyoscyamus, 255 Hyoscyamus albus, 256 Hyoscyamus niger, 257-258 Hyssopus, 157, 159 Hyssopus officinalis, 160 Iberis umbellata, 90 Illecebrum, 97 Illecebrum verticillatum, 99 Inula conyzae, 62 Ipomoea, 105 Ipomoea carnea, 105 Isatis tinctoria, 90 Isolepis, 109, 112 Isolepis setacea, 109-110, 119 Iva xanthifolia, 62 Jacobaea vulgaris, 62 Juglans regia, 11 Juncaceae, 7, 109, 152, 187 Juncus, 153-154 Juncus acutiflorus, 155 Juncus compressus, 154 Juncus effusus, 152-153 Juncus maritimus, 154 Juncus squarrosus, 154 Juncus tenuis, 153 Kickxia, 175-176, 178, 181 Kickxia elatine, 184 Labiatae, 157 Lablab purpureus, 140 Laburnum anagyroides, 141 Lactuca sativa, 63 Lactuca serriola, 51 Lagurus ovatus, 194 Lamiaceae, 7, 70, 156 Lamiastrum galeobdolon ssp. galeobdolon, 161 Lamium, 157, 159 Lamium maculatum, 157-158 Lamium purpureum, 159, 161 Lappula squarrosa, 74 Lapsana communis, 48, 63 Laserpitium latifolium, 33 Lathyrus, 133 Lathyrus ciliolatus, 138 Lathyrus latifolius, 126 Lathyrus sativus, 138, 141 Lathyrus sylvestris, 141 Layia platyglossa, 52 Leersia oryzoides, 188-189 Leguminosae, 123 Lens culinaris, 132, 141 Leontodon saxatilis, 53, 55-56 Leonurus cardiaca, 161 Lepidium coronopus, 77, 79 Lepidium didymum, 90 Lepidium sativum, 91 Leucaena leucocephala, 141 Leucanthemum paludosum, 52-53 Leucanthemum vulgare, 48, 63 Leymus arenarius, 190 Limosella, 175-176, 178 Limosella aquatica, 181, 184 Linaria, 175-176, 178, 181 Linaria arvensis, 181 Linaria vulgaris, 184 Lithospermum arvense, 74 Littorella, 175-176, 178-179, 182 Littorella uniflora, 175, 179, 184 Lobularia maritima, 91 Lolium, 188 Lolium multiflorum, 210 Lolium perenne, 189, 210 Lolium remotum, 210 Lolium temulentum, 198-199 Lotus pedunculatus, 141 Lunaria annua, 76, 81-84, 91 Lupinus, 133 Lupinus albus, 131 Lupinus angustifolius, 142 Luzula, 153-154 Luzula campestris, 154 Luzula multiflora, 155 Lycium, 255 Lycium barbarum, 258 Lycopersicon, 255 Lycopus europaeus, 161 Malcolmia maritima, 91 Malus, 245 Malus sylvestris, 248-249, 251 Malva, 165-166, 170 268 Handboek voor het determineren van zaden en vruchten | R.T.J. Cappers & R.M. Bekker Malva alcea, 172 Malva moschata, 166-167 Malva neglecta, 165-166 Malva parviflora, 166 Malva pusilla, 166-167 Malva verticillata, 166 Malvaceae, 7, 164 Marrubium vulgare, 161 Matricaria chamomilla, 63 Matthiola, 83 Matthiola tricuspidata, 83 Medicago, 129 Medicago falcata, 134 Medicago lupulina, 4 Medicago polymorpha, 129 Medicago sativa, 103, 142 Melilotus, 129, 133 Melilotus altissimus, 125, 142 Melilotus indicus, 134 Melissa, 157 Melissa officinalis, 161 Mentha arvensis, 162 Mespilus, 245 Mespilus germanica, 242, 252 Meum athamanticum, 35 Mimosa pudica, 129-130 Mimosoideae, 123-125, 127, 133-134 Minuartia hybrida, 99 Misopates, 175-176, 178, 181 Misopates orontium, 180-181, 184 Moehringia trinervia, 97, 99 Moenchia erecta, 100 Mucuna, 133 Mucuna pruriens, 133 Mycelis muralis, 48 Myosotis, 67, 71 Myosotis arvensis, 74 Myosotis ramosissima, 68 Myosoton aquaticum, 100 Myosurus, 231 Myosurus minimus, 230, 233, 237 Myrrhis odorata, 35-36 Nardus, 188 Nasturtium microphyllum, 91 Nepeta, 157 Nepeta cataria, 162 Phalaris canariensis, 212 Phalaris minor, 197, 212 Phaseolus coccineus, 143 Phaseolus vulgaris, 135, 143 Phleum, 194 Phleum alpinum, 194 Phragmites australis, 194-195 Physalis, 255 Physalis alkekengi, 259 Picris echioides, 63 Picris hieracioides, 47 Pisum fulvum, 138 Pisum sativum, 136, 143 Plantaginaceae, 7, 174 Plantago, 175-176, 178-179, 181 Plantago arenaria, 176 Plantago coronopus, 182 Plantago lanceolata, 178-179, 181, 185 Plantago major, 174, 176, 181 Plantago maritima, 176 Poa, 187 Poa pratensis, 213 Poaceae, 7, 109, 123, 186 Polycarpon tetraphyllum, 100 Polycnemum arvense, 25 Polygonaceae, 7, 216 Polygonum, 217, 221 Polygonum aviculare, 216, 218, 220, 226 Potentilla, 245-246 Potentilla anserina, 137 Potentilla indica, 11, 245 Potentilla tabernaemontani, 252 Prosopis farcta, 143 Prunella vulgaris, 162 Prunus, 241, 245-246 Prunus avium, 247 Prunus ceracifera, 247 Prunus cerasus, 247 Prunus domestica, 247, 252 Prunus domestica ssp. domestica, 252 Prunus dulcis, 247 Prunus mahaleb, 247 Prunus persica, 247 Pulicaria dysenterica, 229 Pulmonaria officinalis, 71, 75 Pulsatilla, 231 Pulsatilla vulgaris, 232, 238 Pyrus, 245 Pyrus communis, 252 Ranunculaceae, 7, 228 Ranunculus, 229, 231-232 Ranunculus arvensis, 232-233 Ranunculus bulbosus, 238 Ranunculus lingua, 228-229 Ranunculus peltatus, 233 Ranunculus sardous, 232-233 Ranunculus sceleratus, 232-233 Ranunculus subgen. Batrachium, 232 Ranunculus subgen. Ranunculus, 232 Raphanus raphanistrum, 82-85, 92 Rapistrum rugosum, 92 Rheum, 221-222 Rheum x hybridum, 217 Rhynchospora, 112, 114 Rhynchospora alba, 110, 113-114 Rhynchospora fusca, 113, 119 Robinia pseudoacacia, 143 Rorippa palustris, 77, 92 Rosa, 245 Rosa canina, 240 Rosa villosa, 252 Rosaceae, 7, 229, 240 Rosmarinus, 157, 159 Rosmarinus officinalis, 162 Rubus, 241, 245, 248 Rubus fruticosus, 248 Rumex, 217, 220-222 Rumex acetosa, 217, 220 Rumex acetosella, 217, 227 Rumex aquaticus, 221-222 Rumex conglomeratus, 221-222 Rumex crispus, 227 Rumex obtusifolius, 220, 227 Rumex palustris, 221-222 Rumex sanguineus, 221 Saccharum, 187 Sagina nodosa, 100 Index 2 Scientiic names 269 Index 2 Neslia paniculata, 91 Nicandra, 255 Nicandra physalodes, 255, 257-258 Nicotiana, 255 Nicotiana tabacum, 258 Nigella, 231-233 Nigella arvensis, 238 Nonea lutea, 74 Ocimum basilicum, 162 Oenanthe aquatica, 34-35 Oenanthe silaifolia, 35-36 Olea europaea, 11 Omphalodes verna, 74 Onobrychis, 131 Onobrychis viciifolia, 142 Ononis, 133 Ononis repens ssp. spinosa, 142 Origanum vulgare, 162 Ornithopus, 129 Ornithopus sativus, 142 Orobanchaceae, 175 Oryza, 187 Oryza sativa, 189, 211 Panicum, 187, 201 Panicum capillare, 211 Panicum miliaceum, 203, 205 Papilionaceae, 123 Papilionoidae, 123-125, 127, 133-135 Paspalum dilatatum, 211 Pastinaca sativa ssp. sativa, 35, 42 Pelargonium, 147 Pennisetum, 187 Pennisetum glaucum, 212 Pentaglottis sempervirens, 75 Persicaria, 217, 221 Persicaria amphibia, 218, 226 Persicaria bistorta, 219 Persicaria hydropiper, 219, 221 Persicaria lapathifolia, 219, 226 Persicaria maculosa, 220, 222, 224 Persicaria minor, 226 Petasites hybridus, 63 Petrorhagia prolifera, 100 Petroselinum crispum, 43 Phacelia tanacetifolia, 67, 71, 75 Phalaris, 194 Index 2 Salicornia, 19-21 Salicornia europaea, 20, 22, 25 Salsola, 19-21 Salsola kali, 26 Salvia, 157, 159 Salvia pratensis, 159 Salvia verticillata, 163 Sambucus nigra, 11 Sanguisorba, 245 Sanguisorba minor, 253 Sanicula, 31 Saponaria officinalis, 100 Satureja, 157 Satureja hortensis, 159, 163 Scandix pecten-veneris, 35-36 Schismus, 198 Schismus arabicus, 198 Schoenoplectus, 109, 112 Schoenoplectus lacustris, 113 Schoenoplectus tabernaemontani, 119 Schoenus, 112 Schoenus nigricans, 119 Scirpoides, 109, 112 Scirpoides holoschoenus, 119 Scirpus, 109, 112 Scirpus sylvaticus, 119 Scleranthus, 97 Scleranthus annuus, 97, 101 Scleranthus annuus ssp. annuus, 97, 101 Scolymus hispanicus, 64 Scrophulariaceae, 175, 178 Scutellaria columnae, 163 Scuttelaria, 159 Scuttelaria galericulata, 157-158 Secale, 187, 201 Secale cereale, 202, 205, 213 Securigera, 129-130 Securigera varia, 143 Senna occidentalis, 134 Sequoiadendron giganteum, 11 Sesbania sesban, 144 Setaria pumila, 213 Setaria viridis, 214 Silene, 96 Silene baccifera, 97 Silene dioica, 101 Silene latifolia, 95 Silene vulgaris, 94, 96 Silybum marianum, 64 Sinapis arvensis, 79-80, 83-85, 92 Sisymbrium altissimum, 92 Solanaceae, 7, 254 Solanum, 255 Solanum dulcamara, 254, 256 Solanum esculentum, 257 Solanum nigrum, 257, 259 Solanum nigrum ssp. nigrum, 259 Solanum nigrum ssp. schultesii, 257 Sonchus arvensis, 64 Sonchus asper, 64 Sonchus oleraceus, 64 Sorbaria, 244-245 Sorbaria sorbifolia, 253 Sorbaria tomentosa, 242 Sorbus, 245 Sorbus aucuparia, 253 Sorghum, 187, 201 Sorghum bicolor, 204-205, 214 Sorghum halepense, 214 Spartium junceum, 126 Spergula, 97 Spergula arvensis, 101 Spergularia, 97 Spergularia media, 97, 101 Spergularia salina, 97 Spinacia, 19, 21 Spinacia oleracea, 22, 26 Spiraea, 244-245 Spiraea douglasii, 244 Spiraea salicifolia, 253 Stachys, 159 Stachys arvensis, 163 Stellaria media, 101 Sterculiaceae, 165 Suaeda, 21 Suaeda maritima, 26 Subularia aquatica, 92 270 Handboek voor het determineren van zaden en vruchten | R.T.J. Cappers & R.M. Bekker Symphytum officinale, 66-67, 69, 71, 75 Tagetes minuta, 64 Tamarindus indica, 144 Tanacetum vulgare, 47 Taraxacum officinale, 44, 65 Taxus baccata, 11 Tectona grandis, 157 Teesdalia nudicaulis, 93 Tetragonolobus maritimus, 144 Teucrium, 157, 159 Teucrium botrys, 163 Teucrium montanum, 159 Thalictrum, 231 Thalictrum flavum, 238 Thlaspi arvense, 78, 83, 93 Thymus, 157 Thymus vulgaris, 163 Tilia, 166, 170 Tilia platyphyllos, 173 Tilia x vulgaris, 168-169 Tiliaceae, 165 Tordylium apalum, 30 Torilis nodosa, 36-37, 43 Trachyspermum ammi, 31 Trichophorum, 109, 112 Trichophorum cespitosum ssp. germanicum, 109-110, 120 Trifolium, 133 Trifolium dubium, 144 Trifolium glomeratum, 134 Trifolium subterraneum, 137-138 Trigonella foenum-graecum, 125, 144 Tripleurospermum maritimum, 52, 65 Triticum, 187, 195, 201 Triticum aestivum, 196-197, 201, 205, 215 Triticum aestivum ssp. aestivum, 196-197, 201, 205 Triticum aestivum ssp. spelta, 201, 205 Triticum monococcum ssp. monococcum, 195, 201, 205 Triticum turgidum, 194-195, 197, 200-201, 205, 215 Index 2 Triticum turgidum ssp. dicoccoides, 200 Triticum turgidum ssp. dicoccon, 195, 200, 205 Triticum turgidum ssp. durum, 194, 197, 201, 205, 215 Ulex europaeus, 144 Umbelliferae, 29 Vaccaria hispanica, 101 Verbenaceae, 157 Verbesina encelioides, 65 Veronica, 175-176, 178 Veronica arvensis, 176 Veronica montana, 177 Veronica praecox, 185 Veronicaceae, 175, 178 Vicia, 133 Vicia ervilia, 145 Vicia faba, 123 Vicia hirsuta, 131, 133, 137 Vicia sativa, 122, 138, 145 Vicia sativa ssp. amphicarpa, 138 Vicia sativa ssp. sativa, 145 Vicia tetrasperma, 124 Vigna radiata, 145 Vigna umbellata, 145 Vigna unguiculata ssp. unguiculata, 145 Wisteria sinensis, 123 Xanthium, 48, 53 Xanthium spinosum, 65 Xanthium strumarium, 52, 65 Zea, 187, 201 Zea mays, 204-205, 215 Zizania palustris, 215 Index 2 Scientiic names 271 Index 3 Index 3 Glossary of botanical terms Achene, 49, 112, 178-179, 182, 230-233, 235, 241, 245-246, 249 Achenetum, 245 Amphicarpy, 85, 138 Angiosperms, 10 Anthocarp, 244 Aril, 10-11, 133 Awn, 192-193, 196, 200 Beak, 35, 51, 79-80, 82-85, 110, 113-114, 148-149 Berry, 97, 230-231, 233-235, 255, 257 Bilomentum, 82 Bract, 110-111, 114-115, 124, 153, 170, 187-188 Bristle, 109-110, 112-114, 116, 205 Calyx, 33, 50, 68-72, 96-97, 129, 131, 157-159, 166-167, 177, 180, 244, 255-257 Calyx-tooth, 33 Camara, 127, 129, 136 Capitulum, 45-46, 48-50, 54, 109, 124-125, 153, 229 Capsule, 71-72, 96, 103, 166-167, 178-182, 232, 255-257 Carpel, 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, 33 Caryopsis, 194 Chaff, 10-11, 189, 198, 201 Columella, 148-149 Compound fruit, 8, 11-12, 229, 245 Cone, 10-11 Core, 248-249 Corona, 134 Corymbose inflorescence, 77-78, 125, 241 Corymbose infructescence, 243 Cotyledons, 131-132 Craspedium, 127, 129, 136 Cyme, 67, 69-70, 95, 103, 217, 255-256 Cypsela, 113 Dichasial cyme, 241 Dicotyledons, 45 Drupe, 71-72, 245-248 Drupetum, 245 Elaiosome, 71, 97, 154, 159, 232-233 Embryo, 131-132, 194 Endocarp, 10-11, 71-72, 246-248 Epicalyx, 166 Epicarp, 10-11, 71, 247 Exocarp, 10-12, 20, 244, 247-249 False fruit, 244 False septum, 70-71, 81, 128-129, 256 Follicetum, 245 Follicle, 230-233, 235, 244, 249 Fruit wall, 19, 33, 35, 71 Funicle, 79, 83-84, 125, 128, 131, 133-136 Funicular residue, 131, 133 Glandetum, 245 Glandular hair, 148, 157 Glandular scale, 157 Glume, 188-194, 196, 199, 201-202, 204 Gymnosperms, 10 Heterocarpy, 9 Hilar groove, 133-135 Hilar region, 134-135 Hilar rim, 133-135 Hilum, 50, 70-71, 103, 131, 133-135, 159, 194-195, 197-198 Involucral leaf, 46, 53, 55-56 Legume, 127, 129, 131, 136 Lemma, 188-189, 192-194, 196, 199, 201-204 Lens, 134-135 Locule, 70-71, 81, 84, 103, 154, 166, 249, 256 Lodicule, 188 Lomentum, 127, 129, 136 Mericarp, 31-37, 70-72, 148-149, 157159, 166-167, 170, 181-182 272 Handboek voor het determineren van zaden en vruchten | R.T.J. Cappers & R.M. Bekker Mesocarp, 10-11, 71, 247-248 Micropyle, 133-135 Monochasium, 67 Multiple fruit, 8, 11-12, 229, 244-246, 248-249 Ovary, 12, 110, 154, 230, 244, 246-249 Ovule, 10, 70, 82, 126, 131, 248 Palea, 188-189, 193-194, 199, 201, 204 Panicle, 8, 109, 153, 187-188, 230, 241-242 Pappus, 9, 50-51, 53, 55-56 Pedicel, 8, 69, 77, 79, 81-82, 84, 125, 137, 166, 168, 187, 199, 217 Perianth, 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, 244 Pistil, 12, 33, 50, 177, 188, 229-230, 232, 244, 246-249 Placenta, 79, 81-82, 84, 96, 170, 179-180, 255 Pleurogram, 133-134 Pome, 245, 248-249 Pometum, 245, 249 Raceme, 77, 78, 109, 125, 165, 176-177, 187, 217, 230-231, 234, 241, 243, 255 Rachilla, 111, 190-192, 194, 199 Rachis, 114, 187, 190, 192-194, 198, 200, 202 Radicle, 133-134 Receptacle, 11, 20, 21, 45-47, 50, 53, 56, 70, 82, 229-230, 233, 241, 244-246, 248-249 Replum, 129, 136 Schizocarp, 31, 37, 70, 72, 157, 165, 178, 181 Scorpioid cyme, 67, 69-70 Scutellum, 194, 197-198 Index 3 Seedcoat, 83, 105, 131-135, 148, 154, 257 Septum, 70-71, 81, 83-84, 128-129 Silicula, 77 Silique, 77 Simple fruit, 12, 242, 244-248 Spiciform panicle, 187-188 Spike, 109-112, 114, 125, 131, 153, 176, 178, 187-190, 195-198, 201, 217 Spikelet, 187-193, 195-198, 200-202, 204 Split seed, 131-132 Stigma, 33, 50, 81, 83, 178 Style, 33, 35, 50, 81, 111-112, 114, 116, 148, 165-166, 180-181, 218, 230-232, 244, 246, 249 Umbel, 29-31, 36-37, 147-148 Utricle, 112, 114-116 Valve, 79-85, 96, 127, 136-137, 181, 255-257 Wing, 20, 51-52, 83, 97, 125-126, 129, 131, 133, 181, 222 Index 3 Glossary of botanical terms 273