formed by the diploid cells of the par<strong>en</strong>t sporophyte. G<strong>en</strong>era-tion two is the nutritive tissue, which is provi<strong>de</strong><strong>de</strong>ither by the haploid body cells of the female gametophyte (in gymnosperms) or by the triploid <strong>en</strong>dosperm (inangiosperms). The third g<strong>en</strong>eration is the diploid embryo, which combin<strong>es</strong> the g<strong>en</strong>etic material of two differ<strong>en</strong>tindividuals, the mother sporophyte (providing the egg cells) and father sporophyte (providing the poll<strong>en</strong>). Thissophisticated combination of three g<strong>en</strong>etically differ<strong>en</strong>t g<strong>en</strong>erations of tissue in a single organ r<strong>en</strong><strong>de</strong>rs the seedthe most complex structure produced by a seed p<strong>la</strong>nt.... or sometim<strong>es</strong> just oneAlthough it is true that the vast majority of angiosperms produce their seeds sexually, there are a few notableexceptions. Some p<strong>la</strong>nts stopped exploiting the advantag<strong>es</strong> of double fertilization and abstained – either partiallyor totally – from sexual reproduction. There are more than 400 speci<strong>es</strong> in over forty angiosperm famili<strong>es</strong> whichare able produce seeds asexually, by a proc<strong>es</strong>s called apomixis. Curiously, although well repr<strong>es</strong><strong>en</strong>ted amongMonocots and Dicots, apomixis appears to be <strong>en</strong>tirely abs<strong>en</strong>t among the gymnosperms.Apomictic speci<strong>es</strong> are thought to have evolved in<strong>de</strong>p<strong>en</strong>d<strong>en</strong>tly from sexually repro-ducing anc<strong>es</strong>torsmultiple tim<strong>es</strong>. Some of them are only facultative apomicts and can still reproduce sexually, while others areobligate apomicts whose only way of reproduction is apomixis. Although the mechanisms leading to apomicticreproduction are diverse, the un<strong>de</strong>rlying principle is that meiotic division is by-passed, so that a diploid egg cellis produced and <strong>de</strong>velops into an embryo without prior fertilization (parth<strong>en</strong>og<strong>en</strong><strong>es</strong>is). For the <strong>de</strong>velopm<strong>en</strong>t of the<strong>en</strong>dosperm most apomicts still require fertilization of the c<strong>en</strong>tral cell with its two po<strong>la</strong>r nuclei. But someapomictic speci<strong>es</strong> have abandoned the fertilization of the c<strong>en</strong>tral cell, which th<strong>en</strong> initiat<strong>es</strong> the <strong>de</strong>velopm<strong>en</strong>t ofthe <strong>en</strong>dosperm on its own.Sev<strong>en</strong>ty-five per c<strong>en</strong>t of all known apomicts occur in just three famili<strong>es</strong>: the grass<strong>es</strong> (Poaceae), the rosefamily (Rosaceae) and the sunflower family (Asteraceae), which inclu<strong>de</strong> such familiar exampl<strong>es</strong> as dan<strong>de</strong>lion(Taraxacum officinalis), mouse-ear hawkweed (Hieracium pilosel<strong>la</strong>) and cinquefoil (Pot<strong>en</strong>til<strong>la</strong> spp.). As a r<strong>es</strong>ult of thevarious apomictic mechanisms the seeds contain embryos that are g<strong>en</strong>etically id<strong>en</strong>tical to the mother p<strong>la</strong>nt. Suchclonal embryos are useful in the propagation of apomictic crops such as Citrus speci<strong>es</strong> (Rutaceae), mangoste<strong>en</strong>(Garcinia mangostana, Clusiaceae) and b<strong>la</strong>ckberri<strong>es</strong> (Rubus fruticosus, Rosaceae). Their apomic-tically producedseeds yield an exact copy of the mother p<strong>la</strong>nt, thus perpetuating the valuable traits of a particu<strong>la</strong>r race or hybridthrough succ<strong>es</strong>sive seed g<strong>en</strong>erations.Homage to the gymnosperms – at least they look good on paperBeautiful flowers, clever pollination strategi<strong>es</strong>, double fertilization, and more effici<strong>en</strong>t seed production – theangiosperm way of life was, and still is, a great succ<strong>es</strong>s story. The fossil record shows that angiosperms appearedquickly and sudd<strong>en</strong>ly betwe<strong>en</strong> the <strong>en</strong>d of the Jurassic and the beginning of the Cretaceous (c.140 million yearsago). By the <strong>en</strong>d of the Cretaceous (c. 65 million years ago), they had literally explo<strong>de</strong>d into a huge diversity ofspeci<strong>es</strong>, taking over most terr<strong>es</strong>trial p<strong>la</strong>nt communiti<strong>es</strong>. Soon, they had <strong>la</strong>rgely disp<strong>la</strong>ced ferns and gymnosperms,the dominant vegetation during the Permian, Triassic and Jurassic.Neverthel<strong>es</strong>s, ferns and gymnosperms still grow on Earth today. In fact, rec<strong>en</strong>t r<strong>es</strong>earch sugg<strong>es</strong>ts that themajority of living fern speci<strong>es</strong> (80 per c<strong>en</strong>t) diversified only in the Cretaceous, after the angiosperms appeared,which is much <strong>la</strong>ter than one would expect of such a supposedly anci<strong>en</strong>t group. The new and more complexhabitats created by the angiosperms, <strong>es</strong>pecially the for<strong>es</strong>ts they formed, offered new nich<strong>es</strong> for other organisms.Ferns were probably among the opportunists taking advantage of this, which would exp<strong>la</strong>in their Cretaceouscomeback. With more than 10,000 extant speci<strong>es</strong> they outnumber the gymnosperms t<strong>en</strong> tim<strong>es</strong>. D<strong>es</strong>pite theirdiminished diversity, gymnosperms still p<strong>la</strong>y a signi-ficant role today. Conifers in particu<strong>la</strong>r put up some seriouscompetition for angiosperms. Although angiosperms mature more quickly than gymnosperms and g<strong>en</strong>erallyproduce more seeds in the same time, conifers, at least, are better adapted to dry, cool habitats. This is why theydominate for<strong>es</strong>ts in northern <strong>la</strong>titu<strong>de</strong>s, at high elevations and on sandy soils. In fact, coniferous for<strong>es</strong>ts cover about25 per c<strong>en</strong>t of the <strong>la</strong>nd surface and provi<strong>de</strong> most of the cellulose used in papermaking.However, with an <strong>es</strong>timated 422,000 speci<strong>es</strong>, it is true that angiosperms vastly outnumber the thousandspeci<strong>es</strong> of gymnosperms that still exist today. Their incredible versatility and ability to adapt to almost all climat<strong>es</strong>and situations is unrivalled in the p<strong>la</strong>nt kingdom and has allowed them to become the unchall<strong>en</strong>ged rulers of theEarth’s flora. The fantastic diversity of the angiosperms is disp<strong>la</strong>yed in nearly every aspect of their appearance butnowhere more than in their reproductive organs, flowers, seeds and, of course, the containers in which theyproduce their seeds.The seed-bearing organs of the angiospermsThe evolution of animal-pollinated flowers with carpels and more effici<strong>en</strong>t seeds is only the first part of theangiosperm succ<strong>es</strong>s story. The second part covers the magnific<strong>en</strong>t spectrum of strategi<strong>es</strong> and adaptations<strong>de</strong>veloped by angiosperms to disperse their seeds in or<strong>de</strong>r to <strong>en</strong>sure the survival of their speci<strong>es</strong> and to expandinto new territori<strong>es</strong>. Enclosing the ovul<strong>es</strong> within carpels has many advantag<strong>es</strong>. As the seeds rip<strong>en</strong> they ev<strong>en</strong>tuallyhave to leave the mother p<strong>la</strong>nt, and for early angiosperms confinem<strong>en</strong>t in the carpel could have be<strong>en</strong> a significantobstacle. Soon though, the initial difficulty of liberating the seeds from the carpels was overcome. One way tosolve the problem of angiospermy was to <strong>de</strong>velop one-see<strong>de</strong>dn<strong>es</strong>s so that the carpel was incorporated into th<strong>es</strong>eed and both were dispersed together. This condition is still found in most in<strong>de</strong>hisc<strong>en</strong>t fruits such as nuts anddrup<strong>es</strong>. However, during the course of evolution angiosperms have perfected their seed-bearing organs, betterknown as “fruits”, and so turned this initial obstacle into another advantage. They <strong>de</strong>veloped a wi<strong>de</strong> range ofspecific adaptations, which <strong>en</strong>abled them to exploit every possible means of trans-port for their seeds and fruits,including wind, water and, most importantly, animals. Once more, the astonishing ability of the angiosperms todiversify and adapt to every avai<strong>la</strong>ble niche contributed towards their evolutionary succ<strong>es</strong>s. Each dispersal strategyis reflected in a complex syndrome of adaptations that appear after the ovul<strong>es</strong> have be<strong>en</strong> succ<strong>es</strong>sfully fer-tilised.It all starts with the wilting of a pollinated flower.MetamorphosisAs soon as a flower has be<strong>en</strong> pollinated and its ovul<strong>es</strong> fertilised there is no need to attract further pollinators. Th<strong>es</strong>howy petals and the stam<strong>en</strong>s usually wilt and wither away, while the ovary starts to swell and turn into a fruit.Insi<strong>de</strong> the ovary, the ovul<strong>es</strong> grow, the <strong>en</strong>dosperm forms, and the embryo <strong>de</strong>velops, while the soft integum<strong>en</strong>tsturn into the hard seed coat.Whereas gymnosperm seeds take at least twelve to tw<strong>en</strong>ty-four months to rip<strong>en</strong>, most angiosperms producetheir seeds much faster, usually within a few weeks or months. Bananas, for example, take only two to three months,cherri<strong>es</strong> are edible after three to four months, and mangos can be harv<strong>es</strong>ted four to five months (100-130 days) afterflowering. Coconuts take a whole year after pollination to mature and Brazil nuts need ev<strong>en</strong> longer (fifte<strong>en</strong> months).With sev<strong>en</strong> to t<strong>en</strong> years from flower to fruit, the Seychell<strong>es</strong> nut palm (Lodoicea maldivica, Arecaceae) – in many waysan angiosperm extremist – is the slow<strong>es</strong>t fruiter of all. In comparison, the fast<strong>es</strong>t reproducing weeds such as thalecr<strong>es</strong>s (Arabidopsis thaliana, Brassicaceae) cycle from seed to seed in l<strong>es</strong>s than six weeks.Going to extrem<strong>es</strong> – fruits smaller than a sperm, heavier than a millstoneAlthough we are sometim<strong>es</strong> unaware of it, every angiosperm produc<strong>es</strong> some kind of fruit: herb, shrub and treeall produce their own characteristic fruits, and the variety is <strong>en</strong>dl<strong>es</strong>s. The spectrum of siz<strong>es</strong> alone rang<strong>es</strong> from thealmost microscopic fruits of the small<strong>es</strong>t angiosperm, the aquatic watermeal (members of the g<strong>en</strong>us Wolffia), tothe <strong>en</strong>ormous fruits of Cucurbita maxima, better known as giant pumpkins. To put this into perspective: a fullgrownWolffia fruit is just 0.3mm long and smaller than a sperm of the cycad Zamia roezlii, whereas the world’s<strong>la</strong>rg<strong>es</strong>t pumpkins can weigh an incredible 600kg. The angiosperms <strong>de</strong>veloped a trem<strong>en</strong>dous diversity of shap<strong>es</strong>and mo<strong>de</strong>ls, from juicy berri<strong>es</strong> and drup<strong>es</strong>, hard nuts, winged gli<strong>de</strong>rs and helicopters, to exploding capsul<strong>es</strong> andferociously spiny pods. There are so many differ<strong>en</strong>t typ<strong>es</strong> that botanists over the past two c<strong>en</strong>turi<strong>es</strong>, in a <strong>de</strong>sperateattempt to c<strong>la</strong>ssify them, g<strong>en</strong>erated more than 150 technical fruit nam<strong>es</strong>. Exp<strong>la</strong>ining the tantalising intricaci<strong>es</strong> offruit c<strong>la</strong>ssification is beyond the scope of this book. However, to give an impr<strong>es</strong>sion of the agony botanists stillface, the main criteria that are used to bring some <strong>de</strong>gree of or<strong>de</strong>r into the overwhelming diversity of fruit typ<strong>es</strong>will be discussed briefly.A brief introduction to the c<strong>la</strong>ssification of fruitsD<strong>es</strong>pite the difficulti<strong>es</strong> botanists experi<strong>en</strong>ce with increasing att<strong>en</strong>tion to structural <strong>de</strong>tails, the three basicprincipl<strong>es</strong> of fruit c<strong>la</strong>ssification are re<strong>la</strong>tively simple: (1) the un<strong>de</strong>rlying ovary type (apocarpous or syncarpous);(2) the consist<strong>en</strong>cy of the fruit wall (soft or hard); and (3) whether or not a fruit op<strong>en</strong>s up at maturity to releaseits seeds. Since every combination of characters is possible in angiosperms, all three criteria are appliedin<strong>de</strong>p<strong>en</strong>d<strong>en</strong>tly.The three basic typ<strong>es</strong> of fruitsThe most important criterion of fruit c<strong>la</strong>ssification is the structure of the ovary from which a fruit <strong>de</strong>velops.Simple fruits like cherri<strong>es</strong>, tomato<strong>es</strong>, orang<strong>es</strong> and cucumbers <strong>de</strong>velop from one flower with only one pistil, whichcan be either a single carpel or several united carpels. Multiple fruits <strong>de</strong>velop from flowers with several separatepistils. Such flowers are typically found in primitive angiosperm famili<strong>es</strong> such as the Annonaceae, Magnoliaceae,and Winteraceae. The fruit of the tulip tree (Liriod<strong>en</strong>dron tulipifera), a member of the Magnoliaceae family, formsa cone-like structure in which each of the numerous carpels <strong>de</strong>velops into a f<strong>la</strong>t, narrow-winged fruitlet. In theWinter’s bark tree, Drimys winteri (Winteraceae), the apocarpous gynoecium of each flower produc<strong>es</strong> not just onebut a whole cluster of bean-shaped berri<strong>es</strong>. A <strong>de</strong>licious exotic re<strong>la</strong>tive of the Winter’s bark tree is the custardapple (Annona cherimo<strong>la</strong>, Annonaceae). In this case the carpels fuse together as they <strong>de</strong>velop into what appears tobe a simple fruit. The only giveaway of the originally apo-carpous gynoecium is the distinct hexagonal patternon the skin of the fruit, marking the boundari<strong>es</strong> betwe<strong>en</strong> the individual carpels. More familiar exampl<strong>es</strong> of274 Semil<strong>la</strong>s – La <strong>vida</strong> <strong>en</strong> cápsu<strong>la</strong>s <strong>de</strong> <strong>tiempo</strong>
multiple fruits are b<strong>la</strong>ckberri<strong>es</strong> (Rubus fruticosus) and raspberri<strong>es</strong> (Rubus idaeus), both members of the rose family(Rosaceae). Their <strong>de</strong>sc<strong>en</strong>t from flowers with apocarpous ovari<strong>es</strong> exp<strong>la</strong>ins why they r<strong>es</strong>emble a d<strong>en</strong>se cluster oftiny grap<strong>es</strong>, each small globule repr<strong>es</strong><strong>en</strong>ting a carpel.The simi<strong>la</strong>r-looking mulberri<strong>es</strong> (Morus nigra, Moraceae) belong to yet another major category of fruit. Themulberry do<strong>es</strong> not <strong>de</strong>velop from a single flower with several separate pistils but is the joint product of an <strong>en</strong>tireinflor<strong>es</strong>c<strong>en</strong>ce (a group of individual flowers). H<strong>en</strong>ce, each bead or fruitlet of a mulberry was once a tiny flower.Other culinary exampl<strong>es</strong> of compound fruits are pineappl<strong>es</strong> (Ananas comosus, Bromeliaceae) and the fruits ofmembers of the Moraceae family, such as figs (Ficus carica), breadfruits (Artocarpus altilis) and jackfruits (Artocarpusheterophyllus). Incid<strong>en</strong>tally, a good-size jackfruit can be up to 90cm long and weigh 50kg, which mak<strong>es</strong> it the<strong>la</strong>rg<strong>es</strong>t tree-borne fruit in the world.Drup<strong>es</strong> and Berri<strong>es</strong>The two remaining c<strong>la</strong>ssification criteria separate fruits into fl<strong>es</strong>hy and dry on<strong>es</strong>, and into those that remain closed(in<strong>de</strong>hisc<strong>en</strong>t fruits) or op<strong>en</strong> to release their seeds (<strong>de</strong>hisc<strong>en</strong>t fruits). Although the two criteria are appliedin<strong>de</strong>p<strong>en</strong>d<strong>en</strong>tly, soft and fl<strong>es</strong>hy fruits are usually in<strong>de</strong>hisc<strong>en</strong>t and eat<strong>en</strong> by frugivor<strong>es</strong>, which disperse the seeds intheir faec<strong>es</strong>. The two principal typ<strong>es</strong> of fl<strong>es</strong>hy in<strong>de</strong>hisc<strong>en</strong>t fruits are drup<strong>es</strong> and berri<strong>es</strong>.In drup<strong>es</strong> like cherri<strong>es</strong>, plums, peach<strong>es</strong> and mango<strong>es</strong>, the pericarp (the ovary wall in a ripe fruit) isdiffer<strong>en</strong>tiated into three <strong>la</strong>yers: the thin outer epicarp (the skin of the fruit); the fl<strong>es</strong>hy m<strong>es</strong>ocarp (the actual pulpof the fruit); and a hard inner <strong>la</strong>yer called <strong>en</strong>docarp. Epi- and m<strong>es</strong>ocarp are the edible parts, whereas the <strong>en</strong>docarpforms the usually single-see<strong>de</strong>d stone. In berri<strong>es</strong> the pericarp is <strong>en</strong>tirely soft and juicy and remains by <strong>de</strong>finitionin<strong>de</strong>hisc<strong>en</strong>t. In contrast with drup<strong>es</strong>, berri<strong>es</strong> mostly contain several or many seeds. Typical multi-see<strong>de</strong>d berri<strong>es</strong>are tomato<strong>es</strong>, cucumbers, grap<strong>es</strong> and blueberri<strong>es</strong>. The avocado is an example of a single-see<strong>de</strong>d berry. Asb<strong>la</strong>ckberri<strong>es</strong>, raspberri<strong>es</strong> and mulberri<strong>es</strong> have already prov<strong>en</strong>, many fruits we call berri<strong>es</strong> are botanically speakingnot berri<strong>es</strong> at all. Juniper berri<strong>es</strong> are the most <strong>de</strong>ceptive example. Famous for giving Dutch g<strong>en</strong>ever and gintheir characteristic f<strong>la</strong>vour, juniper berri<strong>es</strong> are not berri<strong>es</strong> but the con<strong>es</strong> of a gymnosperm called Juniperuscommunis (Cupr<strong>es</strong>saceae). In the two to three years during which the female con<strong>es</strong> of this dioecious coniferrip<strong>en</strong>, their three uppermost scal<strong>es</strong> <strong>de</strong>velop into a blue, fl<strong>es</strong>hy <strong>la</strong>yer that <strong>en</strong>clos<strong>es</strong> the seeds almost like thepericarp of a true berry.True nutsIn<strong>de</strong>hisc<strong>en</strong>t fruits with an <strong>en</strong>tirely hard, dry pericarp are called nuts. The fruit wall of a nut remains closed, makingthe in<strong>de</strong>p<strong>en</strong>d<strong>en</strong>t distribution of several seeds impossible. For the same reason as drup<strong>es</strong>, nuts are therefore mostlysingle-see<strong>de</strong>d. As the above <strong>de</strong>finition indicat<strong>es</strong>, sci<strong>en</strong>tists use the term nut in a much more precise and slightlydiffer<strong>en</strong>t way from everyday <strong>la</strong>nguage. Botanically, nuts do not inclu<strong>de</strong> only familiar nuts such as hazelnuts,peanuts and cashew nuts but also “winged nuts” (samaras), like those of elms (Ulmus spp.) and ash (Fraxinus spp.),and much smaller fruits which most people wrongly think of as seeds.Nuts that are seeds and seeds that are nutsAccording to the botanical <strong>de</strong>finition of a nut, the small grains of the grass family (which inclu<strong>de</strong>s all our cereals)and the “seeds” of the sunflower are, in fact, miniature nuts. Every grain of rice, wheat or oat – small though itmay be – is a fruit. Likewise, every sunflower “seed” is a small nut that <strong>de</strong>velops from one of the hundreds of tinyflowers arranged spirally across the disc of the inflor<strong>es</strong>c<strong>en</strong>ce. Note that, as is typical for a member of the sunflowerfamily, a sunflower blossom is not a single flower but an <strong>en</strong>tire inflor<strong>es</strong>c<strong>en</strong>ce mimicking the looks of an individualflower. Some re<strong>la</strong>tiv<strong>es</strong> of the sunflower, like dan<strong>de</strong>lion (Taraxacum officinale), meadow salsify (Tragopogon prat<strong>en</strong>sis)and mouse-ear hawkweed (Hieracium pilosel<strong>la</strong>) attach a kind of parachute (a pappus) to their tiny nuts, whichallows them to catch a ri<strong>de</strong> on the wind. With all th<strong>es</strong>e differ<strong>en</strong>c<strong>es</strong> betwe<strong>en</strong> the various kinds of nuts sci<strong>en</strong>tistsbegan to create nam<strong>es</strong> for each type. B<strong>es</strong>i<strong>de</strong>s calling winged nuts samaras, botanists gave the by now traditionalterm caryopsis to the fruit of grass<strong>es</strong>, mainly because their thin pericarp is tightly attached to the seed coat. Smallnuts with a rather soft pericarp that is separate from the seed coat, such as those of the sunflower, were namedach<strong>en</strong><strong>es</strong>, or cypse<strong>la</strong>s if, like the dan<strong>de</strong>lion, they have a parachute attached.All typ<strong>es</strong> of nuts or nutlets have one feature in common: functionally they act in the same way as seeds.The protective role of the seed coat has be<strong>en</strong> tak<strong>en</strong> over by the hard<strong>en</strong>ed fruit wall (pericarp). This exp<strong>la</strong>ins why,in common par<strong>la</strong>nce, we refer to nuts as seeds and to seeds as nuts. Exampl<strong>es</strong> of the <strong>la</strong>tter are treats like (unshelled)pine nuts (Pinus pinea, Pinaceae), Brazil nuts (Bertholletia excelsa, Lecythidaceae) and macadamia nuts (Macadamiaintegrifolia and M. tetraphyl<strong>la</strong>, Proteaceae). While still on the tree, each macadamia seed is wrapped individually ina leathery gre<strong>en</strong> husk (the pericarp), which <strong>la</strong>ter splits along one si<strong>de</strong>; but the split is much too narrow to allowthe seed to <strong>es</strong>cape. Unshelled almonds (Prunus dulcis, Rosaceae) and pistachios (Pistacia vera, Anacardiaceae) are,in fact, the ston<strong>es</strong> of drupaceous fruits.NutletsAn inter<strong>es</strong>ting variation on the simple fruit theme is fruits, which – <strong>de</strong>spite <strong>de</strong>veloping from a syncarpous ovary– disintegrate into their carpel<strong>la</strong>ry constitu<strong>en</strong>ts wh<strong>en</strong> they are ripe. Such schizocarpic fruits break up into two ormore fruitlets, each fruitlet consisting of either a whole or half carpel. The fruitlets themselv<strong>es</strong> are usually dry,remain closed, and contain only a single seed, which mak<strong>es</strong> them almost real nuts. As they are fruitlets rather thanfruits, they are called nutlets instead of nuts. Schizocarpic fruits are typical of the carrot family (Apiaceae). Exampl<strong>es</strong>of this fruit type can be found on any spice shelf – caraway, cumin, corian<strong>de</strong>r, aniseed and f<strong>en</strong>nel. The wingedfruitlets of maple (Acer spp., Sapindaceae), which form a pair in the intact fruit, are another familiar example ofschizocarpic fruits.The fruitlets of both the Apiaceae and Acer repr<strong>es</strong><strong>en</strong>t a single <strong>en</strong>tire carpel. In other famili<strong>es</strong> the divisionof the fruit go<strong>es</strong> one step further. The ovary of the mint family (Lamiaceae) and the borage family (Boraginaceae)g<strong>en</strong>erally consists of two fused carpels, which are <strong>de</strong>eply lobed to form twice as many compartm<strong>en</strong>ts. At maturity,the four compart-m<strong>en</strong>ts separate into single-see<strong>de</strong>d nutlets, each consisting of half a carpel. Ev<strong>en</strong> to theexperi<strong>en</strong>ced eye, the seed-bearing half-carpels of Salvia (sage), Origanum (oregano), Thymus (thyme) and otherLamiaceae look <strong>de</strong>ceptively like true seeds, which is what most people consi<strong>de</strong>r them to be. A bizarre exampleof a schizocarpic fruit is that of Ochna natalitia (Ochnaceae). In the flowers of the g<strong>en</strong>us Ochna, the carpels arejoined only at the base and share a common style. Once pollinated and fertilised, the carpels <strong>de</strong>velop into separatedrupelets that sit, like eggs around the edge of a p<strong>la</strong>te, on an <strong>en</strong><strong>la</strong>rged, fl<strong>es</strong>hy red flower axis.Capsul<strong>es</strong>Fruits in which the pericarp op<strong>en</strong>s (<strong>de</strong>hisc<strong>es</strong>) to expose or release the seeds are called capsul<strong>es</strong>. There are several waysin which the pericarp can op<strong>en</strong>: through por<strong>es</strong>, by a circu<strong>la</strong>r split <strong>de</strong>fining a lid, or by regu<strong>la</strong>r longitudinal slits.Poppy capsul<strong>es</strong>, for example, <strong>de</strong>hisce via a ring of por<strong>es</strong> around the top. The seeds are ejected through th<strong>es</strong>epor<strong>es</strong> like salt from a saltshaker as the fruits sway in the wind on their long, sl<strong>en</strong><strong>de</strong>r stalks. Exampl<strong>es</strong> of capsul<strong>es</strong>that op<strong>en</strong> with a lid are those of the scarlet pimpernel (Anagallis arv<strong>en</strong>sis, Primu<strong>la</strong>ceae), the twinleaf (Jeffersoniadiphyl<strong>la</strong>, Berberidaceae), the squirting cucumber (Ecballium e<strong>la</strong>terium, Cucurbitaceae) and the monkey pot(Lecythis pisonis, Lecythidaceae) from tropical South America. However, most capsul<strong>es</strong> split op<strong>en</strong> along regu<strong>la</strong>rlin<strong>es</strong>, which either follow the septae (the walls betwe<strong>en</strong> the individual carpels of a syncarpous gynoecium) or rundown the middle of each locule (the seed-bearing cavity of a carpel). Capsul<strong>es</strong> of the first type are called septicidalcapsul<strong>es</strong>, those of the second type loculicidal capsul<strong>es</strong>. Septicidal capsul<strong>es</strong> are l<strong>es</strong>s common than loculicidalcapsul<strong>es</strong>; the foxglove (Digitalis spp., P<strong>la</strong>ntaginaceae) and til<strong>la</strong>ndsias (Til<strong>la</strong>ndsia spp., Bromeliaceae) are twoexampl<strong>es</strong> with septicidal capsul<strong>es</strong>. The more common loculicidal capsul<strong>es</strong> are found in many Dicots such as thehorse ch<strong>es</strong>tnut (A<strong>es</strong>culus hippocastanum, Sapindaceae), the Hima<strong>la</strong>yan balsam (Impati<strong>en</strong>s g<strong>la</strong>ndulifera, Balsaminaceae),and in many Monocots such as iris<strong>es</strong> (Iris spp., Iridaceae), alliums (Allium spp., Alliaceae) and fritil<strong>la</strong>ri<strong>es</strong> (Fritil<strong>la</strong>riaspp., Liliaceae).Capsul<strong>es</strong> are mostly <strong>de</strong>scribed as dry fruits but notable exceptions inclu<strong>de</strong> the fl<strong>es</strong>hy capsul<strong>es</strong> of theHima<strong>la</strong>yan balsam (Impati<strong>en</strong>s g<strong>la</strong>ndulifera, Balsaminaceae), the squirting cucumber (Ecballium e<strong>la</strong>terium,Cucurbitaceae) and tropical hedychiums (Hedychium spp., Zingiberaceae).To fully conform to the strict botanical<strong>de</strong>finition of a capsule, a fruit not only has to be <strong>de</strong>hisc<strong>en</strong>t but must also <strong>de</strong>velop from a syncarpous ovarycomposed of two or more united carpels. Thus, the fruit of the twinleaf, which as a member of the barberryfamily (Berberidaceae) has only one carpel per flower, is strictly a capsu<strong>la</strong>r fruit and not a true capsule. The sameappli<strong>es</strong> to the <strong>en</strong>tire legume family (Fabaceae) in which the typical fruit is a single carpel that op<strong>en</strong>s along thedorsal and v<strong>en</strong>tral suture, splitting the fruit in half. Their capsu<strong>la</strong>r fruit, familiar from many ornam<strong>en</strong>tal (e.g.lupins, sweet peas) and edible p<strong>la</strong>nts (e.g. beans, peas, l<strong>en</strong>tils) is traditionally called a legume. If a single carpel op<strong>en</strong>sonly along one line (usually along the v<strong>en</strong>tral si<strong>de</strong>), as in the multiple fruits of the buttercup family(Ranuncu<strong>la</strong>ceae) and star anise (Illicium verum, Schisandraceae) or in the cone-like compound fruits of banksias(Banksia spp., Proteaceae), it is termed a follicle.In<strong>de</strong>hisc<strong>en</strong>t capsul<strong>es</strong>Legum<strong>es</strong> and follicl<strong>es</strong> earned their own nam<strong>es</strong> by not conforming to the <strong>de</strong>finition of a true capsule; they failedon one criterion, the syncarpous ovary. In other cas<strong>es</strong>, many botanists seem willing to turn a blind eye: forexample, they refer to the “fruit” of the baobab (Adansonia digitata, Malvaceae) as an in<strong>de</strong>hisc<strong>en</strong>t capsule. The <strong>la</strong>rgefruit of the baobab tree do<strong>es</strong> give the overall impr<strong>es</strong>sion of a capsule but it do<strong>es</strong> not op<strong>en</strong>, h<strong>en</strong>ce the paradoxicalname in<strong>de</strong>hisc<strong>en</strong>t (non-op<strong>en</strong>ing) capsule. The dry, hard shell of the baobab fruit loosely <strong>en</strong>clos<strong>es</strong> thirty seeds, whichhas led to the name “Judas fruit”. The seeds themselv<strong>es</strong> are embed<strong>de</strong>d in a white mealy pulp, called “cream oftartar”. This pulp is high in vitamin C and used to prepare porridge for newborn babi<strong>es</strong> if the mother hasinsuffici<strong>en</strong>t milk, or it is ma<strong>de</strong> into a refr<strong>es</strong>hing drink to treat fever and diarrhoea; it can also serve as a chol<strong>es</strong>terolfreeParm<strong>es</strong>an substitute. Another “square peg” in the “round hole” <strong>de</strong>finition of a capsule is the famous Brazilnut tree (Bertholletia excelsa). Its <strong>la</strong>rge woody fruits take fourte<strong>en</strong> months to mature and their pericarp is so hardEnglish texts 275
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R O B K E S S E L E R Y W O L F G A
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S E M I L L A SL A V I DA E N C Á
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Erica cinerea (Ericaceae) - brezo;
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Thamnosma africanum (Rutaceae); rec
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INTRODUCCIÓNAntirrhinum coulterian
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LA VIDA EN CÁPSULAS DE TIEMPORO B
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Esta nueva pasión sentó las bases
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20 Semillas - La vida en cápsulas
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22 Semillas - La vida en cápsulas
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Semillas - La vida en cápsulas de
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nadar libremente hasta encontrar un
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Cuando los machos son micro y las h
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página anterior arriba: Archaeospe
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Semillas desnudasLos óvulos de las
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(Ginkgoaceae), propio orden (Ginkgo
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Cuando mega realmente significa meg
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página anterior: Pinus lambertiana
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página anterior: Drimys winteri (W
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vino dado por la combinación de mi
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las plantas en 1664 y en 1672 publi
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especialmente aquellas que oliendo
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página anterior: Angraecum sesquip
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La sorprendente vida sexual de las
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Melocactus zehntneri (Cactaceae) -
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el otro baja hacia la célula centr
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62 Semillas - La vida en cápsulas
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angiospermas en dos grupos, las dic
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Una gran variedad de embriones de d
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sus hojas. Estos embriones almacena
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70 Semillas - La vida en cápsulas
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los aspectos de su apariencia, pero
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abajo: Punica granatum (Lythraceae)
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Una breve introducción a la clasif
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abajo: secciones transversales de u
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página anterior: Scutellaria orien
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página anterior: Ochna natalitia (
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agutí logra perforar un agujero de
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La dispersión de frutos y semillas
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predestinadas a fracasar en su empe
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Centrolobium microchaete (Fabaceae)
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96 Semillas - La vida en cápsulas
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Semillas de espuela de caballero (R
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La dispersión de frutos y semillas
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página anterior: Darlingtonia cali
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página anterior: Clematis tangutic
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izquierda: Blepharis mitrata (Acant
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abajo: Arenaria franklinii (Caryoph
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página anterior: Cistanche tubulos
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La dispersión de frutos y semillas
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específicos requerimientos de germ
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La dispersión de frutos y semillas
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página anterior: : habas de mar -
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Frutos explosivos activosLos frutos
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ecta mientras la parte inferior est
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pero un sentido del olfato poco des
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La dispersión de frutos y semillas
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página anterior y arriba: Afzelia
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Polygala arenaria (Polygalaceae) -
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Dispersión por recolectores y alma
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La dispersión de frutos y semillas
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La dispersión de frutos y semillas
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página anterior: Uncarina spp. (Pe
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Semillas sin ninguna adaptación ob
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Viajar en el tiempo y el espacio 16
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Un año de semillas, siete años de
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página anterior: Nemesia versicolo
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Viajar en el tiempo y el espacio 17
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arriba: Strelitzia reginae (Strelit
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UN PROYECTO ARQUITECTÓNICORO B K E
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página anterior: Cleome sp. (Cappa
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página siguiente: Downland Gridshe
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página siguiente: El Proyecto Edé
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FITOPIARO B K E S S E L E RStellari
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La diferencia entre mirar y ver...
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Trichodesma africanum (Boraginaceae
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página anterior: Crassula pellucid
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Calandrinia eremaea (Portulacaceae)
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Alcea pallida (Malvaceae) - malva p
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Codonocarpus cotinifolius (Gyrostem
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Fitopia 211
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Melocactus neryi (Cactaceae) - melo
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Lophophora williamsii (Cactaceae) -
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Lychnis flos-cuculi (Caryophyllacea
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