grow in muddy swamp<strong>la</strong>nds regu<strong>la</strong>rly inundated by the ti<strong>de</strong>s. Apart from their characteristic stilt roots, the mostsignificant morphological adaptation to their extreme habitat is their method of reproduction. Rather thanshedding ordinary seeds, most of which would be washed away by the ti<strong>de</strong>s and lost, mangrov<strong>es</strong> are viviparous.Vivipary (live birth) in animals means that the embryo grows insi<strong>de</strong> the mother (as in most mammals) not insi<strong>de</strong>an egg (as in most reptil<strong>es</strong> and birds). Viviparous p<strong>la</strong>nts such as mangrov<strong>es</strong> produce seeds that germinate whil<strong>es</strong>till attached to the par<strong>en</strong>t p<strong>la</strong>nt. Once the zygote has be<strong>en</strong> fertilised the embryo simply continu<strong>es</strong> to grow. Byext<strong>en</strong>ding its hypocotyl, it soon p<strong>en</strong>etrat<strong>es</strong> the thin seed coat and breaks through the wall of the berry. A fullygrownembryo of the most common red mangrove (Rhizophora mangle, Rhizophoraceae) may be 25cm long.Ev<strong>en</strong>tually, the club-shaped “seedling” drops off and either p<strong>la</strong>nts itself immediately in the mud un<strong>de</strong>rneath orfloats in the sea until the next ti<strong>de</strong> sets it down elsewhere. The advantag<strong>es</strong> of vivipary in the mangrov<strong>es</strong>’ habitatare clear: by supporting their embryos until they have grown into sizeable, well-differ<strong>en</strong>tiated p<strong>la</strong>ntlets, mangrov<strong>es</strong>give their offspring a head start. Ready to go, and equipped with some r<strong>es</strong>erv<strong>es</strong>, mangrove seedlings root veryquickly once they touch ground.Ocean travellersThe seeds and fruits of many p<strong>la</strong>nts growing on or in the vicinity of coastlin<strong>es</strong> ev<strong>en</strong>tually <strong>en</strong>d up in the sea,where they are carried away by ocean curr<strong>en</strong>ts. Fruits and seeds may be shed directly on the beach or drop intotidal pools and swamps from where the ti<strong>de</strong> collects them. Those originating further in<strong>la</strong>nd reach the sea viastreams and rivers, in many cas<strong>es</strong> accid<strong>en</strong>tally. However, a number of p<strong>la</strong>nts, <strong>es</strong>pecially in the tropics, poss<strong>es</strong>sdiaspor<strong>es</strong> specifically adapted to travel in sea water for months or ev<strong>en</strong> years; one example, the fruit of themangrove tree Cerbera manghas, has already be<strong>en</strong> m<strong>en</strong>tioned. Once such seaworthy diaspor<strong>es</strong> reach the mainocean surface curr<strong>en</strong>ts, their journey can take them thousands of kilometr<strong>es</strong> away from their p<strong>la</strong>ce of origin.Charl<strong>es</strong> Darwin was <strong>en</strong>thralled by the i<strong>de</strong>a that seeds from tropical countri<strong>es</strong> could travel to Europe. Some peoplecollect exotic drift seeds and fruits, popu<strong>la</strong>rly known as “sea beans”, as a hobby.Fascinating though it may be, drifting with ocean curr<strong>en</strong>ts is – like wind dispersal – a very haphazard andwasteful strategy. Many drift fruits and seeds lose their buoyancy and are likely to <strong>en</strong>d up on the bottom of theocean or somewhere with unsuitable living conditions. For example, tropical fruits and seeds from South Americaand the Caribbean are regu<strong>la</strong>rly carried by the Gulf Stream to the rather inhospitable beach<strong>es</strong> of northernEurope. The most frequ<strong>en</strong>t arrivals from the New World are members of the legume family (Fabaceae), whichis probably why people call them “sea beans”. Th<strong>es</strong>e seeds must have appeared strange to people throughouthistory, <strong>es</strong>pecially in the Middle Ag<strong>es</strong>. It is not surprising that so many stori<strong>es</strong>, leg<strong>en</strong>ds and superstitious beliefsare wov<strong>en</strong> around them. Christopher Columbus’s voyage of discovery was allegedly inspired by the exotic seaheart(Entada gigas) and the people of Porto Santo in the Azor<strong>es</strong> still call the seed fava <strong>de</strong> Colom (Columbus’sbean). Today, collectors and creators of botanical jewellery value them for their beautiful shap<strong>es</strong> and colours. Apartfrom the sea-heart, the most famous sea beans are the true sea bean (Mucuna sloanei and M. ur<strong>en</strong>s), the sea purse(Dioclea reflexa), grey and yellow nickernuts (Ca<strong>es</strong>alpinia bonduc, C. major), and Mary’s bean (Merremiadiscoi<strong>de</strong>sperma). Most are legum<strong>es</strong>, although Mary’s bean belongs to the morning glory family (Convolvu<strong>la</strong>ceae).Entada gigas grows as an <strong>en</strong>ormous liana in the tropical for<strong>es</strong>ts of C<strong>en</strong>tral and South America and Africa.Its seeds are one of the most commonly found drift diaspor<strong>es</strong> on European beach<strong>es</strong>. With a diameter of up to5cm, the heart-shaped brown seeds are themselv<strong>es</strong> <strong>la</strong>rge but are also borne in the <strong>la</strong>rg<strong>es</strong>t of all legume pods, upto 1.80m long. Sea- hearts and the <strong>la</strong>rge seeds of the re<strong>la</strong>ted Entada phaseoloi<strong>de</strong>s from Africa and Australia werecommonly carved into snuff box<strong>es</strong> and lockets in Norway and other parts of Europe. In Eng<strong>la</strong>nd, the seeds wereused as teething rings and good luck charms to protect childr<strong>en</strong> at sea. Grey nickernuts were worn as amuletsby the people of the Hebri<strong>de</strong>s to ward off the Evil Eye. The seeds were said to turn b<strong>la</strong>ck wh<strong>en</strong> the wearer wasin danger. The most intriguing sea bean has to be the Mary’s bean. Produced by a woody vine that grows in thefor<strong>es</strong>ts of southern Mexico and C<strong>en</strong>tral America, the b<strong>la</strong>ck or brown, globose to oblong seeds are 20-30mm indiameter and 15-20mm thick. Their hallmark is a cross formed by two groov<strong>es</strong>, h<strong>en</strong>ce their name “crucifixionbean” or “Mary’s bean”. To religious people this seed had a symbolic meaning. Having survived the ocean, it wasbelieved to give protection to anyone who owned it. In the Hebri<strong>de</strong>s, for example, a woman in <strong>la</strong>bour holdinga Mary’s bean was assured of an easy <strong>de</strong>livery. The seeds were han<strong>de</strong>d down as precious talismans from motherto daughter for g<strong>en</strong>erations.Apart from its r<strong>es</strong>ili<strong>en</strong>ce to sea water, the most important pre-requisite of a diaspore to succ<strong>es</strong>sfully travelthe oceans is buoyancy. Most tropical diaspor<strong>es</strong> do not float either in fr<strong>es</strong>h or sea water. It is <strong>es</strong>timated that l<strong>es</strong>sthan one perc<strong>en</strong>t of tropical seed p<strong>la</strong>nts produce fruits or seeds that drift in seawater for at least a month. Thos<strong>es</strong>pecifically adapted to water dispersal poss<strong>es</strong>s various buoyancy <strong>de</strong>vic<strong>es</strong>. Seeds increase their specific gravity bynot filling the <strong>en</strong>tire cavity of the thick, woody seed coat (e.g. kemiri nut, Aleurit<strong>es</strong> moluccana, Euphorbiaceae), byleaving an air-filled gap betwe<strong>en</strong> their two cotyledons (e.g. Entada spp., Mucuna spp., Merremia spp., Moramegistosperma), or by producing lightweight cotyledonary tissue (e.g. legum<strong>es</strong> like Dioclea spp.). Seaworthy fruitsmay or may not combine th<strong>es</strong>e characteristics with air caviti<strong>es</strong> in their pericarp, or a fibrous or corky coat. Withjust the <strong>la</strong>tter, box fruits (Barringtonia asiatica, Lecythidaceae), common flotsam on the beach<strong>es</strong> of Fr<strong>en</strong>chPolyn<strong>es</strong>ia, remain buoyant for at least two years.The coconut (Cocos nucifera, Arecaceae) combin<strong>es</strong> a fibrous, spongy fruit coat with an air bubble in the<strong>en</strong>dosperm cavity. The seed insi<strong>de</strong> is protected by a thick, hard <strong>en</strong>docarp, making the fruit of the coconut a drydrupe rather than a nut. Although no one would call it a sea bean, the coconut is the c<strong>la</strong>ssic example of an oceantraveller. Its excell<strong>en</strong>t adaptation to sea dispersal has spread coconut palms throughout the tropics. The averagemaximum distance that a coconut can travel while still afloat and viable is 5,000 kilometr<strong>es</strong>. Wh<strong>en</strong> it finallybecom<strong>es</strong> stran<strong>de</strong>d on a beach it will germinate slowly once rainfall has washed off the salt collected during itsjourney. Since sea-sand retains hardly any moisture, the liquid <strong>en</strong>dosperm insi<strong>de</strong> the coconut provi<strong>de</strong>s a crucialwater r<strong>es</strong>erve from germination until the roots of the seedling reach fr<strong>es</strong>h groundwater.The most <strong>en</strong>igmatic of drift fruits is the Seychell<strong>es</strong> nut, the fruit with the <strong>la</strong>rg<strong>es</strong>t seed in the world. Althoughnot closely re<strong>la</strong>ted, the Seychell<strong>es</strong> nut is simi<strong>la</strong>r to a coconut, h<strong>en</strong>ce its alternative nam<strong>es</strong> double coconut and coco <strong>de</strong>mer. Unlike the coconut, the Seychell<strong>es</strong> nut is not adapted to ocean dispersal. It cannot float wh<strong>en</strong> fr<strong>es</strong>h and do<strong>es</strong>not survive prolonged contact with sea water. In the fifte<strong>en</strong>th c<strong>en</strong>tury, long before the Seychell<strong>es</strong> were discoveredin 1743, the <strong>en</strong>docarps were washed up on the beach<strong>es</strong> of the Indian Ocean. Since most of them were found onthe Maldiv<strong>es</strong> the speci<strong>es</strong> was giv<strong>en</strong> the somewhat misleading Latin name Lodoicea maldivica. The true distributionof this extraordinary palm tree is limited to two is<strong>la</strong>nds of the Seychell<strong>es</strong>, Praslin and Curieuse. The Seychell<strong>es</strong> nutis famous not only for its size but also for the rather sugg<strong>es</strong>tive shape of its fruits, which gave rise to varioussuperstitions. Ma<strong>la</strong>y and Chin<strong>es</strong>e sailors thought that the double coconut grew on a mysterious un<strong>de</strong>rwater tre<strong>es</strong>imi<strong>la</strong>r to a coconut palm. In Europe, it was thought that the highly-prized fruits had medicinal properti<strong>es</strong> andthat their <strong>en</strong>dosperm was an antidote to poison. Just how valuable the coco <strong>de</strong> mer was before the discovery of theSeychell<strong>es</strong> is <strong>de</strong>scribed by Albert Smith Bickmore in his Travels in the East Indian Archipe<strong>la</strong>go, published in 1869:“To early sailors in the Indian Ocean, the Seychell<strong>es</strong> nut was only known from drift seeds washed ashore throughout the region.The prince of Ceylon, who is said to have giv<strong>en</strong> a whole v<strong>es</strong>sel <strong>la</strong>d<strong>en</strong> with spice for a single specim<strong>en</strong> of the double coconut,could have satisfied his heart’s full<strong>es</strong>t <strong>de</strong>sire if he had only known it was not rare on the Seychell<strong>es</strong>, north of Mauritius.”Self-dispersalRather than <strong>en</strong>trusting their seeds to wind, water or animals, some p<strong>la</strong>nts have <strong>de</strong>veloped mechanisms that <strong>en</strong>ablethem to disperse their seeds themselv<strong>es</strong>, at least for a short distance. Self-dispersal or autochory is either by activelycatapulting the seeds away (ballistic dispersal) or by burying the fruits in the ground (geocarpy).Ballistic dispersalMechanisms used by p<strong>la</strong>nts to expel their seeds from their fruits can be caused either by passive (hygroscopic)movem<strong>en</strong>ts of <strong>de</strong>ad tissu<strong>es</strong> or by active movem<strong>en</strong>ts that are due to high pr<strong>es</strong>sure in living cells.Passive explosiv<strong>es</strong>Dry, <strong>de</strong>hisc<strong>en</strong>t fruits such as capsul<strong>es</strong> and follicl<strong>es</strong> op<strong>en</strong> gradually along pre-formed lin<strong>es</strong> as the pericarp di<strong>es</strong> anddri<strong>es</strong> out. The shrinking of its tissu<strong>es</strong> ev<strong>en</strong>tually caus<strong>es</strong> the fruit wall to rupture. This is usually a slow, gradualproc<strong>es</strong>s but in some fruits the pericarp is specifically adapted to build up a high <strong>de</strong>gree of t<strong>en</strong>sion, which isev<strong>en</strong>tually released by a sudd<strong>en</strong> explosion that expels the seeds. The un<strong>de</strong>rlying principle of this mechanism isbased on the differ<strong>en</strong>t ori<strong>en</strong>tation of elongated cells in neighbouring <strong>la</strong>yers, which oft<strong>en</strong> consist of crossed, thickwalledfibr<strong>es</strong>. As the cells dry out, they contract parallel to their longitudinal axis, causing the neighbouring <strong>la</strong>yersto pull in differ<strong>en</strong>t directions. The t<strong>en</strong>sion is finally released in an explosive torsion movem<strong>en</strong>t of the separatingfruit fragm<strong>en</strong>ts, which usually corr<strong>es</strong>pond to whole or half carpels. In some legum<strong>es</strong> it is the two halv<strong>es</strong> of th<strong>es</strong>ingle carpel which separate along their dorsal and v<strong>en</strong>tral si<strong>de</strong>s. By twisting in opposite directions at lightningspeed, the carpel-halv<strong>es</strong> fling out the seeds with great force. Common broom (Cytisus scoparius), gorse (Ulexeuropaeus), sweet peas (Lathyrus odoratus) and lupins (Lupinus spp.) are familiar exampl<strong>es</strong> disp<strong>la</strong>ying this behaviour.Usually the seeds are dispersed over a short distance. In gorse, they remain within the radius of the mother p<strong>la</strong>ntwith only about 2 per c<strong>en</strong>t travelling 2-2.5m. As is oft<strong>en</strong> the case, the distanc<strong>es</strong> are more impr<strong>es</strong>sive in the tropics.The fruit of Tetraberlinia morelina, an African legume tree at home in the rainfor<strong>es</strong>ts of w<strong>es</strong>t Gabon and southw<strong>es</strong>tCameroon, ai<strong>de</strong>d by its great height, shoots its seeds up to 60m from the mother p<strong>la</strong>nt. This is the long<strong>es</strong>t ballisticdispersal distance ever recor<strong>de</strong>d.A p<strong>la</strong>nt family in which the typical fruit is an explosively <strong>de</strong>hisc<strong>en</strong>t capsule is the Euphorbiaceae. Intemperate herbaceous members such as the petty spurge (Euphorbia peplus), sun spurge (Euphorbia helioscopia), dog’smercury (Mercurialis per<strong>en</strong>nis) and annual mercury (Mercurialis annua) the fruit is composed of three carpels. Wh<strong>en</strong>it explo<strong>de</strong>s, the fruit disintegrat<strong>es</strong> into six half-carpels, which churn out three seeds. The most remarkable examplein this family is native to the tropics of the New World. Not only has the gynoecium of the sandbox tree (Huracrepitans) from South America and the Caribbean many more (5-20) carpels than is usual in spurg<strong>es</strong>. The ribbedfruit the size of a mandarin erupts much more viol<strong>en</strong>tly, with the sound of a loud gunshot (Latin crepare = to burst).280 Semil<strong>la</strong>s – La <strong>vida</strong> <strong>en</strong> cápsu<strong>la</strong>s <strong>de</strong> <strong>tiempo</strong>
The force of the b<strong>la</strong>st catapults the f<strong>la</strong>t discoid seeds, of which there are as many as there are carpels, as far as 14m.The name sandbox tree dat<strong>es</strong> back to the time before the inv<strong>en</strong>tion of blotting paper and fountain p<strong>en</strong>s, wh<strong>en</strong> thefruits served as containers for the sand used to blot the ink that ran profusely from goose quills.Active explosiv<strong>es</strong>Fl<strong>es</strong>hy <strong>de</strong>hisc<strong>en</strong>t fruits are able to build up suffici<strong>en</strong>t pr<strong>es</strong>sure to explo<strong>de</strong> by means of tissu<strong>es</strong> of thin-walled cells,which increase their internal cell pr<strong>es</strong>sure (called turgor) by taking up additional water. As the tissu<strong>es</strong> swell, at somepoint the pr<strong>es</strong>sure against neighbouring ine<strong>la</strong>stic <strong>la</strong>yers is so high that the slight<strong>es</strong>t movem<strong>en</strong>t of the fruit sets offthe explosion that expels the seeds. A passing animal can trigger the mechanism and although the seeds of suchfruits are g<strong>en</strong>erally smooth and non-sticky, they may become <strong>en</strong>tangled in its fur and be carried some distance.C<strong>la</strong>ssic exampl<strong>es</strong> of actively exploding fruits are the touch-me-not (Impati<strong>en</strong>s spp.) in the balsam family(Balsaminaceae), and the squirting cucumber (Ecballium e<strong>la</strong>terium) in the gourd family (Cucurbitaceae).Anyone who has p<strong>la</strong>yed with the fruits of the touch-me-not, g<strong>en</strong>tly squeezing them betwe<strong>en</strong> two fingers,knows that its name is appropriate. Its p<strong>en</strong>dulous fruits are composed of five carpels, with pre-formed <strong>de</strong>hisc<strong>en</strong>celin<strong>es</strong> betwe<strong>en</strong> them. Pr<strong>es</strong>sure builds up in the outer <strong>la</strong>yers of the pericarp and wh<strong>en</strong> it is strong <strong>en</strong>ough to separatethe carpels, the slight<strong>es</strong>t movem<strong>en</strong>t caus<strong>es</strong> the fruits to explo<strong>de</strong>. The outsi<strong>de</strong> of the pericarp expands more thanthe insi<strong>de</strong> causing the fruit valv<strong>es</strong> to curve inwards in a lightning-fast movem<strong>en</strong>t that hurls the seeds up to 5maway. The trigger for the explosion can be a passing animal, raindrops or water dripping off a tree, the wind, orev<strong>en</strong> seeds <strong>la</strong>unched from a neighbouring fruit. As the fruits of the Mediterranean squirting cucumber rip<strong>en</strong>, theinner tissue <strong>la</strong>yers build up turgor pr<strong>es</strong>sure against the thick, rather ine<strong>la</strong>stic outer skin. The pre-<strong>de</strong>terminedbreaking line at the base of the stalk points up in the air as the fruit turns almost 180º against the pedicel.Ev<strong>en</strong>tually, the slight<strong>es</strong>t movem<strong>en</strong>t caus<strong>es</strong> the stalk to pop out like a cork, leaving a hole through which bothwatery juice and seeds squirt out. The pr<strong>es</strong>sure insi<strong>de</strong> the fruit is so high that the seeds can travel more than 10m.The true record breakers are dwarf mistleto<strong>es</strong>, which apply more or l<strong>es</strong>s the same principle as the squirtingcucumber. Predominantly native to North America, they are parasit<strong>es</strong> on pine tre<strong>es</strong> and belong to the g<strong>en</strong>usArceuthobium in the Christmas mistletoe family (Viscaceae; rec<strong>en</strong>tly united with the sandalwood family, Santa<strong>la</strong>ceae).Whereas other mistleto<strong>es</strong> rely upon birds for the dispersal of their seeds, dwarf mistleto<strong>es</strong> (with the exception ofArceuthobium verticilliflorum) have evolved explosively <strong>de</strong>hisc<strong>en</strong>t fruits. Just as in the squirting cucumber, the pedicelb<strong>en</strong>ds downwards as the fruit matur<strong>es</strong> and a <strong>de</strong>hisc<strong>en</strong>ce line forms around its point of attachm<strong>en</strong>t. Once the fruithas reached maturity, the slight<strong>es</strong>t touch dislodg<strong>es</strong> it from the pedicel. The pr<strong>es</strong>sure that builds up insi<strong>de</strong> the darkgre<strong>en</strong>, single-see<strong>de</strong>d berry is so high that it fir<strong>es</strong> the tiny sticky seed through the hole in the rubbery fruit wall overa distance of up to 16m at the remarkable speed of 2m per second (97km per hour). A single pon<strong>de</strong>rosa pineinfected with Arceuthobium campylopodum may be bombar<strong>de</strong>d by more than two million seeds, which overwinter onthe surface where they <strong>la</strong>nd and ger-minate the following spring. If a seed is lucky <strong>en</strong>ough to <strong>la</strong>nd on a compatiblehost tree, its embryo grows, supported by the photosynthetic <strong>en</strong>dosperm tissue, and <strong>en</strong>ters the host’s bark.Aceuthobium speci<strong>es</strong> can live insi<strong>de</strong> their host for months or ev<strong>en</strong> years before producing a p<strong>la</strong>nt.Whether the seeds are catapulted from passively or actively exploding fruits, they are dispersed over nomore than a few metr<strong>es</strong>. The advantage of ballistic dispersal is that it is cheap, requiring no animal reward andusually very little in terms of specialised structur<strong>es</strong>. The preferred strategy of annual p<strong>la</strong>nts is to disperse theirseeds in a way that allows them to maintain and expand an existing popu<strong>la</strong>tion in the same p<strong>la</strong>ce rather thancolonising new territori<strong>es</strong>. For per<strong>en</strong>nial p<strong>la</strong>nts the most important factor is that the seeds should avoidcompetition with the par<strong>en</strong>t p<strong>la</strong>nt. However, the ejection of the seeds from their fruits is oft<strong>en</strong> only the first phaseof their dispersal history. Many seeds, such as the squirting cucumber, pansi<strong>es</strong>, spurg<strong>es</strong> and gorse, have an <strong>en</strong>ergyrich,oily app<strong>en</strong>dage, which functions as an edible bait that lur<strong>es</strong> ants into carrying them away from the par<strong>en</strong>tp<strong>la</strong>nt. Apart from such app<strong>en</strong>dag<strong>es</strong>, ballistically dispersed seeds are usually small, smooth and more or l<strong>es</strong>sspherical, <strong>en</strong>suring low air r<strong>es</strong>istance.Geocarpy – or how do peanuts <strong>en</strong>d up un<strong>de</strong>rground?This is a legitimate qu<strong>es</strong>tion if one has never se<strong>en</strong> a peanut p<strong>la</strong>nt (Arachis hypogaea, Fabaceae). After all, peanuts arefruits and fruits <strong>de</strong>velop from flowers, and flowers need to be pollinated, which mostly happ<strong>en</strong>s in the air. So howcan a fruit <strong>en</strong>d up un<strong>de</strong>rground? The answer is that the p<strong>la</strong>nt itself buri<strong>es</strong> them. After pollination and fertilization, theovary is pushed down into the ground, h<strong>en</strong>ce the name geocarpy for this type of self-dispersal. The single carpel of thepeanut flower sits at the <strong>en</strong>d of a specialised stalk-like organ called the gynophore. As soon as the ovul<strong>es</strong> are fertilised,the gynophore b<strong>en</strong>ds down and elongat<strong>es</strong> until it has pushed the young, pointed ovary un<strong>de</strong>rground. Once the ovaryhas reached its subterranean <strong>de</strong>stination, it swells to produce peanuts. Other legum<strong>es</strong> that bury their fruits in the sameway are the Bambara groundnut from W<strong>es</strong>t Africa (Vigna subterranea) and Astragalus hypogaeus from w<strong>es</strong>t Siberia.Geocarpy is mostly found in annual p<strong>la</strong>nts living in dry, hot climat<strong>es</strong> such as <strong>de</strong>serts, grass<strong>la</strong>nds and savannahhabitats. Here, burying the fruits is not only a safeguard against grazing animals, it primarily <strong>en</strong>sur<strong>es</strong> that the nextg<strong>en</strong>eration is kept in a suitable location in an inhospitable <strong>en</strong>vironm<strong>en</strong>t.One rare temperate example of a p<strong>la</strong>nt that buri<strong>es</strong> its fruits is the ivy-leafed toadf<strong>la</strong>x (Cymba<strong>la</strong>ria muralis,P<strong>la</strong>ntaginaceae), a common wild flower on walls and rocks in Britain, Europe and North America. Wh<strong>en</strong> inbloom, the flowers face the sun. As soon as they have be<strong>en</strong> pollinated, their pedicels turn away from the sunlightand seek out sha<strong>de</strong> by growing longer and searching the substrate for suitably dark cracks and cranni<strong>es</strong> in whichto <strong>de</strong>posit their fruit. Wh<strong>en</strong> ripe, the capsul<strong>es</strong> op<strong>en</strong> to release a number of small, irregu<strong>la</strong>rly ornam<strong>en</strong>ted seeds.Careful p<strong>la</strong>nting by the mother p<strong>la</strong>nt and the rugged surface of the fruits prev<strong>en</strong>t them from rolling out of theirprotected <strong>en</strong>vironm<strong>en</strong>t.Self-burying drillsSome diaspor<strong>es</strong> bury themselv<strong>es</strong> after dispersal. Apart from keeping them hidd<strong>en</strong> from animal predators, this selfburialis regar<strong>de</strong>d as an adaptation to dry soils as it allows the diaspore to reach the more humid <strong>la</strong>yers just un<strong>de</strong>rthe soil surface. The rotating movem<strong>en</strong>t of their hygroscopic app<strong>en</strong>dag<strong>es</strong>, which twist and untwist with chang<strong>es</strong>in humidity, allows the diaspore to drill itself into the ground. This behaviour is most famously found in speci<strong>es</strong>of stork’s bills, e.g. red-stem stork’s bill (Erodium cicutarium), and musky stork’s bill (E. moschatum). Their diaspor<strong>es</strong>consist of fragm<strong>en</strong>ts (fruitlets) of a schizocarpic fruit with a long beak. Once the fruit has come apart, each singl<strong>es</strong>ee<strong>de</strong>dfruitlet retains a share of the beak, which serv<strong>es</strong> as a hygroscopically moving awn. The same mechanismis found in a number of speci<strong>es</strong> of the very distantly re<strong>la</strong>ted grass family (Poaceae). The florets of wild oat (Av<strong>en</strong>afatua), wild barley (Hor<strong>de</strong>um vulgare ssp. spontaneum) and the needle-and-thread grass (Stipa comata) are equippedwith an awn, the basal portion of which twists and untwists with chang<strong>es</strong> in moisture levels. The drilling floretsof the Australian corkscrew spear-grass (Stipa setacea) are so sharp that they are able to p<strong>en</strong>etrate the wool andskin of sheep. Once embed<strong>de</strong>d in the fl<strong>es</strong>h of the animal, the curved hairs on the surface of the florets make themalmost impossible to dislodge. The muscle movem<strong>en</strong>ts of the tortured animals drag the fruits <strong>de</strong>eper and <strong>de</strong>eperinto their body. Florets of Stipa setacea have allegedly be<strong>en</strong> found in the heart muscle of <strong>de</strong>ad sheep.Creeps and jerksHygroscopically moving app<strong>en</strong>dag<strong>es</strong> <strong>en</strong>able other diaspor<strong>es</strong> to creep along the ground for short distanc<strong>es</strong>. Suchcreeping diaspor<strong>es</strong> are found in some grass<strong>es</strong>, and in the sunflower family (Asteraceae) and teasel family(Dipsacaceae). Their fruits have a hygroscopically moving pappus (modified calyx). For example, the fruits(cypse<strong>la</strong>s) of the cornflower (C<strong>en</strong>taurea cyanus, Asteraceae) are crowned by a tuft of short, stiff, scale-like pappussegm<strong>en</strong>ts, which are too small to p<strong>la</strong>y any role in wind dispersal. Their function is rather differ<strong>en</strong>t: with changinghumidity the pappus scal<strong>es</strong> repeatedly move in and out, thus pushing the fruits a few c<strong>en</strong>timetr<strong>es</strong> over theground. The very short, forward-pointing teeth along their margins prev<strong>en</strong>t movem<strong>en</strong>t in the opposite direction.The distanc<strong>es</strong> they creep are short but at least they move away from the par<strong>en</strong>t p<strong>la</strong>nt; wind and rainwater maycarry them further. A more targeted additional strategy has giv<strong>en</strong> them an edible swelling at the base tospecifically attract ants, a mo<strong>de</strong> of dispersal that will be discussed in <strong>de</strong>tail.Specialised awns <strong>en</strong>able the fruits of certain grass<strong>es</strong> (e.g. Arrh<strong>en</strong>atherum e<strong>la</strong>tius, Av<strong>en</strong>a sterilis) to jerk and jumpfor short distanc<strong>es</strong>. The distal part of their long-kneed awns is straight whereas the lower part is helically twistedand extremely hygroscopic. With changing humidity the basal part winds or unwinds, turning the straight distalpart of the awn. Since each fruit has two awns, which turn in opposite directions, their distal parts ev<strong>en</strong>tuallymeet and become <strong>en</strong>tangled. The t<strong>en</strong>sion that builds up betwe<strong>en</strong> them is released wh<strong>en</strong> the pr<strong>es</strong>sure is strong<strong>en</strong>ough to push the distal parts past each other. Within a split second, the jerky movem<strong>en</strong>t of the awns catapultsthe diaspore into the air.Dispersal by animalsAnimal movem<strong>en</strong>ts are l<strong>es</strong>s haphazard than wind and water making animals much more reliable as dispersa<strong>la</strong>g<strong>en</strong>ts. A p<strong>la</strong>nt that manag<strong>es</strong> to <strong>de</strong>velop a re<strong>la</strong>tionship with animals needs fewer seeds to guarantee the survivalof the speci<strong>es</strong>. The evolution of animal-dispersed diaspor<strong>es</strong> clearly has many advantag<strong>es</strong> and so it is not surprisingthat some fifty per c<strong>en</strong>t of gymnosperms (Ephedra, Gnetum, Ginkgo, a few conifers and all cycads) use animals toassist in the dispersal of their seeds. Once again it was the angiosperms that perfected the shift from abiotic tobiotic dispersal ag<strong>en</strong>ts by evolving a fascinating spectrum of strategi<strong>es</strong> <strong>en</strong>abling them to travel either on or insi<strong>de</strong>animals. Th<strong>es</strong>e close, sometim<strong>es</strong> highly specialised re<strong>la</strong>tionships betwe<strong>en</strong> angiosperms and animal dispersersprovi<strong>de</strong> another key to the un<strong>de</strong>rstanding of the evolutionary succ<strong>es</strong>s of this group.The evid<strong>en</strong>ce of such strategi<strong>es</strong> in p<strong>la</strong>nts li<strong>es</strong> in every sweet, juicy fruit we <strong>en</strong>joy. The sweet pulp of thefruit is a bait to lure pot<strong>en</strong>tial dispersers into swallowing the seeds and dispersing them in their faec<strong>es</strong>. If they arefortunate, the seeds will <strong>la</strong>nd in a suitable p<strong>la</strong>ce to grow well away from the shadow of the mother p<strong>la</strong>nt. Thisform of dispersal is called <strong>en</strong>dozoochory, “dispersal insi<strong>de</strong> an animal”. Endozoochory is a frequ<strong>en</strong>tly <strong>en</strong>counteredph<strong>en</strong>om<strong>en</strong>on in many famili<strong>es</strong> with fl<strong>es</strong>hy fruits, e.g. Ericaceae, Rosaceae, So<strong>la</strong>naceae. Endozoochorouslydispersed seeds do not pr<strong>es</strong><strong>en</strong>t any conspicuous adaptations to facilitate their dispersal. They are usually smooth,globose to ovoid, and either covered in a hard <strong>en</strong>docarp (if borne in drup<strong>es</strong>) or with a hard seed coat (if borneEnglish texts 281
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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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116 Semillas - La vida en cápsulas
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118 Semillas - La vida en cápsulas
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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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La dispersión de frutos y semillas
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Cephalophyllum loreum (Aizoaceae) -
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página anterior: Cerbera manghas (
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página anterior: : habas de mar -
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Frutos explosivos activosLos frutos
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142 Semillas - La vida en cápsulas
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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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Euphorbia peplus (Euphorbiaceae) -
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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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Silene gallica (Caryophyllaceae) -
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