The Evolution and Function of Biological Macrostructures [1st ed. 2019] 978-3-662-59290-8, 978-3-662-59291-5

Table of contents :
Front Matter ....Pages i-xi
Shape, Movement, Lever (Georg Glaeser, Werner Nachtigall)....Pages 0-35
Sticking, Filtering, Drilling (Georg Glaeser, Werner Nachtigall)....Pages 36-61
Gripping, Stretching, Folding (Georg Glaeser, Werner Nachtigall)....Pages 62-81
Signaling, Swimming, Flying, Exploding (Georg Glaeser, Werner Nachtigall)....Pages 82-109
Storage, Constructions,Building materials (Georg Glaeser, Werner Nachtigall)....Pages 110-129
Packaging, Primordia, Unfolding Mechanisms (Georg Glaeser, Werner Nachtigall)....Pages 130-149
Brave New World (Georg Glaeser, Werner Nachtigall)....Pages 150-163
Back Matter ....Pages 164-170

Citation preview

Georg Glaeser · Werner Nachtigall

The Evolution and Function of Biological Macrostructures

The Evolution and Function of Biological Macrostructures

Georg Glaeser • Werner Nachtigall

The Evolution and Function of Biological Macrostructures

Georg Glaeser Department of Geometry University of Applied Arts Vienna Vienna, Austria

Werner Nachtigall Saarland University Saarbrücken, Germany

Translation from the German language edition: Die Evolution biologischer Makrostrukturen - ein Fotoshooting by Georg Glaeser and Werner Nachtigall © Springer-Verlag GmbH Deutschland, ein Teil von Springer Nature 2018. ISBN 978-3-662-59290-8 ISBN 978-3-662-59291-5 (eBook) https://doi.org/10.1007/978-3-662-59291-5 © Springer-Verlag GmbH Germany, part of Springer Nature 2019 This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. The publisher, the authors, and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, express or implied, with respect to the material contained herein or for any errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. Cover photo: Georg Glaeser This Springer imprint is published by the registered company Springer-Verlag GmbH DE, part of Springer Nature. The registered company address is: Heidelberger Platz 3, 14197 Berlin, Germany

Preface

Below the left eye of the hermit crab (Clibanarius misanthropus) , you can see a paddle-like leg joint (see p. 56). The appendage acts as a pump with which water from which oxygen has been extracted is expelled through the gill cavity.

v

Different ways of looking at animals and plants Many people are dazzled by the beauty of nature. Those with an interest in biology will probably take out their binoculars or magnifying glass and have a closer look at what nature has to offer. Among nature lovers, nature photographers may be regarded as a “species” of their own: They not only want to document what they see, but strive to create pictures that hold artistic value while simultaneously capturing the biological complexity of their motifs. The goal is to arrive at a photograph that is both informative and aesthetically pleasing. Looking at nature with “innocent joy” was what brought mathematician Georg Glaeser and biologist Werner Nachtigall together. Both are passionate nature photographers, and they have known each other since their previous collaboration on The Evolution of Flight, the second volume in a series of “biological photoshoots.” The work on this project revealed an abundance of macroscopic, and even microscopic, elements that play a significant evolutionary role – just consider the impact of feather microstructures during a bird’s flight – but not all of these elements could be accounted for in the book’s discussion of flight processes. The macroscopic world – little-known but often highly influential The two authors would like to fill this gap with the present book, which will shed light on the astonishing diversity present on the macro level – a hidden world that only few people know about in great detail but that is just as real as the world of big objects that surrounds us. Comparing different formations of organs A first look will reveal the appearance of an animal or a plant. The biological discipline of comparative morphology is concerned with the description of such appearances. The first book of this series (Glaeser/Paulus: The Evolution of the Eye) illustrates how fascinating such comparisons can be by considering the different formations of visual organs. Speed and acceleration Animals and – to some extent – plants have the ability to move, sometimes at great speed and occasionally with astonishing acceleration. This is where high-speed photography comes in. In the book The Evolution of Flight, Glaeser, Paulus, and Nachtigall already explored the thrill of rapid movement, capturing the motion of animals in flight using telephoto lenses and series-production cameras. Macrostructures through the lens of technical biology In biological sciences, the term ‘macrostructure’ refers to any structure that measures several millimetres or only a few centimetres in size and to which a specific function can be assigned. The close examination of macrostructures requires a magnifying glass and occasionally the low magnification of a microscope. Especially on the surfaces of living organisms we can find a confusing and yet fascinating variety of such structures. We have tried to describe a representative selection of these structures from the point of view of technical biology and broaden the reader’s understanding further through photographs. For this purpose, we used macro and micro cameras.

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vii Several examples discussed in this book have already generated bionic innovations, such as the van der Waals adhesion of gecko setae (p. 38), which has inspired adhesive tape that sticks even under water and on oily surfaces; the alula feathers of birds (p. 101) which served as a model for the high-lift slats of aircrafts; and the torsion buckling of bird of paradise flowers (p. 152), which has been adapted for the folding mechanism of self-regulating façade-shading systems. The design of this book is once again based on the tried-and-tested ‘double-page principle’: Each double-page is usually dedicated to one topic. Photographs and/or sketches are used to illustrate key ideas and are often complemented by explanations of the underlying evolutionary process. A benefit of the double-page principle is that the reader is not required to read the book linearly from cover to cover. One can start reading from a random point and then look up the cross-references provided on a second reading. Bibliographical references are given at the end of the book. We tried to keep the texts short to focus the reader’s attention on the pictures, but also strove for readability. For this reason, we avoided using formulas and an abundance of numerical data, and relied instead on concise and informative texts. At several points in the book, we added drawings that were made in a uniform style. The creator of these drawings is Werner Nachtigall; he used a thick eyebrow pencil and coarse paper. The templates that he created with these utensils were then significantly reduced in size. As you will see further on in this book, the result of this process are sketches with a unique graphic quality. Georg Glaeser, who specializes in wildlife photography, was not the only photographer this time. Some of the photographs in this book were taken from Werner Nachtigall’s vast archive of pictures, which include detailed close-ups of preserved animal and plants. Some images of preserved organisms were captured using electronic microscopes. However, we preferred using pictures of living organisms whenever possible. Acknowledgements We wish to thank Hannes F. Paulus, who co-authored the first two books of the “Photoshoots of Evolution” series. For this book, we could count on his expert knowledge and experience in the classification of species. A number of other people deserve special thanks. We are most grateful to (in alphabetical order and without academic degrees) Daniel Abed-Navandi, Peter Calvache, Gudrun Maxam, Tamara Radak and Eugenie Maria Theuer. Stefanie Wolf from Springer Spektrum has also been tremendously helpful and dedicated, offering support throughout this project. The two authors wish you a delightful reading experience with plenty of aha moments!

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© Springer-Verlag GmbH Germany, part of Springer Nature 2019 G. Glaeser, W. Nachtigall, The Evolution and Function of Biological Macrostructures, https://doi.org/10.1007/978-3-662-59291-5_2



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42

Coupling mechanisms Mating insects Especially in the insect world, we can find a great number of coupling mechanisms with which the male specimens of a species mechanically connect to females for copulation. These mechanisms often act like a system of key and lock, and they also “signal” if a potential mating partner is of the same sex. If the coupling of two animals mechanically fails because they are of similar but not identical species, they will abandon their mating attempt. A highly developed form of this key-and-lock principle can be found in dragonflies. Male dragonflies grasp females by the head or the prothorax (see page 148) with claspers at the tip of the male abdomen (page iv). However, hybridization can still occur among closely related species. Photographs can only provide an incomplete image of the complex ways in which these mechanical devices and safety catches interlock. The picture below gives an example of two mating robber flies.

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44

Predetermined breaking points

Predetermined breaking points in animals are quite common and can be found, for instance, in the legs of harvestmen (page 26) and crabs. The many-legged creature on page 61, measuring no more than 2 cm (the house centipede species Scutigera coleoptrata , colloquially known as “hundred-legged”) sheds its legs like harvestmen, but due to the large number of a centipede’s legs, the loss of a limb is less critical in this case. The biological significance of this mechanism probably lies in its facilitation of the centipede’s escape from predators: The cast-off leg or tail continues to twitch for some time, thus distracting the predator’s attention from its prey. Detached lizard tails Lizard tails contain weakly ossified fracture planes that run in the middle of each vertebra. Through violent, spontaneous muscle contractions, the tail will break off at such planes, especially when you try to lift a lizard by its tail. Lizard tails are known to be capable of regenerating (see picture below), but the regenerated tail is just a cartilaginous support structure that contains no real vertebrae and does not undergo any further cell differentiation. The scales of the regenerated tail look different as well. They are simpler and differently coloured, as the photograph below illustrates. The regenerated tail cannot be detached a second time.

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56

Traps and filters

Food traps of black-fly larvae The larvae of black flies (Simuliidae, Melusinidae) primarily live in the fastrunning water of rivers, where they attach themselves to rocks. They have fanlike appendages hanging from their mouth, which act as a food trap. The foldable mouth fan is extended into the stream, where it catches passing particles, such as detritus and bacteria. In irregular intervals, the larvae will retract their two fans and rake them out with the bristles of their mandibles. During an experiment, such larvae were placed in an artificial stream channel that contained a suspension of coloured bacteria. Hungry larvae would take only half an hour to fill their empty stomachs with red bacteria, regardless of their size. The filtering fans of these flies and the swimming bristles of aquatic beetles (page 94), for instance, are very interesting constructions from the viewpoint of fluid mechanics. When they are expanded, they function as a food trap; in their folded state, on the other hand, they act as a paddle. This physical effect is widely exploited by evolution and can be found both in fresh-water and marine species. Black-fly larvae like to sit on the leaves of plants that hang into the water. Their

Food strainer and food-catching broom

large mouth fans generate considerable flow resistance, which could tear the

Shrimps of the genus Atryopsis (image below) – pictured he-

hairs off, but the larvae have special anchorage structures that prevent such

re is the species Atyopsis spinipes (image below) – inhabit

damage. First they spin a patch of silk on the leaf. Then they anchor themselves

tropical streams with strong currents. Like black-fly larvae,

to this patch by means of a circlet of small hooks on their posterior end that

they use fan-like structures as a sieve to filter edible particles

serves as an anchoring organ.

from the passing water.

Female black flies in the imago stage are notorious for causing wounds that

When the current is too weak or when there are not enough

can become infected and heal slowly. In Finland, for instance, black flies are

particles floating in the water, they will repurpose their food

often found in large swarms, which can make life a living hell for humans and

strainer and use it as a food-catching broom with which they

animals.

scan the bottom for edible particles.

57 The filter system of worms Tube worms – the image above shows a specimen of the particularly beautiful species Spirographis spallanzanii ,– live at the bottom of the sea, inside tubes that they build themselves. The worms possess two crowns of tentacles. One of these crowns is fairly long and carries around 200 to 300 tentacles that can twist into five or six spiral coils. The tube, which measures about a centimetre in diameter, is buried in the mud or attached to a stone; the rest of the worm’s body, which is regularly divided into 100 to 300 segments, is hidden inside the tube. Each tentacle has numerous fine branches covered with moving hairs called cilia. Their movement draws water into the funnel of tentacles. During this process, fine particles that float in the water are picked out according to edibility, moistened with sticky mucus, and transported towards the mouth. Cilia crowns acting as food traps and paddles Effective food traps are made up of bristles that are not arranged too closely. They are supposed to let fluids with fine particles pass but catch larger particles. This function is perfectly fulfilled by the cilia crowns of tube worms. They drive the flow of water in such a way that plankton contained in the water is driven towards the wheel of tentacles, where they are caught by the fine cilia. The cilia crowns are not swung to the sides like a fishing net but are held still and only sway slightly with the current. There is always at least a slight current. Paddles work in a different way. When the cilia are arranged close enough to each other so that nothing can pass between them, they act like a solid surface. When this surface is moved rapidly like a paddle, it can generate thrust. This can be observed with the legs of aquatic beetles that are also covered in swimming hairs (see p. 94). It is up to evolution to realize one mechanism or the other by leaving wide or narrow gaps between the individual bristle-like structures.

58

Earth drillers Ovipositors of female crickets The females of bush crickets typically lay their eggs in loose soil. For this purpose, they use slightly curved, powerful ovipositors that are shaped like a sickle and terminate in very hard, chitinous structures. This ovipositor is drilled into the earth, where it releases an egg. For the next egg, a new hole is usually drilled. There are also species that deposit their eggs inside plants by means of very thin ovipositors. The females of grasshoppers, on the other hand, extend their abdomen to twice its normal length by increasing internal pressure. The modified abdomen is then burrowed into the ground, where the animals lay egg pods. Such hydraulic mechanisms were already discussed on pp. 12–21. In the picture on the left, we can see that the female ovipositor is actually made up of four parts.

59 Sprouting great horsetails as earth drillers The strobilus of horsetails breaks through the soil in a compressed state; the subsequent onset of the growth process then causes the individual segments to expand considerably. The “earth driller” of this great horsetail (Equisetum telmateja)measures about a centimetre in diameter, and, fittingly for its purpose, it is shaped like a cone with a sharp tip. Plant cones or leaves that break through the soil can exert significant pressure. This can cause cobblestones and pavement tiles to be lifted and might even damage thick layers of bitumen. The pressure generated by the plant can be many times greater than, for instance, the air pressure inside the tires of large lorries. As opposed to the “sand shoes” of camelids (page 48), the earth drillers of plants are designed in such a way that the forces are concentrated onto as small an area as possible, creating great pressure.

60

Mobile ovipositors and stings

Giant ichneumon The largest ichneumon wasp in Europe is probably the giant ichneumon (Rhyssa persuasoria). With their ovipositors, which are as long as their bodies, females can grow to measure 7 cm in length. Seen in this way, the giant ichneumon also qualifies as one of the largest insects to be found in our regions. In circumference, it is surpassed by the giant woodwasp (Urocerus gigas). By means of the ovipositor, female wood wasps lay their eggs into wood tunnels, where their larvae develop. With their antennae, giant ichneumons are capable of detecting wood-wasp larvae that are chewing on wood inside. They will drill their ovipositor about a centimetre deep into the wood – a process that can take longer than a quarter of an hour. With great precision, they will pierce a larva and inject an egg into the host. To this end, the female ichneumon will lift its abdomen in a characteristic manner and spread the two sheaths of the ovipositor, which is hard as steel but still extremely thin. The driller itself consists of two parts that drill into the wood by moving against each other by means of specialized muscles. As they are driving forward, they glide against each other along the edges of rebate-like grooves and notches. Clearly developed rebate structures can also be found on the ovipositors of female bush crickets, which are discussed on page 58. The sketch shows parts of the ovipositor of a Palpate cave cricket (Dolichopoda palpata): When the cricket lays eggs, the three double-segmented elements slide against each other along finely developed swallowtail joints.

61 Bee and wasp stings The stings of wasps and bees are modified ovipositors; the poison glands of the Aculeata group probably evolved from a functional conversion of glands found in other hymenopterous insects that secrete a lubricant layer around eggs. (The venom glands of snakes have undergone a similar functional shift: The glands of non-venomous species secrete a special kind of saliva that facilitates the swallowing of prey.) The sting apparatuses of hymenopterans are complex constructions. The mechanical function of their extension and the way in which their sharp digging blades insert themselves into skin by moving rapidly against each other in vibrating motions cannot be described in just a few words. The tips of these stings often have a special form that is characteristic for this order of insects. The sketch shows the tip of a wasp’s sting. The two digging blades are serrated and move along the parallel guiding grooves of a shared guiding structure, the stabilizing rod, against each other. Between the two blades, there is a hollow tube enclosed by rebate-like sliding joints through which the venom is pumped. One of the digging blades is slightly emarginated on the side, allowing the venom to be released. The tips of insect stings are finer than the finest injection cannulas; their diameter can measure less than a hundredth of a millimetre. The stinging hairs of stinging nettles (page 68) and the poisonous bristles of some caterpillars are similarly thin.

© Springer-Verlag GmbH Germany, part of Springer Nature 2019 G. Glaeser, W. Nachtigall, The Evolution and Function of Biological Macrostructures, https://doi.org/10.1007/978-3-662-59291-5_3



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67 Pedicellariae of sea stars Sea urchins and sea stars, which belong to the family of echinoderms, possess peculiar claw appendages called pedicellariae. The pedicellariae of most sea stars are composed of two valves, whereas those of sea urchins are made up of three valves. The former type is shown in the photographs on this page. The above picture was taken through the water layer of a dissecting pan; in the picture below, the pedicellariae are smaller and somewhat hidden. The tiny claw appendages are usually no larger than a millimetre; they may sit on the shell’s surface or be mounted on stalks, and their valves are pressed together through muscle tractions. Sensory organs respond to mechanical stimuli. A dirt particle, for instance, may be passed along from pedicellaria to pedicellaria, until it falls sideways off the surface of the animal’s body. In a reversal of this process, the pedicellariae can move food particles to the mouth bit by bit. Some pedicellariae are provided with poison glands. These toxic claws serve as defence against predators. The multi-purpose appendages of sea urchins and sea stars are modified spines (see page 33). This is yet another example of an evolutionary shift in function. Spines are the more primordial forms.

Injection syringes and cannulas

68 M

Stinging hairs of the stinging nettle

While the cell is calcified at the base, it is stiffe-

The stinging hairs, which are clearly visible to the human eye, consist of

ned with silica at the tip. The hair terminates in a

only a single cell. The lower end of this cell is embedded in a multicellular

fine bulb with a predetermined fracture line (see

sheathing that sits on the surface of the leaf. The hair cell tapers out at the

p. 44), where it breaks off at the slightest contact.

top, where its diameter measures less than one hundredth of a millimetre.

The hair cell thus resembles a modern, albeit extremely thin, injection syringe. The movement of the slightly elastic base increases the pressure of the toxic liquid inside the hair, forcing the liquid out of the hair after the tip breaks off. The liquid contains various chemical compounds, such as formic acid. In tropical regions, you can find nettle species that are much more poisonous than the nettles that grow in Europe. In comparison: the sensory hairs of flies An interesting comparison can be drawn between the stinging hair of nettles and the tiny hairs that grow on the back of the fly that is pictured here as well. These hairs are even finer than nettle hairs. The rows of hairs act as a “rake” that makes it possible to manipulate the airflow around the fly’s body. Some of these hairs are actually innervated and can transmit signals about turbulences to the nervous system.

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72

Barb Systems

Fruits of the common agrimony Agrimonies (Agrimonia eupatoria, pictured below), which are found along the edges of woods, grow barbed fruits. The outer half of the fruit terminates in a grouping of powerful inwardly bent hooks, which collectively form tiny Velcro-like patches of barbs. They easily get caught in the fur of animals or the clothes of humans, and are thus disseminated. When the fruits dry, they detach from the stem upon slightest contact along a preformed fracture line (page 44). Even better known are the barbs of the greater burdock Arctium lappa on which the hook-and-loop technology of the Velcro fastener has been modelled. Cleavers (Galium aparine) are covered in a dense layer of finely hooked barbs, which also extends over the double globe of their fruits. That is why they feel sticky even though they do not secrete any adhesive substance. They climb up other plants, and their hooks keep them from falling off.

73 Dandelion seeds Dandelions (Taraxacum officinale) produce seeds that act like pa-

parachute-like pappus – they carry increasingly large, outward-facing

rachutes. The plant belongs to the family of Asteraceae, which grow

hooks. Once the seeds penetrate into the ground, their barbed ends

many florets that are characteristically clustered into flower heads.

keep them from slipping out. The dissemination mechanism of marsh

Each floret produces a seed. These seeds are arranged radially on

valerians (Valeriana dioica) and goatsbeard flowers (Tragopogon ori-

the receptacle, and towards the top – where they transition into the

entalis, page 98) works in a similar manner.



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77 The “Honey pots” of honeypot ants The “honey pots” of honeypot ants in North America are quite peculiar. The abdomen of some specimens swells from the mass of their honey deposit. These animals ingest excessive amounts of honeydew in their stomachs, called gasters. Gorged with honey, they then hang like shining, yellow barrels from the ceilings of their nest chambers, and thus act as living food storage vessels. “Honey pots” of a different kind can be found in the nests of bumblebees. When the young bumblebees have hatched, they leave behind their vessel-like pupae, which are stuck together and line the walls of the nest. The vessels can be used to store nectar and pollen, thus undergoing a functional change. These structures have evolved to perform two functions, which are not parallel processes, as is the case with the adhesive pads of fly feet, but occur in a sequential order, one after another. Chunky oil beetles Black oil beetles (Meloe proscarabaeus) are remarkable for a number of reasons: After emerging from their pupae, female specimens increase to six times their size during socalled “maturation feeding” and then lay several thousand eggs five to six times during their lifetime (the eggs make up from a third to almost a half of their weight). Male specimens are noticeably smaller (the image on the left page shows a pair of male and female beetles) and are recognizable by their kinked antennae (image on the left). Oil beetles get their name from the yellow defensive fluid that they secrete from their knee joints, which contains the toxic substance cantharidin. The beetles look clumsy: The short elytra only cover the swollen base of the abdomen and gape apart at the tips, thus leaving a large part of the abdomen exposed. The beetles are not capable of flight and lay their eggs on the ground, but their larvae develop in an exclusively parasitic manner, especially in the nests of solitary bees. However, only a small portion of the many thousands of potential offspring eventually survives. Another example of extreme stretching can be found in female mosquitos (see 145). In less than half a minute, these insects are able to multiply their weight while sucking blood.



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© Springer-Verlag GmbH Germany, part of Springer Nature 2019 G. Glaeser, W. Nachtigall, The Evolution and Function of Biological Macrostructures, https://doi.org/10.1007/978-3-662-59291-5_4



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The upper left image shows that surface tension is of vital importance for insects, which must keep their wings dry. Just a few drops of cooking oil on the water surface can have lethal consequences for the insect because its weight can then no longer be carried by the water. The chironomid (the large image shows a male, which is easily recognized by its bushy antennae, see page 144) is no longer capable of launching into flight from the water. The upper right image, which shows a mosquito egg raft floating on the water, illustrates the high reproductive rate of this species. Under good conditions, it takes only a few days for a new mosquito to emerge from each egg.

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94

Propulsion Apparatuses

Swimming legs of the great diving beetle While the front legs of water boatmen (genera Corixa, Sigara) have evolved into spoon-shaped sieve apparatuses (page 51) and the middle legs are used as long cantilevers for clinging onto surfaces, the hindlegs of these insects are flattened and fringed with dense swimming hair. During a backward stroke, they are turned like a rudder with the broadside against the water, and during a forward stroke they are held flat and folded towards the ventral side of the trunk. The swimming strokes follow one another in rapid succession. Between two strokes, the trunk is decelerated by its inherent resistance. This produces a “hopping” motion. The males of the great diving beetle Dytiscus marginalis have a widened first leg segment – also called “heel” – with a complex clinging apparatus that uses the suction cup principle (see page 35). The apparatus consists of one big stalked suction cup, two medium-sized and many small ones. The image on the right-hand side shows the paddling leg of a small relative of the great diving beetle, namely that of the grooved diving beetle Acilius sulcatus . The left hindleg has been clamped inside a water tunnel in such a way that it is seen from the direction of the inflow, that is, during a paddling stroke. Due to the water pressure, the leg segments as well as the swimming hairs on these segments are spread apart. They thus form a wide rudder surface. In the picture, the cold water of the tunnel is sparkled with air bubbles that get caught in the hairs; this is not usually the case. Instead of such circular swimming hair, the whirligig beetle (page 89) has extremely thin, flattened swimming lamellae. During a paddling stroke, the flow pressure causes these lamellae to automatically spread and overlap like playing cards. When the leg is brought forward, on the other hand, the lamellae lie flat on the surface, leaving no gap between them. Like the diving beetle’s swimming hair, the lamellae thus generate high thrust with very little counter-thrust. For this reason, the efficiency of the paddling apparatus is very high, that is, favourable.

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98

Parachutes and Gliders

Parachutes of the meadow salsify and the old man’s beard Like dandelions (page 73), the meadow salsify (Tragopogon pratensis) , pictured below) has evolved stalked parachutes on its fruits. These parachutes consist of a relatively large radial structure. The rods of this umbrella-like structure contain fluffs of fine, white hairs on two sides. The two rows of hair overlap slightly. When the fruit is blown by the wind and then slowly starts to descend, air flows through the parachute from the bottom up. From aerodynamics we know that – according to the so-called Reynolds number dependency – the slower the air flows through a circular hair formation and the smaller the formation is, the better it acts as a resistance generator. The fruit of the meadow salsify is thus equipped with the ideal parachute. The fine hairs growing on the fruit of the old man’s beard (Clematis vitalba) act in a similar manner. The slower the fruit descends due to its parachute, the further it can be carried away by lateral winds, thus improving the plant’s dispersal potential.

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103 Explosive fruit of the squirting cucumber The squirting cucumber (Ecballium elaterium) is indigenous to the Mediterranean region. When its fruits are mature, they measure about 5 cm and hang from bent stems. The inside of the fruit with its content, the seeds, is under considerable turgor pressure. This pressure is generated by special cell layers that are under high elastic strain. A preformed fracture line (see page 44) forms at the transition point between the fruit and its bent stem; at this line, the stem is attached to the fruit like a plug in a wine barrel. At the slightest touch – occasionally even under the mature fruit’s own weight – the plug detaches itself and the fruit falls to the ground. As the internal pressure of the fruit is released, the seeds, which are embedded in slimy pulp, explosively shoot out from the “plug hole” and disperse over several metres. The image captures the moment when the fruit is plucked from the stem. The internal pressure of the squirting cucumber can reach more than five atmospheres. The seeds may thus be ejected up to 10 m away from the plant. Explosive fruits are also found in the American exploding cucumber Cyclanthera explodens, which reaches a dispersal distance of up to 10 m, and in the American sandbox tree Hura crepitans, which belongs to the spurge family. When its pods, which are the size of a tennis ball, explode, the 2-cm long seeds are thrown as far as 14 m from the tree.



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106

Cocoons

Gossamer bells of Agroeca brunnea These enchanting structures, which measure about a centimetre, can sometimes be seen hanging in the grass. In German-speaking countries they are commonly known as “fairy lamps”. They are the – not yet fully formed – egg sacs of the spider (Agroeca brunnea). The spider first spins a ribbon-like hanging structure, which is then expanded to take the shape of a cup and eventually sealed at the

bottom. Several dozen eggs are laid into the tapered section at the top of the sac. The larger part at the bottom remains empty and later serves as a shelter for the young spiders. The artfully spun cocoon is finally coated with a layer of soil (see page 115) and cemented closer to its supporting grass stalk.

107 Gossamer cocoons of ichneumonid pupae

The ichneumonid larvae that emerge from these eggs first feed on

Ichneumonids and chalcidoid wasps are among the smallest insects.

the less vital organs of the caterpillar, such as the fat body, and it is

The species that live parasitically from the caterpillars and pupae of

only towards the end of their larval development that they attack the

Pieris brassicae butterflies measure no more than 3 mm. Cotesia

rest of the host. They then crawl outside by drilling a hole through the

glomerata pierces a young caterpillar once it has hatched from its

caterpillar’s skin and pupate by enveloping themselves in cocoons at-

egg, and deposits eggs via a fine, adjustable ovipositor (see p. 15),

tached to their host animal. The holes in the cocoons pictured below

injecting them directly into the host’s body.

signal that the ichneumonids have hatched. The host animal dies soon and its remains shrink into an unsightly lump of skin.



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© Springer-Verlag GmbH Germany, part of Springer Nature 2019 G. Glaeser, W. Nachtigall, The Evolution and Function of Biological Macrostructures, https://doi.org/10.1007/978-3-662-59291-5_5



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Clay constructions The larva room of the potter wasp The potter wasp (genus Eumenes) and the mason wasp of the collective genus Odynerus , which dig tunnels into loess walls, make up the hymenopteran family (Eumenidae). Eumenes build urn-shaped formations out of clay that measure just about a centimetre. They bring the material to the “construction site” in the form of small, wet pellets, which are shaped into a small hemisphere first and then expanded into an almost complete sphere. A small funnel is positioned at the top, resembling an amphora. Having deposited an egg, the wasp carries a number of paralyzed – living yet immobilized – caterpillars inside and seals the entire construct. The hatching wasp larva feeds on the provisions that have been carried inside previously and pupates in the clay jug. A Eumenes female is able to build a number of such casings. Odynerus species store away the dug-up loess material in the form of a prominent faucet-like appendage in front of the entrance hole, which protects it from rain and at the same time offers an approach ramp. The two pictures in the middle on the right-hand side show an oblong construction made by a large Mediterranean mud dauber (Sceliphron destillatorium) after the wasp’s imagines have detached from their pupal cases and left through exit holes. The individual cell formations are thus clearly visible in the right-most picture showing the inside of the burrow. The egg cocoons of brown liocranid sac spiders The Agroeca’s bell-shaped cocoon (also called “fairy light”) was already introduced on page 106. Once the spider has deposited its eggs, it spins a thread from the cocoon to the ground and retrieves tiny portions of wet soil, which it dabs onto the cocoon without leaving any gaps. The cocoon thus looks like a splash of soil sticking to a plant, looking inconspicuous. In addition, the hardened layer of soil protects the clutch from drying up and also – though not always reliably – against small ichneumons seeking to open up the eggs by drilling a hole into them. Barely a month after oviposition, the young spiders hatch and crawl into the lower, bigger cavity of the cocoon, where they spend their first two weeks in a protected environment. Afterwards, they free themselves by biting a hole into the cocoon. The lower series of pictures shows a finished and camouflaged Agroeca bell on the left and the same construction sliced open on the right.

115

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Bone constructions (1) Whale skeletons The image on the right shows the thorax of a medium-sized whale. A regular-sized human is also pictured here for comparison. Baleen whales reach lengths of up to 30 metres and masses of over 100 tons. They feed on microbes, which they sift from the water by means of baleens, cornificated palatal folds. Of the ten existing species, the blue whale (Balaenoptera musculus) is the largest living animal on earth. Its thorax is a kind of giant frame construction in which the supporting ribs are tightly wound by means of muscles and ligament tensions and coated with additional muscles and a final layer of fat. Bones are not consistently built the same way. Thorax ribs, for instance, contain more cartilaginous parts toward the sternum (middle picture). In contrast to these bone elements, which are probably the most massive ones of all currently existent vertebrates, other animals display extraordinarily delicate yet highly stable bone constructions.

The skulls and pelvises of birds, for example, display highly sturdy and widespread, yet hardly 1/10 mm thick bone elements that are melded into thin septa. These are extremely lightweight and at the same time astoundingly sturdy. They are so delicate that they are translucent when they are prepared as sample slices for microscope slides; for the picture on the right-hand side below, blue light was used. Lightweight construction of a leporine skull Bones are not always made of solid hard substances; between them, there are often cavities, with a kind of grid construction – a sponge-like mass in terms of space – emerging. Stability need not suffer as a result of this lightweight construction. Individual bones in the back of the head of European hares (Lepus europaeus), the adhesive lines of which are easily recognizable in the image, are examples of such constructions. By contrast, the bone structure of its auditory region in the form of a ring-shaped base in which the eardrum was expanded is almost completely made of a hard substance. It is therefore also known as the “petrous bone.” Spongy bone substance is also often to be found in the ends of long bones – a solid bone membrane covers the cavernous system of delicate bone clasps and baulks in that case. We also speak of a “sandwich” construction, which is often used for aircraft wings. Here, a stretch-resistant aluminum skin that keeps the construction in shape provides a coating for a pressure-resistant honeycomb structure made of fine aluminum lamellae. The already mentioned membrane-like configuration of the thinnest bone layers is particularly noteworthy; it can, for example, be found in the lamellose arrangement of the elephant skull (p. 122).

121



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124

Lightweight constructions in the plant world

The construction of a field poppy capsule Technically speaking, the capsule of a field poppy (already presented on p. 104) is a construction made of stiff frames with a bendable membrane. The septa are considerably stiff on the outside, forming the loadbearing frame, while the wall of the capsule as such is very delicate and would not be able to hold the system in place on its own. The fuselage of older airplanes Similar frame-membrane constructions of this type are to be found in technology, for example quite often in the context of older airplane fuselages. The “Fieseler Storch,” for instance, was built as a load-bearing tubular construction coated with a non-bearing finish membrane made of waterproofed fabric. The nacelles of older gliders are constructed in a similar manner. Lightweight construction of the bullrush and the calamus The leaves of the lesser bullrush (Typha angustifolia) are formed as water-tight compartments. Parallel cavities inside them are created by way of thin tissue walls arranged in parallel. Outwardly, the entire construction is covered by a – similarly thin – tissue layer. This ensures a certain pressure resistance, even though the leaf needs only little material to uphold its volume. It consists of more than 90% air, which guarantees good buoyancy in case of flooding. The stiffening septa running through it and from outside, the plena in between are recognizable as dark and light green longitudinal ligaments respectively. There are a number of other stiffening constructions in plant leaves. They are either situated inside the leaf, as in the present case, or on the outside of the leaf, as ribs and septa. The giant floating leaves of Queen Victoria’s water lilies (Victoria regia

tain “stems” that are effectively made of leaf sheaths. The banana

or amazonica), which can be admired in botanical gardens all over

tree (genus Musa) and the calamus (Acorus calamus; cross-section

the world, are an example of the latter case. Here, radial stiffening

pictured on the bottom right) belong to this category. The cavities

ribs, connected by concentrically arranged bridges are situated un-

of the calamus ensure that the marsh plant stays afloat in case of

derneath the leaf, giving the system great stability. Other plants con-

flooding.

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126

Above-ground constructions in the plant world

The development of the round-headed leek’s blossom The structure of the round-headed leek blossom Allium sphaerocephalon is initially enclosed by a fine membranous shroud, which later rips. Next, single flowers grow radially out of it. The circular, hollow stem carries a considerable weight with its blossom at a great height and furthermore needs to be resilient towards vibrations caused by wind. In its outward form and constructional demands, the leek at this stage resembles a television tower with the almost ubiquitous sprawling restaurant at a high altitude. Proportionally to its height and the weight of the load it has to carry, the stem of the leek is, however, much more slender than the television tower and its component parts (rendered in terms of the different scale and constructional demands; see the left picture on the right page). Similar ratios to the leek are to be found in the cornstalk, the tube of which only measures a few millimetres and yet carries ears that weigh multiple times the stalk’s weight. A discrepancy in the length-width-ratio Owing to this discrepancy between technical and botanical lightweight constructions, biology was considered to be superior to technology. This assumption is not quite true. Simple laws of physics dictate that the building must have a certain degree of “plumpness.” The taller it is, the smaller its slenderness ratio must be and the stockier is its appearance. A cornstalk displays a length to diameter ratio of about 500 : 1, the Olympia TV tower in Munich of only 17,6 : 1. If Evolution were able to design the rye stalk to be as big as the TV tower, the slenderness ratio of the original could not be upheld – it would approximate that of the technical construction. This observation is validated when we compare the slenderness ratios of “botanical constructions” of different heights: rye stalk (height 1,5 m) 500 : 1, bamboo (height 30 m) 130 : 1, fir (height 70 m) 42 : 1, giant sequoia (higher than 100 m) 15 : 1. Our preference for natural construction is relativised when we take into account the laws of physics and technology. At the same time, these laws present evolution with insurmountable barriers. The closer it approximates the upper limit, the more sophisticated the construction needs to be. This is quite a common experience in biology – and, by the way, in technology, too. The Barba-Kick law of proportional resistance was already formulated in the 19th century. It states that higher tower-like constructions of diameter d and height h should not become wider according to the linear law “d is proportional to h to the power of 1” but according to the exponential law “d is proportional to h to the power of 1,5”. As a result, they must necessarily become stumpier. When constructing a 50-cm-thick, tower-like construction of 10 m height that is 10 times higher than the original (thus, 100 m high), its diameter must not be 10 times 0,5 m = 5 m but it has to be 10 times 3 = 30 m. When comparing shrubbery or flagpoles to TV towers, one quickly arrives at approximately these values.

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128

Flutter stability and tensile strength

The moulding of seaweed The picture shows sugar kelp (Laminaria saccharina), which is common on the Breton coast. Seaweed is exposed to strong tidal movements and must neither tear nor over-expand. The secret of its solidity lies – seemingly paradoxically – in its pliancy. It has the ability to follow any water movement. In flexible, pliant, elongated formations with wavy, tattered, often almost torn-looking contours, they can also be found in marine areas, in which the strong surf appears to make the survival of bigger plants impossible. The flexibility of several species is attributable to a specific fine structure that is similar to the above-described “sandwich” system (p. 122). A faveolate internal structure is encompassed by a highly viscous, slippery outer membrane. However, no part of the plant is hard throughout – at the most, it can be cartilaginous or gelatinous. All elements are malleable to a certain degree, especially in terms of stretching.

top view and cross-section of seaweed (Durvillaea antarctica)

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© Springer-Verlag GmbH Germany, part of Springer Nature 2019 G. Glaeser, W. Nachtigall, The Evolution and Function of Biological Macrostructures, https://doi.org/10.1007/978-3-662-59291-5_6



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134

Casing of mericarp fruits and seeds

Casing of flowers and fruits

A particularly crowded casing can be observed in the blossoms of the

When mericarps are situated on a central cone and take up a cer-

calamus (Acorus calamus). The plant, which originally stems from

tain geometrically defined volume, as in the present example, they

Asia, has been naturalised in Europe since the 16th century and to-

must be shaped in a certain way – in this case conical and a bit ob-

day, it can occasionally be seen as a cohabitant of reeds and water

late at the points of contact. The evolution of such casing does not

flags. Among these, it stands out with its leaves, which are often wa-

define every point of contact and hence not the form of single ele-

vy at the edge. The blossom is club-shaped and sticks out from the

ments either. Rather, these elements develop in specific ways due

sprout slantwise. It is hermaphroditic; male and female florets are

to spatial competition coupled with individual growth. Once again,

narrowly packed and nested into each other. The female florets de-

the laws of physics influence the final shape of a plant. When the

velop first (“protogyny”), with flies acting as pollinators. The form that

available space is used in an optimal way, the casings take the form

is common in Europe does not develop fruits, as it is sterile owing to

of hexahedral prisms. This is visible, for example, in the single ele-

its threefold (triploid) chromosome set. Its dissemination only hap-

ments of pine cones or in the compound eyes of insects, at least as

pens by way of rootstocks that trail in muddy waters.

far as the outer parts are concerned. The thin-walled cells of plantal

On the face of it, the cylindrical blossom of weeping birches

growth tissue – which would grow in an approximately globose shape

(Betula pendula) appears to be very similar. The catkin hangs ho-

without mutual contact – are multiply nested into each other by way

rizontally from the tree in the final phase of its development. After

of ideal packaging. On average, they form geometrical bodies with

maturity, a great number of approximately winged fruits, arranged

12 to 14 surfaces, which approximate an ideal body (a 14-surface

in narrow spirals on a central axis, can be observed. The “wings”

body with 8 octahedron and 6 cube surfaces has the most advanta-

are formed by the two side flaps of the fruit scales, which are bent

geous relationship between surface and volume). This ideal packa-

backwards, while the middle flap is straight and pointed. The latter

ging was discovered when conducting experiments with foams. This

is pictured on p. 99.

technique often employs tetrahedrons that can be stacked gaplessly.

The picture series below shows the casing of fruitlets – around a co-

Maybe you recall the tetrahedral milk cartons from the 60s?

ne (on the left), around the hollow cone of a wild teasel, Dipsacus fullonum (in the middle), and on a slightly convex receptacle.

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136

Casing (2) When mathematics comes into play When looking at sunflowers, daisies and many other flowers (e. g. types of Echinacea), the human eye recognizes spirals that turn in the left or right direction. The spirals approximate logarithmic spirals and correlate with the Fibonacci numbers. The reason for this layout is always the same: distributing as many seeds as possible on the smallest possible surface area. During the growth process of the plant, the best strategy turns out to be choosing adjacent single petals by twisting them according to the golden ratio, while the distance to the center is exponentially enlarged at the same time. Each plant passes on the “chosen” angle to its offspring. The angle changes due to minor mutations. The plant that has chosen the best angle will be able to produce more offspring. Hence, the optimal angle consistently prevails. Optimal packaging Such packaging in the tightest of spaces is highly interesting to mathematicians. When we assume that “a bit of squishing” is allowed in nature, the best and most exact mathematical solutions are even surpassed by evolution (see also p. 133). The picture at the very bottom of this page shows a computer simulation of the arrangement of individual blossoms in a sunflower; in this case, certain overlaps are tolerated. By turning one of the “little screws” (= parameters), we inevitably arrive at phenomena that nature presents to us on a daily basis. The picture on the next page shows a simulation of the optimal layout of spheres on an ellipsoid, which can – with some imagination – be applied to a daisy.





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Pollination Mechanisms (1) Blossom-forming in primroses Many fine adjustments that often extend to astonishing details in the interaction between blossom and insect belong to the most interesting chapters of biology. The mechanisms that plants develop in order to – anthropomorphically speaking – outwit pollinating insects, are legion. On page 5, we already explained the lever mechanism of the sage blossom, with the help of which bumblebees and bees powder themselves with pollen. When cutting the blossoms of a common primrose (Primula vulgaris) lengthwise, two configurations can be found. In one type of flower, the stamens are arranged on top, with the stigma of the carpel below, in the corolla tube. With the other type, it is just the opposite. When comparing the blossoms, it becomes apparent that the stigma and the stamen lie approximately on the same plane in both brachystylous and macrostylous blossoms. When a pollinating insect inserts its proboscis as far as the basis of the corolla tube (where the nectar flows out), it powders the proboscis with pollen at the basis or in the middle, depending on the type of blossom. If it then lands on a flower of the suitable counterpart, the powdered spot lands precisely on the stigma, leading to successful pollination. Additionally, the macrostylous blossom produces small pollen grains and contains long stigma papillae, which are ideally suited to absorbing the small pollen of the other type of blossom. The coupling of the four features “pistil length”, “pollen size”, “stigma papilla length” and “position of the anthers” ensures that a stigma always receives pollen from a different type of blossom, thus avoiding in-breeding.

141

Pollination Mechanisms (2)

142 The snap mechanism of papilionaceae

The blossoms of many members of the papilionaceae family have evolved in such a way that their sturdily developed floral petal stands upright, like a flag, two “wings” protruding on the sides, with the two lower petals partly melded into “carinas.” The carina contains the central carpel. Often, nine or all ten stamens are connate with their filaments, while the carpels remain clear. When an insect lands on the blossom, it is powdered with pollen from below. Papilionaceae have different mechanisms

for

releasing

pollen.

Some look like a noodlesqueezer, pressing out the pollen mass from the adenomatously opened carina in cordlike form (bloomfell, Lotus corniculatus). In others, the pistil contains a fine brush with which it brushes out the pollen (pea Pisum sativum; winterpea, genus Lathyrus). In the case of the common vetch (Vicia sativa) red clover (Trifolium pratense) or the sainfoin (genus Onobrychis ) ), the carina is abruptly folded out from the wings once a certain load is reached, letting the anthers protrude upwards (snap mechanism). In American Papilionaceae, “explosion mechanisms” can be observed (see pp. 102, 103)); these appear to be even stronger than those found in the European gorse or common broom, which will be discussed at a later point in this chapter.

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Antennae

Staggering diversity Insect antennae are extremely polymorphous; however, it is not always possible to deduce a special function from an individual shape. Strongly combed antennae, as those of the Viennese emperor moth, are ideal molecule detectors; they can filter pheromone molecules from an airstream. The antennae of male dorbeetles (see above picture on opposite page), which can be unfolded like a stack of cards and are collapsible, are also used for surface area enlargement. They are spread and set against the airstream during search flights. The unfolding mechanism for the antenna lamellae is hydraulically con-

trolled via a kind of pump system at the base of the antenna. Chemo- and mechanosensors Most antennae, those of honey bees being a case in point, are studded with countless chemo- and mechanosensors, with the chemosensors reacting to odorant molecules from the air or taste molecules from a solution. The sensory organs in and near the second antenna limb, which react to oscillations of the antenna as a whole or to oscillations of the limbs relatively to each other, were already discussed on p. 144.



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© Springer-Verlag GmbH Germany, part of Springer Nature 2019 G. Glaeser, W. Nachtigall, The Evolution and Function of Biological Macrostructures, https://doi.org/10.1007/978-3-662-59291-5_7



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The anteater’s claw The middle finger of the giant anteater (Myrmecophaga tridactyla) bears a sturdy, curved claw with which it lacerates rock-hard termite nests, the occupants of which it enjoys feasting on. In order to succeed in this endeavor, the anteater first folds in the claw, reducing the angle between its axis and the forearm by 90°. It then tucks up its feet while propping up the other one for support. A model for the optimization of tools The anteater’s claw as well as its leg movement and the propping up have found their technical equivalent in hydraulic diggers that dig up roads. In practice, specific recurring (and therefore expensive) fractures occurred on the ripper tooth. In 2012, Sontheim et al. reported how these could be avoided by using nature as a model. The researchers found that the strain on a ripper tooth’s outer contour was particularly strong. The differently curved anteater’s claw only displayed half the strain, the progression of which was much more homogenous. The bionic optimizing process eventually led to a digger claw with the maximum strain on the outer contour reduced by 58%. Below: The armadillo, which is related to the anteater (see p. 9) needs its claws for digging caves in hard soil and when hunting for insect larvae.

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© Springer-Verlag GmbH Germany, part of Springer Nature 2019 G. Glaeser, W. Nachtigall, The Evolution and Function of Biological Macrostructures, https://doi.org/10.1007/978-3-662-59291-5

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168

leafcutter bee, 52 leek, 126 lens eye, 17, 18, 26 leporine skull, 121 Lepus europaeus, 121 lesser bullrush, 124 Leucochloridium paradoxum, 85 lever, 4, 5, 43, 78, 141 light organ, 84 lightweight construction, 27, 33, 121, 122, 124, 125 Liliaceae, 132 Limulus polyphemus, 49 liver fluke, 85 lizard, 44, 78, 160 locust, 66 long bones, 123 looper, 13 Lotus corniculatus, 142 lotus effect, 161 Loxodonta africana, 162 Lucerne grass, 143 Lymnaea stagnalis, 88 maggot, 12, 13 mandibles, 0, 35, 66 mantis, 133 Mantis religiosa, 66 maple, 99 marsh valerian, 73 mason bee, 52 mating, 0, 14, 42, 43, 133 mayfly, 0, 90 meadow horsetail, 23 meadow salsify, 80, 98 meadow spittlebug, 117 mechanical shrimp, 4 mechanosensor, 146 Medicago sativa, 143 Megachilidae, 52 Meloe proscarabaeus, 76, 77 Melolontha melolontha, 81 Melusinidae, 56 membrane construction, 78, 79 mericarp, 135 mericarp fruits, 134 mericarp seeds, 134 Merops apiaster, 100 Mesembryanthemum, 114 micro ornamentation, 160 millipedes, 6 Mobula birostris, 154 mole, 46 mole cricket, 46 mollusc, 25 monitor lizard, 29 morphological structure, 2, 120 mosquito, 86, 144, 145 moss, 114 mouth parts, 64, 120 Musa, 124 Myrmecophaga tridactyla, 153 Myrmeleon formicarius, 69 Nelumbo nucifera, 164 Nepenthes, 41 Notodonta torva, 12 Notonectidae, 51 octopus, 156 Odynerus, 115 Oedipoda germanica, 81 oil beetle, 77 ommatidia, 0, 18 Onobrychis, 142 optimal packaging, 136 orb spider, 40 organpipe corals, 120 oriental garden lizard, 78

Osmia, 52 ovipositor, 15, 58–61, 107 ovules, 132 packaging, 132, 133 paddle fin, 96 paddle leg, 95, 97 Palaemon elegans, 4 Panorpa communis, 15 Panurgus, 52, 53 Papaver rhoeas, 104, 138 papilionaceae, 142 Paracentrotus lividus, 33 parachute, 73, 98 parrot, 28 passive growth, 22 pea, 142 pectoral fin, 154 pedicel, 34 pedicellariae, 67 penguin, 91 Pentatomidae, 118 pericarp, 132 Phallus impudicus, 140 Philaenus spumarius, 117 Pholcus phalangioides, 133 pigeon, 101 pill bug, 20 pincers, 66 pinniped, 91 Pisum sativum, 142 pitcher plant, 41 placoid scales, 92 Planorbarius corneus, 88 Platycnemis pennipes, 148 pleustons, 86 poinsettia, 45 pollen, 5, 52, 53, 77, 112, 141, 142 pollen baskets, 53 pollen-carrying apparatus, 52 pollen-collecting apparatus, 53 pollen-gathering, 52, 53 pollination, 5, 112, 141, 142, 152 pond snail, 88 Portunus, 97 potter wasp, 115 predetermined breaking point, 26, 44, 45, 72 primordia, 138–140 primrose, 141 Primula vulgaris, 141 Priodontes maximus, 9 Prionace glauca, 92 proboscis, 5, 7, 20, 51 propeller, 97 propulsion, 94–97 protective shell, 116, 117 Psychidae, 116 pulvilli, 39 pump system, 146 quinone tanning, 119 raptorial leg, 66 ray, 92, 154, 156 red poppy, 138 red-footed booby, 100 red-winged grasshopper, 81 Rhyssa persuasoria, 60 robber fly, 42 rockpool prawn, 4 rosette, 135 rotang palms, 129 round-headed leek, 126 rove beetle, 80 rush, 125 sage, 5, 141 sainfoin, 142

Left: The wasp spider or jumping spider Salticus scenicus; see also p. 18) has killed a dance fly that is measures about two millimetres (Empididae, see p. 145)



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