Cover

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Title Page, Copyright Page

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pp. i-iv

Contents

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pp. v-vi

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Preface

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pp. vii-viii

...generations, its last edition appearing in 1979 but still structured around the 1966 original. But twenty years on I became frustrated that there had been no single volume since that date which covered the field. This is particularly extraordinary given that the subject itself has expanded and blossomed...

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Acknowledgments

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pp. ix-x

...It is a pleasure to begin by thanking the many generations of students and postgraduates who have let me know when I taught them in ways that inspired their interest and when I merely confused them. From them, I realized where the real fascinations of pollination lay and where my own interests were...

Part I: Essentials of Flower Design and Function

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Chapter 1 Why Pollination Is Interesting

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pp. 3-10

...The flowering plants (angiosperms) account for about one in six of all described species on earth and provide the most obvious visual feature of life on this planet. In the terrestrial environment, their interactions with other living organisms are dominant factors in community structure and function; they underpin all nutrient and energy cycles by...

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Chapter 2 Floral Design and Function

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pp. 11-54

...Flowers are essentially the containers for a plant’s sex organs, but they must perform several interrelated functions. Most obviously, and taking center stage here, they make and mature the gametes and then dispense the male gametes in such a way that they will be transported to another appropriate...

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Chapter 3 Pollination, Mating, and Reproduction in Plants

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pp. 55-87

...Reproduction in plants, as in most organisms, can be either sexual or asexual, but the generation of new variants (which is the underlying necessity for adaptation to new or changing conditions and for evolutionary change) requires that at some point in the life cycle sexual reproduction occurs. Diploid...

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Chapter 4 Evolution of Flowers, Pollination, and Plant Diversity

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pp. 88-102

...Animals of various kinds must have been interacting with green plants—both for shelter and to find food— almost as soon as land plants evolved. Any foliage would provide some shelter, both in terms of alleviating the microclimatic conditions and by offering some possibility of hiding from...

Part II: Floral Advertisements and Floral Rewards

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Chapter 5 Advertisements 1: Visual Signals and Floral Color

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pp. 105-133

...Visual attraction by flowers is substantially related to flower shape and size, and the basic aspects of these were covered in chapter 2. But above all, for most visitors, color and color patterns are attractive. Trichromatic color vision occurs in many terrestrial animals, and in the evolution of insects it...

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Chapter 6 Advertisements 2: Olfactory Signals

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pp. 134-153

...The previous chapter concentrated on visual signaling by flowers, but it is usually not appropriate to look only at flower color and shape as attractants, given the abilities of most flower visitors, and insects in particular, to detect and respond to scents or odors as well; the latter are often the major component...

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Chapter 7 Rewards 1: The Biology of Pollen

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pp. 154-189

...A pollen grain contains the male gamete of the angiosperm plant and is thus the equivalent of a spore in many other plants. In essence, the structure of a pollen grain is adapted to protect and nourish the male gamete during its maturation and subsequent transit between plants, and then to ensure...

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Chapter 8 Rewards 2: The Biology of Nectar

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pp. 190-220

...Nectar is the main secondary floral reward in an evolutionary sense, appearing on the scene probably in the late Cretaceous in angiosperms, later than pollen. But it has very often become the primary offering of a flower (chapter 4), thereby protecting the plant’s investment in the reproductively...

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Chapter 9 Other Floral Rewards

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pp. 221-233

...Occasionally flowers offer neither pollen nor nectar as a foodstuff to their visitors but instead yield other rewards; or they may offer these as “extras” in addition to some pollen. This chapter reviews these possibilities, considering a range of oils, waxes, scents, and resins (on which topics Simpson and Neff [1981, 1983] provided earlier reviews...

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Chapter 10 Rewards and Costs: The Environmental Economics of Pollination

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pp. 234-258

...Pollination is usually thought of as a mutualism, of benefit to both partners, each of which gains in fitness. In such a relationship, both should be trying to maximize their survival and ultimately their reproductive success, which will require balancing their costs against the rewards and hence assessing the net benefits gained. Disentangling the economic...

Part III: Pollination Syndromes?

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Chapter 11 Types of Flower Visitors: Syndromes, Constancy, and Effectiveness

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pp. 261-287

...In the next few chapters, different kinds of flower visitor are reviewed in some depth, using the literature accumulated for over a century documenting their flower visits and floral selection, their color and scent preferences, their food and energy requirements, and aspects of their behavior on flowers...

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Chapter 12 Generalist Flowers and Generalist Visitors

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pp. 288-303

...Many insects that have other core diets (especially entomophagous species, and a range of herbivores) will top up on some floral nectar at times for an easy energy boost, and thus become potential occasional generalist pollinators. However, alongside these there are some rather more regular flower visitors, spending some part of most of their adult...

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Chapter 13 Pollination by Flies

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pp. 304-321

...The flies (order Diptera) constitute a very diverse group of insects, all characterized by just one pair of wings, the ancestral rear pair being modified as flightand balance-control organs termed halteres. Hence flies are often very agile fliers, able to take off and land in any direction and often...

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Chapter 14 Pollination by Butterflies and Moths

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pp. 322-336

...The order Lepidoptera contains the butterflies and moths and represents around 10%–11% of all described insect species. The types that are relevant here are mostly rather large insects (although there are also many thousands of species of small micromoths), but they are usually not particularly...

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Chapter 15 Pollination by Birds

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pp. 337-355

...phenomenon, underappreciated in early literature as it is absent in Europe; but it is familiar through much of the United States and as far north as Alaska and occurs throughout the tropics, in the eastern Mediterranean and Middle East (although not in the more northern...

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Chapter 16 Pollination by Bats

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pp. 356-369

...Flowers visited and pollinated by bats constitute the syndrome termed chiropterophily, as bats were traditionally united in the mammalian order Chiroptera although they are now classified as two distinct and separately evolved orders, Megachiroptera and Microchiroptera. Bats are primarily nocturnal, and as flying endothermic mammals have...

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Chapter 17 Pollination by Nonflying Vertebrates and Other Oddities

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pp. 370-377

...Birds and bats are well known as pollinators, and each has merited its own chapter, but there are also some rather infrequent instances of pollination by other vertebrates, lacking any kind of true flight but able to access flowers either by climbing and gliding among trees or by seeking pendant flowers...

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Chapter 18 Pollination by Bees

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pp. 378-417

...Bees are very special as flower visitors, because almost uniquely they use both nectar and pollen as foods and rely totally on them for both adult and larval nutrition. Adults eat nectar and usually some pollen as well; larvae eat large quantities of both pollen and nectar (converted into honey). Thus any one bee is collecting not just for her own needs but...

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Chapter 19 Wind and Water: Abiotic Pollination

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pp. 418-433

...Abiotic pollination involves the transmission and capture of pollen through a fluid medium, either air or water, and it occurs in at least 60 angiosperm families. Although it was once thought to be a somewhat random process, it is now clear that abiotic pollination is quite sophisticated, with significant...

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Chapter 20 Syndromes and Webs: Specialists and Generalists

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pp. 434-480

...In recent years, many pollination biologists have embarked upon a reassessment of the classical approach to their subject, as established above all by Sprengel, Darwin, Vogel, and van der Pijl. For about 150 years the main emphasis was on showing the adaptive value of floral traits in relation to particular...

Part IV: Floral Ecology

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Chapter 21 The Timing and Patterning of Flowering

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pp. 483-502

...Plants should flower in ways that maximize their own reproductive success. The “flowering pattern” is a composite of the timing and frequency of individual flowers opening, and also of flower longevity. These phenological factors vary between species but also within a species (and often between sexes for dioecious species). They may additionally...

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Chapter 22 Living with Other Flowers: Competition and Pollination Ecology

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pp. 503-523

...Plants rarely occur in isolation, but grow and flower as part of a community of mixed species; and they are rarely visited by just one kind of animal, but receive visits from several potential pollinator types, some of which will be shared with other plants. Thus plantpollinator interactions have a strong...

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Chapter 23 Cheating by Flowers: Cheating the Visitors and Cheating Other Flowers

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pp. 524-541

...Since pollination is not an altruistic exercise, and there is a conflict of needs, both plants and pollinators are liable to cheat to their own benefit, and deception is very common in pollination biology (reviewed by Wiens 1978; Little 1983; Dafni 1984; Renner 2006). For a plant, this essentially means getting pollinated and hence fertilized without...

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Chapter 24 Flower Visitors as Cheats and the Plants’ Responses

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pp. 542-553

...In the last chapter we saw many examples of plants cheating in the plant-pollinator interaction. However, the reverse can also be true—it should at least sometimes pay visitors to gather food from a flower yet resist being manipulated into carrying pollen around, by avoiding the anthers or indeed by...

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Chapter 25 The Interactions of Pollination and Herbivory

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pp. 554-564

...Herbivores in the broadest sense include not just folivores whose diet is green leaves, but also browsers on twigs and bark, seed predators, underground root feeders, florivores, and even nectar robbers. Herbivory is not just due to animals, though: the effects of fungal spores are often very evident on flowers and flowering patterns, and other decomposers...

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Chapter 26 Pollination Using Florivores: From Brood Site Mutualism to Active Pollination

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pp. 565-574

...In chapter 23 various one-sided brood site mimicries were described where the plant benefits from cheating its visitor, whose progeny usually die because their mother has been deceived into laying on a floral tissue that was mimicking the normal egg-laying site. This chapter deals instead with...

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Chapter 27 Pollination in Different Habitats

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pp. 575-604

...Thus far pollination has been dealt with in a collective sense, but it will have been apparent that examples from different habitats have often given rather different impressions of the complexity and level of specialization involved. This chapter therefore dissects the issues that are to the fore in...

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Chapter 28 The Pollination of Crops

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pp. 605-619

...The importance of bee pollination to crops has been well known for at least two millennia, and several ancient civilizations cultivated honeybees or stingless bees in wooden or pottery hives. Likewise, humans have attempted to improve crop productivity for millennia, first by selecting the plants with the more desirable traits but latterly using plant...

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Chapter 29 The Global Pollination Crisis

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pp. 620-638

...For nearly three decades now there have been wellsubstantiated reports of declines in pollinators worldwide, and the problems were explicitly recognized in the UN Sao Paulo declaration (1998–99), so that pollination disruption is at last being emphasized as a major issue (Kearns et al. 1998)...

Appendix

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pp. 639-642

Glossary

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pp. 643-662

References

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pp. 663-750

Subject Index

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pp. 751-767

Index of Animal Genera

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pp. 768-770

Index of Plant Genera

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pp. 771-778