Preface to the Princeton Landmarks in Biology Edition vii Preface xi Symbols Used xiii 1. The Importance of Islands 3 2. Area and Number of Speicies 8 3. Further Explanations of the Area-Diversity Pattern 19 4. The Strategy of Colonization 68 5. Invasibility and the Variable Niche 94 6. Stepping Stones and Biotic Exchange 123 7. Evolutionary Changes Following Colonization 145 8. Prospect 181 Glossary 185 References 193 Index 201

The Theory of Island Biogeography

Herbivorous insects use diverse feeding strategies to obtain nutrients from their host plants. Rather than acting as passive victims in these interactions, plants respond to herbivory with the production of toxins and defensive proteins that target physiological processes in the insect. Herbivore-challenged plants also emit volatiles that attract insect predators and bolster resistance to future threats. This highly dynamic form of immunity is initiated by the recognition of insect oral secretions and signals from injured plant cells. These initial cues are transmitted within the plant by signal transduction pathways that include calcium ion fluxes, phosphorylation cascades, and, in particular, the jasmonate pathway, which plays a central and conserved role in promoting resistance to a broad spectrum of insects. A detailed understanding of plant immunity to arthropod herbivores will provide new insights into basic mechanisms of chemical communication and plant-animal coevolution and may also facilitate new approaches to crop protection and improvement.

Plant Immunity to Insect Herbivores

50 Years of Image Analysis

As the largest class of natural products, terpenes have a variety of roles in mediating antagonistic and beneficial interactions among organisms. They defend many species of plants, animals and microorganisms against predators, pathogens and competitors, and they are involved in conveying messages to conspecifics and mutualists regarding the presence of food, mates and enemies. Despite the diversity of terpenes known, it is striking how phylogenetically distant organisms have come to use similar structures for common purposes. New natural roles undoubtedly remain to be discovered for this large class of compounds, given that such a small percentage of terpenes has been investigated so far.

The function of terpene natural products in the natural world.

What is ecological speciation

Experience and memory of environmental stimuli that indicate future stress can prepare (prime) organismic stress responses even in species lacking a nervous system The process through which such organisms prepare their phenotype for an improved response to future stress has been termed 'priming' However, other terms are also used for this phenomenon, especially when considering priming in different types of organisms and when referring to different stressors Here we propose a conceptual framework for priming of stress responses in bacteria, fungi and plants which allows comparison of priming with other terms, eg adaptation, acclimation, induction, acquired resistance and cross protection We address spatial and temporal aspects of priming and highlight current knowledge about the mechanisms necessary for information storage which range from epigenetic marks to the accumulation of (dormant) signalling molecules Furthermore, we outline possible patterns of primed stress responses Finally, we link the ability of organisms to become primed for stress responses (their 'primability') with evolutionary ecology aspects and discuss which properties of an organism and its environment may favour the evolution of priming of stress responses

Priming and memory of stress responses in organisms lacking a nervous system.

Plants are able to "notice" insect egg deposition and to respond by activating direct and indirect defenses. An overview of these defenses and the underlying mechanisms is given from a tritrophic perspective. First, the interface between plant and eggs is addressed with respect to the mode of attachment of eggs on the plant surface. It is elucidated which plant cells might respond to components from insect eggs or the egg deposition. The scarce knowledge on the elicitors associated with the eggs or the egg-laying female is outlined. Since endosymbiotic microorganisms are often present on the eggs, and microorganisms are also abundant on the leaf surface, the role of these hidden players for eliciting oviposition-induced plant responses is considered. Furthermore, the question of which physiological and molecular processes are induced within the plant in response to egg deposition is addressed. Second, studies on the response of the herbivorous insect to oviposition-induced plant defenses are outlined. Third, the importance of oviposition-induced plant volatiles and contact cues for host and prey location of parasitoids and predators is discussed in the context of other informative chemicals used by carnivores when searching for food. Finally, physiological and ecological costs of oviposition-induced plant responses are addressed.

Early herbivore alert: insect eggs induce plant defense.

The insects that cause or transmit human and animal disease constitute a wide range of taxa sharing a similar feeding habit, but differing in most other aspects of their life histories, including oviposition. This chapter mainly deals with oviposition or larviposition behaviour within the Diptera, since the major groups that exhibit such behaviour and that constitute pests of major medical or veterinary importance are true flies. Aggregation behaviour in bloodsucking Hemiptera is discussed briefly. Bloodfeeding groups generally oviposit at sites remote from their vertebrate hosts: mosquitoes (Culicidae) and blackflies (Simuliidae) oviposit in aquatic environments, and sandflies (Psychodidae), tsetse flies (Glossinidae), bedbugs (Cimicidae) and kissing bugs (Reduviidae) oviposit in terrestrial environments. Facultative or obligatory carnivorous Diptera, where the immature stages can be saprotrophic or parasitic on the same animals on which the adult flies feed, also locate and select oviposition or larviposition sites using chemical cues. These include warble flies, bot flies (Oestridae), blowflies, screwworms (Calliphoridae) and flesh flies (Sarcophagidae). Not surprisingly, given the range of life histories of these diverse groups, the interaction of pheromones with visual cues, odours of environmental origin and the physical properties of the oviposition site, is wide-ranging in nature. Knowledge of the chemoecology of oviposition in these insects lags behind that on herbivorous and parasitic insects and attractant compounds have been identified in only a few cases. A number of plant species have evolved to mimic animal odour cues to attract carnivorous fly species as pollinators. Because of the importance of these insects as vectors of human and animal disease, interest in this field has been driven primarily by a search for chemically baited traps for monitoring and control. Odour baited oviposition traps are currently used in the surveillance of a number of important pest species. Odour baited trapping for the control of pest populations may be feasible for certain species. 5.

Chemoecology of insect eggs and egg deposition

Female parasitic wasps seek hosts for their offspring often in a dynamic environment. Foraging egg parasitoids rely on a variety of chemical cues originating from the adult host, host products, or the host plant rather than from the attacked host stage?the insect egg itself. Besides pupae, insect eggs are the most inconspicuous host stage attacked by parasitic wasps. To overcome the problem of low detectability of host eggs, egg parasitoids have evolved several strategies such as exploiting long-range kairomones of the adult hosts, for example, host aggregation and sex pheromones, plant synomones induced by egg deposition or host feeding, or short-range contact cues derived from the adult host or the host plant. Moreover, egg parasitoids have evolved the ability to use chemical espionage in combination with hitchhiking on the adult host (phoresy) to compensate their limited flight capability and to gain access to freshly laid host eggs. Here, we provide a comprehensive overview on the variety of host-foraging strategies of egg parasitoids exploiting chemical signals. Furthermore, the use of such infochemicals is discussed with respect to the wasps? dietary breadth and their ability to learn

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Foraging behavior of egg parasitoids exploiting chemical information

Plants can respond to insect egg deposition and thus resist attack by herbivorous insects from the beginning of the attack, egg deposition. We review ecological effects of plant responses to insect eggs and differentiate between egg-induced plant defenses that directly harm the eggs and indirect defenses that involve egg parasitoids. Furthermore, we discuss the ability of plants to take insect eggs as warning signals; the eggs indicate future larval feeding damage and trigger plant changes that either directly impair larval performance or attract enemies of the larvae. We address the questions of how egg-associated cues elicit plant defenses, how the information that eggs have been laid is transmitted within a plant, and which molecular and chemical plant responses are induced by egg deposition. Finally, we highlight evolutionary aspects of the interactions between plants and insect eggs and ask how the herbivorous insect copes with egg-induced plant defenses and may avoid them by counteradaptations.

Monika Hilker

Papers

Priming and memory of stress responses in organisms lacking a nervous system.

Early herbivore alert: insect eggs induce plant defense.

Chemoecology of insect eggs and egg deposition

Foraging behavior of egg parasitoids exploiting chemical information

Plant responses to insect egg deposition.