Genetics and analysis of quantitative traits

Beginning with Emlen (1966) and MacArthur and Pianka (1966) and extending through the last ten years, several authors have sought to predict the foraging behavior of animals by means of mathematical models. These models are very similar,in that they all assume that the fitness of a foraging animal is a function of the efficiency of foraging measured in terms of some "currency" (Schoener, 1971) -usually energy- and that natural selection has resulted in animals that forage so as to maximize this fitness. As a result of these similarities, the models have become known as "optimal foraging models"; and the theory that embodies them, "optimal foraging theory." The situations to which optimal foraging theory has been applied, with the exception of a few recent studies, can be divided into the following four categories: (1) choice by an animal of which food types to eat (i.e., optimal diet); (2) choice of which patch type to feed in (i.e., optimal patch choice); (3) optimal allocation of time to different patches; and (4) optimal patterns and speed of movements. In this review we discuss each of these categories separately, dealing with both the theoretical developments and the data that permit tests of the predictions. The review is selective in the sense that we emphasize studies that either develop testable predictions or that attempt to test predictions in a precise quantitative manner. We also discuss what we see to be some of the future developments in the area of optimal foraging theory and how this theory can be related to other areas of biology. Our general conclusion is that the simple models so far formulated are supported are supported reasonably well by available data and that we are optimistic about the value both now and in the future of optimal foraging theory. We argue, however, that these simple models will requre much modification, espicially to deal with situations that either cannot easily be put into one or another of the above four categories or entail currencies more complicated that just energy.

Citation classic - optimal foraging - a selective review of theory and tests

Most empirical and theoretical studies of resource use and population dynamics treat conspecific individuals as ecologically equivalent. This simplification is only justified if interindividual niche variation is rare, weak, or has a trivial effect on ecological processes. This article reviews the incidence, degree, causes, and implications of individual-level niche variation to challenge these simplifications. Evidence for individual specialization is available for 93 species dis- tributed across a broad range of taxonomic groups. Although few studies have quantified the degree to which individuals are specialized relative to their population, between-individual variation can some- times comprise the majority of the population's niche width. The degree of individual specialization varies widely among species and among populations, reflecting a diverse array of physiological, be- havioral, and ecological mechanisms that can generate intrapopu- lation variation. Finally, individual specialization has potentially im- portant ecological, evolutionary, and conservation implications. Theory suggests that niche variation facilitates frequency-dependent interactions that can profoundly affect the population's stability, the amount of intraspecific competition, fitness-function shapes, and the population's capacity to diversify and speciate rapidly. Our collection of case studies suggests that individual specialization is a widespread but underappreciated phenomenon that poses many important but unanswered questions.

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The Ecology of Individuals: Incidence and Implications of Individual Specialization

The evolution of "niche breadth," or "niche width," was a more popular topic in the evolutionary ecological literature of the 1960s and 1970s than it has been recently (109, 118, 120, 134, 155, 156). This review summarizes current hypotheses on the evolution of specialization and generalization and suggests areas in which future research might be rewarding. The topic is so broad that every area of biology bears on it. We cannot hope to offer an exhaustive review of evidence and in particular have slighted much of the ecological literature to emphasize genetic and evolutionary perspectives. We limit our discussion almost entirely to animals. We adopt Hutchinson's (86) representation of a population's ecological niche as an n-dimensional hypervolume, the axes of which are environmental variables or resources. Along each of these, the population displays a wide or narrow tolerance or pattern of utilization, relative to other populations or species. Specialization and generalization must be defined with reference to particular axes (e.g. temperature, range of food particle sizes). Brown (9) suggests that niche breadth along different axes is positively correlated and that this explains positive correlations across species between local abundance and breadth of geographic range. Multidimensional specialization might be expected if species arise in localized regions that differ in several ecological respects from those occupied by parent species. Cody (20), however, suggested that the breadth of habitat is negatively correlated with diet breadth among certain bird species. In practice, quantitative measurement of niche breadth can be difficult (22,

The Evolution of Ecological Specialization

Host plant quality is a key determinant of the fecundity of herbivorous insects. Components of host plant quality (such as carbon, nitrogen, and defensive metabolites) directly affect potential and achieved herbivore fecundity. The responses of insect herbivores to changes in host plant quality vary within and between feeding guilds. Host plant quality also affects insect reproductive strategies: Egg size and quality, the allocation of resources to eggs, and the choice of oviposition sites may all be influenced by plant quality, as may egg or embryo resorption on poor-quality hosts. Many insect herbivores change the quality of their host plants, affecting both inter- and intraspecific interactions. Higher-trophic level interactions, such as the performance of predators and parasitoids, may also be affected by host plant quality. We conclude that host plant quality affects the fecundity of herbivorous insects at both the individual and the population scale.

Host plant quality and fecundity in herbivorous insects

The basic building blocks of communication are signals, assembled in various sequences and combinations, and used in virtually all inter- and intra-specific interactions. While signal evolution has long been a focus of study, there has been a recent resurgence of interest and research in the complexity of animal displays. Much past research on signal evolution has focused on sensory specialists, or on single signals in isolation, but many animal displays involve complex signaling, or the combination of more than one signal or related component, often serially and overlapping, frequently across multiple sensory modalities. Here, we build a framework of functional hypotheses of complex signal evolution based on content-driven (ultimate) and efficacy-driven (proximate) selection pressures (sensu Guilford and Dawkins 1991). We point out key predictions for various hypotheses and discuss different approaches to uncovering complex signal function. We also differentiate a category of hypotheses based on inter-signal interactions. Throughout our review, we hope to make three points: (1) a complex signal is a functional unit upon which selection can act, (2) both content and efficacy-driven selection pressures must be considered when studying the evolution of complex signaling, and (3) individual signals or components do not necessarily contribute to complex signal function independently, but may interact in a functional way.

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Complex signal function: developing a framework of testable hypotheses

To date, learning has been demonstrated in six orders of insects with phytophagous habits. Most examples of learning in these orders have come to light relatively recently [all but a few since the last review of learning in the Annual Review of Entomology 17 years ago (3)], and many more will surely be uncovered. Still, the study of learning in other insects such as the social Hymenoptera remains far more advanced than in phytophagous insects. In this review, work on learning in phytophagous insects is placed into the broader perspective of learning in a variety of insects including bees and parasitic wasps. Although historically the study of learning in phytophagous insects has emphasized the acquisition of host resources for feeding and oviposition, we believe that, as in bees and wasps, learning impinges on almost every aspect of the natural history of a phytophagous insect. Following discussion of the definition of learning, the natural history of learning, and the host stimuli involved in learning, we review the ecological significance of learning with respect to both the traditional categories erected by behaviorists and the more recent concept of programed learning developed by ethologists. Finally, we evaluate critically hypotheses about the evolution of learning and suggest approaches for future research.

Ecological and evolutionary aspects of learning in phytophagous insects

Insect learning: ecological and evolutionary perspectives.

▪ Abstract Oviposition behavior in herbivorous and frugivorous insects and parasitoids is dynamic at the level of the individual, responding to variation in host quality and availability. Patterns of variation in egg load in response to host presence and quality suggest that ovarian development also responds to variation in the host environment. Ovarian dynamics are mediated by feedback from oviposition, by host feeding, and by sensory input from the host. The last of these mechanisms, host sensory cuing, is known to occur in three major orders and provides strong evidence that ovarian dynamics are adaptive by design. Conditions favoring host effects on ovarian development include trade-offs between egg production and either survival or dispersal, uncertainty in the host environment, and a correlation in host conditions between the time that oogenesis is initiated and the time that eggs are laid. Some host defenses block ovarian development, suggesting that ovarian dynamics in hostspecific insects should ...

Ovarian Dynamics and Host Use

Oviposition behavior in phytophagous insects and entomophagous parasitoids is often modified by the presence of conspecific brood (eggs and larvae). Often, females avoid laying eggs on or in hosts bearing brood, a behavior that acts to reduce the level of competition suffered by their offspring. Avoidance of occupied hosts is typically mediated by cues and/or signals associated with brood. In this article, we review the role of Marking Pheromones (MPs) as signals of brood presence in both phytophagous and entomophagous insects. We place information about the function and evolution of MPs in the context of recent theory in the field of animal communication. We highlight the dynamics of host-marking systems and discuss how effects of MPs vary according to factors such as female experience and egg load. We also examine variation in the form and function of MP communication across a variety of insect taxa. While studies of MP communication in phytophagous insects have focused on the underlying behavioral mechanisms and chemistry of MP communication, studies in entomophagous insects have focused on the functional aspects of MPs and their role in ‘decision-making’ in insects. We argue that an approach that incorporates the important contributions of both of these somewhat independent, but complementary areas of research will lead to a more complete understanding of MPs in insects. Finally, we suggest that MP systems are model systems for the study of animal signaling and its evolution.

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Daniel R. Papaj

Papers

Complex signal function: developing a framework of testable hypotheses

Ecological and evolutionary aspects of learning in phytophagous insects

Insect learning: ecological and evolutionary perspectives.

Ovarian Dynamics and Host Use

Host marking behavior in phytophagous insects and parasitoids