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

So far in this course we have dealt entirely with the evolution of characters that are controlled by simple Mendelian inheritance at a single locus. There are notes on the course website about gametic disequilibrium and how allele frequencies change at two loci simultaneously, but we didn’t discuss them. In every example we’ve considered we’ve imagined that we could understand something about evolution by examining the evolution of a single gene. That’s the domain of classical population genetics. For the next few weeks we’re going to be exploring a field that’s actually older than classical population genetics, although the approach we’ll be taking to it involves the use of population genetic machinery. If you know a little about the history of evolutionary biology, you may know that after the rediscovery of Mendel’s work in 1900 there was a heated debate between the “biometricians” (e.g., Galton and Pearson) and the “Mendelians” (e.g., de Vries, Correns, Bateson, and Morgan). Biometricians asserted that the really important variation in evolution didn’t follow Mendelian rules. Height, weight, skin color, and similar traits seemed to

Human biochemical genetics

Pseudoreplication is defined. as the use of inferential statistics to test for treatment effects with data from experiments where either treatments are not replicated (though samples may be) or replicates are not statistically independent. In ANOVA terminology, it is the testing for treatment effects with an error term inappropriate to the hypothesis being considered. Scrutiny of 176 experi- mental studies published between 1960 and the present revealed that pseudoreplication occurred in 27% of them, or 48% of all such studies that applied inferential statistics. The incidence of pseudo- replication is especially high in studies of marine benthos and small mammals. The critical features of controlled experimentation are reviewed. Nondemonic intrusion is defined as the impingement of chance events on an experiment in progress. As a safeguard against both it and preexisting gradients, interspersion of treatments is argued to be an obligatory feature of good design. Especially in small experiments, adequate interspersion can sometimes be assured only by dispensing with strict random- ization procedures. Comprehension of this conflict between interspersion and randomization is aided by distinguishing pre-layout (or conventional) and layout-specifit alpha (probability of type I error). Suggestions are offered to statisticians and editors of ecological j oumals as to how ecologists' under- standing of experimental design and statistics might be improved.

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Pseudoreplication and the Design of Ecological Field Experiments

Predation has long been implicated as a major selective force in the evolution of several morphological and behavioral characteristics of animals. The importance of predation during evolutionary ti...

Behavioral decisions made under the risk of predation: a review and prospectus

http://osenberglab.ecology.uga.edu/wp-content/uploads/2015/09/Charnov-1976-TPB.pdf

Optimal foraging, the marginal value theorem.

Body size is manifestly one of the most important attributes of an organism from an ecological and evolutionary point of view. Size has a predominant influence on an animal's energetic requirements, its potential for resource exploitation, and its susceptibility to natural enemies. A large literature now exists on how physiological, life history, and population parameters scale with body dimensions (24, 131). The ecological literature on species interactions and the structure of animal communities also stresses the importance of body size. Differences in body size are a major means by which species avoid direct overlap in resource use (153), and size-selective predation can be a primary organizing force in some communities (20, 70). Size thus imposes important constraints on the manner in which an organism interacts with its environment and influences the strength, type, and symmetry of interactions with other species (152, 207). Paradoxically, ecologists have virtually ignored the implications of these observations for interactions among species that exhibit size-distributed populations. For instance, it has been often suggested that competing species

https://www.jstor.org/stable/pdf/2096954.pdf

The ontogenetic niche and species interactions in size-structured populations

We present an experiment designed to test the hypothesis that fish respond to both relative predation risk and habitat profitability in choosing habitats in which to feed. Identical populations of three size—classes of bluegill sunfish (Lepomis macrochirus) were stocked on both sides of a divided pond (29 m in diameter), and eight piscivorous largemouth bass (Micropterus salmoides) were introduced to one side. Sizes of both species were chosen such that the small class of bluegills was very vulnerable to the bass, whereas the largest class was invulnerable to bass predation. We then compared mortality, habitat use, and growth of each size—class in the presence and absence of the bass. Only the small size—class suffered significant mortality from the bass (each bass consumed on average about one small bluegill every 3.8 d); the two larger size—classes exhibited similar mortality rates on both sides of the pond. In the absence of the bass, we found that habitat use of all size—classes was similar and that the pattern of habitat use maximized foraging return rates (Werner et al. 1983). In the presence of the bass the two larger size—classes chose habitats to maximize return rates, but the small size—class obtained a greater fraction of its diet from the vegetation habitat, where foraging return rates were only one—third of those in the more open habitats. The small size—class further exhibited a significant depression in individual growth in the presence of the bass; the growth increment during the experiment was 27% less than that for small bluegills in the absence of the bass. Because of the reduced utilization of more open habitats by the small fish in the presence of bass, resources in these habitats were released to the larger size—classes, which showed greater growth in the presence of the bass than in its absence. We develop methods to predict the additional mortality expected on a cohort due to a reduction in growth rate (because individuals are spending a longer time in vunerable sizes), and discuss and potential for predation risk to enforce size—class segregation, which leads de facto to resource partitioning.

An Experimental Test of the Effects of Predation Risk on Habitat Use in Fish

In this paper we review the empirical studies documenting trait-mediated indirect interactions (TMIIs) in food webs. Basic models and empirical approaches that form the foundation of our conceptualization of species interactions generally assume that interactions are an intrinsic property of the two interacting species and therefore are governed by their respective densities. However, if a species reacts to the presence of a second species by altering its phenotype, then the trait changes in the reacting species can alter the per capita effect of the reacting species on other species and, consequently, population density or fitness of the other species. Such trait-mediated indirect interactions can reinforce or oppose density-mediated effects and have been largely overlooked by community ecologists. We first briefly develop the case for the broad mechanistic basis for TMIIs and then review the direct evidence for TMIIs in various permutations of simple three- to four-species food webs. We find strong evid...

A review of trait-mediated indirect interactions in ecological communities

▪ Abstract Lentic freshwater habitats in temperate regions exist along a gradient from small ephemeral ponds to large permanent lakes. This environmental continuum is a useful axis for understanding how attributes of individuals ultimately generate structure at the level of the community. Community structure across the gradient is determined by both (a) physical factors, such as pond drying and winter anoxia, that limit the potential breadth of species distributions, and (b) biotic effects mediated by ecological interactions, principally predation, that determine the realized success of species. Fitness tradeoffs associated with a few critical traits of individuals often form the basis for species turnover along the gradient. Among species that inhabit temporary ponds, distributions are often constrained because traits that enhance developmental rate and competitive ability also increase susceptibility to predators. In permanent ponds, changes in the composition of major predators over the gradient limit ...

/pdf/mechanisms-creating-community-structure-across-a-freshwater-3osigazq5u.pdf

Mechanisms creating community structure across a freshwater habitat gradient

The bluegill sunfish, Lepomis macrochirus, is known to select prey on the basis of size. We present evidence that this size selection is related to the optimal allocation of time spent searching for, and handling prey. A model relating search and handling time to energy return is constructed to determine the optimal breadth of diet. Prey are permitted to differ in size and relative abundance. All elements of the model are estimated from experiments with the bluegill feeding on populations constructed from size classes of Daphnia magna. Relative visibility of the different prey sizes markedly affects relative encounter rates or "effective" proportions. Effective proportions are determined empirically from feeding experiments and theoretically from reaction distance in order to correct for this bias. Search time is then manipulated by varying absolute abundance of prey. At low absolute abundance, prey of different size are eaten as encountered. As prey abundance is increased, size classes are dropped sequentially from the diet in accordance with the theory. Search and handling times are estimated for these experiments and quantitative comparisons with the model indicate these changes in diet maximize return with respect to time spent foraging.

Earl E. Werner

Papers

The ontogenetic niche and species interactions in size-structured populations

An Experimental Test of the Effects of Predation Risk on Habitat Use in Fish

A review of trait-mediated indirect interactions in ecological communities

Mechanisms creating community structure across a freshwater habitat gradient

Optimal Foraging and the Size Selection of Prey by the Bluegill Sunfish (Lepomis Macrochirus)