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

Studies of local adaptation provide important insights into the power of natural selection relative to gene flow and other evolutionary forces. They are a paradigm for testing evolutionary hypotheses about traits favoured by particular environmental factors. This paper is an attempt to summarize the conceptual framework for local adaptation studies. We first review theoretical work relevant for local adaptation. Then we discuss reciprocal transplant and common garden experiments designed to detect local adaptation in the pattern of deme · habitat interaction for fitness. Finally, we review research questions and approaches to studying the processes of local adaptation ‐ divergent natural selection, dispersal and gene flow, and other processes affecting adaptive differentiation of local demes. We advocate multifaceted approaches to the study of local adaptation, and stress the need for experiments explicitly addressing hypotheses about the role of particular ecological and genetic factors that promote or hinder local adaptation. Experimental evolution of replicated populations in controlled spatially heterogeneous environments allow direct tests of such hypotheses, and thus would be a valuable way to complement research on natural populations.

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Conceptual issues in local adaptation

Genetics and analysis of quantitative traits

Summary
1The role of phenotypic plasticity in evolution has historically been a contentious issue because of debate over whether plasticity shields genotypes from selection or generates novel opportunities for selection to act. Because plasticity encompasses diverse adaptive and non-adaptive responses to environmental variation, no single conceptual framework adequately predicts the diverse roles of plasticity in evolutionary change.
2Different types of phenotypic plasticity can uniquely contribute to adaptive evolution when populations are faced with new or altered environments. Adaptive plasticity should promote establishment and persistence in a new environment, but depending on how close the plastic response is to the new favoured phenotypic optimum dictates whether directional selection will cause adaptive divergence between populations. Further, non-adaptive plasticity in response to stressful environments can result in a mean phenotypic response being further away from the favoured optimum or alternatively increase the variance around the mean due to the expression of cryptic genetic variation. The expression of cryptic genetic variation can facilitate adaptive evolution if by chance it results in a fitter phenotype.
3We conclude that adaptive plasticity that places populations close enough to a new phenotypic optimum for directional selection to act is the only plasticity that predictably enhances fitness and is most likely to facilitate adaptive evolution on ecological time-scales in new environments. However, this type of plasticity is likely to be the product of past selection on variation that may have been initially non-adaptive.
4We end with suggestions on how future empirical studies can be designed to better test the importance of different kinds of plasticity to adaptive evolution.

/pdf/adaptive-versus-non-adaptive-phenotypic-plasticity-and-the-55680hih4y.pdf

Adaptive versus non‐adaptive phenotypic plasticity and the potential for contemporary adaptation in new environments

▪ Abstract Changes in ploidy occurred early in the diversification of some animal and plant lineages and represent an ongoing phenomenon in others. While the prevalence of polyploid lineages indicates that this phenomenon is a common and successful evolutionary transition, whether polyploidization itself has a significant effect on patterns and rates of diversification remains an open question. Here we review evidence for the creative role of polyploidy in evolution. We present new estimates for the incidence of polyploidy in ferns and flowering plants based on a simple model describing transitions between odd and even base chromosome numbers. These new estimates indicate that ploidy changes may represent from 2 to 4% of speciation events in flowering plants and 7% in ferns. Speciation via polyploidy is likely to be one of the more predominant modes of sympatric speciation in plants, owing to its potentially broad-scale effects on gene regulation and developmental processes, effects that can produce immed...

/pdf/polyploid-incidence-and-evolution-18k59e8zrk.pdf

Polyploid Incidence and Evolution

Preface Introduction: Themes, objectives, and structure 1. The historical context of differentiation and diversity 2. The biogeography and ecology of endemism 3. The evolution of endemism : from population differentiation to species divergence 4. Trait variation, adaptation, and dispersal in the Mediterranean mosaic 5. Variation and evolution of reproductive traits in the Mediterranean mosaic 6. Ecology and evolution of domesticated and invasive species Conclusions: Endemism, adaptation, and conservation References Species list and nomenclature Index

Plant Evolution in the Mediterranean

The role of polyploidy in the origin of evolutionary novelty and the maintenance of diversity in plant populations has come to be recognized as an integral component of the ecological and evolutionary dynamics of plant species populations. Recent attempts to examine the evolutionary significance of polyploidy have focused on the processes responsible for the origin of polyploid plants and the conditions that favour their establishment and persistence. The importance of these issues is not simply limited to the evolutionary dynamics of polyploidy but is, in fact, central to our understanding of the population biology processes that act on the establishment of new ‘types' and the maintenance of biotic diversity at both the inter- and intraspecific levels.

The evolutionary dynamics of polyploid plants: origins, establishment and persistence.

Phenotypic plasticity has often been assumed to buffer the effects of natural selection and thus act as a constraint on evolutionary change. It has become increasingly clear, however, that phenotypic plasticity actually represents a fundamental component of evolutionary change. Where genetic variation for plasticity exists, a population with a different mean plasticity can evolve. Recent attention has been focused on the conditions necessary for the evolution of phenotypic plasticity, i.e. those under which a generalist strategy, as opposed to a range of genetically differentiated specialists, will be favoured. It is also now clear that genotypes that perform best in one environment usually perform less well than other genotypes in a different environment; hence, their greater response is not an adaptation to environmental variation. A response to environmental variation is only adaptive if it represents a mechanism by which relative fitness is maintained in the face of environmental variation. Adaptive plasticity may thus involve both physiological homeostasis and morphological response.

Phenotypic plasticity as a component of evolutionary change.

The objective of this study is to examine whether habitat, herbivory and traits related to resource acquisition, resource conservation, reproduction and dispersal differ between narrow endemic plant species and their widespread congeners. We undertook pairwise contrasts of 25 ecological characteristics and biological traits in 20 congeneric pairs of narrow endemic and widespread plant species in the French Mediterranean region. Within each pair, the two species had the same life-form, pollination mode and dispersal mode. Endemic species differed significantly from widespread congeners for a number of attributes. Endemic species occur in habitats on steeper slopes, with higher rock cover and in lower and more open vegetation than their widespread congeners. Endemic species are significantly smaller than widespread species, but show no differences in traits related to resource acquisition (specific leaf area, leaf nitrogen content, maximum photosynthetic rate) or resource conservation (leaf dry matter content). After accounting for their smaller stature, endemic species produce fewer and smaller flowers with less stigma-anther separation and lower pollen/ovule ratios and produce fewer seeds per plant than their widespread congeners. No consistent variation in seed mass and propagule structure was found between congeneric species. Herbivory levels did not differ between congeneric species. Ecological characteristics, notably the occupation of rocky habitats with low aboveground competition, may thus have played an important role in the differentiation of narrow endemic species in the Western Mediterranean. Morphological and ecophysiological traits of narrow endemic species indicate that they are not more stress-tolerant than their widespread congeners. Lower investment in pollen transfer and seed production suggest that local persistence is a key feature of the population ecology of narrow endemic species.

The biology and ecology of narrow endemic and widespread plants: a comparative study of trait variation in 20 congeneric pairs

Thymus vulgaris has a chemical polymorphism with six different chemotypes that show marked spatial segregation in nature. Although some populations have a single chemotype in majority, many have two or three chemotypes. In this study we analyze the quantitative variation among T. vulgaris populations in the percentage of oil composed of the dominant monoterpene(s) for each chemotype. In general, phenolic chemotypes (thymol and carvacrol), which occur at the end of the biosynthetic chain, have a significantly lower proportion of their oil composed of their dominant monoterpene than nonphenolic chemotypes (geraniol, α-terpineol, and linalool). This is due to the presence of high amounts of precursors (γ-terpinene and paracymene) in the oil of phenolic chemotypes. The essential oil of the nonphenolic thuyanol chemotype has four characteristic monoterpenes that together make up a lower proportion of the oil than the single dominant monoterpene of the other nonphenolic chemotypes. For all chemotypes, the percentage composition of the dominant monoterpene decreased significantly at sites where the chemotype is not the majority type. This decrease is correlated with a significant increase in either the proportion of the two precursors for the thymol chemotype or the monoterpenes characteristic of the other chemotypes at the site. The latter result suggests that a plant with dominant genes is responsible for the production of different monoterpenes can produce several molecules.

John D. Thompson

Papers

Plant Evolution in the Mediterranean

The evolutionary dynamics of polyploid plants: origins, establishment and persistence.

Phenotypic plasticity as a component of evolutionary change.

The biology and ecology of narrow endemic and widespread plants: a comparative study of trait variation in 20 congeneric pairs

QUALITATIVE AND QUANTITATIVE VARIATION IN MONOTERPENE CO-OCCURRENCE AND COMPOSITION IN THE ESSENTIAL OIL OF Thymus vulgaris CHEMOTYPES