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

Statistical method for testing the neutral mutation hypothesis by DNA polymorphism.

. Much confusion exists in the English-language literature on plant invasions concerning the terms ‘naturalized’ and ‘invasive’ and their associated concepts. Several authors have used these terms in proposing schemes for conceptualizing the sequence of events from introduction to invasion, but often imprecisely, erroneously or in contradictory ways. This greatly complicates the formulation of robust generalizations in invasion ecology.

Based on an extensive and critical survey of the literature we defined a minimum set of key terms related to a graphic scheme which conceptualizes the naturalization/invasion process. Introduction means that the plant (or its propagule) has been transported by humans across a major geographical barrier. Naturalization starts when abiotic and biotic barriers to survival are surmounted and when various barriers to regular reproduction are overcome. Invasion further requires that introduced plants produce reproductive offspring in areas distant from sites of introduction (approximate scales: > 100 m over  6 m/3 years for taxa spreading by roots, rhizomes, stolons or creeping stems). Taxa that can cope with the abiotic environment and biota in the general area may invade disturbed, seminatural communities. Invasion of successionally mature, undisturbed communities usually requires that the alien taxon overcomes a different category of barriers.

We propose that the term ‘invasive’ should be used without any inference to environmental or economic impact. Terms like ‘pests’ and ‘weeds’ are suitable labels for the 50–80% of invaders that have harmful effects. About 10% of invasive plants that change the character, condition, form, or nature of ecosystems over substantial areas may be termed ‘transformers’.

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Naturalization and invasion of alien plants: concepts and definitions

By Matthew James Keelingand Pejman RohaniPrinceton, NJ: Princeton University Press,2008.408 pp., Illustrated. $65.00 (hardcover).Mathematical modeling of infectious dis-eases has progressed dramatically over thepast 3 decades and continues to ﬂourishat the nexus of mathematics, epidemiol-ogy, and infectious diseases research. Nowrecognized as a valuable tool, mathemat-ical models are being integrated into thepublic health decision-making processmore than ever before. However, despiterapid advancements in this area, a formaltraining program for mathematical mod-eling is lacking, and there are very fewbooks suitable for a broad readership. Tosupport this bridging science, a commonlanguage that is understood in all con-tributing disciplines is required.

Modeling Infectious Diseases in Humans and Animals

■ Abstract Contributions from the field of population biology hold promise for understanding and managing invasiveness; invasive species also offer excellent opportunities to study basic processes in population biology. Life history studies and demographic models may be valuable for examining the introduction of invasive species and identifying life history stages where management will be most effective. Evolutionary processes may be key features in determining whether invasive species establish and spread. Studies of genetic diversity and evolutionary changes should be useful for

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The Population Biology of Invasive Species

Although ecologists commonly talk about the impacts of nonindigenous species, little formal attention has been given to defining what we mean by impact, or connecting ecological theory with particular measures of impact. The resulting lack of generalizations regarding invasion impacts is more than an academic problem; we need to be able to distinguish invaders with minor effects from those with large effects in order to prioritize management efforts. This paper focuses on defining, evaluating, and comparing a variety of measures of impact drawn from empirical examples and theoretical reasoning. We begin by arguing that the total impact of an invader includes three fundamental dimensions: range, abundance, and the per-capita or per-biomass effect of the invader. Then we summarize previous approaches to measuring impact at different organizational levels, and suggest some new approaches. Reviewing mathematical models of impact, we argue that theoretical studies using community assembly models could act as a basis for better empirical studies and monitoring programs, as well as provide a clearer understanding of the relationship among different types of impact. We then discuss some of the particular challenges that come from the need to prioritize invasive species in a management or policy context. We end with recommendations about how the field of invasion biology might proceed in order to build a general framework for understanding and predicting impacts. In particular, we advocate studies designed to explore the correlations among different measures: Are the results of complex multivariate methods adequately captured by simple composite metrics such as species richness? How well are impacts on native populations correlated with impacts on ecosystem functions? Are there useful bioindicators for invasion impacts? To what extent does the impact of an invasive species depend on the system in which it is measured? Three approaches would provide new insights in this line of inquiry: (1) studies that measure impacts at multiple scales and multiple levels of organization, (2) studies that synthesize currently available data on different response variables, and (3) models designed to guide empirical work and explore generalities.

Impact: Toward a Framework for Understanding the Ecological Effects of Invaders

▪ Abstract Biological invasions of marine habitats have been common, and many patterns emerge from the existing literature. In North America, we identify 298 nonindigenous species (NIS) of invertebrates and algae that are established in marine and estuarine waters, generating many “apparent patterns” of invasion: (a) The rate of reported invasions has increased exponentially over the past 200 years; (b) Most NIS are crustaceans and molluscs, while NIS in taxonomic groups dominated by small organisms are rare; (c) Most invasions have resulted from shipping; (d) More NIS are present along the Pacific coast than the Atlantic and Gulf coasts; (e) Native and source regions of NIS differ among coasts, corresponding to trade patterns. The validity of these apparent patterns remains to be tested, because strong bias exists in the data. Overall, the emergent patterns reflect interactive effects of propagule supply, invasion resistance, and sampling bias. Understanding the relative contribution of each component re...

Invasion of Coastal Marine Communities in North America: Apparent Patterns, Processes, and Biases

Infectious diseases present ecological and public health challenges that can be addressed with mathematical models. Certain pathogens, however, including the emerging West Nile virus (WN) in North America, exhibit a complex seasonal ecology that is not readily analysed with standard epidemiological methods. We develop a single-season susceptible–infectious–removed (SIR) model of WN cross-infection between birds and mosquitoes, incorporating specific features unique to WN ecology. We obtain the disease reproduction number, R0, and show that mosquito control decreases, but bird control increases, the chance of an outbreak. We provide a simple new analytical and graphical method for determining, from standard public health indicators, necessary mosquito control levels. We extend this method to a seasonally variable mosquito population and outline a multi-year model framework. The model’s numerical simulations predict disease levels that are consistent with independent data.

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An epidemiological model for West Nile virus: invasion analysis and control applications.

Most studies characterizing successful biological invaders emphasize those traits that help a species establish a new population. Invasions are, however, multi-phase processes with at least two phases, dispersal and introduction, that occur before establishment. Characteristics that enhance survival at any of these three phases will contribute to invasion success. Here, we synthesize information on the dispersal, introduction, and establishment of fishes mediated by ship ballast-water transport. We synthesize 54 reports of at least 31 fish species collected from ballast tanks (Phase 1), including 28 new reports from our recent studies (1986 to 1996). Our literature survey revealed 40 reports of 32 fish species whose introductions have been attributed to ballast transport (Phase 2), of which at least 24 survived to establish persistent populations (Phase 3). We detected little overlap at the species level between these two data sets (Phase 1 vs Phases 2 and 3), but patterns emerged at the family level. The Gobiidae (6 species), Clupeidae (4 species), and Gasterosteidae (1 species) were the most commonly found fish families in ballast tanks (Phase 1). The Gobiidae (13 species), Blenniidae (6 species) and Pleuronectidae (2 species) dominated the list of ballast-mediated introductions (Phase 2); gobies and blennies were the families most frequently established (Phase 3). The invasive success of gobies and blennies may be explained in part by their crevicolous nature: both groups seek refuge and lay eggs in small holes, and may take advantage of the ballast-intake holes on ship hulls. This behavior, not typically associated with invasive ability, may contribute to successful introduction and establishment by facilitating the dispersal phase of invasion. The failure of the pleuronectids to invade may reflect poor salinity match between donor and recipient regions. To develop a predictive framework of invasion success, organisms must be sampled at all three phases of the invasion process. Our comparison of two ballast sampling methods suggests that fishes have been undersampled in ballast-water studies, including our own, and that the role of ballast transport in promoting fish invasions has been underestimated.

Fish and ships: relating dispersal frequency to success in biological invasions

▪ Abstract The Exxon Valdez oil spill was the largest in US maritime history. We review post-spill research and set it in its legal context. The Exxon Corporation, obviously responsible for the spill, focused on restoration, whereas the Trustees, a coalition of state and federal entities, focused on damage and its assessment. Despite billions of dollars expended, little new understanding was gained about the recovery dynamics of a high latitude marine ecosystem subject to an anthropogenic pulse perturbation. We discuss a variety of case studies that highlight the limitations to and shortcomings of the research effort. Given that more spills are inevitable, we recommend that future studies address spatial patterns in the intertidal, and focus on the abundances of long-lived species and on organisms that preserve a chronological record of growth. Oil spills, while tragic, represent opportunities to gain insight into the dynamics of marine ecosystems and should not be wasted.

Marjorie J. Wonham

Papers

Impact: Toward a Framework for Understanding the Ecological Effects of Invaders

Invasion of Coastal Marine Communities in North America: Apparent Patterns, Processes, and Biases

An epidemiological model for West Nile virus: invasion analysis and control applications.

Fish and ships: relating dispersal frequency to success in biological invasions

TROUBLE ON OILED WATERS: Lessons from the Exxon Valdez Oil Spill