Marjorie J. Wonham

Impact: toward a framework for understanding the ecological effects of invaders

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.

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

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 models numerical simulations predict disease levels that are consistent with independent data.

Trends in marine biological invasions at local and regional scales: the Northeast Pacific Ocean as a model system

Introduced species are an increasing agent of global change. Biogeographic comparisons of introduced biotas at regional and global scales can clarify trends in source regions, invasion pathways, sink regions, and survey effort. We identify the Northeast Pacific Ocean (NEP; northern California to British Columbia) as a model system for analyzing patterns of marine invasion success in cool temperate waters. We review literature and field surveys, documenting 123 introduced invertebrate, algal, fish, and vascular plant species in the NEP. Major invasion pathways were shipping (hull fouling, solid and water ballast; 1500s-present) and shellfish (particularly oysters) and finfish imports (commonest from the 1870s to mid-1900s). The cumulative number of successful invasions over time increased at linear, quadratic, and exponential rates for different taxa, pathways, and regions within the NEP. Regional analysis of four major NEP estuaries showed that Puget Sound and the contiguous Straits had the most introduced species, followed by Humboldt Bay, Coos Bay and Willapa Bay. Data on cumulative shipping volumes predicted smaller-scale, but not larger-scale spatial patterns in the number of shipping-mediated invasions. We identify the major challenges in scaling up from regional to global invasion analysis in cool temperate regions. Retrospective analyses for distinct biogeographic regions such as the NEP provide insight into vector dynamics and regional invasibility, and are a necessary foundation for monitoring and managing global change caused by biotic invasions.

Invasion pressure to a ballast-flooded estuary and an assessment of inoculant survival

The relationships between invasion pressure, post-transport inoculant survival, and regional susceptibility to invasion are poorly understood. In marine ecosystems, the movement and release of ballast water from ocean-going ships provides a model system by which to examine the interplay among these factors. One of the largest estuaries in North America, the Chesapeake Bay, receives tremendous amounts of foreign ballast water annually and thus should be at high invasion risk. To date, however, few introductions in Chesapeake Bay have been attributed to ballast release. To understand better the dynamics of this invasion process, we (1) characterized and quantified the biota arriving to Chesapeake Bay in foreign ballast water, (2) compared temperatures and salinities of ballast water and harbor water in upper Chesapeake Bay, and (3) tested experimentally survival of organisms collected from ballast water in temperatures and salinities characteristic of the region. From 1993 to 1994, we sampled planktonic and benthic organisms from 60 foreign vessels arriving to Chesapeake Bay. Our data show that the estuary is being inoculated by a diverse assemblage of aquatic organisms from around the world. Furthermore, the short transit time (≤15 d) for most vessels ensured that substantial numbers of larval and post-larval organisms were being deballasted alive. Most of the ballast water discharged into the upper Chesapeake Bay, however, was significantly higher in salinity (>20‰) than that of the receiving harbor. In laboratory tolerance experiments, ballast water organisms perished under such conditions. Thus, a mismatch in physical conditions between donor and receiver regions may explain the dearth of invasions in the upper Bay. It is likely that the lower Chesapeake Bay, which is more saline, remains at higher risk to ballast water invasion. Recognition of such intraregional differences should allow more focused predictions for monitoring and management.

Color Polymorphism and Genetic Structure in the Sea Star Pisaster ochraceus

The sea star Pisaster ochraceus is one of the more striking species on the rocky shores of the Northeast Pacific, in part due to the dramatic color polymorphism of the adults. Along the open Pacific coast, Pisaster populations are 6%–28% orange, with a small percentage of brilliant purple stars and a large percentage of reddish-brown to dull purple ones. However, populations in the San Juan Island Archipelago (Washington, USA) and the southern Strait of Georgia (British Columbia, Canada) are almost entirely brilliant purple. The factors that maintain the color polymorphism, and those that contribute to among-site variation in color frequencies, remain unknown. We examined the relationships between color frequencies and several ecological and morphological variables, and conducted a large-scale phylogeographic survey of Pisaster populations. We found very low population genetic structure, suggesting that gene flow is high and geographic variation in color frequencies is not a vestige of Pleistocene glacial refugia. Color frequencies are also unrelated to adult size and to the frequency of injury within a population. However, there are suggestive relationships between color frequency and diet, and with areas of potentially low salinity. We propose that, although the color polymorphism may have an underlying genetic component, the regional-scale variation in color frequency is ecologically controlled.

Minimizing invasion risk by reducing propagule pressure: a model for ballast-water exchange

Biological invasions are a major and increasing agent of global biodiversity change. Theory and practice indicate that invasion risk can be diminished by reducing propagule pressure, or the quantity, quality, and frequency of introduced individuals. For aquatic invasions, the primary global invasion pathway is ballast-water transport, and the primary risk reduction strategy is currently open-ocean exchange. Exchange was developed with shipping between freshwater ports in mind, but the majority of shipping connects brackish and marine ports. A worldwide convention, adopted in 2004 by the International Maritime Organization, now mandates ballast-water exchange (or equivalent management) for its 164 member states. Will exchange be as effective in reducing invasion risk for euryhaline species (those capable of tolerating a wide range of salinity levels) in salt-water ports? Here we develop a simple mathematical framework for optimizing ballast-water exchange in terms of exchange level, timing, and species sal...

Recovery of the brown alga Fucus gardneri following a range of removal intensities

Abstract The consequences of disturbance are known to be non-linear functions of disturbance intensity. In this study, we investigate the effects of a range of disturbance intensities by experimentally manipulating the cover of Fucus gardneri , a dominant intertidal alga. The effects of disturbance on subsequent Fucus recovery were similar for canopy reductions of 0–80%. However, complete removal of all Fucus individuals delayed Fucus recovery, as measured by canopy cover, by at least several months. Despite changes in the dominant alga, the abundance of invertebrates and other algae did not respond to cover manipulation. Our results suggest that the recovery from small-scale disturbances is enhanced if at least a few individuals survive the event. Only by examining a variety of levels of natural or anthropogenic disturbance will ecologists be able to understand observed recovery patterns and design management strategies.

A Comparative Analysis of Models for West Nile Virus

This chapter describes the steps needed to formulate, analyze and apply epidemiological models to vector-borne diseases. Our models focus on West Nile (WN) virus, an emerging infectious disease in North America, first identified in Africa. We begin by introducing a minimalist model for WN dynamics to illustrate the processes of model formulation, analysis, and application. We then revisit the question of model formulation to examine how two major biological assumptions affect the model structure and therefore its predictions. Next, we briefly compare these different model structures in an introductory exercise of model parameterization, validation, and comparison. Finally, we address model applications in more detail with two examples of how the model output can usefully be connected to public health applications.

Modeling Marine Invasions: Current and Future Approaches

This chapter focuses on how dynamical mathematical modeling has been and could be useful in understanding marine biological invasions. Mathematical models have long been central to the development of general ecological and invasion theory (e.g., Case 1990; Hastings et al. 2005; Lewis and Kareiva 1993; Neubert and Parker 2004; Shigesada and Kawasaki 1997). Although the dynamics of marine systems can be challenging to observe and model (e.g., deYoung et al. 2004; Kinlan et al. 2005), mathematical models are nonetheless beginning to provide insights into invasion dynamics in marine systems.