Anthony Ricciardi

Assisted colonization is not a viable conservation strategy

A potential conservation strategy increasingly discussed by conservation biologists is the translocation of species to favorable habitat beyond their native range to protect them from human-induced threats, such as climate change. Even if preceded by careful risk assessment, such action is likely to produce myriad unintended and unpredictable consequences. Accurate risk assessment is impeded by contingency: the impacts of introduced species vary over time and space under the influence of local environmental variables, interspecific interactions and evolutionary change. Some impacts, such as native species extinctions, are large and irrevocable. Here we argue that conservation biologists have not yet developed a sufficient understanding of the impacts of introduced species to make informed decisions regarding species translocations.

OVERLAND DISPERSAL OF AQUATIC INVASIVE SPECIES: A RISK ASSESSMENT OF TRANSIENT RECREATIONAL BOATING

Predictions of the geographic spread of introduced species are often limited by a lack of data on their mechanisms of dispersal. We interviewed boaters and inspected boating equipment at public boat launches on Lake St. Clair (Michigan, USA) to assess the potential for the zebra mussel, an invasive bivalve, to be dispersed overland to inland waters by transient recreational boating activities. Several mechanisms associated with recreational boating were found to be capable of transporting either larval or adult life stages. Larvae were found in all forms of water carried by boats (i.e., in live wells, bilges, bait buckets, and engines) but were estimated to be 40–100× more abundant in live wells than other locations. Dilution in receiving waters should, however, greatly reduce the risk of establishing new populations by the introduction of larvae. Contrary to common belief, mussel dispersal from these boat launches did not occur by direct attachment to transient boats. Rather, adult and juvenile mussels were transported primarily on macrophytes entangled on boat trailers and, less frequently, on anchors (5.3% and 0.9% of departing boats, respectively). Combining these data with estimates of survival in air and reported boater destinations, we predict that a maximum of 0.12% of the trailered boats departing these access sites delivered live adult mussels to inland waters solely by transport on entangled macrophytes. While this is a small probability, high levels of vector activity resulted in a prediction of a total of 170 dispersal events to inland waters within the summer season from the primary boat launch studied. Many other potential vectors remain to be assessed, but the dispersal of zebra mussels by trailered boats, particularly by “piggybacking” on macrophytes entangled on the trailers, must be controlled in order to limit further range expansion of the zebra mussel within North America.

Progress toward understanding the ecological impacts of nonnative species.

A predictive understanding of the ecological impacts of nonnative species has been slow to develop, owing largely to an apparent dearth of clearly defined hypotheses and the lack of a broad theoretical framework. The context dependency of impact has fueled the perception that meaningful generalizations are nonexistent. Here, we identified and reviewed 19 testable hypotheses that explain temporal and spatial variation in impact. Despite poor validation of most hypotheses to date, evidence suggests that each can explain at least some impacts in some situations. Several hypotheses are broad in scope (applying to plants and animals in virtually all contexts) and some of them, intriguingly, link processes of colonization and impact. Collectively, these hypotheses highlight the importance of the functional ecology of the nonnative species and the structure, diversity, and evolutionary experience of the recipient community as general determinants of impact; thus, they could provide the foundation for a theoretical framework for understanding and predicting impact. Further substantive progress toward this goal requires explicit consideration of within-taxon and across-taxa variation in the per capita effect of invaders, and analyses of complex interactions between invaders and their biotic and abiotic environments.

A unified classification of alien species based on the magnitude of their environmental impacts.

We present a method for categorising and comparing alien or invasive species in terms of how damaging they are to the environment, that can be applied across all taxa, scales, and impact metrics.

Bridging Troubled Waters: Biological Invasions, Transoceanic Shipping, and the Laurentian Great Lakes

Abstract Release of contaminated ballast water by transoceanic ships has been implicated in more than 70% of faunal nonindigenous species (NIS) introductions to the Great Lakes since the opening of the St. Lawrence Seaway in 1959. Contrary to expectation, the apparent invasion rate increased after the initiation of voluntary guidelines in 1989 and mandatory regulations in 1993 for open-ocean ballast water exchange by ships declaring ballast on board (BOB). However, more than 90% of vessels that entered during the 1990s declared no ballast on board (NOBOB) and were not required to exchange ballast, although their tanks contained residual sediments and water that would be discharged in the Great Lakes. Lake Superior receives a disproportionate number of discharges by both BOB and NOBOB ships, yet it has sustained surprisingly few initial invasions. Conversely, the waters connecting lakes Huron and Erie are an invasion hotspot despite receiving disproportionately few ballast discharges. Other vectors, including canals and accidental release, have contributed NIS to the Great Lakes and may increase in relative importance in the future. Based on our knowledge of NIS previously established in the basin, we have developed a vector assignment protocol to systematically ascertain vectors by which invaders enter the Great Lakes.

The invasiveness of an introduced species does not predict its impact

Inconsistent use of terminology plagues the study and management of biological invasions. The term “invasive” has been used to describe inter alia (1) any introduced non-indigenous species; (2) introduced species that spread rapidly in a new region; and (3) introduced species that have harmful environmental impacts, particularly on native species. The second definition in various forms is more commonly used by ecologists, while the third definition is pervasive in policy papers and legislation. We tested the relationship between the invasiveness of an introduced species and its impact on native biodiversity. We quantified a species’ invasiveness by both its rate of establishment and its rate of spread, while its impact was assigned a categorical ranking based on the documented effects of the invader on native species populations. We found no correlations between these variables for introduced plants, mammals, fishes, invertebrates, amphibians and reptiles, suggesting that the mechanisms of invasion and impact are not strongly linked. Our results support the view that the term “invasive” should not be used to connote negative environmental impact.

Invasion risks posed by the aquarium trade and live fish markets on the Laurentian Great Lakes

International trade is an important mechanism for global non-indigenous species introductions, which have had profound impacts on the biodiversity of aquatic ecosystems including the Laurentian Great Lakes. The best-documented vector by which non-indigenous species have entered the Great Lakes is ballast water discharged by transoceanic ships. A variety of potential alternative vectors exist, including the intentional release of aquarium or food organisms. To assess whether these vectors pose a significant invasion risk for the Great Lakes, we surveyed fish sold live in markets and fish, mollusks and macrophytes sold in pet and aquarium stores within the Great Lakes watershed. We evaluated invasion risk using information on species’ thermal tolerance, history of invasion elsewhere, and potential propagule loads as indicated by frequency of occurrence in shops. Our research suggests that both the aquarium industry and live fish markets represent potential sources of future invaders to the Great Lakes, including several aquarium fishes and macrophytes, as well as Asian carp species sold in fish markets. Currently, few regulatory mechanisms exist to control these potential vectors.

Defining the Impact of Non-Native Species

Non-native species cause changes in the ecosystems to which they are introduced. These changes, or some of them, are usually termed impacts; they can be manifold and potentially damaging to ecosystems and biodiversity. However, the impacts of most non-native species are poorly understood, and a synthesis of available information is being hindered because authors often do not clearly define impact. We argue that explicitly defining the impact of non-native species will promote progress toward a better understanding of the implications of changes to biodiversity and ecosystems caused by non-native species; help disentangle which aspects of scientific debates about non-native species are due to disparate definitions and which represent true scientific discord; and improve communication between scientists from different research disciplines and between scientists, managers, and policy makers. For these reasons and based on examples from the literature, we devised seven key questions that fall into 4 categories: directionality, classification and measurement, ecological or socio-economic changes, and scale. These questions should help in formulating clear and practical definitions of impact to suit specific scientific, stakeholder, or legislative contexts. Definiendo el Impacto de las Especies No-Nativas Resumen Las especies no-nativas pueden causar cambios en los ecosistemas donde son introducidas. Estos cambios, o algunos de ellos, usualmente se denominan como impactos; estos pueden ser variados y potencialmente dañinos para los ecosistemas y la biodiversidad. Sin embargo, los impactos de la mayoría de las especies no-nativas están pobremente entendidos y una síntesis de información disponible se ve obstaculizada porque los autores continuamente no definen claramente impacto. Discutimos que definir explícitamente el impacto de las especies no-nativas promoverá el progreso hacia un mejor entendimiento de las implicaciones de los cambios a la biodiversidad y los ecosistemas causados por especies no-nativas; ayudar a entender cuáles aspectos de los debates científicos sobre especies no-nativas son debidos a definiciones diversas y cuáles representan un verdadero desacuerdo científico; y mejorar la comunicación entre científicos de diferentes disciplinas y entre científicos, administradores y quienes hacen las políticas. Por estas razones y basándonos en ejemplos tomados de la literatura, concebimos siete preguntas clave que caen en cuatro categorías: direccionalidad, clasificación y medida, cambios ecológicos o socio-económicos, y escala. Estas preguntas deberían ayudar en la formulación de definiciones claras y prácticas del impacto para encajar mejor con contextos científicos, de las partes interesadas o legislativos específicos.

Global range expansion of the Asian mussel Limnoperna fortunei (Mytilidae): Another fouling threat to freshwater systems

The Asian freshwater mussel Limnoperna fortunei was first documented as a major fouling pest when it colonized Hong Kongs water supply system in the late 1960s. It has since fouled municipal waterworks and power plant cooling systems in Korea, Japan, Taiwan, and most recently, in South America. Dense accumulations of byssally‐attached mussels obstruct flow in water conduits, causing impacts similar to those of the Eurasian zebra mussel (Dreissena polymorpha). Limnoperna has demonstrated potential for global range expansion through the oceanic transport of its planktonic larvae in ship ballast tanks. Therefore, unless effective controls are imposed upon ballast‐water transport, the mussel will continue to invade and impact aquatic systems on other continents. Given that shipping traffic from both Asia and South America has already resulted in recent introductions of exotic bivalves to the USA, a future North American invasion by L. fortunei is highly probable.

Advancing impact prediction and hypothesis testing in invasion ecology using a comparative functional response approach

Invasion ecology urgently requires predictive methodologies that can forecast the ecological impacts of existing, emerging and potential invasive species. We argue that many ecologically damaging invaders are characterised by their more efficient use of resources. Consequently, comparison of the classical ‘functional response’ (relationship between resource use and availability) between invasive and trophically analogous native species may allow prediction of invader ecological impact. We review the utility of species trait comparisons and the history and context of the use of functional responses in invasion ecology, then present our framework for the use of comparative functional responses. We show that functional response analyses, by describing the resource use of species over a range of resource availabilities, avoids many pitfalls of ‘snapshot’ assessments of resource use. Our framework demonstrates how comparisons of invader and native functional responses, within and between Type II and III functional responses, allow testing of the likely population-level outcomes of invasions for affected species. Furthermore, we describe how recent studies support the predictive capacity of this method; for example, the invasive ‘bloody red shrimp’ Hemimysis anomala shows higher Type II functional responses than native mysids and this corroborates, and could have predicted, actual invader impacts in the field. The comparative functional response method can also be used to examine differences in the impact of two or more invaders, two or more populations of the same invader, and the abiotic (e.g. temperature) and biotic (e.g. parasitism) context-dependencies of invader impacts. Our framework may also address the previous lack of rigour in testing major hypotheses in invasion ecology, such as the ‘enemy release’ and ‘biotic resistance’ hypotheses, as our approach explicitly considers demographic consequences for impacted resources, such as native and invasive prey species. We also identify potential challenges in the application of comparative functional responses in invasion ecology. These include incorporation of numerical responses, multiple predator effects and trait-mediated indirect interactions, replacement versus non-replacement study designs and the inclusion of functional responses in risk assessment frameworks. In future, the generation of sufficient case studies for a meta-analysis could test the overall hypothesis that comparative functional responses can indeed predict invasive species impacts.