Hugh J. MacIsaac

Propagule Pressure: A Null Model for Biological Invasions

Invasion ecology has been criticised for its lack of general principles. To explore this criticism, we conducted a meta-analysis that examined characteristics of invasiveness (i.e. the ability of species to establish in, spread to, or become abundant in novel communities) and invasibility (i.e. the susceptibility of habitats to the establishment or proliferation of invaders). There were few consistencies among invasiveness characteristics (3 of 13): established and abundant invaders generally occupy similar habitats as native species, while abundant species tend to be less affected by enemies; germination success and reproductive output were significantly positively associated with invasiveness when results from both stages (establishment/spread and abundance/impact) were combined. Two of six invasibility characteristics were also significant: communities experiencing more disturbance and with higher resource availability sustained greater establishment and proliferation of invaders. We also found that even though ‘propagule pressure’ was considered in only ~29% of studies, it was a significant predictor of both invasiveness and invasibility (55 of 64 total cases). Given that nonindigenous species are likely introduced non-randomly, we contend that ‘propagule biases’ may confound current paradigms in invasion ecology. Examples of patterns that could be confounded by propagule biases include characteristics of good invaders and susceptible habitats, release from enemies, evolution of ‘invasiveness’, and invasional meltdown. We conclude that propagule pressure should serve as the basis of a null model for studies of biological invasions when inferring process from patterns of invasion.Invasion ecology has been criticised for its lack of general principles. To explore this criticism, we conducted a meta-analysis that examined characteristics of invasiveness (i.e. the ability of species to establish in, spread to, or become abundant in novel communities) and invasibility (i.e. the susceptibility of habitats to the establishment or proliferation of invaders). There were few consistencies among invasiveness characteristics (3 of 13): established and abundant invaders generally occupy similar habitats as native species, while abundant species tend to be less affected by enemies; germination success and reproductive output were significantly positively associated with invasiveness when results from both stages (establishment/spread and abundance/impact) were combined. Two of six invasibility characteristics were also significant: communities experiencing more disturbance and with higher resource availability sustained greater establishment and proliferation of invaders. We also found that even though ‘propagule pressure’ was considered in only ~29% of studies, it was a significant predictor of both invasiveness and invasibility (55 of 64 total cases). Given that nonindigenous species are likely introduced non-randomly, we contend that ‘propagule biases’ may confound current paradigms in invasion ecology. Examples of patterns that could be confounded by propagule biases include characteristics of good invaders and susceptible habitats, release from enemies, evolution of ‘invasiveness’, and invasional meltdown. We conclude that propagule pressure should serve as the basis of a null model for studies of biological invasions when inferring process from patterns of invasion.

Biological invasions: recommendations for U.S. policy and management.

The Ecological Society of America has evaluated current U.S. national policies and practices on biological invasions in light of current scientific knowledge. Invasions by harmful nonnative species are increasing in number and area affected; the damages to ecosystems, economic activity, and human welfare are accumulating. Without improved strategies based on recent scientific advances and increased investments to counter invasions, harm from invasive species is likely to accelerate. Federal leadership, with the cooperation of state and local governments, is required to increase the effectiveness of prevention of invasions, detect and respond quickly to new potentially harmful invasions, control and slow the spread of existing invasions, and provide a national center to ensure that these efforts are coordinated and cost effective. Specifically, the Ecological Society of America recommends that the federal government take the following six actions: (1) Use new information and practices to better manage commercial and other pathways to reduce the transport and release of potentially harmful species; (2) Adopt more quantitative procedures for risk analysis and apply them to every species proposed for importation into the country; (3) Use new cost-effective diagnostic technologies to increase active surveillance and sharing of information about invasive species so that responses to new invasions can be more rapid and effective; (4) Create new legal authority and provide emergency funding to support rapid responses to emerging invasions; (5) Provide funding and incentives for cost-effective programs to slow the spread of existing invasive species in order to protect still uninvaded ecosystems, social and industrial infrastructure, and human welfare; and (6) Establish a National Center for Invasive Species Management (under the existing National Invasive Species Council) to coordinate and lead improvements in federal, state, and international policies on invasive species. Recent scientific and technical advances provide a sound basis for more cost-effective national responses to invasive species. Greater investments in improved technology and management practices would be more than repaid by reduced damages from current and future invasive species. The Ecological Society of America is committed to assist all levels of government and provide scientific advice to improve all aspects of invasive-species management.

Characterised and projected costs of nonindigenous species in Canada

Biological invasions by nonindigenous species (NIS) can have adverse effects on economically important goods and services, and sometimes result in an ‘invisible tax’ on natural resources (e.g. reduced yield). The combined economic costs of NIS may be significant, with implications for environmental policy and resource management; yet economic impact assessments are rare at a national scale. Impacts of nuisance NIS may be direct (e.g. loss of hardwood trees) or indirect (e.g. alteration of ecosystem services provided by growing hardwoods). Moreover, costs associated with these effects may be accrued to resources and services with clear ‘market’ values (e.g. crop production) and to those with more ambiguous, ‘non-market’ values (e.g. aesthetic value of intact forest). We characterised and projected economic costs associated with nuisance NIS in Canada, through a combination of case-studies and an empirical model derived from 21 identified effects of 16 NIS. Despite a severe dearth of available data, characterised costs associated with ten NIS in Canadian fisheries, agriculture and forestry totalled

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

187 million Canadian (CDN) per year. These costs were dwarfed by the ‘invisible tax’ projected for sixteen nuisance NIS found in Canada, which was estimated at between

Popularity and propagule pressure: determinants of introduction and establishment of aquarium fish

13.3 and

Identifying the source of species invasions: sampling intensity vs. genetic diversity

34.5 billion CDN per year. Canada remains highly vulnerable to new nuisance NIS, but available manpower and financial resources appear insufficient to deal with this problem.

Invasion risks posed by the aquarium trade and live fish markets on 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.

Regulation of Zooplankton Community Structure of an Acidified Lake by Chaoborus

Propagule pressure is frequently cited as an important determinant of invasion success for terrestrial taxa, but its importance for aquatic species is unclear. Using data on aquarium fishes in stores and historical records of fish introduced and established in Canadian and United States waters, we show clear relationships exist between frequency of occurrence in shops and likelihood of introduction and of establishment. Introduced and established taxa are also typically larger than those available from stores, consistent with the propagule pressure hypothesis in that larger fish may be released more frequently due to outgrowing their aquaria. Attempts to reduce the numbers of introductions may be the most practical mechanism to reduce the number of new successful invasions.

Invasion genetics of the Ciona intestinalis species complex: from regional endemism to global homogeneity.

Population geneticists and community ecologists have long recognized the importance of sampling design for uncovering patterns of diversity within and among populations and in communities. Invasion ecologists increasingly have utilized phylogeographical patterns of mitochondrial or chloroplast DNA sequence variation to link introduced populations with putative source populations. However, many studies have ignored lessons from population genetics and community ecology and are vulnerable to sampling errors owing to insufficient field collections. A review of published invasion studies that utilized mitochondrial or chloroplast DNA markers reveals that insufficient sampling could strongly influence results and interpretations. Sixty per cent of studies sampled an average of less than six individuals per source population, vs. only 45% for introduced populations. Typically, far fewer introduced than source populations were surveyed, although they were sampled more intensively. Simulations based on published data forming a comprehensive mtDNA haplotype data set highlight and quantify the impact of the number of individuals surveyed per source population and number of putative source populations surveyed for accurate assignment of introduced individuals. Errors associated with sampling a low number of individuals are most acute when rare source haplotypes are dominant or fixed in the introduced population. Accuracy of assignment of introduced individuals is also directly related to the number of source populations surveyed and to the degree of genetic differentiation among them (FST). Incorrect interpretations resulting from sampling errors can be avoided if sampling design is considered before field collections are made.

Contrasting patterns in genetic diversity following multiple invasions of fresh and brackish waters.

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.