Study of the impacts of biological invasions, a pervasive component of global change, has generated remarkable understanding of the mechanisms and consequences of the spread of introduced populations. The growing field of invasion science, poised at a crossroads where ecology, social sciences, resource management, and public perception meet, is increasingly exposed to critical scrutiny from several perspectives. Although the rate of biological invasions, elucidation of their consequences, and knowledge about mitigation are growing rapidly, the very need for invasion science is disputed. Here, we highlight recent progress in understanding invasion impacts and management, and discuss the challenges that the discipline faces in its science and interactions with society.

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Impacts of biological invasions: what's what and the way forward

Human-mediated species invasions are a significant component of current global environmental change. There is every indication that the rate at which locations are accumulating non-native species is accelerating as free trade and globalization advance. Thus, the need to incorporate predictive models in the assessment of invasion risk has become acute. However, finding elements of the invasion process that provide consistent explanatory power has proved elusive. Here, we propose propagule pressure as a key element to understanding why some introduced populations fail to establish whereas others succeed. In the process, we illustrate how the study of propagule pressure can provide an opportunity to tie together disparate research agendas within invasion ecology.

https://www.cell.com/trends/ecology-evolution/pdf/S0169-5347(05)00032-7.pdf

The role of propagule pressure in explaining species invasions.

Study of interactions between pairs or larger groups of nonindigenous species has been subordinated in the literature to study of interactions between nonindigenous and native species. To the extent that interactions among introduced species are depicted at all, the emphasis has been on negative interactions, primarily resource competition and interference. However, a literature search reveals that introduced species frequently interact with one another and that facilitative interactions are at least as common as detrimental ones. The population significance of these interactions has rarely been determined, but a great variety of types of direct and indirect interactions among individuals of different nonindigenous species is observed, and many are plausibly believed to have consequences at the population level. In particular, mutualisms between plants and the animals that disperse and/or pollinate them and modification of habitat by both animals and plants seem common and often important in facilitating invasions. There is little evidence that interference among introduced species at levels currently observed significantly impedes further invasions, and synergistic interactions among invaders may well lead to accelerated impacts on native ecosystems ‐ an invasional ‘meltdown’ process.

Positive interactions of nonindigenous species: invasional meltdown?

Humans now strongly influence almost every major aquatic ecosystem, and their activities have dramatically altered the fluxes of growth-limiting nutrients from the landscape to receiving waters. Unfortunately, these nutrient inputs have had profound negative effects upon the quality of surface waters worldwide. This review examines how eutrophication influences the biomass and species composition of algae in both freshwater and costal marine systems. An overview of recent advances in algae-related eutrophication research is presented. In freshwater systems, a summary is presented for lakes and reservoirs; streams and rivers; and wetlands. A brief summary is also presented for estuarine and coastal marine ecosystems. Eutrophication causes predictable increases in the biomass of algae in lakes and reservoirs; streams and rivers; wetlands; and coastal marine ecosystems. As in lakes, the response of suspended algae in large rivers to changes in nutrient loading may be hysteretic in some cases. The inhibitory effects of high concentrations of inorganic suspended solids on algal growth, which can be very evident in many reservoirs receiving high inputs of suspended soils, also potentially may occur in turbid rivers. Consistent and predictable eutrophication-caused increases in cyanobacterial dominance of phytoplankton have been reported worldwide for natural lakes, and similar trends are reported here both for phytoplankton in turbid reservoirs, and for suspended algae in a large river. A remarkable unity is evident in the global response of algal biomass to nitrogen and phosphorus availability in lakes and reservoirs; wetlands; streams and rivers; and coastal marine waters. The species composition of algal communities inhabiting the water column appears to respond similarly to nutrient loading, whether in lakes, reservoirs, or rivers. As is true of freshwater ecosystems, the recent literature suggests that coastal marine ecosystems will respond positively to nutrient loading control efforts. Our understanding of freshwater eutrophication and its effects on algal-related water quality is strong and is advancing rapidly. However, our understanding of the effects of eutrophication on estuarine and coastal marine ecosystems is much more limited, and this gap represents an important future research need. Although coastal systems can be hydrologically complex, the biomass of marine phytoplankton nonetheless appears to respond sensitively and predictably to changes in the external supplies of nitrogen and phosphorus. These responses suggest that efforts to manage nutrient inputs to the seas will result in significant improvements in coastal zone water quality. Additional new efforts should be made to develop models that quantitatively link ecosystem-level responses to nutrient loading in both freshwater and marine systems.

http://globalpnetwork.net/sites/default/files/Smith_2003_ESPR_eutrophication_review.pdf

Eutrophication of freshwater and coastal marine ecosystems a global problem

There has been a dramatic growth in research on biological invasions over the past 20 years, but a mature understanding of the field has been hampered because invasion biologists concerned with different taxa and different environments have largely adopted different model frameworks for the invasion process, resulting in a confusing range of concepts, terms and definitions. In this review, we propose a unified framework for biological invasions that reconciles and integrates the key features of the most commonly used invasion frameworks into a single conceptual model that can be applied to all human-mediated invasions. The unified framework combines previous stage-based and barrier models, and provides a terminology and categorisation for populations at different points in the invasion process.

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A proposed unified framework for biological invasions.

A recent trend in invasion ecology relates the success of non-indigenous species (NIS) to reduced control by enemies such as pathogens, parasites and predators (i.e. the enemy release hypothesis, ERH). Despite the demonstrated importance of enemies to host population dynamics, studies of the ERH are split – biogeographical analyses primarily show a reduction in the diversity of enemies in the introduced range compared with the native range, while community studies imply that NIS are no less affected by enemies than native species in the invaded community. A broad review of the invasion literature implies at least eight non-exclusive explanations for this enigma. In addition, we argue that the ERH has often been accepted uncritically wherever (i) NIS often appear larger, more fecund, or somehow ‘better’ than either congeners in the introduced region, or conspecifics in the native range; and (ii) known enemies are conspicuously absent from the introduced range. However, all NIS, regardless of their abundance or impact, will lose natural enemies at a biogeographical scale. Given the complexity of processes that underlie biological invasions, we argue against a simple relationship between enemy ‘release’ and the vigour, abundance or impact of NIS.

Is invasion success explained by the enemy release hypothesis

The use of simple terms to articulate ecological concepts can confuse ideological debates and undermine management efforts. This problem is particularly acute in studies of nonindigenous species, which alternatively have been called ‘exotic’, ‘introduced’, ‘invasive’ and ‘naturalised’, among others. Attempts to redefine commonly used terminology have proven difficult because authors are often partial to particular definitions. In an attempt to form a consensus on invasion terminology, we synthesize an invasional framework based on current models that break the invasion process into a series of consecutive, obligatory stages. Unlike previous efforts, we propose a neutral terminology based on this framework. This ‘stagebased’ terminology can be used to supplement terms with ambiguous meanings (e.g. invasive, introduced, naturalized, weedy, etc.), and thereby improve clarity of future studies. This approach is based on the concept of ‘propagule pressure’ and has the additional benefit of identifying factors affecting the success of species at each stage. Under this framework, invasions can be more objectively understood as biogeographical, rather than taxonomic, phenomena; and author preferences in the use of existing terminology can be addressed. An example of this recommended protocol might be: ‘We examined distribution data to contrast the characteristics of invasive species (stages IVa and V) and noninvasive species (stages III and IVb)’.

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A neutral terminology to define 'invasive' species

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.

https://link.springer.com/content/pdf/10.1007%2Fs10530-006-9007-7.pdf

Propagule pressure: a null model for biological invasions

The North American Great Lakes have been invaded and dramatically altered by more than 145 alien species. Many invasions have occurred during the past few decades because of the release of Eurasian ballast water from transoceanic ships. Current regulations require ships to exchange foreign ballast with highly saline water before entering the Great Lakes; this procedure should prevent colonization by strictly freshwater species, but species with broad salinity tolerance might survive transport in exchanged water. A recent series of invasions by euryhaline organisms from the Black and Caspian Seas region signals a new phase in the transformation of the Great Lakes - one that supports the concept of an 'invasional meltdown'.

https://www.cell.com/trends/ecology-evolution/pdf/S0169-5347(99)01745-0.pdf

Recent mass invasion of the North American Great Lakes by Ponto-Caspian species.

SYNOPSIS. The expansion of zebra mussel distribution into inland waterways of North America portends significant abiotic and biotic changes mediated either directly or indirectly by Dreissena . Dreissena fouls a wide array of submerged substrates including rock surfaces, macrophytes, native molluscs, canal and dock walls, and watercraft and motor outdrives. Fouling of water intake pipes and associated installations can severely impair water delivery to hydroelectric, municipal and industrial users, necessitating proactive or reactive control measures. Mussels increase water clarity by removing suspended clay, silt, bacteria, phytoplankton, and small zooplankton. Clear water phases associated with Dreissena grazing may exceed in magnitude and duration those generated by zooplankton grazing. Enhanced water clarity increases light transmittance and growth of benthic plants. Some benthic invertebrates {e.g., unionid molluscs) are adversely affected by Dreissena , whereas others, including amphipod crustaceans, exploit structure associated with or wastes generated by zebra mussels. Dreissena is exploited by a host of predators, most notably waterfowl, fish and crayfish. Waterfowl predators that consume contaminated Dreissena have elevated concentrations of organic pesticides and polychlorinated biphenyl compounds. Invasion of shallow lakes and ponds by Dreissena may divert production and biomass from pelagic to benthic foodwebs, shifting ecosystems to an alternative state.

Hugh J. MacIsaac

Papers

Is invasion success explained by the enemy release hypothesis

A neutral terminology to define 'invasive' species

Propagule pressure: a null model for biological invasions

Recent mass invasion of the North American Great Lakes by Ponto-Caspian species.

Potential Abiotic and Biotic Impacts of Zebra Mussels on the Inland Waters of North America