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

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

A major aim in ecology is identifying determinants of invasiveness. We performed a meta-analysis of 117 field or experimental-garden studies that measured pair-wise trait differences of a total of 125 invasive and 196 non-invasive plant species in the invasive range of the invasive species. We tested whether invasiveness is associated with performance-related traits (physiology, leaf-area allocation, shoot allocation, growth rate, size and fitness), and whether such associations depend on type of study and on biogeographical or biological factors. Overall, invasive species had significantly higher values than non-invasive species for all six trait categories. More trait differences were significant for invasive vs. native comparisons than for invasive vs. non-invasive alien comparisons. Moreover, for comparisons between invasive species and native species that themselves are invasive elsewhere, no trait differences were significant. Differences in physiology and growth rate were larger in tropical regions than in temperate regions. Trait differences did not depend on whether the invasive alien species originates from Europe, nor did they depend on the test environment. We conclude that invasive alien species had higher values for those traits related to performance than non-invasive species. This suggests that it might become possible to predict future plant invasions from species traits.

/pdf/a-meta-analysis-of-trait-differences-between-invasive-and-29xa130ydb.pdf

A meta‐analysis of trait differences between invasive and non‐invasive plant species

Summary 1 Once neglected, the role of facilitative interactions in plant communities has received considerable attention in the last two decades, and is now widely recognized It is timely to consider the progress made by research in this field 2 We review the development of plant facilitation research, focusing on the history of the field, the relationship between plant‐plant interactions and environmental severity gradients, and attempts to integrate facilitation into mainstream ecological theory We then consider future directions for facilitation research 3 With respect to our fundamental understanding of plant facilitation, clarification of the relationship between interactions and environmental gradients is central for further progress, and necessitates the design and implementation of experiments that move beyond the clear limitations of previous studies 4 There is substantial scope for exploring indirect facilitative effects in plant communities, including their impacts on diversity and evolution, and future studies should connect the degree of non-transitivity in plant competitive networks to community diversity and facilitative promotion of species coexistence, and explore how the role of indirect facilitation varies with environmental severity 5 Certain ecological modelling approaches (eg individual-based modelling), although thus far largely neglected, provide highly useful tools for exploring these fundamental processes 6 Evolutionary responses might result from facilitative interactions, and consideration of facilitation might lead to re-assessment of the evolution of plant growth forms

/pdf/facilitation-in-plant-communities-the-past-the-present-and-572accn7oh.pdf

Facilitation in plant communities: the past, the present, and the future

Biotic resistance describes the ability of resident species in a community to reduce the success of exotic invasions. Although resistance is a well-accepted phenomenon, less clear are the processes that contribute most to it, and whether those processes are strong enough to completely repel invaders. Current perceptions of strong, competition-driven biotic resistance stem from classic ecological theory, Elton’s formulation of ecological resistance, and the general acceptance of the enemies-release hypothesis. We conducted a meta-analysis of the plant invasions literature to quantify the contribution of resident competitors, diversity, herbivores and soil fungal communities to biotic resistance. Results indicated large negative effects of all factors except fungal communities on invader establishment and performance. Contrary to predictions derived from the natural enemies hypothesis, resident herbivores reduced invasion success as effectively as resident competitors. Although biotic resistance significantly reduced the establishment of individual invaders, we found little evidence that species interactions completely repelled invasions. We conclude that ecological interactions rarely enable communities to resist invasion, but instead constrain the abundance of invasive species once they have successfully established.

https://onlinelibrary.wiley.com/doi/pdf/10.1111/j.1461-0248.2004.00657.x

A meta‐analysis of biotic resistance to exotic plant 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

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

We describe, explain, and "predict" dispersal and ecosystem impacts of six Ponto-Caspian endemic species that recently invaded the Great Lakes via ballast water. The zebra mussel, Dreissena polymorpha, and quagga mussel, Dreissena bugensis, continue to colonize hard and soft substrates of the Great Lakes and are changing ecosystem function through mechanisms of ecosystem engineering (increased water clarity and reef building), fouling native mussels, high particle filtration rate with selective rejection of colonial cyanobacteria in pseudofeces, alteration of nutrient ratios, and facilitation of the rapid spread of their Ponto-Caspian associates, the benthic amphipod Echinogammarus ischnusand the round goby, Neogobius melanostomus , which feeds on zebra mussels. The tubenose goby, Proterorhinus marmoratus , which does not feed on zebra mussels, has not spread rapidly. Impacts of these benthic invaders vary with site: in some shallow areas, habitat changes and the Dreissena → round goby → piscivore food chain have improved conditions for certain native game fishes and waterfowl; in offshore waters, Dreissena is competing for settling algae with the native amphipod Diporeia spp., which are disappearing to the detriment of the native deep-water fish community. The predatory cladoceran Cercopagis pengoi may compete with small fishes for zooplankton and increase food-chain length.

/pdf/dispersal-and-emerging-ecological-impacts-of-ponto-caspian-joc0llwjvd.pdf

Dispersal and emerging ecological impacts of Ponto-Caspian species in the Laurentian Great Lakes.

Since completion of the St. Lawrence Seaway in 1959, at least 43 nonindigenous species (NIS) of animals and protists have established in the Laurentian Great Lakes, of which ~67% were attributed to discharge of ballast water from commercial ships. Twenty-three NIS were first discovered in four "hotspot" areas with a high representation of NIS, most notably the Lake Huron - Lake Erie corridor. Despite implementation of the voluntary (1989, Canada) and mandatory (1993, U.S.A.) ballast water exchange (BWE) regulations, NIS were discovered at a higher rate during the 1990s than in the preceding three decades. Here we integrate knowledge of species' invasion histories, shipping traffic patterns, and physicochemical factors that constrain species' survivorship during ballast-mediated transfer to assess the risk of future introductions to the Great Lakes. Our risk-assessment model identified 26 high-risk species that are likely to survive intercontinental transfer in ballast tanks. Of these, 10 species have already invaded the Great Lakes. An additional 37 lower-risk species, of which six have already invaded, show some but not all attributes needed for successful introduction under current BWE management. Our model indicates that the Great Lakes remain vulnera- ble to ship-mediated NIS invasions.

/pdf/ballast-mediated-animal-introductions-in-the-laurentian-8r1joabs98.pdf

Ballast-mediated animal introductions in the Laurentian Great Lakes: retrospective and prospective analyses

Cercopagis pengoi, a waterflea native to the Ponto-Caspian region, was discovered during 1998 in Lake Ontario. Individuals were found throughout the lake during summer snagged on sportfishing lines...

Igor A. Grigorovich

Papers

Is invasion success explained by the enemy release hypothesis

Propagule pressure: a null model for biological invasions

Dispersal and emerging ecological impacts of Ponto-Caspian species in the Laurentian Great Lakes.

Ballast-mediated animal introductions in the Laurentian Great Lakes: retrospective and prospective analyses

Invasion of Lake Ontario by the Ponto–Caspian predatory cladoceran Cercopagis pengoi