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

Emergence of the metacommunity concept has made a substantial contribution to better understanding of the community composition and dynamics in a regional context. However, long-term field data for testing of available metacommunity models are still scarce, and the extent to which these models apply to the real world remains unknown. Tests conducted so far have largely sought to fit data on the entire regional set of species to one of several metacommunity models, implicitly assuming that all species operate similarly over the same set of sites. However, species differ in their habitat use. These differences can, in the most general terms, be expressed as a gradient of habitat specialization (ranging from habitat specialists to habitat generalists). We postulate that such differences in habitat specialization will have implications for metacommunity dynamics. Specifically, we predict that specialists respond more to local processes and generalists respond to regional spatial processes. We tested these predictions using natural microcosm communities for which long-term (nine-year) environmental and population dynamics data were available. We used redundancy analysis to determine the proportion of variation explained by environmental and spatial factors. We repeated this analysis to explain variation in the entire regional set of species, in generalist species only, and in specialists only. We further used ANOVA to test for differences in the proportions of explained variation. We found that habitat specialists responded primarily to environmental factors and habitat generalists responded mainly to spatial factors. Thus, from the metacommunity perspective, the dynamics of habitat specialists are best explained by a combination of species sorting and mass effects, while that of habitat generalists are best explained by patch dynamics and neutral models. Consequently, we infer that a natural metacommunity can exhibit complicated dynamics, with some groups of species (e.g., habitat specialists) governed according to environmental processes and other groups (e.g., habitat generalists) governed mainly by dispersal processes.

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Contrasts between habitat generalists and specialists: an empirical extension to the basic metacommunity framework

The classical view states that microbial biogeography is not affected by dispersal barriers or historical events, but only influenced by the local contemporary habitat conditions (species sorting). This has been challenged during recent years by studies suggesting that also regional factors such as mass effect, dispersal limitation and neutral assembly are important for the composition of local bacterial communities. Here we summarize results from biogeography studies in different environments, i.e. in marine, freshwater and soil as well in human hosts. Species sorting appears to be the most important mechanism. However, this result might be biased since this is the mechanism that is easiest to measure, detect and interpret. Hence, the importance of regional factors may have been underestimated. Moreover, our survey indicates that different assembly mechanisms might be important for different parts of the total community, differing, for example, between generalists and specialists, and between taxa of different dispersal ability and motility. We conclude that there is a clear need for experimental studies, first, to clearly separate regional and local factors in order to study their relative importance, and second, to test whether there are differences in assembly mechanisms depending on different taxonomic or functional groups.

Local and regional factors influencing bacterial community assembly

1. Temporary aquatic habitats often are inhabited by a unique fauna and flora and contribute significantly to regional diversity. Temporary wetlands around the world are disappearing rapidly. The individual and interacting impacts of factors influencing community structure and dynamics in temporary wetlands are not always well known. 2. Camargue wetlands are mainly characterized by variable salinity and hydroperiod. The individual and combined impacts of these local factors, together with regional variables, on invertebrate communities remain unknown. We therefore characterized and sampled invertebrates in 30 temporary wetlands along salinity and hydroperiod gradients in the Camargue (Southern France) 3, 5 and 7 months after inundation. 3. Over the three sampling occasions, a total of 17 cladoceran species and 49 macroinvertebrate taxa were identified. Hydroperiod and salinity were the most important variables explaining variation in taxonomic composition and can be considered key factors shaping the invertebrate communities in Camargue wetlands. The impact on taxon richness was significantly positive for hydroperiod but significantly negative for salinity. Regional factors had no significant effect on the structure of the studied invertebrate communities, suggesting that dispersal was not limiting and that species sorting was the most important structuring process. 4. The results of this study suggest that the combined and interacting effects of salinization and hydrological modification of Mediterranean temporary wetlands (due to water management, climate change, etc.) can result in reduced diversity in large numbers of Mediterranean wetlands and induce a considerable decline in regional diversity of aquatic invertebrates.

Invertebrate community patterns in Mediterranean temporary wetlands along hydroperiod and salinity gradients

Despite an upsurge of interest in spatial interactions between communities and in the impact of dispersal on ecological and evolutionary processes, dispersal patterns and dynamics in natural metacommunities remain poorly understood. Although passive aerial dispersal of freshwater invertebrates is generally accepted, the frequency and relative importance of wind as a vector is still subject of considerable debate. We assessed the importance of wind dispersal in an invertebrate metacommunity in a cluster of 36 temporary rock pools on an isolated mountaintop in South Africa. Wind dispersal was quantified every four days using nine windsocks (about 1.5 m above rock base), placed in the field during one month. Distance to the nearest pool varied from 2 up to 16 m. Wind direction and speed were monitored for the entire period. About 850 propagules (mostly resting eggs) of 17 taxa were captured. The presence of water in the pools (level of exposure of the dormant propagule bank) and the dominant wind direction were the key factors affecting the yield. Wind speed was much less important. Our results suggest that wind dispersal of propagules from temporary aquatic systems is more frequent than previously thought. This may stabilise the metacommunity by mediating gene flow among populations and facilitating rapid (re)colonisation of patches. On the other hand, wind erosion of the dormant propagule bank may lead to egg bank depletion and local extinction. The measured frequent wind dispersal most likely fuels strong species sorting processes ultimately shaping the structure of the local communities as observed in an earlier study. To elucidate the link between local dispersal rates and their contribution to long range dispersal is a major challenge for future research on aerial dispersal of aquatic invertebrates.

Any way the wind blows ‐ frequent wind dispersal drives species sorting in ephemeral aquatic communities

Explaining the variance of local communities in a spatial-environmental matrix is one of the core interests of ecology today. Recent progress in metacommunity theory has made a substantial contribution to this field, however good empirical data in support of available theories are still relatively scarce. In this study we sampled a cluster of 36 temporary rock pools four times during one season to assess invertebrate metacommunity structure and dynamics and to search for steering processes and variables. Both Mantel tests and redundancy models indicate that local abiotic factors were dominant over spatial factors in explaining community structure and both were acting independently. Spatial variables were only important for passive dispersers and significantly explained 11% of variation in this community component. Pools connected by temporary overflows hosted more similar communities of passive dispersers than unconnected ones while community dissimilarity significantly increased with inter-pool distance. A negative curvilinear relation was discovered between taxon richness and isolation in passive dispersers, providing some support for existing theoretical models of Mouquet and Loreau. Of different metacommunity perspectives, a combination of species sorting and mass effects best explains the observed patterns. Additionally, priority effects and monopolization may buffer against the homogenising effects of dispersal and contribute to the distinctness of isolated communities. This is one of the first studies to present evidence for spatial patterns in aquatic communities on such a small spatial scale (a rock ledge of 99000 m 2 ). Bridging the gap between theory and observed patterns in natural systems is one of the main challenges for future metacommunity research. Small aquatic habitats such as pitcher plants and freshwater rock pools may well have an important role to play as model systems to study ecological processes in a natural spatially explicit environment.

The role of metacommunity processes in shaping invertebrate rock pool communities along a dispersal gradient

Summary
1. Hydroregime (duration, frequency and predictability of the aquatic phase) is a key feature of temporary aquatic habitats that not only moulds community structure and diversity (species sorting) but also life history characteristics of the inhabitants (natural selection). However, since hydroregime is a complex multidimensional entity that cannot be estimated from short term observations, morphometric variables are commonly used as proxies for hydroregime, making it impossible to separate effects of habitat size and hydroregime on biota.

2. We have used a simple hydrological model, validated with recent (average r2 = 96%) and historic water level observations (average r2 = 81%), to accurately reconstruct hydroregime based on historical rainfall and evaporation data in a cluster of 36 temporary rock pools in central South Africa.

3. Using the model output, we demonstrated that both hydroregime and habitat size had unique and shared effects on temporary pool biota and that these effects depended on the dispersal modes of the taxa. Model-generated hydrological data explained more variation in community patterns than short-term observations of hydroperiod. Hydroregime was more important for passive dispersers than for active dispersers that can migrate when pools dry up. Basin morphometry was a relatively poor predictor of pool hydroregime. We concluded that simple modelling may greatly improve the resolution of studies linking hydroregime to biological variables.

4. An accurate account of hydrological variation provides a firm foundation to understand community and population structure and dynamics in temporary aquatic habitats. Since many of these habitats have been destroyed or degraded in recent decades, our findings and tools may contribute to the development of reliable conservation guidelines.

Chris De Vries

Papers

The role of metacommunity processes in shaping invertebrate rock pool communities along a dispersal gradient

Community structure in temporary freshwater pools: disentangling the effects of habitat size and hydroregime