Communities comprising alien species with different residence times are natural experiments allowing the assessment of drivers of community assembly over time. Stochastic processes (such as dispersal and fluctuating environments) should be the dominant factors structuring communities of exotic species with short residence times. In contrast, communities should become more similar, or systematically diverge, if they contain exotics with increasing resident times, due to the increasing importance of deterministic processes (such as environmental filtering). We use zeta diversity (the number of species shared by multiple assemblages) to explore the relationship between the turnover of native species and two categories of alien species with different residence times (archaeophytes [introduced between 4000 BC and 1500 AD] and neophytes [introduced after 1500 AD]) in a network of nature reserves in central Europe. By considering multiple assemblages simultaneously, zeta diversity allows us to determine the contribution of rare and widespread species to turnover. Specifically, we explore the relative effects of assembly processes representing isolation by distance, environmental filtering, and environmental stochasticity (fluctuating environments) on zeta diversity using Multi-Site Generalized Dissimilarity Modelling (MS-GDM). Four clusters of results emerged. First, stochastic processes for structuring plant assemblages decreased in importance with increasing residence time. Environmental stochasticity only affected species composition for neophytes, offering possibilities to predict the spread debt of recent invasions. Second, native species turnover was well explained by environmental filtering and isolation by distance, although these factors did not explain the turnover of archaeophytes and neophytes. Third, native and alien species compositions were only correlated for rare species, whereas turnover in widespread alien species was surprisingly unrelated to the composition of widespread native species. Site-specific approaches would therefore be more appropriate for the monitoring and management of rare alien species, whereas species-specific approaches would suit widespread species. Finally, the size difference of nature reserves influences not only native species richness, but also their richness-independent turnover. A network of reserves must therefore be designed and managed using a variety of approaches to enhance native diversity, while controlling alien species with different residence times and degrees of commonness.

Department of Botany and Zoology Centre for Invasion Biology Stellenbosch University Matieland 7600 South Africa

Department of Ecosystem Biology Faculty of Science University of South Bohemia Branišovská 1760 CZ 370 05 České Budějovice Czech Republic

Department of Invasion Ecology The Czech Academy of Sciences Institute of Botany CZ 252 43 Průhonice Czech Republic Department of Ecology Faculty of Science Charles University Viničná 7 CZ 128 44 Praha2 Czech Republic

Department of Mathematical Sciences Centre for Invasion Biology Stellenbosch University Matieland 7600 South Africa Department of Botany and Zoology Centre for Invasion Biology Stellenbosch University Matieland 7600 South Africa

Department of Mathematical Sciences Centre for Invasion Biology Stellenbosch University Matieland 7600 South Africa Theoretical Ecology Group African Institute for Mathematical Sciences Cape Town 7945 South Africa

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