Patrick Weigelt

Global exchange and accumulation of non-native plants

All around the globe, humans have greatly altered the abiotic and biotic environment with ever-increasing speed. One defining feature of the Anthropocene epoch is the erosion of biogeographical barriers by human-mediated dispersal of species into new regions, where they can naturalize and cause ecological, economic and social damage. So far, no comprehensive analysis of the global accumulation and exchange of alien plant species between continents has been performed, primarily because of a lack of data. Here we bridge this knowledge gap by using a unique global database on the occurrences of naturalized alien plant species in 481 mainland and 362 island regions. In total, 13,168 plant species, corresponding to 3.9% of the extant global vascular flora, or approximately the size of the native European flora, have become naturalized somewhere on the globe as a result of human activity. North America has accumulated the largest number of naturalized species, whereas the Pacific Islands show the fastest increase in species numbers with respect to their land area. Continents in the Northern Hemisphere have been the major donors of naturalized alien species to all other continents. Our results quantify for the first time the extent of plant naturalizations worldwide, and illustrate the urgent need for globally integrated efforts to control, manage and understand the spread of alien species.

No saturation in the accumulation of alien species worldwide

Although research on human-mediated exchanges of species has substantially intensified during the last centuries, we know surprisingly little about temporal dynamics of alien species accumulations across regions and taxa. Using a novel database of 45,813 first records of 16,926 established alien species, we show that the annual rate of first records worldwide has increased during the last 200 years, with 37% of all first records reported most recently (1970–2014). Inter-continental and inter-taxonomic variation can be largely attributed to the diaspora of European settlers in the nineteenth century and to the acceleration in trade in the twentieth century. For all taxonomic groups, the increase in numbers of alien species does not show any sign of saturation and most taxa even show increases in the rate of first records over time. This highlights that past efforts to mitigate invasions have not been effective enough to keep up with increasing globalization.

Bioclimatic and physical characterization of the world’s islands

The Earth’s islands harbor a distinct, yet highly threatened, biological and cultural diversity that has been shaped by geographic isolation and unique environments. Island systems are key natural laboratories for testing theory in ecology and evolution. However, despite their potential usefulness for research, a quantitative description of island environments and an environmental classification are still lacking. Here, we prepare a standardized dataset and perform a comprehensive global environmental characterization for 17,883 of the world’s marine islands >1 km2 (∼98% of total island area). We consider area, temperature, precipitation, seasonality in temperature and precipitation, past climate change velocity, elevation, isolation, and past connectivity—key island characteristics and drivers of ecosystem processes. We find that islands are significantly cooler, wetter, and less seasonal than mainlands. Constrained by their limited area, they show less elevational heterogeneity. Wet temperate climates are more prevalent on islands, whereas desert climates are comparatively rare. We use ordination and clustering to characterize islands in multidimensional environmental space and to delimit island ecoregions, which provides unique insights into the environmental configuration and diversity of the world’s islands. Combining ordination and classification together with global environmental data in a common framework opens up avenues for a more integrative use of islands in biogeography, macroecology, and conservation. To showcase possible applications of the presented data, we predict vascular plant species richness for all 17,883 islands based on statistically derived environment–richness relationships.

Late Quaternary climate change shapes island biodiversity

Island biogeographical models consider islands either as geologically static with biodiversity resulting from ecologically neutral immigration–extinction dynamics, or as geologically dynamic with biodiversity resulting from immigration–speciation–extinction dynamics influenced by changes in island characteristics over millions of years. Present climate and spatial arrangement of islands, however, are rather exceptional compared to most of the Late Quaternary, which is characterized by recurrent cooler and drier glacial periods. These climatic oscillations over short geological timescales strongly affected sea levels and caused massive changes in island area, isolation and connectivity, orders of magnitude faster than the geological processes of island formation, subsidence and erosion considered in island theory. Consequences of these oscillations for present biodiversity remain unassessed. Here we analyse the effects of present and Last Glacial Maximum (LGM) island area, isolation, elevation and climate on key components of angiosperm diversity on islands worldwide. We find that post-LGM changes in island characteristics, especially in area, have left a strong imprint on present diversity of endemic species. Specifically, the number and proportion of endemic species today is significantly higher on islands that were larger during the LGM. Native species richness, in turn, is mostly determined by present island characteristics. We conclude that an appreciation of Late Quaternary environmental change is essential to understand patterns of island endemism and its underlying evolutionary dynamics.

Oceanic island biogeography through the lens of the general dynamic model: Assessment and prospect

The general dynamic model of oceanic island biogeography (GDM) has added a new dimension to theoretical island biogeography in recognizing that geological processes are key drivers of the evolutionary processes of diversification and extinction within remote islands. It provides a dynamic and essentially non‐equilibrium framework generating novel predictions for emergent diversity properties of oceanic islands and archipelagos. Its publication in 2008 coincided with, and spurred on, renewed attention to the dynamics of remote islands. We review progress, both in testing the GDMs predictions and in developing and enhancing ecological–evolutionary understanding of oceanic island systems through the lens of the GDM. In particular, we focus on four main themes: (i) macroecological tests using a space‐for‐time rationale; (ii) extensions of theory to islands following different patterns of ontogeny; (iii) the implications of GDM dynamics for lineage diversification and trait evolution; and (iv) the potential for downscaling GDM dynamics to local‐scale ecological patterns and processes within islands. We also consider the implications of the GDM for understanding patterns of non‐native species diversity. We demonstrate the vitality of the field of island biogeography by identifying a range of potentially productive lines for future research.

Biogeographic, climatic and spatial drivers differentially affect α, β and γ diversities on oceanic archipelagos

Island biogeographic studies traditionally treat single islands as units of analysis. This ignores the fact that most islands are spatially nested within archipelagos. Here, we took a fundamentally different approach and focused on entire archipelagos using species richness of vascular plants on 23 archipelagos worldwide and their 174 constituent islands. We assessed differential effects of biogeographic factors (area, isolation, age, elevation), current and past climate (temperature, precipitation, seasonality, climate change velocity) and intra-archipelagic spatial structure (archipelago area, number of islands, area range, connectivity, environmental volume, inter-island distance) on plant diversity. Species diversity of each archipelago (γ) was additively partitioned into α, β, nestedness and replacement β-components to investigate the relative importance of environmental and spatial drivers. Multiple regressions revealed strong effects of biogeography and climate on α and γ, whereas spatial factors, particularly number of islands, inter-island distance and area range, were key to explain β. Structural equation models additionally suggested that γ is predominantly determined by indirect abiotic effects via its components, particularly β. This highlights that β and the spatial arrangement of islands are essential to understand insular ecology and evolution. Our methodological framework can be applied more widely to other taxa and archipelago-like systems, allowing new insights into biodiversity origin and maintenance.

Climate change will increase the naturalization risk from garden plants in Europe

Abstract Aim Plant invasions often follow initial introduction with a considerable delay. The current non‐native flora of a region may hence contain species that are not yet naturalized but may become so in the future, especially if climate change lifts limitations on species spread. In Europe, non‐native garden plants represent a huge pool of potential future invaders. Here, we evaluate the naturalization risk from this species pool and how it may change under a warmer climate. Location Europe. Methods We selected all species naturalized anywhere in the world but not yet in Europe from the set of non‐native European garden plants. For this subset of 783 species, we used species distribution models to assess their potential European ranges under different scenarios of climate change. Moreover, we defined geographical hotspots of naturalization risk from those species by combining projections of climatic suitability with maps of the area available for ornamental plant cultivation. Results Under current climate, 165 species would already find suitable conditions in > 5% of Europe. Although climate change substantially increases the potential range of many species, there are also some that are predicted to lose climatically suitable area under a changing climate, particularly species native to boreal and Mediterranean biomes. Overall, hotspots of naturalization risk defined by climatic suitability alone, or by a combination of climatic suitability and appropriate land cover, are projected to increase by up to 102% or 64%, respectively. Main conclusions Our results suggest that the risk of naturalization of European garden plants will increase with warming climate, and thus it is very likely that the risk of negative impacts from invasion by these plants will also grow. It is therefore crucial to increase awareness of the possibility of biological invasions among horticulturalists, particularly in the face of a warming climate.

Global patterns and drivers of phylogenetic structure in island floras

Islands are ideal for investigating processes that shape species assemblages because they are isolated and have discrete boundaries. Quantifying phylogenetic assemblage structure allows inferences about these processes, in particular dispersal, environmental filtering and in-situ speciation. Here, we link phylogenetic assemblage structure to island characteristics across 393 islands worldwide and 37,041 vascular plant species (representing angiosperms overall, palms and ferns). Physical and bioclimatic factors, especially those impeding colonization and promoting speciation, explained more variation in phylogenetic structure of angiosperms overall (49%) and palms (52%) than of ferns (18%). The relationships showed different or contrasting trends among these major plant groups, consistent with their dispersal- and speciation-related traits and climatic adaptations. Phylogenetic diversity was negatively related to isolation for palms, but unexpectedly it was positively related to isolation for angiosperms overall. This indicates strong dispersal filtering for the predominantly large-seeded, animal-dispersed palm family whereas colonization from biogeographically distinct source pools on remote islands likely drives the phylogenetic structure of angiosperm floras. We show that signatures of dispersal limitation, environmental filtering and in-situ speciation differ markedly among taxonomic groups on islands, which sheds light on the origin of insular plant diversity.

Plants capable of selfing are more likely to become naturalized.

Many plant species have established self-sustaining populations outside their natural range because of human activities. Plants with selfing ability should be more likely to establish outside their historical range because they can reproduce from a single individual when mates or pollinators are not available. Here, we compile a global breeding-system database of 1,752 angiosperm species and use phylogenetic generalized linear models and path analyses to test relationships between selfing ability, life history, native range size and global naturalization status. Selfing ability is associated with annual or biennial life history and a large native range, which both positively correlate with the probability of naturalization. Path analysis suggests that a high selfing ability directly increases the number of regions where a species is naturalized. Our results provide robust evidence across flowering plants at the global scale that high selfing ability fosters alien plant naturalization both directly and indirectly.

Species richness and biomass explain spatial turnover in ecosystem functioning across tropical and temperate ecosystems

Predicting ecosystem functioning at large spatial scales rests on our ability to scale up from local plots to landscapes, but this is highly contingent on our understanding of how functioning varies through space. Such an understanding has been hampered by a strong experimental focus of biodiversity–ecosystem functioning research restricted to small spatial scales. To address this limitation, we investigate the drivers of spatial variation in multitrophic energy flux—a measure of ecosystem functioning in complex communities—at the landscape scale. We use a structural equation modelling framework based on distance matrices to test how spatial and environmental distances drive variation in community energy flux via four mechanisms: species composition, species richness, niche complementarity and biomass. We found that in both a tropical and a temperate study region, geographical and environmental distance indirectly influence species richness and biomass, with clear evidence that these are the dominant mechanisms explaining variability in community energy flux over spatial and environmental gradients. Our results reveal that species composition and trait variability may become redundant in predicting ecosystem functioning at the landscape scale. Instead, we demonstrate that species richness and total biomass may best predict rates of ecosystem functioning at larger spatial scales.