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Ornamental horticulture is the most important pathway for alien plant introductions worldwide, and consequently, invasive spread of introduced plants often begins in urban areas. Although most introduced ornamental garden-plant species are locally not naturalized yet, many of them have shown invasion potential elsewhere in the world, and might naturalize when climate changes. We inventoried the planted flora of 50 public and 61 private gardens in Radolfzell, a small city in southern Germany, to investigate whether local naturalization success of garden plants is associated with their current planting frequency, climatic suitability (as assessed with climatic niche modelling) and known naturalization status somewhere in the world. We identified 954 introduced garden-plant species, of which 48 are already naturalized in Radolfzell and 120 in other parts of Germany. All currently naturalized garden plants in Radolfzell have a climatic suitability probability of ≥ 0.75 and are naturalized in ≥ 13 out of 843 regions globally. These values are significantly higher than those of garden plants that have not become locally naturalized yet. Current planting frequencies, however, were not related to current naturalization success. Using the identified local naturalization thresholds of climatic suitability and global naturalization frequency, and climate projections for the years 2050 and 2070, we identified 45 garden-plant species that are currently not naturalized in Radolfzell but are likely to become so in the future. Although our approach cannot replace a full risk assessment, it is well-suited and applicable as one element of a screening or horizon scanning-type approach. published

Aim: The general dynamic model (GDM) proposed by Whittaker et al. (2008) is a widely accepted theoretical framework in island biogeography. In this study, we explore whether GDM predictions hold when overall plant diversity is deconstructed into major plant families.Location: 101 islands from 14 oceanic archipelagos worldwide.Methods: Occurrence data for all species of nine large, cosmopolitan flowering plant families were used to test predictions derived from the GDM. We analyzed the effects of island area and age on species richness as well as number and percentage of single-island endemic species per family using mixed-effect models. Results: Total species and endemic richness as well as the percentage of endemic species showed a hump-shaped relationship with island age. The overall pattern was mainly driven by few species-rich plant families. Varying patterns were found for individual families, some of them opposing the general trend. In most cases, native and endemic species richness peaked much earlier in the island life cycle than suggested by the GDM. The contribution of area to the explained variation of all dependent variables was much higher than that of island age.Main conclusions: The results suggest that biodiversity-age relationships reported for large taxonomic groups like plants are driven by only a few species rich families. The way these families influence the overall patterns of species richness and endemism is related to family-specific properties, such as evolutionary history or dispersal strategies. Deviances from the GDM predictions can hence largely be explained by family characteristics and considering taxon-specific traits may help to further improve the explanatory power of the GDM. Families not following the expectations, like Orchidaceae, may be particularly valuable candidates to unveil so far neglected drivers of island biogeographical patterns.

Aims: The 50th anniversary of the publication of the seminal book, The Theory of Island Biogeography, by Robert H. MacArthur and Edward O. Wilson, is a timely moment to review and identify key research foci that could advance island biology. Here, we take a collaborative horizon-scanning approach to identify 50 fundamental questions for the continued development of the field. Location: Worldwide. Methods: We adapted a well-established methodology of horizon scanning to identify priority research questions in island biology, and initiated it during the Island Biology 2016 conference held in the Azores. A multidisciplinary working group prepared an initial pool of 187 questions. A series of online surveys was then used to refine a list of the 50 top priority questions. The final shortlist was restricted to questions with a broad conceptual scope, and which should be answerable through achievable research approaches. Results: Questions were structured around four broad and partially overlapping island topics, including: (Macro)Ecology and Biogeography, (Macro)Evolution, Community Ecology, and Conservation and Management. These topics were then subdivided according to the following subject areas: global diversity patterns (five questions in total); island ontogeny and past climate change (4); island rules and syndromes (3); island biogeography theory (4); immigration–speciation–extinction dynamics (5); speciation and diversification (4); dispersal and colonization (3); community assembly (6); biotic interactions (2); global change (5); conservation and management policies (5); and invasive alien species (4). Main conclusions: Collectively, this cross-disciplinary set of topics covering the 50 fundamental questions has the potential to stimulate and guide future research in island biology. By covering fields ranging from biogeography, community ecology and evolution to global change, this horizon scan may help to foster the formation of interdisciplinary research networks, enhancing joint efforts to better understand the past, present and future of island biotas., B.C.E. was supported by Spanish grant CGL2013-42589-P awarded to B.C.E. by the MINECO and co-financed by FEDER, and Spanish grant S20141203_002597 awarded to B.C.E. by the Organismo Autónomo Parques Nacionales of the MAGRAMA.

Aim Although the increase in species richness with increasing area is considered one of the few laws in ecology, the role of environmental and taxon-specific features in shaping species–area relationships (SARs) remains controversial. Using 421 land-plant floras covering continents, continental islands and oceanic islands, we investigate whether variations in SAR parameters can be interpreted in terms of differences among lineages in speciation mode and dispersal capacities (TAXON), or of geological history and geographical isolation between continents and islands (GEO). Location Global. Methods Linear mixed-effects models describing variation in SARs, depending on the factors GEO and TAXON and controlling for differences between realms (REALM) and biomes (BIOME). Results The best random-effect structure included both random slopes and random intercepts for GEO, TAXON, REALM and BIOME. This accounted for 77% of the total variation in species richness, substantially more than the 27% statistically explained by the model with fixed effects only (i.e. the simple SAR). The slopes of the SARs were higher for oceanic islands than for continental islands and continents, and higher in spermatophytes than in pteridophytes and bryophytes. The intercepts largely exhibited the reverse trend. TAXON was included in best-fit models restricted to oceanic and continental islands, but not continents. Analysing each plant lineage separately, the intercept of GEO was only included in the random structure of spermatophytes. Main conclusions SAR parameters varied considerably depending on geological history and taxon-specific traits. Such differences in SARs among land plants challenge the neutral theory that the accumulation of species richness on islands is controlled exclusively by extrinsic factors. Taxon-specific differences in SARs were, however, confounded by interactions with geological history and geographical isolation. This highlights the importance of applying integrative frameworks that take both environmental context and taxonomic idiosyncrasies into account in SAR analyses.

Recent years have seen an explosion in the availability of biodiversity data describing the distribution, function, and evolutionary history of life on earth. Integrating these heterogeneous data remains a challenge due to large variations in observational scales, collection purposes, and terminologies. Here, we conceptualize widely used biodiversity data types according to their domain (what aspect of biodiversity is described?) and informational resolution (how specific is the description?). Applying this framework to major data providers in biodiversity research reveals a strong focus on the disaggregated end of the data spectrum, whereas aggregated data types remain largely underutilized. We discuss the implications of this imbalance for the scope and representativeness of current macroecological research and highlight the synergies arising from a tighter integration of biodiversity data across domains and resolutions. We lay out effective strategies for data collection, mobilization, imputation, and sharing and summarize existing frameworks for scalable and integrative biodiversity research. Finally, we use two case studies to demonstrate how the explicit consideration of data domain and resolution helps to identify biases and gaps in global data sets and achieve unprecedented taxonomic and geographical data coverage in macroecological analyses., This Essay highlights data resolution as central property of biodiversity data that affects the precision and representativeness of macroecological inferences. It also discusses ways to maximize synergies among data types and showcases the potential of cross-resolution, cross-domain data integration.

Recent evidence is emerging that tropical cloud forests are not restricted to mountains, but may also be found in tropical lowland areas, the so-called ‘tropical lowland cloud forest’ (LCF). LCF occurs in river valleys with high air humidity and morning fog, and is rich in epiphytes. We explored the diversity of bark-inhabiting liverworts in LCF, a group of organisms known to be sensitive indicators of humidity. To test the hypothesis that LCF differs in species richness and species composition from lowland rain forest without fog (LRF), we sampled liverwort diversity in LCF and LRF on 48 whole trees in two sites in French Guiana. Sampling efficiency ( = no. of species found as percentage of estimated total number of species) was about 90% in both forest types. Species richness in LCF was significantly higher than in LRF and species composition differed in all height zones; moreover, LCF had three times more indicator species. Indicators of LCF included shade epiphytes and generalists that occurre...

Isolation is a driving factor of species richness and other island community attributes. Most empirical studies have investigated the effect of isolation measured as distance to the nearest continent. Here we expanded this perspective by comparing the explanatory power of seventeen isolation metrics in sixty-eight variations for vascular plant species richness on 453 islands worldwide. Our objectives were to identify ecologically meaningful metrics and to quantify their relative importance for species richness in a globally representative data set. We considered the distances to the nearest mainland and to other islands, stepping stone distances, the area of surrounding landmasses, prevailing wind and ocean currents and climatic similarity between source and target areas. These factors are closely linked to colonization and maintenance of plant species richness on islands. We tested the metrics in spatial multi-predictor models accounting for area, climate, topography and island geology. Besides area, isolation was the second most important factor determining species richness on the studied islands. A model including the proportion of surrounding land area as the isolation metric had the highest predictive power, explaining 86.1% of the variation. Distances to large islands, stepping stone distances and distances to climatically similar landmasses performed slightly better than distance to the nearest mainland. The effect of isolation was weaker for large islands suggesting that speciation counteracts the negative effect of isolation on immigration on large islands. Continental islands were less affected by isolation than oceanic islands. Our results suggest that a variety of immigration mechanisms influence plant species richness on islands and we show that this can be detected at macro-scales. Although the distance to the nearest mainland is an adequate and easy-to-calculate measure of isolation, accounting for stepping stones, large islands as source landmasses, climatic similarity and the area of surrounding landmasses increases the explanatory power of isolation for species richness.

1. The high biodiversity and biomass of soil communities is crucial for litter decomposition in terrestrial ecosystems such as tropical forests. However, the leaf litter that these communities consume is of particularly poor quality as indicated by elemental stoichiometry. The impact of resource quantity, quality, and other habitat parameters on species richness and biomass of consumer communities is often studied in isolation, although much can be learned from simultaneously studying both community characteristics. 2. Using a data set of 780 macro-invertebrate consumer species across 32 sites in tropical lowland rainforest and agricultural systems on Sumatra, Indonesia, we investigated the effects of basal resource stoichiometry (C:X ratios of N, P, K, Ca, Mg, Na, S in local leaf litter), litter mass (basal resource quantity and habitat space), plant species richness (surrogate for litter habitat heterogeneity), and soil pH (acidity) on consumer species richness and biomass across different consumer groups (i.e., three feeding guilds and ten selected taxonomic groups). 3. In order to distinguish the most important predictors of consumer species richness and biomass, we applied a standardised model averaging approach investigating the effects of basal resource stoichiometry, litter mass, plant species richness, and soil pH on both consumer community characteristics. This standardised approach enabled us to identify differences and similarities in the magnitude and importance of such effects on consumer species richness and biomass. 4. Across consumer groups, we found litter mass to be the most important predictor of both species richness and biomass. Resource stoichiometry had a more pronounced impact on consumer species richness than on their biomass. As expected, taxonomic groups differed in which resource and habitat parameters (basal resource stoichiometry, litter mass, plant species richness, and pH) were most important for modulating their community characteristics. 5. The importance of litter mass for both species richness and biomass indicates that these tropical consumers strongly depend on habitat space and resource availability. Our study supports previous theoretical work indicating that consumer species richness is jointly influenced by resource availability and the balanced supply of multiple chemical elements in their resources.

Aim To provide a mechanistic and probabilistic framework for defining the species pool based on species-specific probabilities of dispersal, environmental suitability and biotic interactions within a specific temporal extent, and to show how probabilistic species pools can help disentangle the geographical structure of different community assembly processes. Innovation Probabilistic species pools provide an improved species pool definition based on probabilities in conjunction with the associated species list, which explicitly recognize the indeterminate nature of species pool membership for a given focal unit of interest and better capture real-world complexity. Probabilistic species pools provide a quantitative assessment of how dispersal, environmental or biotic factors influence estimates of species pool composition and size for a given temporal extent. Conclusions Based on one simulated and two empirical examples we demonstrate that probabilistic species pools allow us to disentangle the geographical variation in dispersal, environmental and biotic assembly processes for species assemblages in focal units. We also show that probabilistic species pools are fully compatible with traditional definitions of species pools and are applicable over a wide range of spatial and temporal extents. Additionally they are robust to missing data and provide a quantified and transparent approach to estimating the size and composition of species pools in a mechanistic way, providing a valuable tool for studies from community ecology to macroecology.

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. published

Island biogeographical models consider islands either as geologically static with biodiversity resulting from ecologically neutral immigration–extinction dynamics1, or as geologically dynamic with biodiversity resulting from immigration–speciation–extinction dynamics influenced by changes in island characteristics over millions of years2. 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 levels3, 4 and caused massive changes in island area, isolation and connectivity5, orders of magnitude faster than the geological processes of island formation, subsidence and erosion considered in island theory2, 6. Consequences of these oscillations for present biodiversity remain unassessed5, 7. 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.

In the light of putatively increasing liana abundances in present-day tropical forests and a persistent lack of understanding of liana abundance patterns and the responsible factors, we attempt to identify the key factors controlling liana abundance along an altitudinal transect in NE Ecuador. At four elevational levels (500, 1000, 1500, and 2000 m), each represented by 10 plots of 400 m2, the abundance and diameters of all lianas (dbh ≥ 1 cm) and trees (dbh ≥ 10 cm) were recorded in old-growth forest stands in the Sumaco Biosphere Reserve (SBR). Results were analysed with available data on soil chemical properties and canopy structure. The mean density of lianas and liana basal area did not differ significantly between the elevational levels; however, the mean liana diameter decreased with increasing elevation. Liana density, basal area and abundance per tree individual were strongly correlated with host tree diameter. The most important determinants of liana abundance in the SBR were structural stand properties (tree basal area and mean tree dbh), but soil nitrogen availability (indicated by C/N ratio) was also found to have some influence. The notably high variation in liana abundance among different old-growth forest stands mainly reflects differences in stand structure between the studied mature forests, whereas the effect of elevation seems to be of less importance.

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 GDM's 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.

All around the globe, humans have greatly altered the abiotic and biotic environment with ever-increasing speed. One defining feature of the Anthropocene epoch(1,2) 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(3). 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.

Aim The aim of this study was to investigate whether the equilibrium theory of island biogeography (ETIB) is equally applicable at regional and local spatial scales, and whether the ‘echo pattern’, the correlation between regional species pool and local diversity by which the diversity of the regional species pool ‘trickles down’ to local scales, influences local diversity equally across a range of sampling scales. Location Twelve mountain regions on islands having different sizes, degrees of isolation and environmental conditions in Indonesia and the Philippines. Methods We sampled ferns in standardized field plots within a fixed spatial design at six spatial scales (with the grain size varying from plot to island). Using ordinary least squares regression and relative variable importance, we tested for the predictive power of area and isolation as well as of local and regional environmental factors for explaining diversity at the chosen spatial scales. Results Consistent with MacArthur and Wilson's ETIB, we found the strongest correlations of area and isolation with diversity at regional scales. The explanatory power of area increased with increasing spatial scale. The strength of the negative relationships among diversity and isolation decreased with increasing spatial scale. We found evidence for an ‘echo pattern’ at large to intermediate but not at small spatial scales. Main conclusions Area and isolation are strong predictors of biodiversity at regional scales. The size of the species pool has a scale-dependent influence on diversity at smaller scales but is unimportant at the most local scale, where environmental conditions dominate as predictors of diversity. To make predictions about local diversity on islands it is therefore important to take the scale dependence of the ‘echo pattern’ into account.

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 km 2 (∼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.

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.

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.

The data and scripts used in analyses for Zizka et al 2022, 'The evolution of Insular woodiness'