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The megadiverse plant family Asteraceae forms an iconic component of island floras including many spectacular radiations, but a global picture of its insular diversity is lacking. Here, we uncover the global biogeographical and evolutionary patterns of Asteraceae on islands to reveal the magnitude and potential causes of their evolutionary success. We compile a global checklist of Asteraceae species native and endemic to islands and combine it with macroecological analyses and a phylogenetic review of island radiations. Asteraceae have a global distribution on islands, comprising approximately 6,000 native island species, with 58% endemics. While diversity of the family on islands is lower than expected given its overall diversity, Asteraceae are the most diverse family on oceanic islands, suggesting an exceptional ability to thrive in isolation. In agreement with island biogeography predictions, native Asteraceae diversity increases with area and decreases with isolation, while endemism increases with both. We identify 39 confirmed island radiations and 69 putative radiations, exceeding numbers for other iconic insular groups. Our results reveal Asteraceae offer immense potential for research in ecology and evolution, given their close tracking of island biogeography expectations, large number of both species and radiations, cosmopolitan distribution, and numerous undiscovered radiations., (© 2024. The Author(s).)

Oceanic island floras are well known for their morphological peculiarities and exhibit striking examples of trait evolution 1-3 . These morphological shifts are commonly attributed to insularity and are thought to be shaped by the biogeographical processes and evolutionary histories of oceanic islands 2,4 . However, the mechanisms through which biogeography and evolution have shaped the distribution and diversity of plant functional traits remain unclear 5 . Here we describe the functional trait space of the native flora of an oceanic island (Tenerife, Canary Islands, Spain) using extensive field and laboratory measurements, and relate it to global trade-offs in ecological strategies. We find that the island trait space exhibits a remarkable functional richness but that most plants are concentrated around a functional hotspot dominated by shrubs with a conservative life-history strategy. By dividing the island flora into species groups associated with distinct biogeographical distributions and diversification histories, our results also suggest that colonization via long-distance dispersal and the interplay between inter-island dispersal and archipelago-level speciation processes drive functional divergence and trait space expansion. Contrary to our expectations, speciation via cladogenesis has led to functional convergence, and therefore only contributes marginally to functional diversity by densely packing trait space around shrubs. By combining biogeography, ecology and evolution, our approach opens new avenues for trait-based insights into how dispersal, speciation and persistence shape the assembly of entire native island floras., (© 2023. The Author(s), under exclusive licence to Springer Nature Limited.)

Insular woodiness (IW)-the evolutionary transition from herbaceousness toward woodiness on islands-is one of the most iconic features of island floras. Since pioneering work by Darwin and Wallace, a number of drivers of IW have been proposed, such as 1) competition for sunlight requiring plants with taller and stronger woody stems and 2) drought favoring woodiness to safeguard root-to-shoot water transport. Alternatively, IW may be the indirect result of increased lifespan related to 3) a favorable aseasonal climate and/or 4) a lack of large native herbivores. However, information on the occurrence of IW is fragmented, hampering tests of these potential drivers. Here, we identify 1,097 insular woody species on 375 islands and infer at least 175 evolutionary transitions on 31 archipelagos, concentrated in six angiosperm families. Structural equation models reveal that the insular woody species richness on oceanic islands correlates with a favorable aseasonal climate, followed by increased drought and island isolation (approximating competition). When continental islands are also included, reduced herbivory pressure by large native mammals, increased drought, and island isolation are most relevant. Our results illustrate different trajectories leading to rampant convergent evolution toward IW and further emphasize archipelagos as natural laboratories of evolution, where similar abiotic or biotic conditions replicated evolution of similar traits.

Island species and habitats are particularly vulnerable to human disturbances, and anthropogenic changes are increasingly overwriting natural island biogeographic patterns. However, quantitative comparisons of how native and alien assemblages respond to human disturbances are scarce. Using data from 6,242 species of vertebrates, invertebrates and plants, from 7,718 sites on 81 islands, we model how land-use change, human population density and distance to the nearest road affect local assemblages of alien and native species on islands. We found that land-use change reduces both richness and abundance of native species, whereas the number and abundance of alien species are high in plantation forests and agricultural or urban sites. In contrast to the long-established pattern for native species (i.e., decline in species number with island isolation), more isolated islands have more alien species across most land uses than do less isolated islands. We show that alien species play a major role in the turnover of island assemblages: our models show that aliens outnumber natives among the species present at disturbed sites but absent from minimally-disturbed primary vegetation. Finally, we found a homogenization pattern for both native and alien assemblages across sites within most land uses. The declines of native species on islands in the face of human pressures, and the particular proneness to invasions of the more remote islands, highlight the need to reduce the intensity of human pressures on islands and to prevent the introduction and establishment of alien species., Competing Interests: The authors have declared that no competing interests exist.

One central concept in island biology is that island assemblages form subsets of the mainland species pool, being disproportionately rich or poor in certain taxonomic groups. This unbalanced composition, termed 'disharmony', is generally explained using a taxon-centred approach, linking the over- or under-representation of taxa to their colonisation abilities. However, islands may also harbour 'functionally' disharmonic flora, being disproportionately rich or poor in species with certain traits, which may offer greater insights into the processes driving island colonisation. Here, we use orchids as a model to illustrate key processes involved in the formation of functionally disharmonic island floras, including filtering effects (for example biotic interactions), and speciation. Our synthesis is based on a comprehensive orchid dataset of 27 637 species and combines both a literature review and simple exploratory analyses to show that orchids are significantly under-represented on islands relative to mainland regions and that insular orchids display shifts in functional traits, from the shortening of nectar spurs to facilitate ornithophily to changes in colour associated with generalist insect pollinators. We highlight that taxa are simply coarse proxies and that we need to consider species traits and interactions to gain a full understanding of the processes constraining plant assembly on islands., (© 2019 The Authors. New Phytologist © 2019 New Phytologist Trust.)

One of the best-known general patterns in island biogeography is the species-isolation relationship (SIR), a decrease in the number of native species with increasing island isolation that is linked to lower rates of natural dispersal and colonization on remote oceanic islands. However, during recent centuries, the anthropogenic introduction of alien species has increasingly gained importance and altered the composition and richness of island species pools. We analyzed a large dataset for alien and native plants, ants, reptiles, mammals, and birds on 257 (sub) tropical islands, and showed that, except for birds, the number of naturalized alien species increases with isolation for all taxa, a pattern that is opposite to the negative SIR of native species. We argue that the reversal of the SIR for alien species is driven by an increase in island invasibility due to reduced diversity and increased ecological naiveté of native biota on the more remote islands., Competing Interests: The authors declare no conflict of interest.

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., (© 2016 Cambridge Philosophical Society.)

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.

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.

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Aim: To investigate spatial variations in the ecological trait structure of breeding bird assemblages on oceanic islands. To test the hypothesis that native and naturalized alien bird species are filtered by different processes, leading to diverging associations between traits and insular environmental gradients. Location: Oceanic islands worldwide. Time Period: Current. Major Taxa Studied: Terrestrial breeding birds. Methods: We assessed the composition of breeding terrestrial bird assemblages from the extent‐of‐occurrence maps of 3170 native and 169 naturalized alien species on 4660 oceanic islands. We quantified their ecological trait structures with respect to diet, mobility and body mass as the standardized distance between a mean pairwise trait distance index and its expectation from a null model. We used spatial generalized additive models to relate trait structures to proxies of environmental conditions and human impact on land, accounting for all species, native species only and alien species only. Results: Diet and mobility traits tended to be more diverse than expected by a null model, while body mass tended towards clustering. Trait‐environment associations were idiosyncratic, but environmental variables tended to explain trait structures better than human impacts on habitats. Islands invaded by alien species had similar trait structures as noninvaded ones, although they hosted assemblages with more clustered body masses. However, trait‐environment relationships diverged when considering all islands and all species, invaded islands only or alien species only. Main Conclusions: Despite their ecosystem‐level influences, alien species have a limited effect on the global patterns of trait structures in the breeding bird assemblages of the world's islands, either because they account for a low proportion of species or because successful invaders and native species have similar trait suites. Trait‐environment associations suggest that filters related to the conditions of alien species' introductions explain their distributions in island assemblages better than the constraints associated with isolated environments. [ABSTRACT FROM AUTHOR]

Assessing the distribution of geographically restricted and evolutionarily unique species and their underlying drivers is key to understanding biogeographical processes and critical for global conservation prioritization. Here, we quantified the geographic distribution and drivers of phylogenetic endemism for ~320,000 seed plants worldwide and identified centers and drivers of evolutionarily young (neoendemism) and evolutionarily old endemism (paleoendemism). Tropical and subtropical islands as well as tropical mountain regions displayed the world’s highest phylogenetic endemism. Most tropical rainforest regions emerged as centers of paleoendemism, while most Mediterranean-climate regions showed high neoendemism. Centers where high neo-and paleoendemism coincide emerged on some oceanic and continental fragment islands, in Mediterranean-climate regions and parts of the Irano-Turanian floristic region. Global variation in phylogenetic endemism was well explained by a combination of past and present environmental factors (79.8 to 87.7% of variance explained) and most strongly related to environmental heterogeneity. Also, warm and wet climates, geographic isolation, and long-term climatic stability emerged as key drivers of phylogenetic endemism. Neo-and paleoendemism were jointly explained by climatic and geological history. Long-term climatic stability promoted the persistence of paleoendemics, while the isolation of oceanic islands and their unique geological histories promoted neoendemism. Mountainous regions promoted both neo-and paleoendemism, reflecting both diversification and persistence over time. Our study provides insights into the evolutionary underpinnings of biogeographical patterns in seed plants and identifies the areas on Earth with the highest evolutionary and biogeographical uniqueness—key information for setting global conservation priorities. [ABSTRACT FROM AUTHOR]

Current models of island biogeography treat endemic and non‐endemic species as if they were functionally equivalent, focussing primarily on species richness. Thus, the functional composition of island biotas in relation to island biogeographical variables remains largely unknown. Using plant trait data (plant height, leaf area and flower length) for 895 native species in the Canary Islands, we related functional trait distinctiveness and climate rarity for endemic and non‐endemic species and island ages. Endemics showed a link to climatically rare conditions that is consistent with island geological change through time. However, functional trait distinctiveness did not differ between endemics and non‐endemics and remained constant with island age. Thus, there is no obvious link between trait distinctiveness and occupancy of rare climates, at least for the traits measured here, suggesting that treating endemic and non‐endemic species as functionally equivalent in island biogeography is not fundamentally wrong. [ABSTRACT FROM AUTHOR]

Island biogeography has classically focused on abiotic drivers of species distributions. However, recent work has highlighted the importance of mutualistic biotic interactions in structuring island floras. The limited occurrence of specialist pollinators and mycorrhizal fungi have been found to restrict plant colonization on oceanic islands. Another important mutualistic association occurs between nearly 15,000 plant species and nitrogen-fixing (N-fixing) bacteria. Here, we look for evidence that N-fixing bacteria limit establishment of plants that associate with them. Globally, we find that plants associating with N-fixing bacteria are disproportionately underrepresented on islands, with a 22% decline. Further, the probability of N-fixing plants occurring on islands decreases with island isolation and, where present, the proportion of N-fixing plant species decreases with distance for large, but not small islands. These findings suggest that N-fixing bacteria serve as a filter to plant establishment on islands, altering global plant biogeography, with implications for ecosystem development and introduction risks. Plants associated with nitrogen-fixing bacterial symbionts are disproportionately underrepresented on islands, suggesting that these bacteria might play a limiting role in island plant biogeography. [ABSTRACT FROM AUTHOR]

The influence of island dynamics and characteristics on taxonomic diversity, particularly species richness, are well studied. Yet, our knowledge on the influence of island dynamics and characteristics on other facets of diversity, namely functional and phylogenetic diversity, is limited, constraining our understanding of assembly processes on islands (e.g. biogeographic history, dispersal and environmental filtering and species interactions). Using barrier islands, a highly dynamic and so far, understudied island type, we investigate how multiple facets of vascular plant diversity (functional, phylogenetic and taxonomic diversity) are shaped by island geomorphology, modern and historic area, and habitat heterogeneity. In line with our expectation, historical dynamics in island geomorphology affected phylogenetic and taxonomic diversity via habitat heterogeneity. However, island area was the best predictor across all facets of diversity. Specifically, larger islands had higher functional and phylogenetic diversity than expected by chance while most of the smaller islands had lower diversity. The influence of area on functional diversity acted via habitat heterogeneity, with habitat heterogeneity influencing negatively functional diversity. Our results suggest that larger islands accumulate functionally and phylogenetically unique species. Further, results for functional diversity pointed towards potential area–heterogeneity trade‐offs, with these trade‐offs likely resulting from increased interspecific competition favoring a specific set of trait values (of stronger competitors), particularly on smaller islands. Together, these results demonstrate that going beyond taxonomic diversity contributes to identifying underlying processes shaping diversity–area relationships. [ABSTRACT FROM AUTHOR]

Island biogeography is the study of the spatio‐temporal distribution of species, communities, assemblages or ecosystems on islands and other isolated habitats. Island diversity is structured by five classes of process: dispersal, establishment, biotic interactions, extinction and evolution. Classical approaches in island biogeography focused on species richness as the deterministic outcome of these processes. This has proved fruitful, but species traits can potentially offer new biological insights into the processes by which island life assembles and why some species perform better at colonising and persisting on islands. Functional traits refer to morphological and phenological characteristics of an organism or species that can be linked to its ecological strategy and that scale up from individual plants to properties of communities and ecosystems. A baseline hypothesis is for traits and ecological strategies of island species to show similar patterns as a matched mainland environment. However, strong dispersal, environmental and biotic‐interaction filters as well as stochasticity associated with insularity modify this baseline. Clades that do colonise often embark on distinct ecological and evolutionary pathways, some because of distinctive evolutionary forces on islands, and some because of the opportunities offered by freedom from competitors or herbivores or the absence of mutualists. Functional traits are expected to be shaped by these processes. Here, we review and discuss the potential for integrating functional traits into island biogeography. While we focus on plants, the general considerations and concepts may be extended to other groups of organisms. We evaluate how functional traits on islands relate to core principles of species dispersal, establishment, extinction, reproduction, biotic interactions, evolution and conservation. We formulate existing knowledge as 33 working hypotheses. Some of these are grounded on firm empirical evidence, others provide opportunities for future research. We organise our hypotheses under five overarching sections. Section A focuses on plant functional traits enabling species dispersal to islands. Section B discusses how traits help to predict species establishment, successional trajectories and natural extinctions on islands. Section C reviews how traits indicate species biotic interactions and reproduction strategies and which traits promote intra‐island dispersal. Section D discusses how evolution on islands leads to predictable changes in trait values and which traits are most susceptible to change. Section E debates how functional ecology can be used to study multiple drivers of global change on islands and to formulate effective conservation measures. Islands have a justified reputation as research models. They illuminate the forces operating within mainland communities by showing what happens when those forces are released or changed. We believe that the lens of functional ecology can shed more light on these forces than research approaches that do not consider functional differences among species. [ABSTRACT FROM AUTHOR]

Aim: The island species–area relationship (ISAR) quantifies how the number of species increases as the area of an island or island‐like habitat gets larger and is one of the most general patterns in ecology. However, studies that measure the ISAR often confound variation in sampling methodology and analyses, precluding appropriate syntheses of its underlying mechanisms. Most ISAR studies use only presence–absence data at the whole‐island scale, whereas we planned to use a framework that applies individual‐based rarefaction to synthesize whether and how the ISAR differs from the null expectation of the passive sampling hypothesis. Location: Five hundred and five islands from 34 different archipelagos across the world, including oceanic islands, lake islands and forest islands. Major taxa studied: Local assemblages of plants, invertebrates, herpetofauna, birds and mammals. Methods: We collated local‐scale species abundance data from multiple archipelagos (median of 12 islands per study) and used a rarefaction‐based approach to synthesize the relationship between island size and (1) sample effort‐controlled rarefied species richness, or (2) an effective number of species derived from the probability of interspecific encounter (an index of community evenness). Results: When we applied rarefaction to control for sampling effort, the numbers of species and their relative abundances across all studies differed from the passive sampling hypothesis. Our measure of evenness also increased with island size, suggesting that the disproportionate effects we observed influenced both rarer and more common species. We found few associations between the slope of this effect and island type or taxon, but we did find that island archipelagos with greater elevational heterogeneity also deviated more from the null expectation than those with less heterogeneity. Main conclusions: Using a synthetic approach across island archipelagos, we reject the null expectation that passive sampling causes the ISAR and instead suggest that ecological mechanisms leading to disproportionate (non‐random) effects on larger relative to smaller islands are predominant. [ABSTRACT FROM AUTHOR]

Aim: Adaptive radiation, in which successful lineages proliferate by exploiting untapped niche space, provides a popular but potentially misleading characterization of evolution on oceanic islands. Here we analyse the respective roles of members of in situ diversified vs. non‐diversified lineages in shaping the main ecosystems of an archipelago to explore the relationship between evolutionary and ecological 'success'. Location: Canary Islands. Taxon: Vascular plants. Methods: We quantified the abundance/rarity of the native flora according to the geographical range (number of islands where present and geographical extent of the range), habitat breadth (climatic niche) and local abundance (cover) using species distribution data based on 500 × 500 m grid cells and 2000 vegetation inventories located all over the archipelago. Results: Species of diversified lineages have significantly smaller geographic ranges, narrower climatic niches and lower local abundances than those of non‐diversified lineages. Species rarity increased with the degree of diversification. The diversified Canarian flora is mainly comprised by shrubs. At both archipelagic and island level, the four core ecosystems (Euphorbia scrub, thermophilous woodlands, laurel forest and pine forest) were dominated by non‐diversified lineages species, with diversified lineages species providing <25% cover. Species of diversified lineages, although constituting 54% of the archipelagic native flora, were only abundant in two rare ecosystems: high mountain scrub and rock communities. Main conclusions: Radiated species, endemic products of in situ speciation, are mostly rare in all three rarity axes and typically do not play an important role in structuring plant communities on the Canaries. The vegetation of the major ecosystem types is dominated by plants representing non‐diversified lineages (species that derive from immigration and accumulation), while species of evolutionarily successful lineages are abundant only in marginal habitats and could, therefore, be considered ecological losers. Within this particular oceanic archipelago, and we posit within at least some others, evolutionary success in plants is accomplished predominantly at the margins. [ABSTRACT FROM AUTHOR]

Aim: Species diversity on islands generally increases with island area. This might arise either from direct effects of island area via neutral assembly processes or from indirect effects via habitat and structural differences between islands that scale positively with island area. Here, we tested whether community‐weighted functional trait means of woody plants are directly or indirectly affected by island area to elucidate how functional traits mediate the assembly on differently sized islands. Location: Twenty‐eight tropical islands (25 m2 – 12,000 m2) in the Raja Ampat archipelago, Indonesia. Taxon: Woody angiosperms. Methods: Studied islands had a shared geological history but differed in terms of area, habitat quality expressed by soil depth, forest structure expressed by tree basal area and degree of isolation. Traits studied were seed and fruit mass, tree height, wood density, leaf mass per area, leaf nitrogen concentration and chlorophyll content (estimated from chlorophyll‐meter units) and summarised as community‐weighted means (CWM) for each island. Using liner regression, we tested whether CWMs were correlated to island area and basal area and structural equation models (SEMs) to test on direct and indirect effects of island area, basal area, soil depth and isolation on trait distributions. Results: CWM of seed mass, tree height and chlorophyll content increased with both island area and basal area, whereas leaf nitrogen concentration decreased with increasing basal area. Fruit mass was not correlated to island area and basal area. SEMs revealed that the shifts in tree height, wood density, leaf nitrogen concentration and chlorophyll content were caused directly by basal area, which in turn was directly and positively affected by both island area and soil depth. Differences in seed mass among islands were explained by combined effects of basal area, island area and isolation, whereas fruit mass was only explained by isolation. Main conclusions: Trait values shifted systematically across islands of different sizes. Being small and having light seeds are prevailing trait combinations for establishing on small islands with simple forest structure. For establishment on larger islands with more complex forest structures, species are taller, have heavier seeds, higher chlorophyll content and lower leaf N concentrations. We conclude that mechanisms affecting CWM on islands directly link to ecological differences between islands like forest structure – and only indirectly to island area. [ABSTRACT FROM AUTHOR]

Questions: Islands harbour a disproportionate amount of global plant diversity, yet their unique native assemblages are particularly vulnerable to biological invasions. It is therefore critical to identify the macroecological constraints that mediate spatial distributions of alien species on islands. Here, we examined abundance–occupancy relationships of native and alien woody plant species, and the role of niche properties and functional traits related to dispersal and competition in shaping occupancy and abundance patterns. Location: Hawaiian Islands. Methods: We calculated relative abundance and occupancy for 64 woody species (42 natives, 22 naturalized aliens), and estimated each species' niche breadth and position. We fitted phylogenetic hierarchical Bayesian models to evaluate abundance– occupancy relationships and the impacts of niche properties and functional traits on occupancy and abundance of native and alien species. Results: Our analyses revealed that locally more abundant native species were also more widespread, but that the abundance of alien species was unrelated to their occupancy. Yet, we found evidence that alien species with longer residence times on the Hawaiian Islands were more widespread. While widespread native and alien woody species both had broad niches, widespread alien woody species exhibited a tendency to occur in more marginal niche positions than widespread native woody species. Niche properties did not affect abundances of either native or alien woody species. Traits associated with dispersal capacity and competitive ability had minimal impacts on either occupancy or abundance of native and alien woody species. Conclusions: We found that niche properties shape the occupancy but not the abundance of native and alien woody species across Hawaiian forests. Our results suggest that, because of substantial invasion debts, the impacts of alien woody invasions in native forests have yet to fully manifest. [ABSTRACT FROM AUTHOR]

The extraordinary diversity of orchids has captivated scientists for more than a century, yet their complex spatial patterns at large scales remain poorly resolved. On islands, orchid diversity patterns are especially puzzling. While some islands are centres of orchid diversity, orchids are underrepresented on most islands. To disentangle such complex patterns, key functional differences among orchids must be considered – a distinction seldom made in biogeographical analyses. Using a global dataset of 454 islands, we tested prominent hypotheses in island biogeography, while simultaneously making the distinction between epiphytes and two terrestrial life forms (geophytes and non‐geophytes). Orchid diversity was unevenly distributed across islands and life forms. Epiphytic orchid diversity strongly increased with temperature, illustrating the near confinement of epiphytes to the tropics. Geophytes became proportionally more important with increasing seasonality, highlighting their ability to withstand harsh climatic conditions. Epiphytes and non‐geophytes both displayed responses (e.g. negative relationship with seasonality) related to their dependence on consistently favourable conditions, possibly because of the absence of subterranean storage organs. This highlights that the factors explaining orchid diversity differ strongly with, and are related to, life form. We suggest that key functional differences within and across plant families be considered in future studies to better understand drivers of complex diversity patterns. [ABSTRACT FROM AUTHOR]

Community assembly processes on islands are often non‐random. The mechanisms behind non‐random assembly, however, are generally difficult to disentangle. Functional diversity in combination with a null model approach that accounts for differences in species richness among islands can be used to test for non‐random assembly processes, but has been applied rarely to island communities. By linking functional diversity of trees on islands with a null model approach, we bridge this gap and test for the role of stochastic versus non‐random trait‐mediated assembly processes in shaping communities by studying functional diversity–area relationships. We measured 11 plant functional traits linked to species dispersal and resource acquisition strategies of 57 tree species on 40 tropical islands. We grouped traits into four life‐history dimensions representing 1) dispersal ability, 2) growth strategy, 3) light acquisition and 4) nutrient acquisition. To test for non‐random assembly processes, we used null models that account for differences in species richness among the islands. Our results reveal contrasting responses of the four life‐history dimensions to island area. The dispersal and the growth strategy dimensions were underdispersed on smaller islands, whereas the light acquisition dimension was overdispersed. The nutrient acquisition dimension did not deviate from null expectations. With increasing island area, shifts in the strength of non‐random assembly processes increased the diversity of dispersal and acquisition strategies in island communities. Our results suggest that smaller islands may be more difficult to colonize and provide more limited niche space compared to larger islands, whose tree communities are likely determined by stochastic processes and higher niche diversity. Our null model approach highlights that analyzing the functional diversity of different life‐history dimensions provides a powerful framework to unravel community assembly processes on islands. These complex, non‐random assembly processes are masked by measures of functional diversity that do not account for differences in species richness between islands. [ABSTRACT FROM AUTHOR]

Aim: Geological processes of island growth and erosion, in addition to eustatic sea‐level fluctuations, alter key physical properties of oceanic islands, such as area, elevation and isolation over time. These dynamics operate over similar time‐scales to the biogeographical processes of colonization, speciation and extinction, and thus should strongly affect island biodiversity patterns as we observe them today. Here, we study the effects of geological history and eustatic sea‐level fluctuations on biodiversity patterns in the Hawaiian, Galapagos and Canary Islands, using an agent‐based simulation model. Location: Hawaiian Islands, Galapagos Islands and Canary Islands. Time period: Geological history up to 20 Ma. Taxa: Spermatophyta, Pteridophyta, Passeriformes, Lepidoptera and Coleoptera. Methods: We analysed the effect of geological, eustatic and biogeographical processes via a spatially explicit agent‐based simulation model. We compared the emergent patterns with observed species richness and the proportion of single‐island endemics across the considered taxa and archipelagos. Results: Our results suggested that both geological and eustatic processes have affected biodiversity assembly in the Hawaiian and Galapagos Islands. In the Canary Islands, we found only the proportion of single‐island endemics to be affected by historical configurations. Main conclusions: Our results highlight the key role of past geological and eustatic processes in affecting modern insular biodiversity patterns. [ABSTRACT FROM AUTHOR]

Island disharmony refers to the biased representation of higher taxa on islands compared to their mainland source regions and represents a central concept in island biology. Here, we develop a generalizable framework for approximating these source regions and conduct the first global assessment of island disharmony and its underlying drivers. We compiled vascular plant species lists for 178 oceanic islands and 735 mainland regions. Using mainland data only, we modelled species turnover as a function of environmental and geographic distance and predicted the proportion of shared species between each island and mainland region. We then quantified the over‐ or under‐representation of families on individual islands (representational disharmony) by contrasting the observed number of species against a null model of random colonization from the mainland source pool, and analysed the effects of six family‐level functional traits on the resulting measure. Furthermore, we aggregated the values of representational disharmony per island to characterize overall taxonomic bias of a given flora (compositional disharmony), and analysed this second measure as a function of four island biogeographical variables. Our results indicate considerable variation in representational disharmony both within and among plant families. Examples of generally over‐represented families include Urticaceae, Convolvulaceae and almost all pteridophyte families. Other families such as Asteraceae and Orchidaceae were generally under‐represented, with local peaks of over‐representation in known radiation hotspots. Abiotic pollination and a lack of dispersal specialization were most strongly associated with an insular over‐representation of families, whereas other family‐level traits showed minor effects. With respect to compositional disharmony, large, high‐elevation islands tended to have the most disharmonic floras. Our results provide important insights into the taxon‐ and island‐specific drivers of disharmony. The proposed framework allows overcoming the limitations of previous approaches and provides a quantitative basis for incorporating functional and phylogenetic approaches into future studies of island disharmony. [ABSTRACT FROM AUTHOR]

Aim: Mountains and islands are both well known for their high endemism. To explain this similarity, parallels have been drawn between the insularity of "true islands" (land surrounded by water) and the isolation of habitats within mountains (so‐called "mountain islands"). However, parallels rarely go much beyond the observation that mountaintops are isolated from one another, as are true islands. Here, we challenge the analogy between mountains and true islands by re‐evaluating the literature, focusing on isolation (the prime mechanism underlying species endemism by restricting gene flow) from a dynamic perspective over space and time. Framework: We base our conceptualization of "isolation" on the arguments that no biological system is completely isolated; instead, isolation has multiple spatial and temporal dimensions relating to biological and environmental processes. We distinguish four key dimensions of isolation: (a) environmental difference from surroundings; (b) geographical distance to equivalent environment [points (a) and (b) are combined as "snapshot isolation"]; (c) continuity of isolation in space and time; and (d) total time over which isolation has been present [points (c) and (d) are combined as "isolation history"]. We evaluate the importance of each dimension in different types of mountains and true islands, demonstrating that substantial differences exist in the nature of isolation between and within each type. In particular, different types differ in their initial isolation and in the dynamic trajectories they follow, with distinct phases of varying isolation that interact with species traits over time to form present‐day patterns of endemism. Conclusions: Our spatio‐temporal definition of isolation suggests that the analogy between true islands and mountain islands masks important variation of isolation over long time‐scales. Our understanding of endemism in isolated systems can be greatly enriched if the dynamic spatio‐temporal dimensions of isolation enter models as explanatory variables and if these models account for the trajectories of the history of a system. [ABSTRACT FROM AUTHOR]

Aim: We tested whether species–area relationships of small islands differ among plant growth forms and whether this influences the prevalence of the small‐island effect (SIE). The SIE states that species richness on small islands is independent of island area or relates to area in a different way compared with larger islands. We investigated whether island isolation affects the limits of the SIE and which environmental factors drive species richness on small islands. Location: Seven hundred islands (< 100 km2) worldwide belonging to 17 archipelagos. Major taxa studied: Angiosperms. Methods: We applied linear and breakpoint species–area models for angiosperm species richness and for herb, shrub and tree species richness per archipelago separately, to test for the existence of SIEs. For archipelagos featuring the SIE, we calculated the island area at which the breakpoints occurred (breakpoint area) and used linear models to test whether the breakpoint areas varied with isolation. We used linear mixed‐effect models to discern the effects of seven environmental variables related to island area, isolation and other environmental factors on the species richness of each growth form for islands smaller than the breakpoint area. Results: For 71% of all archipelagos, we found an SIE for total and herb species richness, and for 59% for shrub species richness and 53% for tree species richness. Shrub and tree species richness showed larger breakpoint areas than total and herb species richness. The breakpoint area was significantly positively affected by the isolation of islands within an archipelago for total and shrub species richness. Species richness on islands within the range of the SIE was differentially affected by environmental factors across growth forms. Main conclusion: The SIE is a widespread phenomenon that is more complex than generally described. Different functional groups have different environmental requirements that shape their biogeographical patterns and affect species–area and, more generally, richness–environment relationships. The complexity of these patterns cannot be revealed when measuring overall plant species richness. [ABSTRACT FROM AUTHOR]

Aim: The decreasing capacity of area to predict species richness on small islands (the small‐island effect; SIE) seems to be one of the few exceptions of the species–area relationship. While most studies have focused on how to detect the SIE, the underlying ecological factors determining this pattern remain largely unexplored. Here, we evaluate one of the few mechanisms proposed to explain the SIE, the subsidized island biogeography hypothesis, which posits that marine productivity around small islands may alter their expected species richness. Location: Seven hundred and ninety islands worldwide, including 420 islands < 1 km2. Time period: Present. Major taxa studied: Angiosperms. Methods: We applied iterative partial regression to determine the effects of island area and marine productivity on plant species richness for islands of varying sizes. We also employed geographically weighted regression to account for non‐stationarity in the marine productivity effects. Lastly, we used estimates of ammonia emissions based on nutrient excretion by seabird colonies from a subset of 66 islands to evaluate the effects of marine resources deposition on angiosperm species richness. Results: We found no effect of marine productivity on insular species richness, at both regional and global scales. In all models, area emerged as the only predictor of plant species richness. A weak contribution of marine productivity was only detectable in models with a low number of islands, but this effect was independent of island size. Although nutrient deposition significantly contributes to explaining plant diversity, this effect was also independent of island size. Main conclusions: Our study demonstrates that marine productivity has no general effect on plant species richness of small islands worldwide. Although marine‐derived resources may still contribute to species richness variation, this effect does not seem to be restricted to small islands. Overall, our results do not provide support for the subsidized island biogeography hypothesis. [ABSTRACT FROM AUTHOR]

Aim: We mapped global patterns of tree phylogenetic endemism (PE) to identify hotspots and test hypotheses about possible drivers. Specifically, we tested hypotheses related to current climate, geographical characteristics and historical conditions and assessed their relative importance in shaping PE patterns. Location: Global. Time period: We used the present distribution of trees, and predictors covering conditions from the mid‐Miocene to present. Major taxa studied: All seed‐bearing trees. Methods: We compiled distributions for 58,542 tree species across 463 regions worldwide, matched these to a recent phylogeny of seed plants and calculated PE for each region. We used a suite of predictor variables describing current climate (e.g., mean annual temperature), geographical characteristics (e.g., isolation) and historical conditions (e.g., tree cover at the Last Glacial Maximum) in a spatial regression model to explain variation in PE. Results: Tree PE was highest on islands, and was higher closer to the equator. All three groups of predictor variables contributed substantially to the PE pattern. Isolation and topographic heterogeneity promoted high PE, as did high current tree cover. Among mainland regions, temperature seasonality was strongly negatively related to PE, while mean annual temperature was positively related to PE on islands. Some relationships differed among the major floristic regions. For example, tree cover at the Last Glacial Maximum was a positive predictor of PE in the Palaeotropics, while tree cover at the Miocene was a negative predictor of PE in the Neotropics. Main conclusions: Globally, PE can be explained by a combination of geographical, historical and current factors. Some geographical variables appear to be key predictors of PE. However, the impact of historic and current climate variables differs considerably among the major floristic regions, reflecting their unique histories. Hence, the current distribution of trees is the result of globally relevant geographical drivers and regional climatic histories. [ABSTRACT FROM AUTHOR]

Understanding how species diversity is related to sampling area and spatial scale is central to ecology and biogeography. Small islands and small sampling units support fewer species than larger ones. However, the factors influencing species richness may not be consistent across scales. Richness at local scales is primarily affected by small‐scale environmental factors, stochasticity and the richness at the island scale. Richness at whole‐island scale, however, is usually strongly related to island area, isolation and habitat diversity. Despite these contrasting drivers at local and island scales, island species–area relationships (SARs) are often constructed based on richness sampled at the local scale. Whether local scale samples adequately predict richness at the island scale and how local scale samples influence the island SAR remains poorly understood. We investigated the effects of different sampling scales on the SAR of trees on 60 small islands in the Raja Ampat archipelago (Indonesia) using standardised transects and a hierarchically nested sampling design. We compared species richness at different grain sizes ranging from single (sub)transects to whole islands and tested whether the shape of the SAR changed with sampling scale. We then determined the importance of island area, isolation, shape and habitat quality at each scale on species richness. We found strong support for scale dependency of the SAR. The SAR changed from exponential shape at local sampling scales to sigmoidal shape at the island scale indicating variation of species richness independent of area for small islands and hence the presence of a small‐island effect. Island area was the most important variable explaining species richness at all scales, but habitat quality was also important at local scales. We conclude that the SAR and drivers of species richness are influenced by sampling scale, and that the sampling design for assessing the island SARs therefore requires careful consideration. [ABSTRACT FROM AUTHOR]

Aims: The general dynamic model (GDM) of oceanic island biogeography predicts how biogeographical rates, species richness and endemism vary with island age, area and isolation. Here, we used a simulation model to assess whether the isolation‐related predictions of the GDM may arise from low‐level process at the level of individuals and populations. Location: Hypothetical volcanic oceanic islands. Methods: Our model considers (a) an idealized island ontogeny, (b) metabolic constraints and (c) stochastic, spatially explicit and niche‐based processes at the level of individuals and populations (plant demography, dispersal, competition, mutation and speciation). Isolation scenarios involved varying the distance to mainland and the dispersal ability of the species pool. Results: For all isolation scenarios, we obtained humped temporal trends for species richness, endemic richness, proportion of endemic species derived from within‐island radiation, number of radiating lineages, number of species per radiating lineage and biogeographical rates. The proportion of endemics derived from mainland–island differentiation and of all endemics steadily increased over time. Extinction rates of endemic species peaked later than for non‐endemic species. Species richness and the number of endemics derived from mainland–island differentiation decreased with isolation as did rates of colonization, mainland–island differentiation and extinction. The proportion of all endemics and of radiated endemics, the number of radiated endemics, of radiating lineages, and of species per radiating lineage and the within‐island radiation rate all increased with isolation. Main conclusions: Our results lend strong support to most of the isolation‐related GDM predictions. New insights include an increasing proportion of endemics, particularly those arising from mainland–island differentiation, across isolation scenarios, as well as extinction trends of endemics differing from the overall extinction rates, with a much later peak. These results demonstrate how simulation models focusing on low ecological levels provide tools to assess biogeographical‐scale predictions and to develop more detailed predictions for further empirical tests. [ABSTRACT FROM AUTHOR]

Questions: Small islands are outstanding model systems to study community assembly. Due to harsher environmental conditions on smaller islands compared to larger ones, environmental filtering may preclude some species, potentially resulting in island size‐dependent species pools. We tested whether the species pool size follows a similar species–area relationship as the observed richness. This can provide new insight into community assembly processes and the elusive small‐island effect (SIE), which states that species richness on smaller islands is less dependent on area than on larger islands. Location: Raja Ampat Archipelago, Indonesia. Methods: We studied the woody vegetation on sixty small islands ranging from 3 m2 to 11,806 m2. For each recorded species, we estimated its area requirements and compared them against random colonization models. We developed a novel method to calculate probabilistic species pools for each island. We compared different species–area models for observed species richness and our index of species pool size to test whether the SIE results from differences in species pool size. Results: We found that most species were restricted to islands significantly larger than expected from random colonization. The occurrence probability of all species increased with island size, indicating a lack of species that are specialized to the conditions on small islands. We found a SIE in observed species richness, but not in species pool size. Conclusion: Woody plants in the studied island system have specific requirements that are linked to island area and determine island‐specific species pools. Lower community completeness on smaller islands compared to larger ones indicated that the SIE is shaped by local limiting processes that have no impact on the species pool, but control how much of it is realized on an island. Together, these results clearly indicate non‐random plant community assembly on small islands. [ABSTRACT FROM AUTHOR]

Aim: To determine the global position of tree line isotherms, compare it with observed local tree limits on islands and mainlands, and disentangle the potential drivers of a difference between tree line and local tree limit. Location: Global. Time period: 1979–2013. Major taxa studied: Trees. Methods: We modelled the potential climatic tree line based on monthly temperatures and precipitation for the period 1979–2013. We then compared the potential tree line based on climate to observed tree limits at 26 oceanic islands, 55 continental islands and 382 mainland locations. The differences between potential tree line and observed tree limits was then analysed by regression with the islands' maximum elevation, age, isolation, and area. Additionally, we estimated growing season temperature niches for 16,041 species known to occur in the vicinity of the studied tree lines, and compared them across mainlands, and islands of continental and oceanic origin. Results: Observed local tree limits differ up to 2,066 m from the potential tree line at the mainland on oceanic islands. Climatic effects are responsible for a difference of up to 1,296 m between tree lines of mainland regions and oceanic islands (but only for 756 m for continental islands). On oceanic islands, a remaining difference of up to 829 m correlates with the isolation and the maximum elevation of an island. Floras of oceanic islands are however depauperate with respect to potential tree line species and species show an affinity to higher growing season temperatures. Main conclusions: Climate can explain about half of the differences between observed local tree limits and potential tree lines between the mainland and continental and oceanic islands. The remaining difference can be attributed to the higher isolation of oceanic islands, especially in the tropics, and as a consequence, a more depauperate flora and a lack of tree species that are able to grow at the tree line. [ABSTRACT FROM AUTHOR]

Islands harbour evolutionary and ecologically unique biota, which are currently disproportionately threatened by a multitude of anthropogenic factors, including habitat loss, invasive species and climate change. Native forests on oceanic islands are important refugia for endemic species, many of which are rare and highly threatened. Long-term monitoring schemes for those biota and ecosystems are urgently needed: (i) to provide quantitative baselines for detecting changes within island ecosystems, (ii) to evaluate the effectiveness of conservation and management actions, and (iii) to identify general ecological patterns and processes using multiple island systems as repeated ‘natural experiments’. In this contribution, we call for a Global Island Monitoring Scheme (GIMS) for monitoring the remaining native island forests, using bryophytes, vascular plants, selected groups of arthropods and vertebrates as model taxa. As a basis for the GIMS, we also present new, optimized monitoring protocols for bryophytes and arthropods that were developed based on former standardized inventory protocols. Effective inventorying and monitoring of native island forests will require: (i) permanent plots covering diverse ecological gradients (e.g. elevation, age of terrain, anthropogenic disturbance); (ii) a multiple-taxa approach that is based on standardized and replicable protocols; (iii) a common set of indicator taxa and community properties that are indicative of native island forests’ welfare, building on, and harmonized with existing sampling and monitoring efforts; (iv) capacity building and training of local researchers, collaboration and continuous dialogue with local stakeholders; and (v) long-term commitment by funding agencies to maintain a global network of native island forest monitoring plots. [ABSTRACT FROM AUTHOR]

Biological invasions are a defining feature of the Anthropocene, but the factors that determine the spatially uneven distribution of alien plant species are still poorly understood. Here, we present the first global analysis of the effects of biogeographic factors, the physical environment and socio-economy on the richness of naturalized and invasive alien plants. We used generalized linear mixed-effects models and variation partitioning to disentangle the relative importance of individual factors, and, more broadly, of biogeography, physical environment and socio-economy. As measures of the magnitude of permanent anthropogenic additions to the regional species pool and of species with negative environmental impacts, we calculated the relative richness of naturalized (= RRN) and invasive (= RRI) alien plant species numbers adjusted for the number of native species in 838 terrestrial regions. Socio-economic factors (per-capita gross domestic product (GDP), population density, proportion of agricultural land) were more important in explaining RRI (~50 % of the explained variation) than RRN (~40 %). Warm-temperate and (sub)tropical regions have higher RRN than tropical or cooler regions. We found that socio-economic pressures are more relevant for invasive than for naturalized species richness. The expectation that the southern hemisphere is more invaded than the northern hemisphere was confirmed only for RRN on islands, but not for mainland regions nor for RRI. On average, islands have ~6-fold RRN, and >3-fold RRI compared to mainland regions. Eighty-two islands (=26 % of all islands) harbour more naturalized alien than native plants. Our findings challenge the widely held expectation that socio-economic pressures are more relevant for plant naturalization than for invasive plants. To meet international biodiversity targets and halt the detrimental consequences of plant invasions, it is essential to disrupt the connection between socio-economic development and plant invasions by improving pathway management, early detection and rapid response. [ABSTRACT FROM AUTHOR]