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SummaryPlant life and growth forms (shortened to ‘plant forms’) represent key functional strategies of plants in relation to their environment and provide important insights into the ecological constraints acting on the distribution of biodiversity. Despite their ecological importance, how the spectra of plant forms contribute to global gradients of plant diversity is unresolved.Using a novel dataset comprising >295,000 species, we quantify the contribution of different plant forms to global gradients of vascular plant diversity. Further, we establish how plant form distributions in different biogeographical regions are associated with contemporary and paleoclimate conditions, environmental heterogeneity, and phylogeny.We find a major shift in representation by woody perennials in tropical latitudes to herb-dominated floras in temperate and boreal regions, following a sharp latitudinal gradient in plant form diversity from the tropics to the poles. We also find significant functional differences between regions, mirroring life and growth form responses to environmental conditions, which is mostly explained by contemporary climate (18-87%), and phylogeny (6-62%), with paleoclimate and heterogeneity playing only a minor role (This research highlights variation in the importance of different plant forms to diversity gradients worldwide, providing a much-needed quantification for long-standing ideas and concepts structuring plant assemblages.

Large-scale biodiversity data, for example, on species distribution and richness information, are being mobilized and becoming available at an increasing rate. Interactive web applications like atlases have been developed to visualize available datasets and make them accessible to a wider audience. Web mapping tools are changing rapidly, and different underlying concepts have been developed to visualize datasets at a high cartographic standard. 2. Here, we introduce the Combined Atlas Framework for the development of interactive web atlases for ecological data visualization. We combine two existing approaches: the five stages of the user-centred design approach for web mapping applications and the three U approach for interface success. 3. Subsequently, we illustrate the use of this framework by developing the Atlas of Plant Invasions based on the Global Naturalized Alien Flora (GloNAF) database. This case study illustrates how the newly developed Combined Atlas Framework with a user-centred design philosophy can generate measurable success through communication with the target user group, iterative prototyping and competitive analysis of other existing web mapping approaches. 4. The framework is useful in creating an atlas that employs user feedback to determine usability and utility features within an interactive atlas system. Finally, this framework will enable a better-informed development process of future visualization and dissemination of biodiversity data through web mapping applications and interactive atlases. published

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.

Assessing phylogenetic endemism, i.e., the distribution of geographically restricted and evolutionarily unique species, is key to understanding biogeographic patterns and processes and critical for global conservation planning. 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 of the Southern Hemisphere as well as tropical mountainous regions displayed the world’s highest phylogenetic endemism. Tropical moist forests (e.g. Amazonia) and continental islands in south-east Asia emerged as centers of paleoendemism, while both high neo- and paleoendemism were found on ancient continental fragment islands (e.g. Madagascar) and in Mediterranean-climate regions. Global variation in phylogenetic endemism was best explained by a combination of past and present environmental factors (80.3% – 88.1% of variance explained). Geographic isolation and environmental heterogeneity emerged as primary drivers promoting high phylogenetic endemism. Also, warm and wet climates with long-term climatic stability showed a significant positive effect. However, environmental effects on phylogenetic endemism varied with geographic isolation, reflecting the unique evolutionary and biogeographic dynamics on oceanic islands. Long-term climatic stability promoted the persistence of paleoendemics, while isolation promoted higher neoendemism leading to oceanic islands and tropical mountainous regions being centers of both neo- and paleoendemism. Our study provides new insights into the evolutionary underpinnings of biogeographic patterns in seed plants, and by identifying areas of high evolutionary and biogeographic uniqueness, it may serve as a key resource for setting global conservation priorities.

Aim:Our current understanding of the causes of global extinction risk is mostly informed by the expert knowledge-based “threats classification scheme” of the IUCN Red List of Threatened Species. Studies based on this dataset came to different conclusions about the relative importance of threats to species, depending on which taxonomic groups and levels of extinction risk were considered, and which version of the database was used. A key reason may lie in data limitations as causes of threat are well known for charismatic and well-studied species, but not for the majority of species assessed. Here, we aim to fill current knowledge gaps about the importance of drivers of global extinction risks by focusing on endemic species.Location:Global.Methods:We examined country-level variation in the proportion of globally threatened and extinct endemic species (Index of Threat, IoT) with a range of spatially explicit information about anthropogenic pressures, mitigation measures and data limitations.Results:IoT coincided with several anthropogenic pressures, with substantial differences among kingdoms, life-forms, levels of extinction risk and geographic locations. IoT of plants, particularly tropical woody plants of moderate extinction risk, was higher in countries with higher GDP and more invasive species. Furthermore, IoT of animals, particularly tropical mammals and invertebrates of moderate extinction risk, was higher in countries with higher GDP and smaller roadless areas.Main conclusions:The extinction crisis for endemic species is associated with a complex network of potential drivers that need to be considered in concert in conservation policy and practice. Although our results require careful interpretation and remain sensitive to data limitations, we encourage similar studies at smaller scales to identify potential drivers of extinction risk at a higher resolution, particularly in regions where species assessments have been conducted consistently or on organisms with a uniform response time to pressures. published

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

Despite the paramount role of plant diversity for ecosystem functioning, biogeochemical cycles, and human welfare, knowledge of its global distribution is still incomplete, hampering basic research and biodiversity conservation.Here, we used machine learning (random forests, extreme gradient boosting, and neural networks) and conventional statistical methods (generalized linear models and generalized additive models) to test environment-related hypotheses of broad-scale vascular plant diversity gradients and to model and predict species richness and phylogenetic richness worldwide. To this end, we used 830 regional plant inventories including c. 300 000 species and predictors of past and present environmental conditions.Machine learning showed a superior performance, explaining up to 80.9% of species richness and 83.3% of phylogenetic richness, illustrating the great potential of such techniques for disentangling complex and interacting associations between the environment and plant diversity. Current climate and environmental heterogeneity emerged as the primary drivers, while past environmental conditions left only small but detectable imprints on plant diversity.Finally, we combined predictions from multiple modeling techniques (ensemble predictions) to reveal global patterns and centers of plant diversity at multiple resolutions down to 7774 km2 . Our predictive maps provide accurate estimates of global plant diversity available at grain sizes relevant for conservation and macroecology. published

With globalisation facilitating the movement of plants and seeds beyond the native range, preventing potentially harmful introductions requires knowledge of what drives the successful establishment and spread of alien plants. Here, we examined global‐scale relationships between naturalisation success (incidence and extent) and invasiveness, soil seed bank properties (type and densities) and key species traits (seed mass, seed dormancy and life form) for 2350 species of angiosperms. Naturalisation and invasiveness were strongly associated with the ability to form persistent (vs. transient) seed banks but relatively weakly with seed bank densities and other traits. Our findings suggest that seed bank persistence is a trait that better captures the ability to become naturalised and invasive compared to seed traits more widely available in trait databases. Knowledge of seed persistence can contribute to our ability to predict global naturalisation and invasiveness and to identify potentially invasive flowering plants before they are introduced., We examine global‐scale relationships between naturalization success and invasiveness, soil seed bank properties (type and density), and key species traits (seed mass, seed dormancy, and life form) for 2350 angiosperm taxa. We found that the ability to form persistent reserves of viable seeds in the soil is a consistent indicator of the incidence and extent of naturalization and of the likelihood of them becoming invasive. We also showed that seed bank persistence is a trait that better captures the ability of angiosperms to become naturalized and invasive compared to seed traits more widely available in trait databases. ​

Biotic interactions are known to play an important role in shaping species geographic distributions and diversity patterns. However, the role of mutualistic interactions in shaping global diversity patterns remains poorly quantified, particularly with respect to interactions with invertebrates. Moreover, it is unclear how the nature of different mutualisms interacts with abiotic drivers and affects diversity patterns of mutualistic organisms. Here, we present a global-scale biogeographic analysis of three different ant-plant mutualisms, differentiating between plants bearing domatia, extrafloral nectaries (EFNs), and elaiosomes, based on comprehensive geographic distributions of ∼15,000 flowering plants and ∼13,000 ant species. Domatia and extrafloral nectaries involve indirect plant defenses provided by ants, while elaiosomes attract ants to disperse seeds. Our results show distinct biogeographic patterns of different ant-plant mutualisms, with domatium- and EFN-bearing plant richness decreasing sharply from the equator towards the poles, while elaiosome-bearing plants prevail at mid-latitudes. Contemporary climate, especially mean annual temperature and precipitation, emerge as the most important predictor of ant-associated plant diversity. In hot and moist regions, typically the tropics, domatium- and EFN-bearing plant richness increases with related ant guild richness, while in warm regions plants with elaiosomes are strongly linked to interacting ants. Our results suggest that ant richness in combination with climate drives the spatial variation of plants bearing domatia, extrafloral nectaries, and elaiosomes, highlighting the importance of mutualistic interactions for understanding plant biogeography and its response to global change.

Despite the paramount role of plant diversity for ecosystem functioning, biogeochemical cycles, and human welfare, knowledge of its global distribution is incomplete, hampering basic research and biodiversity conservation. Here, we used machine learning (random forests, extreme gradient boosting, neural networks) and conventional statistical methods (generalised linear models, generalised additive models) to model species richness and phylogenetic richness of vascular plants worldwide based on 830 regional plant inventories including c. 300,000 species and predictors of past and present environmental conditions. Machine learning showed an outstanding performance, explaining up to 80.9% of species richness and 83.3% of phylogenetic richness. Current climate and environmental heterogeneity emerged as the primary drivers, while past environmental conditions left only small but detectable imprints on plant diversity. Finally, we combined predictions from multiple modelling techniques (ensemble predictions) to reveal global patterns and centres of plant diversity at multiple resolutions down to 7,774 km2. Our predictive maps provide the most accurate estimates of global plant diversity available to date at grain sizes relevant for conservation and macroecology.

Oceanic island floras are well-known for their morphological peculiarities and exhibit striking examples of trait evolution1,2. These morphological shifts are commonly attributed to insularity and thought to be shaped by biogeographical processes and evolutionary histories of oceanic islands1,3. However, the mechanisms through which biogeography and evolution have shaped the distribution and diversity of plant functional traits remain unclear. Here, we describe the functional trait space of an oceanic island flora (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 is concentrated around a functional hotspot dominated by shrubs with a conservative life-history strategy. By dividing the island flora into species groups with distinct biogeographical distributions and diversification histories, our results reveal that long-distance dispersal, and the interplay between inter-island dispersal and archipelago-level speciation processes drive functional divergence and expand trait space. Conversely, speciation via cladogenesis has overall led to functional convergence, densely packing trait space around shrubbiness. Our approach combines ecology, biogeography and evolution and opens avenues for new trait-based insights into how dispersal and speciation shape the assembly of native island floras.

Aim: The General Dynamic Model of island biogeography (GDM) explains patterns of species richness on volcanic hotspot archipelagos with respect to island age (T) and area (A), most commonly employing what we here call semi‐log ATT² and LogATT² models. These can be considered direct extensions of semi‐log and power law species‐area relationships (SARs), with a hump‐shaped function of T modelling drivers of biodiversity over and above area per se. We compare these and related models and investigate their conceptual and mathematical connections. Location: The Azores, Galapagos, Hawaii, Marquesas, Society Islands, Cape Verde Islands, Canary Islands and Aeolian archipelagos. Taxa: Vascular plants, birds, bryophytes, arthropods and molluscs. Methods: We gathered species richness data from databases and published sources, resulting in 34 datasets for various taxa. We used the Akaike information criterion corrected for small sample size (AICc) to compare the relative performance of models. We analysed the data at archipelago level and across archipelagos by using mixed effect models. We also calculated the relative time of expected peak diversity (Tpeak) from parameter estimates. Results: In almost three‐fourth of the cases the best model overall was one that contained only A or T rather than both predictors. There was greater variance between taxa than between archipelagos. Better dispersed taxa tended to reach peak diversity earlier. The most generally applicable of the models including T were the LogATT² and LogTT² models. These have important, previously unrecognized mathematical properties. Main Conclusions: SARs and species–time relationships should always be considered as potentially more parsimonious options. Considering the LogATT² model as a modification of the power law S = cAz, where a lognormal function in T replaces the constant, c allows the construction of an extended framework for the GDM with incorporation of additional predictors such as measures of environmental heterogeneity and application beyond hotspot archipelagos. Refereed/Peer-reviewed

Humans cultivate thousands of economic plants (i.e. plants with economic value) outside their native ranges. To analyze how this contributes to naturalization success, we combine global databases on economic uses and naturalization success of the world’s seed plants. Here we show that naturalization likelihood is 18 times higher for economic than non-economic plants. Naturalization success is highest for plants grown as animal food or for environmental uses (e.g. ornamentals), and increases with number of uses. Taxa from the Northern Hemisphere are disproportionately over-represented among economic plants, and economic plants from Asia have the greatest naturalization success. In regional naturalized floras, the percentage of economic plants exceeds the global percentage and increases towards the equator. Phylogenetic patterns in the naturalized flora partly result from phylogenetic patterns in the plants we cultivate. Our study illustrates that accounting for the intentional introduction of economic plants is key to unravelling drivers of plant naturalization., Understanding why certain alien species become naturalized can shed light on biological invasion patterns. In this global analysis on thousands of taxa, van Kleunen and colleagues show that plant species of economic use are more likely to become naturalized, and that this underlies geographic patterns and phylogenetic signals in naturalization

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 (

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 Refereed/Peer-reviewed

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. publishedVersion

Aim Functional traits can help us to elucidate biogeographical and ecological processes driving assemblage structure. We analysed the functional diversity of plant species of different evolutionary origins across an island archipelago, along environmental gradients and across geological age, to assess functional aspects of island biogeographical theory. Location Canary Islands, Spain. Major taxa studied Spermatophytes. Time period Present day. Methods We collected data for four traits (plant height, leaf length, flower length and fruit length) associated with resource acquisition, competitive ability, reproduction and dispersal ability of 893 endemic, non-endemic native and alien plant species (c. 43% of the Canary Island flora) from the literature. Linking these traits to species occurrences and composition across a 500 m × 500 m grid, we calculated functional diversity for endemic, non-endemic native and alien assemblages using multidimensional functional hypervolumes and related the resulting patterns to climatic (humidity) and island biogeographical (geographical isolation, topographic complexity and geological age) gradients. Results Trait space of endemic and non-endemic native species overlapped considerably, and alien species added novel trait combinations, expanding the overall functional space of the Canary Islands. We found that functional diversity of endemic plant assemblages was highest in geographically isolated and humid grid cells. Functional diversity of non-endemic native assemblages was highest in less isolated and humid grid cells. In contrast, functional diversity of alien assemblages was highest in arid ecosystems. Topographic complexity and geological age had only a subordinate effect on functional diversity across floristic groups. Main conclusions We found that endemic and non-endemic native island species possess similar traits, whereas alien species tend to expand functional space in ecosystems where they have been introduced. The spatial distribution of the functional diversity of floristic groups is very distinct across environmental gradients, indicating that species assemblages of different evolutionary origins thrive functionally in dissimilar habitats. publishedVersion

Islands are hotspots of plant endemism and are particularly vulnerable to the establishment (naturalization) of alien plant species. Naturalized species richness on islands depends on several biogeographical and socioeconomic factors, but especially on remoteness. One potential explanation for this is that the phylogenetically imbalanced composition of native floras on remote islands leaves unoccupied niche space for alien species to colonize. Here, we tested whether the species richness of naturalized seed plants on 249 islands worldwide is related to the phylogenetic composition of their native floras. To this end, we calculated standardized effect size (ses) accounting for species richness for three phylogenetic assemblage metrics (Faith's phylogenetic diversity (PD), PDses; mean pairwise distance (MPD), MPDses; and mean nearest taxon distance (MNTD), MNTDses) based on a phylogeny of 42 135 native island plant species and related them to naturalized species richness. As covariates in generalized linear mixed models, we included native species richness and biogeographical, climatic and socioeconomic island characteristics known to affect naturalized species richness. Our analysis showed an increase in naturalized species richness with increasing phylogenetic clustering of the native assemblages (i.e. native species more closely related than expected by chance), most prominently with MPDses. This effect, however, was smaller than the influence of native species richness and biogeographical factors, e.g. remoteness. Further, the effect of native phylogenetic structure (MPDses) on naturalized species richness was stronger for smaller islands, but this pattern was not consistent across all phylogenetic assemblage metrics. This finding suggests that the phylogenetic composition of native island floras may affect naturalized species richness, particularly on small islands where species are more likely to co-occur locally. Overall, we conclude that the composition of native island assemblages affects their susceptibility to plant naturalizations in addition to other socioeconomic and biogeographical factors, and should be considered when assessing invasion risks on islands., Ecography, 2022 (11), ISSN:0906-7590, ISSN:1600-0587

Aim:Species characteristics and cultivation are both associated with alien plant naturalization and invasiveness. Particular species characteristics are favoured for cultivation, obscuring the relationship between traits and naturalization success. We sought to better understand the drivers of naturalization and invasiveness by analysing relationships with species characteristics and cultivation and by disentangling the direct effects of characteristics from the indirect effects mediated by cultivation.Location:Great Britain.Time period:c. 1000–present.Major taxa studied:Seed plants.Methods:We used a comprehensive dataset of 17,396 alien plant taxa introduced to Great Britain before 1850, a country with one of the most well-documented histories of plant introductions. We integrated this with cultivation data from historical and modern records from botanic gardens and commercial nurseries and with trait data. Accounting for time since introduction, we quantified the influences of cultivation and species characteristics on present-day naturalization and invasiveness in Great Britain.Results:Larger native range size, earlier flowering, long-lived herbaceous growth form, and outdoor cultivated habitat were all associated with naturalization. However, these relationships between characteristics and naturalization largely reflected cultivation patterns. The indirect, mediating influence of cultivation on naturalization varied among species characteristics, and was relatively strong for growth form and weak for native range size. Cultivation variables, particularly availability in present-day nurseries, best explained invasiveness, while species characteristics had weaker associations.Main conclusions:Human influence on species introduction and cultivation is associated with increased probability of naturalization and invasiveness, and it has measurable indirect effects by biasing the distribution of species characteristics in the pool of introduced species. Accounting for human cultivation preferences is necessary to make ecological interpretations of the effects of species characteristics on invasion. published