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Major regional gaps exist in the reporting and accessibility of naturalized plant species distribution data, especially within Southeast Asia. Here, we present the Malesian Naturalized Alien Flora database (MalNAF), the first standardized island-group level checklist of naturalized vascular plant species for the Malesian phytogeographical region. We used MalNAF to investigate the composition, origins, and habitat preferences of the naturalized flora. The naturalized vascular flora of Malesia consists of at least 1177 species. Richness is highest in the Philippines (539 spp.) and lowest in the Maluku Islands (87 spp.). But, the Lesser Sunda Islands had the highest naturalized species richness relative to native richness and Singapore has a higher naturalized plant species richness than would be expected given its size. When comparing the data for Malesia with a global dataset, we found that naturalized richness increased with area for islands but not for continental regions. Across the archipelago, 31 species are widespread, occurring in every island group, but the majority have a limited distribution of 2.4 ± 2.3 (mean ± SD) island groups per naturalized species. The naturalized plant species are representatives of 150 families, twenty of which are newly introduced to the region. Families richest in naturalized plant species in Malesia were Fabaceae (= Leguminosae) (160 spp.), Poaceae (= Gramineae) (138 spp.), and Asteraceae (= Compositae) (96 spp.). Most of these have a native range that includes tropical Asia, closely followed by those from Southern America (inclusive of the Caribbean, Central and South America), although at the island-group level, most have a higher proportion with a Southern American native range. Most naturalized species occur in anthropogenic habitats, but many are present in “natural” habitats with fewer species, such as Leucaena leucocephala, reported from specialized habitats like drylands. MalNAF provides a baseline for future studies of naturalized plant species distributions in the region.

In the United Nations Decade on Ecosystem Restoration1, large knowledge gaps persist on how to increase biodiversity and ecosystem functioning in cash crop-dominated tropical landscapes2. Here, we present findings from a large-scale, 5-year ecosystem restoration experiment in an oil palm landscape enriched with 52 tree islands, encompassing assessments of ten indicators of biodiversity and 19 indicators of ecosystem functioning. Overall, indicators of biodiversity and ecosystem functioning, as well as multidiversity and ecosystem multifunctionality, were higher in tree islands compared to conventionally managed oil palm. Larger tree islands led to larger gains in multidiversity through changes in vegetation structure. Furthermore, tree enrichment did not decrease landscape-scale oil palm yield. Our results demonstrate that enriching oil palm-dominated landscapes with tree islands is a promising ecological restoration strategy, yet should not replace the protection of remaining forests.

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.

Ecosystem stability reveals how ecosystems respond to global change over time. Yet, the focus of past research on small spatial grains and extents overlooks scale dependence and how broad-scale environmental gradients shape stability. Here, we use forest inventory data covering a broad latitudinal gradient from the temperate to the tropical zone to examine cross-scale variation in stability of aboveground biomass and underlying stabilizing mechanisms. While stability did not shift systematically with latitude at either spatial grain, we found evidence that species asynchrony increased towards the tropics at the small spatial grain while species stability decreased at both spatial grains. Moreover, latitude stabilized forest communitiesviaits effects on both stabilizing mechanisms, which compensated for the weak and destabilizing effects of species richness. Yet, the trade-off in the relative importance of species stability and species asynchrony for stability was not mediated by latitude, suggesting that context-dependent factors - to a greater extent than macroecological ones - underlie large-scale patterns of stability. Our results highlight the crucial role of species asynchrony and species stability in determining ecosystem stability across broad-scale environmental gradients, suggesting that conserving biodiversity alone may not be sufficient for stabilizing naturally-assembled forest ecosystems.

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

Background: Tropical montane forests are important reservoirs of carbon and biodiversity but are threatened by deforestation and climate change. It is important to understand how forest structure and aboveground biomass change along gradients of elevation and succession. Questions: What are the interactive effect of elevation and two stages of succession on forest structure parameters? Studied species: Tree communities. Study site and dates: Cofre de Perote, Veracruz, Mexico. August to December 2015. Methods: We studied four sites along an elevational gradient (500, 1,500, 2,500, and 3,500 m). At each elevation and each forest type, we established five 20 × 20 m plots (n = 40 plots). Within each plot, we measured stem density, mean diameter at breast height (dbh), and tree height and derived basal area and aboveground biomass (AGB). Results: AGB peaked at 2,500 m and was significantly related to elevation and succession, with higher values in old-growth forests than in secondary forests at higher altitudes. Lower values of mean dbh and basal area were found at higher elevations. At the lowest elevation, both successional stages had the same values of stem density and AGB. At both lower elevations, secondary forests had higher values of dbh and basal area. There were high biomass stocks in the old-growth forest at 2,500 and 3,500 m. Conclusions: Old-growth forests at higher elevations are threatened by deforestation, consequently these remaining fragments must be preserved because of their storage capacity for biomass and their ability to mitigate climate change.

The area-wide estimation of aboveground biomass (AGB) and its changes as a proxy for the sequestration and emission of carbon are currently associated with high uncertainties. Here we combined interferometric synthetic aperture radar (InSAR) height models derived from TanDEM-X with repeated ground-based inventories from the years 2012 and 2019 to estimate InSAR height and AGB changes in a structurally diverse and dynamic landscape in Sumatra, Indonesia. The results suggested that the InSAR height models were highly accurate and the relationship between InSAR height and AGB change resulted in a coefficient of determination R 2 of 0.65 and a cross-validated root mean square error (RMSE) of 2.38 Mg ha−1 year−1, equivalent to 13.32% of the actual AGB difference range. The estimated AGB changes with TanDEM-X were further related to the initial canopy height and fire activities in the study area. Initial canopy heights and the occurrences of fires had a significant effect on the AGB change. In general, low canopy heights tend to be associated with increasing AGB over time, whereas high canopy heights tend to be associated with stable or decreasing AGB. As expected, fires had a negative impact on the AGB changes being more pronounced in forest areas compared to oil palm concessions. The results of this study are relevant for the utilization of spaceborne InSAR height models and its potential to estimate canopy height and AGB change on large spatial scales. It was demonstrated that these changes can be related to their sources and ecosystem processes. This AGB change estimation technique can be used to model the impacts of fires on AGB change and carbon emissions, which are important for sustainable forest management.

The scientific work of Alexander von Humboldt was influenced by his interaction with the diversity and natural wealth of the Neotropics. He proposed that climate determines plant diversity along elevational gradients based on his observations. Here, we evaluated the most prominent climate-based hypotheses in explaining plant diversity along an elevational gradient that Humboldt himself visited during his journey across Mexico. Specifically, we examined how climatic variables and forest-use intensity affected species richness and phylogenetic structure of major angiosperm life forms (trees, shrubs, epiphytes, herbs and lianas) along the Cofre de Perote mountain, Veracruz, Mexico. We analysed species richness and phylogenetic structure of angiosperms at eight sites between 30 to 3500 m a.s.l. We estimated the phylogenetic structure using a mega-phylogeny of angiosperms and the abundance-weighted net relatedness index. We considered multiple environmental factors’ direct and indirect effects by applying a piecewise structural equation modelling approach. Each life form responds differently to the environmental variables included in our model; however, it is observed that temperature is the main predictor of the taxonomic and phylogenetic diversity of the angiosperms studied, both when the different life forms are grouped and separated. Potential evapotranspiration and precipitation are variables that also influence some life forms’ diversity, especially taxonomic diversity. The forest-use intensity negatively affected only the taxonomic diversity of trees. These results highlight the influence of studying the different life forms of angiosperms in diversity gradient models and show the great influence that temperature has in conjunction with other environmental variables to promote the taxonomic and phylogenetic diversity of plant communities. Given the current global environmental crisis, an integrative biogeographically oriented vision based on Humboldt’s method is necessary. Honouring the work of Humboldt and continuing his legacy demands more research to understand the causes behind elevational diversity gradients.

How to enhance biodiversity in monoculture-dominated landscapes is a key sustainability question that requires considering the spatial organization of ecological communities (beta diversity). Here, we experimentally tested if increasing landscape heterogeneity – through tree islands – is a suitable landscape restoration strategy when aiming to enhance multi-taxa diversity. We found that multi-taxa diversity resulted from islands fostering unique species (turnover: between 0.18 - 0.73) rather than species losses and gains (nestedness: between 0.03 - 0.34), suggesting that tree islands enhance diversity at the landscape scale. Through partial correlation networks, we revealed that landscape heterogeneity is associated with multi-taxa diversity (strength = 0.84). Soil biota were also central to the overall community by connecting beta diversity patterns across taxa. Our results show that increasing landscape heterogeneity enhances multi-taxa diversity in monoculture-dominant landscapes. Furthermore, we highlight that strategies aiming to enhance multi-taxa diversity should consider that spatial distributions of above- and below-ground communities are associated.

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

Agroforestry can contribute to an increase in tree cover in historically forested tropical landscapes with associated gains in biodiversity and ecosystem functioning, but only if established on open land instead of underneath a forest canopy. However, declines in yields with increasing shade are common across agroforestry crops, driving shade-tree removal in forest-derived agroforests and hindering tree regrowth in open-land-derived agroforests. To understand trajectories of change in tree cover in forest- and open-land-derived agroforests, and the impacts of tree cover on vanilla yields, we studied 209 vanilla agroforests along an 88-year chronosequence in Madagascar. Additionally, we used remotely sensed canopy cover data to investigate tree cover change in the agricultural landscape. We found yields to vary widely but independently of canopy cover and land-use history (forest- vs. open-land-derived), averaging at 154.6 kg ha−1 year−1 (SD = 186.9). Furthermore, we found that forest- and open-land-derived vanilla agroforests gained canopy cover over time, but that only open-land-derived agroforests gained canopy height. Canopy cover increased also at the landscape scale: areas in the agricultural landscape with medium initial canopy cover gained 6.4% canopy cover over 10 years, but canopy cover decreased in areas with high initial canopy cover. These opposing trends suggest tree cover rehabilitation across areas covered by vanilla agroforests, whereas remnant forest fragments in the agricultural landscape were transformed or degraded. Our results indicate that yield-neutral tree rehabilitation through open-land-derived agroforestry could, if coupled with effective forest protection, provide benefits for both ecosystem functions and agricultural production in a smallholder-dominated agricultural landscape.

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. ​

Aim It is widely accepted that biodiversity is influenced by both niche‐related and spatial processes from local to global scales. Their relative importance, however, is still disputed, and empirical tests are surprisingly scarce at the global scale. Here, we compare the importance of area (as a proxy for pure spatial processes) and environmental heterogeneity (as a proxy for niche‐related processes) for predicting native mammal species richness world‐wide and within biogeographical regions. Location Global. Time period We analyse a spatial snapshot of richness data collated by the International Union for Conservation of Nature. Major taxa studied All terrestrial mammal species, including possibly extinct species and species with uncertain presence. Methods We applied a spreading dye algorithm to analyse how native mammal species richness changes with area and environmental heterogeneity. As measures for environmental heterogeneity, we used elevation ranges and precipitation ranges, which are well‐known correlates of species richness. Results We found that environmental heterogeneity explained species richness relationships better than did area, suggesting that niche‐related processes are more prevalent than pure area effects at broad scales. Main conclusions Our results imply that niche‐related processes are essential to understand broad‐scale species–area relationships and that habitat diversity is more important than area alone for the protection of global biodiversity.

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.

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The habitat heterogeneity hypothesis predicts that biodiversity increases with increasing habitat heterogeneity due to greater niche dimensionality. However, recent studies have reported that richness can decrease with high heterogeneity due to stochastic extinctions, creating trade-offs between area and heterogeneity. This suggests that greater complexity in heterogeneity-diversity relationships (HDRs) may exist, with potential for group-specific responses to different facets of heterogeneity that may only be partitioned out by a simultaneous test of HDRs of several species groups and several facets of heterogeneity. Here, we systematically decompose habitat heterogeneity into six major facets on ~500 temperate forest plots across Germany and quantify biodiversity of 12 different species groups, including bats, birds, arthropods, fungi, lichens and plants, representing 2,600 species. Heterogeneity in horizontal and vertical forest structure underpinned most HDRs, followed by plant diversity, deadwood and topographic heterogeneity, but the relative importance varied even within the same trophic level. Among substantial HDRs, 53% increased monotonically, consistent with the classical habitat heterogeneity hypothesis but 21% were hump-shaped, 25% had a monotonically decreasing slope and 1% showed no clear pattern. Overall, we found no evidence of a single generalizable mechanism determining HDR patterns.

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

Sustainable land-system transformations are necessary to avert biodiversity and climate collapse. However, it remains unclear where entry points for transformations exist in complex land systems. Here, we conceptualize land systems along land-use trajectories, which allows us to identify and evaluate leverage points, i.e., entry points on the trajectory where targeted interventions have particular leverage to influence land-use decisions. We apply this framework in the biodiversity hotspot Madagascar. In the northeast, smallholder agriculture results in a land-use trajectory originating in old-growth forests and spanning from forest fragments to shifting hill rice cultivation and vanilla agroforests. Integrating interdisciplinary empirical data on seven taxa, five ecosystem services, and three measures of agricultural productivity, we assess trade-offs and cobenefits of land-use decisions at three leverage points along the trajectory. These trade-offs and cobenefits differ between leverage points: Two leverage points are situated at the conversion of old-growth forests and forest fragments to shifting cultivation and agroforestry, resulting in considerable trade-offs, especially between endemic biodiversity and agricultural productivity. Here, interventions enabling smallholders to conserve forests are necessary. This is urgent since ongoing forest loss threatens to eliminate these leverage points due to path dependency. The third leverage point allows for the restoration of land under shifting cultivation through vanilla agroforests and offers cobenefits between restoration goals and agricultural productivity. The co-occurring leverage points highlight that conservation and restoration are simultaneously necessary to avert collapse of multifunctional mosaic landscapes. Methodologically, the framework highlights the importance of considering path dependency along trajectories to achieve sustainable land-system transformations. ispartof: Proceedings Of The National Academy Of Sciences vol:119 issue:7 ispartof: location:United States status: Published online

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