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Trait-based approaches are commonly used to understand ecological phenomena and processes. Trait data are typically gathered by measuring local specimens, retrieving published records, or a combination of the two. Implications of methodological choices in trait-based ecological studies—including source of data, imputation technique, and species selection criteria—are poorly understood. We ask: do different approaches for dataset-building lead to meaningful differences in trait datasets? If so, do these differences influence findings of a trait-based examination of plant invasiveness, measured as abundance and spread rate? We collected on-site (Victoria, Australia) and off-site (TRY database) height and specific leaf area records for as many species as possible out of 157 exotic herbaceous plants. For each trait, we built six datasets of species-level means using records collected on-site, off-site, on-site and off-site combined, and off-site supplemented via imputation based on phylogeny and/or trait correlations. For both traits, the six datasets were weakly correlated (ρ = 0.31–0.95 for height; ρ = 0.14–0.88 for SLA), reflecting differences in species’ trait values from the various estimations. Inconsistencies in species’ trait means across datasets did not translate into large differences in trait-invasion relationships. Although we did not find that methodological choices for building trait datasets greatly affected ecological inference about local invasion processes, we nevertheless recommend: (1) using on-site records to answer local-scale ecological questions whenever possible, and (2) transparency around methodological decisions related to selection of study species and estimation of missing trait values.

Eutrophication usually impacts biodiversity, species composition, and functioning of grassland communities. Whether such effects propagate to influence the stability of these community aspects is unknown. Using standardized experiments across 55 global grasslands, we quantified the effects of nutrient addition on five stability facets (i.e., temporal invariability and resistance during and recovery after dry and wet growing seasons) for three community aspects (i.e., aboveground biomass, community composition, and species richness). Nutrient addition reduced the temporal invariability and resistance of species richness and community composition, but not biomass, during dry and wet growing seasons. Temporal invariability and resistance during, but not recovery after, dry and wet growing seasons were strongly positively correlated in both ambient and eutrophic conditions. This indicates that maintaining and restoring the stability of plant communities requires increasing resistance rather than recovery. Harnessing the complexity of ecological stability provides new insights for grassland ecosystem sustainability in a changing world.

As Earth's climate has varied strongly through geological time, studying the impacts of past climate change on biodiversity helps to understand the risks from future climate change. However, it remains unclear how paleo-climate shapes spatial variation in biodiversity. Here, we assessed the influence of Quaternary climate change on spatial dissimilarity in taxonomic, phylogenetic, and functional composition among neighboring 200-kilometer cells (beta-diversity) for angiosperm trees worldwide. We found that larger glacial-interglacial temperature change was strongly associated with lower spatial turnover (species replacements) and higher nestedness (rich-ness changes) components of beta-diversity across all three biodiversity facets. Moreover, phylogenetic and functional turnover was lower and nestedness higher than random expectations based on taxonomic beta -di-versity in regions that experienced large temperature change, reflecting phylogenetically and functionally se-lective processes in species replacement, extinction, and colonization during glacial-interglacial oscillations. Our results suggest that future human-driven climate change could cause local homogenization and reduction in taxonomic, phylogenetic, and functional diversity of angiosperm trees worldwide. W.-B.X. and A.O. acknowledge support from the Aarhus University Research Foundation (AUFF) Starting Grant (AUFF-F-2018-7-8). J.-C.S. acknowledges support from the Danish Council for Independent Research|Natural Sciences (grant 6108-00078B) to the TREECHANGE Project. J.-C.S. also considers this work a contribution to the Center for Ecological Dynamics in a Novel Biosphere (ECONOVO) funded by Danish National Research Foundation (grant DNRF173) and the VILLUM Investigator project “Biodiversity Dynamics in a Changing World” funded by VILLUM FONDEN (grant 16549). W.L.E. was supported by the VILLUM FONDEN (grant 00025354). M.A. was supported by the Natural Sciences and Engineering Research Council of Canada (NSERC) Discovery grants program. C.B. was supported by the National Research Foundation of Korea (NRF) grant funded by the Korea government (MSIT) (2022R1A2C1003504). A.G.G. was supported by the ANID PIA/BASAL (grant FB210006) and FONDECYT (grant 1200468). R.E.O. was supported by the German Research Foundation to German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig (DFG-FZT 118, 202548816). J.Pe. was supported by the Fundación Ramón Areces project (CIVP20A6621) and Spanish Government project (PID2019-110521GB-I00). J.Pi. was supported by the Estonian Research Council Grants (PUT1355, PRG 1405) and Mobilitas Pluss (MOBERC11). V.D.P. was supported by the National Council of Scientific and Technological Development (CNPq) of Brazil (grant 307689/2014-0).

Changes in some combination of niche availability, niche overlap and the strength of interspecific interactions are thought to drive changes in plant composition along resource gradients. However, because these processes are difficult to measure in the field, their relative importance in driving compositional change in plant communities remains unclear. In an Australian temperate grassland, we added seeds of three native and three exotic grasses to 1875 experimental plots in a way that allowed us to simultaneously estimate niche availability, niche overlap and the strength of pairwise interspecific interactions along a gradient of nutrient availability, obtained by adding 0, 5 or 20 g m−2 each of nitrogen, phosphorous and potassium jointly to plots. Niche availability (the proportion of microsites suitable for establishment and growth) was generally low and did not vary in response to nutrient addition. Most species co-occurred along the nutrient gradient by partitioning the available niche space. Where species interacted due to niche overlap, the abundance of one species, the native Chloris truncata, was usually facilitated by other species, with each of the five other species increasing the niche availability to C. truncata under at least one nutrient treatment. Chloris truncata also competitively excluded two species from some but not all sites they could otherwise have occupied. These outcomes did not clearly differ across nutrient treatments. Our results show that fine-scale spatial heterogeneity in establishment microsites can enable species to co-occur via niche partitioning, and competitive exclusion is rare. This finding contributes to an emerging picture that niche partitioning is common and frequently a stronger influence on recruitment outcomes than interspecific competition. The importance of competition in structuring plant communities may be overestimated if recruitment processes are overlooked.

Many human-introduced alien species economically impact essential industries worldwide. Management prioritization and coordination efforts towards biological invasions are hampered by a lack of comprehensive quantification of costs to key economic activity sectors. Here, we quantify and predict global invasion costs to seven major sectors and unravel the introduction pathways of species causing these costs — focusing mainly on resource-based agriculture, fishery and forestry industries. From 1970 to 2020, costs reported in the InvaCost database as pertaining to Agriculture, Fisheries, and Forestry totaled $509 bn, $1.3 bn, and $134 bn, respectively (in 2017 United States dollars). Pathways of costly species were diverse, arising predominantly from cultural and agricultural activities, through unintentional contaminants with trade, and often impacted different sectors than those for which species were initially introduced. Costs to Agriculture were pervasive and greatest in at least 37% (n = 46/123) of the countries assessed, with the United States accumulating the greatest costs for resource-based industries ($365 bn), followed by China ($101 bn), and Australia ($36 bn). We further identified 19 countries highly economically reliant on Agriculture, Fisheries, and Forestry that are experiencing massive economic impacts from biological invasions, especially in the Global South. Based on an extrapolation to fill cost data gaps, we estimated total global costs ranging at least from $517 − 1,400 bn for Agriculture, $5.7–6.5 bn for Fisheries, and $142–768 bn for Forestry, evidencing substantial underreporting in the Forestry sector in particular. Burgeoning global invasion costs challenge sustainable development and urge for improved management action to reduce future impacts on industry.

Safeguarding Earth?s tree diversity is a conservation priority due to the importance of trees for biodiversity and ecosystem functions and services such as carbon sequestration. Here, we improve the foundation for effective conservation of global tree diversity by analyzing a recently developed database of tree species covering 46,752 species. We quantify range protection and anthropogenic pressures for each species and develop conservation priorities across taxonomic, phylogenetic, and functional diversity dimensions. We also assess the effectiveness of several influential proposed conservation prioritization frameworks to protect the top 17% and top 50% of tree priority areas. We find that an average of 50.2% of a tree species? range occurs in 110-km grid cells without any protected areas (PAs), with 6,377 small-range tree species fully unprotected, and that 83% of tree species experience nonnegligible human pressure across their range on average. Protecting highpriority areas for the top 17% and 50% priority thresholds would increase the average protected proportion of each tree species? range to 65.5% and 82.6%, respectively, leaving many fewer species (2,151 and 2,010) completely unprotected. The priority areas identified for trees match well to the Global 200 Ecoregions framework, revealing that priority areas for trees would in large part also optimize protection for terrestrial biodiversity overall. Based on range estimates for >46,000 tree species, our findings show that a large proportion of tree species receive limited protection by current PAs and are under substantial human pressure. Improved protection of biodiversity overall would also strongly benefit global tree diversity. Made available in DSpace on 2022-06-21T15:23:55Z (GMT). No. of bitstreams: 1 Guo-et-al.-2022-PNAS-High-exposure-of-global-tree-diversity-to-human-pressure.pdf: 2523289 bytes, checksum: 245be3df3fb487ed152d25bf50f4e984 (MD5) Previous issue date: 2022 Proceedings of the National Academy of Sciences of the United States of America.

Riparian forests are structured and maintained by their hydrology. Woody riparian plants typically adapt to the local flood regime to maximise their likelihood of survival and reproductive success. Understanding how extant trees form and reproduce in response to flood disturbance is crucial for predicting vegetation changes and informing restoration. Working in a temperate evergreen riparian forest, we aimed to determine whether disturbance-based responses of plants found in other ecosystems also typify woody plants in riparian forests where disturbances are often mild or chronic, non-lethal, annual events. Using plant surveys and 20-year modelled hydrological data, we examined whether (1) the morphology (main stem diameter, height, crown width, crown extent, stem leaning) and (2) reproduction type (sexual and asexual reproduction) and extent of three dominant woody species (Eucalyptus camphora, Leptospermum lanigerum and Melaleuca squarrosa) vary with flood regime (flood frequency and flood duration); and (3) whether different morphology is associated with different reproductive strategies. Increased flooding generally resulted in increased stem numbers and greater stem leaning—morphologies associated with asexual reproduction—of our study species. More frequent flooding also reduced plant size and sexual reproduction in E. camphora. Sexual reproduction in the studied species was more common in taller plants with single, more upright stems in good condition. Flexible morphology and plastic reproductive strategy may constitute an adaptation of trees to mild or chronic disturbance in floodplains. Our findings suggest that flood regime (i.e. variable frequency and duration of flooding events) is critical to the structural integrity and self-maintenance of species-diverse riparian forests.

1. The stress-gradient hypothesis (SGH) provides a conceptual framework for explaining howenvironmental context determines the nature of biotic interactions. It may be also useful for predicting geographic variability in the effect of management interventions on biological invasions. We aimed to test hypotheses consistent with the SGH to explain context dependency in bamboo invasion of secondary forests in Japan, and establish a predictive understanding of forest management impacts on invasion.2. We use a priori physiological knowledge of invasive giant bamboo, Phyllostachys bambusoides, to generate hypotheses consistent with the SGH. We modelled variation in giant bamboo occupancy within 810 secondary forest plots across the broad environmental gradients of Japan using a national vegetation database.3. Consistent with the SGH, we find that the effect of tree canopy cover on bamboo occupancy depends on interactions between solar radiation and mean annual temperature. In cool regions with high solar radiation – stressful conditions for bamboo – shade cast by dense canopies facilitates invasion. However, in warmer regions that are more benign, dense canopies tend to inhibit spread via competition for light, space and other resources.4. We used our findings to characterise geographic variability in the effect of forest thinning, a widespread management intervention used to enhance forest biodiversity, on the risk of bamboo spread into secondary forests in Japan. Thinning forest canopies to increase understorey light radiation should limit bamboo spread in cooler regions, while tree planting to increase canopy shade should limit bamboo spread in warmer regions. We provide evidence that the SGH can inform practical recommendations for invasive species control.

The effects of altered nutrient supplies and herbivore density on species diversity vary with spatial scale, because coexistence mechanisms are scale dependent. This scale dependence may alter the shape of the species–area relationship (SAR), which can be described by changes in species richness (S) as a power function of the sample area (A): S = cAz, where c and z are constants. We analysed the effects of experimental manipulations of nutrient supply and herbivore density on species richness across a range of scales (0.01–75 m²) at 30 grasslands in 10 countries. We found that nutrient addition reduced the number of species that could co-occur locally, indicated by the SAR intercepts (log c), but did not affect the SAR slopes (z). As a result, proportional species loss due to nutrient enrichment was largely unchanged across sampling scales, whereas total species loss increased over threefold across our range of sampling scales.

A large body of research shows that biodiversity loss can reduce ecosystem functioning. However, much of the evidence for this relationship is drawn from biodiversity-ecosystem functioning experiments in which biodiversity loss is simulated by randomly assembling communities of varying species diversity, and ecosystem functions are measured. This random assembly has led some ecologists to question the relevance of biodiversity experiments to real-world ecosystems, where community assembly or disassembly may be non-random and influenced by external drivers, such as climate, soil conditions or land use. Here, we compare data from real-world grassland plant communities with data from two of the largest and longest-running grassland biodiversity experiments (the Jena Experiment in Germany and BioDIV in the United States) in terms of their taxonomic, functional and phylogenetic diversity and functional-trait composition. We found that plant communities of biodiversity experiments cover almost all of the multivariate variation of the real-world communities, while also containing community types that are not currently observed in the real world. Moreover, they have greater variance in their compositional features than their real-world counterparts. We then re-analysed a subset of experimental data that included only ecologically realistic communities (that is, those comparable to real-world communities). For 10 out of 12 biodiversity-ecosystem functioning relationships, biodiversity effects did not differ significantly between the full dataset of biodiversity experiments and the ecologically realistic subset of experimental communities. Although we do not provide direct evidence for strong or consistent biodiversity-ecosystem functioning relationships in real-world communities, our results demonstrate that the results of biodiversity experiments are largely insensitive to the exclusion of unrealistic communities and that the conclusions drawn from biodiversity experiments are generally robust. By comparing data from real-world grassland communities with data from two of the longest-running grassland biodiversity-ecosystem functioning experiments, the authors show that conclusions derived from experimental systems are robust to the removal of unrealistic experimental communities., Nature Ecology & Evolution, 4 (11), ISSN:2397-334X

Our ability to predict invasions has been hindered by the seemingly idiosyncratic context-dependency of individual invasions. However, we argue that robust and useful generalisations in invasion science can be made by considering “invasion syndromes” which we define as “a combination of pathways, alien species traits, and characteristics of the recipient ecosystem which collectively result in predictable dynamics and impacts, and that can be managed effectively using specific policy and management actions”. We describe this approach and outline examples that highlight its utility, including: cacti with clonal fragmentation in arid ecosystems; small aquatic organisms introduced through ballast water in harbours; large ranid frogs with frequent secondary transfers; piscivorous freshwater fishes in connected aquatic ecosystems; plant invasions in high-elevation areas; tall-statured grasses; and tree-feeding insects in forests with suitable hosts. We propose a systematic method for identifying and delimiting invasion syndromes. We argue that invasion syndromes can account for the context-dependency of biological invasions while incorporating insights from comparative studies. Adopting this approach will help to structure thinking, identify transferrable risk assessment and management lessons, and highlight similarities among events that were previously considered disparate invasion phenomena.

Este artículo contiene 16 páginas, 6 figuras., Invasive species are a global threat to biodiversity and there is a pressing need to better understand why some species become invasive outside of their native range, and others do not. One explanation for invasive species success is their release from concurrent natural enemies upon introduction to the nonnative range. The so-called enemy release hypothesis (ERH) has conflicting support, depending upon the ecosystem and species investigated. To date, most studies testing the generality of the ERH have focused on terrestrial ecosystems. Here, we tested whether enemy release might contribute to the success of the invasive non-native brown seaweeds Undaria pinnatifida and Sargassum muticum in the United Kingdom. We conducted choice and no choice experiments to determine herbivore preference on these invaders relative to six functionally-similar native species. We also measured and compared species traits associated with defence against herbivory (carbon to nitrogen ratio, polyphenolic concentration, tensile strength, and compensatory growth). There were no differences in the biomass consumed between invasive and native species for either choice or no choice tests. The carbon to nitrogen ratio (a measure of nutritional quality) was significantly lower for S. muticum compared to the three native fucoid species, but measures of the other three defence traits were similar or even greater for invasive species compared with native species. Taken together, it is unlikely that the ERH applies to invasive seaweeds in the northeast Atlantic, suggesting that other factors may contribute to the success of invasive species in this system., ALM was supported by the Natural Environment Research Council (Grant number NE/L002531/1) and a School of Biological Sciences Studentship at the University of Southampton. JAC was supported by The Royal Society (RG160911). DAS was supported by a UKRI Future Leaders Fellowship (MR/S032827/1).

Fertilisation experiments have demonstrated that nutrient availability is a key determinant of biomass production and carbon sequestration in grasslands. However, the influence of nutrients in explaining spatial variation in grassland biomass production has rarely been assessed. Using a global dataset comprising 72 sites on six continents, we investigated which of 16 soil factors that shape nutrient availability associate most strongly with variation in grassland aboveground biomass. Climate and N deposition were also considered. Based on theory-driven structural equation modelling, we found that soil micronutrients (particularly Zn and Fe) were important predictors of biomass and, together with soil physicochemical properties and C:N, they explained more unique variation (32%) than climate and N deposition (24%). However, the association between micronutrients and biomass was absent in grasslands limited by NP. These results highlight soil properties as key predictors of global grassland biomass production and point to serial co-limitation by NP and micronutrients.

The total impact of an alien species was conceptualised as the product of its range size, local abundance and per-unit effect in a seminal paper by Parker et al. (Biol Invasions 1:3-19, 1999). However, a practical approach for estimating the three components has been lacking. Here, we generalise the impact formula and, through use of regression models, estimate the relationship between the three components of impact, an approach we term GIRAE (Generalised Impact = Range size × Abundance × per-unit Effect). We discuss how GIRAE can be applied to multiple types of impact, including environmental impacts, damage and management costs. We propose two methods for applying GIRAE. The species-specific method computes the relationship between impact, range size, abundance and per-unit effect for a given species across multiple invaded sites or regions of different sizes. The multi-species method combines data from multiple species across multiple sites or regions to calculate a per-unit effect for each species and is computed using a single regression model. The species-specific method is more accurate, but it requires a large amount of data for each species and assumes a constant per-unit effect for a species across the invaded area. The multi-species method is more easily applicable and data-parsimonious, but assumes the same relationship between impact, range size and abundance for all considered species. We illustrate these methods using data about money spent managing plant invasions in different biomes of South Africa. We found clear differences between species in terms of money spent per unit area invaded, with per-unit expenditure varying substantially between biomes for some species-insights that are useful for monitoring and evaluating management. GIRAE offers a versatile and practical method that can be applied to many different types of data to better understand and manage the impacts of biological invasions.The online version contains supplementary material available at 10.1007/s10530-022-02836-0.

We introduce the AusTraits database - a compilation of values of plant traits for taxa in the Australian flora (hereafter AusTraits). AusTraits synthesises data on 448 traits across 28,640 taxa from field campaigns, published literature, taxonomic monographs, and individual taxon descriptions. Traits vary in scope from physiological measures of performance (e.g. photosynthetic gas exchange, water-use efficiency) to morphological attributes (e.g. leaf area, seed mass, plant height) which link to aspects of ecological variation. AusTraits contains curated and harmonised individual- and species-level measurements coupled to, where available, contextual information on site properties and experimental conditions. This article provides information on version 3.0.2 of AusTraits which contains data for 997,808 trait-by-taxon combinations. We envision AusTraits as an ongoing collaborative initiative for easily archiving and sharing trait data, which also provides a template for other national or regional initiatives globally to fill persistent gaps in trait knowledge., Measurement(s)plant traitTechnology Type(s)digital curationSample Characteristic - OrganismViridiplantaeSample Characteristic - LocationAustralia Machine-accessible metadata file describing the reported data: 10.6084/m9.figshare.14545755

Due to massive energetic investments in woody support structures, trees are subject to unique physiological, mechanical, and ecological pressures not experienced by herbaceous plants. When considering trait relationships across the entire plant kingdom, plant trait frameworks typically must omit traits unique to large woody species, thereby limiting our understanding of how these distinct ecological pressures shape trait relationships in trees. Here, by considering 18 functional traits—reflecting leaf economics, wood structure, tree size, reproduction, and below-ground allocation—we quantify the major axes of variation governing trait expression of trees worldwide. We show that trait variation within and across angiosperms and gymnosperms is captured by two independent processes: one reflecting tree size and competition for light, the other reflecting leaf photosynthetic capacity and nutrient economies. By exploring multidimensional relationships across clusters of traits, we further identify a representative set of seven traits which captures the majority of variation in form and function in trees: maximum tree height, stem conduit diameter, specific leaf area, seed mass, bark thickness, root depth, and wood density. Collectively, this work informs future trait-based research into the functional biogeography of trees, and contributes to our fundamental understanding of the ecological and evolutionary controls on forest biodiversity and productivity worldwide.

Although the high costs of invasion are frequently cited and are a key motivation for environmental management and policy, synthesised data on invasion costs are scarce. Here, we quantify and examine the monetary costs of biological invasions in the United Kingdom (UK) using a global synthesis of reported invasion costs. Invasive alien species have cost the UK economy between US$6.9 billion and $17.6 billion (£5.4 – £13.7 billion) in reported losses and expenses since 1976. Most costs were reported for the entire UK or Great Britain (97%); country-scale cost reporting for the UK's four constituent countries was scarce. Reports of animal invasions were the costliest ($4.7 billion), then plant ($1.3 billion) and fungal ($206.7 million) invasions. Reported damage costs (i.e. excluding management costs) were higher in terrestrial ($4.8 billion) than aquatic or semi-aquatic environments ($29.8 million), and primarily impacted agriculture ($4.2 billion). Invaders with earlier introduction years accrued significantly higher total invasion costs. Invasion costs have been increasing rapidly since 1976, and have cost the UK economy $157.1 million (£122.1 million) per annum, on average. Published information on specific economic costs included only 42 of 520 invaders reported in the UK and was generally available only for the most intensively studied taxa, with just four species contributing 90% of species-specific costs. Given that many of the invasive species lacking cost data are actively managed and have well-recognised impacts, this suggests that cost information is incomplete and that totals presented here are vast underestimates owing to knowledge gaps. Financial expenditure on managing invasions is a fraction (37%) of the costs incurred through damage from invaders; greater investments in UK invasive species research and management are, therefore, urgently required.

1.Non-native species can dominate plant communities by competitively displacing native species, or because environmental change creates conditions favourable to non-native species but unfavourable to native species. We need to disentangle these mechanisms so that management can target competitively dominant species and reduce their impacts.2. Joint-species distribution models (JSDMs) can potentially quantify competitive impacts by simultaneously modelling how species respond to environmental variation and to changes in community composition. We describe a JSDM to model variation in plant cover and show how this can be applied to compositional data to detect dominant competitors that cause other species to decline in abundance.3. We applied the model to an experiment in an invaded grassy-woodland community in Australia where we manipulated biomass removal (through slashing and fencing to prevent grazing by kangaroos) along a fertility gradient. Non-native species dominated plant cover at high fertility sites in the absence of biomass removal. Results from the JSDM identified three of the 72 non-native plant species (Bromus diandrus, Acetosella vulgaris and especially Avena fatua) as having a strong competitive impact on the community, driving changes in composition and reducing the cover of both native and non-native species, particularly in the absence of grazing. The dominant non-native grasses Bromus diandrus and Avena fatua were among the tallest species in the community and had the greatest impact on shorter-statured species, most likely through competition for light under conditions of high fertility and low grazing.4. Synthesis. We demonstrate a method to measure competitive impact using a joint-species distribution model, which allowed us to identify the species driving compositional change through competitive displacement, and where on the landscape competitive impacts were greatest. This information is central to managing plant invasions: by targeting dominant non-native species with large competitive impacts, management can reduce impacts where they are greatest. We provide details of the modelling procedure and reproducible code to encourage further application.

Understanding drivers of success for alien species can inform on potential future invasions. Recent conceptual advances highlight that species may achieve invasiveness via performance along at least three distinct dimensions: 1) local abundance, 2) geographic range size, and 3) habitat breadth in naturalized distributions. Associations among these dimensions and the factors that determine success in each have yet to be assessed at large geographic scales. Here, we combine data from over one million vegetation plots covering the extent of Europe and its habitat diversity with databases on species' distributions, traits, and historical origins to provide a comprehensive assessment of invasiveness dimensions for the European alien seed plant flora. Invasiveness dimensions are linked in alien distributions, leading to a continuum from overall poor invaders to super invaders - abundant, widespread aliens that invade diverse habitats. This pattern echoes relationships among analogous dimensions measured for native European species. Success along invasiveness dimensions was associated with details of alien species' introduction histories: earlier introduction dates were positively associated with all three dimensions, and consistent with theory-based expectations, species originating from other continents, particularly acquisitive growth strategists, were among the most successful invaders in Europe. Despite general correlations among invasiveness dimensions, we identified habitats and traits associated with atypical patterns of success in only one or two dimensions - for example, the role of disturbed habitats in facilitating widespread specialists. We conclude that considering invasiveness within a multidimensional framework can provide insights into invasion processes while also informing general understanding of the dynamics of species distributions., Deutsche Forschungsgemeinschaft (264740629) Grantová Agentura České Republiky (19-28491X) Grantová Agentura České Republiky (19-28807X) Grantová Agentura České Republiky (RVO 67985939) Austrian Science Fund (I 2086 - B29) Bundesministerium für Bildung und Forschung (01LC1807A) Eusko Jaurlaritza (IT299-10) National Research Foundation of Korea (2018R1C1B6005351) University of Latvia (AAp2016/B041//Zd2016/AZ03) Villum Fonden (16549)

Functional traits are commonly used in invasion studies due to their potential to generalise mechanisms to become invasive across species, which can inform both border screening and post-border management tools. Despite decades of research, however, finding general rules of traits that confer invasiveness has proven challenging. Trait-based invasiveness studies often compare discrete groups of invasive and non-invasive (or native) species and attribute their differences in invasiveness to their functional differences. Yet no universal definition of ‘invasiveness’ exists, and thus different criteria can be used to decide which species are invasive and non-invasive. Less often, invasiveness studies present more nuanced, continuous metrics of invasiveness, avoiding the need to define invasive/non-invasive boundaries. Unlike categorical classifications of invasiveness, continuous metrics have the potential to reflect individual demographic processes – i.e. species growth, reproduction and spread. Hence, they acknowledge the possibility that different ecological mechanisms enable invasions and that those mechanisms may relate to traits differently. Initial excitement for the use of functional traits in invasion forecast has turned into realisation that invasions are highly context-dependent processes; i.e. traits promoting invasiveness are strongly linked to the ecological and historical context. Accordingly, risk assessment protocols often include information on human use, species’ known invasion history and species’ environmental preferences, along with species’ traits. Robust prediction of species invasiveness based on functional traits, and its application for biosecurity protocols, requires critical thinking on which traits promote distinct mechanisms of invasion, consideration of the context and validation of findings across contrasting case studies.

Trait-based invasiveness studies typically categorize exotic species as invasive or noninvasive, implicitly assuming species form two homogenous groups. However, species can become invasive in different ways (e.g., high abundance, fast spread), likely relying on different functional traits to do so. As such, binary classification may obscure traits associated with invasiveness. We tested whether (1) the way in which invasiveness is quantified influences its correlation with functional traits and (2) different demography-based metrics are related to different sets of traits. Using a case study of 251 herbs exotic to Victoria, Australia, we quantified species' invasiveness using 10 metrics: four continuous, demography-based dimensions of invasiveness (spread rate, local abundance, geographic and environmental range sizes) and six binary classifications of invasiveness (based on alternative sources and invasion criteria). We examined the correlation between species' invasiveness and a set of four traits known to relate to plant demography and invasion. Then, we examined whether different demographic dimensions of invasiveness were better explained by different sets of traits. We found that the way invasiveness was quantified was important: different traits were linked with different invasiveness metrics, and some traits showed opposite effects across metrics. Species with fast spread were either tall with small seeds (i.e., good colonizers), or had heavy, animal-dispersed seeds. Plants with a large environmental range had greater plasticity for some traits. Locally abundant plants had low SLA and heavy seeds (i.e., strong competitors). Animal dispersal was also key to reach a large geographic range. No traits were consistently related to the six binary classifications. Our results indicate that exotic plants are invasive in different ways and rely on different combinations of traits to be so. Some traits (e.g., seed mass) had complex relationships with invasion: they apparently promote, hampered, or had no influence on different dimensions of invasiveness. Our findings are consistent with the notion that plant species use strategies that may be near optimal under some, but not all, ecological conditions. Compared to binary classifications of invasiveness, the use of invasiveness dimensions advances clearer hypothesis testing in invasion science.

1AbstractThe introduction stage is usually overlooked in trait-based studies of invasiveness, implicitly assuming that species introductions are random. However, human activities promote the movement of specific types of species. Thus, species deliberately introduced for distinct purposes (e.g. gardening, forestry) or as contaminants of human commodities (e.g. stowaway) will likely show particular traits. If species with certain traits have been preferentially introduced (i.e. introduction bias), some traits may have been mistakenly linked to species’ invasion abilities due to their influence on introduction probability.In this work, we propose a theoretical framework with different scenarios of introduction bias. The introduction scenarios are: (1) Random introduction, independent from traits; (2) Biologically biased introduction, following the worldwide distribution of the trait; and (3) Human biased introduction, following a theoretical introduction pathway that favours the introduction of species with high values of the trait. We evaluate how the introduced trait distributions in these scenarios may affect trait distributions in naturalized and invasive species pools under different hypothesized associations between traits and the probabilities of naturalization and invasion. The aim of this work is to identify situations where ignoring introduction bias may lead to spurious correlations being found between species’ traits and species’ ability to become naturalized or invasive.Our framework strongly points to the need to evaluate the traits of species that have become naturalized or invasive along with the traits of species that have failed to do so in order to unravel any existing introduction bias that may confound the correlation between species’ traits and invasion success. Overlooking a possible introduction bias may lead to the overestimation of the correlation between the trait and the species’ invasion ability, especially in cases when the pool of introduced species shows extreme values of the trait distribution (as compared to a random introduction). Trait-based studies that deserve special attention to avoid undesired effects of introduction bias on their findings are: those that investigate naturalization using only the pool of naturalized species, and those studies that examine invasiveness by comparing invasive species with native species.

Much research effort has been invested in understanding ecological impacts of invasive alien species (IAS) across ecosystems and taxonomic groups, but empirical studies about economic effects lack synthesis. Using a comprehensive global database, we determine patterns and trends in economic costs of aquatic IAS by examining: (i) the distribution of these costs across taxa, geographic regions and cost types; (ii) the temporal dynamics of global costs; and (iii) knowledge gaps, especially compared to terrestrial IAS. Based on the costs recorded from the existing literature, the global cost of aquatic IAS conservatively summed to US$345 billion, with the majority attributed to invertebrates (62%), followed by vertebrates (28%), then plants (6%). The largest costs were reported in North America (48%) and Asia (13%), and were principally a result of resource damages (74%); only 6% of recorded costs were from management. The magnitude and number of reported costs were highest in the United States of America and for semi-aquatic taxa. Many countries and known aquatic alien species had no reported costs, especially in Africa and Asia. Accordingly, a network analysis revealed limited connectivity among countries, indicating disparate cost reporting. Aquatic IAS costs have increased in recent decades by several orders of magnitude, reaching at least US$23 billion in 2020. Costs are likely considerably underrepresented compared to terrestrial IAS; only 5% of reported costs were from aquatic species, despite 26% of known invaders being aquatic. Additionally, only 1% of aquatic invasion costs were from marine species. Costs of aquatic IAS are thus substantial, but likely underreported. Costs have increased over time and are expected to continue rising with future invasions. We urge increased and improved cost reporting by managers, practitioners and researchers to reduce knowledge gaps. Few costs are proactive investments; increased management spending is urgently needed to prevent and limit current and future aquatic IAS damages.

1. The establishment of new botanic gardens in tropical regions highlights a need for weed risk assessment tools suitable for tropical ecosystems. The relevance of plant traits for invasion into tropical rainforests has not been well studied.2. Working in and around four botanic gardens in Indonesia where 590 alien species have been planted, we estimated the effect of four plant traits, plus time since species introduction, on: (a) the naturalization probability and (b) abundance (density) of naturalized species in adjacent native tropical rainforests; and (c) the distance that naturalized alien plants have spread from the botanic gardens.3. We found that specific leaf area (SLA) strongly differentiated 23 naturalized from 78 non‐naturalized alien species (randomly selected from 577 non‐naturalized species) in our study. These trends may indicate that aliens with high SLA, which had a higher probability of naturalization, benefit from at least two factors when establishing in tropical forests: high growth rates and occupation of forest gaps. Naturalized aliens had high SLA and tended to be short. However, plant height was not significantly related to species' naturalization probability when considered alongside other traits.4. Alien species that were present in the gardens for over 30 years and those with small seeds also had higher probabilities of becoming naturalized, indicating that garden plants can invade the understorey of closed canopy tropical rainforests, especially when invading species are shade tolerant and have sufficient time to establish.5. On average, alien species that were not animal dispersed spread 78 m further into the forests and were more likely to naturalize than animal‐dispersed species. We did not detect relationships between the measured traits and estimated density of naturalized aliens in the adjacent forests.6. Synthesis: Traits were able to differentiate alien species from botanic gardens that naturalized in native forest from those that did not; this is promising for developing trait‐based risk assessment in the tropics. To limit the risk of invasion and spread into adjacent native forests, we suggest tropical botanic gardens avoid planting alien species with fast carbon capture strategies and those that are shade tolerant., We demonstrate that alien plant species that spread from Indonesian botanic gardens and naturalize in adjacent native tropical rainforests have different traits to alien species that fail to naturalize. Species with high specific leaf area (SLA), small seeds and long residence times were more likely to naturalize than species that lacked these characteristics. Animal‐dispersed species spread over shorter distances and were less likely to naturalize than species not dispersed by animals. These trait‐based differences hold promise for the development of weed risk assessments specific to the tropics.

Phenotypic plasticity can mask population genetic differentiation, reducing the predictability of trait-environment relationships. In short-lived plants, reproductive traits may be more genetically determined due to their direct impact on fitness, whereas vegetative traits may show higher plasticity to buffer short-term perturbations. Combining a multi-treatment greenhouse experiment with global field observations for the short-lived Plantago lanceolata, we 1) disentangled the genetic and plastic responses of functional traits to a set of environmental drivers and 2) assessed the utility of trait-environment relationshisps inferred from observational data for predicting genetic differentiation. Reproductive traits showed distinct genetic differentiation that was highly predictable from observational data, but only when correcting traits for differences in their (labile) biomass component. Vegetative traits showed higher plasticity and contrasting genetic and plastic responses, leading to unpredictable trait patterns. Our study suggests that genetic differentiation may be inferred from observational data only for the traits most closely related with fitness.