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Plants growing outside their native range may be confronted by new regimes of herbivory, but how this affects plant chemical defence profiles has rarely been studied. Using Plantago lanceolata as a model species, we investigated whether introduced populations show significant differences from native populations in several growth and chemical defence traits. Plantago lanceolata (ribwort plantain) is an herbaceous plant species native to Europe and Western Asia that has been introduced to numerous countries worldwide. We sampled seeds from nine native and 10 introduced populations that covered a broad geographic and environmental range and performed a greenhouse experiment, in which we infested half of the plants in each population with caterpillars of the generalist herbivore Spodoptera littoralis. We then measured size‐related and resource‐allocation traits as well as the levels of constitutive and induced chemical defence compounds in roots and shoots of P. lanceolata. When we considered the environmental characteristics of the site of origin, our results revealed that populations from introduced ranges were characterized by an increase in chemical defence compounds without compromising plant biomass. The concentrations of iridoid glycosides and verbascoside, the major anti‐herbivore defence compounds of P. lanceolata, were higher in introduced populations than in native populations. In addition, introduced populations exhibited greater rates of herbivore‐induced volatile organic compound emission and diversity, and similar chemical diversity based on untargeted analyses of leaf methanol extracts. In conclusion, the geographic origin of the populations had a significant influence on morphological and chemical plant traits, suggesting that P. lanceolata populations are not only adapted to different environments in their native range, but also in their introduced range. Read the free Plain Language Summary for this article on the Journal blog.

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.

Rates of species colonisation and extirpation are increasing in plant communities world‐wide. Colonisation could potentially help compensate for, or compound, resident diversity loss that results from global environmental change. We use a multifactorial seed addition grassland experiment to examine relationships between plant colonisation, resident species diversity and key community assembly factors over 3 years. By manipulating colonist seed rate, imposing disturbance and examining abundance and diversity impacts of 14 formerly absent sown colonists in communities that varied in successional stage and time since agricultural abandonment, we were able to disentangle effects of global change factors (species introduction, novel disturbance and land use change) that are usually confounded. Evidence suggested that cover abundance of sown colonists was most strongly influenced by successional stage of recipient communities, though number of growing seasons was also important for the group of seven colonists with resource conservative ‘slow’ life history traits. Colonist type, seed rate and disturbance had weaker relationships with colonist cover. Factors affecting sown colonist cover were highly conditional. A negative relationship between plot‐level disturbance and colonist cover in early successional communities meant that, despite a positive relationship in late succession, colonisation was negatively related to disturbance overall, defying theoretical expectations. Non‐sown resident diversity was negatively related to colonist cover and positively related to successional stage. Resource acquisitive colonists with ‘fast’ life history traits appeared to limit cover of ‘slow colonists’ when the two groups were sown together, likely reflecting niche pre‐emption. Communities at earlier stages of succession had lower resident diversity and experienced higher levels of colonisation than communities at later stages of succession. Elevated colonisation and lower re

The enemy release hypothesis (ERH) is the best-known hypothesis explaining high performance (e.g. rapid population growth) of exotic species. However, the current framing of the ERH does not explicitly link evidence of enemy release with exotic performance. This leads to uncertainty regarding the role of enemy release in biological invasions. Here, we demonstrate that the effect of enemy release on exotic performance is the product of three factors: enemy impact, enemy diversity, and host adaptation. These factors are modulated by seven contexts: time since introduction, resource availability, phylogenetic relatedness of exotic and native species, host-enemy asynchronicity, number of introduction events, type of enemy, and strength of growth-defence trade-offs. ERH-focused studies frequently test different factors under different contexts. This can lead to inconsistent findings, which typifies current evidence for the ERH. For example, over 80% of meta-analyses fail to consider ecological contexts which can alter study findings; we demonstrate this by re-analysing a recent ERH synthesis. Structuring the ERH around factors and contexts promotes generalisable predictions about when and where exotic species may benefit from enemy release, empowering effective management. Our mechanistic factor-context framework clearly lays out the evidence required to support the ERH, unifies many enemy-related invasion hypotheses, and enhances predictive capacity.

Human modification of waterways has reduced flooding in many river systems, leading to the decline of riparian forests, which rely on flooding for their regeneration. Coppicing may help to promote the persistence of riparian trees by triggering resprouting and vegetative regeneration. The vigour of resprouting plants can vary with timing and height of coppicing and may depend on stored non-structural carbohydrate reserves like starch, the availability of which can vary seasonally. However, starch storage dynamics and the resprouting potential of broad-leafed evergreen riparian trees is not well understood. We coppiced two riparian tree species, Eucalyptus camphora and Melaleuca squarrosa, at two different times (autumn, spring) and at two different heights (0 cm and 90 cm). Over 52 weeks, we regularly quantified shoot growth and changes in the starch storage pool size, compared to uncoppiced control trees, in different tree organs (root and stem) and estimated the final shoot volume. The final shoot volume did not differ significantly between coppice treatments. Trees coppiced in autumn had a greater reliance on stored starch while they remained leafless (without shoots) over winter. Trees cut at 90 cm had more starch reserves due to remaining stems but also had higher biomass maintenance costs. Starch storage varied seasonally only in E. camphora, with starch concentrations in control trees increasing over winter and decreasing over summer. Although coppice timing and height affected use of stored starch, resprouting in our study species was not limited by starch availability - both species regenerated vegetatively to recover from physical disturbance. Thus, coppicing may be an efficient means to promote rejuvenation and persistence of tree species where site and tree condition are degraded and no longer support recruitment.

Riparian forest loss and degradation due to river-floodplain disconnection is a global problem. Prospects for the maintenance and recruitment of riparian trees via restored flooding can be uncertain, in part due to competition from understorey vegetation and limited availability of tree propagules. In a field-based trial, we assessed the response of a keystone riparian tree to restored flooding, reduced competition and seed addition. We built diversion weirs to reconnect floodplains supporting riparian forest. Using a multiple control-intervention-reference study design with two restored flooding (intervention) sites, two naturally-engaged (reference) and two dry (control) sites, we assessed seedling recruitment of the dominant tree, Eucalyptus camphora, with and without clearance of understorey vegetation and with and without seed addition. We also assessed the growth of extant trees using dendrometers and seedfall using funnel traps at all six sites. Our weirs resulted in extensive flooding of the adjacent floodplain, while control sites remained dry. Flooding increased seed germination, seedling establishment, tree growth and seedfall. However, seed germination benefits were only realised where understorey vegetation was cleared and seed was added. Seedling establishment was also limited by dry summer conditions, except where flooding duration was longest (∼6 months cf. < 3 months). Restored flooding via river-floodplain reconnection is likely to promote the rehabilitation of riparian forests degraded by flow regulation or stream modification through benefits at multiple tree life-history stages. However, widespread tree recruitment may require complementary works that reduce competition via clearing of understorey vegetation and alleviate seed limitation by direct seeding or planting.

Context dependence is widely invoked to explain disparate results in ecology. It arises when the magnitude or sign of a relationship varies due to the conditions under which it is observed. Such variation, especially when unexplained, can lead to spurious or seemingly contradictory conclusions, which can limit understanding and our ability to transfer findings across studies, space, and time. Using examples from biological invasions, we identify two types of context dependence resulting from four sources: mechanistic context dependence arises from interaction effects; and apparent context dependence can arise from the presence of confounding factors, problems of statistical inference, and methodological differences among studies. Addressing context dependence is a critical challenge in ecology, essential for increased understanding and prediction.

Due to massive energetic investments in woody support structures, trees are subject to unique physiological, mechanical, and ecological pressures not experienced by herbaceous plants. Despite a wealth of studies exploring trait relationships across the entire plant kingdom, the dominant traits underpinning these unique aspects of tree form and function remain unclear. Here, by considering 18 functional traits, encompassing leaf, seed, bark, wood, crown, and root characteristics, we quantify the multidimensional relationships in tree trait expression. We find that nearly half of trait variation is captured by two axes: one reflecting leaf economics, the other reflecting tree size and competition for light. Yet these orthogonal axes reveal strong environmental convergence, exhibiting correlated responses to temperature, moisture, and elevation. By subsequently exploring multidimensional trait relationships, we show that the full dimensionality of trait space is captured by eight distinct clusters, each reflecting a unique aspect of tree form and function. Collectively, this work identifies a core set of traits needed to quantify global patterns in functional biodiversity, and it contributes to our fundamental understanding of the functioning of forests worldwide.

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 non-native 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.

UNLABELLED: 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. SUPPLEMENTARY INFORMATION: The online vers

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 high-priority 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.

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.

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Riparian forest ecosystems are declining globally. Many floodplains no longer flood and thus cease to satisfy the hydrologic requirements for riparian tree maintenance and regeneration. To promote woody riparian plant recruitment where flood regimes have been altered by flow regulation, effective approaches to restoration need to be developed. We implemented a landscape-scale experiment in a remnant, temperate floodplain forest. By constructing two weirs within channelized reaches of a stream, we redirected flows into networks of historic distributary channels, which facilitated widespread floodplain inundation. Using a control-reference-impact study design, we assessed the establishment and growth of planted seedlings of three woody species (Eucalyptus camphora, Leptospermum lanigerum and Melaleuca squarrosa) over 13 months in response to flooding achieved by floodplain reconnection. Planted seedlings had higher height and diameter growth rates at both induced (19–29 cm, 1 mm) and naturally flooded (34–44 cm, 3–5 mm) than at non-flooded (4–10 cm, −5 to −3 mm) sites. However, survival rates and temporal growth patterns differed between species according to variation in flood duration and soil moisture, illustrating the different hydrological requirements of the coexisting species. This highlights that variable flooding and drying patterns are essential to create recruitment niches for different riparian plant species and shows the importance of river-floodplain connectivity for providing adequate flooding regimes. Our study demonstrates the suitability of two complementary restoration approaches – restoring hydrology and active revegetation – for promoting the regeneration of riparian forests.

Community and invasion ecology have mostly grown independently. There is substantial overlap in the processes captured by different models in the two fields, and various frameworks have been developed to reduce this redundancy and synthesize information content. Despite broad recognition that community and invasion ecology are interconnected, a process-based framework synthesizing models across these two fields is lacking. Here we review 65 representative community and invasion models and propose a common framework articulated around six processes (dispersal, drift, abiotic interactions, within-guild interactions, cross-guild interactions, and genetic changes). The framework is designed to synthesize the content of the two fields, provide a general perspective on their development, and enable their comparison. The application of this framework and of a novel method based on network theory reveals some lack of coherence between the two fields, despite some historical similarities. Community ecology models are characterized by combinations of multiple processes, likely reflecting the search for an overarching theory to explain community assembly and structure, drawing predominantly on interaction processes, but also accounting largely for the other processes. In contrast, most models in invasion ecology invoke fewer processes and focus more on interactions between introduced species and their novel biotic and abiotic environment. The historical dominance of interaction processes and their independent developments in the two fields is also reflected in the lower level of coherence for models involving interactions, compared to models involving dispersal, drift, and genetic changes. It appears that community ecology, with a longer history than invasion ecology, has transitioned from the search for single explanations for patterns observed in nature to investigate how processes may interact mechanistically, thereby generating and testing hypotheses. Our framework paves t

The stress‐gradient hypothesis (SGH) provides a conceptual framework for explaining how environmental 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. 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. 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. Synthesis and applications. We provide evidence that the stress‐gradient hypothesis can inform practical recommendations for invasive species control. We characterised 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.

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.

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 prev

Unprecedented rates of introduction and spread of non-native species pose burgeoning challenges to biodiversity, natural resource management, regional economies, and human health. Current biosecurity efforts are failing to keep pace with globalization, revealing critical gaps in our understanding and response to invasions. Here, we identify four priority areas to advance invasion science in the face of rapid global environmental change. First, invasion science should strive to develop a more comprehensive framework for predicting how the behavior, abundance, and interspecific interactions of non-native species vary in relation to conditions in receiving environments and how these factors govern the ecological impacts of invasion. A second priority is to understand the potential synergistic effects of multiple co-occurring stressors— particularly involving climate change—on the establishment and impact of non-native species. Climate adaptation and mitigation strategies will need to consider the possible consequences of promoting non-native species, and appropriate management responses to non-native species will need to be developed. The third priority is to address the taxonomic impediment. The ability to detect and evaluate invasion risks is compromised by a growing deficit in taxonomic expertise, which cannot be adequately compensated by new molecular technologies alone. Management of biosecurity risks will become increasingly challenging unless academia, industry, and governments train and employ new personnel in taxonomy and systematics. Fourth, we recommend that internationally cooperative biosecurity strategies consider the bridgehead effects of global dispersal networks, in which organisms tend to invade new regions from locations where they have already established. Cooperation among countries to eradicate or control species established in bridgehead regions should yield greater benefit than independent attempts by individual countries to exclude these specie

QUESTION: Northern peatlands are increasingly threatened by wildfire. Severe peatland wildfires can provide opportunities for new non‐peatland species to colonise post fire. Changes in plant colonisation could lead to longer‐term shifts in community composition, compromising recovery of peatland structure and function. Understanding the process of post‐fire recovery can thus inform restoration action and help restore peatland vascular plant communities. In this study, we ask: what drives initial vascular plant recovery following a peatland wildfire? LOCATION: Stalybridge moors, England (commonly referred to as the Saddleworth moors). METHODS: We used a series of vegetation surveys and seed germination experiments to identify the composition of vascular plant community one‐year post fire, along with potential propagule sources. We combined this with plant trait data and, using a series of null models, compared observed community trait values against random species assemblages. RESULTS: Our data suggests that plant species are able to arrive at the burned site through multiple non‐exclusive recolonisation pathways. This includes colonisation through the soil seed bank, along with dispersal from surrounding unburned peatland and non‐peatland vegetation. The composition and structure of the recolonised communities was largely determined by the ability of species to reach the post‐fire site from these donor communities. This resulted in a post‐fire community composed of species possessing lower seed masses relative to the wider pool of potential colonisers. CONCLUSIONS: Our results highlight propagule availability as a driver of post‐wildfire vascular plant recovery. This provides opportunities for new non‐peatland species to colonise, potentially driving changes in the direction of vegetation recovery. Ensuring the availability of peatland species following a wildfire could therefore be key to the immediate recovery of these systems.

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.

Plant–soil feedbacks (PSF) and functional traits are two active but not well theoretically integrated areas of research. However, PSF and traits are both affected by life‐history evolution, so the two should theoretically be related. We provide a conceptual framework to link plant functional traits to two types of PSF metrics, and hypothesize that individual PSF (plant performance in conspecific vs. heterospecific soil) should be related to the fast–slow trait spectrum, whereas pairwise PSF (the sum of the individual feedbacks for two species growing in each other's soils) should be related to trait dissimilarity. We performed meta‐analyses to test these hypotheses by compiling two datasets, one dataset consisting of individual PSF values and plant trait values (specific leaf area, SLA; leaf N concentration, LNC; specific root length, SRL; fine root diameter, FRD; plant height; seed mass), and the second consisting of pairwise PSF values and trait dissimilarity. Our meta‐analyses showed that individual PSF values were more negative in faster‐growing species with greater SLA, LNC and SRL, supporting the growth–defence trade‐off hypothesis. Plant height was positively correlated with individual PSF, perhaps because large, long‐lived plants defend against pathogens better than smaller, shorter‐lived plants. We also found that larger‐seeded species had more positive or less negative PSF, likely reflecting greater tolerance of soil pathogens. The direction of relationships between trait dissimilarity and pairwise PSF varied with trait identity. Dissimilarities in SRL and FRD were negatively correlated with pairwise PSF while height dissimilarity was positively correlated with pairwise PSF. The contrasting relationships may reflect distinct links between trait dissimilarity and niche and fitness differences. Synthesis. Our results demonstrate how an integration of PSF and trait‐based approaches can advance plant community ecology.

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.

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.

Genetic differentiation and phenotypic plasticity jointly shape intraspecific trait variation, but their roles differ among traits. In short-lived plants, reproductive traits may be more genetically determined due to their impact on fitness, whereas vegetative traits may show higher plasticity to buffer short-term perturbations. Combining a multi-treatment greenhouse experiment with observational field data throughout the range of a widespread short-lived herb, Plantago lanceolata, we (1) disentangled genetic and plastic responses of functional traits to a set of environmental drivers and (2) assessed how genetic differentiation and plasticity shape observational trait-environment relationships. Reproductive traits showed distinct genetic differentiation that largely determined observational patterns, but only when correcting traits for differences in biomass. Vegetative traits showed higher plasticity and opposite genetic and plastic responses, masking the genetic component underlying field-observed trait variation. Our study suggests that genetic differentiation may be inferred from observational data only for the traits most closely related to fitness.

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 t

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 a

BACKGROUND AND AIMS: Since its emergence in the mid-20th century, invasion biology has matured into a productive research field addressing questions of fundamental and applied importance. Not only has the number of empirical studies increased through time, but also has the number of competing, overlapping and, in some cases, contradictory hypotheses about biological invasions. To make these contradictions and redundancies explicit, and to gain insight into the field's current theoretical structure, we developed and applied a Delphi approach to create a consensus network of 39 existing invasion hypotheses. RESULTS: The resulting network was analysed with a link-clustering algorithm that revealed five concept clusters (resource availability, biotic interaction, propagule, trait and Darwin's clusters) representing complementary areas in the theory of invasion biology. The network also displays hypotheses that link two or more clusters, called connecting hypotheses, which are important in determining network structure. The network indicates hypotheses that are logically linked either positively (77 connections of support) or negatively (that is, they contradict each other; 6 connections). SIGNIFICANCE: The network visually synthesizes how invasion biology's predominant hypotheses are conceptually related to each other, and thus, reveals an emergent structure - a conceptual map - that can serve as a navigation tool for scholars, practitioners and students, both inside and outside of the field of invasion biology, and guide the development of a more coherent foundation of theory. Additionally, the outlined approach can be more widely applied to create a conceptual map for the larger fields of ecology and biogeography.

When plants establish outside their native range, their ability to adapt to the new environment is influenced by both demography and dispersal. However, the relative importance of these two factors is poorly understood. To quantify the influence of demography and dispersal on patterns of genetic diversity underlying adaptation, we used data from a globally distributed demographic research network comprising 35 native and 18 nonnative populations of Plantago lanceolata Species-specific simulation experiments showed that dispersal would dilute demographic influences on genetic diversity at local scales. Populations in the native European range had strong spatial genetic structure associated with geographic distance and precipitation seasonality. In contrast, nonnative populations had weaker spatial genetic structure that was not associated with environmental gradients but with higher within-population genetic diversity. Our findings show that dispersal caused by repeated, long-distance, human-mediated introductions has allowed invasive plant populations to overcome environmental constraints on genetic diversity, even without strong demographic changes. The impact of invasive plants may, therefore, increase with repeated introductions, highlighting the need to constrain future introductions of species even if they already exist in an area.

There is strong evidence for a phylogenetic signal in the degree to which species share co-evolved biotic partners and in the outcomes of biotic interactions. This implies there should be a phylogenetic signal in the outcome of feedbacks between plants and the soil microbiota they cultivate. However, attempts to identify a phylogenetic signal in plant-soil feedbacks have produced mixed results. Here we clarify how phylogenetic signals could arise in plant-soil feedbacks and use a recent compilation of data from feedback experiments to identify: whether there is a phylogenetic signal in the outcome of plant-soil feedbacks; and whether any signal arises through directional or divergent changes in feedback outcomes with evolutionary time. We find strong evidence for a divergent phylogenetic signal in feedback outcomes. Distantly related plant species show more divergent responses to each other's soil microbiota compared with closely related plant species. The pattern of divergence implies occasional co-evolutionary shifts in how plants interact with soil microbiota, with strongly contrasting feedback responses among some plant lineages. Our results highlight that it is difficult to predict feedback outcomes from phylogeny alone, other than to say that more closely related species tend to have more similar responses.

Effective control of an invasive species is frequently used to infer positive outcomes for the broader ecosystem. In many situations, whether the removal of an invasive plant is of net benefit to biodiversity is poorly assessed. We undertook a 10-year study on the effects of invasive shrub management (bitou bush, Chrysanthemoides monilifera ssp. rotundata) on native flora and fauna in a eucalypt forest in south-eastern Australia. Bitou bush eradication is a management priority, yet the optimal control regime (combination of herbicide spray and fire) is difficult to implement, meaning managed sites have complex management histories that vary in effectiveness of control. Here we test the long-term response of common biodiversity indicators (species richness, abundance and diversity of native plants, birds, herpetofauna and small mammals) to both the management, and the post-management status of bitou bush (% cover). While average bitou bush cover decreased with management, bitou bush consistently occurred at around half of our managed sites despite control efforts. The relationship between biodiversity and bitou bush cover following management differed from positive, neutral or negative among species groups and indicators. Native plant cover was lower under higher levels of bitou bush cover, but the abundance of birds and small mammals were positively related to bitou bush cover. Evidence suggests that the successful control of an invader may not necessarily result in beneficial outcomes for all components of biodiversity.

Introduced species threaten native biodiversity, but whether exotic species can competitively displace native species remains contested. Building on theory that predicts multi-species coexistence based on a competition-colonisation tradeoff, we derive a mechanistic basis by which human-mediated species invasions could cause extinctions through competitive displacement. In contrast to past invasions, humans principally introduce modern invaders, repeatedly and in large quantities, and in ways that can facilitate release from enemies and competitors. Associated increases in exotic species' propagule rain, survival and competitive ability could enable some introduced species to overcome the tradeoffs that constrain all other species. Using evidence from metacommunity models, we show how species introductions could disrupt species coexistence, generating extinction debts, especially when combined with other forms of anthropogenic environmental change. Even though competing species have typically coexisted following past biogeographic migrations, the multiplicity and interactive impacts of today's threats could change some exotic species into agents of extinction.

Detecting exotic plant species is essential for invasive species management. By accounting for factors likely to affect species' detection rates (e.g. survey conditions, observer experience), detectability models can help choose search methods and allocate search effort. Integrating information on species' traits can refine detectability models, and might be particularly valuable if these traits can help improve estimates of detectability where data on particular species are rare. Analysing data collected during line transect distance sampling surveys in Indonesia, we used a multi-species hierarchical distance sampling model to evaluate how plant height, leaf size, leaf shape, and survey location influenced plant species detectability in secondary tropical rainforests. Detectability of the exotic plant species increased with plant height and leaf size. Detectability varied among the different survey locations. We failed to detect a clear effect of leaf shape on detectability. This study indicates that information on traits might improve predictions about exotic species detection, which can then be used to optimise the allocation of search effort for efficient species management. The innovation of the study lies in the multi-species distance sampling model, where the distance-detection function depends on leaf traits and height. The method can be applied elsewhere, including for different traits that may be relevant in other contexts. Trait-based multispecies distance sampling can be a practical approach for sampling exotic shrubs, herbs, or grasses species in the understorey of tropical forests.

Summary Restoration of degraded plant communities requires understanding of community assembly processes. Human land use can influence plant community assembly by altering environmental conditions and species’ dispersal patterns. Flooding, including from environmental flows, may counteract land use effects on wetland vegetation. We examined the influence of land use history and flood frequency on the functional composition of wetland plant communities along a regulated river. We applied fourth corner modelling to determine species’ trait‐based responses to flooding and land use by combining data on (i) the occupancy and abundance of species in propagule banks and standing vegetation, (ii) species traits, and (iii) environmental conditions of 22 standing vegetation and 108 soil propagule bank study sites. We used analysis of deviance to test how well each dataset characterised trait–environment interactions, and generalised linear models to identify traits related to species’ responses. The occupancy and abundance of native species in the propagule bank and standing vegetation increased with flood frequency and decreased with duration of agricultural land use. Species in standing vegetation with water‐borne propagule dispersal (hydrochory) showed similar trends. In contrast, species with higher specific leaf area were associated with longer land use duration. Identifying trait‐based differences in the propagule bank and standing vegetation can help disentangle effects of dispersal and environmental filters. The occupancy and abundance of hydrochorous species in standing vegetation were negatively related to land use duration, but hydrochorous species were positively related to land use duration based on their abundance in the propagule bank. This suggests that land use does not limit plant dispersal, but acts as an in situ abiotic filter, limiting species present in standing vegetation. Synthesis and applications. Land use duration and flood frequency have opposite e

QUESTION: How do contrasting influences of inundation and historical land uses affect restoration of soil propagule bank composition in floodplain wetlands? LOCATION: Northern Nature Reserve (large ephemeral floodplain), Macquarie Marshes, New South Wales, Australia. METHODS: We conducted germination assays on soil samples collected from fields with different land‐use histories, stratified along an inundation gradient. We used GLM to determine whether native and exotic species richness and abundance varied along gradients of inundation and land use. RESULTS: Species richness and plant abundance in soil propagule banks were positively related to inundation and negatively related to intense historic land use. The abundance of native species was significantly higher in more frequently inundated areas. Abundances of exotic and ruderal species were higher in areas of intense prior land use. Overall species richness was generally similar across land‐use histories. CONCLUSIONS: Land‐use legacies compromised the ability of propagule banks to rejuvenate native vegetation in this floodplain wetland, especially in less frequently flooded parts of the floodplain, which harboured more ruderal and exotic species. Negative effects of prior land use may be alleviated by increased inundation. Native soil propagule banks were remarkably intact, providing a reservoir for restoration of wetland vegetation, even in soils highly disturbed by up to 20 yr of agricultural cropping. With appropriate inundation, soil propagule banks in less degraded areas of the Macquarie Marshes can provide diverse mixtures of desired species in high abundance but, in highly degraded areas, full restoration may be delayed.

Despite a major research focus on human‐mediated reshuffling of plant communities, no coherent framework unites the numerous types of changes in abundances and distributions of native and non‐native species that are driven by human activities. Human driven vegetation change can occur through: non‐native species introductions; population outbreaks or collapses; range expansions or contractions; and range shifts of both native and non‐native species. Boundaries among these different types of floristic changes are not always distinct because of an overlap in the ecological, climatic, and anthropogenic processes that underpin them. We propose a new framework that connects various human‐mediated causes of vegetation change, highlights the spatial scales at which drivers act and the temporal scale at which plant assemblages respond, and provides critical insights for identifying and appropriately managing these changes.

A definitive list of invasive species traits remains elusive, perhaps due to inconsistent ways of identifying invasive species. Invasive species are typically identified using one or more of four demographic criteria (local abundance, geographic range, environmental range, spread rate), referred to here as the demographic dimensions of invasiveness. In 112 studies comparing invasive and non‐invasive plant traits, all 15 combinations of the four demographic dimensions were used to identify invasive species; 22% of studies identified invasive species solely by high abundance, while 25% ignored abundance. We used demographic data of 340 alien herbs classified as invasive or non‐invasive in Victoria, Australia, to test whether the demographic dimensions are independent and which dimensions influence invasive species listing in practice. Species' abundances, spread rates and range sizes were independent. Relative abundance best explained the invasiveness classification. However, invasive and non‐invasive species each spanned the full range of each demographic dimension, indicating that no dimension clearly separates invasive from non‐invasive species. Graminoids with longer minimum residence times were more frequently classified as invasive, as were forbs occurring near edges of native vegetation fragments. Synthesis. Conflating multiple forms of invasiveness, by not distinguishing invasive species that are identified using different demographic criteria, may obscure traits possessed by particular subsets of invasive species. Traits promoting high abundance likely differ from those enabling fast spread and broad ranges. Examining traits linked with the four demographic dimensions of invasiveness will highlight species at risk of becoming dominant, spreading quickly or occupying large ranges.

Invasive plant management is often justified in terms of conservation goals, yet progress is rarely assessed against these broader goals, instead focussing on short-term reductions of the invader as a measure of success. Key questions commonly remain unanswered including whether invader removal reverses invader impacts and whether management itself has negative ecosystem impacts. We addressed these knowledge gaps using a seven year experimental investigation of Bitou Bush, Chrysanthemoides monilifera subsp. rotundata. Our case study took advantage of the realities of applied management interventions for Bitou Bush to assess whether it is a driver or passenger of environmental change, and quantified conservation benefits relative to management costs of different treatment regimes. Among treatments examined, spraying with herbicide followed by burning and subsequent re-spraying (spray-fire-spray) proved the most effective for reducing the number of individuals and cover of Bitou Bush. Other treatment regimes (e.g. fire followed by spraying, or two fires in succession) were less effective or even exacerbated Bitou Bush invasion. The spray-fire-spray regime did not increase susceptibility of treated areas to re-invasion by Bitou Bush or other exotic species. This regime significantly reduced plant species richness and cover, but these effects were short-lived. The spray-fire-spray regime was the most cost-effective approach to controlling a highly invasive species and facilitating restoration of native plant species richness to levels characteristic of uninvaded sites. We provide a decision tree to guide management, where recommended actions depend on the outcome of post-treatment monitoring and performance against objectives. Critical to success is avoiding partial treatments and treatment sequences that may exacerbate invasive species impacts. We also show the value of taking advantage of unplanned events, such as wildfires, to achieve management objectives at reduced

Aim: Many factors facilitate biological invasions, making it difficult to determine their relative importance, especially when relying on survey data that include confounded variables. Incorporating information about species characteristics can improve inferences drawn from species-environment relationships, which can inform management. We seek to understand why the abundance of exotic, and not native, terrestrial plants is higher in riparian wetlands most impacted by flow regulation. Location: River Murray, SE Australia. Methods: We use variance components analysis and hierarchical generalised linear models to examine whether the positive relationship between flow regulation and proportional cover of exotic plants is driven by altered hydrological regimes, wetland drying and drought, superior colonisation ability of exotic species following disturbance or human-increased propagule pressure. Results: Of the four hypotheses, hydrological modification (indicated by flood magnitude) most likely drives invasion. Flow regulation may inhibit native species adapted to the historical hydrological regime, facilitating exotic species with different environmental ranges. A symptom of environmental change, invasion may have been exacerbated by drought, although it is unclear why. There was no indication that human-increased propagule pressure or colonisation ability facilitated invasion. Exotic cover was unrelated to proximity to towns, recent flood frequency and cattle grazing intensity. Additionally, similar proportions of exotic and native species were used in cultivation and, despite a higher proportion of exotics being known weeds, weed status was unrelated to exotic species occupancy. Overall, colonisation ability was unrelated to species' origin or response to water depth and hydrological change. Although exotics had higher specific leaf area and shorter longevity (indicative of higher colonisation ability), they had heavier (not lighter) seeds and did not differ in heig

Agricultural intensification is critical to meet global food demand, but intensification threatens native species and degrades ecosystems. Sustainable intensification (SI) is heralded as a new approach for enabling growth in agriculture while minimizing environmental impacts. However, the SI literature has overlooked a major environmental risk. Using data from eight countries on six continents, we show that few governments regulate conventionally bred pasture taxa to limit threats to natural areas, even though most agribusinesses promote taxa with substantial weed risk. New pasture taxa (including species, subspecies, varieties, cultivars, and plant-endophyte combinations) are bred with characteristics typical of invasive species and environmental weeds. By introducing novel genetic and endophyte variation, pasture taxa are imbued with additional capacity for invasion and environmental impact. New strategies to prevent future problems are urgently needed. We highlight opportunities for researchers, agribusiness, and consumers to reduce environmental risks associated with new pasture taxa. We also emphasize four main approaches that governments could consider as they build new policies to limit weed risks, including (i) national lists of taxa that are prohibited based on environmental risk; (ii) a weed risk assessment for all new taxa; (iii) a program to rapidly detect and control new taxa that invade natural areas; and (iv) the polluter-pays principle, so that if a taxon becomes an environmental weed, industry pays for its management. There is mounting pressure to increase livestock production. With foresight and planning, growth in agriculture can be achieved sustainably provided that the scope of SI expands to encompass environmental weed risks.