Does disturbance,competition or resource limitation underlie Hieracium lepidulum invasion in New Zealand? Mechanisms of establishment and persistence,and functional differentiation among invasive and native species

The processes underlying plant invasions have been the subject of much ecological research. Understanding mechanisms of plant invasions are difficult to elucidate from observations, yet are crucial for ecological management of invasions. Hieracium lepidulum, an asteraceous invader in New Zealand, is a species for which several explanatory mechanisms can be raised. Alternative mechanisms, including competitive dominance, disturbance of resident vegetation allowing competitive release or nutrient resource limitation reducing competition with the invader are raised to explain invasion. We tested these hypotheses in two field experiments which manipulated competitive, disturbance and nutrient environments in pre‐invasion and post‐invasion vegetation. H. lepidulum and resident responses to environmental treatments were measured to allow interpretation of underlying mechanisms of establishment and persistence. We found that H. lepidulum differed in functional response profile from native species. We also found that other exotic invaders at the sites were functionally different to H. lepidulum in their responses. These data support the hypothesis that different invaders use different invasion mechanisms from one another. These data also suggest that functional differentiation between invaders and native resident vegetation may be an important contributing factor allowing invasion. H. lepidulum appeared to have little direct competitive effect on post‐invasion vegetation, suggesting that competition was not a dominant mechanism maintaining its persistence. There was weak support for disturbance allowing initial establishment of H. lepidulum in pre‐invasion vegetation, but disturbance did not lead to invader dominance. Strong support for nutrient limitation of resident species was provided by the rapid competitive responses with added nutrients despite presence of H. lepidulum. Rapid competitive suppression of H. lepidulum once nutrient limitation was alleviated suggests that nutrient limitation may be an important process allowing the invader to dominate. Possible roles of historical site degradation and/or invader‐induced soil chemical/microbial changes in nutrient availability are discussed.     

Effects of biological invasions on forest carbon sequestration

There has been a rapidly developing literature on the effects of some of the major drivers of global change on carbon (C) sequestration, particularly carbon dioxide (CO₂) enrichment, land use change, nitrogen (N) deposition and climate change. However, remarkably little attention has been given to one major global change driver, namely biological invasions. This is despite growing evidence that invasive species can dramatically alter a range of aboveground and belowground ecosystem processes, including those that affect C sequestration. In this review, we assess the evidence for the impacts of biological invaders on forest C stocks and C sequestration by biological invaders. We first present case studies that highlight a range of invader impacts on C sequestration in forest ecosystems, and draw on examples that involve invasive primary producers, decomposers, herbivores, plant pathogens, mutualists and predators. We then develop a conceptual framework for assessing the effects of invasive species on C sequestration impacts more generally, by identifying the features of biological invaders and invaded ecosystems that are thought to most strongly regulate C in forests. Finally we assess the implications of managing invasive species on C sequestration. An important principle that emerges from this review is that the direct effects of invaders on forest C are often smaller and shorter‐term than their indirect effects caused by altered nutrient availability, primary productivity or species composition, all of which regulate long‐term C pools and fluxes. This review provides a conceptual basis for improving our general understanding of biological invaders on ecosystem C, but also points to a paucity of primary data that are needed to determine the quantitative effects of invaders on ecosystem processes that drive C sequestration.     

Elevated success of multispecies bacterial invasions impacts community composition during ecological succession

Successful microbial invasions are determined by a species’ ability to occupy a niche in the new habitat whilst resisting competitive exclusion by the resident community. Despite the recognised importance of biotic factors in determining the invasiveness of microbial communities, the success and impact of multiple concurrent invaders on the resident community has not been examined. Simultaneous invasions might have synergistic effects, for example if resident species need to exhibit divergent phenotypes to compete with the invasive populations. We used three phylogenetically diverse bacterial species to invade two compositionally distinct communities in a controlled, naturalised in vitro system. By initiating the invader introductions at different stages of succession, we could disentangle the relative importance of resident community structure, invader diversity and time pre‐invasion. Our results indicate that multiple invaders increase overall invasion success, but do not alter the successional trajectory of the whole community.     

Using avatar species to model the potential distribution of emerging invaders

Aim Anticipating the potential distributions of emerging invasive species is complicated by the tendency for species distribution models to perform better when both native and invasive range data are available for model development. If invasive range data are lacking, species models are liable to under‐estimate distributions for emerging invaders, particularly for species that are not at equilibrium with their native range environment due to historical factors, dispersal limitation and/or ecological interactions. We demonstrate the potential to use well‐quantified niche shifts from established ‘avatar’ (i.e. the remote or virtual manifestation of an entity) invaders to develop plausible distributions for data‐poor emerging invaders contingent on niche shifts of similar magnitude or character. Location Global. Methods Using the globally invasive crayfishes Pacifastacus leniusculus and Procambarus clarkii as our avatar invaders, we quantify how niche position, size and structure differs between native and total ranges using Mahalanobis distance (a measure of multivariate similarity) and the climate predictors of annual minimum and maximum air temperature. We then generalize patterns of niche shift from these species to the emerging crayfish invader Cherax quadricarinatus. Results Some patterns of niche shifts were similar for Pacifastacus leniusculus and Procambarus clarkii, but niche shifts were of considerably greater magnitude for P. clarkii. When a native range model for C. quadricarinatus was modified with generalized niche shifts similar to Pacifastacus leniusculus and Procambarus clarkii, the potential global distribution for this species increased considerably, including many areas not identified by the native range model. Main conclusions We illustrate the potential to use avatar invaders to provide cautionary, niche shift‐assuming species distribution models for emerging invaders. Many theoretical and applied implications of the avatar species concept require additional investigation, including the development of frameworks to select appropriate avatar species and evaluate the performance of avatar‐derived models for emerging invaders. Despite these research needs, we believe this concept will have considerable utility for predicting vulnerability to invasion by data‐poor species; this is a critical management need because shifting pathways of introduction and climate change will produce many novel, emerging invasive species in the future.     

Invasive ants as back-seat drivers of native ant diversity decline in New Caledonia

Biological invasions are typically associated with disturbance, which often makes their impact on biodiversity unclear—biodiversity decline might be driven by disturbance, with the invader just being a ‘passenger’. Alternatively, an invader may act as a ‘back-seat driver’, being facilitated by disturbance that has already caused some biodiversity decline, but then causing further decline. Here we examine the interactive effects of anthropogenic fire and invasive ant species (Anoplolepis gracilipes or Wasmannia auropunctata) on native ant diversity in New Caledonia, a globally recognized biodiversity hotspot. We first examined native ant diversity at nine paired burnt and unburnt sites, with four pairs invaded by Anoplolepis, 5 years after an extensive fire. In the absence of invasion, native epigaeic ants were resilient to fire, but native ant richness and the abundance of Forest Opportunists were markedly lower in invaded burnt sites. Second, we examined native ant diversity along successional gradients from human-derived savanna to natural rainforest in the long-term absence of fire, where there was a disconnection between disturbance-mediated variation in microhabitat and the abundance of the disturbance specialist Wasmannia. All native ant diversity responses (total abundance, richness, species composition, functional group richness and the abundance of Forest Opportunists) declined independently of microhabitat variables but in direct association with high Wasmannia abundance. Our results indicate that invasive ants are acting as back-seat drivers of biodiversity decline in New Caledonia, with invasion facilitated by disturbance but then causing further biodiversity decline.     

Shifts in dominance of native and invasive plants in experimental patches of vegetation

While it is widely accepted that disturbance often promotes the process of plant invasion, few data are available on specific relationships between different types of disturbance and success of plant invaders. Nor is much known about variation in invader behaviour in response to changing composition of the dominant native species in the recipient habitats. In the middle Main valley near Würzburg (Germany), two invasive Brassicaceae forbs, Bunias orientalis and Rorippa austriaca, are frequently associated with a small number of dominant native species in the herbaceous vegetation of productive, disturbed sites. Variation in the relative dominance of these species suggests high variability between sites in the factors influencing the local invasion process. In our study, we examined variation in regeneration mode, type and frequency of disturbance, and native species composition, and considered life history differences between the two invasive species as factors contributing to the distinct dominance patterns. These factors were tested in a controlled field experiment in which individuals of the two invasive species and four native species were grown in mixed stands for almost 3 years. The development of the plants was monitored by cover values and all plants were harvested at the end of the study. All factors investigated contributed considerably to the development of distinct dominance patterns in the species mixtures. Relative to the natives, both invasive species were promoted by regeneration from vegetative fragments as compared to development from sexual offspring. While both invasive species were generally promoted by disturbance relative to the natives, they differed in their response to distinct types of disturbance, and these differences were strongly affected by species composition. Our results indicate that invasion success of R. austriaca may depend more on soil disturbance, soil transport and deposition, while B. orientalis is expected to particularly expand at mown sites that do not have dense cover by meadow grasses. Whether the native resident vegetation is mainly composed of successional competitors that suffer from disturbance or of ruderal competitors that compete early and strongly with the invasive species may be of critical importance for invasion by the two species.     

Multiple successional pathways of boreal forest stands in central Canada

Predicting forest composition change through time is a key challenge in forest management. While multiple successional pathways are theorized for boreal forests, empirical evidence is lacking, largely because succession has been inferred from chronosequence and dendrochronological methods. We tested the hypotheses that stands of compositionally similar overstory may follow multiple successional pathways depending on time since last stand‐replacing fire (TSF), edaphic conditions, and presence of intermediate disturbances. We used repeated measurements from combining sequential aerial photography and ground surveys for 361 boreal stands in central Canada. Stands were measured in 8–15 yr intervals over a ~ 60 yr period, covering a wide range of initial stand conditions. Multinomial logistic regression was used to analyze stand type transitions. With increasing TSF, stands dominated by shade‐intolerant Pinus banksiana, Populus sp., and Betula papyrifera demonstrated multiple pathways to stands dominated by shade‐tolerant Picea sp., Abies balsamea, and Thuja occidentalis. Their pathways seemed largely explained by neighborhood effects. Succession of stands dominated by shade‐tolerant species, with an exception of stands dominated by Picea sp., was not related to TSF, but rather dependent on edaphic conditions and presence of intermediate disturbances. Varying edaphic conditions caused divergent pathways with resource limited sites being dominated by nutrient‐poor tolerant species, and richer sites permitting invasion of early successional species and promoting species mixtures during succession. Intermediate disturbances promoted deciduous persistence and species diversity in A. balsamea and mixed‐conifer stands, but no evidence was detected to support “disturbance accelerated succession”. Our results demonstrate that in the prolonged absence of stand‐replacing disturbance boreal forest stands undergo multiple succession pathways. These pathways are regulated by neighborhood effects, resource availability, and presence of intermediate disturbance, but the relative importance of these regulators depends on initial stand type. The observed divergence of successional pathways supports the resource‐ratio hypothesis of plant succession.     

Responses of a native beetle to novel exotic plant species with varying invasion history

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Native species richness buffers invader impact in undisturbed but not disturbed grassland assemblages

Many systems are prone to both exotic plant invasion and frequent natural disturbances. Native species richness can buffer the effects of invasion or disturbance when imposed in isolation, but it is largely unknown whether richness provides substantial resistance against invader impact in the face of disturbance. We experimentally examined how disturbance (drought/burning) influenced the impact of three exotic invaders (Centaurea stoebe, Linaria dalmatica, or Potentilla recta) on native abundance across a gradient of species richness, using previously constructed grassland assemblages. We found that invaders had higher cover in experimentally disturbed plots than in undisturbed plots across all levels of native species richness. Although exotic species varied in cover, all three invaders had significant impacts on native cover in disturbed plots. Regardless of disturbance, however, invader cover diminished with increasing richness. Invader impacts on native cover also diminished at higher richness levels, but only in undisturbed plots. In disturbed plots, invaders strongly impacted native cover across all richness levels, as disturbance favoured invaders over native species. By examining these ecological processes concurrently, we found that disturbance exacerbated invader impacts on native abundance. Although diversity provided a buffering effect against invader impact without disturbance, the combination of invasion and disturbance markedly depressed native abundance, even in high richness assemblages.     

A welcome can of worms? Hypoxia mitigation by an invasive species

Invasive species and bottom‐water hypoxia both constitute major global threats to the diversity and integrity of marine ecosystems. These stressors may interact with unexpected consequences, as invasive species that require an initial environmental disturbance to become established can subsequently become important drivers of ecological change. There is recent evidence that improved bottom‐water oxygen conditions in coastal areas of the northern Baltic Sea coincide with increased abundances of the invasive polychaetes Marenzelleria spp. Using a reactive‐transport model, we demonstrate that the long‐term bioirrigation activities of dense Marenzelleria populations have a major impact on sedimentary phosphorus dynamics. This may facilitate the switch from a seasonally hypoxic system back to a normoxic system by reducing the potential for sediment‐induced eutrophication in the upper water column. In contrast to short‐term laboratory experiments, our simulations, which cover a 10‐year period, show that Marenzelleria has the potential to enhance long‐term phosphorus retention in muddy sediments. Over time bioirrigation leads to a substantial increase in the iron‐bound phosphorus content of sediments while reducing the concentration of labile organic carbon. As surface sediments are maintained oxic, iron oxyhydroxides are able to persist and age into more refractory forms. The model illustrates mechanisms through which Marenzelleria can act as a driver of ecological change, although hypoxic disturbance or natural population declines in native species may be needed for them to initially become established. Invasive species are generally considered to have a negative impact; however, we show here that one of the main recent invaders in the Baltic Sea may provide important ecosystem services. This may be of particular importance in low‐diversity systems, where disturbances may dramatically alter ecosystem services due to low functional redundancy. Thus, an environmental problem in one region may be either exacerbated or alleviated by a single species from another region, with potentially ecosystem‐wide consequences.     

Are invasive species drivers of native species decline or passengers of habitat modification? A case study of the impact of the common myna (Acridotheres tristis) on Australian bird species

Habitat modification and invasive species are significant drivers of biodiversity decline. However, distinguishing between the impacts of these two drivers on native species can be difficult. For example, habitat modification may reduce native species abundance, while an invasive species may take advantage of the new environment. This scenario has been described as the driver‐passenger model, with ‘passengers’ taking advantage of habitat modification and ‘drivers’ causing native species decline. Therefore, research must incorporate both habitat modification and invasive species impact to successfully investigate native species decline. In this paper, we used the common myna (Acridotheres tristis) as a case study to investigate the driver‐passenger model. We investigated changes in bird abundance, over 2 years, in relation to different habitat types and common myna abundance. We hypothesized that the common myna is both a passenger of habitat change and a driver of some bird species decline. Our results indicated that the abundance of many native species is greater in high tree density nature reserves, while the common myna was uncommon in these areas. Common myna abundance was almost three times higher in urban areas than nature reserves and declined rapidly as tree density in nature reserves increased. Our findings indicated that the common myna is primarily a passenger of habitat change. However, we also observed negative associations between common myna abundance and some bird species. We stress the importance of simultaneously investigating both invasive species impact and habitat modification. We suggest habitat restoration could be a useful tool for both native species recovery and invasive species control. Understanding the drivers of native species decline will help inform impact mitigation and direct further research.     

Carbohydrate scarcity increases foraging activities and aggressiveness in the ant Prolasius advenus (Hymenoptera: Formicidae)

1. Behavioural responses to varying macronutrient availability are increasingly studied in highly invasive ant species to better understand their ecological success. However, such work is lacking in relation to native ant species confronted with biological invaders. 2. Here the link between diet and behaviour was examined in Prolasius advenus, a native ant from New Zealand facing intense competition for food with invasive insects, including social wasps. A long‐term laboratory experiment was conducted to assess the impact of protein and carbohydrate restriction on several behavioural parameters. 3. Ants shifted their food‐collecting activity towards the least available macronutrient in both protein‐limited and carbohydrate‐limited colonies. But when lacking carbohydrate, they also strongly increased their efforts to discover resources. After 10 weeks, the proportion of the colony patrolling outside of the nest was eight times higher than at the initiation of the experiment. This high patrolling activity was then maintained for several weeks and resulted in a higher efficiency to explore a novel territory. Moreover, ants fed a carbohydrate‐limited diet engaged in longer aggressive acts towards conspecifics. 4. These behavioural responses to carbohydrate scarcity may, in part, enhance the ability of P. advenus to resist a competing invader under natural conditions. While much of the previous research has linked diet and behavioural dominance in invasive ants, the present study shows that conducting similar investigations in the native species to which they are confronted may shed light on the mechanisms behind biotic resistance and the ability of some native species to coexist with highly abundant invaders.     

Historic land use influences contemporary establishment of invasive plant species

The legacy of agricultural land use can have widespread and persistent effects on contemporary landscapes. Although agriculture can lead to persistent changes in soil characteristics and plant communities, it remains unclear whether historic agricultural land use can alter the likelihood of contemporary biological invasions. To understand how agricultural land-use history might interact with well-known drivers of invasion, we conducted factorial manipulations of soil disturbance and resource additions within non-agricultural remnant sites and post-agricultural sites invaded by two non-native Lespedeza species. Our results reveal that variation in invader success can depend on the interplay of historic land use and contemporary processes: for both Lespedeza species, establishment was greater in remnant sites, but soil disturbance enhanced establishment irrespective of land-use history, demonstrating that contemporary processes can help to overcome legacy constraints on invader success. In contrast, additions of resources known to facilitate seedling recruitment (N and water) reduced invader establishment in post-agricultural but not in remnant sites, providing evidence that interactions between historic and contemporary processes can also limit invader success. Our findings thus illustrate that a consideration of historic land use may help to clarify the often contingent responses of invasive plants to known determinants of invasibility. Moreover, in finding significantly greater soil compaction at post-agricultural sites, our study provides a putative mechanism for historic land-use effects on contemporary invasive plant establishment. Our work suggests that an understanding of invasion dynamics requires knowledge of anthropogenic events that often occur decades before the introduction of invasive propagules.     

Leaf ontogenetic dependence of light acclimation in invasive and native subtropical trees of different successional status

In the Bonin Islands of the western Pacific where the light environment is characterized by high fluctuations due to frequent typhoon disturbance, we hypothesized that the invasive success of Bischofia javanica Blume (invasive tree, mid‐successional) may be attributable to a high acclimation capacity under fluctuating light availability. The physiological and morphological responses of B. javanica to both simulated canopy opening and closure were compared against three native species of different successional status: Trema orientalis Blume (pioneer), Schima mertensiana (Sieb. et Zucc.) Koidz (mid‐successional) and Elaeocarpus photiniaefolius Hook.et Arn (late‐successional). The results revealed significant species‐specific differences in the timing of physiological maturity and phenotypic plasticity in leaves developed under constant high and low light levels. For example, the photosynthetic capacity of T. orientalis reached a maximum in leaves that had just fully expanded when grown under constant high light (50% of full sun) whereas that of E. photiniaefolius leaves continued to increase until 50 d after full expansion. For leaves that had just reached full expansion, T. orientalis, having high photosynthetic plasticity between high and low light, exhibited low acclimation capacity under the changing light (from high to low or low to high light). In comparison with native species, B. javanica showed a higher degree of physiological and morphological acclimation following transfer to a new light condition in leaves of all age classes (i.e. before and after reaching full expansion). The high acclimation ability of B. javanica in response to changes in light availability may be a part of its pre‐adaptations for invasiveness in the fluctuating environment of the Bonin Islands.     

Existing and emerging high impact invasive species are characterized by higher functional responses than natives

Predicting ecological impacts of invasive species and identifying potentially damaging future invaders are research priorities. Since damage by invaders is characterized by their depletion of resources, comparisons of the ‘functional response’ (FR; resource uptake rate as a function of resource density) of invaders and natives might predict invader impact. We tested this by comparing FRs of the ecologically damaging ‘world''s worst’ invasive fish, the largemouth bass (Micropterus salmoides), with a native equivalent, the Cape kurper (Sandelia capensis), and an emerging invader, the sharptooth catfish (Clarias gariepinus), with the native river goby (Glossogobius callidus), in South Africa, a global invasion hotspot. Using tadpoles (Hyperolius marmoratus) as prey, we found that the invaders consumed significantly more than natives. Attack rates at low prey densities within invader/native comparisons reflected similarities in predatory strategies; however, both invasive species displayed significantly higher Type II FRs than the native comparators. This was driven by significantly lower prey handling times by invaders, resulting in significantly higher maximum feeding rates. The higher FRs of these invaders are thus congruent with, and can predict, their impacts on native communities. Comparative FRs may be a rapid and reliable method for predicting ecological impacts of emerging and future invasive species.     

Replacement of a native freshwater macroinvertebrate species by an invader: implications for biological water quality monitoring

The rate of freshwater invasions may be increasing, and macroinvertebrate invaders can have significant impacts on native macroinvertebrate assemblage structure through biotic interactions. More pollution-tolerant invaders can often replace native species. We examined implications of a species replacement for accurate biological monitoring of river systems using biotic indices. Our study uses Northern Ireland and the Isle of Man as examples of countries that possess river networks with many riverine macroinvertebrate assemblages subject to invasion. The introduced amphipod crustacean Gammarus pulex has replaced the native species G. duebeni celticus in many rivers in N. Ireland and the Isle of Man. Extensive seasonal data sets (119 sites) from three river networks, Lough Neagh and the Lagan in N. Ireland, and island-wide in the Isle of Man, were used to investigate the assumed equivalence of the native and invader in biotic indices concerned with the water quality monitoring system. Based on the derivation of the Biological Monitoring Working Party (BMWP) score, the Average Score Per Taxon (ASPT), as an example of a commonly used biotic index of water quality, we found index scores were lower in G. pulex sites compared to G. d. celticus-only sites. This indicated that assemblages were dominated by taxa more tolerant of organic pollution in the invader sites and more sensitive in the native sites. Inclusion of the invader in generation of the ASPT index, overinflated the ASPT values obtained compared to those with the native’s inclusion. This questions the accuracy of the ASPT and similar indices in rivers where the invader had replaced the native. We argue that with invasion pressures increasing, the validity of water quality indices such as the BMWP/ASPT needs to be re-examined in catchments where invaders have replaced natives. Indices such as the BMWP/ASPT are based on family level taxa and are inevitably coarse in their resolution given the wide range of water qualities tolerated by different genera within families. We argue that this resolution is even more compromised by the presence of very pollution-tolerant invaders, who may have replaced natives in disturbed or degraded river systems. The whole structure of water quality indices such as the BMWP/ASPT may need revising to take into account the presence of invasive species within monitored assemblages.     

Native herbivores and environmental heterogeneity as mediators of an exotic grass invasion

Given that many exotic plant species throughout the world are having large ecological and economic effects, it is vital to understand the forces that mediate their success in novel landscapes. Both native herbivores and recipient ecosystems can have substantial effects on the performance of exotic plant species, and may interact with each other or vary in their effects over time. Unfortunately, few studies have evaluated the importance of these kinds of context‐dependent effects. Here, we use a 17‐year‐old exclosure experiment stratified across a coastal grassland in northern California to address the relative importance of a reintroduced mammalian herbivore, tule elk (Cervus canadensis nannodes), and environmental heterogeneity in mediating the growth, abundance, and recruitment of a problematic grass invader, Holcus lanatus. We found that elk reduced Holcus abundance, aboveground biomass, percent cover, frequency, and seedling recruitment, but that these effects often varied among habitat types, with effects being greater in open grasslands than shrub‐dominated grasslands. The performance of Holcus populations also varied significantly among habitat types, with the invader usually having the greatest success in Baccharis‐dominated grasslands. Our results suggest that environmental heterogeneity had much greater influence on Holcus success than elk, and that these effects were due largely to soil pH and moisture. The negative effects of elk on Holcus appeared after 4 years and did not intensify after an additional 13 years. Furthermore, despite their negative effects, these prominent herbivores did not prevent the spread of Holcus into previously uninvaded areas. Our research highlights the importance of assessing the individual and interactive effects of native herbivores and environmental heterogeneity on the success of invasive, exotic plant species. It emphasizes the reality that the negative effects of herbivores on exotic plant species will often vary across heterogeneous landscapes and may be insufficient to prevent the expansion of these invaders.     

Non-random recruitment of invader species in experimental grasslands

To assess potential effects of seed limitation, characteristics of invader species and characteristics of established plant communities on recruitment success, we conducted a split-plot experiment factorially combining three weeding treatments corresponding to increasing successional age (regular weeding相似文献   

Competitive displacement or biotic resistance? Disentangling relationships between community diversity and invasion success of tall herbs and shrubs

Questions: Are negative invasion–diversity relationships due to biotic resistance of the invaded plant community or to post‐invasion displacement of less competitive species? Do invasion–diversity relationships change with habitat type or resident traits? Location/species: Lowlands and uplands of western and southern Germany, Heracleum mantegazzianum; mountain range in central Germany, Lupinus polyphyllus; and coastal dunes of northwest Germany, Rosa rugosa. Methods: We tested the significance and estimated regression slopes of invasion–diversity relationships using generalized linear (mixed effects) models relating invader cover and habitat type to species richness in different plant groups, stratified based on size, life cycle and community association. Results: We found negative, positive and neutral relationships between invader cover and species richness. There were negative linear correlations of invader cover with small plant species throughout, but no negative linear correlation with tall species. Invasion–diversity relationships tended to be more negative in early‐successional habitats, such as dunes or abandoned grasslands, than in late‐successional habitats. Conclusions: Invasion diversity–relationships are complex; they vary among habitat types and among different groups of resident species. Negative invasion–diversity relationships are due to asymmetric competitive displacement of inferior species and not due to biotic resistance. Small species are displaced in early‐successional habitats, while there is little effect on persistence of tall species.     

Biotic resistance buffers the effects of nutrient enrichment on the success of a highly invasive aquatic plant

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