Short-term invasibility patterns in burnt and unburnt experimental Mediterranean grassland communities of varying diversities

This paper reports the findings of a short-term natural invasibility field study in constructed Mediterranean herbaceous communities of varying diversities, under a fire treatment. Three components of invasibility, i.e. species richness, density and biomass of invaders, have been monitored in burnt and unburnt experimental plots with resident diversity ranging from monocultures to 18-species mixtures. In general, species richness, density and biomass of invaders decreased significantly with the increase of resident species richness. Furthermore, the density and biomass of invading species were significantly influenced by the species composition of resident communities. Although aboveground biomass, leaf area index, canopy height and percent bare ground of the resident communities explained a significant part of the variation in the success of invading species, these covariates did not fully explain the effects of resident species richness. Fire mainly influenced invasibility via soil nutrient levels. The effect of fire on observed invasibility patterns seems to be less important than the effects of resident species richness. Our results demonstrate the importance of species richness and composition in controlling the initial stages of plant invasions in Mediterranean grasslands but that there was a lack of interaction with the effects of fire disturbance.     

Productivity, herbivory, and species traits rather than diversity influence invasibility of experimental phytoplankton communities

Biological invasions are a major threat to natural biodiversity; hence, understanding the mechanisms underlying invasibility (i.e., the susceptibility of a community to invasions by new species) is crucial. Invasibility of a resident community may be affected by a complex but hitherto hardly understood interplay of (1) productivity of the habitat, (2) diversity, (3) herbivory, and (4) the characteristics of both invasive and resident species. Using experimental phytoplankton microcosms, we investigated the effect of nutrient supply and species diversity on the invasibility of resident communities for two functionally different invaders in the presence or absence of an herbivore. With increasing nutrient supply, increased herbivore abundance indicated enhanced phytoplankton biomass production, and the invasion success of both invaders showed a unimodal pattern. At low nutrient supply (i.e., low influence of herbivory), the invasibility depended mainly on the competitive abilities of the invaders, whereas at high nutrient supply, the susceptibility to herbivory dominated. This resulted in different optimum nutrient levels for invasion success of the two species due to their individual functional traits. To test the effect of diversity on invasibility, a species richness gradient was generated by random selection from a resident species pool at an intermediate nutrient level. Invasibility was not affected by species richness; instead, it was driven by the functional traits of the resident and/or invasive species mediated by herbivore density. Overall, herbivory was the driving factor for invasibility of phytoplankton communities, which implies that other factors affecting the intensity of herbivory (e.g., productivity or edibility of primary producers) indirectly influence invasions.     

Productivity gradients cause positive diversity–invasibility relationships in microbial communities

Models predict that community invasibility generally declines with species diversity, a prediction confirmed by small‐scale experiments. Large‐scale observations and experiments, however, find that diverse communities tend to be more heavily invaded than simple communities. One hypothesis states that large‐scale environmental heterogeneity, which similarly influences native and invasive species, can cause a positive correlation between diversity and invasibility, overriding the local negative effects of diversity on invasibility. We tested this hypothesis using aquatic microbial communities consisting of protists and rotifers consuming bacteria and nanoflagellates. We constructed a productivity gradient to simulate large‐scale environmental heterogeneity, started communities with the same number of species along this gradient, and subjected equilibrial communities to invasion by non‐resident consumer species. Both invaders and most resident species increased their abundances with resource enrichment, resulting in a positive correlation between diversity and invasibility. Intraspecific interference competition within resident species and the positive effect of enrichment on the number of available resources probably accounted for the higher invasibility with enrichment. Our results provide direct experimental evidence that environmental heterogeneity in productivity can cause a positive diversity–invasibility relationship.     

Community phylogenetic diversity and abiotic site characteristics influence abundance of the invasive plant Rhamnus cathartica L.

Aims Theory predicts that the success of introduced species is related to the diversity of native species through trait-based processes. Abiotic site characteristics may also affect a site's susceptibility to invasion. We quantified resident plant species richness, phylogenetic diversity and several abiotic site characteristics for 24 oak forests in Minnesota, USA, to assess their impact on the abundance of a widespread, introduced terrestrial plant species, common buckthorn (Rhamnus cathartica L.). Specifically, we asked (1) whether resident species richness and phylogenetic diversity affected the abundance of R. cathartica and (2) what site characteristics explained the overall abundance of R. cathartica .Methods Our survey included 24 oak-dominated stands in Minnesota's deciduous forests. In each stand, we identified all species in 16 plots. We also measured a series of environmental site characteristics, including canopy openness (a proxy for light availability), percent bare soil, soil pH, percent sand, an index of propagule availability, duff layer thickness (a proxy for earthworm activity), an index of insolation and slope. For all species present in at least one site, we estimated a community phylogeny. We combined all site-level characteristics, including phylogenetic diversity of the resident plant species, in a multiple regression model to examine site level drivers of community invasibility.Important findings Results indicate that sites with higher overall plant phylogenetic diversity harbor less R. cathartica, even though native species richness was not significantly related to R. cathartica abundance. Regression analyses indicated that, in addition to resident species phylogenetic diversity, the most important predictors of R. cathartica abundance were canopy openness and the amount of bare soil, both positively related to the abundance of the invader. By combining the effects of abiotic site characteristics and resident species phylogenetic diversity in a model that predicted the abundance of R. cathartica, we were able to simultaneously account for a wide range of factors that might influence invasibility. Overall, our results suggest that management strategies aimed at reducing disturbances that lead to increased bare soil and light levels may be more successful if they also maximize phylogenetic diversity of the resident plant community.     

Is biotic resistance enhanced by natural variation in diversity?

Theories linking diversity to ecosystem function have been challenged by the widespread observation of more exotic species in more diverse native communities. Few studies have addressed the underlying processes by dissecting how biotic resistance to new invaders may be shaped by the same environmental influences that determine diversity and other community properties. In grasslands with heterogeneous soils, we added invaders and removed competitors to analyze the causes of invasion resistance. Abiotic resistance was measured using invader success in the absence of the resident community. Biotic resistance was measured as the reduction in invader success in the presence of the resident community. Invaders were most successful where biotic resistance was lowest and abiotic resistance was highest, confirming the dominant role of biotic resistance. Contrary to theory, though, biotic resistance was highest where both species richness and functional diversity were lowest. In the multivariate framework of a structural equation model, biotic resistance was independent of community diversity, and was highest where fertile soils led to high community biomass. Seed predation slightly augmented biotic resistance without qualitatively changing the results. Soil‐related genotypic variation in the invader also did not affect the results. We conclude that in natural systems, diversity may be correlated with invasibility and yet have little effect on biotic resistance to invasion. More generally, the environmental causes of variation in diversity should be considered when examining the potential functional consequences of diversity.     

Species-Rich Scandinavian Grasslands are Inherently Open to Invasion

Invasion of native habitats by alien or generalist species is recognized worldwide as one of the major causes behind species decline and extinction. One mechanism determining community invasibility, i.e. the susceptibility of a community to invasion, which has been supported by recent experimental studies, is species richness and functional diversity acting as barriers to invasion. We used Scandinavian semi-natural grasslands, exceptionally species-rich at small spatial scales, to examine this mechanism, using three grassland generalists and one alien species as experimental invaders. Removal of two putative functional groups, legumes and dominant non-legume forbs, had no effect on invasibility except a marginally insignificant effect of non-legume forb removal. The amount of removed biomass and original plot species richness had no effect on invasibility. Actually, invasibility was high already in the unmanipulated community, leading us to further examine the relationship between invasion and propagule pressure, i.e. the inflow of seeds into the community. Results from an additional experiment suggested that these species-rich grasslands are effectively open to invasion and that diversity may be immigration driven. Thus, species richness is no barrier to invasion. The high species diversity is probably in itself a result of the community being highly invasible, and species have accumulated at small scales during centuries of grassland management.     

Intraspecific genotypic richness and relatedness predict the invasibility of microbial communities

Biological invasions can lead to extinction events in resident communities and compromise ecosystem functioning. We tested the effect of two widespread biodiversity measurements, genotypic richness and genotypic dissimilarity on community invasibility. We manipulated the genetic structure of bacterial communities (Pseudomonas fluorescens) and submitted them to invasion by Serratia liquefaciens. We show that the two diversity measures impact on invasibility via distinct and additive mechanisms. Genotypic dissimilarity of the resident communities linearly increased productivity and in parallel decreased invasion success, indicating that high dissimilarity prevents invasion through niche pre-emption. By contrast, genotypic richness exerted a hump-shaped effect on invasion and was linked to the production of toxins antagonistic to the invader. This effect peaked at intermediate richness, suggesting that high richness levels may increase invasibility. Invasibility could be well predicted by the combination of these two mechanisms, documenting that both genotypic richness and dissimilarity need to be considered, if we are to understand the biotic properties determining the susceptibility of ecosystems to biological invasions.     

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.     

Giant invasive Heracleum persicum: Friend or foe of plant diversity?

The impact of invasion on diversity varies widely and remains elusive. Despite the considerable attempts to understand mechanisms of biological invasion, it is largely unknown whether some communities’ characteristics promote biological invasion, or whether some inherent characteristics of invaders enable them to invade other communities. Our aims were to assess the impact of one of the massive plant invaders of Scandinavia on vascular plant species diversity, disentangle attributes of invasible and noninvasible communities, and evaluate the relationship between invasibility and genetic diversity of a dominant invader. We studied 56 pairs of Heracleum persicum Desf. ex Fisch.‐invaded and noninvaded plots from 12 locations in northern Norway. There was lower native cover, evenness, taxonomic diversity, native biomass, and species richness in the invaded plots than in the noninvaded plots. The invaded plots had nearly two native species fewer than the noninvaded plots on average. Within the invaded plots, cover of H. persicum had a strong negative effect on the native cover, evenness, and native biomass, and a positive association with the height of the native plants. Plant communities containing only native species appeared more invasible than those that included exotic species, particularly H. persicum. Genetic diversity of H. persicum was positively correlated with invasibility but not with community diversity. The invasion of a plant community by H. persicum exerts consistent negative pressure on vascular plant diversity. The lack of positive correlation between impacts and genetic diversity of H. persicum indicates that even a small founder population may cause high impact. We highlight community stability or saturation as an important determinant of invasibility. While the invasion by H. persicum may decrease susceptibility of a plant community to further invasion, it severely reduces the abundance of native species and makes them more vulnerable to competitive exclusion.     

Fungal diversity increases soil fungistasis and resistance to microbial invasion by a non resident species

Biodiversity decline is a major concern for ecosystem functioning. Recent research efforts have been mostly focused on terrestrial plants, while, despite their importance in both natural and artificial ecosystems, little is known about soil microbial communities. This work aims at investigating the effects of fungal species richness on soil invasion by non resident microbes. Synthetic fungal communities with a species diversity ranging from 1 to 8 were assembled in laboratory microcosms and used in three factorial experiments to assess the effect of diversity on soil fungistasis, microbial invasion of soil amended with plant litter and of plant rhizosphere. The capability of different microbes to colonize environments characterized by different resident microbial communities was measured. The number of microbial species in the microcosms positively affected soil fungistasis that was also induced more rapidly in presence of synthetic communities with more species. Moreover, the increase of resident fungal diversity dramatically reduced the invasibility of both soil and plant rhizosphere. We found lower variability of soil fungistasis and invasibility in microcosms with higher species richness of microbial communities. Our study pointed out the existence of negative relationships between fungal diversity and soil invasibility by non resident microbes. Therefore, the loss of microbial species may adversely affect ecosystem functionality under specific environmental conditions.     

Diversity reduces invasibility in experimental plant communities: the role of plant species

Several studies have presented experimental evidence that diversity reduces invasibility in grassland communities. The interpretation of these results has been disputed recently and it was proposed that sampling effects were responsible for the observed decrease of invasibility with diversity. The experiments performed to date were not designed to adequately separate sampling from diversity effects. Using the establishment of native plant species in experimental plant communities as a model of invasibility, we show that the number of invaders decreased with increasing diversity. When the presence of particular species is included, their effects are dominant. Centaurea jacea showed a strong effect at low diversity, whereas Leucanthemum vulgare showed a very strong negative impact at each diversity level. The negative effect of the latter might be related to root‐feeding nematodes that showed far higher abundance in plots with Leucanthemum. However, diversity remained a significant factor in determining the number of invading species and the numbers of an abundant invader.     

Positive diversity–invasibility relationship in species-rich semi-natural grassland at the neighbourhood scale

Background and Aims

Attempts to answer the old question of whether high diversity causes high invasion resistance have resulted in an invasion paradox: while large-scale studies often find a positive relationship between diversity and invasibility, small-scale experimental studies often find a negative relationship. Many of the small-scale studies are conducted in artificial communities of even-aged plants. Species in natural communities, however, do not represent one simultaneous cohort and occur at various levels of spatial aggregation at different scales. This study used natural patterns of diversity to assess the relationship between diversity and invasibility within a uniformly managed, semi-natural community.

Methods

In species-rich grassland, one seed of each of ten species was added to each of 50 contiguous 16 cm² quadrats within seven plots (8 × 100 cm). The emergence of these species was recorded in seven control plots, and establishment success was measured in relation to the species diversity of the resident vegetation at two spatial scales, quadrat (64 cm²) within plots (800 cm²) and between plots within the site (approx. 400 m²) over 46 months.

Key Results

Invader success was positively related to resident species diversity and richness over a range of 28–37 species per plot. This relationship emerged 7 months after seed addition and remained over time despite continuous mortality of invaders.

Conclusions

Biotic resistance to plant invasion may play only a sub-ordinate role in species-rich, semi-natural grassland. As possible alternative explanations for the positive diversity–invasibility relationship are not clear, it is recommended that future studies elaborate fine-scale environmental heterogeneity in resource supplies or potential resource flows from resident species to seedlings by means of soil biological networks established by arbuscular mycorrhizal fungi.     

Species pool size and invasibility of island communities: a null model of sampling effects

The success of alien species on oceanic islands is considered to be one of the classic observed patterns in ecology. Explanations for this pattern are based on lower species richness on islands and the lower resistance of species‐poor communities to invaders, but this argument needs re‐examination. The important difference between islands and mainland is in the size of species pools, not in local species richness; invasibility of islands should therefore be addressed in terms of differences in species pools. Here I examine whether differences in species pools can affect invasibility in a lottery model with pools of identical native and exotic species. While in a neutral model with all species identical, invasibility does not depend on the species pool, a model with non‐zero variation in population growth rates predicts higher invasibility of communities of smaller pools. This is because of species sampling; drawing species from larger pools increases the probability that an assemblage will include fast growing species. Such assemblages are more likely to exclude random invaders. This constitutes a mechanism through which smaller species pools (such as those of isolated islands) can directly underlie differences in invasibility.     

Amphibian richness patterns in Atlantic Forest areas invaded by American bullfrogs

The relationship between invasion success and native biodiversity is central to biological invasion research. New theoretical and analytical approaches have revealed that spatial scale, land‐use factors and community assemblages are important predictors of the relationship between community diversity and invasibility and the negative effects of invasive species on community diversity. In this study we assess if the abundance of Lithobates catesbeianus, the American bullfrog, negatively affects the richness of native amphibian species in Atlantic Forest waterbodies in Brazil. Although this species has been invading Atlantic Forest areas since the 1930s, studies that estimate the invasion effects upon native species diversity are lacking. We developed a model to understand the impact of environmental, spatial and species composition gradients on the relationships between bullfrogs and native species richness. We found a weak positive relationship between bullfrog abundance and species richness in invaded areas. The path model revealed that this is an indirect relationship mediated by community composition gradients. Our results indicate that bullfrogs are more abundant in certain amphibian communities, which can be species‐rich. Local factors describing habitat heterogeneity were the main predictors of amphibian species richness and composition and bullfrog abundance. Our results reinforce the important role of habitats in determining both native species diversity and potential invasibility.     

Dominance not richness determines invasibility of tallgrass prairie

Many recent studies suggest that more diverse communities are more resistant to invasion. Community characteristics that most strongly influence invasion are uncertain, however, due to covariation of diversity with competition and crowding. We examined separately the effects of species richness and dominance on invasion by an exotic legume, Melilotus officinalis , in intact, native Kansas grassland. We manipulated dominance of C₄ grasses by reducing their abundance (i.e. ramet densities) by ∼25 and 50%. In addition, richness was reduced by removing species that were mainly rare and uncommon as might be expected with environmental changes such as drought and fragmentation. In both years of the study (2001–2002), invasibility, measured as peak establishment of Melilotus , was not affected by a 3-fold reduction in species richness, nor was there an interaction between loss of species and reduced dominance on invasion. In contrast, reductions in abundance of the dominants significantly reduced invasibility of the grassland plots in both years. Because the abundance of dominants was highly correlated with measures of competition (i.e. ratio of dominant biomass to total biomass) and crowding (total stem densities), this pattern was opposite to that expected if competition were indeed limiting invasion. Rather, invasion appeared to be facilitated by the dominant species, most likely because reduced dominance increased environmental stress. Our results suggest that dominance is the key community characteristic determining invasibility, because highly competitive and space-filling species can either enhance or reduce susceptibility to invasion depending on whether dominants create a more competitive environment or alleviate stressful conditions.     

Dominant species identity regulates invasibility of old-field plant communities

Dominant species are known to exert strong influence over community dynamics, although little work has addressed how they affect invasibility. In this study, we examined whether dominant species identity and abundance affected invasibility of old-field plant communities. To quantify invasibility, we added seeds of 19 plant species into plots dominated by one of four different herbaceous perennial species ( Andropogon virginicus , Bromus inermis , Centaurea maculosa , or Solidago canadensis ) . We found that, independent of species richness and abiotic variables, plots dominated by Andropogon were the least invasible, while Bromus and Centaurea plots had the highest invasibility. We examined several potential mechanisms by which these dominant species might influence invasibility, and found invasion to increase with decreasing litter biomass and increasing community species richness. The abundance of the dominant species was not a significant predictor of invasion. These results indicate that dominant species identity plays an important role in determining invasibility of plant communities, though exact mechanisms underlying these effects still need to be explored.     

Experimental demonstration that plant diversity reduces invasibility – evidence of a biological mechanism or a consequence of sampling effect?

Several recent studies have claimed to present experimental evidence from synthesised plant communities in which diversity was varied as a treatment that diversity reduces community invasibility by other plant species. This type of result contrasts from that of many observational studies which find diversity and invasibility to be positively correlated in nature, but some recent literature has claimed that these observational studies are confounded by extrinsic covarying factors while experimental studies are not. In this article I evaluate each of eight experiments from six recent publications in which the effect of varying plant diversity on the success of invasive species was investigated. In each case that invasibility was identified by the authors as being adversely affected by plant species richness, the result can be explained by factors that covaried with diversity in the experiment, most notably as a consequence of “sampling effect” (in which the most competitive species or species combination in the total species pool has a greater probability of occurring as species richness is increased), or through the incorrect use of statistical techniques. It is proposed that the apparent discrepancy between the results of many observational and experimental studies at least in grasslands is because: (1) in observational studies competitive dominant species are often associated with the most productive plots, and these dominants both reduce diversity through competitive exclusion of subordinates and competitively suppress invasive species; and (2) in recent experimental studies “sampling effect” results in the most competitive species (and therefore those most likely to suppress the invader) occurring with greater frequency as diversity is increased. Both observational and experimental studies therefore point to the role of competitive dominants in reducing invasibility, and in both situations species richness of the plant community need not be invoked as an explanation for the results.     

Resident plant diversity and introduced earthworms have contrasting effects on the success of invasive plants

Theoretical predictions and empirical studies suggest that resident species diversity is an important driver of community invasibility. Through trait-based processes, plants in communities with high resident species diversity occupy a wider range of ecological niches and are more productive than low diversity communities, potentially reducing the opportunities for invasion through niche preemption. In terrestrial plant communities, biotic ecosystem engineers such as earthworms can also affect invasibility by reducing leaf litter stocks and influencing soil conditions. In a greenhouse experiment, we simultaneously manipulated resident species diversity and earthworm presence to investigate independent and interactive effects of these two variables on the success of several invasive plants. Higher diversity of resident species was associated with lower biomass of invasives, with the effect mediated through resident species biomass. The presence of earthworms had a strong positive effect on the biomass of invasive species across all levels of resident species diversity and a weaker indirect negative effect via decreased soil moisture. Earthworms also weakened the positive correlation between resident species diversity and productivity. We did not observe any interactive effects of resident species biomass and earthworms on invasive species success. Partitioning the net biodiversity effect indicated that selection effects increased with resident species diversity whereas complementarity effects did not. Results suggest that managing for diverse forest communities may decrease the susceptibility of these communities to invasions. However, the presence of introduced earthworms in previously earthworm-free sites may undermine these efforts. Furthermore, future studies of plant community invasibility should account for the effects of introduced earthworms.     

Control of plant species diversity and community invasibility by species immigration: seed richness versus seed density

Immigration rates of species into communities are widely understood to influence community diversity, which in turn is widely expected to influence the susceptibility of ecosystems to species invasion. For a given community, however, immigration processes may impact diversity by means of two separable components: the number of species represented in seed inputs and the density of seed per species. The independent effects of these components on plant species diversity and consequent rates of invasion are poorly understood. We constructed experimental plant communities through repeated seed additions to independently measure the effects of seed richness and seed density on the trajectory of species diversity during the development of annual plant communities. Because we sowed species not found in the immediate study area, we were able to assess the invasibility of the resulting communities by recording the rate of establishment of species from adjacent vegetation. Early in community development when species only weakly interacted, seed richness had a strong effect on community diversity whereas seed density had little effect. After the plants became established, the effect of seed richness on measured diversity strongly depended on seed density, and disappeared at the highest level of seed density. The ability of surrounding vegetation to invade the experimental communities was decreased by seed density but not by seed richness, primarily because the individual effects of a few sown species could explain the observed invasion rates. These results suggest that seed density is just as important as seed richness in the control of species diversity, and perhaps a more important determinant of community invasibility than seed richness in dynamic plant assemblages.     

Reductions in grassland species evenness increase dicot seedling invasion and spittle bug infestation

Previous experiments that tested whether diverse plant communities have lower invasibility have all varied species richness. We experimentally varied evenness of four grassland species (three grasses and one forb) by planting a field experiment in Texas, and monitored the number of unplanted dicot and monocot species that invaded plots for two growing seasons. By varying evenness, we eliminated any sampling effect in our diversity treatment, because all plots contained the same plant species. Experimentally reducing evenness led to a greater number of dicot invaders, which emerged in plots throughout the growing season, but had less of an effect on monocot invaders, which emerged in flushes when experimental plants were semi‐dormant. Frequency of Solidago canadensis (altissima) stems with spittle bugs significantly increased with reductions in evenness during the first year, apparently because the greater number of Solidago stems in high evenness plots diluted the spittle‐bug effect. These results support the view that higher diversity plant communities are more resistant to dicot invaders and insect herbivores.