Native Plant Diversity Generates Microbial Legacies That Either Promote or Suppress Non-Natives, Depending on Drought History.

Diverse native plant communities resist non-native plants more than species-poor communities, in part through resource competition. The role of soil biota in diversity-invasibility relationships is poorly understood, although non-native plants interact with soil biota during invasions. We tested the responses of non-native plants to soil biota generated by different native plant diversities. We applied well-watered and drought treatments in both conditioning and response phases to explore the effects of 'historical' and 'contemporary' environmental stresses. When generated in well-watered soils, the microbial legacies from higher native diversity inhibited non-native growth in well-watered conditions. In contrast, when generated in drought-treated soils, the microbial legacies from higher native diversity facilitated non-native growth in well-watered conditions. Contemporary drought eliminated microbial legacy effects on non-native growth. We provide a new understanding of mechanisms behind diversity-invasibility relationships and demonstrate that temporal variation in environmental stress shapes relationships among native plant diversity, soil biota and non-native plants.

Comparing long-term patterns of spread of native and invasive plants in a successional forest.

A fundamental question in invasive plant ecology is whether invasive and native plants have different ecological roles. Differences in functional traits have been explored, but we lack a comparison of the factors affecting the spread of co-occurring natives and invasives. Some have proposed that to succeed, invasives would colonize a wider variety of sites, would disperse farther, or would be better at colonizing sites with more available light and soil nutrients than natives. We examined patterns of spread over 70 years in a regenerating forest in Connecticut, USA, where both native and invasive species acted as colonizers. We compared seven invasive and 19 native species in the characteristics of colonized plots, variation in these characteristics, and the importance of site variables for colonization. We found little support for the hypotheses that invasive plants succeed by dispersing farther than native plants or by having a broader range of site tolerances. Colonization by invasives was also not more dependent on light than colonization by natives. Like native understory species, invasive plants spread into closed-canopy forest and species-rich communities despite earlier predictions that these communities would resist invasion. The biggest differences were that soil nitrate and the initial land cover being open field increased the odds of colonization for most invasives but only for some natives. In large part, though, the spread of native and invasive plants was affected by similar factors.

Invasional meltdown mediated by plant-soil feedbacks may depend on community diversity.

It has been suggested that establishment of one alien invader might promote further invasions. Such a so-called invasional meltdown could be mediated by differences in soil-legacy effects between alien and native plants. Whether such legacy effects might depend on the diversity of the invaded community has not been explored to date. Here, we conducted a two-phase plant-soil feedback experiment. In a soil-conditioning phase, we grew five alien and five native species as invaders in 21 communities of one, two or four species. In the subsequent test phase, we grew five alien and five native species on the conditioned soils. We found that growth of these test species was negatively affected by soils conditioned by both a community and an invader, and particularly if the previous invader was a conspecific (i.e. negative plant-soil feedback). Alien test species suffered less from soil-legacy effects of previous allospecific alien invaders than from the legacy effects of previous native invaders. However, this effect decreased when the soil had been co-conditioned by a multispecies community. Our findings suggest that plant-soil feedback-mediated invasional meltdown may depend on community diversity and therefore provide some evidence that diverse communities could increase resistance against subsequent alien invasions.

Biotic resistance to invasion is ubiquitous across ecosystems of the United States.

The biotic resistance hypothesis predicts that diverse native communities are more resistant to invasion. However, past studies vary in their support for this hypothesis due to an apparent contradiction between experimental studies, which support biotic resistance, and observational studies, which find that native and non-native species richness are positively related at broad scales (small-scale studies are more variable). Here, we present a novel analysis of the biotic resistance hypothesis using 24 456 observations of plant richness spanning four community types and seven ecoregions of the United States. Non-native plant occurrence was negatively related to native plant richness across all community types and ecoregions, although the strength of biotic resistance varied across different ecological, anthropogenic and climatic contexts. Our results strongly support the biotic resistance hypothesis, thus reconciling differences between experimental and observational studies and providing evidence for the shared benefits between invasive species management and native biodiversity conservation.

Multiple drivers of contrasting diversity-invasibility relationships at fine spatial grains.

The diversity-invasibility hypothesis and ecological theory predict that high-diversity communities should be less easily invaded than species-poor communities, but empirical evidence does not consistently support this prediction. While fine-scale experiments tend to yield the predicted negative association between diversity and invasibility, broad-scale observational surveys generally report a positive correlation. This conflicting pattern between experiments and observational studies is referred to as the invasion paradox and is thought to arise because different processes control species composition at different spatial scales. Here, we test empirically the extent to which the strength and direction of published diversity-invasibility relationships depend on spatial scale and on the metrics used to measure invasibility. Using a meta-analytic framework, we explicitly separate the two components of spatial scale: grain and extent, by focusing on fine-grain studies that vary in extent. We find evidence of multiple drivers of the paradox. When we consider only fine-grain studies, we still observe conflicting patterns between experiments and observational studies. In contrast, when we examine studies that are conducted at both a fine grain and fine extent, there is broad overlap in effect sizes between experiments and observation, suggesting that comparing studies with similar extents resolves the paradox at local scales. However, we uncover systematic differences in the metrics used to measure invasibility between experiments, which use predominantly invader performance, and observational studies, which use mainly invader richness. When we consider studies with the same metric (i.e., invader performance), the contrasting associations between study types also disappear. It is not possible, at present, to fully disentangle the effect of spatial extent and metric on the paradox because both variables are systematically associated in different directions with study type. There is therefore an urgent need to conduct experiments and observational studies that incorporate the full range of variability in spatial extent and invasibility metric.

Effects of native diversity, soil nutrients, and natural enemies on exotic invasion in experimental plant communities.

Many factors can promote exotic plant success. Three of these factors-greater pressure from natural enemies on natives, increased soil nutrient supply, and low native species richness-may interact during invasions. To test for independent and interactive effects of these drivers, we planted herbaceous perennial communities at two levels of native richness (monocultures and five-species polycultures). We then factorially manipulated soil nutrient supply and access to these communities by aboveground foliar enemies (fungal pathogens and insect herbivores), and allowed natural colonization to proceed for four years. We predicted that nutrient addition would increase exotic success, while enemy exclusion and increasing native richness would reduce exotic success. Additionally, we expected that enemy exclusion would reduce the benefits of nutrient addition to exotic species most in species-poor communities, and that this effect would be weaker in species-rich communities. In total, we found no evidence that nutrient supply, enemy access, and native richness interacted to influence exotic success. Furthermore, native richness had no effect on exotic success. Instead, nutrient addition increased, and enemy exclusion decreased, exotic success independently. As predicted, enemy exclusion reduced exotic success, primarily by slowing the decline in abundance of planted native species. Together, these results demonstrate that multiple drivers of exotic success can act independently within a single system.

The native-exotic species richness relationship varies with spatial grain of measurement and environmental conditions.

Biological invasions can have dramatic impacts on communities and biodiversity, and are critical considerations in conservation and management decisions. We present a novel analysis to determine how exotic species success varies with community richness and scale of measurement. Using 5,022 plots representing natural vegetation of the Carolinas, we calculated native and exotic species richness of all vascular plants at five grain sizes. To avoid spatial pseudoreplication, we randomly selected unique subplots from each larger plot, re-selecting 100 times to develop an empirical distribution of the native-exotic richness relationship (NERR). Because observed NERRs vary with spatial scale, we developed separate scale-specific null-model distributions to compare to the empirical data. For each spatial scale, we compared the empirical distribution of 100 slopes to the null distribution containing 99 permutations of species origin per empirical slope. We also analyzed the dataset according to broad assignments corresponding to environmental conditions, using the formation type assigned to each community. The plots followed across most scales the general trend that exotic richness increases with native richness. At the smallest scale, however, the NERR was negative. The slope of the NERR is significantly higher than the null model at the largest observed scale and significantly lower than the null model at the smallest two observed scales. The NERR for most formations follows the general pattern with scale for the entire dataset. Warm temperate forests expressed essentially 0 slope at the largest spatial grain, decreasing to a negative relationship at 1 m2 and smaller. Temperate freshwater marshes and wet meadows and shrublands expressed a positive relationship at all spatial grains, demonstrating that unique environmental and biogeographic conditions differentially affect exotic species. Further, these results indicate that exotic species are unevenly distributed across natural communities and that community assembly processes vary with scale.

Native species dispersal reduces community invasibility by increasing species richness and biotic resistance.

Recent studies indicate that diversity-invasibility relationships can depend on spatial scale, but the contributing role of native species dispersal among local communities in mediating these relationships remains unaddressed. Metacommunity ecology highlights the effects of species dispersal rates on local diversity, thereby suggesting that native species dispersal may influence local biotic resistance to invasion by non-native species. However, the effects of native species dispersal rates on local native diversity and invasibility could depend on any intraspecific differences of the invader that may alter establishment success. Here, I experimentally tested for the influence of native dispersal-diversity relationships on the invasibility of native communities by a non-native species represented by core, midrange and peripheral regions of the introduced geographic range. In mesocosms, native plankton communities were connected by low or moderate rates of dispersal to yield dispersal rate-driven differences in native species richness prior to invasion by a non-native zooplankter, Daphnia lumholtzi. After invasion, establishment success and effects of the non-native species on native community structure and ecosystem properties were evaluated as a function of dispersal rate and invader source region relative to a control without native species. Native species richness was greater at the moderate dispersal rate than the low dispersal rate and yielded a dispersal rate-dependent diversity-invasibility relationship that was robust to invader source region. There was almost no establishment success of the non-native species at moderate dispersal and reduced success at low dispersal relative to the control. Invader population growth rates were negative only at the moderate dispersal rate. Effects of species dispersal on native community and ecosystem response were more influential than effects of invasion and impacts associated with invader source region. The results demonstrate that dispersal-diversity relationships can influence diversity-invasibility relationships at the local spatial scale. These dispersal-driven responses of invasion were unaffected by any ecological differences associated with invasion history-related intraspecific variation of the non-native species. This study emphasizes that dispersal rates of native species in metacommunities can differentially alter local biotic resistance to invasion. Thus, native species dispersal rates have largely been an underappreciated local diversity maintenance mechanism that can confer insurance against biological invasions.

Native-exotic richness relationships: a biogeographic approach using turnover in island plant populations.

Spatial variation in exotic species richness is often correlated with native species richness, for reasons that are poorly understood. To better understand the mechanisms underpinning native-exotic richness relationships, I quantified the colonization and extinction of 18 exotic and 16 native plant species on 39 small islands located off the coast of New Zealand for 8 consecutive yr. Results revealed a positive native-exotic richness relationship, which could be explained by similar demographic responses of native and exotic species to island area. However, native and exotic species showed subtle differences in their response to other island attributes. Turnover in native species declined with island isolation, whereas turnover in exotic species increased with the exposure of islands to ocean-borne disturbances. Overall results illustrate how long-term observations of species turnover can be used to better understand the mechanisms underpinning native-exotic richness relationships, and demonstrate that large, exposed islands can be especially susceptible to invasions by exotic species.