The invasion paradox: reconciling pattern and process in species invasions

The invasion paradox describes the co-occurrence of independent lines of support for both a negative and a positive relationship between native biodiversity and the invasions of exotic species. The paradox leaves the implications of native-exotic species richness relationships open to debate: Are rich native communities more or less susceptible to invasion by exotic species? We reviewed the considerable observational, experimental, and theoretical evidence describing the paradox and sought generalizations concerning where and why the paradox occurs, its implications for community ecology and assembly processes, and its relevance for restoration, management, and policy associated with species invasions. The crux of the paradox concerns positive associations between native and exotic species richness at broad spatial scales, and negative associations at fine scales, especially in experiments in which diversity was directly manipulated. We identified eight processes that can generate either negative or positive native-exotic richness relationships, but none can generate both. As all eight processes have been shown to be important in some systems, a simple general theory of the paradox, and thus of the relationship between diversity and invasibility, is probably unrealistic. Nonetheless, we outline several key issues that help resolve the paradox, discuss the difficult juxtaposition of experimental and observational data (which often ask subtly different questions), and identify important themes for additional study. We conclude that natively rich ecosystems are likely to be hotspots for exotic species, but that reduction of local species richness can further accelerate the invasion of these and other vulnerable habitats.     

Aquatic plant community invasibility and scale-dependent patterns in native and invasive species richness

Invasive species richness often is negatively correlated with native species richness at the small spatial scale of sampling plots, but positively correlated in larger areas. The pattern at small scales has been interpreted as evidence that native plants can competitively exclude invasive species. Large-scale patterns have been understood to result from environmental heterogeneity, among other causes. We investigated species richness patterns among submerged and floating-leaved aquatic plants (87 native species and eight invasives) in 103 temperate lakes in Connecticut (northeastern USA) and found neither a consistently negative relationship at small (3-m2) scales, nor a positive relationship at large scales. Native species richness at sampling locations was uncorrelated with invasive species richness in 37 of the 60 lakes where invasive plants occurred; richness was negatively correlated in 16 lakes and positively correlated in seven. No correlation between native and invasive species richness was found at larger spatial scales (whole lakes and counties). Increases in richness with area were uncorrelated with abiotic heterogeneity. Logistic regression showed that the probability of occurrence of five invasive species increased in sampling locations (3 m2, n = 2980 samples) where native plants occurred, indicating that native plant species richness provided no resistance against invasion. However, the probability of three invasive species' occurrence declined as native plant density increased, indicating that density, if not species richness, provided some resistance with these species. Density had no effect on occurrence of three other invasive species. Based on these results, native species may resist invasion at small spatial scales only in communities where density is high (i.e., in communities where competition among individuals contributes to community structure). Most hydrophyte communities, however, appear to be maintained in a nonequilibrial condition by stress and/or disturbance. Therefore, most aquatic plant communities in temperate lakes are likely to be vulnerable to invasion.     

Landscape connectivity strengthens local-regional richness relationships in successional plant communities

Local species diversity is maintained over ecological time by a balance between dispersal and species interactions. Local-regional species richness relationships are often used to investigate the relative importance of these two processes and the scales at which they operate. For communities undergoing succession, theory predicts a temporal progression in local-regional species richness relationships: from no relationship to positive linear to saturating. However, observational tests have been mixed, and experiments have been rare. Using a replicated large-scale experiment, we evaluate the impact of two dispersal-governing processes at the regional scale, connectivity and shape of the region (i.e., patches), on the progression of local-regional species richness relationships for plant communities undergoing succession. Regional connectivity accelerates the transition from no relationship to a positive linear relationship, while the shape of the region has no consistent effect nine years post-disturbance. Our results experimentally demonstrate the importance of dispersal in structuring local-regional species richness relationships over time and suggest that conservation corridors among regions can increase local diversity through regional enrichment of plant communities undergoing reassembly.     

The importance of quantifying propagule pressure to understand invasion: an examination of riparian forest invasibility

The widely held belief that riparian communities are highly invasible to exotic plants is based primarily on comparisons of the extent of invasion in riparian and upland communities. However, because differences in the extent of invasion may simply result from variation in propagule supply among recipient environments, true comparisons of invasibility require that both invasion success and propagule pressure are quantified. In this study, we quantified propagule pressure in order to compare the invasibility of riparian and upland forests and assess the accuracy of using a community's level of invasion as a surrogate for its invasibility. We found the extent of invasion to be a poor proxy for invasibility. The higher level of invasion in the studied riparian forests resulted from greater propagule availability rather than higher invasibility. Furthermore, failure to account for propagule pressure may confound our understanding of general invasion theories. Ecological theory suggests that species-rich communities should be less invasible. However, we found significant relationships between species diversity and invasion extent, but no diversity-invasibility relationship was detected for any species. Our results demonstrate that using a community's level of invasion as a surrogate for its invasibility can confound our understanding of invasibility and its determinants.     

Scale-Dependent Effects of Habitat Disturbance on Species Richness in Tropical Forests

Abstract: Despite growing concern, no consensus has emerged over the effects of habitat modification on species diversity in tropical forests. Even for comparatively well-studied taxa such as Lepidoptera, disturbance has been reported to increase and decrease diversity with approximately equal frequency. Species diversity within landscapes depends on the spatial scale at which communities are sampled, and the effects of disturbance in tropical forests have been studied at a wide range of spatial scales. Yet the question of how disturbance affects diversity at different spatial scales has not been addressed. We reanalyzed data from previous studies to examine the relationship between spatial scale and effects of disturbance on tropical-forest Lepidoptera. Disturbance had opposite effects on diversity at large and small scales: as scale decreased, the probability of a positive effect of disturbance on diversity increased. We also explicitly examined the relationship between spatial scale and the diversity of butterflies in selectively logged and unlogged forest in Maluku Province, Indonesia. Species richness increased with spatial scale in both logged and unlogged forest, but at a significantly faster rate in unlogged forest, whereas species evenness increased with scale in unlogged forest but did not increase with scale in logged forest. These data indicate that the effects of habitat modification on species diversity are heavily scale-dependent. As a result, recorded effects of disturbance were strongly influenced by the spatial scale at which species assemblages were sampled. Future studies need to account for this by explicitly examining the effects of disturbance at a number of different spatial scales. A further problem arises because the relationship between scale and diversity is likely to differ among taxa in relation to mobility. This may explain to some extent why the measured effects of disturbance have differed between relatively mobile and immobile taxa.     

Compositional and functional stability of arthropod communities in the face of ant invasions

There is a general consensus that the diversity of a biotic community can have an influence on its stability, but the strength, ubiquity, and relative importance of this effect is less clear. In the context of biological invasions, diversity has usually been studied in terms of its effect on a community's invasibility, but diversity may also influence stability by affecting the magnitude of compositional or functional changes experienced by a community upon invasion. We examined the impacts of invasive ants on arthropod communities at five natural area sites in the Hawaiian Islands, and assessed whether differences among sites in community diversity and density variables were related to measures of stability. Ant invasion was usually associated with significant changes in overall community composition, as measured by Bray-Curtis distances, particularly among endemic subsets of the communities. Changes in mean species richness were also strong at three of the five sites. Among sites, diversity was negatively related to stability as measured by resistance to overall compositional change, but this effect could not be separated from the strong negative effect of invasive ant density on compositional stability. When compositional stability was measured as proportional change in richness, the best predictor of stability among endemic community subsets was endemic richness, with richer communities losing proportionately more species than species-poor communities. This effect was highly significant even after controlling for differences in invasive ant density and suggested that communities that had already lost many endemic species were resistant to further species loss upon ant invasion, while more intact communities remained vulnerable to species loss. Communities underwent strong but idiosyncratic functional shifts in association with ant invasion, both in terms of trophic structure and total arthropod biomass. There were no apparent relationships, however, between functional stability and community diversity or density measures. Instead, invasive ant density was the best among-site predictor of the magnitude of functional change. Overall, diversity appeared to be a poor predictor of stability in the face of ant invasion in these communities, possibly because any actual diversity effects were overshadowed by community-specific factors and variation in the magnitude of the ant-mediated perturbation.     

The invasibility of marine algal assemblages: role of functional diversity and identity

The emergence of the biodiversity-ecosystem functioning debate in the last decade has renewed interest in understanding why some communities are more easily invaded than others and how the impact of invasion on recipient communities and ecosystems varies. To date most of the research on invasibility has focused on taxonomic diversity, i.e., species richness. However, functional diversity of the communities should be more relevant for the resistance of the community to invasions, as the extent of functional differences among the species in an assemblage is a major determinant of ecosystem processes. Although coastal marine habitats are among the most heavily invaded ecosystems, studies on community invasibility and vulnerability in these habitats are scarce. We carried out a manipulative field experiment in tide pools of the rocky intertidal to test the hypothesis that increasing functional richness reduces the susceptibility of macroalgal communities to invasion. We selected a priori four functional groups on the basis of previous knowledge of local species characteristics: encrusting, turf, subcanopy, and canopy species. Synthetic assemblages containing one, two, three, or four different functional groups of seaweeds were created, and invasion by native species was monitored over an eight-month period. Cover and resource availability in the assemblages with only one functional group showed different patterns in the use of space and light, confirming true functional differences among our groups. Experimental results showed that the identity of functional groups was more important than functional richness in determining the ability of macroalgal communities to resist invasion and that resistance to invasion was resource-mediated.     

Productivity alters the scale dependence of the diversity-invasibility relationship

At small scales, areas with high native diversity are often resistant to invasion, while at large scales, areas with more native species harbor more exotic species, suggesting that different processes control the relationship between native and exotic species diversity at different spatial scales. Although the small-scale negative relationship between native and exotic diversity has a satisfactory explanation, we lack a mechanistic explanation for the change in relationship to positive at large scales. We investigated the native-exotic diversity relationship at three scales (range: 1-4000 km2) in California serpentine, a system with a wide range in the productivity of sites from harsh to lush. Native and exotic diversity were positively correlated at all three scales; it is rarer to detect a positive relationship at the small scales within which interactions between individuals occur. However, although positively correlated on average, the small-scale relationship between native and exotic diversity was positive at low-productivity sites and negative at high-productivity sites. Thus, the change in the relationship between native and exotic diversity does not depend on spatial scale per se, but occurs whenever environmental conditions change to promote species coexistence rather than competitive exclusion. This occurred within a single spatial scale when the environment shifted from being locally unproductive to productive.     

The natives are restless, but not often and mostly when disturbed

The argument that the threat posed by introduced species is overblown is often buttressed by the observation that native species sometimes also become invasive. An examination of the literature on plant invasions in the United States shows that six times more nonnative species have been termed invasive than native species, and that a member of the naturalized nonnative pool is 40 times more likely than a native species to be perceived as invasive. In the great majority of instances in which a native plant species is seen as invasive, the invasion is associated with an anthropogenic disturbance, especially changed fire or hydrological regime, livestock grazing, and changes wrought by an introduced species. These results suggest that natives are significantly less likely than nonnatives to be problematic for local ecosystems.     

Thinning and burning result in low-level invasion by nonnative plants but neutral effects on natives.

Many historically fire-adapted forests are now highly susceptible to damage from insects, pathogens, and stand-replacing fires. As a result, managers are employing treatments to reduce fuel loadings and to restore the structure, species, and processes that characterized these forests prior to widespread fire suppression, logging, and grazing. However, the consequences of these activities for understory plant communities are not well understood. We examined the effects of thinning and prescribed fire on plant composition and diversity in Pinus ponderosa forests of eastern Washington (USA). Data on abundance and richness of native and nonnative plants were collected in 70 stands in the Colville, Okanogan, and Wenatchee National Forests. Stands represented one of four treatments: thinning, burning, thinning followed by burning, or control; treatments had been conducted 3-19 years before sampling. Multi-response permutation procedures revealed no significant effect of thinning or burning on understory plant composition. Similarly, there were no significant differences among treatments in cover or richness of native plants. In contrast, nonnative plants showed small, but highly significant, increases in cover and richness in response to both thinning and burning. In the combined treatment, cover of nonnative plants averaged 2% (5% of total plant cover) but did not exceed 7% (16% of total cover) at any site. Cover and richness of nonnative herbs showed small increases with intensity of disturbance and time since treatment. Nonnative plants were significantly less abundant in treated stands than on adjacent roadsides or skid trails, and cover within these potential source areas explained little of the variation in abundance within treated stands. Although thinning and burning may promote invasion of nonnative plants in these forests, our data suggest that their abundance is limited and relatively stable on most sites.     

Linking richness, community variability, and invasion resistance with patch size

The influence of community dynamics on the success or failure of an invasion is of considerable interest. What has not been explored is the influence of patch size on the outcomes of invasions for communities with the same species pool. Here we use an empirically validated spatial model of a marine epibenthic community to examine the effects of patch size on community variability, species richness, invasion, and the relationships between these variables. We found that the qualitative form of the relationship between community variability and species richness is determined by the size of the model patch. In small patches, variability decreases with species richness, but beyond a critical patch size, variability increases with increasing richness. This occurs because in large patches large, long-lived colonies attain sufficient size to minimize mortality and dominate the community, leading to decreased species richness and community variability. This mechanism cannot operate on smaller patches where the size of colonies is limited by the patch size and mortality is high irrespective of species identity. Further, invasion resistance is strongly correlated with community variability. Thus, the relationship between species richness and invasion resistance is also determined by patch size. These patterns are generated largely by an inverse relationship between colony size and mortality, and they depend on the spatial nature and patch size of the community. Our results suggest that a continuum of possible relationships can exist between species richness, community variability, invasion resistance, and area. These relationships are emergent behaviors generated by the individual properties of the particular component species of a community.     

Dominant species identity, not community evenness, regulates invasion in experimental grassland plant communities

While there has been extensive interest in understanding the relationship between diversity and invasibility of communities, most studies have only focused on one component of diversity: species richness. Although the number of species can affect community invasibility, other aspects of diversity, including species identity and community evenness, may be equally important. While several field studies have examined how invasibility varies with diversity by manipulating species identity or evenness, the results are often confounded by resource heterogeneity, site history, or disturbance. We designed a mesocosm experiment to examine explicitly the role of dominant species identity and evenness on the invasibility of grassland plant communities. We found that the identity of the dominant plant species, but not community evenness, significantly impacted invasibility. Using path analysis, we found that community composition (dominant species identity) reduced invasion by reducing early-season light availability and increasing late-season plant community biomass. Nitrogen availability was an important factor for the survival of invaders in the second year of the experiment. We also found significant direct effects of certain dominant species on invasion, although the mechanisms driving these effects remain unclear. The magnitude of dominant species effects on invasibility we observed are comparable to species richness effects observed in other studies, showing that species composition and dominant species can have strong effects on the invasibility of a community.     

Rapid assessment of postfire plant invasions in coniferous forests of the western United States.

Fire is a natural part of most forest ecosystems in the western United States, but its effects on nonnative plant invasion have only recently been studied. Also, forest managers are engaging in fuel reduction projects to lessen fire severity, often without considering potential negative ecological consequences such as nonnative plant species introductions. Increased availability of light, nutrients, and bare ground have all been associated with high-severity fires and fuel treatments and are known to aid in the establishment of nonnative plant species. We use vegetation and environmental data collected after wildfires at seven sites in coniferous forests in the western United States to study responses of nonnative plants to wildfire. We compared burned vs. unburned plots and plots treated with mechanical thinning and/or prescribed burning vs. untreated plots for nonnative plant species richness and cover and used correlation analyses to infer the effect of abiotic site conditions on invasibility. Wildfire was responsible for significant increases in nonnative species richness and cover, and a significant decrease in native cover. Mechanical thinning and prescribed fire fuel treatments were associated with significant changes in plant species composition at some sites. Treatment effects across sites were minimal and inconclusive due to significant site and site x treatment interaction effects caused by variation between sites including differences in treatment and fire severities and initial conditions (e.g., nonnative species sources). We used canonical correspondence analysis (CCA) to determine what combinations of environmental variables best explained patterns of nonnative plant species richness and cover. Variables related to fire severity, soil nutrients, and elevation explained most of the variation in species composition. Nonnative species were generally associated with sites with higher fire severity, elevation, percentage of bare ground, and lower soil nutrient levels and lower canopy cover. Early assessments of postfire stand conditions can guide rapid responses to nonnative plant invasions.     

Volatile chemicals from leaf litter are associated with invasiveness of a neotropical weed in Asia

Some invasive plant species appear to strongly suppress neighbors in their nonnative ranges but much less so in their native range. We found that in the field in its native range in Mexico, the presence of Ageratina adenophora, an aggressive Neotropical invader, was correlated with higher plant species richness than found in surrounding plant communities where this species was absent, suggesting facilitation. However, in two nonnative ranges, China and India, A. adenophora canopies were correlated with much lower species richness than the surrounding communities, suggesting inhibition. Volatile organic compound (VOC) signals may contribute to this striking biogeographical difference and the invasive success of A. adenophora. In controlled experiments volatiles from A. adenophora litter caused higher mortality of species native to India and China, but not of species native to Mexico. The effects of A. adenophora VOCs on seedling germination and growth did not differ between species from the native range and species from the nonnative ranges of the invader. Litter from A. adenophora plants from nonnative populations also produced VOCs that differed quantitatively in the concentrations of some chemicals than litter from native populations, but there were no chemicals unique to one region. Biogeographic differences in the concentrations of some volatile compounds between ranges suggest that A. adenophora may be experiencing selection on biochemical composition in its nonnative ranges.     

Invasion in a diversity hotspot: exotic cover and native richness in the Californian serpentine flora

Exotic species have been observed to be more prevalent in sites where the richness of native species is highest, possibly reflecting variation among sites in resources, propagule supply, heterogeneity, or disturbance. However, such a pattern leaves unclear whether natives at species-rich sites are subject to especially severe impacts from exotics as a result. We considered this question using path models in which relationships between exotic cover and native richness were evaluated in the presence of correlated environmental factors. At 109 sites on serpentine soils across California, USA, exotic cover was positively correlated with total native herbaceous richness and was negatively correlated with the richness of both serpentine-endemic and rare native herbs. However, in path models that accounted for the influences of soil chemistry, disturbance, overstory cover, and regional rainfall and elevation, we found no indication that exotic cover reduced any component of native herb richness. Rather, our results indicated similarities and differences in the conditions favoring exotic, native, endemic, and rare species. Our results suggest that, in spite of some localized impacts, exotic species are not exerting a detectable overall effect on the community richness of the unique native flora of Californian serpentine.     

Compositional stability and diversity of vascular plant communities following logging disturbance in Appalachian forests

Human-caused changes in disturbance regimes and introductions of nonnative species have the potential to result in widespread, directional changes in forest community structure. The degree that plant community composition persists or changes following disturbances depends on the balance between local extirpation and colonization by new species, including nonnatives. In this study, we examined species losses and gains, and entry of native vs. exotic species to determine how oak forests in the Appalachian Mountains might shift in species composition following a gradient of pulse disturbances (timber harvesting). We asked (1) how compositional stability of the plant community (resistance and resilience) was influenced by disturbance intensity, (2) whether community responses were driven by extirpation or colonization of species, and (3) how disturbance intensity influenced total and functional group diversity, including the nonnative proportion of the flora through time. We collected data at three spatial scales and three times, including just before, one year post-disturbance, and 10 years post-disturbance. Resistance was estimated using community distance measures between pre- and one year post-disturbance, and resilience using community distance between pre- and 10-year post-disturbance conditions. The number of colonizing and extirpated species between sampling times was analyzed for all species combined and for six functional groups. Resistance and resilience decreased with increasing timber-harvesting disturbance; compositional stability was lower in the most disturbed plots, which was driven by colonization, but not extirpation, of species. Colonization of species also led to increases in diversity after disturbance that was typically maintained after 10 years following disturbance. Most of the community-level responses were driven by post-disturbance colonization of native forbs and graminoids. The nonnative proportion of plant species tended to increase following disturbance, especially at large spatial scales in the most disturbed treatments, but tended to decrease through time following disturbance due to canopy development. The results of this study are consistent with the theory that resources released by disturbance have strong influences on species colonization and community composition. The effects of management activities tested in this study, which span a gradient of timber-harvesting disturbance, shift species composition largely via an increase in species colonization and diversity.     

Spatial scale and species identity influence the indigenous-alien diversity relationship in springtails

Although theory underlying the invasion paradox, or the change in the relationship between the richness of alien and indigenous species from negative to positive with increasing spatial scale, is well developed and much empirical work on the subject has been undertaken, most of the latter has concerned plants and to a lesser extent marine invertebrates. Here we therefore examine the extent to which the relationships between indigenous and alien species richness change from the local metacommunity to the interaction neighborhood scales, and the influences of abundance, species identity, and environmental favorability thereon, in springtails, a significant component of the soil fauna. Using a suite of modeling techniques, including generalized least squares and geographically weighted regressions to account for spatial autocorrelation or nonstationarity of the data, we show that the abundance and species richness of both indigenous and alien species at the metacommunity scale respond strongly to declining environmental favorability, represented here by altitude. Consequently, alien and indigenous diversity covary positively at this scale. By contrast, relationships are more complex at the interaction neighborhood scale, with the relationship among alien species richness and/or density and the density of indigenous species varying between habitats, being negative in some, but positive in others. Additional analyses demonstrated a strong influence of species identity, with negative relationships identified at the interaction neighborhood scale involving alien hypogastrurid springtails, a group known from elsewhere to have negative effects on indigenous species in areas where they have been introduced. By contrast, diversity relationships were positive with the other alien species. These results are consistent with both theory and previous empirical findings for other taxa, that interactions among indigenous and alien species change substantially with spatial scale and that environmental favorability may play a key role in explaining the larger scale patterns. However, they also suggest that the interactions may be affected by the identity of the species concerned, especially at the interaction neighborhood scale.     

Effects of Cattle Grazing on Diversity in Ephemeral Wetlands

Abstract:  Cattle are usually thought of as a threat to biodiversity. In regions threatened by exotic species invasion and lacking native wild grazers, however, cattle may produce the type of disturbance that helps maintain diverse communities. Across 72 vernal pools, I examined the effect of different grazing treatments (ungrazed, continuously grazed, wet-season grazed and dry-season grazed) on vernal-pool plant and aquatic faunal diversity in the Central Valley of California. After 3 years of treatment, ungrazed pools had 88% higher cover of exotic annual grasses and 47% lower relative cover of native species than pools grazed at historical levels (continuously grazed). Species richness of native plants declined by 25% and aquatic invertebrate richness was 28% lower in the ungrazed compared with the continuously grazed treatments. Release from grazing reduced pool inundation period by 50 to 80%, making it difficult for some vernal-pool endemic species to complete their life cycle. My results show that one should not assume livestock and ranching operations are necessarily damaging to native communities. In my central California study site, grazing helped maintain native plant and aquatic diversity in vernal pools.     

Latitudinal patterns of diversity and abundance in North Atlantic intertidal boulder-fields

In order to study taxon richness, biodiversity and abundance patterns in the North Atlantic from temperate latitudes through Arctic to high Arctic latitudes, we recorded the faunas (at ELWS level) colonizing 20 cobbles from three sites at each of seven boulder-field localities (south-west England, 50°N; Wales, 51°N; west Scotland, 56°N; Iceland, 64°N; Tromsø, 70°N; Svalbard, 77°N, 79°N). Inverse correlations were found between latitude and all measures of richness (species, orders, and phyla numbers) and biodiversity (S–W, P). However, these correlations were driven mostly by the consistently impoverished Arctic sites; an even cline of decreasing diversity from south to north did not exist. Multidimensional scaling revealed two communities, temperate–subarctic species-rich and high arctic species-poor. Evenness as measured by Pielou’s index was similar across all latitudes. Abundance data exhibited a similar trend to the species richness and diversity data with a significant negative correlation with latitude, but when Arctic data were excluded the correlation vanished.     

Disturbance influences oyster community richness and evenness, but not diversity

Foundation species in space-limited systems can increase diversity by creating habitat, but they may also reduce diversity by excluding primary space competitors. These contrasting forces of increasing associate diversity and suppressing competitor diversity have rarely been examined experimentally with respect to disturbance. In a benthic marine community in central California, where native oysters are a foundation species, we tested how disturbance influenced overall species richness, evenness, and diversity. Surprisingly, overall diversity did not peak across a disturbance gradient because, as disturbance decreased, decreases in overall species evenness opposed increases in overall species richness. Decreasing disturbance intensity (high oyster abundance) led to increasing species richness of sessile and mobile species combined. This increase was due to the facilitation of secondary sessile and mobile species in the presence of oysters. In contrast, decreasing disturbance intensity and high oyster abundance decreased the evenness of sessile and mobile species. Three factors likely contributed to this decreased evenness: oysters reduced abundances of primary sessile species due to space competition; oysters supported more rare mobile species; and oysters disproportionately increased the relative abundance of a few common mobile species. Our results highlight the need for further studies on how disturbance can differentially affect the evenness and richness of different functional groups, and ultimately how these differences affect the relationship between overall diversity and ecosystem function.