Richness, structure and functioning in metazoan parasite communities

Ecosystem functioning, characterized by components such as productivity and stability, has been extensively linked with diversity in recent years, mainly in plant ecology. The aim of our study was thus to quantify general relationships between diversity, community structure and ecosystem functions in metazoan parasite communities. We used data on parasite communities from 15 species of marine fish hosts from coastal Chile. The volumetric abundance (volume of all parasite species per individual host, in mm³) was used as a surrogate for productivity. Species diversity was measured using both species richness and evenness, while community structure was estimated using the co‐occurrence indices V‐ratio, C‐score and a new C‐score_s index standardized for the number of host replicates. After correcting for fish size, 47% of host species show no relationship, 13% show a hump shaped curve and 40% show positive monotonic relationships between productivity and parasite richness across all host individuals in a sample. We obtained a logarithmically decreasing relationship between evenness and productivity for all fish species, and propose a ‘dominance‐resistance’ hypothesis based on immunity to explain this pattern. The stability of the parasite community, measured as the coefficient of variation in productivity among individual hosts, was strongly and positively related to mean species richness across the 15 host species. The C‐score_s index, based on the number of checkerboard units in the host‐parasite presence/absence matrix, increases linearly with mean productivity across the 15 host species, suggesting that parasite communities tend to be more structured when they are more productive. This is the likely reason why linear relationships between richness and productivity were not observed consistently in all fish species. Parasite communities provide some clear patterns for the diversity–ecosystem functioning debate in ecology, although other factors, such as the history of community assembly, may also influence these patterns.     

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.     

Ecosystem productivity is associated with bacterial phylogenetic distance in surface marine waters

Understanding the link between community diversity and ecosystem function is a fundamental aspect of ecology. Systematic losses in biodiversity are widely acknowledged but the impact this may exert on ecosystem functioning remains ambiguous. There is growing evidence of a positive relationship between species richness and ecosystem productivity for terrestrial macro‐organisms, but similar links for marine micro‐organisms, which help drive global climate, are unclear. Community manipulation experiments show both positive and negative relationships for microbes. These previous studies rely, however, on artificial communities and any links between the full diversity of active bacterial communities in the environment, their phylogenetic relatedness and ecosystem function remain hitherto unexplored. Here, we test the hypothesis that productivity is associated with diversity in the metabolically active fraction of microbial communities. We show in natural assemblages of active bacteria that communities containing more distantly related members were associated with higher bacterial production. The positive phylogenetic diversity–productivity relationship was independent of community diversity calculated as the Shannon index. From our long‐term (7‐year) survey of surface marine bacterial communities, we also found that similarly, productive communities had greater phylogenetic similarity to each other, further suggesting that the traits of active bacteria are an important predictor of ecosystem productivity. Our findings demonstrate that the evolutionary history of the active fraction of a microbial community is critical for understanding their role in ecosystem functioning.     

Newly created ponds complement natural waterbodies for restoration of macroinvertebrate assemblages

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Wild bee community change over a 26‐year chronosequence of restored tallgrass prairie

Restoration efforts often focus on plants, but additionally require the establishment and long‐term persistence of diverse groups of nontarget organisms, such as bees, for important ecosystem functions and meeting restoration goals. We investigated long‐term patterns in the response of bees to habitat restoration by sampling bee communities along a 26‐year chronosequence of restored tallgrass prairie in north‐central Illinois, U.S.A. Specifically, we examined how bee communities changed over time since restoration in terms of (1) abundance and richness, (2) community composition, and (3) the two components of beta diversity, one‐to‐one species replacement, and changes in species richness. Bee abundance and raw richness increased with restoration age from the low level of the pre‐restoration (agricultural) sites to the target level of the remnant prairie within the first 2–3 years after restoration, and these high levels were maintained throughout the entire restoration chronosequence. Bee community composition of the youngest restored sites differed from that of prairie remnants, but 5–7 years post‐restoration the community composition of restored prairie converged with that of remnants. Landscape context, particularly nearby wooded land, was found to affect abundance, rarefied richness, and community composition. Partitioning overall beta diversity between sites into species replacement and richness effects revealed that the main driver of community change over time was the gradual accumulation of species, rather than one‐to‐one species replacement. At the spatial and temporal scales we studied, we conclude that prairie restoration efforts targeting plants also successfully restore bee communities.     

Successional dynamics of soil fungal diversity along a restoration chronosequence post‐coal mining

Soil disruption from open‐cut mining practices can adversely impact microbial communities and the ecosystem services that they mediate. Despite this, assessment of impacts of soil disruption, and the subsequent recovery of microbial communities is rarely studied. Monitoring of ecological restoration success on mine sites has traditionally focused on vegetation; however, most plants rely, at least in part, on associations with soil fungi for enhanced nutrient and water acquisition. Here, we used high‐throughput phylogenetic marker gene sequencing to characterize the diversity of soil fungal communities along a restoration chronosequence ranging from 3 to 23 years at a rehabilitated mine site. We used nonmined analogue sites as a baseline for comparative purposes and examined the associations of soil fungal communities with soil physicochemical and aboveground vegetation variables. Fungal richness on rehabilitated sites was significantly larger than on nonmined sites, suggesting that mixing of topsoil during stockpiling resulted in a composite microbial community. Fungal community composition was significantly influenced by edaphic variables and the length of rehabilitation, with mined sites becoming more similar to nonmined sites over time. Fungal populations associated with ectomycorrhizae were relatively more abundant than those associated with arbuscular mycorrhizae and declined in response to disturbance, but recovered over time on the woody dominated sites indicating a strong coupling of these fungi with aboveground vegetation. Our data indicate that soil fungal diversity is a useful bioindicator of soil restoration in mined sites and may complement more traditional vegetation‐based surveys.     

Evolution of species interactions determines microbial community productivity in new environments

Diversity generally increases ecosystem productivity over short timescales. Over longer timescales, both ecological and evolutionary responses to new environments could alter productivity and diversity–productivity relationships. In turn, diversity might affect how component species adapt to new conditions. We tested these ideas by culturing artificial microbial communities containing between 1 and 12 species in three different environments for ∼60 generations. The relationship between community yields and diversity became steeper over time in one environment. This occurred despite a general tendency for the separate yields of isolates of constituent species to be lower at the end if they had evolved in a more diverse community. Statistical comparisons of community and species yields showed that species interactions had evolved to be less negative over time, especially in more diverse communities. Diversity and evolution therefore interacted to enhance community productivity in a new environment.     

Advanced mine restoration protocols facilitate early recovery of soil microbial biomass,activity and functional diversity

Many ecosystem restoration programmes can take over 15 years to achieve ecosystem functioning comparable to that of an unmodified ecosystem, therefore a reliable shorter-term method of assessing and monitoring ecosystem recovery is needed to ensure that recovery is following an appropriate trajectory. Soil microbes respond to environmental change relatively quickly, and shifts in microbial communities can reflect the current status of their environment. As well as potentially acting as ‘indicator communities’, microbes play an integral role in restoring ecosystem functions. On an active opencast mine on New Zealand's West Coast, three main restoration methods are used, differing in cost and restoration effort. They range from most expensive (1) vegetation direct transfer (VDT), to (2) biosolids-amended stockpiles that are spread and replanted, and (3) untreated stockpiles that are spread and replanted. We assessed the impacts of these methods on soil microbial communities by measuring microbial biomass, dehydrogenase activity, community level physiological profile (CLPP) and functional diversity as measured by carbon substrate utilisation, where restored sites were 5 years old or less. These measures were compared to an unmodified reference ecosystem in the same location. Microbial activity and biomass were highest in pristine habitats, followed by VDT and biosolids-amended soils, then untreated stockpile soil. When compared to all other treatments untreated stockpiled soils had significantly different CLPPs and significantly reduced microbial biomass and activity; microbial biomass was an order of magnitude lower than in pristine soils. Functional diversity and richness did not differ between pristine, VDT and biosolids-amended soils, but were higher than in untreated stockpiled soils. CLPPs did not differ between pristine habitat soil and VDT soil but biosolids-amended and untreated stockpiled soils were significantly different to pristine soil. This study has shown that soil microbial communities are a valuable tool to assess restoration progress, and that ecosystem restoration can begin in a relatively short time following investment in appropriate restoration strategy, ultimately benefiting recolonisation by plants and animals.     

Species and genotype diversity drive community and ecosystem properties in experimental microcosms

Species diversity is important to ecosystems because of the increased probability of including species that are strong interactors and/or because multiple-species communities are more efficient at using resources due to synergisms and resource partitioning. Genetic diversity also contributes to ecosystem function through effects on primary productivity, community structure and resilience, and modulating energy and nutrient fluxes. Lacking are studies investigating the relationship between ecosystem function and diversity where hierarchical levels of biological diversity are systematically varied during experimentation. In this experiment, we manipulated both species and genotypic diversity of two Daphnia species in microcosms initially seeded with Chlamydomonas and measured community- and ecosystem-level properties to determine which level of diversity was most important for explaining variation in the property. Our results show that species diversity alters bacterial community composition while high genotypic diversity reduces bacterial richness and primary productivity. In addition, the highest levels of genotypic and species richness appear to increase community and ecosystem stability. These findings reveal that species and genotypic diversity are significant drivers of community and ecosystem properties and stability.     

Genotypic diversity of a dominant C4 grass across a long-term fire frequency gradient

Aims and Methods Diversity-disturbance research has focused on community diversity, but disturbance frequency could impact diversity within species as well, with important consequences for community diversity and ecosystem function. We examined patterns of genetic diversity of a dominant grass species, Andropogon gerardii, in native North American tallgrass prairie sites located in eastern Kansas that have been subjected to a gradient of fire frequency treatments (burned every 1, 2, 4 or 20 years) since the 1970s. In addition, we were able to assess the relationships between genetic diversity of A. gerardii, species diversity and productivity across this range of fire frequencies.Important findings We found no significant relationships between genetic diversity of A. gerardii at the local scale (1 m 2 plot level) and disturbance frequency (burned 2 to 32 times over a 38-year period). However, at the site level (i.e. across all plots sampled within a site, ~100 m 2) there were differences in genotype richness and composition, as well as genomic dissimilarity among individuals of A. gerardii. Genotype richness was greatest for the site burned at an intermediate (4-year) frequency and lowest for the infrequently (20-year) burned site. In addition, genotypes found in the frequently burned sites were more similar from each other than expected by random chance than those found in the infrequently burned sites. Genotype composition of A. gerardii was not significantly different between the frequently burned sites (annual vs. 2 year) but did differ between frequently burned and infrequently burned sites (1 and 2 year vs. 4 and 20 year, etc.). Together, these results suggest site-level ecological sorting of genotypes in intact prairie across a broad gradient of disturbance frequencies, likely driven by alterations in environmental conditions. Frequent fire promotes the abundance of dominant grass species, reduces plant community diversity and impacts ecosystem processes such as productivity. Our study suggests that genetic diversity within dominant grass species also may be affected by disturbance frequency, which could have important implications for how species are able to respond to disturbance.     

Land-Use History and Contemporary Management Inform an Ecological Reference Model for Longleaf Pine Woodland Understory Plant Communities

Ecological restoration is frequently guided by reference conditions describing a successfully restored ecosystem; however, the causes and magnitude of ecosystem degradation vary, making simple knowledge of reference conditions insufficient for prioritizing and guiding restoration. Ecological reference models provide further guidance by quantifying reference conditions, as well as conditions at degraded states that deviate from reference conditions. Many reference models remain qualitative, however, limiting their utility. We quantified and evaluated a reference model for southeastern U.S. longleaf pine woodland understory plant communities. We used regression trees to classify 232 longleaf pine woodland sites at three locations along the Atlantic coastal plain based on relationships between understory plant community composition, soils (which broadly structure these communities), and factors associated with understory degradation, including fire frequency, agricultural history, and tree basal area. To understand the spatial generality of this model, we classified all sites together and for each of three study locations separately. Both the regional and location-specific models produced quantifiable degradation gradients–i.e., progressive deviation from conditions at 38 reference sites, based on understory species composition, diversity and total cover, litter depth, and other attributes. Regionally, fire suppression was the most important degrading factor, followed by agricultural history, but at individual locations, agricultural history or tree basal area was most important. At one location, the influence of a degrading factor depended on soil attributes. We suggest that our regional model can help prioritize longleaf pine woodland restoration across our study region; however, due to substantial landscape-to-landscape variation, local management decisions should take into account additional factors (e.g., soil attributes). Our study demonstrates the utility of quantifying degraded states and provides a series of hypotheses for future experimental restoration work. More broadly, our work provides a framework for developing and evaluating reference models that incorporate multiple, interactive anthropogenic drivers of ecosystem degradation.     

Is community persistence related to diversity? A test with prairie species in a long-term experiment

Community persistence, or the ability of a community to maintain species composition and diversity through time, is a component of stability that is important to restoration. We ran a biodiversity–ecosystem functioning experiment for three years, and then stopped weeding it for 5–6 years, which allowed us to test whether increased plant species diversity and dissimilarity in height would lead to increased community persistence in the face of high invasion pressure by non-native species. Our approach was unique in that the experiment varied richness (one or four species) and evenness (three levels plus monocultures of the dominant species) using two separate dissimilarity types (having all tall species or having tall and short species combined) in six spatiotemporal blocks. Persistence was quantified as to how well positive productivity–diversity relationships, proportion of planted native species, and species richness remained unchanged over time. Thus, high persistence values indicate low levels of invasion and local extinction. We found that the positive relationship between diversity measures and productivity persisted after cessation of weeding. The proportion of planted species was 32% higher in mixture than in monoculture plots, indicating that monocultures were more heavily invaded by non-native species. Reduced evenness did not affect persistence measures in plots with dissimilar heights, but measures declined linearly with decreased evenness in plots with all tall species. Our results suggest that (1) persistence–diversity relationships are likely to vary with the traits of species becoming rare and going extinct, and (2) it is important to restore higher species diversity in restoration projects to favor the long-term persistence of planted species.     

Variable Effects of Dispersal on Productivity of Bacterial Communities Due to Changes in Functional Trait Composition

Previous studies have shown variable relationships between dispersal rate and ecosystem functioning, but the reasons for and mechanisms behind variable dispersal rate – functioning patterns are currently unknown. In this study we used six bacterial lake water communities in a laboratory experiment in order to investigate how dispersal among communities influences community productivity by evaluating three different mechanisms: 1) changes in taxonomic diversity, 2) changes in phylogenetic diversity or 3) changes in the composition of functional traits. The experiment was conducted in two phases; (A) a dialysis bag experiment where the dispersal rate among six communities was manipulated and the subsequent change in bacterial diversity and growth rate was recorded, and (B) a regrowth experiment where we manipulated available resources to study how well a taxon grows on certain organic carbon resources, i.e. their functional traits. From experiment (B) we could thus estimate changes in functional traits in communities in experiment (A). Bacterial production was affected by dispersal, but not consistently among lakes. Neither change in taxonomic or phylogenetic diversity with dispersal could explain the observed dispersal – productivity relationships. Instead, changes in trait composition with dispersal, especially the communities’ ability to use p-coumaric acid, an aromatic compound, could explain the observed dispersal – productivity relationships. Changes in this trait caused by dispersal seemed especially important for bacterial productivity in waters with a high aromaticity of the organic matter pool. We conclude that the effect of dispersal on bacterial communities can affect ecosystem functioning in different ways, through changes in functional key-traits which are important for the local environment.     

Environmental Controls on Fungal Community Composition and Abundance Over 3 Years in Native and Degraded Shrublands

Soil fungal communities have high local diversity and turnover, but the relative contribution of environmental and regional drivers to those patterns remains poorly understood. Local factors that contribute to fungal diversity include soil properties and the plant community, but there is also evidence for regional dispersal limitation in some fungal communities. We used different plant communities with different soil conditions and experimental manipulations of both vegetation and dispersal to distinguish among these factors. Specifically, we compared native shrublands with former native shrublands that had been disturbed or converted to pasture, resulting in soils progressively more enriched in carbon and nutrients. We tested the role of vegetation via active removal, and we manipulated dispersal by adding living soil inoculum from undisturbed native sites. Soil fungi were tracked for 3 years, with samples taken at ten time points from June 2006 to June 2009. We found that soil fungal abundance, richness, and community composition responded primarily to soil properties, which in this case were a legacy of plant community degradation. In contrast, dispersal had no effect on soil fungi. Temporal variation in soil fungi was partly related to drought status, yet it was much broader in native sites compared to pastures, suggesting some buffering due to the increased soil resources in the pasture sites. The persistence of soil fungal communities over 3 years in this study suggests that soil properties can act as a strong local environmental filter. Largely persistent soil fungal communities also indicate the potential for strong biotic resistance and soil legacies, which presents a challenge for both the prediction of how fungi respond to environmental change and our ability to manipulate fungi in efforts such as ecosystem restoration.     

Vegetation recovery and plant facilitation in a human-disturbed lava field in a megacity: searching tools for ecosystem restoration

Unplanned urban development threatens natural ecosystems. Assessing ecosystem recovery after anthropogenic disturbances and identifying plant species that may facilitate vegetation regeneration are critical for the conservation of biodiversity and ecosystem services in urban areas. At the periphery of Mexico City, illegal human settlements produced different levels of disturbance on natural plant communities developed on a lava field near the Ajusco mountain range. We assessed natural regeneration of plant communities 20 years after the abandonment of the settlements, in sites that received low (manual harvesting of non-timber forest products), medium (removal of aboveground vegetation), and high (removal of substrate and whole vegetation) disturbance levels. We also tested the potential facilitative role played by dominant tree and shrub species. Plant diversity and vegetation biomass decreased as disturbance level increased. Sites with high disturbance level showed poor regeneration and the lowest species similarity compared to the least disturbed sites. Six dominant species (i.e., those with the highest abundance, frequency, and/or basal area) were common to all sites. Among them, three species (the tree Buddleja cordata, and two shrubs, Ageratina glabrata and Sedum oxypetalum) were identified as potential facilitators of community regeneration, because plant density and species richness were significantly higher under their canopies than at open sites. We propose that analyzing community structural traits of the successional vegetation (such as species diversity and biomass) and identifying potential facilitator species are useful steps in assessing the recovery ability of plant communities to anthropogenic disturbances, and in designing restoration strategies.     

Drivers of productivity and its temporal stability in a tropical tree diversity experiment

There is increasing evidence that mixed‐species forests can provide multiple ecosystem services at a higher level than their monospecific counterparts. However, most studies concerning tree diversity and ecosystem functioning relationships use data from forest inventories (under noncontrolled conditions) or from very young plantation experiments. Here, we investigated temporal dynamics of diversity–productivity relationships and diversity–stability relationships in the oldest tropical tree diversity experiment. Sardinilla was established in Panama in 2001, with 22 plots that form a gradient in native tree species richness of one‐, two‐, three‐ and five‐species communities. Using annual data describing tree diameters and heights, we calculated basal area increment as the proxy of tree productivity. We combined tree neighbourhood‐ and community‐level analyses and tested the effects of both species diversity and structural diversity on productivity and its temporal stability. General patterns were consistent across both scales indicating that tree–tree interactions in neighbourhoods drive observed diversity effects. From 2006 to 2016, mean overyielding (higher productivity in mixtures than in monocultures) was 25%–30% in two‐ and three‐species mixtures and 50% in five‐species stands. Tree neighbourhood diversity enhanced community productivity but the effect of species diversity was stronger and increased over time, whereas the effect of structural diversity declined. Temporal stability of community productivity increased with species diversity via two principle mechanisms: asynchronous responses of species to environmental variability and overyielding. Overyielding in mixtures was highest during a strong El Niño‐related drought. Overall, positive diversity–productivity and diversity–stability relationships predominated, with the highest productivity and stability at the highest levels of diversity. These results provide new insights into mixing effects in diverse, tropical plantations and highlight the importance of analyses of temporal dynamics for our understanding of the complex relationships between diversity, productivity and stability. Under climate change, mixed‐species forests may provide both high levels and high stability of production.     

Arid grassland bee communities: associated environmental variables and responses to restoration

In recent years restoration project efforts in arid grasslands of the Pacific Northwest have increased; however, little is known about the bee communities in these areas or how restoration affects them. Native bees provide an essential ecosystem service through pollination of crops and native plants and understanding their response to restoration is a high priority. To address this issue, we conducted a three‐year study in an arid bunchgrass prairie with three objectives: (1) describe the bee community of this unique grassland type and its temporal variability; (2) investigate environmental variables influencing the community; and (3) examine effects of restoration on the community. We identified 62 bee species and found strong seasonal and inter‐annual variation in bee abundance, richness, diversity, and species composition. Unexpectedly, these temporal trends did not correspond with patterns in floral resources; however, several variables were associated with variation in bee abundance, richness, and diversity among sites. Sites with high levels of litter cover had more bees, while sites with taller vegetation or more blooming flowers had greater species richness but lower diversity. We found no detectable effect of restoration on bee abundance, richness, diversity, or composition. Species composition at native sites differed from those in actively and passively restored sites, which did not differ from each other. Restored sites also had fewer flowers and differing floral composition relative to native sites. These results suggest that if grassland restoration is to benefit bees, efforts should focus on both expanding floral resources and enhancing variables that influence nesting habitat.     

Plant trait responses to the environment and effects on ecosystem properties

A combined analysis of plant trait responses to the environment, and their effects on ecosystem properties has recently been proposed. In this study, we related the trait composition of plant communities to soil nutrients and disturbance as environmental drivers and to productivity, decomposition and soil carbon as ecosystem properties. We surveyed two sites, one comprising intensively grazed and fertilized grasslands, the other consisting of semi-natural grassland and open heathland. Species abundance and trait values of 49 species were recorded in 69 plots, as well as parameters describing soil resources, land-use disturbances, and ecosystem properties. Our main goal was to test whether the average or the diversity of the trait values of the vegetation had stronger effects on ecosystem properties (mass ratio vs. diversity hypothesis). Structural equation modeling was used to perform a simultaneous analysis of trait responses and effects. Specific leaf area and leaf nutrient contents were always negatively correlated with stem dry matter content and canopy height, indicating greater investments in supportive and nutrient-conserving tissue as plants increased in size. In the agricultural site, disturbance was the single most important factor decreasing plant height, while leaf traits such as specific leaf area and leaf nutrient contents increased with soil resources in heathlands. Productivity was directly or indirectly driven by leaf traits, and investments in structural tissue increased standing biomass and soil carbon. Different environmental drivers in the two sites produced opposing leaf trait effects on litter decomposition. Ecosystem properties were explained by the community mean trait value as predicted by the mass ratio hypothesis. Evidence for effects of functional diversity on productivity and other ecosystem properties was not detected, suggesting that diversity–productivity relationships depend on the length of the investigated environmental gradients. We conclude that changes in community composition and dominance hierarchies deserve the most attention when ecosystem properties must be maintained.     

Community diversity outweighs effect of warming on plant colonization

Abiotic environmental change, local species extinctions and colonization of new species often co‐occur. Whether species colonization is driven by changes in abiotic conditions or reduced biotic resistance will affect community functional composition and ecosystem management. We use a grassland experiment to disentangle effects of climate warming and community diversity on plant species colonization. Community diversity had dramatic impacts on the biomass, richness and traits of plant colonists. Three times as many species colonized the monocultures than the high diversity 17 species communities (~30 vs. 10 species), and colonists collectively produced 10 times as much biomass in the monocultures than the high diversity communities (~30 vs. 3 g/m²). Colonists with resource‐acquisitive strategies (high specific leaf area, light seeds, short heights) accrued more biomass in low diversity communities, whereas species with conservative strategies accrued most biomass in high diversity communities. Communities with higher biomass of resident C4 grasses were more resistant to colonization by legume, nonlegume forb and C3 grass colonists, but not by C4 grass colonists. Compared with effects of diversity, 6 years of 3°C‐above‐ambient temperatures had little impact on plant colonization. Warmed subplots had ~3 fewer colonist species than ambient subplots and selected for heavier seeded colonists. They also showed diversity‐dependent changes in biomass of C3 grass colonists, which decreased under low diversity and increased under high diversity. Our findings suggest that species colonization is more strongly affected by biotic resistance from residents than 3°C of climate warming. If these results were extended to invasive species management, preserving community diversity should help limit plant invasion, even under climate warming.     

Across species‐pool aggregation alters grassland productivity and diversity

Plant performance is determined by the balance of intra‐ and interspecific neighbors within an individual's zone of influence. If individuals interact over smaller scales than the scales at which communities are measured, then altering neighborhood interactions may fundamentally affect community responses. These interactions can be altered by changing the number (species richness), abundances (species evenness), and positions (species pattern) of the resident plant species, and we aimed to test whether aggregating species at planting would alter effects of species richness and evenness on biomass production at a common scale of observation in grasslands. We varied plant species richness (2, 4, or 8 species and monocultures), evenness (0.64, 0.8, or 1.0), and pattern (planted randomly or aggregated in groups of four individuals) within 1 × 1 m plots established with transplants from a pool of 16 tallgrass prairie species and assessed plot‐scale biomass production and diversity over the first three growing seasons. As expected, more species‐rich plots produced more biomass by the end of the third growing season, an effect associated with a shift from selection to complementarity effects over time. Aggregating conspecifics at a 0.25‐m scale marginally reduced biomass production across all treatments and increased diversity in the most even plots, but did not alter biodiversity effects or richness–productivity relationships. Results support the hypothesis that fine‐scale species aggregation affects diversity by promoting species coexistence in this system. However, results indicate that inherent changes in species neighborhood relationships along grassland diversity gradients may only minimally affect community (meter) – scale responses among similarly designed biodiversity–ecosystem function studies. Given that species varied in their responses to local aggregation, it may be possible to use such species‐specific results to spatially design larger‐scale grassland communities to achieve desired diversity and productivity responses.