The outcomes of ecological restoration are notoriously unpredictable, but we have no general predictive understanding of this contingency. Management decisions can have strong effects on restoration outcomes, but in other cases may be overwhelmed by site characteristics (e.g. soil conditions), landscape context (e.g. abundance of similar habitat) or historical factors (e.g. priority effects). However, we generally cannot predict which of these four classes of drivers will affect restoration outcomes. Disparate aspects of restoration outcomes (e.g. species richness, beta diversity and community composition) and their unique responses further complicate our understanding. Finally, these four classes of drivers might differentially affect subsets of the restored community, where, for example, management might shape the abundance and distribution of species of the target community, while other species are more contingent on site, landscape or historical factors. 2. Here, we used variation partitioning to compare the relative importance of management, site, landscape and historical factors for determining the plant community outcomes of 27 prairie restorations in south-west Michigan. 3. We found that management, especially the composition, diversity and density of seed mixes applied, and history, especially site age, were the most important drivers of prairie restoration species richness, beta diversity and composition. Site and landscape factors were only rarely important for restoration outcomes. 4. Finally, we found that comparing the unique responses of sown and non-sown species typically increased our understanding of the dynamics contributing to community-wide restoration outcomes. 5. Synthesis and applications. This is, to our knowledge, the first quantitative comparison of how four major classes of drivers determine the outcome of restoration. Historical legacies and management decisions, but generally not landscape context or local site conditions, shaped plant communities at restored sites. These findings represent an important step towards developing a more predictive framework for understanding contingency in restoration outcomes.

Summary Recovering biological diversity and ecosystem functioning are primary objectives of ecological restoration, yet these outcomes are often unpredictable. Assessments based on functional traits may help with interpreting variability in both community composition and ecosystem functioning because of their mechanistic and generalizable nature. This promise remains poorly realized, however, because tests linking environmental conditions, functional traits, and ecosystem functioning in restoration are rare. Here, we provide such a test through what is to our knowledge the first empirical application of the ‘response–effect trait framework’ to restoration. This framework provides a trait‐based bridge between community assembly and ecosystem functioning by describing how species respond to environmental conditions based on traits and how the traits of species affect ecosystem functioning. Our study took place across 29 prairies restored from former agricultural fields in southwestern Michigan. We considered how environmental conditions affect ecosystem functioning through and independently of measured functional traits. To do so, we paired field‐collected trait data with data on plant community composition and measures of ecosystem functioning and used structural equation modelling to determine relationships between environmental conditions, community‐weighted means of functional traits and ecosystem functioning. Environmental conditions were predictive of trait composition. Sites restored directly from tillage (as opposed to those allowed to fallow) supported taller species with larger seeds and higher specific leaf area (SLA). Site age and fire frequency were both negatively related to SLA. We also found a positive relationship between soil moisture and SLA. Both trait composition and environmental conditions predicted ecosystem functioning, but these relationships varied among the measured functions. Pollination mode (animal pollination) increased and fire frequency decreased floral resource availability, seed mass had a negative effect on below‐ground biomass production, and vegetative height increased decomposition rate. Soil moisture and fire frequency both increased while site age decreased above‐ground biomass production, and site age and soil moisture both increased decomposition rate. Synthesis and applications. Our results suggest that both trait composition and environmental conditions play a role in shaping ecosystem function during restoration, and the importance of each is dependent on the function of interest. Because of this, environmental heterogeneity will be necessary to promote multiple ecosystem functions across restored landscapes. A trait‐based approach to restoration can aid interpretation of variable outcomes through insights into community assembly and ecosystem functioning.

Community assembly filters, which in theory determine the suite of species that arrive at and establish in a community, have tremendous conceptual relevance to restoration. However, the concept has remained largely theoretical, with a paucity of empirical tests. As such, the applicability of assembly filters theory to ecological restoration remains incompletely known. We tested the relative strengths of dispersal and establishment filters by comparing the plant species composition, measured by species' presence/absence, in 29 restored prairies with the seed mixes used to restore each prairie. We found that both establishment and dispersal filters limited prairie similarity to the seed mix. Sown species responded differentially to filters, with a few species limited only by dispersal (seed density), many others limited only by establishment conditions (i.e. organic matter and sand content of soils, land use history, and fire frequency), and others limited by both dispersal and establishment filters. A few species, typically those sown most often, were not restricted by dispersal or establishment filters, likely because they were sown in high enough densities and all sites had suitable environmental conditions. Finally, one group of species established poorly, but we could not attribute this to either dispersal or establishment filters. This information can help land managers select species likely to establish in restorations when sown at sufficient densities. These results illustrate that dispersal and establishment filters limit the establishment of species in restored communities and these filters are species-dependent. Identifying the most limiting filter(s) for species will inform strategies to increase their establishment success.

The loss of biodiversity at local and larger scales has potentially dramatic effects on ecosystem functioning. Many studies have shown that ecosystem functioning depends on biodiversity, but the role of beta diversity, spatial variation in community composition, is less clear than that of local-scale (alpha) diversity. To test the hypothesis that beta diversity would increase ecosystem multifunctionality through variation in species functional traits, we gathered data on plant community composition, plant functional traits, and seven ecosystem functions across 29 restored prairies. We found that averaged multifunctionality (mean of seven ecosystem functions) increased with both taxonomic beta diversity and functional beta diversity. The abundance of the dominant species, big bluestem, played a more minor role, suggesting a limited role for the selection effect. Neither taxonomic nor functional alpha richness was associated with multifunctionality, though this finding may be sensitive to the identity of the functions included because alpha diversity was associated with some individual functions in opposing directions. These findings suggest that in systems structured largely by natural processes, beta diversity (a patchwork of functionally different plant communities) and dominant species abundance may be more important than alpha diversity in fostering ecosystem multifunctionality. These findings suggest the need for an increased focus on community heterogeneity to reestablish functional ecosystems during restoration.

There is strong evidence for a positive relationship between biodiversity and ecosystem functioning at local spatial scales. However, how different aspects of biodiversity relate to multiple ecosystem functions (multifunctionality) across heterogeneous landscapes, and how the magnitude of biodiversity, dominant species, and environmental effects on functioning compare, remain poorly understood. We compared relationships between plant phylogenetic, functional, and taxonomic diversity and ecosystem multifunctionality across 29 restored grasslands. Functional diversity was positively associated with multifunctionality, more strongly than other diversity measures; however, landscape composition explained nearly four times more variation in multifunctionality than did functional diversity, with plots within human-modified landscapes supporting lower multifunctionality. Individual functions were typically more strongly correlated with environmental variables than with diversity. We also found that abundance of the dominant species, Andropogon gerardii, was positively correlated with multifunctionality. Plant diversity, dominant species, and underlying environmental conditions underpin ecosystem multifunctionality in grasslands, but how biodiversity is measured matters for the strength and direction of biodiversity-ecosystem function relationships. Finally, in natural systems environmental variation unrelated to local biodiversity is important for determining ecosystem functioning.

Forbs comprise most of the plant diversity in North American tallgrass prairie and provide vital ecosystem services, but their abundance in prairie restorations is highly variable. Restoration practitioners typically sow C4 grasses in high abundances because they are inexpensive, provide fuel for prescribed fires, can dominate reference sites, and suppress weeds that suppress sown forbs. However, C4 grasses can also suppress sown forbs, calling this practice into question. We evaluated how C4 grasses influence the abundance and diversity of sown forbs in 78 restored prairies across Illinois, Indiana, and Michigan. We found that the direct negative effects of C4 grasses on sown forbs outweighed indirect positive effects that occurred as C4 grasses suppressed nonsown species, which in turn suppressed sown forbs. This pattern was especially strong for the C4 grass big bluestem (Andropogon gerardii). Therefore, strategies to promote big bluestem and other C4 grasses would not promote sown forbs. Although C4 grass cover was not strongly related to two hypothesized drivers (time since fire or site age), seeding density of C4 grasses increased their cover. Sown forb cover also increased with forb seeding density, increased indirectly with fire (through its negative effect on nonsown species), and decreased indirectly with soil water‐holding capacity (through its positive effect on nonsown species). These results highlight the complex interplay of species groups during grassland restoration and show how managers can promote sown forbs in restored prairies: increasing forb seeding density and reducing time since fire and the abundance of C4 grasses and weeds.

By assisting the recovery of disturbed or destroyed ecosystems, ecological restoration plays an important role in biodiversity conservation. Moreover, restoration has been heralded as an ''acid test'' of ecological understanding, by affording the ability to study community assembly, ecosystem function, and human influence over ecosystems across large spatial and long temporal scales. These data sets report the outcome of community assembly, in terms of plant community composition and structure and one important ecosystem function (aboveground biomass production), in 29 prairie restorations in southwestern Michigan. We also report putative forces shaping the outcome of assembly including the species pools (seed mixes applied during restoration), site conditions, landscape context, and land-use history. Detailed knowledge of each restoration effort, including seed mixes used, is unusual and makes these data sets uniquely suited to addressing questions in community assembly by comparing the sown seeds and resulting assembled plant community. For example, we have used the data to test the role of species pools in determining the diversity of assembling communities. We have also used the data to characterize the relative importance of various drivers of community assembly outcomes during restoration, as a step toward resolving the highly contingent and unpredictable outcomes that plague the field of ecological restoration. We suggest that these data sets may prove useful for addressing additional questions in community ecology through the lens of ecological restoration.

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