A primary goal of restoration ecology is to understand the factors that generate variability in species diversity and composition among restorations. Plant communities may assemble deterministically toward a common community type, or they may assemble stochastically, ending differently because of weather conditions during establishment, soil legacy effects, or exotic species propagule pressure. To test these alternative hypotheses, we sampled plant communities and soil at 93 randomly selected restored prairies distributed throughout Iowa, USA. Five remnant sites were sampled as a reference. We tested our hypotheses using multiple regressions and investigated the strength of direct and indirect effects on species diversity and richness using structural equation models. The prairie restorations were highly variable in their age, size, diversity, soil characteristics, and how they were managed post‐seeding. The strongest predictor of plant species richness and diversity was the degree of invasion, as measured by the abundance of exotic species. Restorations planted with species‐rich seed mixes had reduced exotic species abundance, which led indirectly to higher species richness of restorations. Sites with higher organic matter and a more linear shape had a direct positive effect on exotic abundance, which in turn decreased diversity. We found little support for deterministic assembly, and diversity did not increase with the age of planting. Our results indicate that restored prairie communities tend to assemble into states of high or low diversity, driven by invasion from exotic plant species. Management of exotic species is essential for maximizing species diversity in temperate grassland restorations.

Premise Variation in pollinator effectiveness may contribute to pollen limitation in fragmented plant populations. In plants with multiovulate ovaries, the number of conspecific pollen grains per stigma often predicts seed set and is used to quantify pollinator effectiveness. In the Asteraceae, however, florets are uniovulate, which suggests that the total amount of pollen deposited per floret may not measure pollinator effectiveness. We examined two aspects of pollinator effectiveness—effective pollen deposition and effective pollen movement—for insects visiting Echinacea angustifolia, a composite that is pollen limited in small, isolated populations. Methods We filmed insect visits to Echinacea in two prairie restorations and used these videos to quantify behavior that might predict effectiveness. To quantify effective pollen deposition, we used the number of styles shriveled per visit. To quantify effective pollen movement, we conducted paternity analysis on a subset of offspring and measured the pollen movement distance between mates. Results Effective pollen deposition varied among taxa. Andrena helianthiformis, a Heliantheae oligolege, was the most effective taxon, shriveling more than twice the proportion of styles as all other visitors. Differences in visitor behavior on a flowering head did not explain variation in effective pollen deposition, nor did flowering phenology. On average, visitors moved pollen 16 m between plants, and this distance did not vary among taxa. Conclusions Andrena helianthiformis is an important pollinator of Echinacea. Variation in reproductive fitness of Echinacea in fragmented habitat may result, in part, from the abundance of this species.

The decline in migratory monarch butterflies (Danaus plexippus) over the past 20 years has been attributed to several drivers, including loss of their host plants (milkweeds Asclepias spp.). This has sparked widespread interest in milkweed ecology and restoration. We developed a model on environmental and habitat‐type variables to predict milkweed abundance by sampling 93 prairie plantings (47 conservation plantings and 46 roadsides) and 5 unplowed prairie remnants throughout the state of Iowa, United States. Milkweeds were censused in 10–25 random locations within each site, and data on plant diversity, age of planting, soil characteristics, and management were tested as predictors of abundance. Milkweed densities of all species combined were highest in remnant prairies (8,705 stems/ha), intermediate in roadside plantings (1,274 stems/ha), and lowest in conservation plantings (212 stems/ha). Most milkweeds were common milkweeds Asclepias syriaca, which were more abundant in roadside than conservation plantings. Remnants contained the most milkweed species. Total milkweed and common milkweed abundance were both predicted by higher soil pH, a more linear site shape, and lower soil bulk density across restorations. Our results indicate that common milkweed is maintained by disturbance, and establishes readily in rural roadside habitat. Remnants are important as reservoirs for multiple milkweed species and should be protected.

Seed limitation represents a fundamental constraint to the restoration of native plant communities, and practitioners often apply seed additions to overcome this barrier. However, surprisingly few studies have experimentally tested whether seed additions can increase diversity in herbaceous communities of oak woodlands, which have undergone large‐scale transformation due to logging, altered fire regimes and invasion by non‐native species. Previous studies suggest that structural (thinning of woody biomass) and process‐based (prescribed fire) restoration treatments alone are unlikely to restore the full breadth of taxonomic and functional diversity in the herb layer, which accounts for most species in woodland ecosystems. To explore whether seed additions can improve restoration outcomes in an oak woodland, we sowed high‐diversity seed mixes in paired transects (seeded vs. controls) along a topographic gradient in a degraded site undergoing restoration with non‐native shrub removal, selective tree thinning and prescribed fire. Seed mixes contained native forbs, grasses and sedges from locally sourced material (n = 169 total species) in the regional species pool, and were designed to match species' habitat affinity to appropriate locations along the topographic gradient. The herb flora was sampled pre‐seeding, and for two consecutive years after additions. Seed additions significantly altered community and functional composition, and increased native species richness by 29% (43.0 vs 55.4), and floristic quality by 30% relative to controls. However, fewer than half of the sown species were established 2 years after planting, suggesting that dispersal and establishment limitation are both important barriers to the recovery of the herb flora in oak woodlands. We also tested if species' sown abundance, conservatism or functional group predicted establishment success. Species sown at high abundances and less conservative species recruited the most reliably. Grass and forb establishment rates were more dependent on seeding rate than sedges or legumes, and the mechanisms behind this trend merit further investigation. We found that adding high‐diversity seed mixes in conjunction with non‐native shrub removal, canopy thinning and burning, can accelerate recovery of herbaceous communities in a highly degraded woodland.

Questions Are community‐weighted plant functional traits related to the ‘fast–slow’ growth rate continuum different in seed mixes, restored and reference prairies? If so, then what are the traits that explain differences? Remnant prairies, which serve as reference sites for restorations, often have higher plant species diversity and a lower abundance of exotic (non‐native) species than their restoration counterparts. They may differ in their community‐weighted functional traits as well, although this is poorly studied. Location Tallgrass prairies and abandoned fields in Iowa, United States, North America. Methods We compared functional traits between remnant prairies, restoration seed mixes, and established restorations to determine if community‐weighted trait values (∑pi × traits) are comparable. Relative abundance was determined by previous sampling of 93 restorations, 48 seed mixes and five remnant sites. Functional traits, including leaf specific leaf area (SLA) and leaf dry matter content (LDMC), and plant height, were measured for 145 native and 39 non‐native plant species, 663 plants, and 1326 leaves. Results Seed mixes had greater community‐weighted LDMC and lower SLA than remnants, perhaps because of human selection for high grass abundance in mixes. Established restorations had lower community‐weighted LDMC and higher SLA than remnants and seed mixes, and restorations had a large exotic species component. The proportional abundance of exotics in restorations explained trait differences between remnants and restorations. After taking into account functional groupings, perennial exotic species, on average, had higher SLA and lower LDMC than perennial native species. Remnants and established restorations had similar plant heights, and height was not significantly different between native and exotic species. Conclusions Our results indicate that seed mixes differ from remnants in a manner that was the opposite of the difference between seed mixes and established restorations. Although seed mixes favored species with slower growth rate trait compositions, the species that established in restored prairies tended to have traits associated with faster growth rates. We recommend developing prairie restoration seed mixes with a greater proportion of forbs, especially those with lower LDMC and higher SLA.