Restored native prairie supports abundant and species‐rich native bee communities on conventional farms

Native pollinators are increasingly needed on conventional farms yet rarely fostered via management. One solution is habitat restoration in marginal areas, but colonization may be constrained if resident pollinator richness is low or if restored areas fail to provide sufficient floral or nesting resources. We quantified restoration outcomes for native bees, and associated resources, on three conventional farms with forb‐grass prairie plantings on marginal areas of varying sizes, in a heavily farmed region of central North America. We tested bee abundance and richness in restored prairie versus the dominant habitats of the region—crops, forest remnants, and edges of fields and roads. Restored prairie supported 2× more species (95 of 119 total species) and 3× more bees (72% of captured individuals) compared to the other cover types. All richness and abundance differences among habitat types were associated with higher floral resources in restored prairie. Thirty percent of the bee species were unique to prairie, consistent with long‐distance dispersal but begging the question of origin given the absence of prairie regionally. Our results suggest that road and field edges may be the source, as these areas had more floral and nesting resources than forest or crop fields combined and supported 55% of all species despite covering only approximately 5% of the sampled farms. Habitat scarcity is not the only constraint on native bees in agricultural landscapes, with increasing concern over disease and chemicals. However, we observed that restored areas on marginal lands of conventional farms can support abundant and species‐rich populations of native bees.     

Positive diversity effects on productivity in mixtures of arable weed species as related to density-size relationships

Aims Diversity–productivity relationships among herbaceous species have mostly been studied in grasslands, while less is known about diversity effects among weedy species with a short life cycle.Methods We studied diversity–productivity relationships, shoot density, size and allometry in experimental communities of different species richness (one, three, six, and nine species), functional group number (one to three functional groups: grasses, small herbs and tall herbs) and functional group evenness (even and uneven number of species per functional group) based on a pool of nine arable weed species with a short life cycle in a 2-year experiment.Important findings Higher species richness increased above- and belowground biomass production in both years of the experiment. Additive partitioning showed that positive selection effects increased with increasing species richness and functional group number, while positive complementarity effects were greater when tall herbs were present. Relative yield totals were larger than 1 across all species richness levels but did not increase with species richness, which is consistent with constant positive complementarity effects. Community biomass production and diversity effects increased in the second year of the experiment, when communities achieved greater shoot densities and average shoot sizes. At the community level, varying productivity was mainly attributable to variation in mean shoot sizes. Tall herbs reached greater observed/expected relative yields (=overyielding) due to increased shoot sizes, underyielding of small herbs was mainly attributable to decreased shoot sizes, while grasses partly compensated for reduced shoot sizes by increasing densities. Shifts in community-level density–size relationships and changes in shoot allometry in favour of greater height growth indicated that a greater biomass at a given density was due to increased dimensions of occupied canopy space. We conclude that diversity effects are also possible among short-lived arable weed species, but selection effects through sizes differences among species are key for positive species richness–productivity relationships.     

Overyielding in experimental grassland communities – irrespective of species pool or spatial scale

In a large integrated biodiversity project (‘The Jena Experiment’ in Germany) we established two experiments, one with a pool of 60 plant species that ranged broadly from dominant to subordinate competitors on large 20 × 20 m and small 3.5 × 3.5 m plots (= main experiment), and one with a pool of nine potentially dominant species on small 3.5 × 3.5 m plots (= dominance experiment). We found identical positive species richness–aboveground productivity relationships in the main experiment at both scales. This result suggests that scaling up, at least over the short term, is appropriate in interpreting the implications of such experiments for larger‐scale patterns. The species richness–productivity relationship was more pronounced in the experiment with dominant species (46.7 and 82.6% yield increase compared to mean monoculture, respectively). Additionally, transgressive overyielding occurred more frequently in the dominance experiment (67.7% of cases) than in the main experiment (23.4% of cases). Additive partitioning and relative yield total analyses showed that both complementarity and selection effects contributed to the positive net biodiversity effect.     

Top‐down control of rare species abundances by native ungulates in a grassland restoration

Ecological restorations are predicted to increase in species diversity over time until they reach reference levels. However, chronosequence studies in grasslands often show that diversity peaks after the first few years and then declines over time as grasses become more dominant. We addressed whether bison grazing and seed additions could prevent this decline in diversity. Exclosures that prevented bison grazing were compared with grazed plots over 4 years, and seed additions were conducted inside and outside exclosures to test for seed and microsite limitations. A previous study conducted 4‐months post seeding found that local species richness was primarily seed limited, but that grazing could sometimes increase seedling emergence. Here, we tested whether increased seedling emergence led to longer‐term increases in the species diversity of the plant community. We found that the seed addition effect grew smaller and the grazing effect grew stronger over time, and that seed additions affected the abundance of added species only when plots were grazed. Grazed plots had higher species diversity and lower biomass and litter buildup compared to non‐grazed plots. Our results suggest that moderate grazing by bison or management that mimics grazing can maintain diversity in grass‐dominated situations. Our results also emphasize the need to follow seed additions over several years to assess correctly whether seed limitation exists.     

Functional dominance rather than taxonomic diversity and functional diversity mainly affects community aboveground biomass in the Inner Mongolia grassland

The relationship between biodiversity and productivity has been a hot topic in ecology. However, the relative importance of taxonomic diversity and functional characteristics (including functional dominance and functional diversity) in maintaining community productivity and the underlying mechanisms (including selection and complementarity effects) of the relationship between diversity and community productivity have been widely controversial. In this study, 194 sites were surveyed in five grassland types along a precipitation gradient in the Inner Mongolia grassland of China. The relationships between taxonomic diversity (species richness and the Shannon–Weaver index), functional dominance (the community‐weighted mean of four plant traits), functional diversity (Rao's quadratic entropy), and community aboveground biomass were analyzed. The results showed that (1) taxonomic diversity, functional dominance, functional diversity, and community aboveground biomass all increased from low to high precipitation grassland types; (2) there were significant positive linear relationships between taxonomic diversity, functional dominance, functional diversity, and community aboveground biomass; (3) the effect of functional characteristics on community aboveground biomass is greater than that of taxonomic diversity; and (4) community aboveground biomass depends on the community‐weighted mean plant height, which explained 57.1% of the variation in the community aboveground biomass. Our results suggested that functional dominance rather than taxonomic diversity and functional diversity mainly determines community productivity and that the selection effect plays a dominant role in maintaining the relationship between biodiversity and community productivity in the Inner Mongolia grassland.     

Flowering in grassland predicted by CO2 and resource effects on species aboveground biomass

Continuing enrichment of atmospheric CO₂ may change plant community composition, in part by altering the availability of other limiting resources including soil water, nutrients, or light. The combined effects of CO₂ enrichment and altered resource availability on species flowering remain poorly understood. We quantified flowering culm and ramet production and biomass allocation to flowering culms/ramets for 10 years in C₄‐dominated grassland communities on contrasting soils along a CO₂ concentration gradient spanning pre‐industrial to expected mid‐21st century levels (250–500 μl/L). CO₂ enrichment explained up to 77% of the variation in flowering culm count across soils for three of the five species, and was correlated with flowering culm count on at least one soil for four of five species. In contrast, allocation to flowering culms was only weakly correlated with CO₂ enrichment for two species. Flowering culm counts were strongly correlated with species aboveground biomass (AGB; R² = .34–.74), a measure of species abundance. CO₂ enrichment also increased soil moisture and decreased light levels within the canopy but did not affect soil inorganic nitrogen availability. Structural equation models fit across the soils suggested species‐specific controls on flowering in two general forms: (1) CO₂ effects on flowering culm count mediated by canopy light level and relative species AGB (species AGB/total AGB) or by soil moisture effects on flowering culm count; (2) effects of canopy light level or soil inorganic nitrogen on flowering and/or relative species AGB, but with no significant CO₂ effect. Understanding the heterogeneity in species responses to CO₂ enrichment in plant communities across soils in edaphically variable landscapes is critical to predict CO₂ effects on flowering and other plant fitness components, and species potential to adapt to future environmental changes.     

Biodiversity, productivity and the temporal stability of productivity: patterns and processes

Theory predicts that the temporal stability of productivity, measured as the ratio of the mean to the standard deviation of community biomass, increases with species richness and evenness. We used experimental species mixtures of grassland plants to test this hypothesis and identified the mechanisms involved. Additionally, we tested whether biodiversity, productivity and temporal stability were similarly influenced by particular types of species interactions. We found that productivity was less variable among years in plots planted with more species. Temporal stability did not depend on whether the species were planted equally abundant (high evenness) or not (realistically low evenness). Greater richness increased temporal stability by increasing overyielding, asynchrony of species fluctuations and statistical averaging. Species interactions that favoured unproductive species increased both biodiversity and temporal stability. Species interactions that resulted in niche partitioning or facilitation increased both productivity and temporal stability. Thus, species interactions can promote biodiversity and ecosystem services.     

Feedback from plant species change amplifies CO2 enhancement of grassland productivity

Dynamic global vegetation models simulate feedbacks of vegetation change on ecosystem processes, but direct, experimental evidence for feedbacks that result from atmospheric CO₂ enrichment is rare. We hypothesized that feedbacks from species change would amplify the initial CO₂ stimulation of aboveground net primary productivity (ANPP) of tallgrass prairie communities. Communities of perennial forb and C₄ grass species were grown for 5 years along a field CO₂ gradient (250–500 μL L^?1) in central Texas USA on each of three soil types, including upland and lowland clay soils and a sandy soil. CO₂ enrichment increased community ANPP by 0–117% among years and soils and increased the contribution of the tallgrass species Sorghastrum nutans (Indian grass) to community ANPP on each of the three soil types. CO₂‐induced changes in ANPP and Sorghastrum abundance were linked. The slope of ANPP‐CO₂ regressions increased between initial and final years on the two clay soils because of a positive feedback from the increase in Sorghastrum fraction. This feedback accounted for 30–60% of the CO₂‐mediated increase in ANPP on the upland and lowland clay soils during the final 3 years and 1 year of the experiment, respectively. By contrast, species change had little influence on the ANPP‐CO₂ response on the sandy soil, possibly because Sorghastrum increased largely at the expense of a functionally similar C₄ grass species. By favoring a mesic C₄ tall grass, CO₂ enrichment approximately doubled the initial enhancement of community ANPP on two clay soils. The CO₂‐stimulation of grassland productivity may be significantly underestimated if feedbacks from plant community change are not considered.     

Species reordering,not changes in richness,drives long‐term dynamics in grassland communities

Determining how ecological communities will respond to global environmental change remains a challenging research problem. Recent meta‐analyses concluded that most communities are undergoing compositional change despite no net change in local species richness. We explored how species richness and composition of co‐occurring plant, grasshopper, breeding bird and small mammal communities in arid and mesic grasslands changed in response to increasing aridity and fire frequency. In the arid system, grassland and shrubland plant and breeding bird communities were undergoing directional change, whereas grasshopper and small mammal communities were stable. In the mesic system, all communities were undergoing directional change regardless of fire frequency. Despite directional change in composition in some communities, species richness of all communities did not change because compositional change resulted more from reordering of species abundances than turnover in species composition. Thus, species reordering, not changes in richness, explains long‐term dynamics in these grass and shrub dominated communities.     

一年生和多年生豆禾混播草地超产与多样性效应的比较

为了测度一年生和多年生豆禾混播草地的超产效应与植物多样性效应的关系, 明确一年生和多年生混播草地的高产优势, 探索豆禾混播草地多样性效应和超产效应对其生态功能的响应机制, 于2013-2015年在新疆伊犁地区昭苏盆地开展了3年的牧草产量观测试验。该试验设置3种牧草混播种类和混播比例, 分别为一年生豆禾混播草地(2种牧草混播, AM2)和多年生豆禾混播草地(2种牧草混播, PM2; 4种牧草混播, PM4; 6种牧草混播, PM6), 豆禾混播比例分别为6:4、5:5和4:6。结果表明: 1) 2013、2014年和3年平均值AM2的超产幅度小于PM2和PM6, 2015年AM2的超产幅度大于PM2、PM4和PM6; 混播群落生产力与群落组分中生产力最高产物种单产以及各组分种平均单产的差值表现出相似的规律。2) 2013、2014年和3年平均值AM2的互补效应大于PM2、PM4和PM6, AM2的选择效应则远小于互补效应, PM2、PM4和PM6的互补效应则比较稳定。3)物种丰富度和物种均匀度与牧草产量(群落生产力)大部分情况下呈单峰的“饱和上升型”模式, 分别在4种牧草混播和豆禾混播比例为5:5时, 具有较高生产力。4)多年生豆禾混播草地的互补效应、选择效应和多样性净效应均随生长年限的延长而呈下降趋势, 也导致了超产幅度、超产效应及其稳定性的下降。由此可见, 在建植初期, 互补效应和选择效应共同主导了多年生豆禾混播草地的超产效应, 而随着生长年限的延长, 选择效应则成为主要影响因素; 一年生豆禾混播草地的超产效应则一直受互补效应的影响。     

Interannual variability in responses of belowground net primary productivity (NPP) and NPP partitioning to long‐term warming and clipping in a tallgrass prairie

The dynamics of belowground net primary productivity (BNPP) is of fundamental importance in understanding carbon (C) allocation and storage in grasslands. However, our knowledge of the interannual variability in response of BNPP to ongoing global warming is limited. In this study, we explored temporal responses of BNPP and net primary productivity (NPP) partitioning to warming and clipping in a tallgrass prairie in Oklahoma, USA. Infrared heaters were used to elevate soil temperature by approximately 2 °C since November 1999. Annual clipping was to mimic hay harvest. On average from 2005 to 2009, warming increased BNPP by 41.89% in the unclipped subplots and 66.93% in the clipped subplots, with significant increase observed in wet years. Clipping also had significant positive impact on BNPP, which was mostly found under warming. Overall, f_BNPP, the fraction of BNPP to NPP, increased under both warming and clipping treatments, more in dry years. Water availability (either precipitation or soil moisture) was the most limiting factor for both BNPP and f_BNPP. It strongly dominated the interannual variability in NPP, f_BNPP, and their responses to warming and clipping. Our results suggest that water availability regulates tallgrass prairie's responses to warming and land use change, which may eventually influence the global C cycle. With increasing variability in future precipitation patterns, warming effects on the vegetation in this region may become less predictable.     

Lessons from the fish: a multi‐species analysis reveals common processes underlying similar species‐genetic diversity correlations

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Seed mix design and first year management influence multifunctionality and cost‐effectiveness in prairie reconstruction

Agricultural intensification continues to diminish many ecosystem services in the North American Corn Belt. Conservation programs may be able to combat these losses more efficiently by developing initiatives that attempt to balance multiple ecological benefits. In this study, we examine how seed mix design and first year management influence three ecosystem services commonly provided by tallgrass prairie reconstructions (erosion control, weed resistance, and pollinator resources). We established research plots with three seed mixes, with and without first year mowing. The grass‐dominated “Economy” mix had 21 species and a 3:1 grass‐to‐forb seeding ratio. The forb‐dominated “Pollinator” mix had 38 species and a 1:3 grass‐to‐forb seeding ratio. The grass:forb balanced “Diversity” mix, which was designed to resemble regional prairie remnants, had 71 species and a 1:1 grass‐to‐forb ratio. To assess ecosystem services, we measured native stem density, cover, inflorescence production, and floral richness from 2015 to 2018. The Economy mix had high native cover and stem density, but produced few inflorescences and had low floral richness. The Pollinator mix had high inflorescence production and floral richness, but also had high bare ground and weed cover. The Diversity mix had high inflorescence production and floral richness (comparable to the Pollinator mix) and high native cover and stem density (comparable to the Economy mix). First year mowing accelerated native plant establishment and inflorescence production, enhancing the provisioning of ecosystem services during the early stages of a reconstruction. Our results indicate that prairie reconstructions with thoughtfully designed seed mixes can effectively address multiple conservation challenges.     

Helianthus maximiliani and species fine‐scale spatial pattern affect diversity interactions in reconstructed tallgrass prairies

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Soil microbiome mediates positive plant diversity‐productivity relationships in late successional grassland species

Which processes drive the productivity benefits of biodiversity remain a critical, but unanswered question in ecology. We tested whether the soil microbiome mediates the diversity‐productivity relationships among late successional plant species. We found that productivity increased with plant richness in diverse soil communities, but not with low‐diversity mixtures of arbuscular mycorrhizal fungi or in pasteurised soils. Diversity‐interaction modelling revealed that pairwise interactions among species best explained the positive diversity‐productivity relationships, and that transgressive overyielding resulting from positive complementarity was only observed with the late successional soil microbiome, which was both the most diverse and exhibited the strongest community differentiation among plant species. We found evidence that both dilution/suppression from host‐specific pathogens and microbiome‐mediated resource partitioning contributed to positive diversity‐productivity relationships and overyielding. Our results suggest that re‐establishment of a diverse, late successional soil microbiome may be critical to the restoration of the functional benefits of plant diversity following anthropogenic disturbance.