Phylogenetically and functionally diverse species mixes beget diverse experimental prairies,whether from seeds or plugs

Phylogenetic and functional diversity are relevant for restoration planning, as they influence important ecosystem functions and services. However, it is unknown whether initial phylogenetic and functional diversity of restorations as planned and planted are maintained over time, that is, the extent to which diversity of the restoration planting is reflected in the diversity of the resulting plant community. Furthermore, in the tallgrass prairie, many restorations are planted from seed. Among-species variation in emergence and establishment affects the transition from seed mixes to realized plant communities in these restorations. We evaluated emergence and early establishment of experimental communities in a biodiversity plot experiment designed to test how phylogenetic and functional diversity influence restoration outcomes. We planted the same experimental communities starting from both seeds and plugs to assess differences in establishment. Our results suggest that phylogenetically and functionally diverse species mixes tend to produce phylogenetically and functionally diverse restored plant communities. After 3 years, experimental communities generally maintained their phylogenetic and functional diversity from seed and plug mixes to established vegetation, despite declines in species richness. While plots planted from seeds had on average 1.3 fewer species than plots planted from plugs, phylogenetic and functional diversity did not significantly differ between the two. Furthermore, most species exhibited no significant differences in percent cover when planted from seeds or plugs. Seeds are generally more cost-effective for restoration than plugs, and our results indicate these two establishment methods achieved similar biodiversity outcomes.     

Seeding tallgrass prairie in monospecific patches promotes native species establishment and cover

In tallgrass prairie reconstruction, the way desired seeds are arranged on the landscape may affect species establishment, species persistence, and the establishment and persistence of undesired (nonseeded) species from the local propagule pool. To test effects of species seeding pattern on how grasslands develop spatially, we seeded 20—4 × 4–m bare soil plots with 16 tallgrass prairie species. Treatment plots were divided into 16—1 × 1–m subplots, 64—0.5 × 0.5–m subplots, 256—0.25 × 0.25–m subplots, or 1,024—0.125 × 0.125–m subplots. Each species was hand broadcast into separate subplots (1 m² total area/species) within each plot. An additional treatment included uniformly mixing and broadcasting all seeds across a plot. We recorded species cover at the 0.125 × 0.125–m scale within each plot at the beginning of the second and third growing seasons. While species persistence was greatest within plots seeded with larger subplots, plots with smaller subplots were more spatially diverse and less occupied by nonseeded species over time than larger subplot and mixed plots. As is common in reconstruction efforts, establishment was variable among species and seeding with monospecific subplots enhanced colonization of desired rhizomatous species (e.g., Heliopsis helianthoides, Monarda fistulosa, Elymus virginicus) into unoccupied locations at the expense of species from the local propagule pool. Results suggest that seeding species in smaller, monospecific patches could result in grasslands with a more balanced native species composition than those established with a seed mixture approach.     

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

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Responses of breeding birds in tallgrass prairie to fire and cattle grazing

ABSTRACT.   No other group of North American birds has declined as precipitously and over so large an area as has the grassland assemblage. In the Flint Hills of Kansas, the largest extant region of tallgrass prairie, annual spring burning of rangeland has largely replaced traditional regimes and natural patterns with longer intervals between burns. I examined effects of burning and low-intensity cattle grazing on abundances of seven bird species at Konza Prairie Research Natural Area in June 2002 and 2003. Every species was affected by fire, with Upland Sandpipers ( Bartramia longicauda ) more abundant, and six species—Grasshopper Sparrow ( Ammodramus savannarum ), Henslow's Sparrow ( A. henslowii ), Dickcissel ( Spiza americana ), Eastern Meadowlark ( Sturnella magna ), Brown-headed Cowbird ( Molothrus ater ), and Bell's Vireo ( Vireo bellii )—either less abundant or absent at sites in the breeding season following a fire. These results demonstrate that annual burning limits the potential of much of the Flint Hills prairie to harbor high breeding densities of many grassland birds. On the other hand, I found a trade-off between immediate and longer-term effects of burning for several grass-dependent species. Grasshopper Sparrows, Henslow's Sparrows, and Eastern Meadowlarks, although more numerous in areas that were not burned the preceding spring, were less abundant at sites burned every 4 yrs than those burned at shorter intervals. In contrast, shrub-dependent Bell's Vireos were more abundant at sites burned every 4 yrs. Upland Sandpipers, Grasshopper Sparrows, and Eastern Meadowlarks were more abundant in grazed areas. Use of alternatives to annual burning could increase habitat heterogeneity by transforming the Flint Hills into a mosaic of regularly, but asynchronously, burned pastures that would better meet the diverse habitat needs of the region's grassland birds.     

Responses of plant and bird communities to prescribed burning in tallgrass prairies

Historic losses and fragmentation of tallgrass prairie habitat to agriculture and urban development have led to declines in diversity and abundance of plants and birds associated with such habitat. Prescribed burning is a management strategy that has potential for restoring and rejuvenating prairies in fragmented landscapes, and through such restoration, might create habitat for birds dependent upon prairies. To provide improved data for management decision-making regarding the use of prescribed fire in tallgrass prairies, we compared responses of plant and bird communities on five burned and five unburned tallgrass prairie fragments at the DeSoto National Wildlife Refuge, Iowa, USA, from 1995 to 1997. Overall species richness and diversity were unaffected by burning, but individual species of plants and birds were affected by year-treatment interactions, including northern bobwhite (Colinus virginianus) and ring-necked pheasant (Phasianus colchicus), which showed time-delayed increases in density on burned sites. Analyses of species/area relationships indicated that, collectively, many small sites did make significant contributions to plant biodiversity at landscape levels, supporting the overall conservation value of prairie fragments. In contrast, most birds species were present on larger sites. Thus, higher biodiversity in bird communities which contain area-sensitive species might require larger sites able to support larger, more stable populations, greater habitat heterogeneity, and greater opportunity for niche separation.     

High dissimilarity within a multiyear annual record of pollen assemblages from a North American tallgrass prairie

Grassland vegetation varies in composition across North America and has been historically influenced by multiple biotic and abiotic drivers, including fire, herbivory, and topography. Yet, the amount of temporal and spatial variability exhibited among grassland pollen assemblages, and the influence of these biotic and abiotic drivers on pollen assemblage composition and diversity has been relatively understudied. Here, we examine 4 years of modern pollen assemblages collected from a series of 28 traps at the Konza Prairie Long‐Term Ecological Research Area in the Flint Hills of Kansas, with the aim of evaluating the influence of these drivers, as well as quantifying the amount of spatial and temporal variability in the pollen signatures of the tallgrass prairie biome. We include all terrestrial pollen taxa in our analyses while calculating four summative metrics of pollen diversity and composition – beta‐diversity, Shannon index, nonarboreal pollen percentage, and Ambrosia:Artemisia – and find different roles of fire, herbivory, and topography variables in relation to these pollen metrics. In addition, we find significant annual differences in the means of three of these metrics, particularly the year 2013 which experienced high precipitation relative to the other 3 years of data. To quantify spatial and temporal dissimilarity among the samples over the 4‐year study, we calculate pairwise squared‐chord distances (SCD). The SCD values indicate higher compositional dissimilarity across the traps (0.38 mean) among all years than within a single trap from year to year (0.31 mean), suggesting that grassland vegetation can have different pollen signatures across finely sampled space and time, and emphasizing the need for additional long‐term annual monitoring of grassland pollen.     

Ten years of native seed certification in Germany – a summary

Many renaturation projects and compensation areas are based on the use of seeds from regional indigenous wild plants; in the following: native or regional seeds. Despite this, such seeds make up only a small proportion of the total number of seeds used for greening projects; in Germany, for example, it is only around 1% (=200 t per year). Although the market for regional seeds is small, it is highly competitive. High‐priced native seeds compete with flower mixes of unspecified origin and can only be differentiated from them by reliable quality seals. A quality assurance system based on seed legislation (EU Directive 2010/60, preservation mixtures) has been developed in a few European countries. However, quality assurance ends with the sale of the seeds. Thus, seed use remains unmonitored, and often unsuitable material, or material foreign to the region, is planted in restoration areas. Unfortunately, nature conservation has not made seed‐based restoration one of its key issues, neither at the European nor at the national level. Currently there are many different local and regional standards, methods and private certificates that are confusing for users and which provide little continuity and predictability for producers. We recommend the establishment of an EU directive or a broadly agreed recommendation to the EU member states, spearheaded by nature conservation, which would define the standards for producing and using native seeds (e.g. harmonised regions that cross national borders, quality regulations). At the same time, wild plant interest groups should combine existing structures in order to strengthen seed‐based restoration through international cooperation.     

Restoration of Prairie Community Structure and Ecosystem Function in an Abandoned Hayfield: A Sowing Experiment

Using a multispecies seed sowing experiment, we investigated the roles of seed and microsite limitation in constraining the restoration of native prairie diversity and ecosystem function in an abandoned upland hayfield in northeastern Kansas. Seeds of 32 native and naturalized plant species from the regional pool were sown into undisturbed and experimentally disturbed field plots. After six growing seasons, experimental sowing led to major shifts in species and functional group composition, increases in native species abundance and floristic quality, declines in abundance of non‐native species, and increases in plant diversity. These changes in community structure led to significant changes at the ecosystem level including increases in light capture, peak biomass, primary production, litter biomass, root biomass, and C storage in roots. Our findings reveal the importance of seed limitations in constraining the natural recovery of prairie vegetation, biodiversity, and ecosystem functioning in this grassland and confirm broadcast sowing as a useful tool for the restoration of upland hayfield sites.     

Soil Heterogeneity Effects on Tallgrass Prairie Community Heterogeneity: An Application of Ecological Theory to Restoration Ecology

Spatial heterogeneity of resources can influence plant community composition and diversity in natural communities. We manipulated soil depth (two levels) and nutrient availability (three levels) to create four heterogeneity treatments (no heterogeneity, depth heterogeneity, nutrient heterogeneity, and depth + nutrient heterogeneity) replicated in an agricultural field seeded to native prairie species. Our objective was to determine whether resource heterogeneity influences species diversity and the trajectory of community development during grassland restoration. The treatments significantly increased heterogeneity of available inorganic nitrogen (N), soil water content, and light penetration. Plant diversity was indirectly related to resource heterogeneity through positive relationships with variability in productivity and cover established by the belowground manipulations. Diversity was inversely correlated with the average cover of the dominant grass, Switchgrass (Panicum virgatum), which increased over time in all heterogeneity treatments and resulted in community convergence among the heterogeneity treatments over time. The success of this cultivar across the wide range of resource availability was attributed to net photosynthesis rates equivalent to or higher than those of the native prairie plants in the presence of lower foliar N content. Our results suggest that resource heterogeneity alone may not increase diversity in restorations where a dominant species can successfully establish across the range of resource availability. This is consistent with theory regarding the role of ecological filters on community assembly in that the establishment of one species best adapted for the physical and biological conditions can play an inordinately important role in determining community structure.     

Soil modification by invasive plants: effects on native and invasive species of mixed-grass prairies

Invasive plants are capable of modifying attributes of soil to facilitate further invasion by conspecifics and other invasive species. We assessed this capability in three important plant invaders of grasslands in the Great Plains region of North America: leafy spurge (Euphorbia esula), smooth brome (Bromus inermis) and crested wheatgrass (Agropyron cristatum). In a glasshouse, these three invasives or a group of native species were grown separately through three cycles of growth and soil conditioning in both steam-pasteurized and non-pasteurized soils, after which we assessed seedling growth in these soils. Two of the three invasive species, Bromus and Agropyron, exhibited significant self-facilitation via soil modification. Bromus and Agropyron also had significant facilitative effects on other invasives via soil modification, while Euphorbia had significant antagonistic effects on the other invasives. Both Agropyron and Euphorbia consistently suppressed growth of two of three native forbs, while three native grasses were generally less affected. Almost all intra- and interspecific effects of invasive soil conditioning were dependent upon presence of soil biota from field sites where these species were successful invaders. Overall, these results suggest that that invasive modification of soil microbiota can facilitate plant invasion directly or via ‘cross-facilitation’ of other invasive species, and moreover has potential to impede restoration of native communities after removal of an invasive species. However, certain native species that are relatively insensitive to altered soil biota (as we observed in the case of the forb Linum lewisii and the native grasses), may be valuable as ‘nurse’species in restoration efforts.     

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.     

Physiological and growth responses of switchgrass (Panicum virgatum L.) in native stands under passive air temperature manipulation

In the central Great Plains of North America, climate change predictions include increases in mean annual temperature of 1.5–5.5 °C by 2100. Ecosystem responses to increased temperatures are likely to be regulated by dominant plant species, such as the potential biofuel species Panicum virgatum (switchgrass) in the tallgrass prairie. To describe the potential physiological and whole‐plant responses of this species to future changes in air temperatures, we used louvered open‐sided chambers (louvered OSC; 1 × 1 m, adjustable height) to passively alter canopy temperature in native stands of P. virgatum growing in tallgrass prairie at varying topographic positions (upland/lowland). The altered temperature treatment decreased daily mean temperatures by 1 °C and maximum temperatures by 4 °C in May and June, lowered daytime stomatal conductance and transpiration, decreased tiller density, increased specific leaf area, and delayed flowering. Among topographic contrasts, aboveground biomass, flowering tiller density, and tiller weight were greater in lowland sites compared to upland sites, with no temperature treatment interactions. Differences in biomass production responded more to topography than the altered temperature treatment, as soil water status varied considerably between topographic positions. These results indicate that while water availability as a function of topography was a strong driver of plant biomass, many leaf‐level physiological processes were responsive to the small decreases in daily mean and maximum temperature, irrespective of landscape position. The varying responses of leaf‐level gas exchange and whole‐plant growth of P. virgatum in native stands to altered air temperature or topographic position illustrate that accurately forecasting yields for P. virgatum in mixed communities will require greater integration of physiological responses to simulated climate change (increased temperature) and resource availability over natural environmental gradients (soil moisture).     

Patterns of Species Richness and Composition in Re-Created Grassland

The success of many prairie restorations is not well documented. A restoration begun in 1975 at the Fermi National Accelerator Laboratory near Chicago, Illinois allows assessment of restoration efforts as well as changes through time. Data are presented on species richness and composition for 13 restorations planted in successive years between 1975 and 1990 and two remnant prairies. Presence of species was recorded using a stratified random design. Species richness at several scales and non‐metric multidimensional scaling ordination were used to assess trends in the vegetation. Species richness declined through time at all scales examined and was always less in the restored prairies than that found in the remnant prairies. Species composition changed with time but not in the direction of the composition found in the remnants. Our understanding of the maintenance of species richness is not sufficient to allow the re‐creation of patterns of species found in remnant grassland communities.     

Plant community responses to bison reintroduction on the Northern Great Plains,United States: a test of the keystone species concept

Keystone species restoration, or the restoration of species whose effect on an ecosystem is much greater than their abundance would suggest, is a central justification for many wildlife reintroduction projects globally. Following restoration, plains bison (Bison bison L.) have been identified as a keystone species in the tallgrass prairie ecoregion, but we know of no research to document similar effects in the mixed‐grass prairie where restoration efforts are ongoing. This study addresses whether Northern Great Plains (NGP) mixed‐grass prairie plant communities exhibit traits consistent with four central keystone effects documented for bison in the tallgrass prairie. We collected species composition, diversity, abundance, bare ground cover, and plant height data in three treatments: where livestock (Bos taurus L.) continuously grazed, livestock were removed for 10 years, and bison have been introduced and resident for 10 years. We observed mixed support for bison acting as keystone species in this system. Supporting the keystone role of bison, we observed higher species richness and compositional heterogeneity (β‐diversity) in the bison treatment than either the livestock retention or livestock removal treatments. However, we observed comparable forb, bare ground, and plant height heterogeneity between bison‐restored sites and sites where livestock were retained, contradicting reported keystone effects in other systems. Our results suggest that after 10 years of being restored, bison partially fulfill their role as a keystone species in the mixed‐grass prairie, and we encourage continued long‐term data collection to evaluate their influence in the NGP.     

Woody structure facilitates invasion of woody plants by providing perches for birds

Woody encroachment threatens prairie ecosystems globally, and thus understanding the mechanisms that facilitate woody encroachment is of critical importance. Coastal tallgrass prairies along the Gulf Coast of the US are currently threatened by the spread of several species of woody plants. We studied a coastal tallgrass prairie in Texas, USA, to determine if existing woody structure increased the supply of seeds from woody plants via dispersal by birds. Specifically, we determined if (i) more seedlings of an invasive tree (Tridacia sebifera) are present surrounding a native woody plant (Myrica cerifera); (ii) wooden perches increase the quantity of seeds dispersed to a grassland; and (iii) perches alter the composition of the seed rain seasonally in prairie habitats with differing amounts of native and invasive woody vegetation, both underneath and away from artificial wooden perches. More T. sebifera seedlings were found within M. cerifera patches than in graminoid‐dominated areas. Although perches did not affect the total number of seeds, perches changed the composition of seed rain to be less dominated by grasses and forbs. Specifically, 20–30 times as many seeds of two invasive species of woody plants were found underneath perches independent of background vegetation, especially during months when seed rain was highest. These results suggest that existing woody structure in a grassland can promote further woody encroachment by enhancing seed dispersal by birds. This finding argues for management to reduce woody plant abundance before exotic plants set seeds and argues against the use of artificial perches as a restoration technique in grasslands threatened by woody species.     

Active versus passive restoration of tallgrass prairie in the U.S. Midwest: plant species diversity and assemblage,net primary production and soil carbon sequestration

We investigated the plant species diversity, community assemblage, net primary production and soil carbon sequestration in an old field (OF), which went through passive restoration with spontaneous secondary succession, and a restored prairie (RP), which went through active restoration with reintroduction of native plants. The passive restoration in OF did not attain as high diversity and richness of plant species as did in RP. Our NMS (nonmetric multidimensional scaling) ordination revealed a clear divergence of RP and OF vegetation with dissimilar species compositions to separate paths of succession trajectory. The spontaneous succession in OF has made very little progress due to a potentially strong inhibitory priority effect of invasive plants. The belowground biomass in the high-diversity RP was 1.3× higher than the low-diversity OF. We found no measurable difference in soil C sequestration between RP and OF at this time, possibly due to edaphic factors, lack of legumes, or both. Human intervention is needed for restoring abandoned croplands—highly disturbed, fragmented and isolated habitats—to tallgrass prairies in the U.S. Midwest. Our recommended intervention includes but is not limited to removal of exotic plants, further augmentation of native plants (particularly legumes), and periodic burns.