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.     

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.     

Effects of Nutrient Manipulations and Grass Removal on Cover,Species Composition,and Invasibility of a Novel Grassland in Colorado

Cover and richness of a 5‐year revegetation effort were studied with ,respect to small‐scale disturbance and nutrient manipulations. The site, originally a relict tallgrass prairie mined for gravel, was replanted to native grasses using a seed mixture of tall‐, mixed‐, and short‐grass species. Following one wet and three relatively dry years, a community emerged, dominated by species common in saline soils not found along the Colorado Front Range. A single species, Alkali sacaton (Sporobolus airoides), composed nearly 50% of relative vegetation cover in control plots exhibiting a negative relationship between cover and richness. Seeded species composed approximately 92% of vegetation cover. The remaining 8% was composed of weeds from nearby areas, seed bank survivors, or mix contaminants. Three years of soil nutrient amendments, which lowered plant‐available nitrogen and phosphorus, significantly increased relative cover of seeded species to 97.5%. Fertilizer additions of phosphate enhanced abundance of introduced annual grasses (Bromus spp.) but did not significantly alter cover in control plots. Unmanipulated 4‐m² plots contained an average of 4.7 planted species and 3.9 nonplanted species during the 5‐year period, whereas plots that received grass herbicide averaged 5.4 nonplanted species. Species richness ranged from an average 6.9 species in low‐nutrient, undisturbed plots to 10.9 species in the relatively high‐nutrient, disturbed plots. The use of stockpiled soils, applied sparingly, in conjunction with a native seed mix containing species uncommon to the preexisting community generated a species‐depauperate, novel plant community that appears resistant to invasion by ruderal species.     

Persistence of native and exotic plants 10 years after prairie reconstruction

Prairie reconstructions are a critical component of preservation of the imperiled tallgrass prairie ecosystem in the Midwestern United States. Sustainability of this endeavor depends on establishment of persistent cover of planted native species and resistance to noxious weeds. The goal of this study was to understand the influence of early reconstruction practices on long‐term outcomes. Twelve replicates of three planting methods (dormant‐season broadcast, growing‐season broadcast, and growing‐season drill) and three seed mix richness levels (10, 20, or 34 species), fully crossed in a completely randomized design were planted in 2005 on nine former agricultural fields located in Iowa and Minnesota. Cover by species was estimated in 2005–2007, 2010, and 2015. In 2015, cover of planted species, native nonplanted species, and exotic species were similar to those recorded in 2010. Cover of the noxious weed Cirsium arvense had also declined by an average of 49% without herbicide from a peak in 2007 to low stable levels from 2010 to 2015. Richness of planted forbs, on the other hand, were still increasing in high‐richness broadcast treatments (e.g. 17–59% increase 2010–1015 in Minnesota). Two results in 2015 are reasons for concern: cover of planted species is only slightly over 50% in both Minnesota and Iowa, though with forbs still increasing, this may improve; and the cool‐season exotic grasses Poa pratensis and Bromus inermis are increasing at both Minnesota and Iowa sites. Control of these invasive grasses will be necessary, but care will be needed to avoid negative impacts of control methods on natives.     

Effects of Species Richness on Resident and Target Species Components in a Prairie Restoration

The ecological role of biodiversity in achieving successful restoration has been little explored in restoration ecology. We tested the prediction that we are more likely to create persistent, species‐rich plant communities by increasing the number of species sown, and, to some degree, by varying functional group representation, in experimental prairie plantings. There were 12 treatments consisting of 1‐, 2‐, 3‐, 4‐, 8‐, 12‐, and 16‐species mixtures of native perennials representing four functional groups (C₄ grasses, C₃ grasses, nitrogen‐fixing species, and late‐flowering composites) that predominate within Central Plains tallgrass prairies. In 2000, species were seeded into square plots (6 × 6 m), with five replicates per treatment, on former agricultural land. Annually, we measured total species richness and evenness, target species richness and cover, and richness and cover of resident species (i.e., those emerging from the seed bank). Both target species richness and rate of establishment of target communities were highest in the most species‐rich mixtures, but there was no additional benefit for treatments that contained more than eight species. Richness of resident species did not vary with target species richness; however, cover by resident species was lower in the higher target species treatments. Our results, indicating that establishment of species‐rich prairie mimics can be enhanced by starting with larger numbers of species at the outset, have implications for grassland restoration in which community biodiversity creation and maintenance are key goals.     

Floristic and Soil Organic Matter Changes after Five and Thirty-Five Years of Native Tallgrass Prairie Restoration

We studied two tallgrass prairies and adjacent restoration areas in northeast Kansas to analyze (1) the invasion of native tallgrass prairie species from native prairie source populations into replanted areas; (2) the establishment of planted prairie species five and 35 years after being sown; and (3) the effects of native prairie species on soil organic matter. For the majority of dominant species, composition differed statistically between sampled areas even though seed rain was available from the native tallgrass prairie remnants. Plant community differences were statistically different between each native prairie area and all respective restoration sites according to the Multiple Response Permutation Procedure. In addition, species richness was greatly reduced in replanted areas compared to adjacent native prairie remnants. Soil carbon isotope ratios indicated that the planting of warm-season grasses resulted in substantial replacement of old soil organic matter by the newly replanted grasses but that it did not create substantial increases of soil organic matter beyond replacement. The lack of accumulation reflects a nutrient-poor system (nitrogen-poor in particular), and the relative absence of native or introduced nitrogen-fixing plant species on the replanted areas may be a significant factor. It appears that restoration of the original highly diverse vegetation component of the tallgrass prairie ecosystem, even when aided by seeding and an adjacent prairie seed source, will occur on carbon- and nitrogen-depleted soils only over very long periods of time (perhaps centuries), if at all.     

Associations between avian functional guild response and regional landscape properties for conservation planning

This project facilitates a regional approach to conservation planning in Pennsylvania based on avian breeding habitat selection. The objectives were to: (1) determine the sensitivity of spatial pattern in avian diversity to changing thresholds of intra-guild species richness and (2) relate change of spatial pattern in avian diversity with landscape characteristics of bird Atlas blocks. Two state-wide spatial data layers, based on Landsat satellite data were constructed for this study. These regional landscape data were compared to Breeding Bird Atlas data from 1983 to 1989 using a geographic information system. Breeding bird data were recorded from 4928 blocks that form a grid covering Pennsylvania. Correlation analysis reduced landscape variables to 12 originally derived from forest, urban, roads, streams, and topographic data.Avian functional response guilds were used to analyze associations between breeding bird data and landscape variables. Functional response guilds were created by grouping organisms based on shared habitat preferences or behavioral characteristics. Most of the 18 avian guilds identified for this study were based on shared structural resource characteristics of preferred breeding habitat. Preferred structural resources frequently included the amount and type of forest. For this study, guilds separate resource characteristics by: (1) primary habitat (i.e. forest interior, forest edge), (2) area sensitivity (i.e. forest and grassland), (3) migratory status (i.e. resident, temperate, and tropical), and (4) nest placement (i.e. canopy nester, forest ground nester). Wetland obligate species were treated as a separate guild. Breeding Bird Atlas blocks were tabulated with respect to the number of species present from each guild. For a given guild, the number of its species in a block is termed guild-specific species richness. Sample blocks having high species richness for a given guild often occur adjacent or in close proximity forming spatial clusters in the landscape. Spatial coherence (adjacency/proximity) among the blocks forming these islands is shared guild-specific richness. Spatial clustered blocks of each guild represent areas that presumably possess required resources for members of that guild. Blocks having high intra-guild richness were evaluated with a group of block-level continuous variables using multiple logistic regressions. Logistic regression results indicate that a convincing connection exists between landscape properties of Breeding Bird Atlas blocks and habitat selection characteristics of guild members. Percent of forest cover and mean elevation were the most important habitat characteristics influencing intra-guild richness for most of the guilds tested. Concordance values from logistic regression were used to determine the strength of each guild model. Concordance, the proportion that represents the percent of correct guild richness predictions versus incorrect predictions, suggests a relationship between guild-rich clusters and habitat resources required by each guild. The highest concordance was for the exotics guild at 76.3% and the next highest was 74.8% for the grassland area sensitive guild. This signifies a 75% certainty that landscape variables could predict occurrence of a guild-rich block. Eight more guilds had concordance values greater than 65%.By using a guild approach, this study goes beyond total diversity to the more informative structural and functional diversity of guilds. Spatially clustered blocks of high species richness for a particular guild are more indicative of habitat availability and quality than would be the case for overall species richness. Clusters of blocks having high intra-guild species richness become candidate areas for conservation efforts.     

High Plant Richness in Prairie Reconstructions Support Diverse Leafhopper Communities

A primary reason for restoring plant communities is to increase biodiversity to previous levels. It is expected that restoring land with greater plant diversity will increase biodiversity at higher trophic levels, but high diversity seed mixes are expensive. In this study, we used one insect family, leafhoppers (Hemiptera: Cicadellidae) to assess the difference in leafhopper communities that result from establishing high compared with low plant richness restorations. We tested the hypotheses that: (1) the added effort of a high richness restoration leads to measurable increases in both diversity and richness of leafhoppers; and (2) that leafhopper community composition is more similar to remnant prairies in high richness than in low plant richness restorations. We found that higher plant richness led to 3‐ to 7‐fold increases in leafhopper and prairie‐dependent leafhopper diversity and richness in restorations. Leafhopper communities in high richness restorations were not more similar to remnant prairies, rather they were distinct among high and low richness restorations and prairie interior. Leafhopper richness and diversity correlated with plant richness, and leafhopper community composition differed among plant community assemblages, but not with the occurrence of single plant species. For our sites, species‐rich restorations provided better quality habitat for leafhoppers that was comparable to remnant prairie. Our results suggest that restorations with high plant species richness better support animal food webs.     

Short-Term Temporal Effects on Community Structure of Lepidoptera in Restored and Remnant Tallgrass Prairies

Understanding the degree to which species assemblages naturally vary over time will be critically important when assessing whether direct management effects or contingency is responsible for species gain or loss. In this study, we tested three predictions related to short‐term variation in prairie moth communities: (1) communities would only exhibit significant temporal variation in newly restored sites (1–3 years old); (2) prairie size and age would positively influence community reassembly, with larger, older restorations sampling a greater proportion of the regional species pool; and (3) older restorations (7–10 years old) would have yet to converge on the community composition of prairie remnants. Moths were sampled from 13 Tallgrass prairie restorations and remnants in central Iowa in 2004–2005. Repeated measures analysis of variance revealed significant effects of sampling year on moth species richness and abundance as well as on the richness of two functional groups, but difference among prairie types was only observed in 2005. Rarefaction analysis revealed that older restorations and prairie remnants supported higher species richness compared to recently planted sites, and nonmetric, multidimensional scaling ordination indicated that restorations older than 7 years were clearly converging on the species composition of remnants. These results suggest that moth communities in restorations and remnants are highly variable in time but that as restorations age, they appear to reaccumulate moth species found in prairie remnants. The long‐term persistence of a particular species assemblage within a given site, however, might be a difficult endpoint to attain in central Iowa prairies because of significant annual variation in species occurrence.     

Dominant Grasses Suppress Local Diversity in Restored Tallgrass Prairie

Warm‐season (C₄) grasses commonly dominate tallgrass prairie restorations, often at the expense of subordinate grasses and forbs that contribute most to diversity in this ecosystem. To assess whether the cover and abundance of dominant grass species constrain plant diversity, we removed 0, 50, or 100% of tillers of two dominant species (Andropogon gerardii or Panicum virgatum) in a 7‐year‐old prairie restoration. Removing 100% of the most abundant species, A. gerardii, significantly increased light availability, forb productivity, forb cover, species richness, species evenness, and species diversity. Removal of a less abundant but very common species, P. virgatum, did not significantly affect resource availability or the local plant community. We observed no effect of removal treatments on critical belowground resources, including inorganic soil N or soil moisture. Species richness was inversely correlated with total grass productivity and percent grass cover and positively correlated with light availability at the soil surface. These relationships suggest that differential species richness among removal treatments resulted from treatment induced differences in aboveground resources rather than the belowground resources. Selective removal of the dominant species A. gerardii provided an opportunity for seeded forb species to become established leading to an increase in species richness and diversity. Therefore, management practices that target reductions in cover or biomass of the dominant species may enhance diversity in established and grass‐dominated mesic grassland restorations.     

Adaptation of plant‐mycorrhizal interactions to moisture availability in prairie restoration

The strength and direction of plant response to inoculation with arbuscular mycorrhizal fungi (AM fungi) is dependent on both abiotic and biotic contexts, often generating patterns of AM fungal mediation of plant adaptation. However, knowledge of plant‐community level effects of these interactions in grassland restoration is limited. We conducted a field inoculation experiment by inoculating five plant species native to a drier prairie and five plant species native to a moister prairie with mycorrhizal fungal communities from each prairie type. Species were paired by genus or family to account for phylogenetic effects. The inoculated plants were transplanted to study plots seeded with a restoration seed mix. Plots were manipulated to create either moister or drier conditions similar to environments of the plant species and mycorrhizal communities. In both transplanted and seeded plant species, we found that only drier prairie‐range species benefited from moisture‐regime matched AM fungal inoculum. Other seeded prairie plant species demonstrated a negative response to inoculation, likely due to the earlier successional stage of these species. Additionally, nonseeded plants benefited from inoculation in different ways: native nonseeded plants had highest cover with drier prairie inoculum in drier conditions, while nonnative plants had highest cover with moister prairie‐origin inoculum. These results suggest that use of local AM fungi may be particularly important in restorations at drier sites, even at relatively small differences in moisture availability. Further, specific knowledge of relative responsiveness of seeded plant species and nonseeded plant species to AM fungal inoculation will be useful in planning restorations.     

Recovery of Native Plant Community Characteristics on a Chronosequence of Restored Prairies Seeded into Pastures in West‐Central Iowa

Restored grasslands comprise an ever‐increasing proportion of grasslands in North America and elsewhere. However, floristic studies of restored grasslands indicate that our ability to restore plant communities is limited. Our goal was to assess the effectiveness of restoration seeding for recovery of key plant community components on former exotic, cool‐season pastures using a chronosequence of six restoration sites and three nearby remnant tallgrass prairie sites in West‐Central Iowa. We assessed trends in Simpson's diversity and evenness, richness and abundance of selected native and exotic plant guilds, and mean coefficient of conservatism (mean C). Simpson's diversity and evenness and perennial invasive species abundance all declined with restoration site age. As a group, restoration sites had greater richness of native C₃ species with late phenology, but lower richness and abundance of species with early phenology relative to remnant sites. Total native richness, total native abundance (cover), mean C, and abundance of late phenology C₃ plants were similar between restoration and remnant sites. Observed declines in diversity and evenness with restoration age reflect increases in C₄ grass abundance rather than absolute decreases in the abundance of perennial C₃ species. In contrast to other studies, restoration seeding appears to have led to successful establishment of tallgrass prairie species that were likely to be included in seeding mixtures. While several floristic measures indicate convergence of restoration and remnant sites, biodiversity may be further enhanced by including early phenology species in seeding mixes in proportion to their abundance on remnant prairies.     

Prairie plant guilds: a multivariate analysis of prairie species based on ecological and morphological traits

An ecomorphological analysis of the tallgrass prairie of central North America divided representative species of the native grassland flora into eight guilds or groups of species with similar life-form, phenology, and ecology. The guilds, segregated by multivariate analysis, are: (1) warm-season graminoids with Kranz anatomy and the Hatch-Slack photosynthetic pathway (C4 grasses); (2) cool-season graminoids without Kranz anatomy, but with the common Calvin or C3 photosynthetic pathway (C3 grasses and sedges); (3) annuals and biennial forbs; (4) ephemeral spring forbs; (5) spring forbs; (6) summer/fall forbs; (7) legumes; and (8) woody shrubs. The study was based on 158 plant species indigenous to three upland prairie sites in northeastern Kansas. Each species was scored for 32 traits which fall into five broad categories: plant habit, leaf characteristics, stem structures, root structures, and reproductive traits, including phenology. A multivariate, detrended correspondence analysis sorted the 158 species into the eight principal groups or guilds. These groups were further supported by a cluster analysis and discriminant function analysis of the same data set. The discriminant function analysis determined that 94.3% of the species were correctly classified in their respective guilds, and that the guilds were statistically different. Results indicate that guild analysis offers a basis for detailed classification of grassland vegetation that is more ecologically focused than species composition, as the myriad of species (about 1,000 prairie species on the central plains of North America) vary in presence, cover, and importance with their individualistic distribution.Abbreviations C3= C3 photosynthesis - C4= C4 photosynthesis - LSD= least significant difference     

Habitat patchiness and plant species richness

The pattern of woody species richness decline with a decrease in woody vegetation cover was studied within a tallgrass prairie. The decline in species richness is highly non-linear, with a well-defined threshold below which species richness collapses. This relationship can be understood after considering information on how landscape structure changes with woody vegetation cover, and how species richness is related to landscape structure.     

No interaction between competition and herbivory in limiting introduced Cirsium vulgare rosette growth and reproduction

Both competition and herbivory have been shown to reduce plant survival, growth, and reproduction. Much less is known about whether competition and herbivory interact in determining plant performance, especially for introduced, weedy plant species in the invaded habitat. We simultaneously evaluated both the main and interactive effects of plant neighbors and insect herbivory on rosette growth and seed reproduction in the year of flowering for Cirsium vulgare (bull thistle, spear thistle), an introduced Eurasian species, in tallgrass prairie in 2 years. Effects of insect herbivory were strong and consistent in both years, causing reduced plant growth and seed production, whereas the effects of competition with established vegetation were weak. The amount of herbivore damage inflicted on rosettes did not depend on the presence of neighbor plants. We also found no interaction between competition and herbivory on key parameters of plant growth and fitness. The results of this study contradict the hypothesis that competitive context interacts with insect herbivory in limiting the invasiveness of this introduced thistle. Further, the results provide additional, experimental evidence that high levels of herbivory on established rosettes by native insects exert significant biotic resistance to the invasiveness of C. vulgare in western tallgrass prairie.     

Mycorrhizal suppression alters plant productivity and forb establishment in a grass-dominated prairie restoration

A fundamental goal of restoration is the re-establishment of plant diversity representative of native vegetation. However, many prairie restorations or Conservation Reserve Program sites have been seeded with warm-season grasses, leading to grass-dominated, low-diversity restorations not representative of native grasslands. These dominant grasses are strongly mycotrophic, while many subordinate forb species appear to be less dependent on mycorrhizal symbiosis. Therefore, manipulating arbuscular mycorrhizal fungi (AMF) may be useful in promoting establishment and growth of forb species in grass-dominated prairie restorations. To assess the potential role of mycorrhizae in affecting the productivity and community composition of restored tallgrass prairie, we conducted a 4-year field experiment on an 8-year-old grassland restoration at the Konza Prairie in northeastern Kansas, USA. At the initiation of our study, seeds of 12 forb species varying in degree of mycorrhizal dependence were added to established grass-dominated plots. Replicate plots were treated bi-weekly with a soil drench of fungicide (Topsin-M^®) over four growing seasons and compared to non-treated control plots to assess the role of AMF in affecting plant species composition, productivity, leaf tissue quality, and diversity in restored tallgrass prairie. Topsin applications successfully reduced mycorrhizal colonization of grass roots to approximately 60–80% relative to roots in control plots. Four years of mycorrhizal suppression reduced productivity of the dominant grasses and increased plant species richness and diversity. These results highlight the importance of mycorrhizae as mediators of plant productivity and community dynamics in restored tallgrass prairie and indicate that temporarily suppressing AMF decreases productivity of the dominant C4 grasses and allows for establishment of seeded forb species.     

In tallgrass prairie restorations,relatedness influences neighborhood-scale plant invasion while resource availability influences site-scale invasion

Attempting to control invasive plant species in tallgrass prairie restorations is time-consuming and costly, making improved approaches for predicting and reducing invasion imperative. Both biotic and abiotic factors mediate plant invasions, and can potentially be used by restoration managers to reduce invasion rates. Biotic factors such as plant species richness and phylogenetic diversity of the native community may impact invasion. Relatedness of invading species to those in recipient communities has also been shown to influence invasion success. However, the direction of this influence is variable, reflecting Darwin’s Naturalization Conundrum. Abiotic factors such as fire regime and soil factors may impact invasion by selecting against invasive species or indicating suitable habitats for them. We surveyed 17 tallgrass prairie restorations in Illinois, USA, to investigate the effects of biotic and abiotic factors on invasion by non-native plant species at two different scales. We predicted we would find support for Darwin’s Naturalization Hypothesis at the plot (neighborhood) scale with invasion by distantly related species, and find support for the Pre-adaptation Hypothesis at the site scale. We hypothesized that biotic factors would exert more influence at the neighborhood scale, while abiotic factors would be more influential at a coarser site scale. Contrary to our expectations, at the neighborhood scale we found that closely related invasive species are more likely to invade, supporting the Pre-adaptation Hypothesis. We found that native species richness and age of restoration were negatively correlated with invasion. At the site scale, soil organic matter [SOM] concentrations and heterogeneity in SOM were positively associated with the number of invasive species while pH heterogeneity was negatively associated. Restoration practitioners may be able to reduce plant invasions by increasing native species richness, and non-native species most closely related to the resident community should potentially be prioritized as those most likely to be highly invasive.     

Can the Results of Biodiversity-Ecosystem Productivity Studies Be Translated to Bioenergy Production?

Biodiversity experiments show that increases in plant diversity can lead to greater biomass production, and some researchers suggest that high diversity plantings should be used for bioenergy production. However, many methods used in past biodiversity experiments are impractical for bioenergy plantings. For example, biodiversity experiments often use intensive management such as hand weeding to maintain low diversity plantings and exclude unplanted species, but this would not be done for bioenergy plantings. Also, biodiversity experiments generally use high seeding densities that would be too expensive for bioenergy plantings. Here we report the effects of biodiversity on biomass production from two studies of more realistic bioenergy crop plantings in southern Michigan, USA. One study involved comparing production between switchgrass (Panicum virgatum) monocultures and species-rich prairie plantings on private farm fields that were managed similarly to bioenergy plantings. The other study was an experiment where switchgrass was planted in monoculture and in combination with increasingly species-rich native prairie mixtures. Overall, we found that bioenergy plantings with higher species richness did not produce more biomass than switchgrass monocultures. The lack of a positive relationship between planted species richness and production in our studies may be due to several factors. Non-planted species (weeds) were not removed from our studies and these non-planted species may have competed with planted species and also prevented realized species richness from equaling planted species richness. Also, we found that low seeding density of individual species limited the biomass production of these individual species. Production in future bioenergy plantings with high species richness may be increased by using a high density of inexpensive seed from switchgrass and other highly productive species, and future efforts to translate the results of biodiversity experiments to bioenergy plantings should consider the role of seeding density.     

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.     

Experimental analysis of patch dynamics and community heterogeneity in tallgrass prairie

Effects of fire and small-scale soil disturbances on species richness, community heterogeneity, and microsuccession were investigated in a central Oklahoma tallgrass prairie. In the fall of 1985, 0.2 m² soil disturbances were created on burned and unburned tallgrass prairie. Vegetation on and off disturbances was sampled at monthly intervals over two growing seasons. During the first growing season, the cover of forbs and annuals, and species richness were significantly greater on versus off disturbances, but these differences did not persist through the second year. The variation in species composition among disturbed plots (heterogeneity) was significantly greater compared to undisturbed areas throughout the study. Fire had no consistent effect on richness and heterogeneity of vegetation on soil disturbances but fire reduced heterogeneity on undisturbed vegetation. Rate of succession, based on an increase in cumulative cover of perennial grasses over time, did not differ among treatments during the first growing season. During the second year, rate of succession was significantly greater on burned soil disturbances compared to unburned soil disturbances. These results suggest that while small-scale soil disturbances have primarily short-lived effects on grassland community structure, disturbances do help to maintain spatial and temporal variation in tallgrass prairie communities. Unlike in undisturbed vegetation, however, species richness and heterogeneity on soil disturbances were little effected by fire, but the rate of colonization onto disturbances appeared to be enhanced by fire.