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.     

Productivity and Subordinate Species Response to Dominant Grass Species and Seed Source during Restoration

Grasses can be important regulators of species diversity and ecosystem processes in prairie systems. Although C₄ grasses are usually assumed to be ecologically similar because they are in the same functional group, there may be important differences among species or between seed sources that could impact restorations. I tested whether C₄ grass species identity, seed source, or grass species richness scales to influence aboveground net primary productivity (ANPP), resistance to weed invasion, or establishment of subordinate prairie species during restoration. Plots in western Iowa, United States, were planted with equal‐sized transplants of one of five common grass species (Panicum virgatum L., Sorghastrum nutans (L.) Nash, Andropogon gerardii Vitman, Schizachyrium scoparium (Michx.) Nash, and Bouteloua curtipendula (Michx.) Torrey) either from local seed or from cultivar seed sources. These plots were compared to plots containing all five species in mixture and to nonplanted plots. Differences in ANPP were found among species but not between cultivars and noncultivars or between monocultures and mixtures. Panicum virgatum, S. nutans, and S. scoparium were more productive than A. gerardii and B. curtipendula. Weed invasion was much higher when plots were not planted with grasses. Schizachyrium scoparium allowed greater establishment of subordinant prairie species than all other focal grass species. There were two separate mechanisms by which grasses suppressed prairie species establishment either (1) by growing tall and capturing light or (2) by quickly filling in bare space by spreading horizontally through rhizome growth in short species. These results suggest that high ANPP can be found with noncultivar plantings during the first 2 years after planting and that subordinate species establishment is most likely when shorter bunchgrasses such as S. scoparium are dominant.     

Tallgrass prairie restoration: implications of increased atmospheric nitrogen deposition when site preparation minimizes adventive grasses

Site preparation designed to exhaust the soil seedbank of adventive species can improve the success of tallgrass prairie restoration. Despite these efforts, increased rates of atmospheric nitrogen (N) deposition over the next century could potentially promote the growth of nitrophilic, adventive species in tallgrass restoration projects. We used a field experiment to examine how N addition affected species composition and plant productivity over the first 3 years of a tallgrass prairie restoration that was preceded by the planting of glyphosate‐resistant crops and multiple applications of glyphosate to exhaust the pre‐existing seedbank. We predicted that N addition would increase the percent cover of adventive plant species not included in the original seeding. Contrary to our prediction, only the cover of native species increased with N addition; native non‐leguminous forbs increased substantially, with Conyza canadensis (a weedy native species not part of the restoration seed mix) exploiting the combination of high N and bare ground in the first year, and non‐leguminous forbs (in particular Monarda fistulosa) and native C₃ grasses, all of which were seeded, increasing with N addition by the third year. Native legumes was the only functional group that exhibited lower cover in N addition plots than in control plots. There was no significant response by native C₄ grasses to N addition, and adventive grasses remained mostly absent from the plots. Overall, our results suggest that site pre‐treatment with herbicide may continue to be effective in minimizing adventive grasses in restored tallgrass prairie, despite future increases in atmospheric N deposition.     

Contrasting above‐ and belowground sensitivity of three Great Plains grasslands to altered rainfall regimes

Intensification of the global hydrological cycle with atmospheric warming is expected to increase interannual variation in precipitation amount and the frequency of extreme precipitation events. Although studies in grasslands have shown sensitivity of aboveground net primary productivity (ANPP) to both precipitation amount and event size, we lack equivalent knowledge for responses of belowground net primary productivity (BNPP) and NPP. We conducted a 2‐year experiment in three US Great Plains grasslands – the C₄‐dominated shortgrass prairie (SGP; low ANPP) and tallgrass prairie (TGP; high ANPP), and the C₃‐dominated northern mixed grass prairie (NMP; intermediate ANPP) – to test three predictions: (i) both ANPP and BNPP responses to increased precipitation amount would vary inversely with mean annual precipitation (MAP) and site productivity; (ii) increased numbers of extreme rainfall events during high‐rainfall years would affect high and low MAP sites differently; and (iii) responses belowground would mirror those aboveground. We increased growing season precipitation by as much as 50% by augmenting natural rainfall via (i) many (11–13) small or (ii) fewer (3–5) large watering events, with the latter coinciding with naturally occurring large storms. Both ANPP and BNPP increased with water addition in the two C₄ grasslands, with greater ANPP sensitivity in TGP, but greater BNPP and NPP sensitivity in SGP. ANPP and BNPP did not respond to any rainfall manipulations in the C₃‐dominated NMP. Consistent with previous studies, fewer larger (extreme) rainfall events increased ANPP relative to many small events in SGP, but event size had no effect in TGP. Neither system responded consistently above‐ and belowground to event size; consequently, total NPP was insensitive to event size. The diversity of responses observed in these three grassland types underscores the challenge of predicting responses relevant to C cycling to forecast changes in precipitation regimes even within relatively homogeneous biomes such as grasslands.     

The effects of fire frequency and grazing on tallgrass prairie productivity and plant composition are mediated through bud bank demography

Periodic fire, grazing, and a variable climate are considered the most important drivers of tallgrass prairie ecosystems, having large impacts on the component species and on ecosystem structure and function. We used long-term experiments at Konza Prairie Biological Station to explore the underlying demographic mechanisms responsible for tallgrass prairie responses to two key ecological drivers: fire and grazing. Our data indicate that belowground bud banks (populations of meristems associated with rhizomes or other perennating organs) mediate tallgrass prairie plant response. Fire and grazing altered rates of belowground bud natality, tiller emergence from the bud bank, and both short-term (fire cycle) and long-term (>15 year) changes in bud bank density. Annual burning increased grass bud banks by 25% and decreased forb bud banks by 125% compared to burning every 4 years. Grazing increased the rate of emergence from the grass bud bank resulting in increased grass stem densities while decreasing grass bud banks compared to ungrazed prairie. By contrast, grazing increased both bud and stem density of forbs in annually burned prairie but grazing had no effect on forb bud or stem density in the 4-year burn frequency treatment. Lastly, the size of the reserve grass bud bank is an excellent predictor of long-term ANPP in tallgrass prairie and also of short-term interannual variation in ANPP associated with fire cycles, supporting our hypothesis that ANPP is strongly regulated by belowground demographic processes. Meristem limitation due to management practices such as different fire frequencies or grazing regimes may constrain tallgrass prairie responses to interannual changes in resource availability. An important consequence is that grasslands with a large bud bank may be the most responsive to future climatic change or other global change phenomena such as nutrient enrichment, and may be most resistant to exotic species invasions.     

Plant–microbe interactions change along a tallgrass prairie restoration chronosequence

Soil microbial communities are critical in determining the performance and density of species in plant communities. However, their role in regulating the success of restorations is much less clear. This study assessed the ability of soil microbial communities to regulate the growth and performance of two potentially dominant grasses and two common forbs in tallgrass prairie restorations. Specifically, we examined the effects of soil microbial communities along a restoration chronosequence from agricultural fields to remnant prairies using experimentally inoculated soils. The two grass species, Andropogon gerardii and Sorghastrum nutans, grew best with the agricultural inoculates and experienced a decline in performance in later stages of the chronosequence, indicating that the microbial community shifted from being beneficial to grasses in the early stages to inhibiting grasses in the later stages of restoration. Growth of the forb, Silphium terebinthinaceum, varied little with inoculation or position along the restoration chronosequence. Growth of Baptisia leucantha, a legume, appeared limited by nodule formation in agricultural soils, peaked in young restoration soils along with nodule formation, but decreased in older soils as the microbial community became more antagonistic. Overall, negative feedbacks tended to be less important early in restoration, but appeared important in remnant and older restored prairies. Our results provide evidence that it may be advantageous for management practices to take negative soil feedbacks into consideration when trying to recreate the diversity of tallgrass prairies.     

The Role of Seed and Vegetative Reproduction in Plant Recruitment and Demography in Tallgrass Prairie

Recruitment, establishment and survivorship of seed- and vegetatively-derived shoots were quantified biweekly in annually burned and infrequently burned tallgrass prairie to investigate the contributions of seed and vegetative reproduction to the maintenance and dynamics of tallgrass prairie plant populations, the demography of seedlings and ramets, and the influence of fire on the demography of grasses and forbs. Clonally produced grass and forb ramets comprised >99%of all established shoots present at the end of the growing season, whereas established seedlings accounted for <1%,emphasizing the rarity of successful seedling establishment and the importance of vegetative reproduction in driving the annual regeneration and dynamics of aboveground plant populations in tallgrass prairie. Most recruitment from vegetative reproduction occurred early in the growing season and was higher in annually burned than infrequently burned sites, although low levels of new stem recruitment occurred continuously throughout the growing season. Peak recruitment on annually burned prairie coincided with peak recruitment of the dominant C₄ grasses Andropogon gerardii and Sorghastrum nutans prior to prescribed spring fire, with a second peak in recruitment occurring following fire. On infrequently burned prairie, grass and forb recruitment was highest in early April and declined steadily through May. The naturalized C₃ grass, Poa pratensis, was responsible for most of the early recruitment on unburned sites, whereas A. gerardii contributed most to recruitment later in May. Infrequently burned prairie was dominated by these two grasses and contained a larger forb component than annually burned prairie. The principal demographic effect of fire was on ramet natality rather than mortality. Fire regime, plant functional group, or timing of cohort emergence before or after fire did not affect ramet survivorship. C₄ grass shoots that emerged early and were damaged by fire showed similar survivorship patterns to tillers that emerged after fire. Differences in species composition between annually burned and infrequently burned prairie are driven by fire effects on vegetative reproduction and appear to be related principally to the effect of fire and detritus accumulation on the development of belowground vegetative meristems of C₄ grasses and their emergence dynamics.     

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.     

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.     

A Test of Diversity–Productivity Models in Natural,Degraded, and Restored Wet Prairies

Conceptual restoration models depict strong correlations between structure and function, with both decreasing as an ecosystem is degraded and increasing during restoration. We evaluated the “linear” and “asymptotic” models by measuring diversity and annual net primary productivity (NPP) within four states of a southern Wisconsin floodplain: a remnant (unplowed) wet prairie, two degraded sites (soybean field and invaded prairie), and a restored prairie. Neither model fit our data for aboveground (ANPP), belowground (BNPP), or total (TNPP) productivity. ANPP declined as species richness increased (r = 0.998, df = 2), with highest values for soybeans (1,024 g/m²; two species in 30 0.25‐m² plots) and invaded prairie (937 g/m²; nine species, 99% cover of Phalaris arundinacea), intermediate for restored prairie (712 g/m²; 28 species), and lowest for diverse remnant prairie (571 g/m²; 36 species). In contrast, BNPP was lowest for soybeans (225 g/m²) and highest for remnant prairie (571 g/m²). TNPP in restored prairie (990 g/m²) matched that of the remnant (1,147 g/m²) within 7 years, but root:shoot NPP ratios were quite different (0.39 and 0.99, respectively). Overall, results suggest that the relationship between species diversity and productivity can differ with the component measured (ANPP, BNPP, or TNPP) and that diversity does not ensure high productivity. Because measuring ANPP does not fully test ecosystem‐function theory, we recommend assessing BNPP and additional ecosystem processes in future attempts to determine whether adding species will restore more function to degraded ecosystems.     

The role of hemiparasitic plants: influencing tallgrass prairie quality,diversity, and structure

Wood betony, Orobanchaceae (Pedicularis canadensis) and bastard toadflax, Santalaceae (Comandra umbellata) are two root‐hemiparasitic plant species found in tallgrass prairie communities. Natural resource managers are interested in utilizing these species as “pseudograzers” in grasslands to reduce competitively dominant grasses and thereby increase ecological diversity and quality in prairie restorations and urban plantings. We performed an observational field study at 5 tallgrass prairie sites to investigate the association of hemiparasite abundance with metrics of phylogenetic and ecological diversity, as well as floristic quality. Although no reduction in C₄ grasses was detected, there was a significant association between hemiparasite abundance and increased floristic quality at all 5 sites. Hemiparasite abundance and species richness were positively correlated at one restoration site. In a greenhouse mesocosm experiment, we investigated response to parasitism by P. canadensis in 6 species representing different plant functional groups of the tallgrass prairie. The annual legume partridge pea, Fabaceae (Chamaecrista fasciculata) had the greatest significant dry biomass reduction among 6 host species, but the C₄ grass big bluestem, Poaceae (Andropogon gerardii) had significantly greater aboveground biomass when grown with the hemiparasite. Overall, host species biomass as a total community was significantly reduced in mesocosms, consistent with other investigations that demonstrate influence on community structure by hemiparasitic plant species. Although hemiparasites were not acting as pseudograzers, they have the potential to influence community structure in grassland restorations and remnants.     

Net primary productivity and rain‐use efficiency as affected by warming,altered precipitation,and clipping in a mixed‐grass prairie

Grassland productivity in response to climate change and land use is a global concern. In order to explore the effects of climate change and land use on net primary productivity (NPP), NPP partitioning [f_BNPP, defined as the fraction of belowground NPP (BNPP) to NPP], and rain‐use efficiency (RUE) of NPP, we conducted a field experiment with warming (+3 °C), altered precipitation (double and half), and annual clipping in a mixed‐grass prairie in Oklahoma, USA since July, 2009. Across the years, warming significantly increased BNPP, f_BNPP, and RUE_BNPP by an average of 11.6%, 2.8%, and 6.6%, respectively. This indicates that BNPP was more sensitive to warming than aboveground NPP (ANPP) since warming did not change ANPP and RUE_ANPP much. Double precipitation stimulated ANPP, BNPP, and NPP but suppressed RUE_ANPP, RUE_BNPP, and RUE_NPP while half precipitation decreased ANPP, BNPP, and NPP but increased RUE_ANPP, RUE_BNPP, and RUE_NPP. Clipping interacted with altered precipitation in impacting RUE_ANPP, RUE_BNPP, and RUE_NPP, suggesting land use could confound the effects of precipitation changes on ecosystem processes. Soil moisture was found to be a main factor in regulating variation in ANPP, BNPP, and NPP while soil temperature was the dominant factor influencing f_BNPP. These findings suggest that BNPP is critical point to future research. Additionally, results from single‐factor manipulative experiments should be treated with caution due to the non‐additive interactive effects of warming with altered precipitation and land use (clipping).     

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.     

Effects of Soil Carbon Amendment on Nitrogen Availability and Plant Growth in an Experimental Tallgrass Prairie Restoration

Restoration of tallgrass prairie on former agricultural land is often impeded by failure to establish a diverse native species assemblage and by difficulties with nonprairie, exotic species. High levels of available soil nitrogen (N) on such sites may favor fast‐growing exotics at the expense of more slowly growing prairie species characteristic of low‐N soils. We tested whether reducing N availability through soil carbon (C) amendments could be a useful tool in facilitating successful tallgrass prairie restoration. We added 6 kg/m² hardwood sawdust to experimental plots on an abandoned agricultural field in the Sandusky Plains of central Ohio, United States, increasing soil C by 67% in the upper 15 cm. This C amendment caused a 94% reduction in annual net N mineralization and a 27% increase in soil moisture but had no effect on total N or pH. Overall, plant mass after one growing season was reduced by 64% on amended compared with unamended soil, but this effect was less for prairie forbs (?34%) than for prairie grasses (?67%) or exotics (?62%). After the second growing season, only exotics responded significantly to the soil C amendment, with a 40% reduction in mass. The N concentration of green‐leaf tissue and of senescent leaf litter was also reduced by the soil C treatment in most cases. We conclude that soil C amendment imparts several immediate benefits for tallgrass prairie restoration––notably reduced N availability, slower plant growth, and lower competition from exotic species.     

Growth and Arbuscular Mycorrhizal Fungal Colonization of Two Prairie Grasses Grown in Soil from Restorations of Three Ages

I compared growth and arbuscular mycorrhizal fungal (AMF) colonization of two prairie grasses (Wild rye [Elymus canadensis] and Little bluestem [Schizachyrium scoparium]), an early‐ and a late‐dominating species in prairie restorations, respectively, grown in soil from restored prairies of differing age, soil characteristics, and site history. There were no consistent patterns between restoration age and soil inorganic nutrients or organic matter. The oldest restoration site had higher soil mycorrhizal inoculum potential (MIP) than 2‐ and 12‐year‐old restorations. However, MIP did not translate into actual colonization for two species grown in soils from the three restorations, nor did MIP relate to phosphorus availability. There were significant differences in root mass and colonization among Wild rye plants but not among Little bluestem plants grown in soils from the three restorations. Wild rye grown in 2‐year‐old restoration soil had significantly higher AMF colonization than when it was grown in soils from the 12‐ and 17‐year‐old restorations. Wild rye grown in 2‐year‐old restoration soil also had higher colonization than Little bluestem grown in 2‐ and 12‐year‐old restoration soils. Little bluestem had no significant correlations between shoot biomass, root biomass or colonization, and concentrations of soil P, total N, or N:P. However, for Wild rye, total soil N was positively correlated with root mass and negatively correlated with colonization, suggesting that in this species, mycorrhizae may affect N availability. Collectively, these results suggest that soil properties unrelated to restoration age were important in determining differences in growth and AMF colonization of two species of prairie grasses.     

Diminishing Spatial Heterogeneity in Soil Organic Matter across a Prairie Restoration Chronosequence

Habitat restoration resulting in changes in plant community composition or species dominance can affect the spatial pattern and variability of soil nutrients. Questions about how these changes in soil spatial heterogeneity develop over time at restoration sites, however, remain unaddressed. In this study, a geostatistical approach was used to quantify changes over time in the spatial heterogeneity of soil organic matter (SOM) across a 26‐year chronosequence of tallgrass prairie restoration sites at FermiLab, outside of Chicago, Illinois. We used total soil N and C as an index of the quantity of SOM. We also examined changes in C:N ratio, which can influence the turnover of SOM. Specifically, the spatial structure of total N, total C, and C:N ratio in the top 10 cm of soil was quantified at a macroscale (minimum spacing of 1.5 m) and a microscale (minimum spacing of 0.2 m). The magnitude of spatial heterogeneity (MSH) was characterized as the proportion of total sample variation explained by spatially structured variation. At the macroscale, the MSH for total N decreased with time since restoration (r²= 0.99, p < 0.001). The decrease in spatial heterogeneity over time corresponded with a significant increase in the dominance of the C₄ grasses. At the microscale, there was significant spatial structure for total N at the 4‐year‐old, 16‐year‐old, and 26‐year‐old sites, and significant spatial structure for total C at the 16‐year‐old and 26‐year‐old sites. These results suggest that an increase in dominance of C₄ grasses across the chronosequence is homogenizing organic matter variability at the field scale while creating fine‐scale patterns associated with the spacing of vegetation. Areas of higher soil moisture were associated with higher soil N and C at the two oldest restoration sites and at the native prairie site, potentially suggesting patches of increased belowground productivity in areas of higher soil moisture. This study is one of the first to report significant changes over time in the spatial structure of organic matter in response to successional changes initiated by restoration.     

Resource manipulation effects on net primary production,biomass allocation and rain-use efficiency of two semiarid grassland sites in Inner Mongolia,China

Productivity of semiarid grasslands is affected by soil water and nutrient availability, with water controlling net primary production under dry conditions and soil nutrients constraining biomass production under wet conditions. In order to investigate limitations on plants by the response of root–shoot biomass allocation to water and nitrogen (N) availability, a field experiment, on restoration plots with rainfed, unfertilized control plots, fertilized plots receiving N (25 kg urea-N ha⁻¹) and water (irrigation simulating a wet season), was conducted at two sites with different grazing histories: moderate (MG) and heavy (HG) grazing. Irrigation and N addition had no effect on belowground biomass. Irrigation increased aboveground (ANPP) and belowground net primary production (BNPP) and rain-use efficiency based on ANPP (RUE_ANPP), whereas N addition on rainfed plots had no effect on any of the measured parameters. N fertilizer application on irrigated plots increased ANPP and RUE_ANPP and reduced the root fraction (RF: root dry matter/total dry matter), resulting in smaller N effects on total net primary production (NPP) and rain-use efficiency based on NPP. This suggests that BNPP should be included in evaluating ecosystem responses to resource availability from the whole-plant perspective. N effects on all measured parameters were similar on both sites. However, site HG responded to irrigation with higher ANPP and a lower RF when compared to site MG, indicating that species composition had a pronounced effect on carbon allocation pattern due to below- and aboveground niche complementarity.     

Genetic Structure of Remnant Populations and Cultivars of Switchgrass (Panicum virgatum) in the Context of Prairie Conservation and Restoration

Switchgrass (Panicum virgatum L.) is a dominant, perennial C₄ grass of North American tallgrass prairies with cultivars that are widely used in grassland restoration, pastures, and landscaping. However, these cultivars may be genetically dissimilar to small, remnant populations, raising concerns about altered genetic composition of native populations through gene flow. To address this issue on a local scale in Ohio and Illinois, we used microsatellite markers to characterize genetic diversity and differentiation of 10 remnant prairie populations (5 in each state) and 8 common cultivars. The bulk of genetic variation was found to reside within rather than among wild populations, consistent with the outcrossing breeding system of switchgrass. Genetic diversity was similar among the remnant populations despite large differences in area (approximately 2–2,590 ha), highlighting the importance of small native populations as reservoirs of variation and potential seed sources for prairie restoration. Cultivars generally had similar levels of variation to the wild populations, but we found clear genetic dissimilarity between wild and cultivated gene pools (especially for Kanlow, but also Trailblazer, Blackwell, Dacotah, Summer, and Sunburst cultivars). This suggests that using cultivars in local prairie restoration efforts may alter the genetic composition of wild populations. Whether such changes are deemed as negative depends on the cultivar under consideration and specific conservation goals for preserving native switchgrass populations. Patterns of genetic variation in remnant prairie populations and potential cultivar sources can be used to develop guidelines for restoration as well as future planting of cultivars for biofuels.     

CO2‐caused change in plant species composition rivals the shift in vegetation between mid‐grass and tallgrass prairies

Atmospheric CO₂ enrichment usually changes the relative contributions of plant species to biomass production of grasslands, but the types of species favored and mechanisms by which change is mediated differ among ecosystems. We measured changes in the contributions of C₃ perennial forbs and C₄ grasses to aboveground biomass production of tallgrass prairie assemblages grown along a field CO₂ gradient (250–500 μmol mol^?1) in central Texas USA. Vegetation was grown on three soil types and irrigated each season with water equivalent to the growing season mean of precipitation for the area. We predicted that CO₂ enrichment would increase the forb contribution to community production, and favor tall‐grasses over mid‐grasses by increasing soil water content and reducing the frequency with which soil water fell below a limitation threshold. CO₂ enrichment favored forbs over grasses on only one of three soil types, a Mollisol. The grass fraction of production increased dramatically across the CO₂ gradient on all soils. Contribution of the tall‐grass Sorghastrum nutans to production increased at elevated CO₂ on the two most coarse‐textured of the soils studied, a clay Mollisol and sandy Alfisol. The CO₂‐caused increase in Sorghastrum was accompanied by an offsetting decline in production of the mid‐grass Bouteloua curtipendula. Increased CO₂ favored the tall‐grass over mid‐grass by increasing soil water content and apparently intensifying competition for light or other resources (Mollisol) or reducing the frequency with which soil water dipped below threshold levels (Alfisol). An increase in CO₂ of 250 μmol mol^?1 above the pre‐industrial level thus led to a shift in the relative production of established species that is similar in magnitude to differences observed between mid‐grass and tallgrass prairies along a precipitation gradient in the central USA. By reducing water limitation to plants, atmospheric CO₂ enrichment may alter the composition and even structure of grassland vegetation.     

Belowground bud production is linked to population establishment in Sorghastrum nutans (Poaceae)

Belowground bud production mediates many responses of tallgrass prairie grasses to temporal environmental variation, and clonal spread contributes to dominance in successional plant communities. However, we know little about the importance of these regenerative traits for vegetation establishment in grassland restoration and about their regional variability. Further, geographic seed or transplant sources affect plant performance. We investigated the effects of seed source on the number of belowground buds per tiller, clonal spread (total rhizome length adjusted for plant biomass), and population establishment in the native C₄ grass Sorghastrum nutans sown into tallgrass prairie restorations using reciprocal common gardens in three sites (in Nebraska, Kansas, and Oklahoma). We characterized buds per tiller and clonal spread at the plant level and related both traits to population establishment at the restoration plot level. Plant-level bud production significantly differed among seed sources and common garden sites, and was positively associated with plot-level establishment (density and canopy cover), but plant-level clonal spread did not differ among seed sources or common garden sites and was not associated with the plot-level establishment. Our results are among the first to link bud production, which varied among source populations, to differences in plant population establishment. These results suggest that we should more often consider differences among sources in traits closely tied to post-seeding population establishment. The use of sources that possess such traits may facilitate successful establishment in restoration settings.