Soil water partitioning contributes to species coexistence in tallgrass prairie

The majority of tallgrass prairie root biomass is located in the upper soil layers (0–25 cm), but species differences exist in reliance on soil water at varying depths. These differences have led to the hypothesis that resource partitioning belowground facilitates species co‐existence in this mesic grassland. To determine if plant water relations can be linked to soil water partitioning as a potential mechanism allowing C₃ species to persist among the more dominant C₄ grasses, we measured differences in the source of water‐use using the isotopic signature of xylem water, volumetric soil water content at 4 depths, and leaf water potentials. Data were collected for seven species representing C₄ grasses, C₃ forbs and C₃ shrubs over three growing seasons at the Konza Prairie (Kansas, USA) to encompass a range of natural climatic conditions. C₄ grasses relied on shallow soil water (5 cm) across the growing season and had midday leaf water potentials that were highly correlated with shallow soil water regardless of soil water availability at other portions of the soil profile (20, 40 and 90 cm). In contrast, C₃ species only used shallow soil water when plentiful at this depth; these species increased their dependence on soil water from greater depths as the upper soil layers dried. Structural equation models describing plant water relations were very similar for the three C₄ species, whereas a unique set of models and drivers were identified for each of the C₃ species. These results support soil water partitioning as a mechanism for species coexistence, as C₄ species in this grassland have relatively consistent dependence on water in shallow soil layers, whereas C₃ species show niche differentiation in water use strategies to avoid competition with C₄ grasses for water in shallow soil layers when this resource is limiting and leaf water stress is high.     

Root plasticity of native and invasive Great Basin species in response to soil nitrogen heterogeneity

Soil nutrients are heterogeneously distributed in natural systems. While many species respond to this heterogeneity through root system plasticity, little is known about how the magnitude of these responses may vary between native and invasive species. We quantified root morphological and physiological plasticity of co-occurring native and invasive Great Basin species in response to soil nitrogen heterogeneity and determined if trade-offs exist between these foraging responses and species relative growth rate or root system biomass. The nine study species included three perennial bunchgrasses, three perennial forbs, and three invasive perennial forbs. The plants were grown in large pots outdoors. Once a week for 4 weeks equal amounts of ¹⁵NH₄ ¹⁵NO₃ were distributed in the soil either evenly through the soil profile, in four patches, or in two patches. All species acquired more N in patches compared to when N was applied evenly through the soil profile. None of the species increased root length density in enriched patches compared to control patches but all species increased root N uptake rate in enriched patches. There was a positive relationship between N uptake rate, relative growth rate, and root system biomass. Path analysis indicated that these positive interrelationships among traits could provide one explanation of how invasive forbs were able to capture 2 and 15-fold more N from enriched patches compared to the native grasses and forbs, respectively. Results from this pot study suggest that plant traits related to nutrient capture in heterogeneous soil environments may be positively correlated which could potentially promote size-asymmetric competition belowground and facilitate the spread of invasive species. However, field experiments with plants in different neighbor environments ultimately are needed to determine if these positive relationships among traits influence competitive ability and invader success.     

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.     

The dependence of root system properties on root system biomass of 10 North American grassland species

Dependence of the properties of root systems on the size of the root system may alter conclusions about differences in plant growth in different environments and among species. To determine whether important root system properties changed as root systems aged and accumulated biomass, we measured three important properties of fine roots (tissue density, diameter, and C:N) and three biomass ratios (root:shoot, fine:coarse, and shallow:deep) of monocultures of 10 North American grassland species five times during their second and third years of growth. With increasing belowground biomass, root tissue density increased and diameter decreased. This may reflect cortical loss associated with the aging of roots. For non-legumes, fine root C:N decreased with increasing root biomass, associated with decreases in soil solution NO^{3 –} concentrations. No changes in fine root C:N were detected with increasing belowground biomass for the two legumes we studied. Among all 10 species, there were generally no changes in the relative amounts of biomass in coarse and fine roots, root:shoot, or the depth placement of fine roots in the soil profile as belowground biomass increased. Though further research is needed to separate the influence of root system size, age of the roots, and changes in nutrient availability, these factors will need to be considered when comparing root functional traits among species and treatments.     

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.     

Aboveground productivity and root–shoot allocation differ between native and introduced grass species

Plant species in grasslands are often separated into groups (C₄ and C₃ grasses, and forbs) with presumed links to ecosystem functioning. Each of these in turn can be separated into native and introduced (i.e., exotic) species. Although numerous studies have compared plant traits between the traditional groups of grasses and forbs, fewer have compared native versus introduced species. Introduced grass species, which were often introduced to prevent erosion or to improve grazing opportunities, have become common or even dominant species in grasslands. By virtue of their abundances, introduced species may alter ecosystems if they differ from natives in growth and allocation patterns. Introduced grasses were probably selected nonrandomly from the source population for forage (aboveground) productivity. Based on this expectation, aboveground production is predicted to be greater and root mass fraction to be smaller in introduced than native species. We compared root and shoot distribution and tissue quality between introduced and native C₄ grass species in the Blackland Prairie region of Central Texas, USA, and then compared differences to the more well-studied divergence between C₄ grasses and forbs. Comparisons were made in experimental monocultures planted with equal-sized transplants on a common soil type and at the same density. Aboveground productivity and C:N ratios were higher, on average, in native grasses than in native forbs, as expected. Native and introduced grasses had comparable amounts of shallow root biomass and tissue C:N ratios. However, aboveground productivity and total N were lower and deep root biomass and root mass fraction were greater in native than introduced grasses. These differences in average biomass distribution and N could be important to ecosystems in cases where native and introduced grasses have been exchanged. Our results indicate that native–introduced status may be important when interpreting species effects on grassland processes like productivity and plant N accumulation.     

Maintenance of leaf N controls the photosynthetic CO2 response of grassland species exposed to 9 years of free‐air CO2 enrichment

Determining underlying physiological patterns governing plant productivity and diversity in grasslands are critical to evaluate species responses to future environmental conditions of elevated CO₂ and nitrogen (N) deposition. In a 9‐year experiment, N was added to monocultures of seven C₃ grassland species exposed to elevated atmospheric CO₂ (560 μmol CO₂ mol^?1) to evaluate how N addition affects CO₂ responsiveness in species of contrasting functional groups. Functional groups differed in their responses to elevated CO₂ and N treatments. Forb species exhibited strong down‐regulation of leaf N_mass concentrations (?26%) and photosynthetic capacity (?28%) in response to elevated CO₂, especially at high N supply, whereas C₃ grasses did not. Hence, achieved photosynthetic performance was markedly enhanced for C₃ grasses (+68%) in elevated CO₂, but not significantly for forbs. Differences in access to soil resources between forbs and grasses may distinguish their responses to elevated CO₂ and N addition. Forbs had lesser root biomass, a lower distribution of biomass to roots, and lower specific root length than grasses. Maintenance of leaf N, possibly through increased root foraging in this nutrient‐poor grassland, was necessary to sustain stimulation of photosynthesis under long‐term elevated CO₂. Dilution of leaf N and associated photosynthetic down‐regulation in forbs under elevated [CO₂], relative to the C₃ grasses, illustrates the potential for shifts in species composition and diversity in grassland ecosystems that have significant forb and grass components.     

Root Dynamics of Cultivar and Non‐Cultivar Population Sources of Two Dominant Grasses during Initial Establishment of Tallgrass Prairie

Dominance of warm‐season grasses modulates tallgrass prairie ecosystem structure and function. Reintroduction of these grasses is a widespread practice to conserve soil and restore prairie ecosystems degraded from human land use changes. Seed sources for reintroduction of dominant prairie grass species include local (non‐cultivar) and selected (cultivar) populations. The primary objective of this study was to quantify whether intraspecific variation in developing root systems exists between population sources (non‐cultivar and cultivar) of two dominant grasses (Sorghastrum nutans and Schizachyrium scoparium) widely used in restoration. Non‐cultivar and cultivar grass seedlings of both species were isolated in an experimental prairie restoration at the Konza Prairie Biological Station. We measured above‐ and belowground net primary production (ANPP and BNPP, respectively), root architecture, and root tissue quality, as well as soil moisture and plant available inorganic nitrogen (N) in soil associated with each species and source at the end of the first growing season. Cultivars had greater root length, surface area, and volume than non‐cultivars. Available inorganic N and soil moisture were present in lower amounts in soil proximal to roots of cultivars than non‐cultivars. Additionally, soil NO₃–N was negatively correlated with root volume in S. nutans cultivars. While cultivars had greater BNPP than non‐cultivars, this was not reflected aboveground root structure, as ANPP was similar between cultivars and non‐cultivars. Intraspecific variation in belowground root structure and function exists between cultivar and non‐cultivar sources of the dominant prairie grasses during initial reestablishment of tallgrass prairie. Population source selection should be considered in setting restoration goals and objectives.     

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.     

Effects of plant species and soil history on root morphology,arbuscular mycorrhizal colonization of roots,and biomass in four tallgrass prairie species

Although dominant C₄ grasses in tallgrass prairie are highly mycotrophic, for many non-dominant species neither extent of mycorrhizal colonization nor root morphology effects on plant–soil feedback interactions are known. In a laboratory study conducted from November 2013 to February 2014 at Governors State University (University Park, IL), we grew plants of a dominant C₄ grass (Andropogon gerardii) and three non-dominant forbs (Symphyotrichum laeve var. laeve, Symphyotrichum novae-angliae and Parthenium integrifolium) individually in soil collected in the field beneath a conspecific, collected beneath another study species, or in sterilized soil. The study addressed the following questions: (1) Is extent of mycorrhizal colonization of roots related to root structure? (2) How does soil history interact with plant root traits to influence plant–soil feedback? (3) How might plant–soil feedback patterns influence competitive interactions among study species? We found that proportion arbuscular mycorrhizal fungi (AMF) colonization decreased with increasing specific root length. Soil history had a stronger influence than plant species on total biomass of plants, with all species having highest total biomass when grown in Andropogon soil. Consequently, net, or heterospecific, feedback did not differ among pairwise species combinations, and was not different from 0. While these results suggest that no study species should have a competitive advantage in the field, Andropogon might still have an advantage through mechanisms such as competition for light. Future work in the field and including less mycotrophic species is needed to better understand AMF effects on competitive interactions.

     

Controls of nitrogen limitation in tallgrass prairie

Summary The relationship between fire frequency and N limitation to foliage production in tallgrass prairie was studied with a series of fire and N addition experiments. Results indicated that fire history affected the magnitude of the vegetation response to fire and to N additions. Sites not burned for over 15 years averaged only a 9% increase in foliage biomass in response to N enrichment. In contrast, foliage production increased an average of 68% in response to N additions on annually burned sites, while infrequently burned sites, burned in the year of the study, averaged a 45% increase. These findings are consistent with reports indicating that reduced plant growth on unburned prairie is due to shading and lower soil temperatures, while foliage production on frequently burned areas is constrained by N availability. Infrequent burning of unfertilized prairie therefore results in a maximum production response in the year of burning relative to either annually burned or long-term unburned sites.Foliage biomass of tallgrass prairie is dominated by C₄ grasses; however, forb species exhibited stronger production responses to nitrogen additions than did the grasses. After four years of annual N additions, forb biomass exceeded that of grass biomass on unburned plots, and grasses exhibited a negative response to fertilizer, probably due to competition from the forbs. The dominant C₄ grasses may out-compete forbs under frequent fire conditions not only because they are better adapted to direct effects of burning, but because they can grow better under low available N regimes created by frequent fire.     

Belowground responses of <Emphasis Type="Italic">Picea asperata</Emphasis> seedlings to warming and nitrogen fertilization in the eastern Tibetan Plateau

The impacts of global climatic change on belowground ecological processes of terrestrial ecosystems are still not clear. We therefore conducted an experiment in the subalpine coniferous forest ecosystem of the eastern edges of the Tibetan Plateau to study roots of Picea asperata seedlings and rhizosphere soil responses to soil warming and nitrogen availability from April 2007 to December 2008. The seedlings were subjected to two levels of temperature (ambient; infrared heater warming) and two nitrogen levels (0 or 25 g m⁻²year⁻¹ N). We used a free air temperature increase from an overhead infrared heater to raise both air and soil temperature by 2.1 and 2.6°C, respectively. The results showed that warming alone significantly increased total biomass, coarse root biomass and fine root biomass of P. asperata seedlings. Both total biomass and fine root biomass were increased, but coarse root biomass was significantly decreased by nitrogen fertilization and warming combined with nitrogen fertilization. Warming induced a prominent increase in soil organic carbon (SOC) and NO₃ ⁻-N of rhizosphere soil, while nitrogen fertilization significantly decreased SOC and NH₄ ⁺-N of rhizosphere soil. The warming, fertilization and warming × N fertilization interaction decreased soil microbial C significantly, but substantially increased soil microbial N. These results suggest that nitrogen deposition combined with warmer temperatures under future climatic change possibly will have no effect on fine root production of P. asperata seedlings, but could enhance the nitrification process of their rhizosphere soils in subalpine coniferous forests.     

Granivory reduces biomass and lignin concentrations of plant tissue during grassland assembly

Small mammals can influence grassland assembly by selecting against palatable plants – the community can become dominated by the plants they avoid. This predation-based selection could have indirect effects on community biomass and tissue quality, especially given how untasty plants may have higher concentrations of recalcitrant carbon compounds including lignin. We tested small mammal effects on biomass and tissue quality of roots and shoots in a two-year-old 18 ha restored tallgrass prairie with established zones of high and low plant predation. We focused on the three dominant herbaceous functional groups of tallgrass prairie (perennial forbs, C₃ and C₄ grasses), and targeted the early stages of assembly given that plant predation by small animals can unfold quickly and is difficult to subsequently quantify. We predicted rodent predation to create communities with reduced biomass but an increased abundance of lignin-rich plants; we only observed the former. Rodents reduced aboveground biomass by 46% but preferentially targeted lignin-rich plants, with the latter result explained by the predominance of granivory over herbivory – there was no opportunity for selection based on tissue palatability. Based strictly on aboveground biomass, we estimated small mammals reduced standing stocks of recalcitrant carbon by 65 kg ha⁻¹, with reductions in belowground stocks almost certainly higher given that root:shoot ratios averaged 21:1. Given that the quantity and quality of plant production can affect ecosystem functions including decomposition and the regulation of soil carbon stocks, our work suggests that non-random plant predation may substantially affect rates of soil carbon accumulation in the early stages of grassland development.     

Grazing and Ecosystem Carbon Storage in the North American Great Plains

Isotopic signatures of ¹³C were used to quantify the relative contributions of C₃ and C₄ plants to whole-ecosystem C storage (soil+plant) in grazed and ungrazed sites at three distinct locations (short-, mid- and tallgrass communities) along an east–west environmental gradient in the North American Great Plains. Functional group composition of plant communities, the source and magnitude of carbon inputs, and total ecosystem carbon storage displayed inconsistent responses to long-term livestock grazing along this gradient. C₄ plants [primarily Bouteloua gracilis (H.B.K.) Lag ex Steud.] dominated the long-term grazed site in the shortgrass community, whereas the ungrazed site was co-dominated by C₃ and C₄ species; functional group composition did not differ between grazed and ungrazed sites in the mid- and tallgrass communities. Above-ground biomass was lower, but the relative proportion of fine root biomass was greater, in grazed compared to ungrazed sites at all three locations. The grazed site of the shortgrass community had 24% more whole-ecosystem carbon storage compared to the ungrazed site (4022 vs. 3236 g C m⁻²). In contrast, grazed sites at the mid- and tallgrass communities had slightly lower (8%) whole-ecosystem carbon storage compared to ungrazed sites (midgrass: 7970 vs. 8683 g C m⁻²; tallgrass: 8273 vs. 8997 g C m⁻²). Differential responses between the shortgrass and the mid- and tallgrass communities with respect to grazing and whole-ecosystem carbon storage are likely a result of: (1) maintenance of larger soil organic carbon (SOC) pools in the mid- and tallgrass communities (7476–8280 g C m⁻²) than the shortgrass community (2517–3307 g C m⁻²) that could potentially buffer ecosystem carbon fluxes, (2) lower root carbon/soil carbon ratios in the mid- and tallgrass communities (0.06–0.10) compared to the shortgrass community (0.20–0.27) suggesting that variation in root organic matter inputs would have relatively smaller effects on the size of the SOC pool, and (3) the absence of grazing-induced variation in the relative proportion of C₃ and C₄ functional groups in the mid- and tallgrass communities. We hypothesize that the magnitude and proportion of fine root mass within the upper soil profile is a principal driver mediating the effect of community composition on the biogeochemistry of these grassland ecosystems.     

Biomass production and nitrogen content of C3- and C4- grasses in pure and mixed culture with different nitrogen supply

Summary Two C₃ grasses (Hordeum vulgare L., Avena sativa L.) and two C₄ grasses (Panicum miliaceum L., Panicum crus-galli L.) were cultivated in standard soil in the open air in pure cultures and in various mixed cultures at low and high nitrogen fertilization levels. After three months the dry weight, length and nitrogen content of the aboveground and below-ground parts of the plants and the shoot/root ratios were determined. Hordeum vulgare was the most successful species irrespective of the nitrogen fertilization level, and also exhibited in most cases the highest nitrogen concentrations. Panicum miliaceum, on the other hand, was the species least able to compete. The production of biomass was reduced in cultures growing under nitrogen starvation conditions, this phenomenon being more pronounced with respect to the C₄ than to the C₃ species. The decrease in the production of biomass at low N conditions was most drastic with Panicum crus-galli, the species with the lowest nitrogen content and thus assumed to be best adapted to nitrogen starvation conditions. In cultures growing at low nitrogen fertilization levels the shoot/root ratios of all species.shifted in favour of an increasing root proportion. The extent of this shift, however, differed from species to species.     

Plant nitrogen concentration, use efficiency, and contents in a tallgrass prairie ecosystem under experimental warming

Plant nitrogen (N) relationship has the potential to regulate plant and ecosystem responses strongly to global warming but has not been carefully examined under warmed environments. This study was conducted to examine responses of plant N relationship (i.e. leaf N concentration, N use efficiency, and plant N content in this study) to a 4‐year experimental warming in a tallgrass prairie in the central Great Plains in USA. We measured mass‐based N and carbon (C) concentrations of stem, green, and senescent leaves, and calculated N resorption efficiency, N use efficiency, plant N content, and C : N ratios of five dominant species (two C₄ grasses, one C₃ grass, and two C₃ forbs). The results showed that warming decreased N concentration of both green and senescent leaves, and N resorption efficiency for all species. N use efficiencies and C : N ratios were accordingly higher under warming than control. Total plant N content increased under warming because of warming‐induced increases in biomass production that are larger than the warming‐induced decreases in tissue N concentration. The increases in N contents in both green and senescent plant tissues suggest that warming enhanced both plant N uptake and return through litterfall in the tallgrass ecosystem. Our results also suggest that the increased N use efficiency in C₄ grasses is a primary mechanism leading to increased biomass production under warming in the grassland ecosystem.     

Differences in rhizosphere carbon-partitioning among plant species of different families

Interspecific variations in carbon (C) allocation and partitioning in the rhizosphere were investigated on 12 Mediterranean species belonging to different family groups (grasses, legumes, non-legume forbs) and having different life cycles. Plants grown individually in artificial soil, in a greenhouse and inoculated with rhizosphere microflora were labelled with ¹⁴CO₂ for 3 h at the vegetative stage. Rhizosphere respiration was measured during 6 days after which labelled C partitioning between shoots, roots, soil, root washing solution and respiration was estimated. The percentage of assimilated ¹⁴C allocated below ground differed significantly between species (41 – 76%) but no significant difference was found between grasses, legumes and non-legume forbs. When expressed as percentage of below-ground ¹⁴C, rhizosphere respiration was significantly smaller for non-legume forbs (42%) than for grasses (46%) and legumes (51%). Consequently more ¹⁴C was incorporated into root biomass in the former. Half-life of ¹⁴CO₂ evolution through respiration ranged from 23 h in legumes to 27 h for non-legume forbs and 37 h for grasses. This suggested differences in microbial activities due to quantities and quality of root exuded C. Rhizosphere respiration was positively correlated with the amount of ¹⁴C in the solution used to wash the roots on one hand, and root N concentration on the other hand. This led to a functional hierarchy between plant family groups of the overall rhizosphere activity. It went from non-legume forbs being the less active (except Crepis sancta)in terms of respiration and exudation, to grasses and then legumes, the most active but also the richest in nitrogen.     

Effects of Frequent Mowing on Survival and Persistence of Forbs Seeded into a Species-Poor Grassland

Many early attempts at tallgrass prairie reconstruction failed to achieve the high species diversity of remnant prairies, and instead consist primarily of C₄ grasses. We hypothesized that frequent mowing of established prairie grasses could create sufficient gaps in the aboveground and belowground environment to allow for the establishment of native forbs from seed. We studied forb seedling establishment in a 25‐year‐old prairie planting in northern Iowa that was dominated by native warm‐season grasses. In winter 1999, 23 species of native forbs were broadcast into the recently burned sod at a rate of 350 viable seeds/m². Treatment plots were mowed weekly for either one or two growing seasons, and control plots were unmowed. Mowed plots had greater light availability than controls, especially when warm‐season grasses began to flower. Overwinter seedling mortality was 3% in mowed treatments compared to 29% in the controls. Forbs in mowed plots had significantly greater root and shoot mass than those in control plots in the first and second growing seasons but were not significantly more abundant. By the fourth growing season, however, forbs were twice as abundant in the mowed treatments. No lasting negative impacts of frequent mowing on the grass population were observed. Mowing a second year influenced species composition but did not change total seedling establishment. Experimental evidence is consistent with the idea that mowing reduced competition for light from large established grasses, allowing forb seedlings the opportunity to reach sufficient size to establish, survive, and flower in the second and subsequent years.     

Septate endophyte colonization and host responses of grasses and forbs native to a tallgrass prairie

Native tallgrass prairies support distinct dark septate endophyte (DSE) communities exemplified by Periconia macrospinosa and Microdochium sp. that were recently identified as common root symbionts in this system. Since these DSE fungi were repeatedly isolated from grasses and forbs, we aimed to test their abilities to colonize different hosts. One Microdochium and three Periconia strains were screened for colonization and growth responses using five native grasses and six forbs in an in vitro system. Previously published data for an additional grass (Andropogon gerardii) were included and reanalyzed. Presence of indicative inter- and intracellular structures (melanized hyphae, microsclerotia, and chlamydospores) demonstrated that all plant species were colonized by the DSE isolates albeit to varying degrees. Microscopic observations suggested that, compared to forbs, grasses were colonized to a greater degree in vitro. Host biomass responses varied among the host species. In broad comparisons, more grass species than forbs tended to respond positively to colonization, whereas more forb species tended to be non-responsive. Based on the suspected differences in the levels of colonization, we predicted that tallgrass prairie grasses would support greater DSE colonization than forbs in the field. A survey of field-collected roots from 15 native species supported this hypothesis. Our study supports the “broad host range” of DSE fungi, although the differences in the rates of colonization in the laboratory and in the field suggest a greater compatibility between grasses and DSE fungi. Furthermore, host responses to DSE range from mutualism to parasitism, suggesting a genotype-level interplay between the fungi and their hosts that determines the outcome of this symbiosis.     

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.