Comparison of damage to native and exotic tallgrass prairie plants by natural enemies

We surveyed the prevalence and amount of leaf damage related to herbivory and pathogens on 12 pairs of exotic (invasive and noninvasive) and ecologically similar native plant species in tallgrass prairie to examine whether patterns of damage match predictions from the enemy release hypothesis. We also assessed whether natural enemy impacts differed in response to key environmental factors in tallgrass prairie by surveying the prevalence of rust on the dominant C₄ grass, Andropogon gerardii, and its congeneric invasive exotic C₄ grass, A. bladhii, in response to fire and nitrogen fertilization treatments. Overall, we found that the native species sustain 56.4% more overall leaf damage and 83.6% more herbivore-related leaf damage when compared to the exotic species. Moreover, we found that the invasive exotic species sustained less damage from enemies relative to their corresponding native species than the noninvasive exotic species. Finally, we found that burning and nitrogen fertilization both significantly increased the prevalence of rust fungi in the native grass, while rust fungi rarely occurred on the exotic grass. These results indicate that reduced damage from enemies may in part explain the successful naturalization of exotic species and the spread of invasive exotic species in tallgrass prairie.     

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.     

Effects of the soil microbiome on the demography of two annual prairie plants

1. Both mutualistic and pathogenic soil microbes are known to play important roles in shaping the fitness of plants, likely affecting plants at different life cycle stages.
2. In order to investigate the differential effects of native soil mutualists and pathogens on plant fitness, we compared survival and reproduction of two annual tallgrass prairie plant species (Chamaecrista fasciculata and Coreopsis tinctoria) in a field study using 3 soil inocula treatments containing different compositions of microbes. The soil inocula types included fresh native whole soil taken from a remnant prairie containing both native mutualists and pathogens, soil enhanced with arbuscular mycorrhizal (AM) fungi derived from remnant prairies, and uninoculated controls.
3. For both species, plants inoculated with native prairie AM fungi performed much better than those in uninoculated soil for all parts of the life cycle. Plants in the native whole prairie soil were either generally similar to plants in the uninoculated soil or had slightly higher survival or reproduction.
4. Overall, these results suggest that native prairie AM fungi can have important positive effects on the fitness of early successional plants. As inclusion of prairie AM fungi and pathogens decreased plant fitness relative to prairie AM fungi alone, we expect that native pathogens also can have large effects on fitness of these annuals. Our findings support the use of AM fungi to enhance plant establishment in prairie restorations.

     

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.     

Tallgrass prairie ants: their species composition,ecological roles,and response to management

Ants are highly influential organisms in terrestrial ecosystems, including the tallgrass prairie, one of the most endangered ecosystems in North America. Through their tunneling, ants affect soil properties and resource availability for animals and plants. Ants also have important ecological roles as consumers of plant tissue and seeds. In the last several decades, various organizations, agencies, and agricultural producers have attempted to create wildlife habitat or reduce soil erosion by seeding thousands of hectares of bare cropland in the central United States with tallgrass prairie seed mixes. Although initially, monitoring of these restorations and of unplowed prairie remnants focused on plants and birds, in recent years the response of invertebrates such as ants has increasingly been the subject of research. An understanding of tallgrass prairie ant communities can help land managers and scientists better monitor the ecological condition of tallgrass prairie and guide management and restoration efforts. Here I review our current knowledge of ant species found within tallgrass prairie, their ecological roles, and their response to management.     

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

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

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

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

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

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

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

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

Patch dynamics of a native grass in relation to the spread of invasive smooth brome (Bromus inermis)

The effects of invasive species on the patch dynamics (establishment, growth, and local extinction) of native species are not well studied, owing to the need for relatively fine-scale data on the distribution of species. Within the prairie pothole region of the United States and Canada, the grass, Bromus inermis (smooth brome) has become established by invading disturbed prairies, and through repeated introductions for soil retention and animal graze. In this study, the impact of smooth brome on the patch dynamics of a dominant native grass species, Spartina pectinata (prairie cordgrass), was assessed using fine-scale (sub-meter) mapping of the distribution of cordgrass and brome in three prairie fragments from 2000 to 2006. Using GIS spatial analyses, we determined that cordgrass patch growth was two times greater in areas not invaded by smooth brome versus areas that were heavily infested with smooth brome. Among sites and time periods, there was a consistent significant negative relationship between the amount of smooth brome surrounding a patch of cordgrass and the growth of that cordgrass patch. The probability of establishment of a new patch of cordgrass averaged 1.3 times higher in areas of low brome coverage (<25%) than areas of high brome coverage (>75%). Conversely, existing cordgrass patches were 7.8 times more likely to go extinct in areas of high than low brome coverage. This is one of only a few field studies to provide evidence of the negative impact of smooth brome on native flora and hopefully will serve as justification for the development of a formal management plan to limit the distribution of this species in tallgrass prairie ecosystems.     

Comparison of butterfly communities and abundances between marginal grasslands and conservation lands in the eastern Great Plains

The alteration and fragmentation of native tallgrass prairie in the Midwestern United States has created a need to identify other land types with the ability to support grassland butterfly species. This study examines butterfly usage of marginal grasslands, which consist of semi-natural grasslands existing within in a larger agricultural matrix, compared to grasslands managed for conservation of prairie species. Using generalized linear mixed models we analyzed how land purpose (marginal vs. conservation grasslands) affected butterfly abundance. We found grassland butterfly species to be significantly more common on conservation grasslands, whereas generalist species were significantly more common on marginal grasslands. Results of ordination analyses indicated that while many species used both types of habitats, butterfly species assemblages were distinct between habitat types and that edge to interior ratio and the floristic quality index of sites were important habitat characteristics driving this distinction. Within conservation grasslands we examined the relationship between butterfly abundance and the planting diversity used in restoring each site. We found higher diversity restorations hosted more individuals of butterflies considered habitat generalists, as well as species considered to be of conservation concern.     

Nitrogen fertilization increases diversity and productivity of prairie communities used for bioenergy

Using prairie biomass as a renewable source of energy may constitute an important opportunity to improve the environmental sustainability of managed land. To date, assessments of the feasibility of using prairies for bioenergy production have focused on marginal areas with low yield potential. Growing prairies on more fertile soil or with moderate levels of fertilization may be an effective means of increasing yields, but increased fertility often reduces plant community diversity. At a fertile site in central Iowa with high production potential, we tested the hypothesis that nitrogen fertilization would increase aboveground biomass production but would decrease diversity of prairies sown and managed for bioenergy production. Over a 3 year period (years 2–4 after seeding), we measured aboveground biomass after plant senescence and species and functional‐group diversity in June and August for multispecies mixtures of prairie plants that received no fertilizer or 84 kg N ha^?1 year^?1. We found that nitrogen fertilization increased aboveground biomass production, but with or without fertilization, the prairies produced a substantial amount of biomass: averaging (±SE) 12.2 ± 1.3 and 9.1 ± 1.0 Mg ha^?1 in fertilized and unfertilized prairies, respectively. Unfertilized prairies had higher species diversity in June, whereas fertilized prairies had higher species diversity in August at the end of the study period. Functional‐group diversity was almost always higher in fertilized prairies. Composition of unfertilized prairies was characterized by native C₄ grasses and legumes, whereas fertilized prairies were characterized by native C₃ grasses and forbs. Although most research has found that nitrogen fertilization reduces prairie diversity, our results indicate that early‐spring nitrogen fertilization, when used with a postsenescence annual harvest, may increase prairie diversity. Managing prairies for bioenergy production, including the judicious use of fertilization, may be an effective means of increasing the amount of saleable products from managed lands while potentially increasing plant diversity.     

Relatedness of Macrophomina phaseolina isolates from tallgrass prairie, maize, soybean and sorghum

Agricultural and wild ecosystems may interact through shared pathogens such as Macrophomina phaseolina , a generalist clonal fungus with more than 284 plant hosts that is likely to become more important under climate change scenarios of increased heat and drought stress. To evaluate the degree of subdivision in populations of M. phaseolina in Kansas agriculture and wildlands, we compared 143 isolates from maize fields adjacent to tallgrass prairie, nearby sorghum fields, widely dispersed soybean fields and isolates from eight plant species in tallgrass prairie. Isolate growth phenotypes were evaluated on a medium containing chlorate. Genetic characteristics were analysed based on amplified fragment length polymorphisms and the sequence of the rDNA-internal transcribed spacer (ITS) region. The average genetic similarity was 58% among isolates in the tallgrass prairie, 71% in the maize fields, 75% in the sorghum fields and 80% in the dispersed soybean fields. The isolates were divided into four clusters: one containing most of the isolates from maize and soybean, two others containing isolates from wild plants and sorghum, and a fourth containing a single isolate recovered from Solidago canadensis in the tallgrass prairie. Most of the sorghum isolates had the dense phenotype on media containing chlorate, while those from other hosts had either feathery or restricted phenotypes. These results suggest that the tallgrass prairie supports a more diverse population of M. phaseolina per area than do any of the crop species. Subpopulations show incomplete specialization by host. These results also suggest that inoculum produced in agriculture may influence tallgrass prairie communities, and conversely that different pathogen subpopulations in tallgrass prairie can interact there to generate 'hybrids' with novel genetic profiles and pathogenic capabilities.     

Interspecific nutrient transfer in a tallgrass prairie plant community

Interplant nutrient transfer may be an important ecological process in grasslands, and may significantly influence plant neighborhood interactions. We investigated the potential for phosphorus transfer between the dominant grass Andropogon gerardii and several neighboring plant species in tallgrass prairie via a field ³²PO₄ labelling experiment. The mean amount of ³²P received from donor shoots differed significantly among neighboring species and decreased with increasing distance from the donor. In general, forbs and cool-season C₃ grasses received more labelled ³²P than warm-season C₄ grasses. Phosphorus transfer occurred over distances up to 0.5 m. The effects of species and distance on movement of phosphorus changed with increasing time after labelling. The relative mass of receiver and donor shoots did not affect amounts of ³²P transfer. A benomyl fungicide treatment, applied to suppress mycorrhizal activity, likely did not affect existing vegetative hyphae and did not affect the amount of ³²P transferred. These studies demonstrate that: (1) phosphorus is transferred among neighboring species in tallgrass prairie plant communities, (2) phosphorus may be transferred over significantly greater distances than reported in other grasslands, and (3) there is differential transfer among co-occurring species. Hypothesized mechanisms accounting for these patterns in tallgrass prairie include mycorrhizal hyphal interconnections and/or extensive and differential root and rhizosphere overlap among neighboring species.     

POST-BURN DIFFERENCES IN SOLAR RADIATION,LEAF TEMPERATURE AND WATER STRESS INFLUENCING PRODUCTION IN A LOWLAND TALLGRASS PRAIRIE

The influence of standing dead biomass on available solar radiation, leaf temperature (T_leaf) and leaf water potential (ѱ_leaf) of Andropogon gerardii in unburned tallgrass prairie was compared to burned prairie in eastern Kansas. The standing dead reduced photosynthetically active radiation incident on emerging shoots by 58.8% in unburned compared to burned prairie during the initial 30 days of the growing season. Aboveground production in unburned prairie was similarly reduced during this period (55.4%) compared to burned prairie. Leaf temperatures in A. gerardii were greater in unburned prairie than in burned early in the season, but were nearly equal by the end of the growing season. The maximum elevation of T_leaf in unburned prairie above burned was 9.5 C. The maximum unburned T_leaf measured was 41.5 C compared to 39.4 C in burned prairie. Lower windspeed adjacent to leaves in unburned prairie resulting in reduced convective cooling may have caused higher T_leaf in unburned prairie. Leaf water potential was significantly lower in unburned prairie than in burned prairie early in the season but was higher in unburned prairie by late season. The seasonal minimum ѱ_leaf in burned prairie was — 1.60 MPa compared to —1.45 MPa in unburned prairie. The combined effect of these post-burn differences in solar radiation, T_leaf and ѱ_leaf may be significant in contributing to the lower production in unburned compared to burned tallgrass prairie.     

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.     

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.     

Diversity–productivity relationships in two ecologically realistic rarity–extinction scenarios

To develop a better understanding of how biodiversity loss and productivity are related, we need to consider ecologically realistic rarity (i.e. reduced evenness and increased dominance) and extinction (i.e. reduced richness) scenarios. Furthermore, we need to identify and better understand the factors that influence species and community yielding behaviors because the general conditions for overyielding are the same as those for coexistence. We established experimental tallgrass prairie plots in Iowa to determine how two ecologically realistic rarity–extinction scenarios influenced aboveground net primary productivity (ANPP) and disassembly. Equal‐mass seedlings of six tallgrass prairie species were transplanted into field plots to establish realistic declining species evenness (high, medium, low) and richness (4, 1) treatments. Across declining evenness treatments, the relative abundance of the ubiquitous tall species Andropogon gerardii increased, the relative abundance of the tall species Salvia azurea was constant, and the relative abundance of two short (dissimilar height scenario) or two tall species (tall scenario) decreased. Monocultures of Andropogon represented a continuation of this trend until there was complete dominance by Andropogon and extinction of all other species. Our treatments also allowed us to test if variation in plant height contributes to the complementarity effect. Niche partitioning in plant height was not positively related to complementarity. The effects of declining species evenness and richness on the diversity–productivity relationship were different for these two ecologically realistic rarity–extinction scenarios. Specifically, as diversity declined across treatments, ANPP and the selection effects decreased in tall communities, but not in dissimilar communities. Additionally, differences between these two scenarios revealed that decreased species yielding behavior is associated with two tallgrass prairie extinction risk factors, rarity and short height. The differences between these scenarios demonstrate the importance of incorporating the known patterns of diversity declines into future studies.     

Determinants of C3 forb growth and production in a C4 dominated grassland

Forbs are the most abundant species within the vascular flora of tallgrass prairie and they make the greatest contribution to biodiversity of all growth forms. However, little is known about the factors that determine their productivity and growth rates. The objective of this study was to assess the controls of forb growth (absolute and relative) and production in tallgrass prairie from a long-term burning experiment at the Konza Prairie in NE Kansas. Over the 14-year study, forbs comprised 9% of the total biomass production on sites with a high fire frequency vs. 29% on the low fire frequency site, with gramminoids accounting for the remainder. Although interannual variations in peak biomass of the grasses was strongly correlated with environmental variables related to water availability, there were no similar relationships for forbs, suggesting that production of forbs and grasses responded to interannual variations in climate in different ways. Multivariate analysis of climatic controls on growth rates of grasses and forbs yielded similar results. Although forbs had low biomass and absolute growth per unit ground area in frequently burned prairie, their relative growth rates were highest in such sites. Thus, it appears that reduced growth rates of individual forbs per se do not limit forb success in annually burned prairie. Instead, direct negative effects of fire on forbs (increased mortality) may be responsible. Determinants of forb growth and productivity in unburned prairie remain unresolved.     

Using prairie restoration to curtail invasion of Canada thistle: the importance of limiting similarity and seed mix richness

Theory has predicted, and many experimental studies have confirmed, that resident plant species richness is inversely related to invisibility. Likewise, potential invaders that are functionally similar to resident plant species are less likely to invade than are those from different functional groups. Neither of these ideas has been tested in the context of an operational prairie restoration. Here, we tested the hypotheses that within tallgrass prairie restorations (1) as seed mix species richness increased, cover of the invasive perennial forb, Canada thistle (Cirsium arvense) would decline; and (2) guilds (both planted and arising from the seedbank) most similar to Canada thistle would have a larger negative effect on it than less similar guilds. Each hypothesis was tested on six former agricultural fields restored to tallgrass prairie in 2005; all were within the tallgrass prairie biome in Minnesota, USA. A mixed-model with repeated measures (years) in a randomized block (fields) design indicated that seed mix richness had no effect on cover of Canada thistle. Structural equation models assessing effects of cover of each planted and non-planted guild on cover of Canada thistle in 2006, 2007, and 2010 revealed that planted Asteraceae never had a negative effect on Canada thistle. In contrast, planted cool-season grasses and non-Asteraceae forbs, and many non-planted guilds had negative effects on Canada thistle cover. We conclude that early, robust establishment of native species, regardless of guild, is of greater importance in resistance to Canada thistle than is similarity of guilds in new prairie restorations.