The effects of eastern red cedar (Juniperus virginiana) invasion and removal on a dry bluff prairie ecosystem

Eastern red cedar (Juniperus virginiana) establishment increased dramatically in the tallgrass prairie biome of North America during the last 30 years. Since many of the remaining remnant prairies occur on steep, dry, and nutrient poor sites, threatened by the invasion of native and non-native woody species, it is important to understand how an invasive species such as eastern red cedar influences key environmental factors that may determine the future trajectory of these systems and whether abiotic and biotic components of the system are resilient following cedar removal. To address these issues we: (1) investigated the influence of eastern red cedar on micro-environmental factors; (2) evaluated how these micro-environmental factors responded to eastern red cedar removal; and (3) assessed the effect of eastern red cedar on herbaceous species germination and distribution. The invasion of eastern red cedar was associated with lower surface light availability and soil temperature, as seen in prior studies, but otherwise had effects distinct from those observed in prior studies. There was no effect of cedar on soil pH, and unlike prior studies, cedar patches had higher soil moisture compared to native C₄ prairie grass plots. Moreover, these effects had strong spatial signatures, with impacts of invasion on micro-environment and native vegetation differing dramatically with slope position and aspect. Three years after eastern red cedar was removed, micro-environmental factors and species composition became similar to the tree-free grass-dominated plots, indicating a significant capacity for recovery following possible cedar control. In a broader context, this study sheds light on the pathways and mechanisms driving the impacts of this biological invasion on dry, steep, nutrient poor systems and illustrates the capability of these systems to recover once the invading species is removed.     

Growth dynamics of oak seedlings (Quercus macrocarpa Michx. and Quercus muhlenbergii Engelm.) from gallery forests: implications for forest expansion into grasslands

Seedling growth dynamics of Quercus macrocarpa Michx. and Quercus muhlenbergii Engelm. were compared over a 3-month period under optimal growth conditions. These two species are the dominant trees at the western limit of the eastern deciduous forest, and are typically confined to gallery forests along stream beds in tallgrass prairie. Since tallgrass prairie is characterized by a highly variable climate and is prone to periodic drought, we hypothesized that these oaks would have rapid root growth and produce deep taproots as seedlings, enabling them to avoid drought stress and persist in this region. These traits may also facilitate forest expansion into the more xeric tallgrass prairie if fires are suppressed. Taproots of Q. macrocarpa and Q. muhlenbergii grew to approximately 140 cm and 100 cm in length, respectively, after 104 days. In both species, 65% or more of seedling biomass was allocated below ground, and root/total biomass was significantly greater in Q. muhlenbergii at 0-20 and 21-40 days after germination. The seedling taproot elongation rates reported here are much greater than rates reported in other eastern deciduous forest trees. Long-term precipitation data and soil moisture patterns from tallgrass prairie, when combined with rapid taproot elongation rates, suggest that soil moisture may not limit oak establishment or growth in tallgrass prairie in most years, although water uptake by roots was not measured in this study. Other factors, such as fire, herbivory, and seed predation and dispersal may be equally important in constraining the distribution of these species to gallery forests.     

The role of seed predation in the maintenance of the Cross Timbers ecotone of Oklahoma,USA

I investigated how seed predation differed among tree species and among microhabitats across the Cross Timbers and what that variation may tell us about how this ecotone is maintained. The ecotone is located in Oklahoma, USA, between the eastern deciduous forest and tallgrass prairie where seeds of eight common tree species were placed in three microhabitats (oak forest, tallgrass prairie, and sumac shrub/small-tree/grass mix). After nine days in the field, percent seeds remaining were scored for each of the 120 (8 species×3 microhabitats×5 replicates) dishes. I found for both wind-dispersed tree species, (ash, elm) there was significantly more predation in the prairie microsite, with similar small predation levels in the shrub and forest. For two of the three bird-dispersed species (dogwood, hackberry), there was significantly more predation in the prairie and shrub microsites compared to the forest. Red cedar, however, was not taken by predators very much anywhere. Finally, all three mammal-dispersed tree species (two oaks, pecan) showed significantly more predation in the shrub and forest microsites compared to the prairie. Whereas wind- and bird-dispersed species suffered less predation as microsites became more woody and dark, the dominant oaks showed the opposite trend. Consequently, seed predators are not preventing oaks from advancing across this ecotone, but yearly fluctuations in predator population density, especially in the shrub transitional zone, could be helping to maintain it.     

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.     

Carbon and water relations of juvenile Quercus species in tall‐grass prairie

Abstract. In ecosystems where environments are extreme, such as deserts, adult plant species may facilitate the establishment and growth of seedlings and juveniles. Because high temperatures and evaporative demand characterize tall‐grass prairies of the central United States (relative to forests), we predicted that the grassland‐forest ecotone, by minimizing temperature extremes and moderating water stress, may function to facilitate the expansion of Quercus species into undisturbed tall‐grass prairie. We assessed the carbon and water relations of juvenile Quercus macrocarpa and Q. muhlenbergii, the dominant tree species in gallery forests of northeast Kansas, in ecotone and prairie sites. To evaluate the potentially competitive effects of neighboring herbaceous biomass on these oaks, juveniles (< 0.5 m tall) of both species also were subjected to either: (1) removal of surrounding above‐ground herbaceous biomass, or (2) control (prairie community intact) treatments. Herbaceous biomass removal had no significant effect on gas exchange or water relations in these oak species in either the prairie or the ecotone environment. Although the ecotone did alleviate some environmental extremes, photosynthetic rates and stomatal conductance were ca. 20 % higher (p < 0.05) in both oaks in prairie sites vs. the ecotone. Moreover, although leaf temperatures on average were higher in oaks in the prairie, high leaf temperatures in the ecotone had a greater negative effect on photosynthesis. These data suggest that the grassland‐forest ecotone did not facilitate the growth of Quercus juveniles expanding into this grassland. Moreover, the carbon and water relations of juvenile oaks in the prairie appeared to be unaffected by the presence of the dominant C₄ grasses.     

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.     

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.     

Tallgrass prairie pollen assemblages in mid-continental North America

The mid-continent of North America has experienced dramatic and abrupt climate change during the Holocene, but the response of grassland vegetation to past climate change has been difficult to quantify. To improve interpretation of tallgrass prairie vegetation from pollen assemblages, we acquired and analysed a surface sample set collected from 25 small ponds (less than 10 ha surface area) in the largest contiguous remnant of tallgrass prairie in the USA. We compared these tallgrass prairie assemblages to 476 modern pollen samples classified as “prairie” in the North American Surface Sample database. We then compared the surface pollen assemblages with fossil pollen assemblages from sediment cores at two sites in Kansas—Cheyenne Bottoms and Muscotah Marsh—using the modern analog technique. Pollen assemblages in the Flint Hills surface samples were very similar to each other, with an average squared chord distance of 0.19. They were different than other modern grassland pollen assemblages mainly due to higher percentages of pollen from six woody taxa: Carya, Cornus, Juniperus, Juglans, Maclura, and Platanus. Arboreal pollen percentages ranged from 17 to 62 % and did not correlate with woody cover among sites. Cheyenne Bottoms was open grassland for the past 25,000 years, but it did not have many tallgrass prairie analogs. Muscotah Marsh did not have many grassland analogs over the past 30,000 years, possibly due to its position on the prairie-forest border or its surrounding wetland vegetation.     

The influence of savanna trees on nutrient,water and light availability and the understorey vegetation

In an East African savanna herbaceous layer productivity and species composition were studied around Acacia tortilis trees of three different age classes, as well as around dead trees and in open grassland patches. The effects of trees on nutrient, light and water availability were measured to obtain an insight into which resources determine changes in productivity and composition of the herbaceous layer. Soil nutrient availability increased with tree age and size and was lowest in open grassland and highest under dead trees. The lower N:P ratios of grasses from open grassland compared to grasses from under trees suggested that productivity in open grassland was limited by nitrogen, while under trees the limiting nutrient was probably P. N:P ratios of grasses growing under bushes and small trees were intermediate between large trees and open grassland indicating that the understorey of Acacia trees seemed to change gradually from a N-limited to a P-limited vegetation. Soil moisture contents were lower under than those outside of canopies of large Acacia trees suggesting that water competition between trees and grasses was important. Species composition of the herbaceous layer under Acacia trees was completely different from the vegetation in open grassland. Also the vegetation under bushes of Acacia tortilis was different from both open grassland and the understorey of large trees. The main factor causing differences in species composition was probably nutrient availability because species compositions were similar for stands of similar soil nutrient concentrations even when light and water availability was different. Changes in species composition did not result in differences in above-ground biomass, which was remarkably similar under different sized trees and in open grassland. The only exception was around dead trees where herbaceous plant production was 60% higher than under living trees. The results suggest that herbaceous layer productivity did not increase under trees by a higher soil nutrient availability, probably because grass production was limited by competition for water. This was consistent with the high plant production around dead trees because when trees die, water competition disappears but the high soil nutrient availability remains. Hence, in addition to tree soil nutrient enrichment, below-ground competition for water appears to be an important process regulating tree-grass interactions in semi-arid savanna.     

Soil physiochemical changes following 12 years of annual burning in a humid–subtropical tallgrass prairie: a hypothesis

Burning is known to stimulate growth of grassland vegetation, promote species diversity, and inhibit natural invasion by woody plants. However, the frequency at which grasslands are burned as part of their management can affect soil nutrient content and, ultimately, productivity. The objective of this study was to characterize changes in soil physical and chemical properties in a native tallgrass prairie after 12 years of annual burning. In 1989, five soil samples from the 0 to 10 cm depth were collected along a transect through a 3 ha parcel of native tallgrass prairie in central Arkansas. Soil sampling was repeated in 2001 to assess changes over time. Results showed that soil bulk density, electrical conductivity, extractable P, Na, Fe, and Mn decreased significantly (P < 0.05), while soil organic matter, total N and C, and the C/N ratio increased significantly (P < 0.05) within the 12-year period during which annual burning was the only imposed management practice. Mean extractable K, Ca, Mg, S, and Zn levels were all lower in 2001 than in 1989, but differences were not significant, while soil pH did not change. The results of this study indicate that annual export of several essential plant nutrients during prescribed burning of relatively small, remnant prairie fragments exceeds annual imports from atmospheric deposition and/or organic matter mineralization. Annual prescribed burning may be too frequent to maintain optimal ecosystem functioning and productivity. Decreasing the frequency of prescribed burning for native grassland management may help to retain more soil nutrients to sustain a higher level of productivity.     

Abiotic constraints on the establishment of Quercus seedlings in grassland

High evaporative demand and periodic drought characterize the growing season in midwestern grasslands relative to deciduous forests of the eastern US, and predicted climatic changes suggest that these climatic extremes may be exacerbated. Despite this less than optimal environment for tree seedling establishment, deciduous trees have expanded into adjacent tallgrass prairie within the last century leading to a dramatic land cover change. In order to determine the role of light and temperature on seedling establishment, we assessed carbon and water relations and aboveground growth of first‐year Quercus macrocarpa seedlings exposed to one of three conditions: (1) intact tallgrass prairie communities (control), (2) aboveground herbaceous biomass removed (grass removal), and (3) shade plus biomass removal to reduce light (PFD) to levels typical of the grassland‐forest ecotone (shade). In the 2000 growing season, precipitation was 35% below the long‐term average, which had a significant negative effect on oak seedling carbon gain at midseason (photosynthesis declined to 10% of maximum rates). However, net photosynthesis and stomatal conductance in the shade treatment was ca. 2.5 and 1.5 times greater, respectively, than in control treatment seedlings during this drought. During this period, leaf and air temperatures in control seedlings were similar whereas leaf temperatures in the shade treatment remained below air temperature. A late‐season recovery period, coincident with decreased air temperatures, resulted in increased net photosynthesis for all seedlings. Increased photosynthetic rates and water relations in shaded seedlings compared to seedlings in full sun suggest that, at least in dry years, high light and temperature may negatively impact oak seedling performance. However, high survival rates for all seedlings indicate that Q. macrocarpa seedlings are capable of tolerating both present‐day and future climatic extremes. Unless historic fire regimes are restored, forest expansion and land cover change are likely to continue.     

Restoration Efforts for Plant and Bird Communities in Tallgrass Prairies Using Prescribed Burning and Mowing

Recent losses and fragmentation of tallgrass prairie habitat to agriculture and urban development have led to corresponding declines in diversity and abundance of plants and birds associated with such habitat. Mowing and burning are alternative management strategies for restoring and rejuvenating prairies in fragmented landscapes, but their specific, comparative effects are the subjects of ongoing evaluation. We compared the responses of plant and bird communities on four sets of mowed, burned, and untreated sites of small (3–10 ha), fragmented tallgrass prairies at the DeSoto National Wildlife Refuge (DNWR), Iowa, U.S.A., during May–July in 1998 and 1999. Species richness and diversity of plants, resident grassland birds, and communities of birds associated with grassland edges (edge species) were independent of treatment. Although not affecting species richness and diversity in plant communities, mowed sites ranked lower in total plant coverage and total forb coverage than burned sites or untreated sites. In contrast, untreated sites had more coverage by shrubs, suggesting that mowing and burning did retard shrub encroachment. Overall, abundance and diversity of plants and birds were generally insensitive to management strategies. Small, fragmented sites of rare habitat may not respond in the short term to management treatments and may not be capable of supporting highly diverse communities, no matter how intensively manipulated. It is more probable that diversity of native prairie communities can be enhanced and restored only through long‐term efforts, acquisition of large land units capable of supporting stable populations, and deliberate reintroduction of species of high conservation value.     

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.     

Feeding ecology and emergence production of annual cicadas (Homoptera: Cicadidae) in tallgrass prairie

The emergence phenology and feeding ecology of annual cicadas in tallgrass prairie are poorly documented. However, these large insects are abundant, and their annual emergence represents a potentially important flux of energy and nutrients from belowground to aboveground. We conducted a study at Konza Prairie Research Natural Area in eastern Kansas to characterize and quantify cicada emergence and associated energy and nutrient fluxes. We established emergence trap transects in three habitat types (upland prairie, lowland prairie, and riparian forest), and collected cicadas every 3 days from May to September. A subset of trapped cicadas was used for species- and sex-specific mass, nutrient, and stable isotope analyses. Five species were trapped during the study, of which three were dominant. Cicadetta calliope and Tibicen aurifera exhibited significantly higher emergence production in upland prairie than in lowland prairie, and were not captured in forested sites at all. T. dorsata emerged from all three habitat types, and though not significant, showed a trend of greater abundance in lowland grasslands. Two less abundant species, T. pruinosa and T. lyricen, emerged exclusively from forested habitats. Nitrogen fluxes associated with total cicada emergence were estimated to be ∼4 kg N ha^–1 year^–1 in both grassland habitats, and 1.01 kg N ha^–1 year^–1 in forested sites. Results of stable isotope analyses showed clear patterns of resource partitioning among dominant cicada species emerging from grassland sites. T. aurifera and C. calliope had δ¹³C and δ¹⁵N signatures indicative of feeding on shallowly rooted C₄ plants such as the warm-season grasses dominant in tallgrass prairie ecosystems, whereas T. dorsata signatures suggested preferential feeding on more deeply rooted C₃ plants. Received: 20 September 1999 / Accepted: 9 December 1999     

RESPONSE OF ANDROPOGON GERARDII (POACEAE) TO FIRE-INDUCED HIGH VS. LOW IRRADIANCE ENVIRONMENTS IN TALLGRASS PRAIRIE: LEAF STRUCTURE AND PHOTOSYNTHETIC PIGMENTS

Photosynthetic pigments and several structural characteristics were measured in leaves of Andropogon gerardii from tallgrass prairie populations in an unburned, low-irradiance site and a burned, high-irradiance site to determine if these species displayed sun/shade differences similar to those documented in forest species. Early in the growing season, leaves of A. gerardii in the low-irradiance, unburned site had significantly lower stomatal density, pore length, and conductance, as well as specific leaf mass and thickness than leaves from the high-irradiance, burned site. Moreover, the chlorophyll a:b ratio, carotenoid content, and bundle sheath-vascular complex area were significantly lower, but chlorophyll content (mass/mass) was greater in leaves in unburned vs. burned sites. These differences are consistent with sun/shade adaptations reported for forest understory plants and may contribute to the low productivity of A. gerardii in unburned tallgrass prairie.     

Effects of mycorrhizae on growth and demography of tallgrass prairie forbs

The effects of arbuscular mycorrhizal (AM) symbiosis on ramet and genet densities, vegetative growth rates, and flowering of three forb species were studied in native tallgrass prairie in northeastern Kansas. Mycorrhizal activity was experimentally suppressed for six growing seasons on replicate plots in an annually burned and an infrequently burned watershed with the fungicide benomyl. Benomyl reduced mycorrhizal root colonization to an average of 4.2%, approximately a two-thirds reduction relative to controls (13.7% colonization). Mycorrhizae influenced the population structure of these forbs. Although mycorrhizal suppression had no long-term effect on genet densities and no effect on ramet survivorship throughout the growing season, the number of ramets per individual was significantly increased such that ramet densities of all three species were approximately doubled in response to long-term mycorrhizal suppression. Effects of mycorrhizae on ramet growth and reproduction varied among species. Ramet growth rates, biomass, and flowering of Salvia azurea were greater in plots with active mycorrhizal symbiosis, whereas mycorrhizae reduced ramet growth rates and biomass of Artemesia ludoviciana. Aster sericeus ramet growth rates and biomass were unaffected by the fungicide applications, but its flowering was reduced.The pattern of responses of these three species to mycorrhizae differed considerably between the two sites of contrasting fire regime, indicating that the interaction of fire-induced shifts in resource availability and mycorrhizal symbiosis together modulates plant responses and the intensity and patterns of interspecific competition between and among tallgrass prairie grass and forb species. Further, the results indicate that effects of mycorrhizae on community structure are a result of interspecific differences in the balance between direct positive effects of the symbiosis on host plant performance and indirect negative effects mediated through altered competitive interactions.     

Using Auchenorrhyncha (Insecta: Hemiptera) to develop a new insect index in measuring North American tallgrass prairie quality

Auchenorrhyncha (i.e., leafhoppers, treehoppers, spittlebugs, and planthoppers) represent some of the most diverse groups of herbivorous insects in the tallgrass prairie biome, they have close associations with many native prairie grasses and forbs, and respond in predictable ways to changes in native grassland degradation. These attributes make Auchenorrhyncha ideal candidates in the development of a habitat quality index to measure tallgrass prairie quality. In this study we propose the development of a species-based habitat quality index called the Auchenorrhyncha quality index or AQI as a useful method in tracking the condition of tallgrass prairie quality. The AQI is computed by summing six ecological characteristics (i.e., host plant specificity, voltinism, overwintering microhabitat, wing length, habitat fidelity, and origin) for each Auchenorrhynchan insect encountered, yielding coefficient of conservatism (CC) values that range from 0 (habitat generalist/tolerant to disturbance) to 18 (habitat specific/intolerant to disturbance). These CC values are averaged and combined with species richness producing un-weighted by abundance AQI (AQI_w/outN) and weighted by abundance AQI (AQI_w/N). The performance of the AQI was evaluated by examining the effects of sampling intensity on this index using a sweep net and a vacuum apparatus from 10 sites located on the three main North American tallgrass prairie communities, wet-mesic, sand, and loess hill. Scientists and land managers can adequately sample Auchenorrhyncha from four transects using a vacuum. Also, the highest AQI values were found from loess hill and sand prairies, indicating that conservation efforts should focus on these prairie communities. Additional applications of the AQI may include: (1) discriminating prairie quality at various spatial scales; (2) testing hypotheses about the effects of disturbance on prairie habitat (e.g. prescribed burning); (3) using the AQI as a model in developing habitat quality indices based on other diverse groups of grassland insects; and (4) the AQI has the capacity to be readily modified in assessing the quality of other biomes. Ultimately, the AQI should be used in combination with other habitat quality indices based on other diverse groups of organisms, such as plants and other insects, to provide a more complete assessment of native habitat quality.     

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.     

Beneath the veil: plant growth form influences the strength of species richness–productivity relationships in forests

Aim Species richness has been observed to increase with productivity at large spatial scales, though the strength of this relationship varies among functional groups. In forests, canopy trees shade understorey plants, and for this reason we hypothesize that species richness of canopy trees will depend on macroclimate, while species richness of shorter growth forms will additionally be affected by shading from the canopy. In this study we test for differences in species richness–productivity relationships (SRPRs) among growth forms (canopy trees, shrubs, herbaceous species) in small forest plots. Location We analysed 231 plots ranging from 34.0° to 48.3° N latitude and from 75.0° to 124.2° W longitude in the United States. Methods We analysed data collected by the USDA Forest Inventory and Analysis program for plant species richness partitioned into different growth forms, in small plots. We used actual evapotranspiration as a macroclimatic estimate of regional productivity and calculated the area of light‐blocking tissue in the immediate area surrounding plots for an estimate of the intensity of local shading. We estimated and compared SRPRs for different partitions of the species richness dataset using generalized linear models and we incorporated the possible indirect effects of shading using a structural equation model. Results Canopy tree species richness increased strongly with regional productivity, while local shading primarily explained the variation in herbaceous plant richness. Shrub species richness was related to both regional productivity and local shading. Main conclusions The relationship between total forest plant species richness and productivity at large scales belies strong effects of local interactions. Counter to the pattern for overall richness, we found that understorey herbaceous plant species richness does not respond to regional productivity gradients, and instead is strongly influenced by canopy density, while shrub species richness is under multivariate control.     

Responses of two bunchgrasses to nitrogen addition in tallgrass prairie: the role of bud bank demography

Growth of tallgrass prairie plants, many of which maintain substantial bud banks, can be limited by nitrogen (N), water, and/or light. We hypothesized that tallgrass prairie plants respond to increases in N through demographic effects on the bud bank. We tested the effects of a pulse of N on (1) bud bank demography, (2) plant reproductive allocation, and (3) ramet size. We parameterized matrix models, considering each genet as a population of plant parts. Nitrogen addition significantly impacted bud bank demography in two subdominant species of bunchgrass: Sporobolus heterolepis (a C(4) grass) and Koeleria macrantha (a C(3) grass), but had no effect on the size of individual ramets. Emergence from the bud bank and ramet population growth rates (λ) were significantly higher in S. heterolepis genets that received supplemental N. Nitrogen addition also affected the bud demography of K. macrantha, but N addition decreased rather than increased λ. Prospective and retrospective demographic analyses indicated that bud bank dynamics were the most important demographic processes driving plant responses to nutrient availability. Thus, the variation in productivity in these tallgrass prairie species is driven principally by the demography of the bud bank rather than by the physiology and growth of aboveground tillers. Improved understanding of bud bank dynamics may lead to improved predictive models of grassland responses to environmental changes such as altered N deposition and precipitation.