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.     

Effects of management practices on annual net N-mineralization in a restored prairie and maize agroecosystems

Nitrogen (N) mineralization is a spatially variable and difficult component of the N cycle to quantify accurately under field conditions. Net N-mineralization was compared by direct measurement, indirect estimate, and laboratory incubation for a restored tallgrass prairie and for deficiently and optimally N-fertilized, no-tillage (NT) and chisel-plowed (CP) maize (Zea mays L.) agroecosystems on Plano silt loam soil (fine-silty, mixed, superactive, mesic Typic Argiudoll) in Wisconsin, USA. Four years of in-situ field measurements using an incubated-soil-core/ion-exchange-resin-bag technique showed that land use significantly affected net N-mineralization. Net N-mineralization was consistently smaller in the restored prairie than in the maize agroecosystems and typically larger in the CP than in the NT maize agroecosystems. Three independent methods for indirectly estimating annual net N-mineralization (i.e., N budget residual, deficiently N-fertilized plant N uptake, and profile-scaled in-situ field measurements) were relatively consistent at capturing land-use and tillage effects on net N-mineralization. Laboratory incubation and periodic leaching of Fall-sampled soils demonstrated that both mineralized N and labile C were co-limiting factors influencing N-mineralization in agricultural soils and generally supported field measurements by showing a significant difference in net N-mineralization with and without added fertilizer-N.     

Plankton respiration and biomass as functional indicators of recovery in restored prairie wetlands

Reliable ecological indicators of wetland integrity are necessary for assessing recovery of restored wetlands; yet, little consensus currently exists on which indicators are most appropriate. We employed indicators derived from simple, standard measures of ecosystem function selected on the basis of ecological succession theory developed by [Science 164 (1969) 262; Bioscience 35 (1985) 419], which suggests that respiration:biomass ratios should increase in disturbed systems due to the diversion of energy from growth to maintenance. This hypothesis holds potential for the development of a simple ecological indicator and therefore was tested among prairie wetlands restored after drainage disturbance. No difference was observed in respiration:biomass ratios in restored wetlands and reference wetlands designated as controls. Plankton respiration or biomass may be poor indicators of disturbance because plankton responds quickly to re-establishment of a wetland hydrology regime and/or because different plankton species may have redundant function. We suggest employing more revealing assessment techniques that employ simultaneous examination of ecosystem structure and function to better characterize subtle or lingering effects of wetland disturbance after restoration.     

Vegetation and environmental conditions in recently restored wetlands in the prairie pothole region of the USA

How closely the vegetation of restored wetlands resembles that of comparable natural wetlands is a function of the probability of propagules of wetland species reaching reflooded wetlands and how similar environmental conditions in the restored wetland are those in the natural wetlands. Three years after reflooding, we examined the vegetation composition, water level fluctuations, soil organic carbon content, and soil bulk density as well as surface water pH, alkalinity, conductivity, and calcium and magnesium concentrations of 10 restored and 10 natural wetlands. In the restored wetlands, more species of submersed aquatics colonized than were found in natural wetlands, and they rapidly spread to form extensive beds that were larger than those found in natural wetlands. Emergent and wet meadow species in restored wetlands, however, were found in only sparse stands as were a variety of annuals. The vegetation of natural wetlands was predominantly large stands of emergent species. Fluctuations in water storage volume and basin surface area were similar for both restored and natural wetlands. The surface water in restored wetlands had higher pH and lower alkalinity, conductivity, and calcium and magnesium concentrations than that in natural wetlands. Soils of restored wetlands have a lower organic carbon content and higher bulk density than do those of natural wetlands. Our results suggest that for submersed aquatics, dispersal of propagules to restored wetlands is rapid and environmental conditions in restored wetlands are very suitable for their establishment. For other guilds of wetland species, e.g., sedges and other wet meadow species, dispersal to restored wetlands is likely much slower and may pose a serious problem for the re-establishment of these species in restored wetlands. Even if dispersal is not limiting, low surface organic carbon and high bulk density may prevent the establishment of these species in restored wetlands.     

Greenhouse gas emissions and carbon sequestration potential in restored wetlands of the Canadian prairie pothole region

North American prairie pothole wetlands are known to be important carbon stores. As a result there is interest in using wetland restoration and conservation programs to mitigate the effects of increasing greenhouse gas concentration in the atmosphere. However, the same conditions which cause these systems to accumulate organic carbon also produce the conditions under which methanogenesis can occur. As a result prairie pothole wetlands are potential hotspots for methane emissions. We examined change in soil organic carbon density as well as emissions of methane and nitrous oxide in newly restored, long-term restored, and reference wetlands across the Canadian prairies to determine the net GHG mitigation potential associated with wetland restoration. Our results indicate that methane emissions from seasonal, semi-permanent, and permanent prairie pothole wetlands are quite high while nitrous oxide emissions from these sites are fairly low. Increases in soil organic carbon between newly restored and long-term restored wetlands supports the conclusion that restored wetlands sequester organic carbon. Assuming a sequestration duration of 33 years and a return to historical SOC densities we estimate a mean annual sequestration rate for restored wetlands of 2.7 Mg C ha⁻¹year⁻¹ or 9.9 Mg CO₂ eq. ha⁻¹ year⁻¹. Even after accounting for increased CH₄ emissions associated with restoration our research indicates that wetland restoration would sequester approximately 3.25 Mg CO₂ eq. ha⁻¹year⁻¹. This research indicates that widescale restoration of seasonal, semi-permanent, and permanent wetlands in the Canadian prairies could help mitigate GHG emissions in the near term until a more viable long-term solution to increasing atmospheric concentrations of GHGs can be found.     

Resource limitation and recruitment patterns in a coral reef fish assemblage

The potential effects of food and shelter availability on the recruitment and early survivorship of coral reef fishes were studied on St. Croix, U.S. Virgin Islands. The faunal assemblage studied included diurnally active fishes found in the “rubble/sand” habitat. The most abundant members were: beaugregory, Stegastes leucostictus (Muller & Troschel), goldspotted goby, Gnatholepis thompsoni Jordan, bridled goby, Coryphopterus glaucofraenum Gill, surgeonfishes, Acanthurus bahianus Castelnau and A. chirurgus (Bloch), and French grunt, Haemulon flavolineatum (Desmarest). Comparisons of recruitment to reefs constructed from substrata that varied in morphological characteristics showed that there were differences in the relative abundances of recruits attracted to and/or surviving on the different reef types. Juveniles of most species appeared to prefer the branching coral Porites porites (Pallas), which provided a large number of small crevices between the branches.Manipulations of the availability of shelter sites for fishes demonstrated that recruitment and/or early survivorship were strongly limited by the number of refuges. This result was found in six separate carried out during different years and in different seasons. Shelter site availability presumably limits fish populations through its effects on prédation rates.Experimental manipulations of food availability indicated that food does not directly influence settlement or early survivorship of coral reef fishes. However, it is probable that correlations between habitat characteristics and food availability have influenced the evolution of settling preferences.     

Resource limitation,predation risk and compensatory growth in a damselfly

Periods of poor nutrition during early development may have negative fitness consequences in subsequent periods of ontogeny. In insects, suppression of growth and developmental rate during the larval stage are likely to affect size and timing of maturity, which in turn may lead to reduced reproductive success or survivorship. In light of these costs, individuals may achieve compensatory growth via behavioural or physiological mechanisms following food limitation. In this study, we examined the effects of a temporary period of food restriction on subsequent growth and age and size at maturity in the larval damselfly Ischnura verticalis (Odonata: Coenagrionidae). We also asked whether this temporary period of reduced nutrition affected subsequent foraging behaviour under predation risk. I. verticalis larvae exposed to a temporary food shortage suffered from a reduced growth rate during this period relative to a control group that was fed ad libitum. However, increased growth rates later in development ensured that adult body size measurements (head and pronotum widths) did not differ between the treatments upon emergence. In contrast, adult dry mass did not catch up to that of the controls, indicating that the increased growth rates for size dimensions occur at the cost of similar gains in mass. Predators reduced foraging effort of larvae, but this reduction did not differ between control larvae and those previously exposed to poor nutrition.     

Seed size and recruitment limitation influence seedling establishment in three tallgrass prairie species

A seed size–seed number tradeoff predicts that large numbers of seeds have an establishment advantage under low levels of competition, while large seed size is advantageous under asymmetric competition. Testing these predictions in the context of tallgrass prairie restoration should increase predictability of outcomes of species composition—restoration technique combinations. I conducted field and greenhouse experiments comparing seedling establishment among three tallgrass prairie species with different seed mass-seed number combinations, but with similar mass of seeds sown. Species were prairie dock [Silphium terebinthinaceum, mass = 33.2 + 2.1 mg ( \(\overline{x}\)  + SE, n = 100)], wild quinine (Parthenium integrifolium, mass = 3.1 + 0.3 mg), and smooth blue aster (Symphyotrichum laeve var. laeve, mass = 0.53 + 0.02 mg). Seeds were sown into mowed or tilled plots with shade and mulch treatments in an old field in northeastern Illinois USA, and seeds were sown into containers with or without vegetative cover in a greenhouse. In mowed plots, species with larger numbers of seeds sown had higher seedling densities. In tilled plots, seedling densities of the large-seeded Silphium were reduced less by shade than the small-seeded Parthenium. Symphyotrichum, however, did not respond to treatments as expected based on its seed size. High relative growth rate possibly increased Symphyotrichum seedling establishment in competitive environments. Results of this study demonstrated recruitment limitation when seedlings established into existing vegetation, and suggest that species composition resulting from specific seeding rates during prairie restoration may be more predictable when species are interseeded into existing vegetation, than when seeds are sown into tilled fields.     

Interactive effects of mycorrhizae and a root hemiparasite on plant community productivity and diversity

Plant communities can be affected both by arbuscular mycorrhizal fungi (AMF) and hemiparasitic plants. However, little is known about the interactive effects of these two biotic factors on the productivity and diversity of plant communities. To address this question, we set up a greenhouse study in which different AMF inocula and a hemiparasitic plant (Rhinanthus minor) were added to experimental grassland communities in a fully factorial design. In addition, single plants of each species in the grassland community were grown with the same treatments to distinguish direct AMF effects from indirect effects via plant competition. We found that AMF changed plant community structure by influencing the plant species differently. At the community level, AMF decreased the productivity by 15–24%, depending on the particular AMF treatment, mainly because two dominant species, Holcus lanatus and Plantago lanceolata, showed a negative mycorrhizal dependency. Concomitantly, plant diversity increased due to AMF inoculation and was highest in the treatment with a combination of two commercial AM strains. AMF had a positive effect on growth of the hemiparasite, and thereby induced a negative impact of the hemiparasite on host plant biomass which was not found in non-inoculated communities. However, the hemiparasite did not increase plant diversity. Our results highlight the importance of interactions with soil microbes for plant community structure and that these indirect effects can vary among AMF treatments. We conclude that mutualistic interactions with AMF, but not antagonistic interactions with a root hemiparasite, promote plant diversity in this grassland community. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Resource limitation,competition and the influence of life history in a freshwater snail community

Summary Previous work on a snail community occurring throughout lakes in southwestern Michigan showed that predation by molluscivorous sunfish had large impacts on only the rarest snail species. Thus, competition might play a major role in population limitation because dominant members of the snail community are relatively immune to predation. The present experiments were conducted to determine the extent to which the snail community depleted the abundance of food resources (epiphytes) and the extent to which epiphyte abundances limited snail production. An experimental gradient in snail densities showed that removal of snails increased epiphyte biomass by approximately 3-fold relative to that observed at natural snail densities. Enrichment of the environment with phosphorus fertilizer increased epiphyte biomass by approximately 20-fold and provided a test of food limitation in the snail community. All snail taxa exhibited positive numerical or growth responses to enrichment. The observations that snails depleted resources and that resources limited snail production demonstrated that snails competed exploitatively for epiphytes. The response of each snail species to increased food abundance differed depending on the timing of fertilization relative to the snails' life histories. Snails hatched before the experiment began were larger in fertilized treatments, due to increased growth, but their densities were similar among treatments. On the other hand, densities of snails born during the experiment were up to 15-fold greater in fertilized treatments, due in part to increased survival of newborn snails. Comparison of the responses of snails to food addition and to predator removals (based on prior experiments) suggested that food availability limits snail production more than predators do. Additionally, the large responses by algae and snails to fertilization demonstrated that both the producers and herbivores in this simplified food chain were strongly resource limited.     

Evaluation of indicators of nitrogen limitation in deep prairie lakes with laboratory bioassays and limnocorrals

A comparison of Kenyan reefs of different historical and observed levels of fishing exploitation showed that more exploited reef lagoons had greater sea urchin densities and sizes, fewer and smaller fish and less coral cover. In the most exploited lagoon the biomass of the burrowing sea urchin Echinometra mathaei increased five fold during the previous 15 years. An ecological study of the three most common omnivorous sea urchin species inhabiting hard substrate within these reef lagoons (E. mathaei, Diadema savignyi and D. setosum) suggests that they are ecologically separated by predation and avoid predators and competitors by occupying different size burrows or crevices within the lagoon. Predator removal through fishing activities may result in ecological release of the sea urchins and result in competitive exclusion of weaker competitors. The most exploited reef had a nearly monospecific barren of E. mathaei living outside burrows suggesting that E. mathaei may be the top competitor. Its ecological release appears to lead to a decrease in live coral cover, increased substrate bioerosion and eventually a loss of topographic complexity, species diversity, fish biomass and utilizable fisheries productivity. Data from the outer reef edge were more difficult to interpret but may indicate similar patterns. Within this area, physical stresses such as waves and currents may be a greater controlling force in regulating fishing activities and coral reef community structure.     

Spatial variation in seed limitation of plant species richness and population sizes in floodplain tallgrass prairie

The relative importance of seed availability versus biotic interactions that affect early life stages in limiting plant population sizes and determining composition of plant communities is a central debate in plant ecology. We conducted a seed addition experiment in restored tallgrass prairie in central Kansas to determine (1) whether addition of seed of 18 native forb species produced persistent (three growing seasons) increases in the species' population sizes and plant species richness, (2) what properties of recipient communities best explained spatial variation in added species' establishment, and (3) whether seed size explained interspecific patterns in establishment success. Adding seed led to persistent increases in the number of added species present and in plant species richness at one of three sites. Increased species richness at the one site where community composition was structured by seed availability largely resulted from greater densities of four species. Seed size did not predict species' establishment success. Pre-existing plant species richness was correlated with added species' establishment success, but the direction of the relationship (positive vs. negative) varied among sites. Living aboveground plant biomass in experimental plots in the year of seed addition was negatively correlated with the number of added species established three years later. Our results provide further evidence for large spatial variation in seed limitation of plant community composition. Surprisingly, mean light availability and heterogeneity in light, both important parameters in conceptual models of grassland plant coexistence, did not predict the response of the recipient plant community to seed addition as well as pre-existing plant species richness and living aboveground biomass.     

Rate of succession in restored wetlands and the role of site context

Question: Are changes in plant species composition, functional group composition and rates of species turnover consistent among early successional wetlands, and what is the role of landscape context in determining the rate of succession? Location: Twenty‐four restored wetlands in Illinois, USA. Methods: We use 4 years of vegetation sampling data from each site to describe successional trends and rates of species turnover in wetlands. We quantify: (1) the rate at which composition changes from early‐successional to late‐successional species and functional groups, as indicated by site movement in ordination space over time, and (2) the rate of change in the colonization and local extinction of individual species. We correlate the pace of succession to site area, isolation and surrounding land cover. Results: Some commonalities in successional trends were evident among sites. Annual species were replaced by clonal perennials, and colonization rates declined over time. However, differences among sites outweighed site age in determining species composition, and the pace of succession was influenced by a site's landscape setting. Rates of species turnover were higher in smaller wetlands. In addition, wetlands in agricultural landscapes underwent succession more rapidly, as indicated by a rapid increase in dominance by late‐successional plants. Conclusions: Although the outcome of plant community succession in restored wetlands was somewhat predictable, species composition and the pace of succession varied among sites. The ability of restoration practitioners to accelerate succession through active manipulation may be contingent upon landscape context.     

Effects of ungulates and prairie dogs on seed banks and vegetation in a North American mixed-grass prairie

The relationship between vegetation cover and soil seed banks was studied in five different ungulate herbivore-prairie dog treatment combinations at three northern mixed-grass prairie sites in Badlands National Park, South Dakota. There were distinct differences in both the seed bank composition and the aboveground vegetation between the off-prairie dog colony treatments and the on-colony treatments. The three on-colony treatments were similar to each other at all three sites with vegetation dominated by the forbs Dyssodia papposa, Hedeoma spp., Sphaeralcea coccinea, Conyza canadensis, and Plantago patagonica and seed banks dominated by the forbs Verbena bracteata and Dyssodia papposa. The two off-colony treatments were also similar to each other at all three sites. Vegetation at these sites was dominated by the grasses Pascopyrum smithii, Bromus tectorum and Bouteloua gracilis and the seed banks were dominated by several grasses including Bromus tectorum, Monroa squarrosa, Panicum capillare, Sporobolus cryptandra and Stipa viridula. A total of 146 seedlings representing 21 species germinated and emerged from off-colony treatments while 3069 seedlings comprising 33 species germinated from on-colony treatments. Fifteen of the forty species found in soil seed banks were not present in the vegetation, and 57 of the 82 species represented in the vegetation were not found in the seed banks. Few dominant species typical of mixed-grass prairie vegetation germinated and emerged from seed banks collected from prairie dog colony treatments suggesting that removal of prairie dogs will not result in the rapid reestablishment of representative mixed-grass prairie unless steps are taken to restore the soil seed bank.     

Plant distributions on a saskatchewan prairie

Summary An intensive study of plant species distributions on a Saskatchewan prairie is reported here. Frequency data for fifty-nine vascular species and varieties are presented in association with environmental data relating to soil moisture, water retaining capacity, soil pH and slope position. Secondary consideration is given to light, which is important in the distribution of the understory in dense shrub stands, and soil texture. Soil moisture is shown to be the main determinant of plant species distributions.     

Assessing bee (Hymenoptera: Apoidea) diversity of an Illinois restored tallgrass prairie: methodology and conservation considerations

Bee species diversity and the effectiveness of four sampling methods were investigated in a west-central Illinois restored tallgrass prairie. Bees were sampled using malaise traps, ground-level pan traps, elevated pan traps, and vane traps. A total of 4,622 bees representing 31 genera and 111 species were collected. Malaise traps collected the greatest number of bees and species, and ground-level pan traps the least. Among the pan traps and vane traps, blue-colored traps collected the greatest abundance and species richness, and yellow traps the least. Chao1 estimator and rarefaction analyses showed that substantial increases in sample sizes would be necessary to achieve asymptotic species richness levels, particularly if ground-level pan traps alone were used. Elevated pan traps and vane traps collected relatively similar species composition. Different colored pan traps at the same height collected more similar species composition than did those at different heights, but species composition of blue ground-level pan traps was relatively similar to elevated pan traps, regardless of color. Indicator species analysis revealed 22 species that were significantly associated with a specific trap type, and 11 species that were associated with a particular pan trap color/elevation. Results of this study show that elevated traps can increase the effectiveness of bee surveys in tallgrass prairie, and that a combination of trap types gives a more complete picture of the bee fauna than does a single survey method. These results should be considered along with cost, ease of use, and goals when planning and designing bee inventories.     

Resource limitation and the lethal and sublethal effects of a viral pathogen in the Indian meal moth, Plodia interpunctella

Abstract.

• 1 The effects of resource limitation and the lethal and sublethal effects of a granulosis virus on a lepidopteran host, the Indian meal moth, Plodia interpunctella, were examined.
• 2 The food quality was manipulated by the addition of an inert bulking agent (methyl cellulose) which caused the size, development rate and fecundity of the moths to be reduced.
• 3 The resource quality had no effect on the mortality due to the virus. In contrast, sublethal effects of the virus on pupal weight were more apparent under conditions of resource limitation.
• 4 Considerable variation between the sublethal effects after challenge with different doses of the virus was found. The balance between deleterious sublethal effects of the virus and the selection of more robust individuals by the bioassays is proposed as a mechanism to explain this variation.
• 5 Implications for the dynamics of insect hosts and their pathogens are discussed.