Nitrogen fertilization and water supply affect germination and plant establishment of the soil seed bank present in a semi-arid Mediterranean scrubland

Anthropogenic nitrogen (N) inputs in terrestrial ecosystems are higher than those that occur naturally and have been related to global biodiversity loss and altered ecosystem functioning. However, its effects on Mediterranean-type ecosystems, where production is water-limited and N regulated, remain unclear. We conducted a green-house experiment where we evaluated the effects of four simulated scenarios of N pollution (0, 10, 20 and 50 kg N ha⁻¹ year⁻¹) and two differential water supply regimes on the germination (experiment 1) and early plant establishment (experiment 2) of a seed bank from a semi-arid Mediterranean ecosystem of central Spain. Seed bank density was estimated as 62,374 ± 3,279 seeds m⁻². Approximately 99.5% of emerged seeds corresponded to only 14 species of a total of 52, the majority of which were the annual forb Sagina apetala. The responses for N treatments were species-specific, mainly positive or unimodal, with watering treatments having some interactive effects. N and water supply also affected total and specific productivity; the responses found for N treatments were mainly humpback-shaped and an increased water supply had additive effects on community establishment in terms of total plant biomass. This response was linked to forb responsiveness. Contrary to predictions, grass biomass did not change with N supply; however, grass to forb ratio was affected because of changes in the latter. Overall, these experiments suggest a critical load for plant biomass production and conclude that N and water availability and supply can modify germination and plant establishment. This should be taken into account when analysing the effects of global change on the dynamics of plant communities where annuals are dominant or vegetation must establish from seed following a natural or anthropogenic disturbance regime.     

Nitrogen limitation, 15N tracer retention, and growth response in intact and Bromus tectorum-invaded Artemisia tridentata ssp. wyomingensis communities

Annual grass invasion into shrub-dominated ecosystems is associated with changes in nutrient cycling that may alter nitrogen (N) limitation and retention. Carbon (C) applications that reduce plant-available N have been suggested to give native perennial vegetation a competitive advantage over exotic annual grasses, but plant community and N retention responses to C addition remain poorly understood in these ecosystems. The main objectives of this study were to (1) evaluate the degree of N limitation of plant biomass in intact versus B. tectorum-invaded sagebrush communities, (2) determine if plant N limitation patterns are reflected in the strength of tracer ¹⁵N retention over two growing seasons, and (3) assess if the strength of plant N limitation predicts the efficacy of carbon additions intended to reduce soil N availability and plant growth. Labile C additions reduced biomass of exotic annual species; however, growth of native A. tridentata shrubs also declined. Exotic annual and native perennial plant communities had divergent responses to added N, with B. tectorum displaying greater ability to use added N to rapidly increase aboveground biomass, and native perennials increasing their tissue N concentration but showing little growth response. Few differences in N pools between the annual and native communities were detected. In contrast to expectations, however, more ¹⁵N was retained over two growing seasons in the invaded annual grass than in the native shrub community. Our data suggest that N cycling in converted exotic annual grasslands of the northern Intermountain West, USA, may retain N more strongly than previously thought.     

Soil drying and nitrogen availability modulate carbon and water exchange over a range of annual precipitation totals and grassland vegetation types

Increased intensity in precipitation events and longer periods of water deficit are predicted as a general trend under future climate scenarios with potentially large effects on terrestrial ecosystem function. The primary objective of this study was to understand how variation in the intensity of precipitation inputs followed by intermittent soil drying events influence leaf and ecosystem carbon dioxide (CO₂) and water exchange in a California annual grassland mesocosm experiment. We further examined how nitrogen (N) availability, and differences in plant community composition (grass-forb combinations) affected gas exchange responses to the precipitation treatments. Net ecosystem CO₂ exchange (NEE) and evapotranspiration (ET) increased significantly with greater precipitation and were positively correlated with soil moisture. A repeated 10-day soil drying period, following 11 days of watering, strongly depressed NEE over a range of annual precipitation totals (297, 657 and 987 mm), and plant community types. Ecosystem dark respiration ( R _e) and leaf level photosynthesis ( A _max) showed greater sensitivity to periods of soil drying in the low precipitation plots (297 mm). N additions significantly increased NEE and R _e, particularly as water availability was increased. Across the range of precipitation totals and plant community types, intermittent periods of soil moisture deficit and native soil N availability constrained leaf and ecosystem level CO₂ exchange, while the influence on water vapor exchange was less pronounced.     

Repeated mowing to restore remnant native grasslands invaded by nonnative annual grasses: upsides and downsides above and below ground

California grasslands have been severely impacted by the invasion of nonnative annual grasses, which often limit restoration of this important ecosystem. In this study, we explored the use of mowing as a restoration tool for native perennial grasslands at the Santa Rosa Plateau Ecological Reserve in southern California. We sought to evaluate if, over time, mowing would reduce nonnative annual grass cover and benefit native species, especially the native bunchgrass Stipa pulchra. We hypothesized that repeated mowing, carefully timed to target nonnative annual grasses prior to seed maturation, would reduce nonnative seed inputs into the soil and eventually lead to diminished abundance of these species. We monitored vegetation in mowed and unmowed plots for 4 years, and conducted a seed bank study after 5 years to better understand the cumulative effects of mowing on native and nonnative seed inputs. Consistent with our hypotheses, we found that mowing successfully reduced nonnative annual grass cover and benefitted some native species, including S. pulchra. However, we also found that nonnative forb species showed progressive increases in mowed plots over time. We observed similar patterns of species composition in the soil seed bank. Together, these results suggest that mowing can be used to control nonnative annual grasses and increase the abundance of native bunchgrasses, but that this method may also have the unintended consequence of increasing nonnative forb species.     

Combined effects of precipitation and nitrogen deposition on native and invasive winter annual production in California deserts

Primary production in deserts is limited by soil moisture and N availability, and thus is likely to be influenced by both anthropogenic N deposition and precipitation regimes altered as a consequence of climate change. Invasive annual grasses are particularly responsive to increases in N and water availabilities, which may result in competition with native forb communities. Additionally, conditions favoring increased invasive grass production in arid and semi-arid regions can increase fire risk, negatively impacting woody vegetation that is not adapted to fire. We conducted a seeded garden experiment and a 5-year field fertilization experiment to investigate how winter annual production is altered by increasing N supply under a range of water availabilities. The greatest production of invasive grasses and native forbs in the garden experiment occurred under the highest soil N (inorganic N after fertilization = 2.99 g m^?2) and highest watering regime, indicating these species are limited by both water and N. A classification and regression tree (CART) analysis on the multi-year field fertilization study showed that winter annual biomass was primarily limited by November–December precipitation. Biomass exceeded the threshold capable of carrying fire when inorganic soil N availability was at least 3.2 g m^?2 in piñon-juniper woodland. Due to water limitation in creosote bush scrub, biomass exceeded the fire threshold only under very wet conditions regardless of soil N status. The CART analyses also revealed that percent cover of invasive grasses and native forbs is primarily dependent on the timing and amount of precipitation and secondarily dependent on soil N and site-specific characteristics. In total, our results indicate that areas of high N deposition will be susceptible to grass invasion, particularly in wet years, potentially reducing native species cover and increasing the risk of fire.     

Germination,survival, and growth of grass and forb seedlings: Effects of soil moisture variability

Seed germination and seedling growth, survivorship, and final biomass and their responses to watering interval were studied in two grass and six forb species to assess germination and seedling growth responses to increased soil moisture variability as might occur with future increases in precipitation variability. Seeds were planted in prairie soil and watered at 1, 2, 4, or 7 d intervals (I). Seed germination peaked at I = 4 d whereas leaf growth in grasses and forbs, and final biomass in grasses peaked at I = 7 d, suggesting that growth and biomass were favored at greater soil moisture variability than seed germination. Biomass responses to I were stronger than the germination responses, suggesting that soil moisture variability more strongly influenced post germination growth. Individual species responses to I fell into three groups; those with responses to I for: (1) seed germination and seedling survival, (2) biomass, or (3) both germination and biomass production. These species groups may be more useful than life form (i.e., grass/forb) for understanding seed germination and seedling dynamics in grasslands during periods of soil moisture variability. Seed germination and early growth may assume more importance in grassland plant community dynamics under more variable precipitation patterns.     

Invertebrate,not small vertebrate,herbivory interacts with nutrient availability to impact tallgrass prairie community composition and forb biomass

The effects of herbivores and their interactions with nutrient availability on primary production and plant community composition in grassland systems is expected to vary with herbivore type. We examined the effects of invertebrate and small vertebrate herbivores and their interactions with nutrient availability on grassland plant community composition and aboveground biomass in a tallgrass prairie ecosystem. The abundance of forbs relative to grasses increased with invertebrate herbivore removals. This increase in forb abundance led to a shift in community composition, where invertebrate removals resulted in greater plant species evenness as well as a divergence in composition among plots. In contrast, vertebrate herbivore removals did not affect plant community composition or aboveground biomass. Nutrient additions alone resulted in a decrease in plant species richness and an increase in the abundance of the dominant grass, but the dominant grass species did not greatly increase in abundance when nutrient additions were combined with invertebrate removals. Rather, several subdominant forbs came to dominate the plant community. Additionally, the combined nutrient addition and invertebrate herbivore removal treatment increased forb biomass, suggesting that invertebrate herbivores suppress the responses of forb species to chronic nutrient additions. Overall, the release of forbs from invertebrate herbivore pressure may result in large shifts in species composition, with consequences for aboveground biomass and forage quality due to altered grass:forb ratios in grassland systems.     

Nutrient Limitation in a Fire‐derived,Nitrogen‐rich Hawaiian Grassland1

Grasslands created by grass invasions into shrublands or woodlands followed by fire are now a dominant feature of many seasonally dry environments. In Hawaii Volcanoes National Park, introduced perennial grasses dominate grasslands created by fire in grass‐invaded woodlands. In a previous study, we found that net primary production in these grasslands is substantially lower than in unburned woodlands. Yet, our estimates of annual net nitrogen (N) mineralization showed higher rates in these savannas than in the unburned woodlands, rates that appear to greatly exceed annual N demand by the vegetation. We therefore hypothesized that N should not be limited to the plants growing in these sites. We tested this hypothesis with a 2‐yr fertilization experiment. At peak biomass, we found a 30 percent increase in live biomass in plots with N added and no increase in production with only phosphorus (P) added. N and P together were synergistic, suggesting that co‐limitation or P limitation becomes important when N is more available. Plants responded to added N by increasing individual leaf area and shoot length by over 50 percent. Tissue N was higher with added N; hence, biomass N was substantially higher. Tissue P concentrations declined with N addition but were elevated by P addition despite lack of a growth response to P alone. Overall, N limitation exists despite high annual rates of net N mineralization, and co‐limitation of production by P may occur when N is abundant. Here, asynchrony between plant nutrient demand and N availability may contribute to N limitation.     

Soil microbial properties and plant growth responses to carbon and water addition in a temperate steppe: the importance of nutrient availability

Background

Global climatic change is generally expected to stimulate net primary production, and consequently increase soil carbon (C) input. The enhanced C input together with potentially increased precipitation may affect soil microbial processes and plant growth.

Methodology/Principal Findings

To examine the effects of C and water additions on soil microbial properties and plant growth, we conducted an experiment lasting two years in a temperate steppe of northeastern China. We found that soil C and water additions significantly affected microbial properties and stimulated plant growth. Carbon addition significantly increased soil microbial biomass and activity but had a limited effect on microbial community structure. Water addition significantly increased soil microbial activity in the first year but the response to water decreased in the second year. The water-induced changes of microbial activity could be ascribed to decreased soil nitrogen (N) availability and to the shift in soil microbial community structure. However, no water effect on soil microbial activity was visible under C addition during the two years, likely because C addition alleviated nutrient limitation of soil microbes. In addition, C and water additions interacted to affect plant functional group composition. Water addition significantly increased the ratio of grass to forb biomass in C addition plots but showed only minor effects under ambient C levels. Our results suggest that soil microbial activity and plant growth are limited by nutrient (C and N) and water availability, and highlight the importance of nutrient availability in modulating the responses of soil microbes and plants to potentially increased precipitation in the temperate steppe.

Conclusions/Significance

Increased soil C input and precipitation would show significant effects on soil microbial properties and plant growth in the temperate steppe. These findings will improve our understanding of the responses of soil microbes and plants to the indirect and direct climate change effects.     

Patch identity and the spatial heterogeneity of woody encroachment in exotic-dominated old-field grasslands

Most grassland communities in agricultural landscapes comprise a mix of exotic and native plants, where grasses and forbs are disposed in low diversity patches conforming a heterogeneous matrix of vegetation. Within these “novel” ecosystems, woody encroachment is one of the principal causes of ecosystem degradation. Here, we examined the resistance to exotic woody establishment (Gleditsia triacanthos) into four different monospecific patches characteristics of old-field grasslands in Inland Pampa: an annual forb (Conium maculatum), an annual grass (Lolium multiflorum), and two perennial grasses (Cynodon dactylon and Festuca arundinacea). We evaluated the filter to tree recruitment by rodent seed removal and survival and growth of Gleditsia seedlings transplanted into undisturbed and disturbed microsites, within each patch. Beneath intact vegetation seed removal was an important biotic filter to woody establishment whereas disturbances facilitated seed survival in patches of perennial grasses. Patch identity affected tree growth, and Cynodon reduced the final biomass compared to forbs. Disturbance enhanced tree performance independently of patch type. After 2 years, tree survival was independent of disturbance and patch identity. As patch identity may regulate granivory and growth of tree saplings, community susceptibility or resistance to woody invasion rather than representing a static community attribute could vary according to the dynamic changes in the proportion of susceptible-resistant patches. Broadly, our work reinforces the concept that mechanisms regulating vegetation heterogeneity add a component of stochasticity to biotic resistance to community plant invasion.     

Response of ecosystem carbon exchange to warming and nitrogen addition during two hydrologically contrasting growing seasons in a temperate steppe

A large remaining source of uncertainty in global model predictions of future climate is how ecosystem carbon (C) cycle feedbacks to climate change. We conducted a field manipulative experiment of warming and nitrogen (N) addition in a temperate steppe in northern China during two contrasting hydrological growing seasons in 2006 [wet with total precipitation 11.2% above the long‐term mean (348 mm)] and 2007 (dry with total precipitation 46.7% below the long‐term mean). Irrespective of strong intra‐ and interannual variations in ecosystem C fluxes, responses of ecosystem C fluxes to warming and N addition did not change between the two growing seasons, suggesting independence of warming and N responses of net ecosystem C exchange (NEE) upon hydrological variations in the temperate steppe. Warming had no effect on NEE or its two components, gross ecosystem productivity (GEP) and ecosystem respiration (ER), whereas N addition stimulated GEP but did not affect ER, leading to positive responses of NEE. Similar responses of NEE between the two growing seasons were due to changes in both biotic and abiotic factors and their impacts on ER and GEP. In the wet growing season, NEE was positively correlated with soil moisture and forb biomass. Negative effects of warming‐induced water depletion could be ameliorated by higher forb biomass in the warmed plots. N addition increased forb biomass but did not affect soil moisture, leading to positive effect on NEE. In the dry growing season, NEE showed positive dependence on grass biomass but negative dependence on forb biomass. No changes in NEE in response to warming could result from water limitation on both GEP and ER as well as little responses of either grass or forb biomass. N addition stimulated grass biomass but reduced forb biomass, leading to the increase in NEE. Our findings highlight the importance of changes in abiotic (soil moisture, N availability) and biotic (growth of different plant functional types) in mediating the responses of NEE to climatic warming and N enrichment in the semiarid temperate steppe in northern China.     

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.     

Mechanisms underlying the impacts of exotic annual grasses in a coastal California meadow

Biological invasions can impact the abundance and diversity of native species, but the specific mechanisms remain poorly discerned. In California grasslands, invasion by European annual grasses has severely reduced the quality of habitat for native forb species. To understand how introduced grasses suppress native and exotic forbs, we examined the response of a Southern California grassland community to factorial removals of live grass and the litter produced in previous seasons. To examine the role that belowground competition for water plays in mediating the impact of grasses, we crossed grass and litter removal treatments with water addition. Our results show that forbs were almost equally suppressed by both competition from live grass and direct interference by litter. Water addition did not ameliorate the effect of grass competition, suggesting that water was not the resource for which plants compete. This evidence is consistent with the susceptibility of forbs to light limitation, especially considering that litter does not consume water or nutrients. Interestingly, despite different histories of co-occurrence with annual grass dominants, native and exotic forbs were comparably suppressed by exotic grasses. Our results indicate that suppression by both live and dead stems underlie the influence of exotic grasses on forb competitors.     

Dominant Grasses Suppress Local Diversity in Restored Tallgrass Prairie

Warm‐season (C₄) grasses commonly dominate tallgrass prairie restorations, often at the expense of subordinate grasses and forbs that contribute most to diversity in this ecosystem. To assess whether the cover and abundance of dominant grass species constrain plant diversity, we removed 0, 50, or 100% of tillers of two dominant species (Andropogon gerardii or Panicum virgatum) in a 7‐year‐old prairie restoration. Removing 100% of the most abundant species, A. gerardii, significantly increased light availability, forb productivity, forb cover, species richness, species evenness, and species diversity. Removal of a less abundant but very common species, P. virgatum, did not significantly affect resource availability or the local plant community. We observed no effect of removal treatments on critical belowground resources, including inorganic soil N or soil moisture. Species richness was inversely correlated with total grass productivity and percent grass cover and positively correlated with light availability at the soil surface. These relationships suggest that differential species richness among removal treatments resulted from treatment induced differences in aboveground resources rather than the belowground resources. Selective removal of the dominant species A. gerardii provided an opportunity for seeded forb species to become established leading to an increase in species richness and diversity. Therefore, management practices that target reductions in cover or biomass of the dominant species may enhance diversity in established and grass‐dominated mesic grassland restorations.     

Invasion by <Emphasis Type="Italic">Aegilops triuncialis</Emphasis> (Barb Goatgrass) Slows Carbon and Nutrient Cycling in a Serpentine Grassland

Invasive plant species alter plant community composition and ecosystem function. In the United States, California native grasslands have been displaced almost completely by invasive annual grasses, with serpentine grasslands being one of the few remaining refugia for California grasslands. This study examined how the invasive annual grass, Aegilops triuncialis, has altered decomposition processes in a serpentine annual grassland. Our objectives were to (1) assess howA. triuncialis alters primary productivity and litter tissue chemistry, (2) determine whether A. triuncialis litter is more recalcitrant to decomposition than native litter, and (3) evaluate whether differences in the soil microbial community in A. triuncialis-invaded and native-dominated areas result in different decomposition rates of invasive and/or native plant litter. In invaded plant patches, A. triuncialis was approximately 50% of the total plant cover, in contrast to native plant patches in which A. triuncialis was not detected and native plants comprised over 90% of the total plant cover. End-of-season aboveground biomass was 2-fold higher in A. triuncialis dominated plots compared to native plots; however, there was no significant difference in belowground biomass. Both above- and below-ground plant litter from A. triuncialis plots had significantly higher lignin:N and C:N ratios and lower total N, P, and K than litter from native plant plots. Aboveground litter from native plots decomposed more rapidly than litter from A. triuncialis plots, although there was no difference in decomposition of belowground tissues. Soil microbial community composition associated with different soil patch types had no effect on decomposition rates. These data suggest that plant invasion impacts decomposition and nutrient cycling through changes in plant community tissue chemistry and biomass production.     

环青海湖地区天然草地和退耕恢复草地植物群落生物量对氮、磷添加的响应

植物群落生物量反映了植被的初级生产能力, 是陆地生态系统碳(C)输入的最主要来源, 往往受到自然界中氮(N)、磷(P)元素供应的限制。该试验以青藏高原环青海湖地区的高寒草原为研究对象, 探讨了天然草地和退耕恢复草地植被群落生物量对N (10 g·m^-2)、P (5 g·m^-2)养分添加的响应。N、P添加显著增加了天然草地禾草的生物量, 进而促使地上总生物量显著提高。退耕恢复草地禾草和杂类草的生物量对N添加均有一致的正响应, 从而促使地上总生物量显著增加174%, 群落地上和地下总生物量显著增加34%; 而P添加对恢复草地生物量各项参数均无显著影响。回归分析显示: 天然草地植物群落地上生物量随土壤中NO₃^--N含量的增加而增加(p < 0.05), 退耕恢复草地植被地上、地下和总生物量均与土壤NO₃^--N含量显著正相关(p < 0.01), 说明环湖地区高寒草原植物生长主要受N供应的限制, P的限制作用随土地利用方式的转变和群落演替阶段的不同而变化; 相比天然草地, 恢复草地在现阶段植被初级生产力受N的限制作用更强烈, 土壤中可利用N含量是限制其植被自然恢复和重建的关键因子。     

Responses of plant community biomass to nitrogen and phosphorus additions in natural and restored grasslands around Qinghai Lake Basin

Aims Plant biomass reflects the primary productivity of community vegetation, and is the main resource of carbon input in the terrestrial ecosystem. It is usually limited by nitrogen (N) and phosphorus (P) availability in the soil. Alpine grassland around Qinghai Lake Basin has experienced extensive land-use changes due to the cultivation of native grassland and vegetation recovery on cropped land. In this experiment, two grassland types were chosen, natural alpine grassland (NG) and its adjacent restored grassland (RG), to determine the responses of plant community biomass to N and P additions with different land-use. Methods NH₄NO₃ and Ca(H₂PO₄)₂·H₂O were added in a completely randomized block design, with medium levels of 10 g N·m^-2 and 5 g P·m^-2. Soil NO₃^--N and available P contents, and the plant community biomass were measured in the two grasslands. Two-way ANOVA was used to determine the effects of nutrient additions on all measured indicators, and regression analysis was used to analyze the correlations between plant biomass and soil NO₃^--N and available P contents.Important findings Results showed: (1) N and P additions both increased grass biomass in the NG, and significantly elevated the total aboveground biomass, with the promoting effect of N addition higher than that of P addition; N addition significantly increased both grass and forb biomass in the RG, and markedly promoted the total aboveground biomass, while P addition had no effects on the functional groups and total aboveground biomass (p > 0.05). (2) N and P additions both had no effects on the belowground and total biomass in the NG, whereas N addition significantly increased the total biomass by 34% in the RG, which suggested that the effect of N limitation on the vegetation primary productivity was stronger in the RG at present stage. (3) The aboveground biomass in the NG increased with soil NO₃^--N content (p < 0.05), and the above- and below-ground as well as the total biomass were all positively correlated with soil NO₃^--N content in the RG (p < 0.01). These results indicated that the plant growth in alpine grassland around Qinghai Lake Basin was prone to N limitation, and the effect of P limitation changed with land-use. Soil available N might be the key limiting factor for vegetation restoration and reconstruction in the RG. The “Grain for Green” project (the land-use policy) and atmospheric N deposition are benefiting both plant growth and C accumulation in the alpine grassland ecosystem around Qinghai Lake Basin.     

Effects of drought and N level on the interactions of the root hemiparasite Rhinanthus alectorolophus with a combination of three host species

• Increasing nitrogen deposition and more frequent drought events are likely to change plant interactions in natural grasslands. Both factors may also influence the interactions between hemiparasitic plants, regarded as keystone species in many grasslands, and their host species.
• We grew a combination of three suitable hosts, a grass, a forb and a legume, with and without the hemiparasite Rhinanthus alectorolophus at three levels of nitrogen (N) and two levels of water availability in a factorial design.
• Biomass of the hemiparasite and host community increased with N level and was reduced by drought to a similar degree. Larger plants in fertilised pots started to wilt earlier, and the presence of a hemiparasite further increased drought sensitivity. The hemiparasite strongly reduced biomass of the host community and overall productivity, and affected the competitive balance among host plants because it particularly reduced biomass of the dominant grass. These effects were the opposite of those of high N. The hemiparasite increased the root mass fraction of the hosts at all levels of N and water availability, indicating that the effect of the hemiparasite on the hosts was mainly due to loss of belowground resources.
• Our results indicate that hemiparasites will not always respond more strongly to increased N availability and drought than autotrophic plants, and that hemiparasites can have similarly strong effects on grassland communities as soil fertility and drought. By preferentially attacking dominant species the hemiparasites might alleviate the negative effects of nutrient enrichment on grassland diversity.

     

Frequency and Extent of Water Limitation to Primary Production in a Mesic Temperate Grassland

The frequency and extent of water limitation to aboveground net primary production (ANPP) in a mesic grassland in NE Kansas (Konza Prairie, USA) was assessed with an 8-year irrigation experiment. Since 1991, transects spanning upland and lowland sites in annually burned, ungrazed tallgrass prairie were provided with supplemental water to satisfy evapotranspirational demands. This protocol minimized water limitations during the growing season, as well as interannual variability in water stress. Irrigation of this mesic grassland increased ANPP in 6 of 8 years by an average of 26% when compared to control transects. Although interannual variation in ANPP was greater in uplands than lowlands at nominal levels of precipitation, reducing interannual variability in water availability via irrigation eliminated topographic differences; the irrigation protocol also reduced interannual variability in ANPP by as much as 40%. The addition of supplemental water enabled us to extend the relationship between annual precipitation and ANPP in grasslands to precipitation levels (average, 1153 mm; maximum, 1346 mm) similar to those experienced by more mesic grasslands that today exist only as remnants several hundred kilometers east of Kansas. This relationship was linear (r ²= 0.81), with maximum ANPP (738 g/m²) similar to values reported for sites in Illinois and Wisconsin. After 8 years of irrigation, production of the C₃ forb component was twice that in control sites. These results indicate that water limitations in grasslands at the western edge of the presettlement extent of tallgrass prairie affect ANPP in most years and that this high frequency of water limitation may lead to greater dominance of the C₄ grasses than is seen in more eastern grassland sites. Received 18 January 2000; accepted 19 July 2000.