Species effects on nitrogen cycling: a test with perennial grasses

Summary To test for differing effects of plant species on nitrogen dynamics, we planted monocultures of five perennial grasses (Agropyron repens, Agrostis scabra, Poa pratensis, Schizachyrium scoparium, and Andropogon gerardi) on a series of soils ranging from sand to black soil. In situ net N mineralization was measured in the monocultures for three years. By the third year, initially identical soils under different species had diverged up to 10-fold in annual net mineralization. This divergence corresponded to differences in the tissue N concentrations, belowground lignin concentrations, and belowground biomasses of the species. These results demonstrate the potential for strong feedbacks between the species composition of vegetation and N cycling. If individual plant species can affect N mineralization and N availability, then competition for N may lead to positive or negative feedbacks between the processes controlling species composition and ecosystem processes such as N and C cycling. These feedbacks create the potential for alternative stable states for the vegetation-soil system given the same initial abiotic conditions.     

Capture and allocation of nitrogen byQuercus douglasii seedlings in competition with annual and perennial grasses

Summary The spatial overlap of woody plant root systems and that of annual or perennial grasses promotes competition for soil-derived resources. In this study we examined competition for soil nitrogen between blue oak seedlings and either the annual grassBromus mollis or the perennial grassStipa pulchra under controlled outdoor conditions. Short-term nitrogen competition was quantified by injecting¹⁵N at 30 cm depth in a plane horizontal to oak seedling roots and that of their neighbors, and calculating¹⁵N uptake rates, pool sizes and¹⁵N allocation patterns 24 h after labelling. Simultaneously, integrative nitrogen competition was quantified by examining total nitrogen capture, total nitrogen pools and total nitrogen allocation.Stipa neighbors reduced inorganic soil nitrogen content to a greater extent than didBromus plants. Blue oak seedlings responded to lower soil nitrogen content by allocating lower amounts of nitrogen per unit of biomass producing higher root length densities and reducing the nitrogen content of root tissue. In addition, blue oak seedlings growing with the perennial grass exhibited greater rates of¹⁵N uptake, on a root mass basis, compensating for higher soil nitrogen competition inStipa neighborhoods. Our findings suggest that while oak seedlings have lower rates of nitrogen capture than herbaceous neighbors, oak seedlings exhibit significant changes in nitrogen allocation and nitrogen uptake rates which may offset the competitive effect annual or perennial grasses have on soil nitrogen content.     

Plant and soil nitrogen dynamics in California annual grassland

Seasonal changes in soil water and nitrogen availability were related to the phenology and growth of plants in California annual grassland. Plant accumulation of nitrogen was mainly confined to two short periods of the year: fall and early spring. At these times, plants were in the vegetative growth phase, roots were growing rapidly and soil moisture was high. During these periods, soil nitrate was low or depleted. High flux of nitrogen in this ecosystem, however, is indicated by the rapid disappearance of the previous year's detrital material, high microbial biomass, and high mineralizable nitrogen and nitrification potential.At the end of the summer drought, significant amounts of the previous year's detrital material had disappeared, chloroform-labile N (expressed as microbial biomass N) was at its seasonal maximum, and soil inorganic nitrogen pools were high. This suggests inorganic nitrogen flux during the drought period. The drought escaper life history characteristics of annual grasses in California annual grassland, however, may prevent plants from utilizing available nitrogen during a large part of the year.     

Reproductive effort and phenology of seed production of savanna grasses with different growth form and life history

The phenology of seed production in natural savanna grasslands was studied in the grass speciesAristida congesta, Cymbopogon plurinodis, Cynodon dactylon, Digitaria eriantha ssp.pentzii, Eragrostis rigidior, Eragrostis superba, Panicum coloratum, Schmidtia pappophoroides, Tragus berteronianus andUrochloa panicoides. Maximum seed production varied according to life history strategy and growth form from 0.03 mg seed g^-1 shoot dry weight in the perennialD. eriantha ssp.pentzii which produces long stolons and 14.8 mg seed g^-1 shoot inE rigidior, which produces short geniculate stolons, to 169.1 mg g^-1 in the annualT. berteronianus. Seed production was in most species divided over several peaks during the season. Peaks of seed production were observed 3 to 7 months after the onset of the growth season depending on the start of the rains and the life history strategy and growth form of the species. Seed production varied from maxima of 180 seeds m^-2 inD. eriantha ssp.pentzii to 47000 seeds m^-2 in annual stands ofT. berteronianus. Except for annual grasslands withU. panicoides, seedling emergence data reported are smaller by at least a factor of 10 than the observed seed production. Among other factors, a low quality of produced seeds, predation by birds and insects and previous grazing by livestock may have contributed to this difference.     

Rainfall seasonality determines annual/perennial grass balance in vegetation of Mediterranean Iberian

Much recent attention has been focused on the invasion and dominance of annual grass species in areas thought to have been historically dominated by perennial life forms. Explanations of this phenomenon in the literature have focused on two mechanisms favoring annuals: ruderal strategy associated with disturbance, and stress escaping associated with dry sites or deserts. Here I present evidence from vegetation surveys at 50 sites across a 1,200 km band of the Iberian Peninsula—a source region for many invasive annuals—showing that relative annual versus perennial grass composition is not well correlated with degree of disturbance or average annual precipitation. However, annual dominance is strongly and significantly linked to the seasonality of precipitation, in particular the relative intensity of summer drought. Disturbance was significantly associated with annual grass dominance in Iberia, but with much less explanatory power than summer drought intensity. Slope, aspect, and soil parent material were not significantly correlated with annual versus perennial grass dominance. These results suggest that subtle differences in rainfall seasonality largely drive grass composition in herbaceous Mediterranean vegetation. Furthermore, the patterns of annual grass invasion observed in the world’s other Mediterranean climate regions may be associated with similar climatic drivers.     

Contrasting impacts of grass species on nitrogen cycling in a grazed Sudanian savanna

We investigated the impact of perennial and annuals grass species on nitrogen cycling in a Sudanian savanna of Burkina Faso. We also analysed how the local context in terms of grazing and soil properties modifies these impacts. We selected four plots differing both by the intensity of grazing by cattle and soil depth, and used soil and grass biomass ¹⁵N as integrative indicators of N cycle. If perennials are able to foster a more efficient nitrogen cycling there should be lower ¹⁵N abundances in their biomass and soil. If soil depth and cattle pressure significantly modify nitrogen fluxes, soil depth and cattle pressure should influence ¹⁵N signatures. Our results suggest that perennial grasses are more conservative for nitrogen (inhibition of nitrification, less leaching via a perennial root system, slower cycling). The increase in leaf δ¹⁵N with N concentration is steeper in Loudetia togoensis than in the three other grasses. No significant difference was found between the ¹⁵N signatures of the four plots. Our results on ¹⁵N signatures and the fact that perennial grasses are much more abundant in the plots that are less grazed and have deeper soils, confirm that the switch from perennial to annual grasses is linked to a degradation in soil fertility and pasture quality. This suggests that ¹⁵N signatures can be used as indicators of fertility.     

Effect of soil nitrogen stress on the relative growth rate of annual and perennial grasses in the Intermountain West

A high relative growth rate (RGR) is thought to be an important trait allowing invasive annual grasses to exploit brief increases in nitrogen (N) supply following disturbance in the Intermountain West. Managing soils for low N availability has been suggested as a strategy that may reduce this growth advantage of annual grasses and facilitate establishment of desirable perennials grasses. The objective of this study was to examine the degree to which soil N availability affects RGR and RGR components of invasive annual and desirable perennial grasses. It was hypothesized that (1) invasive annual grasses would demonstrate a proportionately greater reduction in RGR than perennial grasses as soil N stress increased, and (2) the mechanism by which low N availability decreases RGR of annual and perennial grasses would depend on the severity of N stress, with moderate N stress primarily affecting leaf mass ratio (LMR) and severe N stress primarily affecting net assimilation rate (NAR). Three annual and three perennial grasses were exposed to three levels of N availability. RGR and components of RGR were quantified over four harvests. Moderate N stress reduced RGR by decreasing LMR and severe N stress lowered RGR further by decreasing NAR. However, reduction in RGR components was similar between invasive and natives, and as a consequence, annual grasses did not demonstrate a proportionately greater reduction in RGR than perennials under low N conditions. These results suggest managing soil N will do little to reduce the initial growth advantage of annual grasses. Once perennials establish, traits not captured in this short-term study, such as high tissue longevity and efficient nutrient recycling, may allow them to compete effectively with annuals under low N availability. Nevertheless, if soil N management does not facilitate the initial establishment of perennials in annual grass infested communities, then there is little likelihood that such techniques will provide a long-term benefit to restoration projects in these systems.     

Simulated nitrogen dynamics and nitrate leaching in a perennial grass ley

A soil nitrogen model was used for a 4-year simulation of nitrogen dynamics and nitrate leaching, both during grass ley growth and after ploughing a grass ley. Model results were compared with field measurements of soil mineral-N status and leaching. A soil water and heat model provided daily values for abiotic conditions, which were used as driving variables in the nitrogen simulation. Simulated values for mineral-N levels in the soil agreed well with field data for the first 3 years of the simulation. During the final year the model predicted considerably higher levels of soil mineral-N content compared with measurements. To reach the mineral-N level measured at the time of ploughing the ley, the simulated N-uptake by plants had to be increased by 8 g N m⁻². Simulations of nitrate leaching suggested that estimates of leaching based on measurements in tile-drained plots can be considerably underestimated. Accurate quantification of leaching in tile-drained plots often requires additional information on water-flow paths. A substantial increase in simulated and measured values for the mineral-N content of the soil occurred after ploughing the ley. In the simulation, most of the increase was due to a high crop residue input and the absence of a growing crop after ploughing. Litter accumulations in the soil during the 4-year period contributed little to the increase in soil mineral-N.     

The influence of earthworms (Lumbricidae) on the nitrogen dynamics in the soil litter system of a deciduous forest

Summary The influence of earthworms (Aporrectodea caliginosa (Savigny) and Lumbricus castaneus (Savigny)) on the rate of nitrogen net mineralization of the soil was studied in the laboratory and in the field. The additional mineralization of nitrogen cause by the burrowing activity of the substrat feeding earthworm A. caliginosa (N _L )was directly correlated to the biomass of the lumbricids independently of their number. A rise in temperature caused an exponential increase in N _L values. The Q ₁₀ value of this process (2.18) was found to be much higher than that of the nitrogen mineralization without earthworms (Q ₁₀=1.22). At 15°C the N _L value caused by A. caliginosa was calculated to be about 250 g N g^-1 fresh body wt d^-1. Using the experimentally determined exponential relationship between temperature and N _L values, the additional nitrogen mineralization caused by a population of A. caliginosa in a beechwood on limestone was calculated to be 4.23 kg ha^-1 a^-1.In contrast to A. caliginosa the litter dwelling species L. castaneus lost considerable amounts of biomass (56%) during the 4 week incubation period. Only 1/3 of the nitrogen equivalent to the weight loss of the animals was recovered in the mineral nitrogen pool.The addition of litter (old beech leaf litter, freshly fallen beech and ash leaf litter) had a pronounced effect on both nitrogen net mineralization and N _L values of the soil. Presence of old beech leaves caused an increase in both values, wheres the other litter types effected a decrease in nitrogen net mineralization. Fragmented ash litter was found to have the most distinct effect on N _L values (-69%) and nitrogen net mineralization (-74%).     

Ungulate stimulation of nitrogen cycling and retention in Yellowstone Park grasslands

We studied how ungulates and a large variation in site conditions influenced grassland nitrogen (N) dynamics in Yellowstone National Park. In contrast to most grassland N studies that have examined one or two soil N processes, we investigated four rates, net N mineralization, nitrification, denitrification, and inorganic N leaching, at seven paired sites inside and outside long-term (33+ year) exclosures. Our focus was how N fluxes were related to one another among highly variable grasslands and how grazers influenced those relationships. In addition, we examined variation in soil δ¹⁵N among grasslands and the relationships between soil ¹⁵N abundance and N processes. Previously, ungulates were reported to facilitate net N mineralization across variable Yellowstone grasslands and denitrification at mesic sites. In this study, we found that herbivores also promoted nitrification among diverse grasslands. Furthermore, net N mineralization, nitrification, and denitrification (kg N ha^–1 year^–1, each variable) were postively and linearly related to one another among all grasslands (grazed and fenced), and grazers reduced the nitrification/net N mineralization and denitrification/net N mineralization ratios, indicating that ungulates inhibited the proportion of available NH₄ ⁺ that was nitrified and denitrified. There was no relationship between net N mineralization or nitrification with leaching (indexed by inorganic N adsorbed to resin buried at the bottom of rooting zones) and leaching was unaffected by grazers. Soil δ¹⁵N was positively and linearly related to in situ net N mineralization and nitrification in ungrazed grasslands; however, there was no relationship between isotopic composition of N and those rates among grazed grasslands. The results suggested that grazers simultaneously increased N availability (stimulated net N mineralization and nitrification per unit area) and N conservation (reduced N loss from the soil per unit net N mineralization) in Yellowstone grasslands. Grazers promoted N retention by stimulating microbial productivity, probably caused by herbivores promoting labile soil C. Process-level evidence for N retention by grazers was supported by soil δ¹⁵N data. Grazed grassland with high rates of N cycling had substantially lower soil δ¹⁵N relative to values expected for ungrazed grassland with comparable net N mineralization and nitrification rates. These soil ¹⁵N results suggest that ungulates inhibited N loss at those sites. Such documented evidence for consumer control of N availability to plants, microbial productivity, and N retention in Yellowstone Park is further testimony for the widespread regulation of grassland processes by large herbivores. Received: 5 May 1999 / Accepted: 1 November 1999     

Soil nitrogen dynamics in a holm oak forest

Soil nitrogen (N) dynamics were studied in a dense, holm oak (Quercus ilex ssp. ilex) stand in the Montseny mountains to determine annual and seasonal patterns of N availability and uptake in an undisturbed Mediterranean forest on acidic soil. Soil mineral N content, net N mineralization (NNM), and net nitrification (NN) were determined by monthly sampling at two soil depths followed by in situ incubation in polyethylene bags. NNM per unit of soil mass was much higher at 0–5 cm than at 5–20 cm (annual means 24 and 2.5 mg N/kg, respectively) but on an area basis NNM was similar at both depths. A total of 80 kg N/ha/yr were mineralized from the first 20 cm of soil. NN amounted to only 9% of the annual NNM (7.5 kg N/ha/yr) and it occurred only in the upper 5 cm. NNM was maximum in June and July, while the NN peaked in May. Despite favourable soil temperature and moisture, NNM was negative in autumn because of microbial immobilization. Seasonal and depth variations of NNM appeared to be controlled more by substrate quality than by organic matter quantity, temperature or moisture. NN was not limited by ammonium availability. Calculated N uptake amounted to 91 kg/ha yr, peaking in June and July. The investigated stand showed a moderately high N availability, but ammonium was the major form of mineral N supply for holm oak.     

Organic carbon and nitrogen losses influenced by vegetation removal in a semiarid mediterranean soil

A reduction in plant cover can lead to an increase in the erosionprocesses that diminish soil quality. Any rise in temperature resulting frompredicted climate changes may aggravate this effect, particularly in semiaridMediterranean areas. Bearing this in mind, the capacity of a soil to preserveorganic matter becomes very important if the soil is to maintain its physicaland chemical properties. Soil organic carbon and nitrogen changes wereevaluatedin a non-disturbed (with natural vegetation) and a disturbed (all vegetationmanually clipped to ground level) pine system. Nine years after vegetationremoval significant differences (p < 0.01) were found in the soil organiccarbon content between plots (top 20 cm), but not in totalnitrogen. In the disturbed plot 0.0232 Mg ha^–1y^–1 of soil organic carbon were lost through erosionand4.30 Mg ha^–1 y^–1 throughmineralization. In the first 48 months after vegetation removal the soilorganiccarbon content fell from 40.3 to 28.0 g kg^–1. Inthe last 60 months of the experiment the amount of organic carbon in the soilfell from 28.0 to 27.7 g kg^–1. This result wasmainly attributable to the intense oxidization, which took place during thefirst 60 months, of organic matter linked to the coarse soil mineral fraction.Up to the 72^nd month the losses by erosion were a total of 532.7g, which rose to 1284.4 g by the end of theexperiment(108 months). The effect of vegetation removal in a Mediterranean semiarid arealeads to a rapid decline in the amount of organic carbon stored in the soil.Such perturbation is irreversible if left to nature.     

Herbivore influence on soil microbial biomass and nitrogen mineralization in a northern grassland ecosystem: Yellowstone National Park

Microorganisms are largely responsible for soil nutrient cycling and energy flow in terrestrial ecosystems. Although soil microorganisms are affected by topography and grazing, little is known about how these two variables may interact to influence microbial processes. Even less is known about how these variables influence microorganisms in systems that contain large populations of free-roaming ungulates. In this study, we compared microbial biomass size and activity, as measured by in situ net N mineralization, inside and outside 35- to 40-year exclosures across a topographic gradient in northern Yellowstone National Park. The objective was to determine the relative effect of topography and large grazers on microbial biomass and nitrogen mineralization. Microbial C and N varied by almost an order of magnitude across sites. Topographic depressions that contained high plant biomass and fine-textured soils supported the greatest microbial biomass. We found that plant biomass accurately predicted microbial biomass across our sites suggesting that carbon inputs from plants constrained microbial biomass. Chronic grazing neither depleted soil C nor reduced microbial biomass. We hypothesize that microbial populations in grazed grasslands are sustained mainly by inputs of labile C from dung deposition and increased root turnover or root exudation beneath grazed plants. Mineral N fluxes were affected more by grazing than topography. Net N mineralization rates were highest in grazed grassland and increased from dry, unproductive to mesic, highly productive communities. Overall, our results indicate that topography mainly influences microbial biomass size, while mineral N fluxes (microbial activity) are affected more by grazing in this grassland ecosystem. Received: 4 June 1997 / Accepted: 16 December 1997     

Possibilities for dispersal in annual and perennial grasses in a savanna in Botswana

Dispersal of 11 dominant grass species in the savanna of southern Botswana was investigated. The dispersal is autochorous, anemochorous and epizoochorous independent of the life-cycle (annuals, perennials). Dispersal distances were estimated experimentally. Anemochorous species with a plume-like spikelet such as Chloris virgata and Enneapogon cenchroides have a low rate of descent (<1 m s^-1) and a low Reynold number (100–110). Nevertheless they can only be transported up to 13 m from the originating infructescence at a wind velocity of 10 m s^-1. Therefore, the majority of the disseminules remained near the parent plant. By analysing the seed pool under the canopy of trees of Dichrostachys cinerea epizoochorous species such as Tragus berteronianus were dominant at the cattle resting sites under trees. The results are discussed in relation to the three seed dispersal hypotheses and the model of selective interaction of dispersal, dormancy, and seed site as adaptions to variable environments.     

California native and exotic perennial grasses differ in their response to soil nitrogen,exotic annual grass density,and order of emergence

Early emergence of plant seedlings can offer strong competitive advantages over later-germinating neighbors through the preemption of limiting resources. This phenomenon may have contributed to the persistent dominance of European annual grasses over native perennial grasses in California grasslands, since the former species typically germinate earlier in the growing season than the latter and grow rapidly after establishing. Recently, European perennial grasses have been spreading into both non-native annual and native perennial coastal grass stands in California. These exotic perennials appear to be less affected by the priority effects arising from earlier germination by European annual grasses. In addition, these species interactions in California grasslands may be mediated by increasing anthropogenic or natural soil nitrogen inputs. We conducted a greenhouse experiment to test the effects of order of emergence and annual grass seedling density on native and exotic perennial grass seedling performance across different levels of nitrogen availability. We manipulated the order of emergence and density of an exotic annual grass (Bromus diandrus) grown with either Nassella pulchra (native perennial grass), Festuca rubra (native perennial grass), or Holcus lanatus (exotic perennial grass), with and without added nitrogen. Earlier B. diandrus emergence and higher B. diandrus density resulted in greater reduction in the aboveground productivity of the perennial grasses. However, B. diandrus suppressed both native perennials to a greater extent than it did H. lanatus. Nitrogen addition had no effect on the productivity of native perennials, but greatly increased the growth of the exotic perennial H. lanatus, grown with B. diandrus. These results suggest that the order of emergence of exotic annual versus native perennial grass seedlings could play an important role in the continued dominance of exotic annual grasses in California. The expansion of the exotic perennial grass H. lanatus in coastal California may be linked to its higher tolerance of earlier-emerging annual grasses and its ability to access soil resources amidst high densities of annual grasses.     

Below-ground carbon and nitrogen accumulation in perennial grasses: A comparison of caespitose and rhizomatous growth forms

An experiment was conducted to compare below-ground soil organic carbon and total nitrogen accumulation between caespitose and rhizomatous perennial grasses in long-term (<25 yrs) grazed and ungrazed sites in semi-arid and mesic communities in the North American Great Plains. Development of greater nutrient pools beneath than between clones occurred at minimal clone basal areas (<60 cm²) for both caespitose species. Caespitose grasses accumulated substantially greater pools of carbon (20–200 fold) and nitrogen (50–500 fold) in soils to a depth of 10 cm beneath clones than rhizomatous grasses accumulated in rhizomes in both communities. Carbon and nitrogen pools in soils beneath caespitose clones exceeded combined (soil + rhizome) pools for rhizomatous grasses for a majority of the clone basal areas (>90 cm²) in the mesic community. In contrast, both pool sizes were smaller beneath the caespitose grass at all clone basal areas than the combined pools for the rhizomatous grass in the semi-arid community. The occurrence of larger soil nutrient pools beneath the rhizomatous species in the semi-arid community was largely a consequence of niche separation for microsites characterized by soils with higher nutrient concentrations, rather than plant-induced increases in nutrient concentrations. Although nutrient islands do not occur beneath rhizomatous grasses, their distribution in the semi-arid community was restricted to microsites characterized by soils with higher SOC and N concentrations. A greater efficiency of nutrient accumulation per unit rhizome mass and the maintenance of rhizome nutrient pools of similar magnitude to those of the rhizomatous grass in the mesic community may also contribute to the distribution of rhizomatous grasses in semi-arid communities. The existence of nutrient islands beneath a wide range of clone sizes in both mesic and semi-arid communities provides circumstantial evidence to suggest that nutrient islands beneath caespitose grasses may contribute to clone fitness in this growth form.     

Gopher mound soil reduces growth and affects ion uptake of two annual grassland species

Summary Portions of an annual serpentine grassland community in California are subject to frequent gopher mound formation. Consequently, studies were undertaken to characterize the effects of mound soils on plant growth and ion uptake. For two of the dominant annual species (Bromus mollis L. and Plantago erecta Morris), growth was reduced in gopher mound soil relative to that in inter-mound soil. A similar reduction in growth was found for plants grown in soils collected at a depth corresponding to the depth of gopher burrowing. This reduction in growth was associated with lower total P and N contents of the soil which were reflected in lower shoot contents of N and P. Additional experiments, however, showed that reduced N and P availabilities in mound soil were not entirely responsible for the growth reduction. Similarly, shoot Ca/Mg ratios were reduced in mound soil but additions of Ca improved the Ca/Mg ratio without improving growth. Growth reductions were associated with altered shoot concentrations of microelements, particularly elevated levels of Mn. A competition experiment between Plantago and Bromus showed that Bromus was more competitive than Plantago in mound and inter-mound soils and that soil type had only small affects on the nature of the interaction between the two species.     

Established native perennial grasses out-compete an invasive annual grass regardless of soil water and nutrient availability

Competition and resource availability influence invasions into native perennial grasslands by non-native annual grasses such as Bromus tectorum. In two greenhouse experiments we examined the influence of competition, water availability, and elevated nitrogen (N) and phosphorus (P) availability on growth and reproduction of the invasive annual grass B. tectorum and two native perennial grasses (Elymus elymoides, Pascopyrum smithii). Bromus tectorum aboveground biomass and seed production were significantly reduced when grown with one or more established native perennial grasses. Conversely, average seed weight and germination were significantly lower in the B. tectorum monoculture than in competition native perennial grasses. Intraspecific competition reduced per-plant production of both established native grasses, whereas interspecific competition from B. tectorum increased production. Established native perennial grasses were highly competitive against B. tectorum, regardless of water, N, or P availability. Bromus tectorum reproductive potential (viable seed production) was not significantly influenced by any experimental manipulation, except for competition with P. smithii. In all cases, B. tectorum per-plant production of viable seeds exceeded parental replacement. Our results show that established plants of Elymus elymoides and Pascopyrum smithii compete successfully against B. tectorum over a wide range of soil resource availability.