Relationships between root density of the African grass Hyparrhenia diplandra and nitrification at the decimetric scale: an inhibition-stimulation balance hypothesis

Previous studies have shown that Lamto savannah exhibits two different types of nitrogen cycle with high and low nitrification sites and suggested that the perennial grass Hyparrhenia diplandra is responsible for this duality at a subpopulation level, with one ecotype being thought to be able to inhibit nitrification. The present work aimed to investigate the relationships between nitrification and the roots of H. diplandra at two scales. (i) Site-scale experiments gave new insight into the hypothesized control of nitrification by H. diplandra tussocks: the two ecotypes exhibited opposite influences, inhibition in a low nitrification site (A) and stimulation in a high nitrification site (B). (ii) Decimetric-scale experiments demonstrated close negative or positive relationships (in sites A or B, respectively) between the roots and nitrification (in the 0-10 cm soil layer), showing an unexpectedly high sensitivity of the nitrification process to root density. In both soils, the correlation between the roots and nitrification decreased with depth and practically disappeared in the 20-30 cm soil layer (where the nitrification potential was found to be very low). Therefore, the impact of H. diplandra on nitrification may be viewed as an inhibition-stimulation balance.     

Nitrification potentials in early successional black locust and in mixed hardwood forest stands in the southern Appalachians,USA

Soil nitrogen mineralisation and nitrification potentials, and soil solution chemistry were measured in black locust (Robinia pseudo-acacia L.), in pine-mixed hardwood stands on an early successional watershed (WS6), and in an older growth oak-hickory forest located on an adjacent, mixed hardwood watershed (WS14) at Coweeta Hydrologic laboratory, in the southern Appalachian mountains, U.S.A. Nitrification potentials were higher in black locust and pine-mixed hardwood early successional stands than in the oak-hickory forest of the older growth watershed. Ammonification rates were the main factor controlling nitrification in the early successional stands. There was no evidence of inhibition of nitrification in soils from the older growth oak-hickory forest site.Within the early successional watershed, black locust sites had net mineralisation and nitrification rates at least twice as high as those in the pine mixed-hardwood stands. Concentrations of exchangeable nitrate in the soil of black locust stands were higher than in pine-mixed hardwoods at 0–15 cm in March and they were also higher at 0–15, 16–30 and 31–45 cm depth in the black locust dominated sites in July. Soil solution nitrate concentrations were higher under black locust than under pine-mixed hardwoods. Areas dominated by the nitrogen fixing black locust had greater nitrogen mineralisation and nitrification rates, resulting in higher potential for leaching losses of nitrate from the soil column in the early successional watershed.     

Trees improve grass quality for herbivores in African savannas

The tree–grass interactions of African savannas are mainly determined by varying rainfall patterns and soil fertility. Large savanna trees are known to modify soil nutrient conditions, but whether this has an impact on the quality of herbaceous vegetation is unclear. However, if this were the case, then the removal of trees might also affect the structure and quality of the grass layer. We studied the impact of large nitrogen- and non-nitrogen fixing trees on the sub-canopy (SC) grass layer in low- and high-rainfall areas of differing soil fertility in eastern and southern Africa. We compared the structure and nutrient levels of SC grasses with those outside the canopy. Grass leaf nitrogen and phosphorus contents beneath tree canopies were elevated at all study sites and were up to 25% higher than those outside the canopy in the site of lowest rainfall and soil fertility. Grass leaf fibre and organic matter (OM) contents were slightly enhanced beneath tree canopies. At the site of highest rainfall and soil fertility, grasses beneath the canopy had significantly lower ratios of stem:leaf biomass and dead:living leaf material. Grass species composition differed significantly, with the highly nutritious Panicum spp. being most abundant underneath tree crowns. In the two drier study sites, soil nitrogen and OM contents were enhanced by 30% beneath trees. N-fixation capacity of trees did not contribute to the improved quality of grass under the canopy. We conclude that trees improve grass quality, especially in dry savannas. In otherwise nutrient-poor savanna grasslands, the greater abundance of high-quality grass species with higher contents of N and P and favourable grass structure beneath trees could attract grazing ungulates. As these benefits may be lost with tree clearance, trees should be protected in low fertility savannas and their benefits for grazing wildlife recognised in conservation strategies.     

Spatial pattern of soil nitrogen availability and its relationship to stand structure in a coniferous-broadleaved mixed forest with a dense dwarf bamboo understory in northern Japan

Natural disturbances create spatial patterns of the ecosystem processes and functions in natural forests. However, how dynamics and the spatial structure of forests relate to soil nitrogen dynamics is not well understood. We examined the spatial relationship between the distributions of canopy and understory species, and soil nitrogen dynamics in a natural coniferous-broadleaved mixed forest with a dense understory of Sasa dwarf bamboo in northern Japan. The O horizon was thick where coniferous litter predominated, and it was thin where broadleaved litter predominated. The soil water content was low in areas with a thick O horizon and a high abundance of coniferous trees. The soil nitrate content was low where the soil water content was low, and the soil nitrate content increased linearly with increasing net nitrification potential. These results suggest that the soil nitrate content under the coniferous canopy was lower because of the low nitrification potential of soil microbes in soils with low water contents. The soil nitrate content and nitrification potential were higher in the canopy gap than under the canopy. Our results suggest that forest structure, specifically the thickness of the forest floor, significantly affects the spatial pattern of the soil water content, thereby creating a spatial pattern of soil nitrogen availability at a relatively small scale with flat topography. The higher nitrification potential under the canopy gap could pose a long-term risk of nitrate leaching because of the suppression of the natural regeneration of canopy species by dense Sasa dwarf bamboo in this forest ecosystem.     

Soil nitrogen patterns induced by colonization of Polygonum cuspidatum on Mt. Fuji

Summary The spatial pattern of soil nitrogen was analyzed for a patchy vegetation formed by the colonization of Polygonum cuspidatum in a volcanic desert on Mt. Fuji. Soils were sampled radially from the bare ground to the center of the patch, and analyses were done for bulk density, water content, soil acidity, organic matter, organic nitrogen, and ammonium and nitrate nitrogen. The soils matured with succession from the bare ground through P. cuspidatum to Miscanthus oligostachyus and Aster ageratoides sites: bulk density decreased, and water content, organic matter, organic nitrogen, and ammonium nitrogen increased. Nitrate nitrogen showed the highest values at the P. cuspidatum site. Application of principal component analysis to the soil data discriminated two component factors which control the variation of soil characteristics: the first factor is related to soil formation and the second factor to nitrogen mineralization and nitrification. The effect of soil formation on nitrogen mineralization and nitrification was analyzed with a first-order kinetic model. The decreasing trends with soil formation in the ratios of mineral to organic nitrogen and of nitrate to ammonium nitrogen could be accounted for by the higher activity of immobilization by microorganisms and uptake by plants in the more mature ecosystem.     

Repression of nitrification in soils under a climax grassland vegetation

Abstract Two hypotheses on repression of nitrification in climax vegetations (i.e. nitrogen immobilization and allelopathy) were investigated. In this study the potential nitrification activities and numbers of ammonium-oxidizing bacteria were established in a nature reserve with a series of natural grasslands with vegetational different stages of succession of plants species. The pastures had not been fertilized for 3, 7, 20 and 46 years, respectively, and the gradual decrease in availability of nutrients had led to pastures dominated by different grass species. In each field soil parameters, potential nitrification activities (PNA) and numbers of ammonium-oxidizing bacteria were determined in the root zone of Holcus lanatus as well as in that of a grass species characteristic of the stage of succession. In the rhizosphere of H. lanatus decreasing PNA and numbers of ammonium-oxidizing bacteria were observed as the period of non fertilization increased. Within each field no significant differences in PNA were observed between the root zones of H. lanatus and those of the dominant grass species. From these results it is concluded that, in these fields, decreasing nitrification was related only to decreasing ammonium availability and not to species composition. No indications were obtained that allelochemicals were involved in the flow nitrification potentials of late stages of succession. The optimum pH of the ammonium-oxidizing community, measured as PNA, decreased as the period of non fertilization increased. It is suggested that impoverishment of the grassland soil with respect to nitrogen availability selects against ammonium-oxidizing bacteria with a relatively high pH optimum.     

Soil organic carbon content at a range of north Australian tropical savannas with contrasting site histories

Soils play an important role in the global carbon cycle, and can be major source or sink of CO₂ depending upon land use, vegetation type and soil management practices. Natural and human impact on soil carbon concentration and storage is poorly understood in native north Australian savanna, yet this represents the largest carbon store in the ecosystem. To gain understanding of possible management impacts on this carbon pool, soil organic carbon (SOC) of the top 1m of red earth sands and sandy loams common in the region was sampled at 5 sites with different vegetation cover and site history (fire regime and tree removal). SOC was high when compared to other published values for savannas and was more comparable with dry-deciduous tropical forests. Sites sampled in this study represent high rainfall savannas of northern Australia (> 1700 mm annual rainfall) that feature frequent burning (2 in 3 years or more frequent) and a cycle of annual re-growth of tall C₄ grasses that dominate the savanna understorey. These factors may be responsible for the higher than expected SOC levels of the surface soils, despite high respiration rates. Medium term fire exclusion (15–20 years) at one of the sampled sites (Wildlife Park) dramatically reduced the grassy biomass of the understorey. This site had lower SOC levels when compared to the grass dominated and frequently burnt sites, which may be due to a reduction in detrital input to surface (0–30 cm) soil carbon pools. Exclusion of trees also had a significant impact on both the total amount and distribution of soil organic carbon, with tree removal reducing observed SOC at depth (100 cm). Soil carbon content was higher in the wet season than that in the dry season, but this difference was not statistically significant. Our results indicated that annual cycle of grass growth and wildfire resulted in small carbon accumulation in the upper region of the soil, and removal of woody plants resulted in significant carbon losses to recalcitrant, deep soil horizons greater than 80 cm depth.     

Browsing herbivores improve the state and functioning of savannas: A model assessment of alternative land‐use strategies

Changing climatic conditions and unsustainable land use are major threats to savannas worldwide. Historically, many African savannas were used intensively for livestock grazing, which contributed to widespread patterns of bush encroachment across savanna systems. To reverse bush encroachment, it has been proposed to change the cattle‐dominated land use to one dominated by comparatively specialized browsers and usually native herbivores. However, the consequences for ecosystem properties and processes remain largely unclear. We used the ecohydrological, spatially explicit model EcoHyD to assess the impacts of two contrasting, herbivore land‐use strategies on a Namibian savanna: grazer‐ versus browser‐dominated herbivore communities. We varied the densities of grazers and browsers and determined the resulting composition and diversity of the plant community, total vegetation cover, soil moisture, and water use by plants. Our results showed that plant types that are less palatable to herbivores were best adapted to grazing or browsing animals in all simulated densities. Also, plant types that had a competitive advantage under limited water availability were among the dominant ones irrespective of land‐use scenario. Overall, the results were in line with our expectations: under high grazer densities, we found heavy bush encroachment and the loss of the perennial grass matrix. Importantly, regardless of the density of browsers, grass cover and plant functional diversity were significantly higher in browsing scenarios. Browsing herbivores increased grass cover, and the higher total cover in turn improved water uptake by plants overall. We concluded that, in contrast to grazing‐dominated land‐use strategies, land‐use strategies dominated by browsing herbivores, even at high herbivore densities, sustain diverse vegetation communities with high cover of perennial grasses, resulting in lower erosion risk and bolstering ecosystem services.     

植物氮形态利用策略及对外来植物入侵性的影响

氮是影响外来植物入侵性的重要因素之一, 但相关研究多关注土壤氮水平的效应, 较少考虑氮形态的作用。为从土壤氮形态利用的角度阐释外来植物的入侵机制, 本文在植物氮形态利用策略分析的基础上, 综述了外来植物氮形态利用的偏好性及其对入侵性的影响。植物的氮形态利用策略有偏好性和可塑性两种, 这可能与植物对土壤氮形态特性的长期适应有关; 植物不仅可以对土壤氮形态做出响应, 而且还能改造土壤氮形态, 并对改变后的土壤氮形态做出反馈响应。很多外来植物入侵硝态氮占优势的干扰生境, 偏好硝态氮的外来植物与本地植物竞争硝态氮; 而偏好铵态氮的外来植物通过抑制土壤硝化作用, 营造铵态氮环境, 促进自身生长, 同时抑制偏好硝态氮的本地植物生长。然而, 植物氮形态利用策略不是一成不变的, 而是受多种生物和非生物因素共同作用影响的复杂过程, 今后应加强多因素交互作用对外来入侵植物氮形态利用策略的影响及机制研究, 更好地揭示氮形态利用策略, 尤其是氮形态利用的可塑性与外来植物入侵性的关系。     

Nitrogen Fixation and Nitrogen Assimilation in a Temperate Saline Ecosystem

As nitrogen is known to be a limiting factor for plant growth, we were interested in the relationship between soil microbial activity and the nitrogen assimilation of 5 different halophytes from 4 saline sites near the lake “Neusiedlersee”, Austria. The following were studied between May and October 1985: nitrogen fixation (¹⁵N₂ and acetylene reduction): N-mineralization; several soil characteristics and in vivo nitrate reductase activity of roots and shoots of these plants. NO^?₃, org. N- and carboxylate contents of both roots and shoots, as well as the effect of NO^?₃-fertilization on the amounts of these substances, were determined on plants growing in the field during a 3-day period in September 1985. Fertilization led to a decrease in acetylene reduction activity at most sites, and an increase in the nitrate reductase activity of the shoots of all plants. Overall, carboxylate and organic nitrogen contents of these halophytes did not change in response to fertilization. Only in the roots of Aster tripolium and Atriplex hastata was there a marked increase in the nitrate reductase activity in response to fertilization. Species growing at the same site, such as Plantago maritima and Lepidium crassifolium showed contrasting levels of assimilatory activity. Apparent low rates of ammonification and nitrification were detected in soils from the 4 sites. The results are discussed in relation to the nitrogen and carbon economies of the microorganisms and plants.     

Nitrate nutrition ofDeschampsia flexuosa (L.) Trin. in relation to nitrogen deposition in Sweden

Summary Current and maximally induced nitrate reductase activity (NRA), total-N, nitrate, K, P, Ca, Mg, Mo and sucrose in leaves ofDeschampsia flexuosa was measured three times during the vegetation period in forests along a deposition gradient (150 km) in south Sweden, in north Sweden where the nitrogen deposition is considerably lower, and at heavily N-fertilized plots. In addition, the interaction between nitrogen nutrition and light was studied along transects from clearings into forest in both south and north Sweden. Plants from sites with high nitrogen deposition had elevated current NRA compared to plants from less polluted sites, indicating high levels of available soil nitrate at the former. Current NRA and total N concentration in grass from sites with high deposition resembled those found at heavily N-fertilized plots. Under such circumstances, the ratio current NRA: maximally induced NRA as well as the concentration of nitrate was high, while the concentration of sucrose was low. This suggests that the grass at these sites was already utilizing a large portion of its capacity to assimilate nitrate. Light was found to play an important role in the assimilation of nitrate; leaf concentration of sucrose was found to be negatively correlated with both nitrate and total N. Consequently, grass growing under dense canopies in south Sweden is not able to dilute N by increasing growth. The diminished capacity of the grass to assimilate nitrate will increase leaching losses of N from forests approaching N saturation.     

Carbon and nitrogen in soil and vegetation at sites differing in successional age

We studied vegetation and soil development during primary succession in an inland drift sand area in the Netherlands. We compared five sites at which primary succession had started at different moments in the past, respectively 0, 10, 43 and 121 years ago, and a site at which succession had not yet started. In the three younger sites the vegetation was herbaceous, whereas in the two older sites a pine forest had formed. Forest formation was accompanied by the development of an FH-layer in the soil, an increase in the amount of soil organic matter, and an increase in nitrogen mineralisation rate from 1.9 to 18 g N m^–2 yr^–1. Soil moisture content also increased, whereas pH showed a steady decrease with site age. The vegetation changed from a herbaceous vegetation dominated by mosses and lichens and the grass species Corynephorus canescens and Festuca ovina towards a pine forest with an understorey vegetation dominated by Deschampsia flexuosa and, at the oldest site, with dwarf shrubs Empetrum nigrum and Vaccinium myrtillus. At the same time the total amounts of carbon and nitrogen of the ecosystem increased, with a relatively stronger increase of the carbon pool. The establishment of trees during succession greatly affects the dynamics of the ecosystem, especially its carbon dynamics.     

Potential Nitrification as an Indicator of Preferential Uptake of Ammonium or Nitrate by Plants in an Oak Woodland Understorey

The preferences of some woodland understorey species for ammoniumand nitrate were investigated by measuring the potential nitrification(conversion of ammonium to nitrate) in the rhizosphere comparedwith the bulk soil. Less acid-tolerant species, which usuallyprefer nitrate or a mixture of ammonium and nitrate in hydroponicculture, should have a higher potential nitrification in therhizosphere compared to the bulk soil due to a low uptake ofammonium (since ammonium is relatively immobile). Acid-tolerantspecies should have a high uptake of ammonium and thereby loweror equal potential nitrification in the rhizosphere comparedto the bulk soil. The hypothesis was tested in a field investigationof five understorey herb species, Deschampsia flexuosa, Convallariamajalis, Poa nemoralis, Geum urbanum andAegopodium podagrariaperformed in oak forests in southern Sweden. Overall, the twoless acid-tolerant species, Geum urbanum and Aegopodium podagraria,had high potential nitrification in the rhizosphere comparedto the bulk soil (indicating a relatively low uptake of ammonium),whilst the acid tolerant species, Deschampsia flexuosa andConvallariamajalis , had approximately equal potential nitrification inthe rhizosphere compared to the bulk soil (indicating a relativelyhigh uptake of ammonium). In the case of Poa nemoralis, a specieswhich grows in both acid and less acid soils, we found the potentialnitrification in the rhizosphere and in the bulk soil to besimilar at low inorganic nitrogen concentrations, but the difference(rhizosphere > bulk) increased when nitrification in thebulk soil was enhanced (i.e. when the nitrogen availabilityincreased). The potential nitrification in the bulk soil variedbetween 0 and 16 nmol g^-1h^-1and was positively correlated withpH. When species occurred at the same site, the potential nitrificationin the bulk soil tended to be lower for the acid tolerant species.Despite a large variation in potential nitrification, the methodoffers a possibility of measuring the preference of plants forammonium/nitrate in a soil system, under natural conditions.Copyright 2000 Annals of Botany Company Ammonium uptake, nitrate uptake, nitrogen preference, potential nitrification, rhizosphere, Deschampsia flexuosa, Convallaria majalis, Poa nemoralis, Geum urbanum, Aegopodium podagraria     

Biogeochemical and Microbial Legacies of Non‐Native Grasses Can Affect Restoration Success

The restoration of disturbed ecosystems is challenging and often unsuccessful, particularly when non‐native plants are abundant. Ecosystem restoration may be hindered by the effects of non‐native plants on soil biogeochemical characteristics and microbial communities that persist even after plants are removed. To examine the importance of soil legacy effects, we used experimental restorations of Florida shrubland habitat that had been degraded by the introduction of non‐native grasses coupled with either mechanical disturbance or pasture conversion. We removed non‐native grasses and inoculated soils with native microbial communities at each degraded site, then examined how habitat structure, soil nitrogen, soil microbial abundances, and native seed germination responded over two years compared to undisturbed native sites. Grass removal treatments effectively restored some aspects of native habitat structure, including decreased exotic grass cover, increased bare ground, and reduced litter cover. Soil fungal abundance was also somewhat restored by grass removals, but soil algal abundance was unaffected. In addition, grass removal and microbial inoculation improved seed germination rates in degraded sites, but these remained quite low compared to native sites. High soil nitrogen persisted throughout the experiment regardless of treatment. Many treatment effects were site‐specific, however, with legacies in the more degraded vegetation type tending to be more difficult to overcome. These results support the need for context‐dependent restoration approaches and suggest that the degree of soil legacy effects may be a good indicator of restoration potential.     

Positive feedbacks to growth of an invasive grass through alteration of nitrogen cycling

Understanding the mechanisms by which invasive plants maintain dominance is essential to achieving long-term restoration goals. While many reports have suggested invasive plants alter resource availability, experimental tests of feedbacks between invasive plants and soil resources are lacking. We used field observations and experimental manipulations to test if the invasive grass Microstegium vimineum both causes and benefits from altered soil nitrogen (N) cycling. To quantify M. vimineum effects on N dynamics, we compared inorganic N pools and nitrification rates in 20 naturally invaded and uninvaded plots across a range of mixed hardwood forests, and in experimentally invaded and uninvaded common garden plots. Potential nitrification rates were 142 and 63?% greater in invaded than uninvaded plots in forest and common garden soils, respectively. As a result, soil nitrate was the dominant form of inorganic N during peak M. vimineum productivity in both studies. To determine the response of M. vimineum to altered nitrogen availability, we manipulated the dominant N form (nitrate or ammonium) in greenhouse pots containing M. vimineum alone, M. vimineum with native species, and native species alone. M. vimineum productivity was highest in monocultures receiving nitrate; in contrast, uninvaded native communities showed no response to N form. Notably, the positive response of M. vimineum to nitrate was not apparent when grown in competition with natives, suggesting an invader density threshold is required before positive feedbacks occur. Collectively, our results demonstrate that persistence of invasive plants can be promoted by positive feedbacks with soil resources but that the magnitude of feedbacks may depend on interspecific interactions.     

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.     

Increased inorganic nitrogen leaching from a mountain grassland ecosystem following grazing removal: a hangover of past intensive land-use?

Heathlands and grasslands occur in montane regions, naturally or due to anthropogenic land-use. These are typically nutrient-poor but exposure to elevated nitrogen deposition and intensive livestock grazing causes large-scale ecological change. We studied the long-term implications of grazing removal on soil and drainage water biogeochemistry and the implications for nitrogen cycling in 50-year replicated grazing exclosures on a montane grassland exposed to high rates of ambient nitrogen deposition. Evidence of ‘ecosystem recovery’ represented by successional change from graminoid to shrub-dominance after cessation of grazing was not reflected in the soil biogeochemistry. Cessation of grazing had a negative impact, with increased soil extractable and soil solution nitrate concentrations; an apparent shift towards a more nitrogen-rich, bacterially dominated microbial community; and the acidification of soils and leachate. The increase in nitrate leaching appears to have been counterbalanced by a decrease in dissolved organic nitrogen leaching, approximately maintaining the overall nitrogen balance of the system, whilst apparently altering ecosystem functioning. High rates of organic matter cycling and inorganic nitrogen uptake in grazed grassland may have sustained ecosystem N limitation under elevated nitrogen deposition. Grazing removal caused long-term over-supply of nitrogen from mineralisation of enriched organic matter, exacerbated by continued high nitrogen deposition, exceeding the uptake demand of heath vegetation and resulting in nitrification and nitrate leaching. This disequilibrium between vegetation and soil following grazing removal has implications for restoration after periods of intensive grazing. Grazing may not simply leave a legacy of nutrient enrichment but its cessation may trigger nitrogen saturation and soil and freshwater eutrophication and acidification which counteract the immediate benefits of natural vegetation recovery. Long term, nitrogen saturation of abandoned grasslands is likely to reduce ecosystem resilience to invasion by nitrophilous species, pathogen attack and vulnerability to environmental pressures such as climate change. We conclude that partial and/or phased reduction in grazing levels may permit the more synchronised recovery of soils and vegetation, thereby avoiding imbalances between nitrogen supply and nitrogen demand and detrimental ecological effects.     

The invasive grass,Melinis minutiflora,inhibits tree regeneration in a Neotropical savanna

Abstract Exotic grasses are becoming increasingly abundant in Neotropical savannas, with Melinis minutiflora Beauv. being particularly invasive. To better understand the consequences for the native flora, we performed a field study to test the effect of this species on the establishment, survival and growth of seedlings of seven tree species native to the savannas and forests of the Cerrado region of Brazil. Seeds of the tree species were sown in 40 study plots, of which 20 were sites dominated by M. minutiflora, and 20 were dominated by native grasses. The exotic grass had no discernable effect on initial seedling emergence, as defined by the number of seedlings present at the end of the first growing season. Subsequent seedling survival in plots dominated by M. minutiflora was less than half that of plots dominated by native species. Consequently, at the end of the third growing season, invaded plots had only 44% as many seedlings as plots with native grasses. Above‐ground grass biomass of invaded plots was more than twice that of uninvaded plots, while seedling survival was negatively correlated with grass biomass, suggesting that competition for light may explain the low seedling survival where M. minutiflora is dominant. Soils of invaded plots had higher mean Ca, Mg and Zn, but these variables did not account for the higher grass biomass or the lower seedling survival in invaded plots. The results indicate that this exotic grass is having substantial effects on the dynamics of the tree community, with likely consequences for ecosystem structure and function.     

Soil Nitrogen Conditions Approach Preinvasion Levels following Restoration of Nitrogen-Fixing Black Locust (Robinia pseudoacacia) Stands in a Pine–Oak Ecosystem

Restoring native plant communities on sites formerly occupied by invasive nitrogen‐fixing species poses unique problems due to elevated soil nitrogen availability. Mitigation practices that reduce available nitrogen may ameliorate this problem. We evaluated the effects of tree removal followed by soil preparation or mulching on native plant growth and soil nitrogen transformations in a pine–oak system formerly occupied by exotic nitrogen‐fixing Black locust (Robinia pseudoacacia) trees. Greenhouse growth experiments with native grasses, Andropogon gerardii and Sorghastrum nutans, showed elevated relative growth rates in soils from Black locust compared with pine–oak stands. Field soil nutrient concentrations and rates of net nitrification and total net N‐mineralization were compared 2 and 4 years since Black locust removal and in control sites. Although soil nitrogen concentrations and total net N‐mineralization rates in the restored sites were reduced to levels that were similar to paired pine–oak stands after only 2 years, net nitrification rates remained 3–34 times higher in the restored sites. Other nutrient ion concentrations (Ca, Mg) and organic matter content were reduced, whereas phosphorus levels remained elevated in restored sites. Thus, 2–4 years following Black locust tree removal and soil horizon mixing achieved through site preparation, the concentrations of many soil nutrients returned to preinvasion levels. However, net nitrification rates remained elevated; cover cropping or carbon addition during restoration of sites invaded by nitrogen fixers could increase nitrogen immobilization and/or reduce nitrate availability, making sites more amenable to native plant establishment.