Plant Effects on Spatial and Temporal Patterns of Nitrogen Cycling in Shortgrass Steppe

Because of the water-limited nature and discontinuous plant cover of shortgrass steppe, spatial patterns in ecosystem properties are influenced more by the presence or absence of plants than by plant type. However, plant type may influence temporal patterns of nutrient cycling between plant and soil. Plants having the carbon-3 (C₃) or carbon-4 (C₄) photosynthetic pathway differ in phenology as well as other attributes that affect nitrogen (N) cycling. We estimated net N mineralization rates and traced nitrogen-15 (¹⁵N) additions among plant and soil components during May, July, and September of 1995 in native plots of C₃ plants, C₄ plants, or mixtures of C₃ and C₄. Net N mineralization was significantly greater in C₃ plots than in C₄ plots during both July and September. C₃ plots retained significantly more ¹⁵N in May than did mixed and C₄ plots; these differences in ¹⁵N retention were due to greater ¹⁵N uptake by C₃ plants than by C₄ plants during May. There were no significant differences in total ¹⁵N retention among plant communities for July and September. Soil ¹⁵N was influenced more by presence or absence of plants than by type of plant; greater quantities of ¹⁵N remained in soil interspaces between plants than in soil directly under plants for July and September. Our results indicate that plant functional type (C₃ versus C₄) can affect both the spatial and the temporal patterns of N cycling in shortgrass steppe. Further research is necessary to determine how these intraseasonal differences translate to longer-term and coarser-scale effects of plants on N cycling, retention, and storage. Received 8 December 1997; accepted 6 May 1998.     

Nitrogen Oxide Fluxes and Nitrogen Cycling during Postagricultural Succession and Forest Fertilization in the Humid Tropics

The effects of changes in tropical land use on soil emissions of nitrous oxide (N₂O) and nitric oxide (NO) are not well understood. We examined emissions of N₂O and NO and their relationships to land use and forest composition, litterfall, soil nitrogen (N) pools and turnover, soil moisture, and patterns of carbon (C) cycling in a lower montane, subtropical wet region of Puerto Rico. Fluxes of N₂O and NO were measured monthly for over 1 year in old (more than 60 years old) pastures, early- and mid-successional forests previously in pasture, and late-successional forests not known to have been in pasture within the tabonuco (Dacryodes excelsa) forest zone. Additional, though less frequent, measures were also made in an experimentally fertilized tabonuco forest. N₂O fluxes exceeded NO fluxes at all sites, reflecting the consistently wet environment. The fertilized forest had the highest N oxide emissions (22.0 kg N · ha⁻¹· y⁻¹). Among the unfertilized sites, the expected pattern of increasing emissions with stand age did not occur in all cases. The mid-successional forest most dominated by leguminous trees had the highest emissions (9.0 kg N · ha⁻¹· y⁻¹), whereas the mid-successional forest lacking legumes had the lowest emissions (0.09 kg N · ha⁻¹· y⁻¹). N oxide fluxes from late-successional forests were higher than fluxes from pastures. Annual N oxide fluxes correlated positively to leaf litter N, net nitrification, potential nitrification, soil nitrate, and net N mineralization and negatively to leaf litter C:N ratio. Soil ammonium was not related to N oxide emissions. Forests with lower fluxes of N oxides had higher rates of C mineralization than sites with higher N oxide emissions. We conclude that (a) N oxide fluxes were substantial where the availability of inorganic N exceeded the requirements of competing biota; (b) species composition resulting from historical land use or varying successional dynamics played an important role in determining N availability; and (c) the established ecosystem models that predict N oxide loss from positive relationships with soil ammonium may need to be modified. Received 22 February 2000; accepted 6 September 2000.     

Original Articles: Differential Influence of Plant Species on Soil Nitrogen Transformations Within Moist Meadow Alpine Tundra

Plant species can influence nitrogen (N) cycling indirectly through the feedbacks of litter quality and quantity on soil N transformation rates. The goal of this research was to focus on small-scale (within-community) variation in soil N cycling associated with two community dominants of the moist meadow alpine tundra. Within this community, the small-scale patchiness of the two most abundant species (Acomastylis rossii and Deschampsia caespitosa) provides natural variation in species cover within a relatively similar microclimate, thus enabling estimation of the effects of plant species on soil N transformation rates. Monthly rates of soil N transformations were dependent on small-scale variation in both soil microclimate and species cover. The relative importance of species cover compared with soil microclimate increased for months 2 and 3 of the 3-month growing season. Growing-season net N mineralization rates were over ten times greater and nitrification rates were four times greater in Deschampsia patches than in Acomastylis patches. Variability in litter quality [carbon:nitrogen (C:N) and phenolic:N], litter quantity (aboveground and fine-root production), and soil quality (C:N) was associated with three principal components. Variability between the species in litter quality and fine-root production explained 31% of the variation in net N mineralization rates and 36% of net nitrification rates. Site variability across the landscape in aboveground production and soil C:N explained 33% of the variation in net N mineralization rates and 21% of net nitrification rates. Within the moist meadow community, the high spatial variability in soil N transformation rates was associated with differences in the dominant species' litter quality and fine-root production. Deschampsia-dominated patches consistently had greater soil N transformation rates than did Acomastylis-dominated patches across the landscape, despite site variability in soil moisture, soil C:N, and aboveground production. Plant species appear to be an important control of soil N transformation in the alpine tundra, and consequently may influence plant community structure and ecosystem function.     

Nitrogen availability and nitrification during succession: Primary,secondary, and old-field seres

Suggestions that nutrient cycles become more strongly regulated and that nitrification is progressively inhibited in the course of ecological succession have stimulated numerous field measurements. Results of these are inconsistent; in some cases nitrogen turnover and nitrification decrease during succession, while in others both increase substantially.Consideration of the nature of disturbance which initiates each succession explains much of the difference in nitrogen dynamics. Primary succession (the development of ecosystems on wholly new substrates) invariably involves a low nitrogen availability and nitrification early in succession. In contrast, destructive disturbance followed by immediate regrowth (the pure case of secondary succession) invariably increases nitrogen availability (and generally nitrate production) in recently disturbed sites; it is followed by a decline during later stages of succession. Succession following a period of chronic disturbance (i.e. prolonged agriculture) does not follow such clear patterns; the duration and intensity of disturbance may control whether nitrogen availability and potential nitrification increase or decrease early in such seres.     

Original Articles: Dynamic Redistribution of Isotopically Labeled Cohorts of Nitrogen Inputs in Two Temperate Forests

We compared simulated time series of nitrogen-15 (¹⁵N) redistribution following a large-scale labeling experiment against field recoveries of ¹⁵NH₄ ⁺ and ¹⁵NO₃ ⁻ in vegetation tissues. We sought to gain insight into the altered modes of N cycling under long-term, experimentally elevated N inputs. The study took place in two contrasting forests: a red pine stand and a mixed deciduous stand (predominantly oak) at the Harvard Forest, Massachusetts, USA. We used TRACE, a dynamic simulation model of ecosystem biogeochemistry that includes ¹⁵N/¹⁴N ratios in N pools and fluxes. We simulated input–output and internal fluxes of N, tracing the labeled cohorts of N inputs through ecosystem pools for one decade. TRACE simulated the peaks and timing of ¹⁵N recovery in foliage well, providing a key link between modeling and field studies. Recovery of tracers in fine roots was captured less well. The model was structured to provide rapid, initial sinks for ¹⁵NO₃ ⁻ and ¹⁵NH₄ ⁺ in both forests, as indicated by field data. In simulations, N in litter turned over rapidly, even as humus provided a long-term sink for rapidly cycling N. This sink was greater in the oak forest. Plant uptake fluxes of N in these fertilized plots were on the same order of magnitude as net assimilation fluxes in forest-floor humus. A striking result was the small rate of incorporation of N in humus resulting from the transfer of litter material to humus, compared with large fluxes of N into humus and its associated microorganisms through direct transfers from pools of inorganic N in soils. Received 19 May 1998; accepted 30 September 1998     

Response of the Shortgrass Steppe to Changes in Rainfall Seasonality

Studies in temperate grassland ecosystems have shown that differences in composition of C₃ and C₄ plant functional types can have important influences on ecosystem pools and processes. We used a plant community dynamics model (STEPPE) linked to a biogeochemical cycling model (CENTURY) to determine how ecosystem properties in shortgrass steppe are influenced by plant functional type composition. Because of phenological differences between C₃ and C₄ plants, we additionally simulated the effects of precipitation seasonality on plant communities and examined how C₃ and C₄ composition interacts with precipitation to affect ecosystems. The model output suggests that differences in C₃ and C₄ composition can lead to differences in soil organic carbon (C) and nitrogen (N) within 1000 simulation years. Soil organic C and N (g C and N m^{− 2} to 0.2-m depth) were least in a 100% C₄ community compared with a 100% C₃ community and a mixed C₃–C₄ community. A change in the time of maximum precipitation from summer to spring in a simulated shortgrass steppe slightly favored C₃ plants over C₄ plants. The proportion of total net primary production accounted for by C₃ plants increased from 21% to 25% after 200 years, when 90 mm of precipitation was switched from summer to spring. Soil organic matter (SOM) was relatively stable in the C₄-dominated communities with respect to changes in precipitation seasonality, whereasSOM in the C₃ community was sensitive to precipitation seasonality changes. These results suggest an important interaction between plant community composition and precipitation seasonality on SOM, with phenology playing a key role. Received 9 June 1998; accepted 6 January 1999     

Original Articles: Nitrogen Mineralization in Two Unpolluted Old-Growth Forests of Contrasting Biodiversity and Dynamics

Studies in unpolluted, old-growth forests in the coastal range of southern Chile (42°30′S) can provide a baseline for understanding how forest ecosystems are changing due to the acceleration of nitrogen (N) inputs that has taken place over the last century. Chilean temperate forests, in contrast to their northern hemisphere counterparts, exhibit extremely low losses of inorganic N to stream waters. The objectives of this study were (a) to determine whether low inorganic N outputs in these forests were due to low rates of N mineralization or nitrification, and (b) to examine how biodiversity (defined as number of dominant tree species) and forest structure influence N mineralization and overall patterns of N cycling. Studies were conducted in a species-poor, conifer-dominated (Fitzroya cupressoides) forest with an even-aged canopy, and in a mixed-angiosperm (Nothofagus nitida) forest with a floristically more diverse and unstable canopy. Nitrogen mineralization rates measured in laboratory assays varied seasonally, reaching 6.0 μg N/g DW/day in both forests during late summer. Higher values were related to higher microbial activity, larger pools of labile inorganic N, and increased fine litter inputs. Field assays, conducted monthly, indicated positive net flux from N mineralization mainly from December to January in both forests. Annual net flux of N from mineralization varied from 20 to 23 kg/ha/year for the Fitzroya forest and from 31 to 37 kg/ha/year for the Nothofagus forest. Despite low losses of inorganic N to streams, N mineralization and nitrification are not inhibited in these forests, implying the existence of strong sinks for NO₃ ⁻ in the ecosystem. Field N mineralization rates were two times higher in the Nothofagus forest than in the Fitzroya forest, and correlated with greater N input via litterfall, slightly higher soil pH, and narrower carbon (C)–nitrogen ratios of soils and litter in the former. Differences in N mineralization between the two forest types are attributed to differences in biotic structure, stand dynamics, and site factors. Median values of net N mineralization rates in these southern hemisphere forests were lower than median rates for forests in industrialized regions of North America, such as the eastern and central USA. We suggest that these high N mineralization rates may be a consequence of enhanced atmospheric N deposition.     

Effects of Succession on Nitrogen Export in the West-Central Cascades, Oregon

This study examined impacts of succession on N export from 20 headwater stream systems in the west central Cascades of Oregon, a region of low anthropogenic N inputs. The seasonal and successional patterns of nitrate (NO₃−N) concentrations drove differences in total dissolved N concentrations because ammonium (NH₄−N) concentrations were very low (usually < 0.005 mg L⁻¹) and mean dissolved organic nitrogen (DON) concentrations were less variable than nitrate concentrations. In contrast to studies suggesting that DON levels strongly dominate in pristine watersheds, DON accounted for 24, 52, and 51% of the overall mean TDN concentration of our young (defined as predominantly in stand initiation and stem exclusion phases), middle-aged (defined as mixes of mostly understory reinitiation and older phases) and old-growth watersheds, respectively. Although other studies of cutting in unpolluted forests have suggested a harvest effect lasting 5 years or less, our young successional watersheds that were all older than 10 years still lost significantly more N, primarily as NO₃−N, than did watersheds containing more mature forests, even though all forest floor and mineral soil C:N ratios were well above levels reported in the literature for leaching of dissolved inorganic nitrogen. The influence of alder may contribute to these patterns, although hardwood cover was quite low in all watersheds; it is possible that in forested ecosystems with very low anthropogenic N inputs, even very low alder cover in riparian zones can cause elevated N exports. Only the youngest watersheds, with the highest nitrate losses, exhibited seasonal patterns of increased summer uptake by vegetation as well as flushing at the onset of fall freshets. Older watersheds with lower N losses did not exhibit seasonal patterns for any N species. The results, taken together, suggest a role for both vegetation and hydrology in N retention and loss, and add to our understanding of N cycling by successional forest ecosystems influenced by disturbance at various spatial and temporal scales in a region of relatively low anthropogenic N input.     

Effects of Soil Carbon Amendment on Nitrogen Availability and Plant Growth in an Experimental Tallgrass Prairie Restoration

Restoration of tallgrass prairie on former agricultural land is often impeded by failure to establish a diverse native species assemblage and by difficulties with nonprairie, exotic species. High levels of available soil nitrogen (N) on such sites may favor fast‐growing exotics at the expense of more slowly growing prairie species characteristic of low‐N soils. We tested whether reducing N availability through soil carbon (C) amendments could be a useful tool in facilitating successful tallgrass prairie restoration. We added 6 kg/m² hardwood sawdust to experimental plots on an abandoned agricultural field in the Sandusky Plains of central Ohio, United States, increasing soil C by 67% in the upper 15 cm. This C amendment caused a 94% reduction in annual net N mineralization and a 27% increase in soil moisture but had no effect on total N or pH. Overall, plant mass after one growing season was reduced by 64% on amended compared with unamended soil, but this effect was less for prairie forbs (?34%) than for prairie grasses (?67%) or exotics (?62%). After the second growing season, only exotics responded significantly to the soil C amendment, with a 40% reduction in mass. The N concentration of green‐leaf tissue and of senescent leaf litter was also reduced by the soil C treatment in most cases. We conclude that soil C amendment imparts several immediate benefits for tallgrass prairie restoration––notably reduced N availability, slower plant growth, and lower competition from exotic species.     

Interactions between Litter Lignin and Nitrogenitter Lignin and Soil Nitrogen Availability during Leaf Litter Decomposition in a Hawaiian Montane Forest

Previous work in a young Hawaiian forest has shown that nitrogen (N) limits aboveground net primary production (ANPP) more strongly than it does decomposition, despite low soil N availability. In this study, I determined whether (a) poor litter C quality (that is, high litter lignin) poses an overriding constraint on decomposition, preventing decomposers from responding to added N, or (b) high N levels inhibit lignin degradation, lessening the effects of added N on decomposition overall. I obtained leaf litter from one species, Metrosideros polymorpha, which dominates a range of sites in the Hawaiian Islands and whose litter lignin concentration declines with decreasing precipitation. Litter from three dry sites had lignin concentrations of 12% or less, whereas litter from two wet sites, including the study site, had lignin concentrations of more than 18%. This litter was deployed 2.5 years in a common site in control plots (receiving no added nutrients) and in N-fertilized plots. Nitrogen fertilization stimulated decomposition of the low-lignin litter types more than that of the high-lignin litter types. However, in contrast to results from temperate forests, N did not inhibit lignin decomposition. Rather, lignin decay increased with added N, suggesting that the small effect of N on decomposition at this site results from limitation of decomposition by poor C quality rather than from N inhibition of lignin decay. Even though ANPP is limited by N, decomposers are strongly limited by C quality. My results suggest that anthropogenic N deposition may increase leaf litter decomposition more in ecosystems characterized by low-lignin litter than in those characterized by high-lignin litter. Received 26 October 1999; accepted 2 June 2000.     

Nitrogen Addition Shapes Soil Phosphorus Availability in Two Reforested Tropical Forests in Southern China

Scant information is available on how soil phosphorus (P) availability responds to atmospheric nitrogen (N) deposition, especially in the tropical zones. This study examined the effect of N addition on soil P availability, and compared this effect between forest sites of contrasting land‐use history. Effects of N addition on soil properties, litterfall production, P release from decomposing litter, and soil P availability were studied in a disturbed (reforested pine forest with previous understory vegetation and litter harvesting) and a rehabilitated (reforested mixed pine/broadleaf forest with no understory vegetation and litter harvesting) tropical forest in southern China. Experimental N‐treatments (above ambient) were the following: Control (no N addition), N50 (50 kg N ha^?1 yr^?1), and N100 (100 kg N ha^?1 yr^?1). Results indicated that N addition significantly decreased soil P availability in the disturbed forest. In the rehabilitated forest, however, soil P availability was significantly increased by N addition. Decreases in soil P availability may be correlated with decreases in rates of P release from decomposing litter in the N‐treated plots, whereas the increase in soil P availability was correlated with an increase in litterfall production. Our results suggest that response of soil P availability to N deposition in the reforested tropical forests in southern China may vary greatly with temporal changes in tree species composition and soil nutrient status, caused by different land‐use practices.     

The Influence of Soil Inoculum and Nitrogen Availability on Restoration of High-Elevation Steppe Communities Invaded by Bromus tectorum

Cheatgrass ( Bromus tectorum L.) is an exotic annual grass that has invaded approximately 40,000,000 ha of rangelands in the United States, including montane ecosystems that are important habitats for wildlife and livestock. In addition to well-understood mechanisms by which Cheatgrass gains competitive advantage, recent studies have shown that Cheatgrass may also change the associated soil microbial community to impact native perennial plants and promote the persistence of Cheatgrass. Furthermore, reducing plant-available N represents a tool for initiating conditions that accelerate successional change from annual- to perennial-dominated communities. At a montane, mixed shrub–grassland Cheatgrass-dominated site in Colorado, we applied sucrose to reduce available N, and we added soil from a native plant community in order to reestablish the microbial community. This approach tested the idea that intact native soil microbial communities may enhance the beneficial effect of reducing soil N availability in a restoration setting. By the end of the experiment, reduced N availability decreased Cheatgrass by 9.8%, non-native annual/biennial plant cover by 15.0%, and increased relative perennial plant cover by 13.4%; soil inoculation reduced Cheatgrass by 7.6% and increased perennial abundance by 11.3%. Soil inoculum additions and reduced N availability both contributed toward restoring a perennial-dominated community and demonstrates that addition of native soil inoculum may be a useful tool for restoration efforts.     

崇明东滩湿地高等植被演替特征

研究了崇明东滩湿地植被演替过程中生态特征以及重要值、地上部分生物量、植被地上部分氮含量以及氮容量的季节性变化.结果表明：随着高程的升高,演替早期的代表物种海三棱藨草（Scirpus mariqueter）的重要值逐渐减小,互花米草（Spartina alterniflora）和芦苇（Phragmites australis）的重要值增大.演替过程中植物的生物多样性逐渐增加.3种植物地上部分生物量在生长季节内（3—10月）均呈单峰变化,一般在夏季（7、8月）生物量最大.3种植物地上部分氮含量在生长季节内均呈降低趋势,氮容量在生长季节内呈单峰变化.估算每年崇明东滩保护区内芦苇地上部分蓄积的氮为383.4 t,互花米草为39.3 t,海三棱藨草为50.5 t.     

森林土壤氮素可利用性的影响因素研究综述

近几十年来 ,人类对木材、纤维和其他森林资源需求的急剧增加 ,对森林的集约化经营管理成为必然趋势。由于大部分森林生态系统缺乏Ｎ素 ,因此施肥成为经济有效的途径。但是 ,由于森林中的Ｎ肥利用效率低于农业系统 ,且Ｎ肥生产成本较高 ,易造成环境中多余Ｎ素的污染 ,所以需要更有效的经营管理方法。要改进这类方法 ,则必须很好地理解全球各种森林生态系统的Ｎ素循环和Ｎ素可利用性[3 1] 。可利用性养分 (ａｖａｉｌａｂｌｅｎｕｔｒｉｅｎｔ)是指土壤中易被植物吸收同化的养分元素或化合物的数量[4 0 ] ,可以理解为植物利用土壤中易吸收和…     

Measurements and Modeling of Carbon and Nitrogen Cycling in Agroecosystems of Southern Wisconsin: Potential for SOC Sequestration during the Next 50 Years

Landmanagement practices such as no-tillage agriculture and tallgrass prairie restoration have been proposed as a possible means to sequester atmospheric carbon, helping to refurbish soil fertility and replenish organic matter lost as a result of previous agricultural management practices. However, the relationship between land-use changes and ecosystem structure and functioning is not yet understood. We studied soil and vegetation properties over a 4-year period (1995–98), and assembled measurements of microbial biomass, soil organic carbon (SOC) and nitrogen (N), N-mineralization, soil surface carbon dioxide (CO₂) flux, and leached C and N in managed (maize; Zea mays L.) and natural (prairie) ecosystems near the University of Wisconsin Agricultural Research Station at Arlington. Field data show that different management practices (tillage and fertilization) and ecosystem type (prairie vs maize) have a profound influence on biogeochemistry and water budgets between sites. These measurements were used in conjunction with a dynamic terrestrial ecosystem model, called IBIS (the Integrated Biosphere Simulator), to examine the long-term effects of land-use changes on biogeochemical cycling. Field data and modeling suggest that agricultural land management near Arlington between 1860 and 1950 caused SOC to be depleted by as much as 63% (native SOC approximately 25.1 kg C m⁻²). Reductions in N-mineralization and microbial biomass were also observed. Although IBIS simulations depict SOC recovery in no-tillage maize since the 1950s and also in the Arlington prairie since its restoration was initiated in 1976, field data suggest otherwise for the prairie. This restoration appears to have done little to increase SOC over the past 24 years. Measurements show that this prairie contained between 28% and 42% less SOC (in the top 1 m) than the no-tillage maize plots and 40%–47% less than simulated potential SOC for the site in 1999. Because IBIS simulates competition between C3 and C4 grass species, we hypothesized that current restored prairies, which include many forbs not characterized by the model, could be less capable of sequestering C than agricultural land planted entirely in monocultural grass in this region. Model output and field measurements show a potential 0.4 kg C m⁻² y⁻¹ difference in prairie net primary production (NPP). This study indicates that high-productivity C4 grasslands (NPP = 0.63 kg C m⁻² y⁻¹) and high-yield maize agroecosystems (10 Mg ha⁻¹) have the potential to sequester C at a rate of 74.5 g C m⁻² y⁻¹ and 86.3 g C m⁻² y⁻¹, respectively, during the next 50 years across southern Wisconsin. Received 28 December 1999; accepted 11 December 2000.     

Quantification of the Nitrogen Cycle in a Prairie Stream

Nitrogen (N) was added for 35 days in the form of ¹⁵NH₄Cl to Kings Creek on Konza Prairie, Kansas. Standing stocks of N in key compartments (that is, nutrients, detritus, organisms) were quantified, and the amount of labeled N entering the compartments was analyzed. These data were used to calculate turnover and flux rates of N cycling through the food web, as well as nutrient transformation rates. Inorganic N pools turned over much more rapidly in the water column of this stream than in pelagic systems where comparable measurements have been made. As with other systems, the mass of ammonium was low but it was the key compartment mediating nutrient flux through the ecosystem, whereas dissolved organic N, the primary component of N flux through the system, is not actively cycled. Nitrification was also a significant flux of N in the stream, with rates in the water column and surface of benthos accounting for approximately 10% of the total ammonium uptake. Primary consumers assimilated 67% of the inorganic N that entered benthic algae and microbes. Predators acquired 23% of the N that consumers obtained. Invertebrate collectors, omnivorous crayfish (Orconectes spp.), and invertebrate shredders dominated the N flux associated with primary consumers. Mass balance calculations indicated that at least 23% of the 309 mg of ¹⁵N added during the 35 days of release was retained within the 210-m stream reach during the release. Overall, the rates of turnover of N in organisms and organic substrata were significantly greater when C:N was low. This ratio may be a surrogate for biological activity with regard to N flux in streams. Received 2 August 1999; accepted 18 July 2000.     

Spatial Variability of Soil Properties in the Shortgrass Steppe: The Relative Importance of Topography, Grazing, Microsite, and Plant Species in Controlling Spatial Patterns

We conducted a study to evaluate the relative importance of topography, grazing, the location of individual plants (microsite), and plant species in controlling the spatial variability of soil organic matter in shortgrass steppe ecosystems. We found that the largest spatial variation occurs in concert with topography and with microsite-scale heterogeneity, with relatively little spatial variability due to grazing or to plant species. Total soil C and N, coarse and fine particulate organic matter C and N, and potentially mineralizable C were significantly affected by topography, with higher levels in toeslope positions than in midslopes or summits. Soils beneath individual plants (Bouteloua gracilis and Opuntia polyacantha) were elevated by 2–3 cm relative to surrounding soils. All pools of soil organic matter were significantly higher in the raised hummocks directly beneath plants than in the soil surface of interspaces or this layer under plants. High levels of mineral material in the hummocks suggest that erosion is an important process in their formation, perhaps in addition to biotic accumulation of litter beneath individual plants. Over 50 y of heavy grazing by cattle did not have a significant effect on most of the soil organic matter pools we studied. This result was consistent with our hypothesis that this system, with its strong dominance of belowground organic matter, is minimally influenced by aboveground herbivory. In addition, soils beneath two of the important plant species of the shortgrass steppe, B. gracilis and O. polyacantha, differed little from one another. The processes that create spatial variability in shortgrass steppe ecosystems do not affect all soil organic matter pools equally. Topographic variability, developing over pedogenic time scales (centuries to thousands of years), has the largest effect on the most stable pools of soil organic matter. The influence of microsite is most evident in the pools of organic matter that turn over at time scales that approximate the life span of individual plants (years to decades and centuries).     

Restoration of a Landfill Site in Berlin, Germany by Spontaneous and Directed Succession

We report on spontaneous and directed succession on a dry sandy landfill site of low fertility at Berlin‐Malchow, Germany. Changes in species composition and cover were followed on unmown and mown permanent plots of 2 × 2 m size through 5 years of vegetation development. Species richness on unmown plots was relatively constant during the time of observation, with 20 to 25 species per 4 m². Total cover of unmown plots continuously increased from approximately 10% in the first year to 80% in the fifth year. There are no clearly discernible sequential successional stages until present. The species composition includes species of all life forms, which colonized the site immediately after the initiation of the succession process representing the initial floristic composition type of vegetation development. However, perennial grasses and herbs gradually increased in cover up to approximately 40%. Woody plants were also present from the first year of succession and increased up to more than 20% cover in the fifth year, forming a shrub layer (>0.5 m) after the second year. Mowing significantly increased species richness, which was evident from the third year onward. This effect was mainly due to the reduction of the tall perennial grass Calamagrostis epigejos. Solidago canadensis and woody species were also significantly affected (lower cover and height), whereas short perennial herbs like Plantago lanceolata and Trifolium repens benefited from mowing.