Effect of Sheep Urine Deposition on the Bacterial Community Structure in an Acidic Upland Grassland Soil

The effect of the addition of synthetic sheep urine (SSU) and plant species on the bacterial community composition of upland acidic grasslands was studied using a microcosm approach. Low, medium, and high concentrations of SSU were applied to pots containing plant species typical of both unimproved (Agrostis capillaris) and agriculturally improved (Lolium perenne) grasslands, and harvests were carried out 10 days and 50 days after the addition of SSU. SSU application significantly increased both soil pH (P < 0.005), with pH values ranging from pH 5.4 (zero SSU) to pH 6.4 (high SSU), and microbial activity (P < 0.005), with treatment with medium and high levels of SSU displaying significantly higher microbial activity (triphenylformazan dehydrogenase activity) than treatment of soil with zero or low concentrations of SSU. Microbial biomass, however, was not significantly altered by any of the SSU applications. Plant species alone had no effect on microbial biomass or activity. Bacterial community structure was profiled using bacterial automated ribosomal intergenic spacer analysis. Multidimensional scaling plots indicated that applications of high concentrations of SSU significantly altered the bacterial community composition in the presence of plant species but at different times: 10 days after application of high concentrations of SSU, the bacterial community composition of L. perenne-planted soils differed significantly from those of any other soils, whereas in the case of A. capillaris-planted soils, the bacterial community composition was different 50 days after treatment with high concentrations of SSU. Canonical correspondence analysis also highlighted the importance of interactions between SSU addition, plant species, and time in the bacterial community structure. This study has shown that the response of plants and bacterial communities to sheep urine deposition in grasslands is dependent on both the grass species present and the concentration of SSU applied, which may have important ecological consequences for agricultural grasslands.     

Burning and Grazing Management in a California Grassland: Effect on Bunchgrass Seed Viability

Prescribed fire is an important management tool for reducing the dominance of non‐native species in annual grasslands; both annual and perennial native species show strong vegetative responses in the subsequent growing season. However, although the post‐fire contribution of native species to the seed bank is assumed to be larger than in pretreatment years, the effects on seed quality, particularly viability and longevity, are not well understood. In this study, I germinated Nassella pulchra (purple needlegrass) seed that had been stored for 10 years after collection from target plants receiving treatment combinations of summer burning and grazing by sheep. Seeds from burned plants were larger and had higher germinability than seed from unburned plants. Seeds from plants that were both burned and grazed had the highest germination. The strong relationship between long‐term viability and seed size suggests greater maternal provisioning and increased seed quality subsequent to burning and grazing. I conclude that managing for seed quality may be a useful approach for conservation of native species in California's critically endangered grassland habitats.     

Grazing and Ecosystem Carbon Storage in the North American Great Plains

Isotopic signatures of ¹³C were used to quantify the relative contributions of C₃ and C₄ plants to whole-ecosystem C storage (soil+plant) in grazed and ungrazed sites at three distinct locations (short-, mid- and tallgrass communities) along an east–west environmental gradient in the North American Great Plains. Functional group composition of plant communities, the source and magnitude of carbon inputs, and total ecosystem carbon storage displayed inconsistent responses to long-term livestock grazing along this gradient. C₄ plants [primarily Bouteloua gracilis (H.B.K.) Lag ex Steud.] dominated the long-term grazed site in the shortgrass community, whereas the ungrazed site was co-dominated by C₃ and C₄ species; functional group composition did not differ between grazed and ungrazed sites in the mid- and tallgrass communities. Above-ground biomass was lower, but the relative proportion of fine root biomass was greater, in grazed compared to ungrazed sites at all three locations. The grazed site of the shortgrass community had 24% more whole-ecosystem carbon storage compared to the ungrazed site (4022 vs. 3236 g C m⁻²). In contrast, grazed sites at the mid- and tallgrass communities had slightly lower (8%) whole-ecosystem carbon storage compared to ungrazed sites (midgrass: 7970 vs. 8683 g C m⁻²; tallgrass: 8273 vs. 8997 g C m⁻²). Differential responses between the shortgrass and the mid- and tallgrass communities with respect to grazing and whole-ecosystem carbon storage are likely a result of: (1) maintenance of larger soil organic carbon (SOC) pools in the mid- and tallgrass communities (7476–8280 g C m⁻²) than the shortgrass community (2517–3307 g C m⁻²) that could potentially buffer ecosystem carbon fluxes, (2) lower root carbon/soil carbon ratios in the mid- and tallgrass communities (0.06–0.10) compared to the shortgrass community (0.20–0.27) suggesting that variation in root organic matter inputs would have relatively smaller effects on the size of the SOC pool, and (3) the absence of grazing-induced variation in the relative proportion of C₃ and C₄ functional groups in the mid- and tallgrass communities. We hypothesize that the magnitude and proportion of fine root mass within the upper soil profile is a principal driver mediating the effect of community composition on the biogeochemistry of these grassland ecosystems.     

土地利用变化对草原生态系统土壤碳贮量的影响

本文概述了长期非持续性土地利用（草原开垦和过度放牧）对草原生态系统土壤碳贮量影响的有关研究成果和进展,指出了今后这方面研究的重点内容和对策。     

Effects of Burning and Grazing on a Coastal California Grassland

We tested the effects of fall burning and protection from livestock grazing as management to enhance native grasses on a coastal grassland in central California. Plants from the Mediterranean, introduced beginning in the late 1700s, have invaded and now dominate most of California's grasslands. Coastal grasslands are generally less degraded than those inland and have higher potential for restoration and conservation. Productivity of the experimental plots varied annually and declined over the course of the study because of rainfall patterns. Foliar cover of the native Danthonia californica (California oatgrass) increased more under grazing than grazing exclusion and did not respond to burning. Two other natives, Nassella pulchra (purple needlegrass) and Nassella lepida (foothill needlegrass), responded variably to treatments. The response of N. pulchra differed from that reported on more inland sites in California. Restoring these grasslands is complicated by differing responses of target species to protection from grazing and burning. The current practice of managing to enhance single species of native plants (e.g., N. pulchra) may be detrimental to other equally important native species.     

Managing Semi-Arid Rangelands for Carbon Storage: Grazing and Woody Encroachment Effects on Soil Carbon and Nitrogen

High grazing intensity and wide-spread woody encroachment may strongly alter soil carbon (C) and nitrogen (N) pools. However, the direction and quantity of these changes have rarely been quantified in East African savanna ecosystem. As shifts in soil C and N pools might further potentially influence climate change mitigation, we quantified and compared soil organic carbon (SOC) and total soil nitrogen (TSN) content in enclosures and communal grazing lands across varying woody cover i.e. woody encroachment levels. Estimated mean SOC and TSN stocks at 0–40 cm depth varied across grazing regimes and among woody encroachment levels. The open grazing land at the heavily encroached site on sandy loam soil contained the least SOC (30 ± 2.1 Mg ha^-1) and TSN (5 ± 0.57 Mg ha^-1) while the enclosure at the least encroached site on sandy clay soil had the greatest mean SOC (81.0 ± 10.6 Mg ha^-1) and TSN (9.2 ± 1.48 Mg ha^-1). Soil OC and TSN did not differ with grazing exclusion at heavily encroached sites, but were twice as high inside enclosure compared to open grazing soils at low encroached sites. Mean SOC and TSN in soils of 0–20 cm depth were up to 120% higher than that of the 21–40 cm soil layer. Soil OC was positively related to TSN, cation exchange capacity (CEC), but negatively related to sand content. Our results show that soil OC and TSN stocks are affected by grazing, but the magnitude is largely influenced by woody encroachment and soil texture. We suggest that improving the herbaceous layer cover through a reduction in grazing and woody encroachment restriction are the key strategies for reducing SOC and TSN losses and, hence, for climate change mitigation in semi-arid rangelands.     

Burning and Grazing Management in a California Grassland: Growth, Mortality, and Recruitment of Nassella pulchra

Abstract Annual grasslands in California are often managed with seasonal grazing and prescribed burning on the assumption that such practices have long‐term benefits for native species. Mature native perennial bunchgrasses, particularly Nassella pulchra (purple needlegrass), are often the focal species, although very little is known about responses at different life history stages. Thus, important questions remain about long‐term population dynamics of both mature plants and seedling recruitment. In plots receiving repeated grazing and burning events over 7 years, mortality of mature plants was threefold higher on mounds than on intermounds and likely reflected increased competition intensity associated with increased resource availability in deeper soil. Burning and grazing treatments had strong positive effects on basal area of mature N. pulchra. However, plants in grazed plots that were not burned contained considerable standing dead biomass. Topographic location strongly influenced growth as intermound plants grew relatively more than mound plants, but the effects on growth of burning and grazing did not vary with topographic location. In mapped plots N. pulchra recruitment was very low, and overall density dropped an average of 31%. However, a significant time‐by‐burning effect indicated that survival was significantly higher in burned plots. After 7 years of repeated treatments, effects of burning and grazing management on mature N. pulchra were positive but not for all phenological stages. Understanding long‐term influence of management on bunchgrass populations may not be easy to determine because short‐term results may not reflect long‐term responses and some life cycle dynamics may be observed only over very long periods.     

Seasonal Invasion Dynamics in a Spatially Heterogeneous River with Fluctuating Flows

A key problem in environmental flow assessment is the explicit linking of the flow regime with ecological dynamics. We present a hybrid modeling approach to couple hydrodynamic and biological processes, focusing on the combined impact of spatial heterogeneity and temporal variability on population dynamics. Studying periodically alternating pool-riffle rivers that are subjected to seasonally varying flows, we obtain an invasion ratchet mechanism. We analyze the ratchet process for a caricature model and a hybrid physical–biological model. The water depth and current are derived from a hydrodynamic equation for variable stream bed water flows and these quantities feed into a reaction-diffusion-advection model that governs population dynamics of a river species. We establish the existence of spreading speeds and the invasion ratchet phenomenon, using a mixture of mathematical approximations and numerical computations. Finally, we illustrate the invasion ratchet phenomenon in a spatially two-dimensional hydraulic simulation model of a meandering river structure. Our hybrid modeling approach strengthens the ecological component of stream hydraulics and allows us to gain a mechanistic understanding as to how flow patterns affect population survival.     

Foraging Behavior Patterns of Sheep and Horses Under a Mixed Species Grazing System

The research objective was to assess the behavior patterns of the Polish Konik horse breed and the Uhruska variety of the Polish Lowland Sheep breed under a mixed-grazing system, and their relationship with climatic factors. The observation included 4 adult horses, 27 ewes with lambs and 10 primiparous ewes. The behavior of the animals and the weather conditions were recorded at 60-min intervals. Horses and sheep displayed similarities in both species-specific behavioral patterns and timing of grazing activity, and the duration of foraging sessions was mostly influenced by time of day, lower temperature and relative humidity. In addition, the two species showed more interest in the watering place in the afternoon, and drinking frequency was mainly dependent on air temperature. The animals that pastured together gradually began to mix, but at first, they remained within their species group. Both animal species can be used for environmental protection and landscape conservation making rational use of pastures within protected areas.     

Sheep Grazing Decreases Organic Carbon and Nitrogen Pools in the Patagonian Steppe: Combination of Direct and Indirect Effects

We explored the net effects of grazing on soil C and N pools in a Patagonian shrub–grass steppe (temperate South America). Net effects result from the combination of direct impacts of grazing on biogeochemical characteristics of microsites with indirect effects on relative cover of vegetated and unvegetated microsites. Within five independent areas, we sampled surface soils in sites subjected to three grazing intensities: (1) ungrazed sites inside grazing exclosures, (2) moderately grazed sites adjacent to them, and (3) intensely grazed sites within the same paddock. Grazing significantly reduced soil C and N pools, although this pattern was clearest in intensely grazed sites. This net effect was due to the combination of a direct reduction of soil N content in bare soil patches, and indirect effects mediated by the increase of the cover of bare soil microsites, with lower C and N content than either grass or shrub microsites. This increase in bare soil cover was accompanied by a reduction in cover of preferred grass species and standing dead material. Finally, stable isotope signatures varied significantly among grazed and ungrazed sites, with δ¹⁵N and δ¹³C significantly depleted in intensely grazed sites, suggesting reduced mineralization with increased grazing intensity. In the Patagonian steppe, grazing appears to exert a negative effect on soil C and N cycles; sound management practices must incorporate the importance of species shifts within life form, and the critical role of standing dead material in maintaining soil C and N stocks and biogeochemical processes. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Author Contributions  RAG designed study, performed research, analyzed data, wrote the paper; ATA designed study, wrote the paper; CGGM designed study, performed research, analyzed data; MGP performed research; OES designed study; RBJ designed study, contributed new methods.     

Interactive Effects of Time, CO2, N, and Diversity on Total Belowground Carbon Allocation and Ecosystem Carbon Storage in a Grassland Community

Predicting if ecosystems will mitigate or exacerbate rising CO₂ requires understanding how elevated CO₂ will interact with coincident changes in diversity and nitrogen (N) availability to affect ecosystem carbon (C) storage. Yet achieving such understanding has been hampered by the difficulty of quantifying belowground C pools and fluxes. Thus, we used mass balance calculations to quantify the effects of diversity, CO₂, and N on both the total amount of C allocated belowground by plants (total belowground C allocation, TBCA) and ecosystem C storage in a periodically burned, 8-year Minnesota grassland biodiversity, CO₂, and N experiment (BioCON). Annual TBCA increased in response to elevated CO₂, enriched N, and increasing diversity. TBCA was positively related to standing root biomass. After removing the influence of root biomass, the effect of elevated CO₂ remained positive, suggesting additional drivers of TBCA apart from those that maintain high root biomass. Removing root biomass effects resulted in the effects of N and diversity becoming neutral or negative (depending on year), suggesting that the positive effects of diversity and N on TBCA were related to treatment-driven differences in root biomass. Greater litter production in high diversity, elevated CO₂, and enhanced N treatments increased annual ecosystem C loss in fire years and C gain in non-fire years, resulting in overall neutral C storage rates. Our results suggest that frequently burned grasslands are unlikely to exhibit enhanced C sequestration with increasing atmospheric CO₂ levels or N deposition.     

The Success of a Rotational Grazing System in Conserving the Diversity of Chalk Grassland Auchenorrhyncha

A complex rotational grazing trial on a south-facing slope of chalk grassland at the Old Winchester Hill National Nature Reserve is briefly introduced. The responses of 23 numerous species of Auchenorrhyncha, and of species richness (S) and total abundance (N), from 1981 to 1985 are described. The greatest effects were those of variation between years, between positions on the hillside (top, middle and bottom) and between grazing plots within these positions. 10 (of the total of 23) species favoured the top of the slope, where the vegetation was significantly taller than in the middle or at the bottom. S, N and the numbers of 8 species were significantly lower on plots grazed in the year of sampling compared with ungrazed plots. Early (vs. late) grazing significantly reduced S, N and the abundance of two species, but increased the numbers of Macrosteles laevis. S, N and the abundance of 13 species was significantly and positively correlated with vegetation height measured early (May–June) and late (July–October); the numbers of 4 other species were so correlated with the latter height only. The significance of the results is discussed in relation to the management of grassland nature reserves for the maintenance of high invertebrate diversity. It is concluded that rotational management is an important and valuable system, but suggested that such systems should be as simple as possible whilst remaining adequate to achieve conservation objectives.     

Traditional Cattle Grazing in a Mosaic Alkali Landscape: Effects on Grassland Biodiversity along a Moisture Gradient

Extensively managed pastures are of crucial importance in sustaining biodiversity both in local- and landscape-level. Thus, re-introduction of traditional grazing management is a crucial issue in grassland conservation actions worldwide. Traditional grazing with robust cattle breeds in low stocking rates is considered to be especially useful to mimic natural grazing regimes, but well documented case-studies are surprisingly rare on this topic. Our goal was to evaluate the effectiveness of traditional Hungarian Grey cattle grazing as a conservation action in a mosaic alkali landscape. We asked the following questions: (i) How does cattle grazing affect species composition and diversity of the grasslands? (ii) What are the effects of grazing on short-lived and perennial noxious species? (iii) Are there distinct effects of grazing in dry-, mesophilous- and wet grassland types? Vegetation of fenced and grazed plots in a 200-ha sized habitat complex (secondary dry grasslands and pristine mesophilous- and wet alkali grasslands) was sampled from 2006–2009 in East-Hungary. We found higher diversity scores in grazed plots compared to fenced ones in mesophilous- and wet grasslands. Higher cover of noxious species was typical in fenced plots compared to their grazed counterparts in the last year in every studied grassland type. We found an increasing effect of grazing from the dry- towards the wet grassland types. The year-to-year differences also followed similar pattern: the site-dependent effects were the lowest in the dry grassland and an increasing effect was detected along the moisture gradient. We found that extensive Hungarian Grey cattle grazing is an effective tool to suppress noxious species and to create a mosaic vegetation structure, which enables to maintain high species richness in the landscape. Hungarian Grey cattle can feed in open habitats along long moisture gradient, thus in highly mosaic landscapes this breed can be the most suitable livestock type.     

Optimizing Radio Retention and Minimizing Radio Impacts in a Field Study of Upland Sandpipers

Abstract: Two challenges in wildlife telemetry are optimizing the duration of transmitter attachment and minimizing the impacts of radios on the behavior and demography of the study animal. We tested 4 methods of radio attachment for a breeding population of upland sandpipers (Bartramia longicauda) under natural conditions at a tallgrass prairie site in Kansas, USA. To estimate radio retention and weekly survival rates, we used the nest survival model of Program MARK. Radio retention was lowest at the start and the end of the breeding period. The expected duration of radio retention was 1.8 years for a leg-loop harness, 40 days for radios glued to clipped feathers, 26 days for radios glued directly to feathers, and 7 days for radios glued to bare skin. Few radiomarked birds died during our study, but 4 of 8 mortality events were discovered within one week of radiomarking. Both glue and harnesses increased predation risk immediately after radio attachment. The weekly probability of survival was high after a 1-week acclimation period, and the expected survival for a 10-week breeding period was similar in males and females. Attachment of radios with glue had no effect on annual return rates. However, attachment of radios with leg harnesses resulted in lower return rates among radiomarked birds than birds without radios. Radios attached with glue were shed in <1 year but radios attached with harnesses were retained for up to 1-2 years. Our results indicate a tradeoff between optimizing radio retention and minimizing impacts on demography. Glue techniques had retention rates that were suitable for only short-term studies, but attachment with glue had no long-term effect on annual return rates. Leg harnesses provided effective radio retention that had little effect on survival rates during the stationary breeding period, but resulted in lower annual return rates. Robust estimates of radio retention and survival will assist researchers in selecting attachment techniques that best meet the study goals of future telemetry projects.     

Methane-Oxidizing Bacteria in a California Upland Grassland Soil: Diversity and Response to Simulated Global Change

We investigated the diversity of methane-oxidizing bacteria (i.e., methanotrophs) in an annual upland grassland in northern California, using comparative sequence analysis of the pmoA gene. In addition to identifying type II methanotrophs commonly found in soils, we discovered three novel pmoA lineages for which no cultivated members have been previously reported. These novel pmoA clades clustered together either with clone sequences related to “RA 14” or “WB5FH-A,” which both represent clusters of environmentally retrieved sequences of putative atmospheric methane oxidizers. Conservation of amino acid residues and rates of nonsynonymous versus synonymous nucleotide substitution in these novel lineages suggests that the pmoA genes in these clades code for functionally active methane monooxygenases. The novel clades responded to simulated global changes differently than the type II methanotrophs. We observed that the relative abundance of type II methanotrophs declined in response to increased precipitation and increased atmospheric temperature, with a significant antagonistic interaction between these factors such that the effect of both together was less than that expected from their individual effects. Two of the novel clades were not observed to respond significantly to these environmental changes, while one of the novel clades had an opposite response, increasing in relative abundance in response to increased precipitation and atmospheric temperature, with a significant antagonistic interaction between these factors.     

Trampling and Spatial Heterogeneity Explain Decomposer Abundances in a Sub-Arctic Grassland Subjected to Simulated Reindeer Grazing

Mammal grazing is composed of three mechanisms—removal of foliar tissue (defoliation), return of nutrients via dung and urine (fertilization), and trampling. To evaluate the relative role of these mechanisms in the effect of reindeer grazing on soil biota in northern grasslands, we subjected experimental plots in a sub-arctic alpine meadow to defoliation, fertilization (using NPK-solution), simulated trampling, and their factorial combinations once a year from 2002 to 2004 and measured the response of plants and decomposers (including microbes, nematodes, collembolans, and enchytraeids) in 2004. Trampling affected both plant and decomposer communities: the coverage of the moss Pleurozium schreberi and the sedge Carex vaginata, as well as the abundance of collembolans and enchytraeids were reduced in trampled plots. Trampling and fertilization also interacted significantly, with fertilization increasing the abundance of bacteria and bacterial-feeding and omnivorous nematodes in trampled plots only, and trampling decreasing fungal biomass in non-fertilized plots only. Defoliation had no overall effects on plants or decomposers. Nematode genera were not affected by the experimental treatments, but nematode and plant communities were significantly associated, and all decomposer biota, except collembolans, were strongly affected by the spatial heterogeneity of the study site. Our results indicate that trampling may have larger and defoliation and fertilization smaller roles than anticipated in explaining reindeer grazing effects in sub-arctic grasslands. However, even the effects of trampling seem to be outweighed by the spatial heterogeneity of decomposer abundances. This suggests that in sub-arctic grasslands spatial variation in abiotic factors can be a more important factor than grazing in controlling soil biota abundances. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Author contributions  LIS was involved in planning of the study, fieldwork, lab work, analysis, and wrote the article with contributions from all other authors; JM and MMK conceived the study and contributed to the data analyses; MMK further contributed to the field work and JM refined the final appearance of the text; JO started and was responsible for managing the field experiment, collecting the plant data and gave advice on statistical analysis.     

The Response of a Heterogeneous Upland Boreal Shield Catchment to a Short Term NO3 - Addition

Boreal Shield rocky ridges at the Experimental Lakes Area, northwestern Ontario, contain two plant/soil communities with contrasting N cycles. Picea mariana–Pinus banksiana”forest islands” are N limited whereas the lichen, moss, and grass community (or “lichen patches”) on the surrounding bedrock outcrops appear intrinsically N saturated. The potential for this landscape to retain a N input of eightfold ambient levels was tested with a 2-y addition of 40 kg N ha^-1 y^-1 as NaNO₃ to one small catchment (0.40 ha). The elevated N input was poorly retained by the whole catchment during snowmelt. However, during the growing season, N retention in the treated catchment remained as efficient as in references. Forest islands and bedrock surfaces responded in opposite fashions to the elevated N input. By the second year of N addition, bedrock surfaces no longer retained additional N inputs. In contrast, N-amended and reference forest islands retained a similar proportion of N inputs, indicating that forest islands did not become N saturated. The response of the whole catchment to N addition was more similar to forest islands than bedrock surfaces. Even if forest islands only cover a small proportion of catchment area, they can have a strong impact on whole catchment element export because most of the water must move through at least one island before leaving the system. Because the different components of the boreal shield landscape are hydrologically connected, N saturation may occur as a cascading effect in this ecosystem. Monitoring boreal shield landscapes by using outlets at the lower end of the hydrological cascade can fail to detect the impacts of perturbations such as increased N deposition on upper components. Received 1 December 1998; accepted 8 April 1999.     

草地生态系统碳循环及其对全球变化的响应I 碳循环的分室模型、碳输入与贮量

概述了草地生态系统碳素循环的一般特征,介绍了一个草地碳循环的分室模型;对世界草地生态系统的碳贮量和碳输入量的有关数据进行了归纳整理,并初步讨论了草地生态系统在全球碳循环中的作用。     

Long-Term Nitrogen Addition Does Not Increase Soil Carbon Storage or Cycling Across Eight Temperate Forest and Grassland Sites on a Sandy Outwash Plain

Ecosystems - Experimental nitrogen (N) deposition generally inhibits decomposition and promotes carbon (C) accumulation in soils, but with substantial variation among studies. Differences in...