Soil C and N responses to woody plant expansion in a mesic grassland

Changes in land management and reductions in fire frequency have contributed to increased cover of woody species in grasslands worldwide. These shifts in plant community composition have the potential to alter ecosystem function, particularly through changes in soil processes and properties. In semi-arid grasslands, the invasion of shrubs and trees is often accompanied by increases in soil resources and more rapid N and C cycling. We assessed the effects of shrub encroachment in a mesic grassland in Kansas (USA) on soil CO₂ flux, extractable inorganic N, and N mineralization beneath shrub communities (Cornus drummondii) and surrounding undisturbed grassland sites. In this study, a shift in plant community composition from grassland to shrubland resulted in a 16% decrease in annual soil CO₂ flux(4.78 kg CO₂ m^–2 year^–1 for shrub dominated sites versus 5.84 kg CO₂ m^–2 year^–1 for grassland sites) with no differences in total soil C or N or inorganic N. There was considerable variability in N mineralization rates within sites, which resulted in no overall difference in cumulative N mineralized during this study (4.09 g N m^–2 for grassland sites and 3.03 g N m^–2 for shrub islands). These results indicate that shrub encroachment into mesic grasslands does not significantly alter N availability (at least initially), but does alter C cycling by decreasing soil CO₂ flux.     

Consequences of shrub expansion in mesic grassland: Resource alterations and graminoid responses

Abstract. In the mesic grasslands of the central United States, the shrub Cornus drummondii has undergone widespread expansion in the absence of recurrent fire. We quantified alterations in light, water and N caused by C. drummondii expansion in tall‐grass prairie and assessed the hypothesis that these alterations are consistent with models of resource enrichment by woody plants. Responses in graminoid species, particularly the dominant C₄ grass Andropogon gerardii, were concurrently evaluated. We also removed established shrub islands to quantify their legacy effect on resource availability and assess the capability of this grassland to recover in sites formerly dominated by woody plants. The primary effect of shrub expansion on resource availability was an 87% reduction in light available to the herbaceous understorey. This reduced C uptake and N use efficiency in A. gerardii and lowered graminoid cover and ANPP at the grass‐shrub ecotone relative to undisturbed grassland. Shrub removal created a pulse in light and N availability, eliciting high C gain in A. gerardii in the first year after removal. By year two, light and N availability within shrub removal areas returned to levels typical of grassland, as had graminoid cover and ANPP were similar to those in open grassland. Recovery within central areas of shrub removal sites lagged behind that at the former grass‐shrub ecotone. These results indicate that the apparent alternative stable state of C. drummondii dominance in tall‐grass prairie is biotically maintained and driven by reductions in light, rather than resource enrichment. Within areas of shrub removal, the legacy effect of C. drummondii dominance is manifest primarily through the loss of rhizomes of the dominant grasses, rather than any long‐term changes in resource availability. C. drummondii removal facilitates grassland recovery, but the effort required to initiate this transition is a significant cost of woody plant expansion in mesic grasslands. Prevention of woody plant expansion in remnant tall‐grass prairies is, therefore, a preferred management option.     

Organic vs. Conventional Grassland Management: Do 15N and 13C Isotopic Signatures of Hay and Soil Samples Differ?

Distinguishing organic and conventional products is a major issue of food security and authenticity. Previous studies successfully used stable isotopes to separate organic and conventional products, but up to now, this approach was not tested for organic grassland hay and soil. Moreover, isotopic abundances could be a powerful tool to elucidate differences in ecosystem functioning and driving mechanisms of element cycling in organic and conventional management systems. Here, we studied the δ¹⁵N and δ¹³C isotopic composition of soil and hay samples of 21 organic and 34 conventional grasslands in two German regions. We also used Δδ¹⁵N (δ¹⁵N plant - δ¹⁵N soil) to characterize nitrogen dynamics. In order to detect temporal trends, isotopic abundances in organic grasslands were related to the time since certification. Furthermore, discriminant analysis was used to test whether the respective management type can be deduced from observed isotopic abundances.Isotopic analyses revealed no significant differences in δ¹³C in hay and δ¹⁵N in both soil and hay between management types, but showed that δ¹³C abundances were significantly lower in soil of organic compared to conventional grasslands. Δδ¹⁵N values implied that management types did not substantially differ in nitrogen cycling. Only δ¹³C in soil and hay showed significant negative relationships with the time since certification. Thus, our result suggest that organic grasslands suffered less from drought stress compared to conventional grasslands most likely due to a benefit of higher plant species richness, as previously shown by manipulative biodiversity experiments. Finally, it was possible to correctly classify about two third of the samples according to their management using isotopic abundances in soil and hay. However, as more than half of the organic samples were incorrectly classified, we infer that more research is needed to improve this approach before it can be efficiently used in practice.     

Shrub expansion stimulates soil C and N storage along a coastal soil chronosequence

Expansion of woody vegetation in grasslands is a worldwide phenomenon with implications for C and N cycling at local, regional and global scales. Although woody encroachment is often accompanied by increased annual net primary production (ANPP) and increased inputs of litter, mesic ecosystems may become sources for C after woody encroachment because stimulation of soil CO₂ efflux releases stored soil carbon. Our objective was to determine if young, sandy soils on a barrier island became a sink for C after encroachment of the nitrogen‐fixing shrub Morella cerifera, or if associated stimulation of soil CO₂ efflux mitigated increased litterfall. We monitored variations in litterfall in shrub thickets across a chronosequence of shrub expansion and compared those data to previous measurements of ANPP in adjacent grasslands. In the final year, we quantified standing litter C and N pools in shrub thickets and soil organic matter (SOM), soil organic carbon (SOC), soil total nitrogen (TN) and soil CO₂ efflux in shrub thickets and adjacent grasslands. Heavy litterfall resulted in a dense litter layer storing an average of 809 g C m^?2 and 36 g N m^?2. Although soil CO₂ efflux was stimulated by shrub encroachment in younger soils, soil CO₂ efflux did not vary between shrub thickets and grasslands in the oldest soils and increases in CO₂ efflux in shrub thickets did not offset contributions of increased litterfall to SOC. SOC was 3.6–9.8 times higher beneath shrub thickets than in grassland soils and soil TN was 2.5–7.7 times higher under shrub thickets. Accumulation rates of soil and litter C were highest in the youngest thicket at 101 g m^?2 yr^?1 and declined with increasing thicket age. Expansion of shrubs on barrier islands, which have low levels of soil carbon and high potential for ANPP, has the potential to significantly increase ecosystem C sequestration.     

Aboveground litter quality changes may drive soil organic carbon increase after shrub encroachment into mountain grasslands

Shrub encroachment into grasslands is ubiquitous but its impact on soil organic C (SOC) remains unclear. In previous work we had observed that shrub encroachment into mesic mountain grasslands increased SOC content. Here we sought the mechanisms of this increase. To this end, we assessed aboveground and belowground production for a conifer shrub (Juniperus communis L), a legume shrub (Cytisus balansae ssp. europaeus (G. López & Jarvis) Muñoz Garmendia) and grass (Festuca eskia Ramond ex DC), together with decomposition rates for both aboveground litter and roots. Belowground C net inputs do not clearly explain SOC increase: grass root production was higher than that of either shrub and the decomposition rate of grass roots was the lowest. Aboveground C net inputs were only slightly greater in shrubs than in grass, but the decomposition rate of litter of both shrubs was much lower than that of grass. The decomposition of conifer litter was N-limited, whereas that of legume shrub litter was P-limited. Thus we conclude that the SOC increases after shrub encroachment into mesic grasslands probably as a result of higher recalcitrance of shrub aboveground litter relative to grass litter.     

Priming of Soil Carbon Decomposition in Two Inner Mongolia Grassland Soils following Sheep Dung Addition: A Study Using 13C Natural Abundance Approach

To investigate the effect of sheep dung on soil carbon (C) sequestration, a 152 days incubation experiment was conducted with soils from two different Inner Mongolian grasslands, i.e. a Leymus chinensis dominated grassland representing the climax community (2.1% organic matter content) and a heavily degraded Artemisia frigida dominated community (1.3% organic matter content). Dung was collected from sheep either fed on L. chinensis (C₃ plant with δ¹³C = −26.8‰; dung δ¹³C = −26.2‰) or Cleistogenes squarrosa (C₄ plant with δ¹³C = −14.6‰; dung δ¹³C = −15.7‰). Fresh C₃ and C₄ sheep dung was mixed with the two grassland soils and incubated under controlled conditions for analysis of ¹³C-CO₂ emissions. Soil samples were taken at days 17, 43, 86, 127 and 152 after sheep dung addition to detect the δ¹³C signal in soil and dung components. Analysis revealed that 16.9% and 16.6% of the sheep dung C had decomposed, of which 3.5% and 2.8% was sequestrated in the soils of L. chinensis and A. frigida grasslands, respectively, while the remaining decomposed sheep dung was emitted as CO₂. The cumulative amounts of C respired from dung treated soils during 152 days were 7–8 times higher than in the un-amended controls. In both grassland soils, ca. 60% of the evolved CO₂ originated from the decomposing sheep dung and 40% from the native soil C. Priming effects of soil C decomposition were observed in both soils, i.e. 1.4 g and 1.6 g additional soil C kg⁻¹ dry soil had been emitted as CO₂ for the L. chinensis and A. frigida soils, respectively. Hence, the net C losses from L. chinensis and A. frigida soils were 0.6 g and 0.9 g C kg⁻¹ soil, which was 2.6% and 7.0% of the total C in L. chinensis and A. frigida grasslands soils, respectively. Our results suggest that grazing of degraded Inner Mongolian pastures may cause a net soil C loss due to the positive priming effect, thereby accelerating soil deterioration.     

Climatic Control on Plant and Soil δ13C along an Altitudinal Transect of Lushan Mountain in Subtropical China: Characteristics and Interpretation of Soil Carbon Dynamics

Decreasing temperature and increasing precipitation along altitude gradients are typical mountain climate in subtropical China. In such a climate regime, identifying the patterns of the C stable isotope composition (δ¹³C) in plants and soils and their relations to the context of climate change is essential. In this study, the patterns of δ¹³C variation were investigated for tree leaves, litters, and soils in the natural secondary forests at four altitudes (219, 405, 780, and 1268 m a.s.l.) in Lushan Mountain, central subtropical China. For the dominant trees, both leaf and leaf-litter δ¹³C decreased as altitude increased from low to high altitude, whereas surface soil δ¹³C increased. The lower leaf δ¹³C at high altitudes was associated with the high moisture-related discrimination, while the high soil δ¹³C is attributed to the low temperature-induced decay. At each altitude, soil δ¹³C became enriched with soil depth. Soil δ¹³C increased with soil C concentrations and altitude, but decreased with soil depth. A negative relationship was also found between O-alkyl C and δ¹³C in litter and soil, whereas a positive relationship was observed between aromatic C and δ¹³C. Lower temperature and higher moisture at high altitudes are the predominant control factors of δ¹³C variation in plants and soils. These results help understand C dynamics in the context of global warming.     

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

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

Spatial patterns of leaf δ13C and δ15N of aquatic macrophytes in the arid zone of northwestern China

Analysis of stable isotope composition is an important tool in research on plant physiological ecology. However, large‐scale patterns of leaf‐stable isotopes for aquatic macrophytes have received considerably less attention. In this study, we examined the spatial pattern of stable isotopes of carbon (δ¹³C) and nitrogen (δ¹⁵N) of macrophytes leaves collected across the arid zone of northwestern China (approximately 2.4 × 10⁶ km²) and attempted to illustrate its relationship with environmental factors (i.e., temperature, precipitation, potential evapotranspiration, sediment total carbon and nitrogen). Our results showed that the mean values of the leaf δ¹³C and δ¹⁵N in the macrophytes sampled from the arid zone were −24.49‰ and 6.82‰, respectively, which were far less depleted than those measured of terrestrial plants. The order of averaged leaf δ¹³C from different life forms was as follows: submerged > floating‐leaved > emergent. Additionally, our studies indicated that the values of foliar δ¹³C values of all the aquatic macrophytes were only negatively associated with precipitation, but the foliar δ¹⁵N values were mainly associated with temperature, precipitation, and potential evapotranspiration. Therefore, we speculated that water‐relation factors are the leaf δ¹³C determinant of macrophytes in the arid zone of northwestern China, and the main factors affecting leaf δ¹⁵N values are the complex combination of water and energy factors.     

Relationships between C3 Plant Foliar Carbon Isotope Composition and Element Contents of Grassland Species at High Altitudes on the Qinghai-Tibet Plateau,China

Relationships of foliar carbon isotope composition (δ¹³C) with foliar C, N, P, K, Ca, Mg contents and their ratios of 219 C₃ species leaf samples, obtained in August in 2004 to 2007 from 82 high altitude grassland sites on the Qinghai-Tibet Plateau China, were examined. This was done with reference to the proposition that foliar δ¹³C increases with altitude and separately for the life-form groups of graminoids, forbs and shrubs and for the genera Stipa and Kobresia. For all samples, foliar δ¹³C was negatively related to foliar K, P and ∑_{K+ Ca+ Mg}, and positively correlated to foliar C, C/N and C/P. The significance of these correlations differed for the taxonomic and life-form groups. Lack of a relationship of foliar δ¹³C with foliar N was inconsistent with the majority of studies that have shown foliar δ¹³C to be positively related to foliar N due to a decrease of C_i/C_a (the ratio between intercellular and atmospheric concentration of CO₂) and explained as a result of greater photosynthetic capacity at higher foliar N concentration. However this inconsistency relates to other high altitude studies that have found that photosynthetic capacity remains constant as foliar N increases. After accounting for the altitudinal relationship with foliar δ¹³C, of the elements only the K effect was significant and was most strongly expressed for Kobresia. It is concluded that factors critical to plant survival and growth at very high altitudes, such as low atmospheric pressure and low temperatures, may preclude expression of relationships between foliar δ¹³C and foliar elements that have been observed at lower altitudes.     

Effects of Air Pollutants on the Composition of Stable Carbon Isotopes, deltaC, of Leaves and Wood, and on Leaf Injury

Air pollutants are known to cause visible leaf injury as well as impairment of photosynthetic CO₂ fixation. Here we evaluate whether the effects on photosynthesis are large enough to cause changes in the relative composition of stable carbon isotopes, δ¹³C, of plant tissue samples, and, if so, how the changes relate to visual leaf injury. For that purpose, several woody and herbaceous plant species were exposed to SO₂ + O₃ and SO₂ + O₃ + NO₂ for one month (8 hours per day, 5 days per week). At the end of the fumigations, the plants were evaluated for visual leaf lesions, and δ¹³C of leaf tissue was determined. Woody plants generally showed less visual leaf injury and smaller effects on δ¹³C of pollutant exposure than did herbaceous plants. If δ¹³C was affected by pollutants, it became, with few exceptions, less negative. The data from the fumigation experiments were consistent with δ¹³C analyses of whole wood of annual growth rings from two conifer tree species, Pseudotsuga menziesii and Pinus strobus. These trees had been exposed until 1977 to exhaust gases from a gas plant at Lacq, France. Wood of both conifer species formed in the polluted air of 1972 to 1976 had less negative δ¹³C values than had wood formed in the much cleaner air in 1982 to 1986. No similar, time-dependent differences in δ¹³C of wood were observed in trees which had been continuously growing in clean air. Our δ¹³C data from both relatively short-term artificial exposures and long-term natural exposure are consistent with greater stomatal limitation of photosynthesis in polluted air than in clean air.     

North Atlantic minke whale (Balaenoptera acutorostrata) feeding habits and migrations evaluated by stable isotope analysis of baleen

Isotopic analyses of the incrementally growing baleen in Mysticeti have been used to learn about their feeding and movement patterns. Using methods previously applied to Pacific minke whales, stable δ¹⁵N and δ¹³C isotope values were measured along the baleen plates of male and female minke whales from two locations in the Northeast Atlantic. The sample sizes used in this study are comparable to those previously used in the literature, and, although limited in size, the evidence suggests differences in isotopic signatures between whales caught at different locations. Both the δ¹⁵N and δ¹³C data suggest whales at the higher latitude site of Svalbard have a narrower diet than the whales from Lofoten/Vesterålen in Norway. Across all whales, the δ¹⁵N data indicate the whales primarily prey on fish for much of the year, only switching to zooplankton during the spring bloom. The δ¹³C data fail to confirm whether the whales migrate over long distances.     

Carbon Isotope Composition of Nighttime Leaf-Respired CO2 in the Agricultural-Pastoral Zone of the Songnen Plain,Northeast China

Variations in the carbon isotope signature of leaf dark-respired CO₂ (δ¹³C_R) within a single night is a widely observed phenomenon. However, it is unclear whether there are plant functional type differences with regard to the amplitude of the nighttime variation in δ¹³C_R. These differences, if present, would be important for interpreting the short-term variations in the stable carbon signature of ecosystem respiration and the partitioning of carbon fluxes. To assess the plant functional type differences relating to the magnitude of the nighttime variation in δ¹³C_R and the respiratory apparent fractionation, we measured the δ¹³C_R, the leaf gas exchange, and the δ¹³C of the respiratory substrates of 22 species present in the agricultural-pastoral zone of the Songnen Plain, northeast China. The species studied were grouped into C₃ and C₄ plants, trees, grasses, and herbs. A significant nocturnal shift in δ¹³C_R was detected in 20 of the studied species, with the magnitude of the shift ranging from 1‰ to 5.8‰. The magnitude of the nighttime variation in δ¹³C_R was strongly correlated with the daytime cumulative carbon assimilation, which suggests that variation in δ¹³C_R were influenced, to some extent, by changes in the contribution of malate decarboxylation to total respiratory CO₂ flux. There were no differences in the magnitude of the nighttime variation in δ¹³C_R between the C₃ and C₄ plants, as well as among the woody plants, herbs and graminoids. Leaf respired CO₂ was enriched in ¹³C compared to biomass, soluble carbohydrates and lipids; however the magnitude of enrichment differed between 8 pm and 4 am, which were mainly caused by the changes in δ¹³C_R. We also detected the plant functional type differences in respiratory apparent fractionation relative to biomass at 4 am, which suggests that caution should be exercised when using the δ¹³C of bulk leaf material as a proxy for the δ¹³C of leaf-respired CO₂.     

deltaC Values in Maize Leaf Correlate with Phosphoenolpyruvate Carboxylase Levels

Values of δ¹³C and levels of phosphoenolpyruvate carboxylase and ribulose 1,5-bisphosphate carboxylase/oxygenase were analyzed in segments from the fourth leaf of young maize (Zea mays L.) plants. The δ¹³C values became significantly more negative from the base to the tip of the leaves. Phosphoenolpyruvate carboxylase levels and ribulose bisphosphate carboxylase levels both increased from the base to the tip. The principal effect of phosphoenolpyruvate carboxylase levels or δ¹³C should arise through its effect on the carboxylation/diffusion balance in the mesophyll. In this case, δ¹³C values should become more negative as phosphoenolpyruvate carboxylase levels increase, unless there are offsetting changes in stomatal aperture. The principal effect of ribulose bisphosphate carboxylase/oxygenase on δ¹³C should occur through its effect on the extent of leakage of CO₂ from the bundle sheath cells. In this case, δ¹³C values should become more positive as ribulose bisphosphate carboxylase levels increase. Accordingly, the variation in δ¹³C values seen in maize leaves appears to be the result of variations in the level of phosphoenolpyruvate carboxylase.     

Seedling Responses to Water Pulses in Shrubs with Contrasting Histories of Grassland Encroachment

Woody plant encroachment into grasslands has occurred worldwide, but it is unclear why some tree and shrub species have been markedly more successful than others. For example, Prosopis velutina has proliferated in many grasslands of the Sonoran Desert in North America over the past century, while other shrub species with similar growth form and life history, such as Acacia greggii, have not. We conducted a glasshouse experiment to assess whether differences in early seedling development could help explain why one species and not the other came to dominate many Sonoran Desert grasslands. We established eight watering treatments mimicking a range of natural precipitation patterns and harvested seedlings 16 or 17 days after germination. A. greggii had nearly 7 times more seed mass than P. velutina, but P. velutina emerged earlier (by 3.0±0.3 d) and grew faster (by 8.7±0.5 mg d⁻¹). Shoot mass at harvest was higher in A. greggii (99±6 mg seedling⁻¹) than in P. velutina (74±2 mg seedling⁻¹), but there was no significant difference in root mass (54±3 and 49±2 mg seedling⁻¹, respectively). Taproot elongation was differentially sensitive to water supply: under the highest initial watering pulse, taproots were 52±19 mm longer in P. velutina than in A. greggii. Enhanced taproot elongation under favorable rainfall conditions could give nascent P. velutina seedlings growth and survivorship advantages by helping reduce competition with grasses and maintain contact with soil water during drought. Conversely, A. greggii''s greater investment in mass per seed appeared to provide little return in early seedling growth. We suggest that such differences in recruitment traits and their sensitivities to environmental conditions may help explain ecological differences between species that are highly similar as adults and help identify pivotal drivers of shrub encroachment into grasslands.     

The sensitivity of ecosystem carbon exchange to seasonal precipitation and woody plant encroachment

Ongoing, widespread increases in woody plant abundance in historical grasslands and savannas (woody encroachment) likely will interact with future precipitation variability to influence seasonal patterns of carbon cycling in water-limited regions. To characterize the effects of woody encroachment on the sensitivity of ecosystem carbon exchange to seasonal rainfall in a semi-arid riparian setting we used flux-duration analysis to compare 2003-growing season NEE data from a riparian grassland and shrubland. Though less seasonally variable than the grassland, shrubland NEE was more responsive to monsoon rains than anticipated. During the 2004-growing season we measured leaf gas exchange and collected leaf tissue for δ¹³C and nitrogen content analysis periodically among three size classes of the dominant woody-plant, Prosopis velutina and the dominant understory species, Sporobolus wrightii, a C₄ bunchgrass, present at the shrubland. We observed size-class and plant functional type independent patterns of seasonal plant performance consistent with greater-than-anticipated sensitivity of NEE in the shrubland. This research highlights the complex interaction between growing-season precipitation, plant-available alluvial groundwater and woody plant abundance governing ecosystem carbon balance in this semi-arid watershed.     

Ungulate stimulation of nitrogen cycling and retention in Yellowstone Park grasslands

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

Fractionation of the stable isotopes of inorganic carbon by seagrasses

The δ¹³C values for seagrasses collected along the Texas Gulf Coast range from −10.9 to −11.4‰. These values are similar to the δ¹³C values of terrestrial C₄ plants, but seagrasses lack bundle sheath cells which are important in determining the δ¹³C values of C₄ plants. This work attempts to explain the reason the δ¹³C values of seagrasses resemble the δ¹³C values of C₄ plants.     

Plant Trait Variation along an Altitudinal Gradient in Mediterranean High Mountain Grasslands: Controlling the Species Turnover Effect

Assessing changes in plant functional traits along gradients is useful for understanding the assembly of communities and their response to global and local environmental drivers. However, these changes may reflect the effects of species composition (i.e. composition turnover), species abundance (i.e. species interaction), and intra-specific trait variability (i.e. species plasticity). In order to determine the relevance of the latter, trait variation can be assessed under minimal effects of composition turnover. Nine sampling sites were established along an altitudinal gradient in a Mediterranean high mountain grassland community with low composition turnover (Madrid, Spain; 1940 m–2419 m). Nine functional traits were also measured for ten individuals of around ten plant species at each site, for a total of eleven species across all sites. The relative importance of different sources of variability (within/between site and intra-/inter-specific functional diversity) and trait variation at species and community level along the considered gradients were explored. We found a weak individual species response to altitude and other environmental variables although in some cases, individuals were smaller and leaves were thicker at higher elevations. This lack of species response was most likely due to greater within- than between-site species variation. At the community level, inter-specific functional diversity was generally greater than the intra-specific component except for traits linked to leaf element content (leaf carbon content, leaf nitrogen content, δ¹³C and δ¹⁵N). Inter-specific functional diversity decreased with lower altitude for four leaf traits (specific leaf area, leaf dry matter content, δ¹³C and δ¹⁵N), suggesting trait convergence between species at lower elevations, where water shortage may have a stronger environmental filtering effect than colder temperatures at higher altitudes. Our results suggest that, within a vegetation type encompassing various environmental gradients, both, changes in species abundance and intra-specific trait variability adjust for the community functional response to environmental changes.     

Carabid beetles as indicators for shrub encroachment in dry grasslands

Semi-natural grasslands are key habitats for biodiversity conservation in Central Europe. Shrub encroachment is one of the most threatening drivers of grassland degradation and affects soil properties, microclimate, and vegetation with possible impacts on higher trophic levels. We aimed to analyse the impact of shrub encroachment with broom (Cytisus scoparius) on carabid beetle diversity, species composition, and functional traits. In a field study on dry grasslands on the island of Hiddensee (Germany) we studied 15 sites along a gradient of increasing broom encroachment and classified them into three dry grassland types with low, medium, and high shrub cover. Our results provide evidence that shrub encroachment initially has positive effects on species richness and activity densities of dry grassland carabids. Carabid species composition differed among differently shrub-covered dry grassland types, and sites with low and high shrub cover were each characterised by unique carabid assemblages. The species composition of sites with a medium shrub biomass had a transitional character and contained species which are typical for open dry grassland, but also shared species with sites with a high shrub cover. Among functional trait parameters investigated, especially the body size of carabid beetles was related to environmental parameters associated with shrub encroachment. Body size was positively correlated to shrub biomass and soil humidity, but negatively to temperature. Eurytopy values of carabids were related to high litter cover, i.e. habitat generalist (eurytopic) species mainly occurred in densely shrub-encroached sites. In order to preserve unique carabid assemblages of open dry grasslands with stenotopic and smaller species, it is most important to prevent a shrub encroachment higher than about 60% cover. For management we suggest extensive grazing (by cattle, sheep or horses) to prevent shrub encroachment on dry grasslands. In areas with high shrub cover additionally the use of goats or mechanical removal of shrubs might be necessary.