Contrasting nitrogen and phosphorus dynamics across an elevational gradient for subarctic tundra heath and meadow vegetation

Aims

This study explores soil nutrient cycling processes and microbial properties for two contrasting vegetation types along an elevational gradient in subarctic tundra to improve our understanding of how temperature influences nutrient availability in an ecosystem predicted to be sensitive to global warming.

Methods

We measured total amino acid (Amino-N), mineral nitrogen (N) and phosphorus (P) concentrations, in situ net N and P mineralization, net Amino-N consumption, and microbial biomass C, N and P in both heath and meadow soils across an elevational gradient near Abisko, Sweden.

Results

For the meadow, NH₄ ⁺ concentrations and net N mineralization were highest at high elevations and microbial properties showed variable responses; these variables were largely unresponsive to elevation for the heath. Amino-N concentrations sometimes showed a tendency to increase with elevation and net Amino-N consumption was often unresponsive to elevation. Overall, PO₄-P concentrations decreased with elevation and net P immobilization mostly occurred at lower elevations; these effects were strongest for the heath.

Conclusions

Our results reveal that elevation-associated changes in temperature can have contrasting effects on the cycling of N and P in subarctic soils, and that the strength and direction of these effects depend strongly on dominant vegetation type.     

Linkages between N turnover and plant community structure in a tundra landscape

The spatial distribution of organic soil nitrogen (N) in alpine tundra was studied along a natural environmental gradient, covering five plant communities, at the Latnjajaure Field Station, northern Swedish Lapland. The five communities (mesic meadow, meadow snowbed, dry heath, mesic heath, and heath snowbed) are the dominant types in this region and are differentiated by soil pH. Net N mineralization, net ammonification, and net nitrification were measured using 40-day laboratory incubations based on extractable NH₄⁺ and NO₃⁻. Nitrification enzyme activity (NEA), denitrification enzyme activity (DEA), amino acid concentrations, and microbial respiration were measured for soils from each plant community. The results show that net N mineralization rates were more than three times higher in the meadow ecosystems (mesic meadow 0.7 μg N g⁻¹ OM day⁻¹ and meadow snowbed 0.6 μg N g⁻¹ OM day⁻¹) than the heath ecosystems (dry heath 0.2 μg N g⁻¹ OM day⁻¹, mesic heath 0.1 μg N g⁻¹ OM day⁻¹ and heath snowbed 0.2 μg N g⁻¹ OM day⁻¹). The net N mineralization rates were negatively correlated to organic soil C/N ratio (r = −0.652, P < 0.001) and positively correlated to soil pH (r = 0.701, P < 0.001). Net nitrification, inorganic N concentrations, and NEA rates also differed between plant communities; the values for the mesic meadow were at least four times higher than the other plant communities, and the snowbeds formed an intermediate group. Moreover, the results show a different pattern of distribution for individual amino acids across the plant communities, with snowbeds tending to have the highest amino acid N concentrations. The differences between plant communities along this natural gradient also illustrate variations between the dominant mycorrhizal associations in facilitating N capture by the characteristic functional groups of plants. Responsible Editor: Bernard Nicolardot     

Interactive effects of vegetation type and elevation on aboveground and belowground properties in a subarctic tundra

An improved knowledge of how contrasting types of plant communities and their associated soil biota differ in their responses to climatic variables is important for better understanding the future impacts of climate change on terrestrial ecosystems. Elevational gradients serve as powerful study systems for answering questions on how ecological processes can be affected by changes in temperature and associated climatic variables. In this study, we evaluated how plant and soil microbial communities, and abiotic soil properties, change with increasing elevation in subarctic tundra in northern Sweden, for each of two dominant but highly contrasting vegetation types, namely heath (dominated by woody dwarf shrubs) and meadow (dominated by herbaceous species). To achieve this, we measured plant community characteristics, microbial community properties and several soil abiotic properties for both vegetation types across an elevation gradient of 500 to 1000 m. We found that the two vegetation types differed not only in several above‐ and belowground properties, but also in how these properties responded to elevation, pointing to important interactive effects between vegetation type and elevation. Specifically, for the heath, available soil nitrogen and phosphorus decreased with elevation whereas fungal dominance increased, while for the meadow, idiosyncratic responses to elevation for these variables were found. These differences in belowground responses to elevation among vegetation types were linked to shifts in the species and functional group composition of the vegetation. Our results highlight that these two dominant vegetation types in subarctic tundra differ greatly not only in fundamental aboveground and belowground properties, but also in how these properties respond to elevation and are therefore likely to be influenced by temperature. As such they highlight that vegetation type, and the soil abiotic properties that determine this, may serve as powerful determinants of how both aboveground and belowground properties respond to strong environmental gradients.     

Carbon and Nitrogen Cycling in Soils from Acidic and Nonacidic Tundra with Different Glacial Histories in Northern Alaska

Moist acidic and nonacidic tundra are two of the most common vegetation types of the tundra in the northern foothills of the Brooks Range, Alaska, and they differ considerably in vegetation, soil nutrient availability, and soil pH. Both occur on mesic, gentle slopes, but acidic tundra is more common on older glacial surfaces whereas nonacidic tundra is more common on younger surfaces. Although much prior research has focused on moist acidic tundra, nonacidic tundra is still relatively unstudied. We compared rates of soil carbon (C) and nitrogen (N) cycling and their response to warming and changes in moisture in moist acidic tundra on Itkillik I glacial drift (50,000–120,000 years old, pH = 3–4) and moist nonacidic tundra on Itkillik II glacial drift (11,500–60,000 year old, pH = 6–7). We hypothesized that rates of soil C and N cycling would be faster at the nonacidic site because it has a more favorable pH for microbial activity and higher-quality organic matter inputs arising from its greater herbaceous plant production relative to the acidic site. However, in contrast to our hypothesis, in situ soil respiration, as well as respiration, dissolved organic C production, and net N mineralization in laboratory incubations, was greater for soils from the acidic site. Nevertheless, the sites responded similarly to manipulations of temperature and moisture, exhibiting exponential increases in respiration with warming between 4°C and 15°C but surprisingly little sensitivity to changes in moisture between 300% and 700%. Slower soil organic matter decomposition at the nonacidic site likely results from the stabilization of soil organic matter by high concentrations of calcium. Received 27 August 2001; accepted 3 April 2002.     

Coordinated responses of soil communities to elevation in three subarctic vegetation types

Global warming has begun to have a major impact on the species composition and functioning of plant and soil communities. However, long‐term community and ecosystem responses to increased temperature are still poorly understood. In this study, we used a well‐established elevational gradient in northern Sweden to elucidate how plant, microbial and nematode communities shift with elevation and associated changes in temperature in three highly contrasting vegetation types (i.e. heath, meadow and Salix vegetation). We found that responses of both the abundance and composition of microbial and nematode communities to elevation differed greatly among the vegetation types. Within vegetation types, changes with elevation of plant, microbial and nematode communities were mostly linked at fine levels of taxonomic resolution, but this pattern disappeared when coarser functional group levels were considered. Further, nematode communities shifted towards more conservative nutrient cycling strategies with increasing elevation in heath and meadow vegetation. Conversely, in Salix vegetation microbial communities with conservative strategies were most pronounced at the mid‐elevation. These results provide limited support for increasing conservative nutrient cycling strategies at higher elevation (i.e. with a harsher climate). Our findings indicate that climate‐induced changes in plant community composition may greatly modify or counteract the impact of climate change on soil communities. Therefore, to better understand and predict ecosystem responses to climate change, it will be crucial to consider vegetation type and its specific interactions with soil communities.     

Responses of moist non-acidic arctic tundra to altered environment: productivity, biomass, and species richness

In arctic Alaska, researchers have manipulated air temperature, light availability, and soil nutrient availability in several tundra communities over the past two decades. These communities responded quite differently to the same manipulations, and species responded individualistically within communities and among sites. For example, moist acidic tundra is primarily nitrogen (N)‐limited, whereas wet sedge tundra is primarily phosphorus (P)‐limited, and the magnitude of growth responses varies across sites within communities. Here we report results of four years of manipulated nutrients (N and/or P) and/or air temperature in an understudied, diverse plant community, moist non‐acidic tussock tundra, in northern Alaska. Our goals were to determine which factors limit above‐ground net primary productivity (ANPP) and biomass, how community composition changes may affect ecosystem attributes, and to compare these results with those from other communities to determine their generality. Although relative abundance of functional groups shifted in several treatments, the only significant change in community‐level ANPP and biomass occurred in plots that received both N and P, driven by an increase in graminoid biomass and production resulting from a positive effect of adding N. There was no difference in community biomass among any other treatments; however, some growth forms and individual species did respond. After four years no one species has come to dominate the treatment plots and species richness has not changed. These results are similar to studies in dry heath, wet sedge, and moist acidic tundra where community biomass had the greatest response to both N and P and warming results were more subtle. Unlike in moist acidic tundra where shrub biomass increased markedly with fertilization, our results suggest that in non‐acidic tundra carbon sequestration in plant biomass will not increase substantially under increased soil nutrient conditions because of the lack of overstory shrub species.     

Plant community response to nitrogen and phosphorus enrichment varies across an alpine tundra moisture gradient

Background: The extent to which nutrient availability influences plant community composition and dynamics has been a focus of ecological enquiry for decades.

Aims: Results from a long-term nitrogen (N) and phosphorus (P) addition experiment in alpine tundra were used to evaluate the importance of the two nutrients in structuring plant communities in three communities that differed in their snow cover amounts and duration and soil moisture characteristics.

Methods: A factorial N and P experiment was established in three meadows differing in initial vegetation composition and soil moisture. Plant and soil characteristics were measured after 20 years, and the dissimilarity among meadows and treatments were measured using permutational analysis of variance.

Results: Plant species richness declined uniformly across the three meadow types and in response to N and N + P additions, while both evenness and the Shannon diversity index finding indicated that nutrient additions had the highest impact on moister habitats. Overall, N impacts overshadowed changes attributed to P additions, and the N and N + P plots in wet meadow sites were the least diverse and scored the lowest dissimilarity averages among treatments. Dissimilarity estimates indicated that the control and P plots in the dry meadow community were more distinct in composition than all other plots, and especially those in the moist or wet meadows. Above-ground biomass of grasses and sedges (graminoids) increased with N additions while forbs appeared to show responses dictated in part by the graminoid responses. The most abundant grass species of moist and wet meadow, Deschampsia cespitosa, dominated N and N + P plots of the wet sites, but did not show a N response in moist areas in spite of its general abundance in moist meadow. Competition from other plant species in the moist areas likely diminished the D. cespitosa response and contributed to the resilience of the community to nutrient enrichment.

Conclusions: Initial community composition, as influenced by the specific moisture regime, appears to control the extent to which changes in nutrient resources can alter plant community structure. Long-term fertilization tends to support most but not all findings obtained from shorter-termed efforts, and wet meadows exhibit the largest changes in plant species numbers and composition when chronically enriched with N.     

Interactive Effects Between Reindeer and Habitat Fertility Drive Soil Nutrient Availabilities in Arctic Tundra

Herbivores impact nutrient availability and cycling, and the net effect of herbivory on soil nutrients is generally assumed to be positive in nutrient-rich environments and negative in nutrient-poor ones. This is, however, far from a uniform pattern, and there is a recognized need to investigate any interactive effects of herbivory and habitat fertility (i.e., plant C/N ratios) on soil nutrient availabilities. We determined long-term effects of reindeer on soil extractable nitrogen (N) and phosphorus (P) and their net mineralization rates along a fertility gradient of plant carbon (C) to N and P ratios in arctic tundra. Our results showed that reindeer had a positive effect on soil N in the more nutrient-poor sites and a negative effect on soil P in the more nutrient-rich sites, which contrasts from the general consensus. The increase in N availability was linked to a decrease in plant and litter C/N ratios, suggesting that a shift in vegetation composition toward more graminoids favors higher N cycling. Soil P availability was not as closely linked to the vegetation and is likely regulated more by herbivore-induced changes in soil physical and chemical properties. The changes in soil extractable N and P resulted in higher soil N/P ratios, suggesting that reindeer could drive the vegetation toward P-limitation. This research highlights the importance of including both the elements N and P and conducting studies along environmental gradients in order to better understand the interactive effects of herbivory and habitat fertility on nutrient cycling and primary production.     

Soil fauna communities and microbial respiration in high Arctic tundra soils at Zackenberg, Northeast Greenland

The soil fauna communities were described for three dominant vegetation types in a high arctic site at Zackenberg, Northeast Greenland. Soil samples were extracted to quantify the densities of mites, collembolans, enchytraeids, diptera larvae, nematodes and protozoa. Rates of microbial respiration were also assessed. Collembolans were found in highest densities in dry heath soil, about 130,000 individuals m⁻², more than twice as high as in mesic heath soils. Enchytraeids, diptera larvae and nematodes were also more abundant in the dry heath soil than in mesic heath soils, whereas protozoan densities (naked amoeba and heterotrophic flagellates) were equal. Respiration rate of unamended soil was similar in soil from the three plots. However, a higher respiration rate increase in carbon + nutrient amended soil and the higher densities of soil fauna (with the exception of mites and protozoa) in dry heath compared to the mesic heath soils indicated a higher decomposition rate here.     

Controls of Bedrock Geochemistry on Soil and Plant Nutrients in Southeastern Utah

The cold deserts of the Colorado Plateau contain numerous geologically and geochemically distinct sedimentary bedrock types. In the area near Canyonlands National Park in Southeastern Utah, geochemical variation in geologic substrates is related to the depositional environment with higher concentrations of Fe, Al, P, K, and Mg in sediments deposited in alluvial or marine environments and lower concentrations in bedrock derived from eolian sand dunes. Availability of soil nutrients to vegetation is also controlled by the formation of secondary minerals, particularly for P and Ca availability, which, in some geologic settings, appears closely related to variation of CaCO₃ and Ca-phosphates in soils. However, the results of this study also indicate that P content is related to bedrock and soil Fe and Al content suggesting that the deposition history of the bedrock and the presence of P-bearing Fe and Al minerals, is important to contemporary P cycling in this region. The relation between bedrock type and exchangeable Mg and K is less clear-cut, despite large variation in bedrock concentrations of these elements. We examined soil nutrient concentrations and foliar nutrient concentration of grasses, shrubs, conifers, and forbs in four geochemically distinct field sites. All four of the functional plant groups had similar proportional responses to variation in soil nutrient availability despite large absolute differences in foliar nutrient concentrations and stoichiometry across species. Foliar P concentration (normalized to N) in particular showed relatively small variation across different geochemical settings despite large variation in soil P availability in these study sites. The limited foliar variation in bedrock-derived nutrients suggests that the dominant plant species in this dryland setting have a remarkably strong capacity to maintain foliar chemistry ratios despite large underlying differences in soil nutrient availability.     

Links between plant community composition,soil organic matter quality and microbial communities in contrasting tundra habitats

Plant communities, soil organic matter and microbial communities are predicted to be interlinked and to exhibit concordant patterns along major environmental gradients. We investigated the relationships between plant functional type composition, soil organic matter quality and decomposer community composition, and how these are related to major environmental variation in non-acid and acid soils derived from calcareous versus siliceous bedrocks, respectively. We analysed vegetation, organic matter and microbial community compositions from five non-acidic and five acidic heath sites in alpine tundra in northern Europe. Sequential organic matter fractionation was used to characterize organic matter quality and phospholipid fatty acid analysis to detect major variation in decomposer communities. Non-acidic and acidic heaths differed substantially in vegetation composition, and these disparities were associated with congruent shifts in soil organic matter and microbial communities. A high proportion of forbs in the vegetation was positively associated with low C:N and high soluble N:phenolics ratios in soil organic matter, and a high proportion of bacteria in the microbial community. On the contrary, dwarf shrub-rich vegetation was associated with high C:N and low soluble N:phenolics ratios, and a high proportion of fungi in the microbial community. Our study demonstrates a strong link between the plant community composition, soil organic matter quality, and microbial community composition, and that differences in one compartment are paralleled by changes in others. Variation in the forb-shrub gradient of vegetation may largely dictate variations in the chemical quality of organic matter and decomposer communities in tundra ecosystems. Soil pH, through its direct and indirect effects on plant and microbial communities, seems to function as an ultimate environmental driver that gives rise to and amplifies the interactions between above- and belowground systems. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Within- and across-species responses of plant traits and litter decomposition to elevation across contrasting vegetation types in subarctic tundra

Elevational gradients are increasingly recognized as a valuable tool for understanding how community and ecosystem properties respond to climatic factors, but little is known about how plant traits and their effects on ecosystem processes respond to elevation. We studied the response of plant leaf and litter traits, and litter decomposability across a gradient of elevation, and thus temperature, in subarctic tundra in northern Sweden for each of two contrasting vegetation types, heath and meadow, dominated by dwarf shrubs and herbaceous plants respectively. This was done at each of three levels; across species, within individual species, and the plant community using a community weighted average approach. Several leaf and litter traits shifted with increasing elevation in a manner consistent with greater conservation of nutrients at all three levels, and the most consistent response was an increase in tissue N to P ratio. However, litter decomposition was less directly responsive to elevation because the leaf and litter traits which were most responsive to elevation were not necessarily those responsible for driving decomposition. At the community level, the response to elevation of foliar and litter traits, and decomposability, varied greatly among the two vegetation types, highlighting the importance of vegetation type in determining ecological responses to climatic factors such as temperature. Finally our results highlight how understanding the responses of leaf and litter characteristics of functionally distinct vegetation types, and the processes that they drive, to temperature helps provide insights about how future climate change could affect tundra ecosystems.     

Plant community responses to 5 years of simulated climate change in meadow and heath ecosystems at a subarctic-alpine site

Climate change was simulated by increasing temperature and nutrient availability in an alpine landscape. We conducted a field experiment of BACI-design (before/after control/impact) running for five seasons in two alpine communities (heath and meadow) with the factors temperature (increase of ca. 1.5–3.0°C) and nutrients (5 g N, 5 g P per m²) in a fully factorial design in northern Swedish Lapland. The response variables were abundances of plant species and functional types. Plant community responses to the experimental perturbations were investigated, and the responses of plant functional types were examined in comparison to responses at the species level. Nutrient addition, exclusively and in combination with enhanced temperature increase, exerted the most pronounced responses at the species-specific and community levels. The main responses to nutrient addition were increases in graminoids and forbs, whereas deciduous shrubs, evergreen shrubs, bryophytes, and lichens decreased. The two plant communities of heath or meadow showed different vegetation responses to the environmental treatments despite the fact that both communities were located on the same subarctic-alpine site. Furthermore, we showed that the abundance of forbs increased in response to the combined treatment of temperature and nutrient addition in the meadow plant community. Within a single-plant functional type, most species responded similarly to the enhanced treatments although there were exceptions, particularly in the moss and lichen functional types. Plant community structure showed BACI responses in that vegetation dominance relationships in the existing plant functional types changed to varying degrees in all plots, including control plots. Betula nana and lichens increased in the temperature-increased enhancements and in control plots in the heath plant community during the treatment period. The increases in control plots were probably a response to the observed warming during the treatment period in the region. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Effects of heterogeneous habitat use by cattle on nutrient availability and litter decomposition in soils of an Alpine pasture

Grazing by free-ranging cattle on Alpine pastures in southern Switzerland creates sharp contrasts in plant species composition between small camp areas, which are grazed intensely and receive most cattle excreta, and surrounding pasture dominated by Nardus stricta, which is only lightly grazed. We hypothesised that these contrasts are maintained by positive feedbacks related to nutrient availability in soil, in that (a) plant material with rapid decomposition and nutrient release decomposes in camp areas and (b) litter decomposition is further stimulated by enhanced nutrient availability in soil. We compared nutrient availability at three camp areas with that in surrounding Nardus vegetation and investigated how the decomposition of plant material from both vegetation types responds to nutrient availability in soil, both in the field (during 14 weeks) and in the laboratory (during 4, 10, and 16 weeks). At all three field sites P availability was significantly enhanced in camp areas, whereas differences in N availability were inconsistent among the three sites. Laboratory incubations indicated that microbial activity after the addition of labile C (cellulose) was limited by P availability in the Nardus vegetation but not in camp areas. The camp-area plant substrate decomposed much faster (81.5% vs. 27.1% ash-free dry mass loss in the field) and released more N and P than the Nardus substrate, which tended to immobilise soil nutrients. However, the decomposition rate of neither substrate was influenced by nutrient availability in soil, both in the field (comparing camp areas and Nardus vegetation) and in the laboratory (comparing incubations with and without N or P fertilisation). We conclude that the contrasting quality of plant substrates contributes to the greater nutrient availability in camp areas (feedback a) but that the latter does not influence the decomposition of in situ plant material (feedback b) because the latter is not nutrient-limited.     

Nutrient fluxes via litterfall and leaf litter decomposition vary across a gradient of soil nutrient supply in a lowland tropical rain forest

The extent to which plant communities are determined by resource availability is a central theme in ecosystem science, but patterns of small-scale variation in resource availability are poorly known. Studies of carbon (C) and nutrient cycling provide insights into factors limiting tree growth and forest productivity. To investigate rates of tropical forest litter production and decomposition in relation to nutrient availability and topography in the absence of confounding large-scale variation in climate and altitude we quantified nutrient fluxes via litterfall and leaf litter decomposition within three distinct floristic associations of tropical rain forest growing along a soil fertility gradient at the Sepilok Forest Reserve (SFR), Sabah, Malaysia. The quantity and nutrient content of small litter decreased along a gradient of soil nutrient availability from alluvial forest (most fertile) through sandstone forest to heath forest (least fertile). Temporal variation in litterfall was greatest in the sandstone forest, where the amount of litter was correlated negatively with rainfall in the previous month. Mass loss and N and P release were fastest from alluvial forest litter, and slowest from heath forest litter. All litter types decomposed most rapidly in the alluvial forest. Stand-level N and P use efficiencies (ratios of litter dry mass to nutrient content) were greatest for the heath forest followed by the sandstone ridge, sandstone valley and alluvial forests, respectively. We conclude that nutrient supply limits productivity most in the heath forest and least in the alluvial forest. Nutrient supply limited productivity in sandstone forest, especially on ridge and hill top sites where nutrient limitation may be exacerbated by reduced rates of litter decomposition during dry periods. The fluxes of N and P varied significantly between the different floristic communities at SFR and these differences may contribute to small-scale variation in species composition.     

Plant responsiveness to variation in precipitation and nitrogen is consistent across the compositional diversity of a California annual grassland

Question: How does responsiveness to water and Nitrogen (N) availability vary across the compositional and functional diversity that exists in a mesic California annual grassland plant community? Location: Northern California annual grassland. Methods: A mesocosm system was used to simulate average annual precipitation totals and dry and wet year extremes observed in northern California mesic grasslands. The effects of precipitation and N availability on biomass and fecundity were measured on three different vegetation types, a mixed grass forb community, and a forb and a grass monoculture. The treatment effects on plant community composition were examined in the mixed species community. Results: While growth and seed production of the three vegetation types was inherently different, their responses to variation in precipitation and N were statistically similar. Plant density, shoot biomass, and seed production tended to increase with greater water availability in all vegetation types, with the exception of a consistent growth reduction in high precipitation (1245 mm) plots in the first year of the study. Shoot biomass responded positively to N addition, an effect that increased with greater water availability. Nitrogen addition had little effect on plant density or seed production. In the mixed grass‐forb community, biomass responsiveness to water and N treatments were consistently driven by the shoot growth of Avena barbata, the dominant grass species. Conclusions: Vegetation responses to changes in precipitation and N availability were consistent across a range of composition and structural diversity in this study. Plant growth and seed production were sensitive to both increased and decreased precipitation totals, and the magnitude of these responses to N availability varied depending on soil moisture conditions. Our results suggest the impacts of changing precipitation regimes and N deposition on annual productivity of California grasslands may be predictable under different climate scenarios across a range of plant communities.     

In situ mineralization of nitorgen and phosphorus of arctic soils after perturbations simulating climate change

Seasonal net nitrogen (N) and phosphorus (P) mineralization was investigated at Abisko, Swedish Lapland in soils of a subarctic heath and in soils of a colder (by about 4° C), high altitude fellfield by (a) using in situ soil incubation in soils which had been shaded or subjected to two levels of increased temperature, combined with (b) reciprocal transplantation of soils between the two sites. Proportionally large and significant net seasonal mineralization of N, in contrast to non-significant P mineralization, was found in untransplanted and transplanted fellfield soil. In contrast, P was mineralized in proportionally large amounts, in contrast to low N mineralization, in the transplanted and untransplanted heath soil. The differences indicate that P was strongly immobilized in relation to N at the fellfield and that N was more strongly immobilized than P in the heath soil. The immobilization in both soils remained high even after a temperature change of 4–5° C experienced by transplanted soils. Air temperature increases of up to 4–5° C in greenhouses resulted in a soil temperature increase of 1–2° C and did not cause any extra increase of net N and P mineralization. The results suggest that soil temperature increases of up to 2° C, which are likely to occur by the end of the next century as an effect of a predicted 4–5° C rise in air temperature, have only small effects on net mineralization in at least two characteristic tundra soils. These effects are probably smaller than the natural fluctuation of plant available nutrients from site to site, even within the same plant community. A further soil temperature increase of up to 4–5° C may enhance decomposition and gross mineralization, but the rate of net mineralization, and hence the change of nutrient availability to the plants, depends on the extent of microbial immobilization of the extra nutrients released.     

The consequences of multiple resource shifts on the productivity and composition of alpine tundra communities: inferences from a long-term snow and nutrient manipulation experiment

Background: Gradients in the amounts and duration of snowpack and resulting soil moisture gradients have been associated with different plant communities across alpine landscapes.

Aims: The extent to which snow additions could alter plant community structure, both alone and in combination with nitrogen (N) and phosphorus (P) additions, provided an empirical assessment of the strength of these variables on structuring the plant communities of the alpine tundra at Niwot Ridge, Colorado Front Range.

Methods: A long-term snow fence was used to study vegetation changes in responses to snowpack, both alone and in conjunction with nutrient amendments, in plots established in dry and moist meadow communities in the alpine belt. Species richness, diversity, evenness and dissimilarity were evaluated after 20 years of treatments.

Results: Snow additions, alone, reduced species richness and altered species composition in dry meadow plots, but not in moist meadow; more plant species were found in the snow-impacted areas than in nearby controls. Changes in plant community structure to N and N + P additions were influenced by snow additions. Above-ground plant productivity in plots not naturally affected by snow accumulation was not increased, and the positive responses of plant species to nutrient additions were reduced by snow addition. Plant species showed individualistic responses to changes in snow and nutrients, and indirect evidence suggested that competitive interactions mediated responses. A Permanova analysis demonstrated that community dissimilarity was affected by snow, N, and P additions, but with these responses differing by community type for snow and N. Snow influenced community patterns generated by N, and finally, the communities impacted by N + P were significantly different than those affected by the individual nutrients.

Conclusions: These results show that changes in snow cover over a 20-year interval produce measureable changes in community composition that concurrently influence and are influenced by soil nutrient availability. Dry meadow communities exhibit more sensitivity to increases in snow cover whereas moist meadow communities appear more sensitive to N enrichment. This study shows that the dynamics of multiple limiting resources influence both the productivity and composition of alpine plant communities, with, species, life form, and functional traits mediating these responses.     

黄土丘陵区植被与地形特征对土壤和土壤微生物生物量生态化学计量特征的影响

研究黄土丘陵区植被与地形特征对土壤和土壤微生物生物量生态化学计量特征影响有助于深入理解黄土丘陵区不同植被带下土壤和土壤微生物相互作用及养分循环规律.选择黄土丘陵区延河流域3个植被区(森林区、森林草原区、草原区)和5种地形部位(阴/阳沟坡、阴/阳梁峁坡、峁顶)的土壤作为研究对象,利用生态化学计量学理论研究植被和地形对土壤和土壤微生物生物量生态化学计量特征的影响.结果表明: 土壤及土壤微生物生物量碳、氮、磷含量在不同地形之间的差别主要表现在沟坡位置和阴坡高于其他坡位和阳坡.植被类型的变化对两个土层(0～10、10～20 cm)土壤和土壤微生物生物量碳、氮、磷的影响均达到显著水平,坡向对表层(0～10 cm)土壤和土壤微生物生物量碳、氮、磷的影响强于坡位,而在10～20 cm土层,坡位对土壤和土壤微生物生物量碳、氮、磷影响更显著.植被类型显著影响土壤C∶N、C∶P、N∶P和土壤微生物生物量C∶N、C∶P,坡向和坡位仅影响土壤C∶P和N∶P,植被类型的变化是影响土壤C∶N的主要因素.同时,植被类型对土壤养分和微生物生物量碳、氮、磷含量及其生态化学计量特征的影响大于地形因子.标准化主轴分析结果表明,黄土丘陵区不同植被带土壤微生物具有内稳性,特别在草原带,土壤微生物生物量生态化学计量学特征具有更加严格的约束比例.在黄土丘陵区,土壤微生物生物量N∶P或许可以作为判断养分限制的另一个有力工具,若将土壤微生物生物量N∶P与植物叶片N∶P配合使用可能有助于我们更加精确地判断黄土丘陵区的土壤养分限制情况.     

Factors determining plant species richness in Alaskan arctic tundra

Abstract. We studied the relationship between plant N:P ratio, soil characteristics and species richness in wet sedge and tussock tundra in northern Alaska at seven sites. We also collected data on soil characteristics, above‐ground biomass, species richness and composition. The N:P ratio of the vegetation did not show any relationship with species richness. The N:P ratio of the soil was related with species richness for both vegetation types. Species richness in the tussock tundra was most strongly correlated with soil calcium content and soil pH, with a strong correlation between these two factors. N:P ratio of the soil was also correlated with soil pH. Other factors correlated with species richness were soil moisture and Sphagnum cover. Organic matter content was the factor most strongly correlated with species richness in the wet sedge vegetation. N:P ratio of the soil was strongly correlated with organic matter content. We conclude that N:P ratio in the vegetation is not an important factor determining species richness in arctic tundra and that species richness in arctic tundra is mainly determined by pH and flooding. In tussock tundra the pH, declining with soil age, in combination with Sphagnum growth strongly decreases species richness, while in wet sedge communities flooding over long periods of time creates less favourable conditions for species richness.