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Background and aims: Current knowledge of soil heterogeneity-diversity relationships (HDR) is largely based on studies manipulating single factor, but the advancements in HDR may require a comprehensive experiment incorporating multiple factors. Methods: We conducted a three-year field experiment in which a seed mixture of 16 common grassland species was sown in plots with heterogeneous soils consisting of small (10 cm × 10 cm) or large patches (30 cm × 30 cm) of low and high nutrients or low and high pH, and homogeneous soils with an even mixture of low and high nutrient/pH soils. Soil nutrients and pH were manipulated in separate treatments. We determined plant species richness and diversity at two focal scales (40 cm × 40 cm plot-scale and 10 cm × 10 cm patch-scale). Results: Plot-scale richness and diversity were not influenced by soil heterogeneity, but patch-scale richness was lower in plots with heterogeneous nutrients than in plots where nutrients were distributed homogeneously. There was no difference between the two heterogeneous nutrient soils with different grain sizes. Patch-scale diversity was higher in heterogeneous pH soils of large patch size than in heterogeneous pH soils of small patch size or the homogeneous pH soil at the final harvest. Species richness and diversity quantified at both plot and patch scales declined in all soils over time. Conclusions: The influence of soil heterogeneity on plant species diversity depends on whether the soil varies in nutrients or pH, and on the temporal-spatial scale at which species diversity and soil heterogeneity are measured. These results indicate that soil heterogeneity has the potential to promote plant coexistence and future HDR studies should consider multiple soil factors at various temporal-spatial scales. [ABSTRACT FROM AUTHOR]

Backgrounds and aims: Negative plant-soil feedbacks (PSFs) are thought to promote species coexistence, but most evidence is derived from theoretical models and data from plant monoculture experiments.Methods: We grew Anthoxanthum odoratum and Centaurea jacea in field plots in monocultures and in mixtures with three ratios (3:1, 2:2 and 1:3) for three years. We then tested in a greenhouse experiment the performance of A. odoratum and C. jacea in pots planted with monocultures and 1:1 mixtures and filled with live and sterile soils collected from the field plots.Results: In the greenhouse experiment, C. jacea produced less aboveground biomass in soil conditioned by C. jacea monocultures than in soil conditioned by A. odoratum monocultures, while the aboveground biomass of A. odoratum in general did not differ between the two monospecific soils. The negative PSF effect was greater in the 1:1 plant mixture than in plant monocultures for A. odoratum but did not differ for C. jacea. In the greenhouse experiment, the performance of C. jacea relative to A. odoratum in the 1:1 plant mixture was negatively correlated to the abundance of C. jacea in the field plot where the soil was collected from. This relationship was significant both in live and sterile soils. However, there was no relationship between the performance of A. odoratum relative to C. jacea in the 1:1 plant mixture in the greenhouse experiment and the abundance of A. odoratum in the field plots.Conclusions: The response of a plant to PSF depends on whether the focal species grows in monocultures or in mixtures and on the identity of the species. Interspecific competition can exacerbate the negative plant-soil feedbacks compared to intraspecific competition when a plant competes with a stronger interspecific competitor. Moreover, the abundance of a species in mixed plant communities, via plant-soil feedback, negatively influences the relative competitiveness of that species when it grows later in interspecific competition, but this effect varies between species. This phenomenon may contribute to the coexistence of competing plants under natural conditions through preventing the dominance of a particular plant species. [ABSTRACT FROM AUTHOR]

Background and aims: Permafrost degradation has the potential to change the Arctic tundra landscape. We observed rapid local thawing of ice-rich permafrost resulting in thaw pond formation, which was triggered by removal of the shrub cover in a field experiment. This study aimed to examine the rate of permafrost thaw and the initial vegetation succession after the permafrost collapse. Methods: In the experiment, we measured changes in soil thaw depth, plant species cover and soil subsidence over nine years (2007-2015). Results: After abrupt initial thaw, soil subsidence in the removal plots continued indicating further thawing of permafrost albeit at a much slower pace: 1 cm y over 2012-2015 vs. 5 cm y over 2007-2012. Grass cover strongly increased after the initial shrub removal, but later declined with ponding of water in the subsiding removal plots. Sedges established and expanded in the wetter removal plots. Thereby, the removal plots have become increasingly similar to nearby 'natural' thaw ponds. Conclusions: The nine years of field observations in a unique shrub removal experiment at a Siberian tundra site document possible trajectories of small-scale permafrost collapse and the initial stage of vegetation recovery, which is essential knowledge for assessing future tundra landscape changes. [ABSTRACT FROM AUTHOR]

Aims: Shrub expansion is common in the tundra biome and has been linked to climate warming. However, the underlying mechanisms are still not fully understood. This study aimed to investigate the seasonal and vertical rooting patterns of different plant functional types, which is important for predicting tundra vegetation dynamics. Methods: We harvested root samples by soil coring and investigated seasonal changes in root biomass and vertical root distribution across a vegetation gradient, focusing on the differences between graminoids and dwarf shrubs, at a northeastern Siberian tundra. Results: Graminoid fine root biomass increased significantly during the growing season, whereas that of shrubs was already high at the beginning and did not change later on. Shrubs had a much shallower rooting pattern than graminoids. Also, shrub roots did not respond to increases in permafrost thawing depth over the growing season, whereas graminoids grew fine roots in deeper, recently thawed soil layers during the growing season. Conclusions: Our results show that shrubs are predominantly shallow-rooted and grow roots earlier than graminoids, which allows shrubs to take advantage of the nutrient pulse after snowmelt in the early growing season. In contrast, the deep-rooted graminoids can access the nutrients in deeper soil and may profit from increasing permafrost thawing depth. The outcome of the competitive interactions between graminoids and shrubs in tundra may depend on the balance between the benefits associated with earlier root growth and deeper root distribution, respectively. The shrub expansion with climate warming observed in recent decades suggests that earlier root growth in the upper soil layer may be more important than increased rooting depth later in the growing season. [ABSTRACT FROM AUTHOR]

It is widely believed that deciduous tundra-shrub dominance is increasing in the pan-Arctic region, mainly due to rising temperature. We sampled dwarf birch ( Betula nana L.) at a northeastern Siberian tundra site and used dendrochronological methods to explore the relationship between climatic variables and local shrub dominance. We found that establishment of shrub ramets was positively related to summer precipitation, which implies that the current high dominance of B. nana at our study site could be related to high summer precipitation in the period from 1960 to 1990. The results confirmed that early summer temperature is most influential to annual growth rates of B. nana. In addition, summer precipitation stimulated shrub growth in years with warm summers, suggesting that B. nana growth may be co-limited by summer moisture supply. The dual controlling role of temperature and summer precipitation on B. nana growth and establishment is important to predict future climate-driven vegetation dynamics in the Arctic tundra. [ABSTRACT FROM AUTHOR]

In many estuarine areas around the world, the safety of human societies depends on the functioning of embankments (dikes) that provide protection against river floods and storm tides. Vegetation on land-side slopes protects these embankments from erosion by heavy rains or overtopping waves. We carried out a field experiment to investigate the effect of plant species diversity on soil loss through erosion on a simulated dike. The experiment included four diversity treatments (1, 2, 4, and 8 species). In the third year of the experiment, we measured net annual soil loss by measuring erosion losses every 2 weeks. We show that loss of plant species diversity reduces erosion resistance on these slopes: net annual soil loss increased twofold when diversity declines fourfold. The different plant species had strongly diverging effects on soil erosion, both in the single-species and in the multi-species plots. Analysis of the dynamics of the individual species revealed that the main mechanism explaining the strong effects of plant species diversity on soil erosion is the compensation or insurance effect, that is, the capacity of diverse communities to supply species to take over the functions of species that went extinct as a consequence of fluctuating environmental conditions. We conclude that the protection and restoration of diverse plant communities on embankments and other vegetated slopes are essential to minimize soil erosion, and can contribute to greater safety in the most densely populated areas of the world. [ABSTRACT FROM AUTHOR]

Arctic tundra ecosystems are warming almost twice as fast as the global average. Permafrost thaw and the resulting release of greenhouse gases from decomposing soil organic carbon have the potential to accelerate climate warming. In recent decades, Arctic tundra ecosystems have changed rapidly, including expansion of woody vegetation, in response to changing climate conditions. How such vegetation changes contribute to stabilization or destabilization of the permafrost is unknown. Here we present six years of field observations in a shrub removal experiment at a Siberian tundra site. Removing the shrub part of the vegetation initiated thawing of ice-rich permafrost, resulting in collapse of the originally elevated shrub patches into waterlogged depressions within five years. This thaw pond development shifted the plots from a methane sink into a methane source. The results of our field experiment demonstrate the importance of the vegetation cover for protection of the massive carbon reservoirs stored in the permafrost and illustrate the strong vulnerability of these tundra ecosystems to perturbations. If permafrost thawing can more frequently trigger such local permafrost collapse, methane-emitting wet depressions could become more abundant in the lowland tundra landscape, at the cost of permafrost-stabilizing low shrub vegetation. [ABSTRACT FROM AUTHOR]

Agricultural intensification may result in important shifts in insect community composition and function, but this remains poorly explored. Studying how groups of species with shared traits respond to local and landscape scale land-use management can reveal mechanisms behind such observed impacts. We tested if ground beetles (Coleoptera: Carabidae) divided into trait groups based on body sizes, wing morphologies and dietary preferences respond differently to farming practise (organic and conventional), farming intensity (measured as yield) and landscape complexity (measured as the proportion of arable land within a 1,000 m radius) across Europe. We used data from 143 farms in five regions in northern and central Europe. Organic farms did not differ in abundance or richness of any trait group compared to conventional farms. As farm scale intensity (yield) increased, overall abundance of beetles decreased, but abundances of small and medium sized beetles, as well as that of wingless beetles, were unaffected. Overall species richness was not affected by yield, whereas consideration of traits revealed that phytophagous and omnivorous beetles were less species rich on farms with high yields. Increasing the proportion of arable land in the landscape increased overall beetle abundance. This was driven by an increase in omnivorous beetles. The total species richness was not affected by an increase in the proportion arable land, although the richness of wingless beetles was found to increase. Potential effects on ecosystem functioning need to be taken into account when designing schemes to maintain agricultural biodiversity, because species with different ecological traits respond differently to local management and landscape changes. [ABSTRACT FROM AUTHOR]

The decomposition of dead organic matter is a major determinant of carbon and nutrient cycling in ecosystems, and of carbon fluxes between the biosphere and the atmosphere. Decomposition is driven by a vast diversity of organisms that are structured in complex food webs. Identifying the mechanisms underlying the effects of biodiversity on decomposition is critical given the rapid loss of species worldwide and the effects of this loss on human well-being. Yet despite comprehensive syntheses of studies on how biodiversity affects litter decomposition, key questions remain, including when, where and how biodiversity has a role and whether general patterns and mechanisms occur across ecosystems and different functional types of organism. Here, in field experiments across five terrestrial and aquatic locations, ranging from the subarctic to the tropics, we show that reducing the functional diversity of decomposer organisms and plant litter types slowed the cycling of litter carbon and nitrogen. Moreover, we found evidence of nitrogen transfer from the litter of nitrogen-fixing plants to that of rapidly decomposing plants, but not between other plant functional types, highlighting that specific interactions in litter mixtures control carbon and nitrogen cycling during decomposition. The emergence of this general mechanism and the coherence of patterns across contrasting terrestrial and aquatic ecosystems suggest that biodiversity loss has consistent consequences for litter decomposition and the cycling of major elements on broad spatial scales. [ABSTRACT FROM AUTHOR]

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To comprehend the potential consequences of biodiversity loss on the leaf litter decomposition process, a better understanding of its underlying mechanisms is necessary. Here, we hypothesize that positive litter mixture effects occur via complementary resource use, when litter species complement each other in terms of resource quality for detritivores. To investigate this, monocultures and mixtures of two leaf litter species varying in quality were allowed to decompose with and without a single macro-detritivore species (the terrestrial woodlice Oniscus asellus). Resource quality of the mixture was assessed by the mean concentration, the dissimilarity in absolute and relative concentrations, and the covariance between nitrogen (N), phosphorus (P) and calcium (Ca) supply. Our results clearly show that litter mixing effects were driven by differences in their resource quality for detritivores. In particular, complementary supply of N and P was a major driver of litter mixing effects. Interestingly, litter mixing effects caused by the addition of woodlice were predominantly driven by N dissimilarity, whereas in their absence, increased P concentration was the main driver of litter mixing effects. These results show that ultimately, litter diversity effects on decomposition may be driven by complementary resource use of the whole decomposer community (i.e., microbes and macro-detritivores). [ABSTRACT FROM AUTHOR]

In eight European study sites (in Spain, Ireland, Netherlands, Germany, Poland, Estonia and Sweden), abundance of breeding farmland bird territories was obtained from 500 × 500 m survey plots (30 per area, N = 240) using the mapping method. Two analyses were performed: (I) a Canonical Correspondence Analysis of species abundance in relation to geographical location and variables measuring agricultural intensification at field and farm level to identify significant intensification variables and to estimate the fractions of total variance in bird abundance explained by geography and agricultural intensification; (II) several taxonomic and functional community indices were built and analysed using GLM in relation to the intensification variables found significant in the CCA. The geographical location of study sites alone explains nearly one fifth (19.5%) of total variation in species abundance. The fraction of variance explained by agricultural intensification alone is much smaller (4.3%), although significant. The intersection explains nearly two fifths (37.8%) of variance in species abundance. Community indices are negatively affected by correlates of intensification like farm size and yield, whereas correlates of habitat availability and quality have positive effects on taxonomic and functional diversity of assemblages. Most of the purely geographical variation in farmland bird assemblage composition is associated to Mediterranean steppe species, reflecting the bio-geographical singularity of that assemblage and reinforcing the need to preserve this community. Taxonomic and functional diversity of farmland bird communities are negatively affected by agricultural intensification and positively affected by increasing farmland habitat availability and quality. [ABSTRACT FROM AUTHOR]

A large proportion of northern peatlands consists of Sphagnum-dominated ombrotrophic bogs. In these bogs, peat mosses ( Sphagnum) and vascular plants occur in an apparent stable equilibrium, thereby sustaining the carbon sink function of the bog ecosystem. How global warming and increased nitrogen (N) deposition will affect the species composition in bog vegetation is still unclear. We performed a transplantation experiment in which mesocosms with intact vegetation were transplanted southward from north Sweden to north-east Germany along a transect of four bog sites, in which both temperature and N deposition increased. In addition, we monitored undisturbed vegetation in control plots at the four sites of the latitudinal gradient. Four growing seasons after transplantation, ericaceous dwarf shrubs had become much more abundant when transplanted to the warmest site which also had highest N deposition. As a result ericoid aboveground biomass in the transplanted mesocosms increased most at the southernmost site, this site also had highest ericoid biomass in the undisturbed vegetation. The two dominant Sphagnum species showed opposing responses when transplanted southward; Sphagnum balticum height increment decreased, whereas S. fuscum height increment increased when transplanted southward. Sphagnum production did not differ significantly among the transplanted mesocosms, but was lowest in the southernmost control plots. The dwarf shrub expansion and increased N concentrations in plant tissues we observed, point in the direction of a positive feedback toward vascular plant-dominance suppressing peat-forming Sphagnum in the long term. However, our data also indicate that precipitation and phosphorus availability influence the competitive balance between Sphagnum, dwarf shrubs and graminoids. [ABSTRACT FROM AUTHOR]

Soil respiration is an important pathway of the C cycle. However, it is still poorly understood how changes in plant community diversity can affect this ecosystem process. Here we used a long-term experiment consisting of a gradient of grassland plant species richness to test for effects of diversity on soil respiration. We hypothesized that plant diversity could affect soil respiration in two ways. On the one hand, more diverse plant communities have been shown to promote plant productivity, which could increase soil respiration. On the other hand, the nutrient concentration in the biomass produced has been shown to decrease with diversity, which could counteract the production-induced increase in soil respiration. Our results clearly show that soil respiration increased with species richness. Detailed analysis revealed that this effect was not due to differences in species composition. In general, soil respiration in mixtures was higher than would be expected from the monocultures. Path analysis revealed that species richness predominantly regulates soil respiration through changes in productivity. No evidence supporting the hypothesized negative effect of lower N concentration on soil respiration was found. We conclude that shifts in productivity are the main mechanism by which changes in plant diversity may affect soil respiration. [ABSTRACT FROM AUTHOR]

Methane (CH4) emissions were compared for an intensively and extensively managed agricultural area on peat soils in the Netherlands to evaluate the effect of reduced management on the CH4 balance. Chamber measurements (photoacoustic methods) for CH4 were performed for a period of three years in the contributing landscape elements in the research sites. Various factors influencing CH4 emissions were evaluated and temperature of water and soil was found to be the main driver in both sites. For upscaling of CH4 fluxes to landscape scale, regression models were used which were specific for each of the contributing landforms. Ditches and bordering edges were emission hotspots and emitted together between 60% and 70% of the total terrestrial CH4 emissions. Annual terrestrial CH4 fluxes were estimated to be 203 (±48%), 162 (±60%) and 146 (±60%) kg CH4 ha−1 and 157 (±63%), 180 (±54%) and 163 (±59%) kg CH4 ha−1 in the intensively managed site and extensively managed site, for 2006, 2007 and 2008 respectively. About 70% of the CH4 was emitted in the summer period. Farm based emissions caused per year an additional 257 kg CH4 ha−1 and 172 kg CH4 ha−1 for the intensively managed site and extensively managed site, respectively. To further evaluate the effect of agricultural activity on the CH4 balance, the annual CH4 fluxes of the two managed sites were also compared to the emissions of a natural peat site with no management and high ground water levels. By comparing the terrestrial and additional farm based emissions of the three sites, we finally concluded that transformation of intensively managed agricultural land to nature development will lead to an increase in terrestrial CH4 emission, but will not by definition lead to a significant increase in CH4 emission when farm based emissions are included. [ABSTRACT FROM AUTHOR]

We hypothesized that the outcome of competition between ericaceous plants and grasses is strongly affected by the concentrations of phenolics in the litter that they produce. To test the effect of phenolic-rich litter on soluble soil nitrogen concentrations, plant nitrogen uptake and inter-specific competition, we conducted a greenhouse experiment with the shrub Calluna vulgaris and the grass Deschampsia flexuosa and their leaf litters . Two litters of C. vulgaris were used, with equal nitrogen concentration but different (high and low) concentrations of total phenolics. The D. flexuosa leaf litter contained lower concentrations of phenolics, but higher concentrations of nitrogen than the C. vulgaris litters. The plants were grown in monocultures and in mixed cultures. Inorganic and dissolved organic nitrogen were measured monthly during the experiment. After four months, we measured above- and belowground biomass and the nutrient concentrations in above- and belowground plant parts. In monocultures, C. vulgaris produced more shoot and root biomass on its own litter than with no litter. Growth of Calluna was reduced on grass litter . D. flexuosa plants produced most biomass on their own litter type, whether in monocultures or in mixed cultures. Addition of Calluna litter stimulated the growth of D. flexuosa both in monoculture and in mixtures. The grass plants outcompeted Calluna both on shrub litter and on grass litter but not when grown without litter. The two C. vulgaris litter types that differed in their concentration of phenolics did not differ in their effects on the competition between the two species or on the production of inorganic and dissolved organic nitrogen. We conclude that the nitrogen content of the litter is more important as a plant feature driving competition between shrubs and grasses than the concentrations of phenolics. [ABSTRACT FROM AUTHOR]

To predict the role of ombrotrophic bogs as carbon sinks in the future, it is crucial to understand how Sphagnum vegetation in bogs will respond to global change. We performed a greenhouse experiment to study the effects of two temperature treatments (17.5 and 21.7°C) and two N addition treatments (0 and 4 g N m−2 year−1) on the growth of four Sphagnum species from three geographically interspersed regions: S. fuscum, S. balticum (northern and central Sweden), S. magellanicum and S. cuspidatum (southern Sweden). We studied the growth and cover change in four combinations of these Sphagnum species during two growing seasons. Sphagnum height increment and production were affected negatively by high temperature and high N addition. However, the northern species were more affected by temperature, while the southern species were more affected by N addition. High temperature depressed the cover of the ‘wet’ species, S. balticum and S. cuspidatum. Nitrogen concentrations increased with high N addition. N:P and N:K ratios indicated P-limited growth in all treatments and co-limitation of P and K in the high N treatments. In the second year of the experiment, several containers suffered from a severe fungal infection, particularly affecting the ‘wet’ species and the high N treatment. Our findings suggest that global change can have negative consequences for the production of Sphagnum species in bogs, with important implications for the carbon sequestration in these ecosystems. [ABSTRACT FROM AUTHOR]

(1) Increased atmospheric nitrogen deposition has shifted plant dominance from ericaceous plants to grass species. To elucidate the reduced competitiveness of heather, we tested the hypothesis that additions of nitrogen reduce the concentrations of phenolics and condensed tannins in ericaceous leaves and retard mycorrhizal colonisation in ericaceous plants. We also tested the negative effects of reduced light intensity on carbon-based secondary compounds and mycorrhizal colonisation in ericaceous plants. (2) We performed a field inventory at three heathland sites in the Netherlands varying in nutrient supply and light intensity. Leaves of ericaceous plants and grasses were collected and analysed for concentrations of tannins, phenolics and nutrients. Similarly, we took root samples to record mycorrhizal colonisation and soil samples to measure the soil mineralisation. In addition, we conducted two-factorial experiments with Calluna vulgaris plants, in which we varied fertiliser and shade levels under greenhouse and field conditions. (3) The field inventory revealed that nitrogen addition and shading both negatively affected the concentration of total phenolics. The total phenolics and condensed tannin concentrations were positively correlated ( P < 0.001), but in the field experiment, the condensed tannins were not significantly affected by the treatments. Our results provide the first evidence that the carbon nutrient balance can be used to predict the amount of total phenolics in the dwarf shrub C. vulgaris. (4) In the field experiments, shading of plants resulted in significantly less mycorrhizal colonisation. Only in the greenhouse experiment did addition of nitrogen negatively affect mycorrhizal colonisation. (5) Our results imply that increased atmospheric nitrogen deposition can depress the tannin concentrations in ericaceous plants and the mycorrhizal colonisation in roots, thereby reducing the plants’ competitiveness with respect to grasses... [ABSTRACT FROM AUTHOR]

We investigated how shoot and root allocation in plants responds to increasing levels of competitive stress at different levels of soil fertility. In addition, we analyzed whether different responses were due to adaptive plasticity or should be attributed to ontogenetic drift. Plantago lanceolata plants were grown during 18 weeks at five plant densities and four nutrient supply levels in pots in the greenhouse. Thereafter root and shoot biomass was measured. There were clear negative effects of increasing plant densities on plant weights revealing strong intraspecific competition. At the lower N-treatments, the proportional allocation to root mass increased with increasing competitive stress, indicating the important role of belowground competition. At the higher N-supply rate, the relationship between competitive stress and shoot to root ratio was neutral. These responses could not be attributed to ontogenetic drift, but could only be explained by assuming adaptive plasticity. It was concluded that at lower N-supplies belowground competition dominates and leads to increased allocation to roots, while at the higher N-supply competition for soil resources and light had balanced impacts on shoot and root allocation. An alternative hypothesis explaining the observed pattern is that light competition has far less pronounced impacts on root–shoot allocation than nutrient deprival. [ABSTRACT FROM AUTHOR]

Increased N deposition in Europe has affected mire ecosystems. However, knowledge on the physiological responses is poor. We measured photosynthetic responses to increasing N deposition in two peatmoss species ( Sphagnum balticum and Sphagnum fuscum) from a 3-year, north–south transplant experiment in northern Europe, covering a latitudinal N deposition gradient ranging from 0.28 g N m−2 year−1 in the north, to 1.49 g N m−2 year−1 in the south. The maximum photosynthetic rate (NPmax) increased southwards, and was mainly explained by tissue N concentration, secondly by allocation of N to the photosynthesis, and to a lesser degree by modified photosystem II activity (variable fluorescence/maximum fluorescence yield). Although climatic factors may have contributed, these results were most likely attributable to an increase in N deposition southwards. For S. fuscum, photosynthetic rate continued to increase up to a deposition level of 1.49 g N m−2 year−1, but for S. balticum it seemed to level out at 1.14 g N m−2 year−1. The results for S. balticum suggested that transplants from different origin (with low or intermediate N deposition) respond differently to high N deposition. This indicates that Sphagnum species may be able to adapt or physiologically adjust to high N deposition. Our results also suggest that S. balticum might be more sensitive to N deposition than S. fuscum. Surprisingly, NPmax was not ( S. balticum), or only weakly ( S. fuscum) correlated with biomass production, indicating that production is to a great extent is governed by factors other than the photosynthetic capacity. [ABSTRACT FROM AUTHOR]

Peat bogs play a large role in the global sequestration of C, and are often dominated by different Sphagnum species. Therefore, it is crucial to understand how Sphagnum vegetation in peat bogs will respond to global warming. We performed a greenhouse experiment to study the effect of four temperature treatments (11.2, 14.7, 18.0 and 21.4°C) on the growth of four Sphagnum species: S. fuscum and S. balticum from a site in northern Sweden and S. magellanicum and S. cuspidatum from a site in southern Sweden. In addition, three combinations of these species were made to study the effect of temperature on competition. We found that all species increased their height increment and biomass production with an increase in temperature, while bulk densities were lower at higher temperatures. The hollow species S. cuspidatum was the least responsive species, whereas the hummock species S. fuscum increased biomass production 13-fold from the lowest to the highest temperature treatment in monocultures. Nutrient concentrations were higher at higher temperatures, especially N concentrations of S. fuscum and S. balticum increased compared to field values. Competition between S. cuspidatum and S. magellanicum was not influenced by temperature. The mixtures of S. balticum with S. fuscum and S. balticum with S. magellanicum showed that S. balticum was the stronger competitor, but it lost competitive advantage in the highest temperature treatment. These findings suggest that species abundances will shift in response to global warming, particularly at northern sites where hollow species will lose competitive strength relative to hummock species and southern species. [ABSTRACT FROM AUTHOR]

Understanding how complex food webs assemble through time is fundamental both for ecological theory and for the development of sustainable strategies of ecosystem conservation and restoration. The build-up of complexity in communities is theoretically difficult, because in random-pattern models complexity leads to instability. There is growing evidence, however, that nonrandom patterns in the strengths of the interactions between predators and prey strongly enhance system stability. Here we show how such patterns explain stability in naturally assembling communities. We present two series of below-ground food webs along natural productivity gradients in vegetation successions. The complexity of the food webs increased along the gradients. The stability of the food webs was captured by measuring the weight of feedback loops of three interacting ‘species’ locked in omnivory. Low predator–prey biomass ratios in these omnivorous loops were shown to have a crucial role in preserving stability as productivity and complexity increased during succession. Our results show the build-up of food-web complexity in natural productivity gradients and pin down the feedback loops that govern the stability of whole webs. They show that it is the heaviest three-link feedback loop in a network of predator–prey effects that limits its stability. Because the weight of these feedback loops is kept relatively low by the biomass build-up in the successional process, complexity does not lead to instability. [ABSTRACT FROM AUTHOR]

Kauri ( Agathis australis (D. Don) Lindl.) occurs naturally in the warm temperate forest of northern New Zealand where it grows mixed with angiosperm tree species. Below mature kauri trees thick organic layers develop in which large amounts of nitrogen are accumulated. This nitrogen seems to be inaccessible to plants. While litter quality can explain the low decomposition rate below kauri, it is not known what causes the accumulation of nitrogen. We hypothesised that kauri tannins reduce nitrogen mineralisation and litter decomposition below kauri. We further hypothesised that high tannin concentrations in the soil would increase the availability of dissolved organic nitrogen relative to the availability of inorganic nitrogen. To test these hypotheses a laboratory incubation was carried out for 1 year. Purified tannins of kauri and of two other common New Zealand tree species were added to samples of the soil organic layer from under a kauri tree. The results suggest that during the first month of incubation the added tannins reduced nitrogen availability by sequestering proteins or by stimulating nitrogen immobilisation. In the long-term, the reduced nitrogen release, which was found following tannin addition, seems attributable to the complexation of proteins by tannins. It further appeared that the addition of tannins did not change the ratio of dissolved organic nitrogen to inorganic nitrogen in the long-term. We conclude that the effect of kauri tannins on nitrogen release offers a good explanation for the accumulation of nitrogen below kauri trees. [ABSTRACT FROM AUTHOR]

When perennial herbs face the risk of being outcompeted in the course of succession, they are hypothesized to either increase their biomass allocation to flowers and seeds or to invest more in vegetative growth. We tested these hypotheses in a 3-year garden experiment with four perennials ( Hypochaeris radicata, Cirsium dissectum, Succisa pratensis and Centaurea jacea) by growing them in the midst of a tall tussock-forming grass ( Molinia caerulea) that may successionally replace them in their natural habitat. In all species except for the short-lived H. radicata, costs of sexual reproduction were significant over the 3 years, since continuous bud removal enhanced total biomass or rosette number. To mimic succession we added nutrients, which resulted in a tripled grass biomass and higher death rates in the shorter-lived species. The simulated succession resulted also in a number of coupled growth responses in the survivors: enhanced plant size as well as elevated seed production. The latter was partly due to larger plant sizes, but mostly due to higher reproductive allocation, which in turn could be partly explained by lower relative somatic costs and by lower root–shoot ratios in the high-nutrient plots. Our results suggest that perennial plants can increase both their persistence and their colonization ability by simultaneously increasing their vegetative size and reproductive allocation in response to enhanced competition and nutrient influxes. These responses can be very important for the survival of a species in a metapopulation context. [ABSTRACT FROM AUTHOR]

To elucidate the impact of high nitrogen (N) deposition on intact bog vegetation, as affected by phosphorus (P) availability, we conducted a 4-year fertilization experiment with N and P at six sites, one with moderate N deposition and five with high N deposition. During the growing season, N (40 kg ha-1 ya-1), P (3 kg ha-1 y-1), or a combination of both elements was applied to the vegetation. The fertilization effects turned out to be additive in nature. Adding P decreased the inorganic N concentration and increased the P concentration in the rhizosphere at two sites. Furthermore, P stimulated Sphagnum growth and Sphagnum net primary productivity (NPP) at two sites; it also seemed lo encourage growth at two other sites including the moderate-deposition site. Vascular plant growth remained largely unaffected but was depressed at one high-deposition site. Adding N increased the concentration of inorganic N in the rhizosphere at the moderate-deposition site and at two of the three high-deposition sites; vascular plant growth and litter production were encouraged at three high-deposition sites. The addition of N depressed Sphagnum height increment at four high deposition sites and reduced Sphagnum NPP at two sites. Shading by vascular plants was of minor importance in explaining the negative effects of N on Sphagnum. We conclude that because P alleviates the negative impact N has on Sphagnum by enhancing its capability to assimilate the deposited N, P availability is a major factor determining the impact of N deposition on Sphagnum production and thus on carbon sequestration in bogs. [ABSTRACT FROM AUTHOR]

Agricultural intensification, greatly accelerated as a result of the EU Common Agricultural Policy (CAP), has led to drastic reductions in the populations of many wild plant and animal species that used to be characteristic of farmland. In 1992, the EU provided the member states with its Agri-environment Regulation 2078/92 to help member states reverse these developments by means of agri-environment schemes. The question is: will the implementation of these schemes be sufficient to restore the biological diversity on farmland? Most studies that have examined the effectiveness of agri-environment schemes have focussed on farmland birds in Great Britain and The Netherlands. So far, the positive effects appear to be limited. Continuous evaluation and adaptation of these schemes is needed to enable the biodiversity on farmland to recover from the EU's former policy. [ABSTRACT FROM AUTHOR]

In an East African savanna herbaceous layer productivity and species composition were studied around Acacia tortilis trees of three different age classes, as well as around dead trees and in open grassland patches. The effects of trees on nutrient, light and water availability were measured to obtain an insight into which resources determine changes in productivity and composition of the herbaceous layer. Soil nutrient availability increased with tree age and size and was lowest in open grassland and highest under dead trees. The lower N:P ratios of grasses from open grassland compared to grasses from under trees suggested that productivity in open grassland was limited by nitrogen, while under trees the limiting nutrient was probably P. N:P ratios of grasses growing under bushes and small trees were intermediate between large trees and open grassland indicating that the understorey of Acacia trees seemed to change gradually from a N-limited to a P-limited vegetation. Soil moisture contents were lower under than those outside of canopies of large Acacia trees suggesting that water competition between trees and grasses was important. Species composition of the herbaceous layer under Acacia trees was completely different from the vegetation in open grassland. Also the vegetation under bushes of Acacia tortilis was different from both open grassland and the understorey of large trees. The main factor causing differences in species composition was probably nutrient availability because species compositions were similar for stands of similar soil nutrient concentrations even when light and water availability was different. Changes in species composition did not result in differences in above-ground biomass, which was remarkably similar under different sized trees and in open grassland. The only exception was around dead trees where herbaceous plant production was 60% higher than under living trees. The results suggest that herbaceous layer productivity did not increase under trees by a higher soil nutrient availability, probably because grass production was limited by competition for water. This was consistent with the high plant production around dead trees because when trees die, water competition disappears but the high soil nutrient availability remains. Hence, in addition to tree soil nutrient enrichment, below-ground competition for water appears to be an important process regulating tree-grass interactions in semi-arid savanna. [ABSTRACT FROM AUTHOR]

The response of plant growth to rising CO2 levels appears todepend on nutrient availability, but it is not known whether the growth of bogplants reacts similarly. We therefore studied the effects of elevatedCO2 in combination with N supply on the growth of Sphagnum mosses and vascular plants in ombrotrophic bogvegetation. Because the growth of Sphagnum is lessnutrient-limited than that of vascular plants, we hypothesized that Sphagnum would benefit from elevated CO2. In ourgreenhouse experiment, peat monoliths (34 cm diameter, 40cm deep) with intact bog vegetation were exposed to ambient (350ppmv) or elevated (560 ppmv) atmosphericCO2 combined with low (no N addition) or high (5 g Nm−2 yr−1 added) N deposition for twogrowing seasons. Elevated atmospheric CO2 had unexpected deleterious effectson the growth of Sphagnum magellanicum, the dominant Sphagnumspecies. Growth was greatly reduced, particularly in the second growing seasonwhen, regardless of N supply, the mosses looked unhealthy. The negativeCO2 effect was strongest in the warmest months, suggesting a combinedeffect of elevated CO2 and the raised temperatures in the greenhouse.High N deposition favored Rhynchospora alba, which became the dominant vascular plant speciesduring the experiment. Biomass increased more when N supply was high. There wereno significant effects of elevated CO2 on vascular plants, althoughelevated CO2 combined with high N supply tended to increase theaboveground vascular plant biomass. As Sphagnum is the maincarbon-sequestrating species in bogs and rising atmospheric CO2levels are likely to be followed by increases in temperature, there is an urgentneed for further research on the combined effects of elevated CO2 andincreased temperature on Sphagnum growth in bog ecosystems. [ABSTRACT FROM AUTHOR]

To elucidate the sensitivity of bog ecosystems to high levels of nitrogen (N) deposition, we investigated the fate of 15N-labeled N deposition in bog vegetation in the Netherlands, both at ambient and increased N deposition. We doubled N deposition by adding 5 g N m−2 y−1 as dissolved NH4NO3 during three growing seasons to large peat monoliths (1.1 m diameter) with intact bog vegetation kept in large outdoor containers. A small amount of 15N tracer was applied at the start of the second growing season, and its distribution among Sphagnum, vascular plant species, and peat was determined at the end of the third growing season. The 15N tracer was also applied to additional plots at the untreated field site to check for initial distribution. One week after addition, 79% of the total amount of 15N retrieved was found in the living Sphagnum layer and less than 10% had been captured by vascular plants. Fifteen months later, 63% of the total amount of 15N retrieved was still present in the living Sphagnum layer at ambient N deposition. Increased N deposition significantly reduced the proportion of 15N in Sphagnum and increased the amount of 15N in vascular plants. Deep-rooting vascular plant species were significantly more 15N enriched, suggesting that at higher atmospheric inputs N penetrates deeper into the peat. Our results provide the first direct experimental evidence for that which has often been suggested: Increased atmospheric N deposition will lead to increased N availability for vascular plants in ombrotrophic mires. [ABSTRACT FROM AUTHOR]

We tested the prediction that the successional replacement of plant species during succession on inland sand dunes results from the effects of an increase in nitrogen mineralization on competitive interactions. The growth and competitive strength of Festuca ovina and Deschampsia flexuosa on soil substrates with different amounts of soil organic matter or nitrogen supply were measured. Small tillers of Festuca ovina and Deschampsia flexuosa were grown in monocultures and 1:1-mixtures on soil columns with undisturbed layers of soil organic matter from different successional age. There was (a) no visible soil organic matter, (b) a thin soil organic layer (0.5 cm) and (c) a thicker soil organic layer (6.0 cm) present on the soil columns. The species were also grown on columns with no visible soil organic matter (bare sand) with two different levels of N fertilization to mimic the increased N mineralization in the older successional stages. In monoculture, Festuca produced more biomass on the substrates with a soil organic layer compared to the unfertilized sand substrate. It also produced more biomass on sand substrates with N fertilization. Deschampsia produced more biomass in treatments with a soil organic layer compared to the bare sand treatments, but did not respond to the ammonium-nitrate addition. In competition, Festuca seemed to be the stronger competitor on the unfertilized sand substrate. Festuca was also the better competitor on the N fertilized sand treatments, while on the treatments with a soil organic layer Deschampsia was the winning species. Our results do not support the hypothesis that an increase in N supply is responsible for the replacement of Festuca by Deschampsia that concur with the accumulation of soil organic matter during succession in inland dunes. [ABSTRACT FROM AUTHOR]

Herbivore grazing is increasingly used as a management tool to prevent the dominance of vegetation by tall grasses or trees. In this report, a model is described that is used to analyze plant-herbivore interactions and their scaling up to landscape scale. The model can be used to predict effects of herbivory on vegetation development. The model is an ecosystem model including modules for carbon and nitrogen cycling through plants, soil organic matter, and atmosphere. Plants compete for light and nitrogen. An herbivory module is included that implements selective foraging by a herbivore in a spatially heterogeneous area. Simulations were done to analyze the effects of herbivore density on vegetation dynamics, to analyze the impact of soil fertility on maximum herbivore density, and to analyze effects of herbivore density on landscapes. Two important points come forward from the model. Maximum herbivore abundance shows a hump-shaped curve along a soil fertility gradient. At higher soil fertility, light competition becomes more important. Herbivory interferes with plant competition, giving the tall, less palatable species a competitive advantage and thereby reducing the food quality and availability and hence the carrying capacity of the area. At a landscape scale, herbivory leads to increased heterogeneity. This increased heterogeneity may increase carrying capacity. The implications of these points for nature management are discussed. [ABSTRACT FROM AUTHOR]

To evaluate the effects of climate change on boreal forest ecosystems, both atmospheric CO2 (to 560 ppmv) and air temperature (by 3°–5°C above ambient) were increased at a forested headwater catchment in southern Norway. The entire catchment (860 m2) is enclosed within a transparent greenhouse, and the upper 20% of the catchment area is partitioned such that it receives no climate treatment and serves as an untreated control. Both the control and treatment areas inside the greenhouse receive deacidified rain. Within 3 years, soil nitrogen (N) mineralization has increased and the growing season has been prolonged relative to the control area. This has helped to sustain an increase in plant growth relative to the control and has also promoted increased N export in stream water. Photosynthetic capacity and carbon–nitrogen ratio of new leaves of most plant species did not change. While the ecosystem now loses N, the long-term fate of soil N is a key uncertainty in predicting the future response of boreal ecosystems to climate change. [ABSTRACT FROM AUTHOR]

The concept of the relative nutrient requirement ( L) that was introduced in the first paper of this series is used to analyse the effects of the dominant plant population on nutrient cycling and nutrient mineralization in wet heathland ecosystems. A distinction is made between the effect that the dominant plant species has on (1) the distribution of nutrients over the plant biomass and the soil compartment of the ecosystem and (2) the recirculation rate of nutrients. The first effect of the dominant plant species can be calculated on the basis of the δ/ k ratio (which is the ratio of the relative mortality to the decomposition constant). The second effect can be analysed using the relative nutrient requirement ( L). The mass loss and the changes in the amounts of N and P in decomposing above-ground and below-ground litter produced by Erica tetralix and Molinia caerulea were measured over three years. The rates of mass loss from both above-ground and below-ground litter of Molinia were higher than those from Erica litter. After an initial leaching phase, litter showed either a net release or a net immobilization of nitrogen or phosphorus that depended on the initial concentrations of these nutrients. At the same sites, mineralization of nitrogen and phosphorus were measured for two years both in communities dominated by Molinia and in communities dominated by Erica. There were no clear differences in the nitrogen mineralization, but in one of the two years, phosphate mineralization in the Molinia-community was significantly higher. On the basis of the theory that was developed, mineralization rates and ratios between amounts of nutrients in plant biomass and in the soil were calculated on the basis of parameters that were independently measured. There was a reasonable agreement between predicted and measured values in the Erica-communities. In the Molinia-communities there were large differences between calculated and measured values, which was explained by the observation that the soil organic matter in these ecosystems still predominantly consisted of Erica-remains. [ABSTRACT FROM AUTHOR]

The model proposed in the first paper in this series predicts that in mixtures of plant species with different rooting depths there will be an inverse correlation between the relative crowding coefficient of the deep rooting species with respect to the shallow rooting one and the frequency of the deep rooting plants. Two field experiments are reported in which this phenomenon was observed. The first experiment involved Plantago lanceolata L. and Lolium perenne L. and the second involved Plantago lanceolata L. and Anthoxanthum odoratum L. In a third competition experiment between Plantago lanceolata L. and Anthoxanthum odoratum L. it was tested whether this observed phenomenon was indeed caused by the differences between the rooting depths of these species. The experiment was divided into one series where different rooting depths were possible and one series where gauze with a narrow mesh width prevented differences between rooting depths. In the series without gauze the predicted frequency-dependence of the relative crowding coefficient was observed, whereas in the series with gauze the relative crowding coefficient was independent of plant frequency. A comparison between the yields in the series with and without gauze suggests that in mixture Plantago is forced to utilize the nutrients from deeper soil layers, whereas this is not the case in monoculture. Although such phenomena complicate the description of the process investigated, it is concluded that the model proposed provides an approximate prediction of most results of the experiments which were carried out. [ABSTRACT FROM AUTHOR]

In a previous paper in this series a model was proposed lor the competition between plant populations with different rooting depths. This model predicts that in mixtures of plant populations with different rooting depths the Relative Yield Total will exceed unity. Secondly it predicts that in these mixtures the relative crowding coefficient of the deep rooting population with respect to the shallow rooting one will decrease with increasing plant frequency. Two competition experiments in deep pots are carried out to test these predictions. In the first experiment a comparison is made between a series in which nutrients limited plant growth severely and a series in which nutrients were applied in a quantity that allowed high productivity. In the nutrient poor series the Relative Yield Total exceeded unity by about twenty per cent, while in the nutrient rich series only a small deviation from one was observed. The second experiment was divided into a series of deep pots in which different rooting depths were possible and a series of shallow pots in which the two species were forced to share the same space. In the series of shallow pots the relative crowding coefficient of the two species was found to be independent of plant frequency. In the series of deep pots the relative corowding coefficient of the two species with respect to each other did decrease significantly with increasing frequency. The observed frequency-dependence for the shallow rooting species could be explained by an extension of the theory presented previously. In the spacing series used in the same experiment a decrease of the shoot to root ratio with increasing plant density was observed. Furthermore, in the replacement series the shoot to root ratio in mixture was found to be lower than in monoculture. The consequences of these phenomena for the estimation of the relative crowding coefficient and the Relative Yield Total on the basis of shoot weights are discussed. However, it is argued that the fact that shoot to root ratios change with plant frequency does not provide an alternative explanation for the frequency-dependence observed. [ABSTRACT FROM AUTHOR]

As an extension of De Wit's competition theory a theoretical description has been developed of competition between plant populations with different rooting depths. This model shows that in mixtures of plants with different rooting depths the value of the Relative Yield Total can be expected to exceed one. Moreover, it predicts the frequency-dependence of the relative crowding coefficient of the deep rooting population with respect to the shallow rooting population. The relationship between properties of plant species and the environment required to establish a stable equilibrium turns out to be surprisingly simple. The shallow rooting species has to have a larger competitive ability sufficient to compensate for the extra nutrients that are exploited by the deeper rooting plants. The dependence of equilibrium plant frequencies on the properties of plant species and the characteristics of the environment is discussed. [ABSTRACT FROM AUTHOR]

We studied the root foraging ability and its consequences for the nutrient acquisition of five grass species that differ in relative growth rate and that occur in habitats that differ widely in nutrient availability. Foraging responses were quantified, based on the performance of the plants in homogeneous and heterogeneous soil environments of the same overall nutrient availability. Although all species tended to produce a significantly higher root length density in a nutrient-rich patch, this response was significant only for the faster-growing species. The increased root length density resulted from small, though not significant, changes in root biomass and specific root length. The effectiveness of root proliferation was determined by quantifying the total amount of nutrients (N and P) accumulated by the plants over the course of the experiment. Plants acquired more N in a heterogeneous environment than in a homogeneous environment, although the total nutrient availability was the same. The ability to acquire nutrients (N or P) in the heterogeneous environment was not related to the ability of species to increase root length density in response to local nutrient enrichment. In contrast to other studies, our results suggest that the role of morphological plasticity of roots in acquiring patchily distributed resources is limited. Possible reasons for this discrepancy are discussed. [ABSTRACT FROM AUTHOR]

An experiment was carried out in two heathland ecosystems, one dominated by Calluna vulgaris and the other by Molinia caerulea, to analyse the effects of soil organic matter accumulation and nutrient mineralization on plant species dynamics during succession. The experiment included one treatment that received nutrient solution and two treatments where the rate of soil organic matter accumulation was reduced by removing litter or accelerated by adding litter. In a fourth treatment the C. vulgaris litter produced in the C. vulgaris-dominated plots was replaced by litter of M. caerulea and vice versa. Treatments were applied over 8 years. Addition of nutrient solution caused C. vulgaris to decline, and grass species to increase sharply, compared to the control plots. Addition of litter enhanced both N mineralization and the biomass of M. caerulea and Deschampsia flexuosa but reduced the biomass of C. vulgaris. The effects of replacing C. vulgaris litter by M. caerulea litter, or vice versa, on N mineralization and species dynamics could not be attributed to differences between the decomposability of the different litter materials that were transferred. The results confirm the hypothesis that increased litter inputs accelerate the rate of species replacement during succession. [ABSTRACT FROM AUTHOR]

The effects of different litter input rates and of different types of litter on soil organic matter accumulation and net N mineralization were investigated in plant communities dominated by Erica tetralix L. or Molinia caerulea (L.) Moench. Plots in which the litter on the soil had repeatedly been removed were compared with plots in the same plant community in which litter had been added to the soil. In another treatment, litter was removed and replaced by litter from the other plant community. Net N mineralization was measured in situ after 5 years. Less soil organic matter and soil N was found in plots in which litter had been removed, compared with control plots, or plots to which litter had been added, but these differences were significant for the Erica sp. soils only. Plots in which litter had been replaced and control plots did not differ significantly in the amount of soil organic matter. However, in both plant communities, the differences agreed with the faster decomposition rate of Molinia sp. litter compared with Erica sp. litter. The gravimetric soil moisture content was correlated positively with the amount of soil organic matter, both in the Erica sp. soils and the Molinia sp. soils. Net N mineralization rates (g N m) differed significantly between treatments for Erica sp. soils but no for Molinia sp. soils. For Erica sp. soils, net N mineralization rates increased with increasing amounts of soil organic matter and soil N. Replacing the litter with Molinia sp. litter (which differs in chemical composition) had no clear additional effect on the net N mineralization rate. [ABSTRACT FROM AUTHOR]

Assesses the impacts of modern agricultural schemes in protecting the biodiversity of agricultural landscapes in Netherlands. Inadequacy of management agreements; Importance of the motivation and expertise of farmers; Absence of positive effects on the conservation of biological diversity.

An addendum is presented concerning the article "Reconciling complexity with stability in naturally assembling food webs," which was published in the volume 449 (2007) issue, pages 599 - 602.

This study focuses on the influence of nitrogen (N) availability andspecies on rhizodeposition and on decomposition of rhizodeposits, roots and soil organic matter. Four perennial grass species were studied that are characteristic of grassland habitats that differ in nutrient availability. These perennial grass species, Holcus lanatus L., Festuca rubra L., Anthoxanthum odoratum L. and Festuca ovina L., were homogeneously labeled with 14CO2. Plants were grown on soil without N addition and with N addition (14 g N m-2). After 8 weeks, plants were harvested and root production and the remaining amount of rhizodeposits in the soil were measured. 14C-labelled roots were incubated in fresh soil. Decomposition was measured of 1) the labeled rhizodeposits in the soil in which theplants had been growing and 2) the labeled dead roots incubated in fresh soil, by trapping the evolved 14CO2, over 69 days. In general, decomposability of both roots and rhizodeposits increased when nitrogen availability increased. Moreover, the speciesdiffered in their response to N. Higher N supply increased total rhizodeposition of H. lanatus and the decomposability of rhizodeposited carbon compounds of this high fertility species was greater than of the low fertility species F. ovina, but lower than of A. odoratum. Thepresented study gives no evidence for a relation between root decomposition rate and the nutrient availability of the habitat of the fourspecies. Overall, we suggest on the basis of the results that species can affect nutrient cycling by differences in rates of rhizodeposition and litter production. This offers a mechanism whereby species can influence species replacement during succession. [ABSTRACT FROM AUTHOR]

During the initial phases of succession on nutrient-poor, mineral substrates dead plant material accumulates rapidly in the soil. This accumulation of soil organic matter can result in a more than 10-fold increase in nitrogen mineralization within a few decades. These changes in soil features have important consequences for plant growth and the competition between plant species, During succession in heathlandsan increase in nutrient mineralization leads to species with low maximum growth rates and low biomass loss rates being replaced by species with high potential growth rates and high biomass losses. The plantproperties responsible for reduced biomass loss rates appear to result in the litter produced being poorly decomposable, whereas the litter from plants with high potential growth rates decomposes more easily. Model simulations suggest that such combinations of plant featuresgreatly influence the increase in mineralization and the change in plant species composition during ecosystem development. Studies in thefield and garden plot experiments confirmed this hypothesis. [ABSTRACT FROM AUTHOR]