The role of above-ground competition and nitrogen vs. phosphorus enrichment in seedling survival of common European plant species of semi-natural grasslands.
ABSTRACT: Anthropogenic activities have severely altered fluxes of nitrogen and phosphorus in ecosystems worldwide. In grasslands, subsequent negative effects are commonly attributed to competitive exclusion of plant species following increased above-ground biomass production. However, some studies have shown that this does not fully account for nutrient enrichment effects, questioning whether lowering competition by reducing grassland productivity through mowing or herbivory can mitigate the environmental impact of nutrient pollution. Furthermore, few studies so far discriminate between nitrogen and phosphorus pollution. We performed a full factorial experiment in greenhouse mesocosms combining nitrogen and phosphorus addition with two clipping regimes designed to relax above-ground competition. Next, we studied the survival and growth of seedlings of eight common European grassland species and found that five out of eight species showed higher survival under the clipping regime with the lowest above-ground competition. Phosphorus addition negatively affected seven plant species and nitrogen addition negatively affected four plant species. Importantly, the negative effects of nutrient addition and higher above-ground competition were independent of each other for all but one species. Our results suggest that at any given level of soil nutrients, relaxation of above-ground competition allows for higher seedling survival in grasslands. At the same time, even at low levels of above-ground competition, nutrient enrichment negatively affects survival as compared to nutrient-poor conditions. Therefore, although maintaining low above-ground competition appears essential for species' recruitment, for instance through mowing or herbivory, these management efforts are likely to be insufficient and we conclude that environmental policies aimed to reduce both excess nitrogen and particularly phosphorus inputs are also necessary.
Project description:Plant functional traits reflect individual and community ecological strategies. They allow the detection of directional changes in community dynamics and ecosystemic processes, being an additional tool to assess biodiversity than species richness. Analysis of functional patterns in plant communities provides mechanistic insight into biodiversity alterations due to anthropogenic activity. Although studies have consi-dered of either anthropogenic management or nutrient availability on functional traits in temperate grasslands, studies combining effects of both drivers are scarce. Here, we assessed the impacts of management intensity (fertilization, mowing, grazing), nutrient stoichiometry (C, N, P, K), and vegetation composition on community-weighted means (CWMs) and functional diversity (Rao's Q) from seven plant traits in 150 grasslands in three regions in Germany, using data of 6 years. Land use and nutrient stoichiometry accounted for larger proportions of model variance of CWM and Rao's Q than species richness and productivity. Grazing affected all analyzed trait groups; fertilization and mowing only impacted generative traits. Grazing was clearly associated with nutrient retention strategies, that is, investing in durable structures and production of fewer, less variable seed. Phenological variability was increased. Fertilization and mowing decreased seed number/mass variability, indicating competition-related effects. Impacts of nutrient stoichiometry on trait syndromes varied. Nutrient limitation (large N:P, C:N ratios) promoted species with conservative strategies, that is, investment in durable plant structures rather than fast growth, fewer seed, and delayed flowering onset. In contrast to seed mass, leaf-economics variability was reduced under P shortage. Species diversity was positively associated with the variability of generative traits. Synthesis. Here, land use, nutrient availability, species richness, and plant functional strategies have been shown to interact complexly, driving community composition, and vegetation responses to management intensity. We suggest that deeper understanding of underlying mechanisms shaping community assembly and biodiversity will require analyzing all these parameters.
Project description:Vachellia sieberiana fixes atmospheric nitrogen (N) and distributes it back into ecosystems. We hypothesize that biological nitrogen fixation in this plant species is limited by competition from the invasive shrub, Chromolaena odorata. Competition would therefore result in the legume plant switching its limited nitrogen (N) sources in phosphorus-poor soils in savannah ecosystems when resources have to be shared. This study investigated the different patterns of N use and growth costs by a native and an introduced leguminous shrubby species. We propose that the two species sharing the same environment might result in competition. The competitive effect would induce in the indigenous legume to better utilize atmospheric-derived N modifying plant growth kinetics and plant mineral concentrations. Seedlings of V. sieberiana were cultivated in natural soil inoculum with low levels of phosphorus (mg L-1 ± SE) of 3.67 ± 0.88. The experiments were divided into two treatments where (i) seedlings of V. sieberiana were subjected to competition by cultivating them together with seedlings of C. odorata, and (ii) seedlings of V. sieberiana were cultivated independently. Although V. sieberiana was subjected to competition, the N2-fixing bacteria that occupied the nodules was Mesorhizobium species, similar to plants not subjected to competition. Total plant biomass was similar between treatments although V. sieberiana plants subjected to competition accumulated more below-ground biomass and showed higher carbon construction costs than plants growing individually. Total plant phosphorus and nitrogen decreased in seedlings of V. sieberiana under competition, whereas no differences were observed in percent N derived from the atmosphere (%NDFA) between treatments. The specific nitrogen utilization rate (SNUR) was higher in V. sieberiana plants subjected to competition while specific nitrogen absorption rate (SNAR) showed the opposite response. Vachellia sieberiana is highly adapted to nutrient-poor savannah ecosystems and can withstand competition from invasive shrubs by utilizing both atmospheric and soil nitrogen sources.
Project description:While mowing-induced changes in plant traits and their effects on ecosystem functioning in semi-arid grassland are well studied, the relations between plant size and nutrient strategies are largely unknown. Mowing may drive the shifts of plant nutrient limitation and allocation. Here, we evaluated the changes in nutrient stoichiometry and allocation with variations in sizes of Leymus chinensis, the dominant plant species in Inner Mongolia grassland, to various mowing frequencies in a 17-yr controlled experiment. Affected by mowing, the concentrations of nitrogen (N), phosphorus (P), and carbon (C) in leaves and stems were significantly increased, negatively correlating with plant sizes. Moreover, we found significant trade-offs between the concentrations and accumulation of N, P, and C in plant tissues. The N:P ratios of L. chinensis aboveground biomass, linearly correlating with plant size, significantly decreased with increased mowing frequencies. The ratios of C:N and C:P of L. chinensis individuals were positively correlated with plant size, showing an exponential pattern. With increased mowing frequencies, L. chinensis size was correlated with the allocation ratios of leaves to stems of N, P, and C by the tendencies of negative parabola, positive, and negative linear. The results of structure equation modeling showed that the N, P, and C allocations were co-regulated by biomass allocation and nutrient concentration ratios of leaves to stems. In summary, we found a significant decoupling effect between plant traits and nutrient strategies along mowing frequencies. Our results reveal a mechanism for how long-term mowing-induced changes in concentrations, accumulations, ecological stoichiometry, and allocations of key elements are mediated by the variations in plant sizes of perennial rhizome grass.
Project description:While bryophytes greatly contribute to plant diversity of semi-natural grasslands, little is known about the relationships between land-use intensity, productivity, and bryophyte diversity in these habitats. We recorded vascular plant and bryophyte vegetation in 85 agricultural used grasslands in two regions in northern and central Germany and gathered information on land-use intensity. To assess grassland productivity, we harvested aboveground vascular plant biomass and analyzed nutrient concentrations of N, P, K, Ca and Mg. Further we calculated mean Ellenberg indicator values of vascular plant vegetation. We tested for effects of land-use intensity and productivity on total bryophyte species richness and on the species richness of acrocarpous (small & erect) and pleurocarpous (creeping, including liverworts) growth forms separately. Bryophyte species were found in almost all studied grasslands, but species richness differed considerably between study regions in northern Germany (2.8 species per 16 m(2)) and central Germany (6.4 species per 16 m(2)) due environmental differences as well as land-use history. Increased fertilizer application, coinciding with high mowing frequency, reduced bryophyte species richness significantly. Accordingly, productivity estimates such as plant biomass and nitrogen concentration were strongly negatively related to bryophyte species richness, although productivity decreased only pleurocarpous species. Ellenberg indicator values for nutrients proved to be useful indicators of species richness and productivity. In conclusion, bryophyte composition was strongly dependent on productivity, with smaller bryophytes that were likely negatively affected by greater competition for light. Intensive land-use, however, can also indirectly decrease bryophyte species richness by promoting grassland productivity. Thus, increasing productivity is likely to cause a loss of bryophyte species and a decrease in species diversity.
Project description:BACKGROUND AND AIMS:During their domestication, maize, bean and squash evolved in polycultures grown by small-scale farmers in the Americas. Polycultures often overyield on low-fertility soils, which are a primary production constraint in low-input agriculture. We hypothesized that root architectural differences among these crops causes niche complementarity and thereby greater nutrient acquisition than corresponding monocultures. METHODS:A functional-structural plant model, SimRoot, was used to simulate the first 40 d of growth of these crops in monoculture and polyculture and to determine the effects of root competition on nutrient uptake and biomass production of each plant on low-nitrogen, -phosphorus and -potassium soils. KEY RESULTS:Squash, the earliest domesticated crop, was most sensitive to low soil fertility, while bean, the most recently domesticated crop, was least sensitive to low soil fertility. Nitrate uptake and biomass production were up to 7 % greater in the polycultures than in the monocultures, but only when root architecture was taken into account. Enhanced nitrogen capture in polycultures was independent of nitrogen fixation by bean. Root competition had negligible effects on phosphorus or potassium uptake or biomass production. CONCLUSIONS:We conclude that spatial niche differentiation caused by differences in root architecture allows polycultures to overyield when plants are competing for mobile soil resources. However, direct competition for immobile resources might be negligible in agricultural systems. Interspecies root spacing may also be too large to allow maize to benefit from root exudates of bean or squash. Above-ground competition for light, however, may have strong feedbacks on root foraging for immobile nutrients, which may increase cereal growth more than it will decrease the growth of the other crops. We note that the order of domestication of crops correlates with increasing nutrient efficiency, rather than production potential.
Project description:Populations of Arnica montana, a characteristic species of nutrient poor grasslands in Central Europe, have been deteriorating over the last decades, especially in lowland regions. Population size has been declining and signs of sexual reproduction are scarce. To start a long-term regeneration program, we investigated the major habitat specific drivers for the decline in Hesse, Germany. Firstly, we conducted a field study to analyze habitat characteristics of 32 Hessian lowland sites, comparing those on which this species has become extinct during the last 15 years with sites of small and declining, as well as large, stable populations. We compared habitat traits focusing on soil parameters, nutrients, and vegetation characteristics. Secondly, we set up a greenhouse experiment to study the response of A. montana seedlings to competition and nutrient input to assess the effects of competition pressure and fertilization. The results show lower carbon-to-nitrogen ratios and higher Ellenberg nitrogen indicator values on sites with extinct populations compared to existing populations. Both pH and Ellenberg soil reaction indicator values were higher on sites with extinct populations. In the greenhouse, the combination of nitrogen addition and competition resulted in lower seedling numbers. While rosette size was not dependent on fertilization, growth was strongly enhanced in the plots lacking vegetation. Both studies suggest that soil nutrient enrichment followed by competition pressure diminishes the number of safe sites for A. montana seedling recruitment and establishment and negatively impacts the growth of existing rosettes, thus leading to the continuous decline of populations. There is an urgent need for actions to reduce unintentional nitrogen deposition in the remaining nutrient poor areas as well as to modify land use to withdraw nutrients from enriched soils in order to preserve the remaining A. montana populations and to create bare ground for the safekeeping and enhancement of self-sustainable populations.
Project description:In suburban regions, vacant lots potentially offer significant opportunities for biodiversity conservation. Recently, in Japan, due to an economic recession, some previously developed lands have become vacant. Little is known, however, about the legacy of earlier earthmoving, which involves topsoil removal and ground leveling before residential construction, on plant community composition in such vacant lots. To understand (dis)assembly processes in vacant lots, we studied 24 grasslands in a suburban region in Japan: 12 grasslands that had experienced earthmoving and 12 that had not. We surveyed plant community composition and species richness, and clarified compositional turnover (replacement of species) and nestedness (nonrandom species loss) by distance-based ?-diversities, which were summarized by PCoA analysis. We used piecewise structural equation modeling to examine the effects of soil properties, mowing frequency, past and present habitat connectivities on compositional changes. As a result, past earthmoving, mowing frequency, soil properties, and past habitat connectivity were found to be the drivers of compositional turnover. In particular, we found legacy effects of earthmoving: earthmoving promoted turnover from native grassland species to weeds in arable lands or roadside by altering soil properties. Mowing frequency also promoted the same turnover, implying that extensive rather than intensive mowing can modify the negative legacy effects and maintain grassland species. Decrease in present habitat connectivity marginally enhanced nonrandom loss of native grassland species (nestedness). Present habitat connectivity had a positive effect on species richness, highlighting the important roles of contemporary dispersal. Our study demonstrates that community assembly is a result of multiple processes differing in spatial and temporal scales. We suggest that extensive mowing at local scale, as well as giving a high conservation priority to grasslands with high habitat connectivity at regional scale, is the promising actions to maintain endangered native grassland species in suburban landscapes with negative legacy effects of earthmoving.
Project description:BACKGROUND AND AIMS:Alpine cushion plants can initially facilitate other species during ecological succession, but later on can be negatively affected by their development, especially when beneficiaries possess traits allowing them to overrun their host. This can be reinforced by accelerated warming favouring competitively strong species over cold-adapted cushion specialists. However, little empirical research has addressed the trait-based mechanisms of these interactions. The ecological strategies of plants colonizing the cushion plant Thylacospermum caespitosum (Caryophyllaceae), a dominant pioneer of subnival zones, were studied in the Western Himalayas. METHODS:To assess whether the cushion colonizers are phylogenetically and functionally distinct, 1668 vegetation samples were collected, both in open ground outside the cushions and inside their live and dead canopies, in two mountain ranges, Karakoram and Little Tibet. More than 50 plant traits related to growth, biomass allocation and resource acquisition were measured for target species, and the phylogenetic relationships of these species were studied [or determined]. KEY RESULTS:Species-based trait-environment analysis with phylogenetic correction showed that in both mountain ranges Thylacospermum colonizers are phylogenetically diverse but functionally similar and are functionally different from species preferring bare soil outside cushions. Successful colonizers are fast-growing, clonal graminoids and forbs, penetrating the cushion by rhizomes and stolons. They have higher root-to-shoot ratios, leaf nitrogen and phosphorus concentrations, and soil moisture and nutrient demands, sharing the syndrome of competitive species with broad elevation ranges typical of the late stages of primary succession. In contrast, the species from open ground have traits typical of stress-tolerant specialists from high and dry environments. CONCLUSION:Species colonizing tight cushions of T. caespitosum are competitively strong graminoids and herbaceous perennials from alpine grasslands. Since climate change in the Himalayas favours these species, highly specialized subnival cushion plants may face intense competition and a greater risk of decline in the future.
Project description:Nutrient enrichment, particularly nitrogen, is an important determinant of plant community productivity, diversity and invasibility in a wetland ecosystem. It may contribute to increasing colonization and dominance of invasive species, such as Phragmites australis, especially during wetland restoration. Providing native species a competitive advantage over invasive species, manipulating soil nutrients (nitrogen) may be an effective strategy to control the invasive species and that management tool is essential to restore the degraded ecosystems. Therefore, we examined competition between Phragmites australis and Melaleuca ericifolia in a greenhouse setting with activated carbon (AC) treatments, followed by cutting of Phragmites shoots in nutrient-rich soils. Additionally, we evaluated the effect of AC on plant-free microcosms in the laboratory, to differentiate direct effects of AC on soil microbial functions from indirect effects. Overall, the objective was to test whether lowering nitrogen might be an effective approach for reducing Phragmites invasion in the wetland. The AC reduced Phragmites total biomass more significantly in repeated cut regime (57%) of Phragmites shoots compared to uncut regime (39%). Conversely, it increased Melaleuca total biomass by 41% and 68% in uncut and repeated cut regimes, respectively. Additionally, AC decreased more total nitrogen in above-ground biomass (41 to 55%) and non-structural carbohydrate in rhizome (21 to 65%) of Phragmites, and less total nitrogen reduction in above-ground biomass (25 to 24%) of Melaleuca in repeated cut compared to uncut regime. The significant negative correlation between Phragmites and Melaleuca total biomass was observed, and noticed that Phragmites acquired less biomass comparatively than Melaleuca in AC-untreated versus AC-treated pots across the cutting frequency. AC also caused significant changes to microbial community functions across Phragmites populations, namely nitrogen mineralization, nitrification, nitrogen microbial biomass and dehydrogenase activity (P???0.05) that may potentially explain changes in plant growth competition between Phragmites and Melaleuca. The overall effects on plant growth, however, may be partially microbially mediated, which was demonstrated through soil microbial functions. Results support the idea that reducing community vulnerability to invasion through nutrient (nitrogen) manipulations by AC with reducing biomass of invasive species may provide an effective strategy for invasive species management and ecosystem restoration.
Project description:Approximately 94% of the land area of the Northern Tibetan Plateau is covered by grasslands, which comprise one of five key livestock producing regions in China. In contrast to most other regions worldwide, these alpine grasslands are much more sensitive to global climate change, thus they are under intense study. The differences in species diversity, plant biomass, and soil properties of five representative's alpine grassland types in the Northern Tibetan Plateau were investigated in this research. The results revealed that 11 community types were identified according to the importance of dominant species and constructive species. There were significant differences in the Margalef index (H), Simpson diversity index (D), Shannon-wiener diversity index (H'), and Pielou evenness index (J) indices between these five alpine grasslands. Further, the above-ground biomass (AGB), below-ground biomass (BGB), total biomass (TB), root:shoot (R/S) ratio, and coverage showed significant differences in 5 alpine grasslands. There were also considerable variations in the pH, total nitrogen concentration (TN), total phosphorus concentration (TP), soil organic carbon (SOC) and C-to-N ratio (C:N) among the five alpine grasslands. The highest value of biomass and soil characteristics was always in the alpine steppe (AS), or AM, while the lowest of that was in the alpine desert steppe (ADS), or alpine desert (AD). Moreover, there were significant differences in the soil particle size fractions between the five alpine grasslands. In the AM and AS, the dominant soil particle was clay, while in the alpine meadow-steppe (AMS), ADS, and AD it was fine and medium sand. Substantial correlations were found between the biomass and species diversity indices H, D or H' and soil TN, TP, or SOC. Moreover, silt had a significantly positive correlation with soil C:N, BGB, TB, and R/S, while medium sand and coarse sand was significant negatively correlated. With regard to these grassland types, it is proposed that the AM or AS may be an actively changing grassland types in the Northern Tibetan Plateau.