No abstract

We evaluated whether the kind of nutrient limitation (N, P, or K) may affect species richness-productivity patterns and subsequently may explain variation in species richness and in richness of threatened species. We present a data set from previous studies in wetlands in Poland, Belgium, and The Netherlands and examine species richness-productivity patterns for vascular plants in all 150 sites together as well as for N-, P-, and Klimited sites separately. The kind of nutrient limitation was assessed by N:P, N:K, and K: P ratios in the vegetation. Critical values for these ratios were derived from a literature review of fertilization experiments. The kind of nutrient limitation influenced species richness-productivity patterns in our 150 sites through large differences in productivity. P (co)limitation occurred only at low productivity, K (co)-limitation up to intermediate productivity, and N limitation along the entire productivity gradient. There was a decreasing trend in species richness with increasing productivity for K (co)-limited sites, whereas for both the N-limited sites and P (co)-limited sites a sort of ''filled hump-shaped curve'' was observed. The species richness-productivity relationship for threatened species was restricted to a much narrower productivity range than that for all species. Richness of threatened species was higher in P (co)-limited sites than in N-limited sites, suggesting that increased P availabilities in wetlands may be particularly important in causing disappearance of threatened species in western Europe. The role of nutrient limitation in species richnessproductivity relationships not only reveals mechanisms that may explain variation in species richness and occurrence of threatened species, but it also may be important for nature management practice.

Plant species diversity in Eurasian wetlands and grasslands depends not only on productivity but also on the relative availability of nutrients, particularly of nitrogen and phosphorus. Here we show that the impacts of nitrogen:phosphorus stoichiometry on plant species richness can be explained by selected plant life-history traits, notably by plant investments in growth versus reproduction. In 599 Eurasian sites with herbaceous vegetation we examined the relationship between the local nutrient conditions and community-mean life-history traits. We found that compared with plants in nitrogen-limited communities, plants in phosphorus-limited communities invest little in sexual reproduction (for example, less investment in seed, shorter flowering period, longer lifespan) and have conservative leaf economy traits (that is, a low specific leaf area and a high leaf dry-matter content). Endangered species were more frequent in phosphorus-limited ecosystems and they too invested little in sexual reproduction. The results provide new insight into how plant adaptations to nutrient conditions can drive the distribution of plant species in natural ecosystems and can account for the vulnerability of endangered species.

Soils are the product of a complex suite of chemical, biological, and physical processes. In spite of the importance of soils for society and for sustaining life on earth, our knowledge of soil formation rates and of the influence of biological activity on mineral weathering and geochemical cycles is still limited. In this paper we provide a description of the Damma Glacier Critical Zone Observatory and present a first synthesis of our multidisciplinary studies of the 150-yr soil chronosequence. The aim of our research was to improve our understanding of ecosystem development on a barren substrate and the early evolution of soils and to evaluate the influence of biological activity on weathering rates. Soil pH, cation exchange capacity, biomass, bacterial and fungal populations, and soil organic matter show clear gradients related to soil age, in spite of the extreme heterogeneity of the ecosystem. The bulk mineralogy and inorganic geochemistry of the soils, in contrast, are independent of soil age and only in older soils (>100 yr) is incipient weathering observed, mainly as a decreasing content in albite and biotite by coincidental formation of secondary chlorites in the clay fraction. Further, we document the rapid evolution of microbial and plant munities along the chronosequence.

Summary1. Different grass species dominate grasslands fertilized with nitrogen (N) or phosphorus (P), possibly due to the impact of N : P stoichiometry on competitive interactions. How species compete for nutrients, and whether the mechanisms are similar for N and P, is still not fully understood. 2. We investigated whether the outcome of competition between Alopecurus pratensis and Agrostis capillaris depends on N : P stoichiometry, and on the ability to acquire supplied N or P. Monocultures and mixtures of the two species were grown in pots at nine combinations of N and P supply (N : P ratios 1AE7, 15 and 135, all at three supply levels). After 3 months, we determined plant biomass, morphological traits and nutrient concentrations of plant tissues. 3. N : P supply ratios had similar effects on the growth of the two species but contrasting effects on their competitive strength, as assessed by comparing growth in competition to growth in monoculture: Alopecurus was the stronger competitor under N limitation (N : P ratio 1AE7), whereas Agrostis was an equal or stronger competitor under P limitation (N : P ratio 135). This result resembled patterns of species distribution observed in the field. 4. The strong competitive response of Alopecurus at the low N : P supply ratio was associated with a high investment in root biomass and root length, and a high nitrogen productivity. The competitive response of Agrostis at the high N : P supply ratio was associated with low root mortality and high root phosphatase activity. 5. Our results obtained under N limitation support the pre-emption theory in which plants with the highest root length are able to acquire more N from the soil than their competitors, and therefore can suppress their growth. Under P limitation, however, plant investment in root length could not explain competitor suppression. Here, other factors important in competition for P, such as mycorrhizal hyphal length, root longevity, or exudation rates of P releasing compounds, merit investigation.

Abstract. The above‐ground standing crop and nutrient concentrations in plant material were examined in 45 stands of mire vegetation in the Biebrza peatland, Poland. The stands included flood‐plains, rich fens, transitional fens and bogs. The pattern in nutrient concentrations in the above‐ground plant material resembled the pattern in nutrient concentrations in peatwater and peat which had been investigated in an earlier study. Concentrations of N were quite uniform along the gradient. P‐concentrations were highest in the transitional fen. Critical nutrient concentrations were defined on the basis of a review of nutrient concentrations in plant material from peatlands in which a fertilization experiment had been carried out. Defined critical values for phanerogams were: 13‐14 and 0.7 mg/g dry wt for N and P respectively. Concentrations lower than these values indicate deficiency. P/N ratios ≥ 0.07 indicate N‐deficiency and P/N ratios ≤ 0.04 — 0.05 indicate P‐deficiency. According to these values the Biebrza fens and bogs appear to be primarily deficient in N. The growth of the flood‐plain vegetation does not appear to be restricted by nutrients.

The quantities and spatial distribution of nutrients in savanna ecosystems are affected by many factors, of which fire, herbivory and symbiotic N 2fixation are particularly important. We measured soil nitrogen (N) pools and the relative abundance of N and phosphorus (P) in herbaceous vegetation in five vegetation types in a humid savanna in Tanzania. We also performed a factorial fertilization experiment to investigate which nutrients most limit herbaceous production. N pools in the top 10 cm of soil were low at sites where fires were frequent, and higher in areas with woody legume encroachment, or high herbivore excretion. Biomass production was co-limited by N and P at sites that were frequently burnt or heavily grazed by native herbivores. In contrast, aboveground production was limited by N in areas receiving large amounts of excreta from livestock. N 2 -fixation by woody legumes did not lead to P-limitation, but did increase the availability of N relative to P. We conclude that the effects of fire, herbivory and N 2fixation upon soil N pools and N:P-stoichiometry in savanna ecosystems are, to a large extent, predictable.Electronic supplementary material: The online version of this article (

During the past century, the biomass of woody species has increased in many grassland and savanna ecosystems. As many of these species fix nitrogen symbiotically, they may alter not only soil nitrogen (N) conditions but also those of phosphorus (P). We studied the N-fixing shrub Dichrostachys cinerea in a mesic savanna in Zambia, quantifying its effects upon pools of soil N, P, and carbon (C), and availabilities of N and P. We also evaluated whether these effects induced feedbacks upon the growth of understory vegetation and encroaching shrubs. Dichrostachys cinerea shrubs increased total N and P pools, as well as resin-adsorbed N and soil extractable P in the top 10-cm soil. Shrubs and understory grasses differed in their foliar N and P concentrations along gradients of increasing encroachment, suggesting that they obtained these nutrients in different ways. Thus, grasses probably obtained them mainly from the surface upper soil layers, whereas the shrubs may acquire N through symbiotic fixation and probably obtain some of their P from deeper soil layers. The storage of soil C increased significantly under D. cinerea and was apparently not limited by shortages of either N or P. We conclude that the shrub D. cinerea does not create a negative feedback loop by inducing P-limiting conditions, probably because it can obtain P from deeper soil layers. Furthermore, C sequestration is not limited by a shortage of N, so that mesic savanna encroached by this species could represent a C sink for several decades.We studied the effects of woody encroachment on soil N, P, and C pools, and availabilities of N and P to Dichrostachys cinerea shrubs and to the understory vegetation. Both N and P pools in the soil increased along gradients of shrub age and cover, suggesting that N fixation by D. cinerea did not reduce the P supply. This in turn suggests that continued growth and carbon sequestration in this mesic savanna ecosystems are unlikely to be constrained by nutrient limitation and could represent a C sink for several decades.