Aboveground nutrient cycling and forest development on poor sandy soil

The aboveground nutrient turnover of three ecosystems representing the main stages of heathland forest succession in NW Germany was investigated in a comparative study with regard to nutrient availability of the soil and light availability below the canopy. It was expected that nutrient availability would play a decisive role in forest development on nutrient poor acidic soil. The results show that the input of nutrients into the Calluna heathland is higher than the annual aboveground turnover of N, P, Ca, Mg, and K via litterfall. Compared to the pioneering birch-pine forest, the annual aboveground turnover of nutrients within the Calluna heathland, and therefore the nutrient availability is very low, while the light availability below the canopy is high. The increasing productivity of the growing successional forest is combined with an increasing nutrient turnover, mainly via litter fall. As a result, the increasing nutrient availability favours shade tolerant species with a higher nutrient demand at the late stages of succession. Consequently, the presumed terminal stage of succession, the oak-beech forest, is characterized by low light availability below the canopy and higher nutrient supply according to the resource-ratio hypothesis of Tilman (1985, 1986, 1988).     

Resource availability at three presumed stages of a heathland succession on the Lüneburger Heide,Germany

Abstract. The availability of maj or plant resources was investigated in three vegetation types that were assumed to represent different stages of a secondary succession on heathland on the Lüneburger Heide, northwestern Germany. Canopy transmission and absorption of photosynthetically active radiation (PAR), soil-water availability, and nutrient (Ca, K, Mg, N, P) availability were monitored with high spatial and temporal resolution in (1) a Calluna vulgaris heathland, (2) a pioneer birch-pine forest and (3) a late-successional oak-beech forest, situated close to each other on comparable geological substrate (diluvial). Mean fractional transmission of PAR during summer decreased from 0.48 in the heathland to 0.04 in the oak-beech forest while the fractional canopy absorption increased from 0.49 to 0.92. Soil-water availability as indicated by the soil-water potential, was significantly influenced by differential canopy interception loss and characteristic rooting patterns in the three vegetation types. Annual mean nutrient concentrations in the equilibrium soil solution were similar or, for some elements, increased from the heathland to the birch-pine and the oak-beech forest despite a growing demand. A marked increase was found for the total nutrient pools in the soil-organic layer between early and late successional vegetation types. On the poor glacial parent material, nutrient pools seem to be strongly dependent on stand productivity and litter production which increased with succession. Thus, for nutrients, facilitation seems to be important in this type of succession.     

Litter and forest-floor dynamics in aPinus resinosa plantation in Wisconsin

Summary Breakdown of dry matter and release of nutrients from decomposing leaf litter and forest-floor material were measured in a 34-year-old red pine (Pinus resinosa Ait.) plantation in central Wisconsin using (1) leaf-litter bags (2) litterfall and forest-floor nutrient data and an exponential decay function, and (3) nutrient flux data and a mass balance equation. After one year of decomposition, 77% of the original dry matter in leaf-litter bags remained. The release of macronutrients in decomposing leaf litter was K>Mg>P, S>N>Ca, and the release of micronutrients and aluminum was Mn, B>Al>Cu>Zn. Nitrogen in decomposing leaf litter showed the leaching, accumulation, and final release phases delineated by Berg and Staff⁴. Half-lives of dry matter and nutrients in the forest floor ranged from 0.5 (K) to 39 (Al) yr. Forest-floor turnover rates of the various elements followed the same trends as in leaf-litter bags except that Ca turned over more readily than P, S, and N and Zn turned over more readily than the other micronutrients. A forest-floor nutrient balance sheet confirmed that the macronutrients N and Ca are accumulating most readily in the forest floor. The overall implications of these trends for tree nutrition are discussed.     

The role of plant resources in forest succession: changes in radiation, water and nutrient fluxes, and plant productivity over a 300-yr-long chronosequence in NW-Germany

In the past insufficient attention has been paid to quantitative measurements of resource fluxes in ecosystems that undergo successional change. In this study, simultaneous changes in seven plant resources (photosynthetically active radiation (PAR), water, nitrogen, phosphorus, calcium, magnesium and potassium) are quantified by a chronosequence approach for a 300-yr-long secondary succession on poor soil from Calluna vulgaris heathland to Fagus sylvatica-Quercus petraea late-successional forest (heathland-to-forest succession).Above-ground net primary production increases sevenfold, and total above-ground phytomass about fortyfold during heathland-to-forest succession. Plant organs that capture resources increase much more slowly (leaf area index: threefold; fine root biomass: 1.3-fold). The increase in productivity is based both on higher absorptivity and conversion efficiency of PAR by the canopies of the successional plants.Accumulation of organic material on the forest floor significantly improves soil water availability. Evapotranspiration losses increase early in succession as the growing vegetation increases in both height and leaf area but tend to decrease again in the late-successional community. Drainage losses are at their minimum at the conifer-dominated pioneer forest stage.Accumulation of available nutrients in the soil is a key process in heathland-to-forest succession that significantly improves plant nutrient availability but leads to only minor changes in carbon/nutrient ratios and humus quality. Litter decomposition rates increase and result in a more rapid nutrient turnover in late successional stages. External nutrient inputs (from the atmosphere and soil weathering) significantly contribute to plant nutrient supply early in succession, whereas the internal cycling of nutrients through litter fall and nutrient mineralisation by far exceeds external inputs at the late stages.Vitousek & Reiners' (1975) ecosystem nutrient loss hypothesis is supported by the heathland-to-forest succession data. Odum's (1969) hypotheses on how nutrient cycles change during the course of succession is, in one part, rejected, in part supported. Tilman's (1988) hypothesis on nutrient limitation early, and light limitation late in primary succession is rejected.     

The effect of increased nutrient availability on leaf turnover and aboveground productivity of two evergreen ericaceous shrubs

Summary Leaf turnover and aboveground productivity in relation to nutrient availability were studied in the evergreen shrubs Erica tetralix and Calluna vulgaris. In monospecific stands of these species four levels of nutrient (NPK) availability were created during three growing seasons. Percentage survival and life expectancy of Erica leaves decreased with increasing nutrient availability. For Calluna there was no effect. Winter mortality of Erica leaves was smaller than growing season mortality. These was no difference for Calluna. The timing of leaf mortality of both species was not affected by nutrient treatment. At the end of the experimental period current year leaf biomass, total biomass and current year second year and third year biomass of both species showed a significant increase with increasing nutrient availability. The relative increase was greater for Calluna, except for second and third year biomass. Stem production and stem mortality of both species increased with increasing nutrient availability. The increased stem mortality resulted also for Calluna in an increased leaf turnover (per unit ground area) with increasing nutrient availability. Nutrient cycling in ecosystems dominated by these species will increase with increasing nutrient availability, because of increased leaf and stem turnover and productivity. This phenotypic effect is similar to the effect of the shift in dominance between different species which occurs along natural gradients of nutrient availability.     

Shifts in plant nutrient use strategies under secondary forest succession

In evergreen broad-leaved forests (EBLFs) in Tiantong National Forest Park, Eastern China, we studied the soil chemistry and plant leaf nutrient concentration along a chronosequence of secondary forest succession. Soil total N, P and leaf N, P concentration of the most abundant plant species increased with forest succession. We further examined leaf lifespan, leaf nutrient characteristics and root–shoot attributes of Pinus massoniana Lamb, the early-successional species, Schima superba Gardn. et Champ, the mid-successional species, and Castanopsis fargesii Franch, the late-successional species. These species showed both intraspecific and interspecific variability along succession. Leaf N concentration of the three dominant species increased while N resorption tended to decrease with succession; leaf P and P resorption didn’t show a consistent trend along forest succession. Compared with the other two species, C. fargesii had the shortest leaf lifespan, largest decay rate and the highest taproot diameter to shoot base diameter ratio while P. massoniana had the highest root–shoot biomass ratio and taproot length to shoot height ratio. Overall, P. massoniana used ‘conservative consumption’ nutrient use strategy in the infertile soil conditions while C. fargesii took up nutrients in the way of ‘resource spending’ when nutrient supply increased. The attributes of S. superba were intermediate between the other two species, which may contribute to its coexistence with other species in a wide range of soil conditions.     

Rapid leaf decay and nutrient release in a chinese tallow forest

Summary The Chinese tallow tree, Sapium sebiferum, was introduced to the Texas Gulf Coast in the early 1900's and has spread into coastal prairie habitats. Since coastal prairie contains few deciduous trees, we studied leaf fall dynamics, rate of decomposition, and the quantity and rate of nutrient input from decomposing tallow leaves. Among-year leaf fall averaged 382.6 g·m^-2·yr^-1, similar to other south temperat deciduous forests and about as predicted by Meentemeyer et al.'s (1982) AE-lignin model. Decay of tallow leaves (k=-4.33) was much more rapid than native black willow (k=-0.35) and than other temperate deciduous trees (k=-0.77). The ratio of lignin to initial nitrogen concentration, highly correlated with rate of decomposition for hardwood trees, was low for Chinese tallow and may contribute to rapid leaf decay. Taking AE and lignin content into account, Meentemeyer's (1984) model predicted k=-1.39 for Chinese tallow and k=-0.88 for black willow. Decay of tallow was much faster but decay of willow was slower than predicted, suggesting that decay on the coastal prairie may be controlled by factors other than lignin content and climate. N, P, and K characteristically accumulate as leaves decay. However, these elements did not accumulate as tallow leaves decayed, possibly because high densities of Armadillidium vulgare, a detritivore, reduced immobilization of elements by microbes. This would result in increased turnover of these elements. Accumulation of Al, Fe, Zn, and S in decaying tallow litter may be related to flood-drain cycles on coastel prairie clay soils. Ca, N, K, Mg, and S were added to forest soil in greatest amounts from decaying tallow leaves. Concentrations of P, K, NO₃–N, Zn, Mn, and Fe were significantly higher and Mg and Na were significantly lower in forest than in prairie soil, raising the possibility that Chinese tallow trees altered the distribution of nutrients in the soil profile. We conclude that the Chinese tallow tree may enhance productivity in ecosystems to which it has been introduced by addition of nutrients from rapid decay of leaves.     

黄土高原子午岭林区典型树种叶片N、P再吸收特征

为揭示黄土高原子午岭林区不同演替阶段和植被类型主要树种养分再吸收特征,研究选取4种次生植被树种(白桦、山杨、辽东栎和油松)和2种人工植被树种(刺槐和侧柏),测定其成熟叶、凋落叶和林下土壤碳(C)、氮(N)、磷(P)含量,研究了叶片N、P再吸收率及其与养分指标的关系。结果表明:(1)不同树种叶片养分和林下土壤养分含量存在显著差异,土壤C、N含量和C∶N∶P计量比均表现为演替后期林地(辽东栎和油松)演替前期林地(山杨和白桦)人工林(侧柏和刺槐);(2)不同树种叶片N、P再吸收率分别为17.18%—43.34%和27.13%—58.12%,均表现为演替后期林地人工林演替前期林地,且P的再吸收率总体高于N的再吸收率;(3)不同树种叶片N、P再吸收率与叶片养分指标的关系强于土壤,与养分计量比的相关性大于养分含量的相关性。说明子午岭典型植被会通过叶片N、P再吸收来适应养分限制环境,尤其是演替后期植被再吸收能力更强,研究可为黄土高原植被恢复提供理论依据。     

Is Calluna vulgaris a suitable bio-monitor of management-mediated nutrient pools in heathland ecosystems?

In the face of ongoing atmospheric nutrient loads the employment of management measures to remove nutrients from heathland ecosystems has increased in importance. The present study is the first to analyse whether Calluna vulgaris is a suitable bio-monitor of management-mediated nutrient pools in heathland ecosystems. If Calluna vulgaris proves to be an appropriate indicator, its bio-indicative usage may prove to be a helpful tool for an assessment of management success in heathland ecosystems. In the Lüneburger Heide nature reserve (NW Germany) we analysed the impacts of grazing, mowing, prescribed burning, choppering and sod-cutting on the nutritional status of Calluna vulgaris by measuring nutrient contents (N, P, Ca, Mg, K) of current year's shoots 1 and 5 years after application of management measures. Results were related to management-induced nutrient flows and nutrient pools at the focal heath sites. Our results indicate that the less the physical environment of a heath site was affected by management measures the better the nutrient contents of current year's shoots of Calluna vulgaris mirrored changes in nutrient pools. For low-intensity measures (i.e. grazing, mowing, prescribed burning), shoot nutrient contents were a suitable indicator for changes in nutrient pools, particularly for nutrients with conservative cycles such as P. At grazed and mown sites high output rates of P caused by these measures were well reflected by decreased shoot P content. At burned sites, Calluna vulgaris proved to be a good indicator of changes in nutrient pools of the organic layer, mainly attributable to the deposition of nutrients with ash. In contrast, at sites subjected to high-intensity measures, shoot nutrient contents did not reflect management-mediated shifts in nutrient pools, despite the high nutrient losses caused by choppering and sod-cutting. At these sites, shoot nutrient contents mirrored only the effects of altered mineralisation rates attributable to changes in the physical environment following high-intensity measures. As plant growth and competition in heathlands is considered to be controlled by N or P, shoot N:P ratios are recommended as a tool to indicate whether plant growth tends to be limited by N, by P or by N and P. This, in turn, allows for an assessment of long-term effects of both atmospheric nutrient loads and management-mediated shifts in N and P pools at a focal heath site.     

Nutrient release from decomposing leaf litter of Banksia ornata,Dark Island heathland,South Australia

Distinct O₁ and O₂ layers, representing annual litter fall, enabled the sequential loss of biomass and nutrients (phosphorus and nitrogen) to be reconstructed in undisturbed litter layers of Banksia ornata in the Dark Island heathland, South Australia. Apart from an initial loss in biomass and nitrogen, the dry weight and nutrient content of the O₁ layer, exposed to the desiccating influence of the atmosphere, remained relatively constant until covered by the following year's leaf fall. Under the blanket of newly fallen leaves, biomass decomposition proceeded continuously through autumn, winter, spring, into the dry summer season. Even though the biomass of the decomposing leaf (O₂) layer decreased continuously, its nutrient content remained relatively constant until the summer season was reached when total decomposition and nutrient loss occurred. During spring, fine rootlets invaded the decomposing litter layer (O₂) and, together with decomposer fungi, bacteria and soil fauna, maintained the total nutrient content of the decomposing leaf at a constant level. By late spring-early summer shoot growth of the dominant heath species was initiated, inducing the mobilization of the nutrients stored in the decomposing litter layer.     

Coastal heathland succession influences butterfly community composition and threatens endangered butterfly species

Succession has a strong influence on species diversity and composition of semi-natural open terrestrial ecosystems. While several studies examined the effects of succession on butterflies in grassland and forest ecosystems, the response of heathland butterflies to succession had not been investigated so far. To address this issue we sampled butterfly abundance and environmental parameters on the Baltic island of Hiddensee (NE Germany) along a gradient of coastal heathland succession from grey dunes to birch forest. Our results provide evidence that succession of coastal heathland has a strong influence on butterfly diversity, abundance, and species composition. Thereby grass and tree encroachment present the main threats for heathland butterflies. Diversity and abundance of butterflies were highest in shrub-encroached heath directly followed by early stages of coastal heathland succession (dwarf-shrub heath, grey dune). Both observed threatened species (Hipparchia semele, Plebeius argus) were negatively affected by succession: abundance decreased with increasing vegetation density (both species) and grass cover (P. argus); consequently, the two later successional stages (shrub, birch forest) were not occupied. Our findings highlight the importance of the preservation of early stages of coastal heathland succession for endangered butterfly species. For coastal heathland management we therefore suggest to maintain early successional stages by sheep grazing, mowing or, in case of high nutrient contents, intensive techniques such as sod-cutting or choppering. To a lower extend shrub-encroached sites should also be present, which might be beneficial for overall species richness.     

Do leaf-litter decomposers control biofilm primary production and benthic algal community structure in forest streams? Insights from an outdoor mesocosm experiment

• 1. Forested headwater streams are generally considered to be light-limited ecosystems where primary production is reduced, and the main source of energy and nutrients is composed of allochthonous detritus. We hypothesised that in these ecosystems, the development of primary producers might also be limited by (1) competition for nutrients with leaf-litter decomposers (e.g. bacteria and fungi), and (2) leaf-litter leachates or allelopathic compounds produced by aquatic fungi.
• 2. To test these hypotheses, a 48-day mesocosm experiment was performed in 12 artificial streams containing stream water inoculated with epilithic biofilm suspensions collected from a forested headwater stream. Three different treatments were applied: control without leaf litter (C), microbially conditioned leaf litter added at the beginning of the experiment and left to decompose throughout the experiment (L), or leaf litter renewed three times during the experiment (RL).
• 3. We predicted that (1) the presence of litter, through microbial nutrient immobilisation and allelopathy, would reduce primary production and that (2) this effect would be amplified by litter renewal. We also predicted that nutrient competition would mean that (3) leaf-litter decomposers will alter primary producer community composition and physiology. These predictions were tested by analysing biofilm development, physiology, stoichiometry, and benthic algal community structure. To distinguish between the effects of nutrient immobilisation and allelopathy, the biofilm responses to leaf-litter leachates collected after different microbial conditioning durations were also measured in a parallel laboratory experiment.
• 4. Contrary to our expectations, by day 28, primary producer growth was higher in the mesocosms containing leaf litter (L and RL) despite the rapid decrease in dissolved nutrients when leaf litter was present. After 48 days, the lowest phototrophic biofilm development was observed when leaf litter was renewed (RL), whereas phototrophic biofilm development was similar in the C and L treatments. Biofilm stoichiometry indicated that this effect was most probably related to greater nitrogen limitation in the RL treatment. The presence of leaf litter also affected primary producers' photophysiology, which could be attributed to changes in taxonomic composition and to physiological adjustments of primary producers.
• 5. Laboratory measurements showed that despite a strong inhibition of primary producer growth by unconditioned leaf-litter leachates, microbially conditioned leaf litter had either low or no effects on the development of primary producers.
• 6. These results reveal that leaf-litter decomposers can have both positive and negative effects on primary producers underlining the need to consider microbial interactions when investigating the functioning of forested headwater streams.

     

Do climate and soil influence phenotypic variability in leaf litter,microbial decomposition and shredder consumption?

We tested the hypothesis that water stress and soil nutrient availability drive leaf-litter quality for decomposers and detritivores by relating chemical and physical leaf-litter properties and decomposability of Alnus glutinosa and Quercus robur, sampled together with edaphic parameters, across wide European climatic gradients. By regressing principal components analysis of leaf traits [N, P, condensed tannins, lignin, specific leaf area (SLA)] against environmental and soil parameters, we found that: (1) In Q. robur the condensed tannin and lignin contents increased and SLA decreased with precipitation, annual range of temperature, and soil N content, whereas leaf P increased with soil P and temperature; (2) In A. glutinosa leaves N, P, and SLA decreased and condensed tannins increased with temperature, annual range of temperature, and decreasing soil P. On the other hand, leaf P and condensed tannins increased and SLA decreased with minimum annual precipitation and towards sites with low temperature. We selected contrasting leaves in terms of quality to test decomposition and invertebrate consumption. There were intraspecific differences in microbial decomposition rates (field, Q. robur) and consumption by shredders (laboratory, A. glutinosa). We conclude that decomposition rates across ecosystems could be partially governed by climate and soil properties, affecting litter quality and therefore decomposers and detritivores. Under scenarios of global warming and increased nutrients, these results suggest we can expect species-specific changes in leaf-litter properties most likely resulting in slow decomposition with increased variance in temperatures and accelerated decomposition with P increase.     

Small-scale spatial heterogeneity in the vertical distribution of soil nutrients has limited effects on the growth and development of Prosopis glandulosa seedlings

We examined the above- and below-ground responses of seedlings of the woody shrub Prosopis glandulosa to the spatial heterogeneity of soil nutrients within the root zone. We performed a microcosm experiment where seedlings were grown with different combinations of nutrients [nitrogen (N), phosphorus (P), and both combined (NP)] and under different levels of nutrient heterogeneity (nutrients supplied as patches located in the bottom and/or upper portion of rooting zone versus homogeneous distribution). Seedling morphology and biomass did not show a strong response to changes in nutrient ion or spatial heterogeneity. Height, number of leaves, and specific leaf area did not vary significantly between treatments. The number of leaves, foliar biomass, stem biomass and biomass allocation to stems of seedlings showed more responsiveness to the addition of N and NP than to the addition of P. The spatial heterogeneity of nutrients affected the diameter, root biomass and leaf N content. Seedlings had higher diameter and root biomass when the nutrients were homogeneously distributed as compared to their placement as patches in the bottom of the microcosms. Their leaf N concentration increased in those treatments where the nutrient patch was located in the lower half as compared to the upper half of the microcosms. Root foraging responses to nutrient patches varied with their location. Significant root proliferation was observed when patches of N, P and NP were located in the upper portion of the rooting zone; when they were located in the lower portion such a response was observed only for P. Despite our findings that Prosopis seedlings have a low overall responsiveness to small-scale vertical differences in soil nutrient heterogeneity, our results suggest that these differences may modify the growth dynamics of the secondary roots of this ecologically important dryland species during the early stages of its development.     

A comparative study on nutrient cycling in wet heathland ecosystems

Summary The concept of the relative nutrient requirement (L _n) 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 /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 _n). 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.     

Evaluating a sustainability index for nutrients in a short rotation energy cropping system

In dryland environments 3–5 year rotations of tree crops and agriculture represent a major potential bioenergy feedstock and a means to restore landscape hydrologic balances and phytoremediate sites, while maintaining food production. In soils with low natural fertility, the long‐term viability of these systems will be critically affected by site nutrient status and subsequent cycling of nutrients. A nutrient assimilation index (NAI) was developed to allow comparison of species and tree component nutrient assimilation and to optimize nutrient management, by quantifying different strategies to manage site nutrients. Biomass, nutrient export and nutrient use efficiency were assessed for three short rotation tree crop species. Nutrient exports following harvest at 3 years of high density (4000 trees ha^?1) were consistently higher in Pinus radiata, with values of 85 kg ha^?1 of N, 11kg ha^?1 of P, and 62 kg ha^?1 of K, than Eucalyptus globulus and Eucalyptus occidentalis. Component NAI was generally in the order of leaf?1 for N in leaves of P. radiata to 4.7 Mg kg^?1 for P in stem‐wood of E. occidentalis, indicating higher sustainability of wood biomass compared with leaf biomass. The leaves for each species contained between 40 and 60% of the total nutrient contents while comprising around 25–30% of the total biomass. These nutrient exports via biomass removal are similar to those that follow 3 years of wheat production in the same region, indicating there is no additional drawdown of nutrient reserves during the tree cropping phase of the rotation.     

Biomass partitioning,architecture and turnover of six herbaceous species from habitats with different nutrient supply

Three grasses (Holcus lanatus, Anthoxanthum odoratum and Festuca ovina) and three herbs (Rumex obtusifolius, Plantago lanceolata and Hieracium pilosella) were grown in a greenhouse at 3 nutrient levels in order to evaluate plant allocation, architecture and biomass turnover in relation to fertility level of their habitats.Four harvests were done at intervals of 4 weeks. Various plant traits related to biomass partitioning, plant architecture, biomass turnover and performance were determined. Differences in nutrient supply induced a strong functional response in the species shoot:root allocation, but architecture and turnover showed little or no response. Architectural parameters like specific leaf area and specific root length, however, in general decreased during plant development.Species from more nutrient-rich successional stages were characterized by a larger specific leaf area and longer specific shoot height (height/shoot biomass), resulting in a higher RGR and total biomass in all nutrient conditions. There was no evidence that species from nutrient-poor environments had a longer specific root length or any other superior growth characteristic. The only advantage displayed by these species was a lower leaf turnover when expressed as the fraction of dead leaves and a shorter specific shoot height (SSH) which might prevent herbivory and mowing losses.The dead leaf fraction, which is a good indicator for biomass and nutrient loss, appeared to be not only determined by the leaf longevity, but was also found to be directly related to the RGR of the species. This new fact might explain the slow relative growth rates in species from a nutrient-poor habitat and should be considered in future discussions about turnover.     

Effects of prescribed burning on plant available nutrients in dry heathland ecosystems

Heathland management is an important tool with which to modify ecosystem impacts caused by atmospheric nutrient deposition. Since changes in nutrient availability as a result of management measures affect the outcomes of heathland succession and species competition, studies on this issue are important from both a nature conservation and management point of view. This study reports the effects of prescribed burning on nutrient availability in dry heathland soils and the nutrient content of the two competing heathland species Calluna vulgaris and Deschampsia flexuosa, with particular reference to N and P. We hypothesise that winter prescribed burning leads to additional N availability, which enhances the importance of P in the context of nutrient limitation in heathland ecosystems. In the nature reserve “Lueneburg Heath” (NW Germany) we examined the availability of nutrients in the humus horizons and in the leachate as well as the relevant C:element ratios in Calluna and Deschampsia before and after a burning experiment. Our results show that prescribed burning resulted in drastically increased NH₄⁺ availability in the O-horizon. We observed only short-term effects (for NO₃⁻, PO₄³⁻, Mg) and insignificant effects on the availability of other nutrients (K, Ca). As a consequence of an increased nutrient availability in the humus horizons and a limited nutrient uptake by plants after burning, leaching increased significantly for N, Ca, K, and Mg after burning treatment. No significant changes were found in the foliar C:N ratios for either species after prescribed burning, although Deschampsia showed an increased deficiency for all the other nutrients, particularly for P, as expressed by increased foliar C:P and N:P ratios. By contrast, the nutrient content of Calluna did not change significantly, suggesting that prescribed burning favours the competitive capacity of Calluna as against Deschampsia. We assume that water shortage as a result of changes in the microclimate was mainly responsible for the deterioration of the nutrient content of Deschampsia. This gives Calluna a competitive advantage, enabling it to out-compete Deschampsia on burned heathlands, with respect to the key factor P-limitation.     

VARIATION IN THE NUTRIENT CONTENT OF LEAVES OF QUERCUS ALBA,QUERCUS COCCINEA,AND PINUS RIGIDA IN THE BROOKHAVEN FOREST FROM BUD-BREAK TO ABSCISSION

Leaves of Quercus coccinea, Q. alba, and Pinus rigida were collected at six dates during the growing season and analyzed for N, P, K, Ca, Mg, Fe, S, and Na. Leaf weights per unit of leaf area (or length) were determined for the same period. Quercus coccinea and Q. alba leaves increased in weight per unit area by about 30 % and 50 %, respectively. First-year pine leaves increased in weight per unit length by about 65 %. During the second year the weight of pine leaves changed little. Two broad patterns in the nutrient content of leaves were apparent when nutrient content was expressed on the basis of leaf area rather than leaf weight. N, P, and K concentrations increased to a peak in mid- or late summer and declined abruptly just prior to abscission. Concentrations of other elements tended to rise slowly throughout the life of the leaves in all three species. The differences among nutrients and among species support the hypothesis that differential partitioning of the nutrient pool occurs as a result of evolutionary adaptation. The changes in weight of leaves per unit area and in nutrient content during the growing season are important for studies of net primary production and in appraisals of the cycling of nutrients. Least distortion of nutrient relationships occurs when area or length of leaf is used as the basis for expression of nutrient content.     

Effects of a nutrient pulse supply on nutrient status of the Mediterranean trees Quercus ilex subsp. ballota and Pinus halepensis on different soils and under different competitive pressure

Nutrient availability is a key factor in Mediterranean ecosystems that affects the primary productivity and the community structure. The great variability of its natural availability is now increasing due to frequent fires, pollution events and changes in rainfall regime associated to climate change. Quercus ilex ssp. ballota and Pinus halepensis are the most abundant tree species in the NW Mediterranean basin. They frequently compete in the early and middle successional stages. We investigated the effects of N and P pulse supplies on nutrient uptake capacity in these two species in an after-fire field area and in nursery conditions on different soil types and competing conditions. In the field, N fertilisation had weak effects on nutrient concentration and mineralomass likely as a consequence of this nutrient not being limiting in this field site whereas P fertilisation increased the P mineralomass and the Mg, S, Fe, K and Ca concentrations and mineralomass in the different biomass fractions of both species 1 and 3 years after fertilisation application. In the nursery experiments, P fertilisation increased the mineralomass and concentrations of P, Mg, S, Fe, K and Ca in all biomass fractions including the roots in both species and in different soils and competition conditions. The increment of nutrient mineralomass was due to both the increase of growth and of nutrient concentrations. Both species were able to absorb significant amounts of the P applied by fertilisation (between 5 and 20%) in short time (18 months). Competing vegetation decreased the positive effects of P fertilisation, and in many cases the negative effect of competing vegetation on nutrient mineralomass was stronger when P availability was increased by fertilisation. Q. ilex subsp ballota showed a greater competitive ability for P than the more pioneer species Pinus halepensis in the field but not in the nursery conditions. Pinus halepensis had greater nutrient mineralomass in calcareous than in siliceous soils. Q. ilex subsp. ballota had a higher root biomass allocation and root nutrient allocation than P. halepensis, but both species showed a high capacity to increase their nutrient uptake when its availability increased by fertilisation, thus assuring a great nutrient reserve for future growth periods and contributing to retain nutrients in the ecosystem.