Time‐dependent effects of fertilization on plant biomass in floating fens

Abstract. A cross‐over fertilization experiment was carried out in Dutch floating fens to investigate effects on biomass production in the same and the following years. In total 16 fertilizer treatments were applied, combining four treatments in 1999 with four treatments in 2000 (addition of 20 g.m^?2 N, 5 g.m^?2 P, both elements and unfertilized control). The above‐ground biomass production of vascular plants was co‐limited by N and P in both years. However, in plots that were only fertilized in 1999 the effects of individual nutrients differed between the two years: N‐fertilization slightly increased the amount of biomass produced in the same year (1999), whereas P‐fertilization did so in the following year (2000). Fertilizer applied in 1999 also influenced the effects of fertilizer applied in 2000. One year after N‐fertilization vascular plant growth was still co‐limited by N and P, but one year after P‐fertilization, vascular plant growth was only limited by N. Bryophyte biomass responded weakly to fertilization. Nutrient concentrations in plant biomass, nutrient standing crops and measurements of N and P availability in the soil indicated that one year after fertilization, the N‐fertilizer had mostly ‘disappeared’ from N‐fertilized plots, whereas the availability of P remained markedly enhanced in P‐fertilized plots. In addition, P‐fertilization enhanced the uptake of N by plants the following year. The time‐dependence of fertilizer effects was probably caused by (1) higher addition of P than of N relative to the requirements of plants; (2) longer retention of P than of N in the system; (3) positive effect of P‐fertilization on the availability of N; (4) contrasting effects of N‐ and P‐fertilization on nutrient losses by plants and/or on their responses to subsequent nutrient addition; (5) changing interactions between vascular plants and mosses (mainly Sphagnum spp.); (6) nutrient export through the repeated harvest of above‐ground biomass. To determine which nutrient limits plant growth fertilization experiments should be short, avoiding that indirect effects of a non‐limiting nutrient influence results. To indicate how changed nutrient supply will affect an ecosystem longer‐term experiments are needed, so that indirect effects have time to develop and be detected.     

Nutrient limitation in boreal rich‐fen vegetation: A fertilization experiment

Abstract. Rich‐fen vegetation influenced by hay‐making in the Sølendet Nature Reserve, Central Norway, was fertilized with N, P and K in a full‐factorial fertilization experiment to investigate the nutrient limitation of plant growth at both community and species levels. Above‐ground biomass, shoot density and nutrient concentration were measured in several species and groups of species at three sites after two years of fertilization. At the community level, the results indicate multiple limitation by N and P in the two least productive rich‐fen communities: one characterized by small sedges and herbs, and the other by high abundance of Menyanthes trifoliata and tall sedges. Increased nutrient availability had no effect on a more highly productive, tall‐growing, spring‐influenced community, indicating no nutrient limitation. The results at the species level correspond well with those at the community level, indicating multiple limitation by N and P in most of the dominant and sub‐dominant species. However, P seems to limit growth more than N in Succisa pratensis, and N seems to limit growth more than P in Carex panicea. Furthermore, Eriophorum angustifolium seems to be limited by K. The results did not show which nutrient limits the growth of Carex dioica, C. lasiocarpa and Trichophorum cespitosum. Indications that growth in low‐productive, boreal rich‐fen communities is generally limited by P was not confirmed.     

Soil fertility and fine root dynamics in response to 4 years of nutrient (N,P, K) fertilization in a lowland tropical moist forest,Panama

The question of how tropical trees cope with infertile soils has been challenging to address, in part, because fine root dynamics must be studied in situ. We used annual fertilization with nitrogen (N as urea, 12.5 g N m^?2 year^?1), phosphorus (P as superphosphate, 5 g P m^?2 year^?1) and potassium (K as KCl, 5 g K m^?2 year^?1) within 38 ha of old‐growth lowland tropical moist forest in Panama and examined fine root dynamics with minirhizotron images. We expected that added P, above all, would (i) decrease fine root biomass but, (ii) have no impact on fine root turnover. Soil in the study area was moderately acidic (pH = 5.28), had moderate concentrations of exchangeable base cations (13.4 cmol kg^?1), low concentrations of Bray‐extractable phosphate (PO₄ = 2.2 mg kg^?1), and modest concentrations of KCl‐extractable nitrate (NO₃ = 5.0 mg kg^?1) and KCl‐extractable ammonium (NH₄ = 15.5 mg kg^?1). Added N increased concentrations of KCl‐extractable NO₃ and acidified the soil by one pH unit. Added P increased concentrations of Bray‐extractable PO₄ and P in the labile fraction. Concentrations of exchangeable K were elevated in K addition plots but reduced by N additions. Fine root dynamics responded to added K rather than added P. After 2 years, added K decreased fine root biomass from 330 to 275 g m^?2. The turnover coefficient of fine roots <1 mm diameter ranged from 2.6 to 4.4 per year, and the largest values occurred in plots with added K. This study supported the view that biomass and dynamics of fine roots respond to soil nutrient availability in species‐rich, lowland tropical moist forest. However, K rather than P elicited root responses. Fine roots smaller than 1 mm have a short lifetime (<140 days), and control of fine root production by nutrient availability in tropical forests deserves more study.     

Combined effects of nitrogen enrichment, sulphur pollution and climate change on fen meadow vegetation N:P stoichiometry and biomass

Nitrogen (N) and sulphur (S) deposition, as well as altered soil moisture dynamics due to climate change can have large effects on fen meadow biogeochemistry and vegetation. Their combined effects may differ strongly from their separate effects, since each process affects different nutrients through different mechanisms. However, the impacts of these environmental problems are rarely studied in combination. We therefore investigated the separate and interactive effects of current levels of N- and S-deposition and changes in soil moisture dynamics on fen meadow vegetation. We focused on vegetation biomass and N:P stoichiometry, including access to soil P through root surface phosphatase activity, in a 3-year factorial addition experiment in an N-limited rich fen meadow in the Biebrza valley in Poland. We applied 29.5 kg N ha^?1 year^?1 and 32.1 kg S ha^?1 year^?1, which correspond to current deposition levels in Western Europe. Changes in soil moisture dynamics due to climate change were mimicked by amplified drying of the soil in summer. This level of N-deposition had limited effects on plant biomass production in this rich fen, despite low foliar N:P ratios that suggest N limitation. This level of S-deposition, however, resulted in decreased vegetation P-uptake and biomass. We also showed that increased summer drought resulted in considerable increases in vegetation biomass. We found no interactive effects on vegetation biomass or N:P stoichiometry, possibly as a result of the limited main effects of the separate processes.     

Chemical oxygen demand,nitrogen and phosphorus removal by subsurface wetlands with Phragmites vegetation in different models

To improve the removal efficiency of subsurface wetlands vegetated mainly by Phragmites, pilot‐scale gravel‐based wetlands were used to treat sewage characterized by chemical oxygen demand (COD), total nitrogen (TN) and total phosphorus (TP) pollution. For Phragmites vegetation, COD, TP and TN removal loads of wetland vegetation with Phragmites australis–Typha angustata–Scirpus validus as main species reached 0.517 g m^?2 d^?1, 0.277 g P m^?2 d^?1 and 0.023 g N m^?2 d^?1. The COD removal loads in pilot‐scale and medium‐scale (260 m² in area) wetlands with Phragmites‐monoculture vegetation were 0.62–0.64 g m^?2 d^?1, while that of P. australis–T. angustata–S. validus wetland reached 0.974 g m^?2 d^?1. Thus, the preferable poly‐culture model for Phragmites wetland vegetation was P. australis, T. angustata, S. validus and Zizania latifolia with stem density ratio of 8:1:5:1. After harvest, nitrogen and phosphorus standing stocks of wetland vegetations ranged only 2.2–9.93 g N m^?2 and 5.39–13.5 g P m^?2, respectively, as both the above ground biomass and the nitrogen and phosphorus contents of the wetland vegetation harvested in late autumn were low.     

Differential effects of ammonium and nitrate deposition on fen phanerogams and bryophytes

Question: High atmospheric nitrogen (N) deposition has been shown to affect productivity and species composition of terrestrial ecosystems. This study focused on the differential effects of the two inorganic N forms in atmospheric deposition (i.e. ammonium and nitrate). Methods and location: Nutrient addition experiments were carried out during 4 years in a mesotrophic fen in a low‐deposition area in Ireland. In a factorial design, plots were fertilized with ammonium and/or nitrate, in two doses comparable with 35 and 70 kg N ha^?1 y^?1 and compared with an unfertilized control. Results: Vascular plant biomass as well as bryophyte biomass were not affected by N dose but showed significantly different responses to the N form. In the ammonium‐fertilized plots, vascular plant biomass was higher and moss biomass was lower than the control, while nitrate additions had no effect. Vascular plant species density was high (16 species per 0.49 m²) and was not affected by any of the treatments; bryophyte species density was also high (seven species per 0.04 m²) but showed a significant decrease upon ammonium fertilization. Conclusion: The vulnerability of the mesotrophic vegetation to enhanced atmospheric N deposition depends strongly on the N form. If N would be mainly deposited as NO_x, no detrimental effects on the vegetation will occur. If, however, the deposition is mainly in the form of NH_y, the bryophyte vegetation will be seriously damaged, while the vascular plant vegetation will show an increased biomass production with possible shifts in dominance from Carex and herb species to grasses and shrubs.     

Fen‐meadow succession in relation to spatial and temporal differences in hydrological and soil conditions

Question: In fen meadows with Junco‐Molinion plant communities, falling groundwater levels may not lead to a boosted above‐ground biomass production if limitation of nutrients persists. Instead, depending on drainage intensity and micro‐topography, acidification may trigger a shift into drier and more nutrient‐poor plant communities. Location: Nature reserve, central Netherlands, 5 m a.s.l. Methods: Long‐term study (1988‐1997) in a fen meadow along a gradient in drainage intensity at different scales. Results: Above‐ground biomass increased only slightly over ten years, despite a lower summer groundwater table. The accountable factors were probably a limited availability of nutrients (K in the higher well‐drained plots, P in the intermediate plots and N in the lower hardly drained plots), plus removal of hay. Junco‐Molinion species increased in dry sites and Parvo‐caricetea species increased in wet sites, presumably primarily because of soil acidification occurring when rainwater becomes more influential than base‐rich groundwater. The extent of the shift in species composition depends primarily on the drainage intensity and secondarily on microtopography. Local hydrological measures have largely failed to restore wetter and more basic‐rich conditions. Conclusions: Acidification and nutrient removal, leaching and immobilization resulted in the succession towards Junco‐Molinion at the cost of Calthion palustris elements. Lower in the gradient this change was reduced by the presence of buffered groundwater in slightly drained sites. To conserve the typical plant communities of the Junco‐Molinion to Calthion gradient in the long term, further acidification must be prevented, for example by inundation with base‐rich surface water.     

Non‐destructive allometric estimates of above‐ground and below‐ground biomass of high‐mountain vegetation in the Andes

Aim

Studies that monitor high‐mountain vegetation, such as paramo grasslands in the Andes, lack non‐destructive biomass estimation methods. We aimed to develop and apply allometric models for above‐ground, below‐ground and total biomass of paramo plants.

Location

The paramo of southern Colombia between 1°09′N and 077°50′W, at 3,400 and 3,700 m a.s.l.

Methods

We established 61 1‐m² plots at random locations, excluding disturbed, inaccessible and peat bog areas. We measured heights and basal diameters of all vascular plants in these plots and classified them into seven growth forms. Near each plot, we sampled the biomass from plants of abundant genera, after having measured their height and basal diameter. Hence, we measured the biomass of 476 plants (allometric set). For each growth form we applied power‐law functions to develop allometric models of biomass against basal diameter, height, height x basal diameter and height × basal area. The best models were selected using AIC_c weights. Using the observed and predicted plant biomass of the allometric set we calculated absolute percentage errors using cross‐validation. The biomass of a plot was estimated by summing the predicted biomass of all plants in a plot. Confidence limits around these sums were calculated by bootstrapping.

Results

For groups of <20 plants the biomass predictions yielded large (>15%) errors. Applying groups that resembled the 1‐m² plots in density and composition, the errors for above‐ground and total biomass estimates were <15%. Across all plots, we obtained an above‐ground, below‐ground and total plot biomass of 329 ± 190, 743 ± 486 and 1011 ± 627 g/m² (mean ± SD), respectively. These values were within the range of biomass estimates obtained destructively in the tropical Andes.

Conclusions

In new applications, if target vegetation samples are similar regarding growth forms and genera to our allometric set, their biomass might be predicted applying our equations, provided they contain at least 50–100 plants. In other situations, we would recommend gathering additional biomass measurements from local plants to evaluate new regression equations.     

Below‐ground biomass and productivity of a grazed site and a neighbouring ungrazed exclosure in a grassland in central Argentina

Abstract We estimated the below‐ground net plant productivity (BNPP) of different biomass components in an intensively and continuously 45‐ha grazed site and in a neighbouring exclosure ungrazed for 16 years for a natural mountain grassland in central Argentina. We measured approximately twice as much dead below‐ground biomass in the grazed site as in the ungrazed site, with a strong concentration of total below‐ground biomass towards the upper 10 cm of the soil layer in both sites. The main contribution to total live biomass was accounted for by very fine (<0.5 mm) and fine roots (0.5–1.0 mm) both at the grazed (79%) and at the ungrazed (81%) sites. We measured more dead biomass for almost all root components, more live biomass of rhizomes, tap roots and bulbs, and less live biomass of thicker roots (>1 mm) in the grazed site. The seasonal variation of total live below‐ground biomass mainly reflected climate, with the growing season being limited to the warmer and wetter portion of the year, but such variation was higher in the grazed site. Using different methods of estimation of BNPP, we estimated maximum values of 1241 and 723 g m^?2 year^?1 for the grazed and ungrazed sites, respectively. We estimated that very fine root productivity was almost twice as high at the grazed site as at the ungrazed one, despite the fact that both sites had similar total live biomass, and root turnover rate was twofold at the grazed site.     

Modeling to discern nitrogen fertilization impacts on carbon sequestration in a Pacific Northwest Douglas‐fir forest in the first‐postfertilization year

This study investigated how nitrogen (N) fertilization with 200 kg N ha^?1 of urea affected ecosystem carbon (C) sequestration in the first‐postfertilization year in a Pacific Northwest Douglas‐fir (Pseudotsuga menziesii) stand on the basis of multiyear eddy‐covariance (EC) and soil‐chamber measurements before and after fertilization in combination with ecosystem modeling. The approach uses a data‐model fusion technique which encompasses both model parameter optimization and data assimilation and minimizes the effects of interannual climatic perturbations and focuses on the biotic and abiotic factors controlling seasonal C fluxes using a prefertilization 9‐year‐long time series of EC data (1998–2006). A process‐based ecosystem model was optimized using the half‐hourly data measured during 1998–2005, and the optimized model was validated using measurements made in 2006 and further applied to predict C fluxes for 2007 assuming the stand was not fertilized. The N fertilization effects on C sequestration were then obtained as differences between modeled (unfertilized stand) and EC or soil‐chamber measured (fertilized stand) C component fluxes. Results indicate that annual net ecosystem productivity in the first‐post‐N fertilization year increased by～83%, from 302 ± 19 to 552 ± 36 g m^?2 yr^?1, which resulted primarily from an increase in annual gross primary productivity of～8%, from 1938 ± 22 to 2095 ± 29 g m^?2 yr^?1 concurrent with a decrease in annual ecosystem respiration (R_e) of～5.7%, from 1636 ± 17 to 1543 ± 31 g m^?2 yr^?1. Moreover, with respect to respiration, model results showed that the fertilizer‐induced reduction in R_e (～93 g m^?2 yr^?1) principally resulted from the decrease in soil respiration R_s (～62 g m^?2 yr^?1).     

Nutrient limitation and nutrient‐driven shifts in plant species composition in a species‐rich fen meadow

Question: We studied the development and persistence of the effects of nutrient pulses on biomass production and species composition in a fen meadow. Location: Nature reserve, central Netherlands, 5 m a.s.l. Methods: Single pulse fertilization with N and P in a factorial design on an undrained central and a drained margin site in a species‐rich fen meadow (Cirsio dissecti‐Molinietum). Biomass production and species composition were monitored during four years. Results: At the central site, N addition boosted biomass production, but only during one year. The species composition was not changed. P fertilization increased the biomass production and changed the species composition from a vegetation dominated by Carex panicea to a grassland community with abundant Holcus lanatus, but not before the second year. At the margin site, P fertilization changed the species composition in a similar way, but biomass production was not increased. N fertilization had no effect. At both sites the P induced shift in species composition persisted for four years although the P effect declined during the experiment. Conclusions: The biomass responses show that N was limiting in the central site. Another nutrient, besides N and P (probably K) must have been limiting in the marginal site. The fast decline of the N effect on biomass is ascribed to increased denitrification and biomass removal. The delay in the P effect on biomass and species composition and the persistence of the P effect on species composition are ascribed to fast immobilisation and subsequent slow release of fertilizer P in the peat soil. Recurrence of the P pulses is expected to cause permanent changes in species composition.     

Yield and species composition of a mesic grassland savanna in South Africa are influenced by long‐term nutrient addition

Species composition and productivity of natural grasslands are influenced by soil nutrient status. With high resource availability, productivity is expected to increase, and competition is assumed to gain prominence with predicted exclusion of species of lower competitive ability. During 2010 and 2011 we used the dry weight rank method to measure above‐ground phytomass production of herbage in 96 plots (9 m × 2.7 m) fertilized for 60 years with two forms of nitrogen (N as limestone ammonium nitrate or ammonium sulphate at four levels: 0, 7.1, 14.1, 21.2 g m^?2), phosphorus (P as superphosphate at two levels: 0, 33.6 g m^?2), and lime (two levels: 0, 225 g m^?2). Light attenuation was measured as the proportion of photosynthetically active radiation reaching the lower leaf layers of the grasses and the ground surface. Light conditions beneath the grass layer were reduced by nutrient addition to 30% of full sunlight but remained above 60% in non‐fertilized plots. Grass total above‐ground phytomass production increased with nutrient addition. The strongest yield responses were attained with N plus P addition. Species responses showed that Themeda triandra and Hyparrhenia hirta decreased in above‐ground phytomass production with nutrient addition while Panicum maximum, Eragrostis curvula and E. plana increased. These findings are discussed in terms of competitive interactions among species, their position in the grass canopy and their physiological tolerances to high nitrogen environments.     

Arbuscular mycorrhizal fungi benefit from 7 years of free air CO2 enrichment in well-fertilized grass and legume monocultures

Rising atmospheric carbon dioxide partial pressure (pCO₂) and nitrogen (N) deposition are important components of global environmental change. In the Swiss free air carbon dioxide enrichment (FACE) experiment, the effect of altered atmospheric pCO₂ (35 vs. 60 Pa) and the influence of two different N‐fertilization regimes (14 vs. 56 g N m^?2 a^?1) on root colonization by arbuscular mycorrhizal fungi (AMF) and other fungi (non‐AMF) of Lolium perenne and Trifolium repens were studied. Plants were grown in permanent monoculture plots, and fumigated during the growth period for 7 years. At elevated pCO₂ AMF and non‐AMF root colonization was generally increased in both plant species, with significant effects on colonization intensity and on hyphal and non‐AMF colonization. The CO₂ effect on arbuscules was marginally significant (P=0.076). Moreover, the number of small AMF spores (≤100 μm) in the soils of monocultures (at low‐N fertilization) of both plant species was significantly increased, whereas that of large spores (>100 μm) was increased only in L. perenne plots. N fertilization resulted in a significant decrease of root colonization in L. perenne, including the AMF parameters, hyphae, arbuscules, vesicles and intensity, but not in T. repens. This phenomenon was probably caused by different C‐sink limitations of grass and legume. Lacking effects of CO₂ fumigation on intraradical AMF structures (under high‐N fertilization) and no response to N fertilization of arbuscules, vesicles and colonization intensity suggest that the function of AMF in T. repens was non‐nutritional. In L. perenne, however, AM symbiosis may have amended N nutrition, because all root colonization parameters were significantly increased under low‐N fertilization, whereas under high‐N fertilization only vesicle colonization was increased. Commonly observed P‐nutritional benefits from AMF appeared to be absent under the phosphorus‐rich soil conditions of our field experiment. We hypothesize that in well‐fertilized agricultural ecosystems, grasses benefit from improved N nutrition and legumes benefit from increased protection against pathogens and/or herbivores. This is different from what is expected in nutritionally limited plant communities.     

Fine Root Production and Turnover in a Norway Spruce Stand in Northern Sweden: Effects of Nitrogen and Water Manipulation

Fine root length production, biomass production, and turnover in forest floor and mineral soil (0–30 cm) layers were studied in relation to irrigated (I) and irrigated-fertilized (IL) treatments in a Norway spruce stand in northern Sweden over a 2-year period. Fine roots (<1 mm) of both spruce and understory vegetation were studied. Minirhizotrons were used to estimate fine root length production and turnover, and soil cores were used to estimate standing biomass. Turnover was estimated as both the inverse of root longevity (RTL) and the ratio of annual root length production to observed root length (RTR). RTR values of spruce roots in the forest floor in I and IL plots were 0.6 and 0.5 y⁻¹, respectively, whereas the corresponding values for RTL were 0.8 and 0.9 y⁻¹. In mineral soil, corresponding values for I, IL, and control (C) plots were 1.2, 1.2, and 0.9 y⁻¹ (RTR) and 0.9, 1.1, and 1 y⁻¹ (RTL). RTR and RTL values of understory vegetation roots were 1 and 1.1 y⁻¹, respectively. Spruce root length production in both the forest floor and the mineral soil in I plots was higher than in IL plots. The IL-treated plots gave the highest estimates of spruce fine root biomass production in the forest floor, but, for the mineral soil, the estimates obtained for the I plots were the highest. The understory vegetation fine root production in the I and IL plots was similar for both the forest floor and the mineral soil and higher (for both layers) than in C plots. Nitrogen (N) turnover in the forest floor and mineral soil layers (summed) via spruce roots in IL, I, and C plots amounted to 2.4, 2.1, and 1.3 g N m⁻² y⁻¹, and the corresponding values for field vegetation roots were 0.6, 0.5, and 0.3 g N m⁻² y⁻¹. It was concluded that fertilization increases standing root biomass, root production, and N turnover of spruce roots in both the forest floor and mineral soil. Data on understory vegetation roots are required for estimating carbon budgets in model studies.     

Partitioning of ecosystem respiration of CO2 released during land‐use transition from temperate agricultural grassland to Miscanthus × giganteus

Conversion of large areas of agricultural grassland is inevitable if European and UK domestic production of biomass is to play a significant role in meeting demand. Understanding the impact of these land‐use changes on soil carbon cycling and stocks depends on accurate predictions from well‐parameterized models. Key considerations are cultivation disturbance and the effect of autotrophic root input stimulation on soil carbon decomposition under novel biomass crops. This study presents partitioned parameters from the conversion of semi‐improved grassland to Miscanthus bioenergy production and compares the contribution of autotrophic and heterotrophic respiration to overall ecosystem respiration of CO₂ in the first and second years of establishment. Repeated measures of respiration from within and without root exclusion collars were used to produce time‐series model integrations separating live root inputs from decomposition of grass residues ploughed in with cultivation of the new crop. These parameters were then compared to total ecosystem respiration derived from eddy covariance sensors. Average soil surface respiration was 13.4% higher in the second growing season, increasing from 2.9 to 3.29 g CO₂‐C m^?2 day^?1. Total ecosystem respiration followed a similar trend, increasing from 4.07 to 5.4 g CO₂‐C m^?2 day^?1. Heterotrophic respiration from the root exclusion collars was 32.2% lower in the second growing season at 1.20 g CO₂‐C m^?2 day^?1 compared to the previous year at 1.77 g CO₂‐C m^?2 day^?1. Of the total respiration flux over the two‐year time period, aboveground autotrophic respiration plus litter decomposition contributed 38.46% to total ecosystem respiration while belowground autotrophic respiration and stimulation by live root inputs contributed 46.44% to soil surface respiration. This figure is notably higher than mean figures for nonforest soils derived from the literature and demonstrates the importance of crop‐specific parameterization of respiration models.     

Contrasting impacts of a native and an invasive exotic shrub on flood‐plain succession

Abstract. Question: How do Coriaria arborea, an N‐fixing native shrub, and Buddleja davidii, a non‐N‐fixing exotic shrub, affect N:P stoichiometry in plants and soils during early stages of primary succession on a flood‐plain? Location: Kowhai River Valley, northeast South Island, New Zealand. Methods: We measured soil and foliar nutrient concentrations, light levels, plant community composition and the above‐ground biomass of Coriaria and Buddleja in four successional stages: open, young, vigorous and mature. Results: Coriaria occurred at low density but dominated above‐ground biomass by the vigorous stage. Buddleja occurred at 5.3 ± 1.0 stems/m² in the young stage and reached a maximum biomass of 520–535 g.m^‐2 during the young and vigorous stages. Mineral soil N increased with above‐ground Coriaria biomass (r²= 0.45), but did not vary with Buddleja biomass. In contrast, soil P increased with Buddleja biomass (r²= 0.35), but not with Coriaria biomass. In early successional stages, 70–80% of the species present were exotic, but this declined to about 15% by the mature stage. Exotic plant species richness declined with increasing Coriaria biomass, but no other measures of diversity varied with either Coriaria or Buddleja biomass. Conclusion: These results demonstrate that Buddleja dominates early succession and accumulates P whereas Coriaria dominates later succession and accumulates N. A key ecosystem effect of the invasive exotic Buddleja is alteration of soil N:P stoichiometry.     

Long‐term productivity of Mediterranean herbaceous vegetation after a single phosphorus application

Question: What is the mechanism that underlies long‐term maintenance of high herbaceous productivity after a single application of phosphorus (4.5 gP m^?2 and 9 gP m^?2) in a hilly Mediterranean environment in a phosphorus‐deficient ecosystem? Location: Inland, 15 km E of the Mediterranean coast, W Galilee, Israel (35°15′E, 33°01′N; 500 m asl). Methods: The experiment was established in 1988. Multi‐year data on above‐ground biomass, botanical composition, P content of vegetation and soil, and the grazing management context of the experiment were integrated to construct a feasible account of the P dynamics of the ecosystem. Results: The productivity of the herbaceous component already responded to P application in the first year. The effect on the shrubby component of the ecosystem was marginal. The available (bicarbonate extractable) P in the upper soil layer peaked in the year after application of P and then declined to the original level within 7 years. Despite the decline in available soil P, a high, fluctuating level of herbaceous biomass production was maintained for 20 years. Legume species (Fabaceae) became a prominent constituent of the herbaceous vegetation after the P pulse. Conclusions: The long‐term shift in productivity of the herbaceous component of the grazed ecosystem was triggered by a nutritional pulse that induced a feedback loop based on changes in botanical composition of the herbaceous vegetation, the animal–vegetation interaction, grazing and supplementary feeding regimen of the cattle.     

Above‐ and below‐ground plant biomass response to experimental warming in northern Alaska

Question: Does experimental warming, designed to simulate future warming of the Arctic, change the biomass allocation and mycorrhizal infection of tundra plants? Location: High Arctic tundra near Barrow, Alaska, USA (71°18′N 156°40′W). Methods: Above and below ground plant biomass of all species was harvested following 3–4 yr of 1‐2°C of experimental warming. Biomass allocation and arbuscular mycorrhizal infection were also examined in the two dominant species, Salix rotundifolia and Carex aquatilis. Results: Above‐ground biomass of graminoids increased in response to warming but there was no difference in total plant biomass or the ratio of above‐ground to below‐ground biomass for the community as a whole. Carex aquatilis increased above‐ground biomass and proportionally allocated more biomass above ground in response to warming. Salix rotundifolia increased the amount of above‐ and below‐ground biomass allocated per leaf in response to warming. Mycorrhizal infection rates showed no direct response to warming, but total abundance was estimated to have likely increased in response to warming owing to increased root biomass of S. rotundifolia. Conclusions: The community as a whole was resistant to short‐term warming and showed no significant changes in above‐ or below‐ground biomass despite significant increases in above‐ground biomass of graminoids. However, the patterns of biomass allocation for C. aquatilis and S. rotundifolia did change with warming. This suggests that long‐term warming may result in changes in the above‐ground to below‐ground biomass ratio of the community.     

Fine root production and nutrient content in wet and moist arctic tundras as influenced by chronic fertilization

We used ingrowth cores to estimate fine root production in organic soils of wet sedge and moist tundra ecosystems near Toolik Lake on Alaska's North Slope. Root-free soil cores contained in nylon mesh tubes (5 cm diameter, 20–30 cm long) were placed in control and chronically fertilized (N plus P) plots in mid-August 1994 and were retrieved 1 year later. Estimated fine root production in control plots was 75 g m^–2 year^–1 in wet sedge and 56 g m^–2 year^–1 in moist tussock tundra. Fine root production in fertilized plots was 85 g m^–2 year^–1 in wet sedge and 67 g m^–2 year^–1 in moist tussock tundra. Although our estimates of fine root production were higher on fertilized than control plots, differences were not statistically significant within either tundra type. Comparisons between our estimates of fine root production and other estimates of aboveground (plus rhizome) production on the same (wet sedge tundra) or similar (moist tussock tundra) plots suggest that fine root production was about one-third of total net primary production (NPP) under non-fertilized conditions and about one-fifth of total NPP under chronic fertilization. Fine root N and P concentrations increased with fertilization in both tundra types, but P concentrations increased more than N concentrations in wet sedge tundra, whereas relative increases in N and P concentrations in moist tundra roots were similar. These data are consistent with other studies suggesting that NPP in wet sedge tundra is often P limited and that co-limitation by N and P is more important in moist tussock tundra.     

Effects of nutrient additions on ecosystem carbon cycle in a Puerto Rican tropical wet forest

Wet tropical forests play a critical role in global ecosystem carbon (C) cycle, but C allocation and the response of different C pools to nutrient addition in these forests remain poorly understood. We measured soil organic carbon (SOC), litterfall, root biomass, microbial biomass and soil physical and chemical properties in a wet tropical forest from May 1996 to July 1997 following a 7‐year continuous fertilization. We found that although there was no significant difference in total SOC in the top 0–10 cm of the soils between the fertilization plots (5.42±0.18 kg m^?2) and the control plots (5.27±0.22 kg m^?2), the proportion of the heavy‐fraction organic C in the total SOC was significantly higher in the fertilized plots (59%) than in the control plots (46%) (P<0.05). The annual decomposition rate of fertilized leaf litter was 13% higher than that of the control leaf litter. We also found that fertilization significantly increased microbial biomass (fungi+bacteria) with 952±48 mg kg^?1soil in the fertilized plots and 755±37 mg kg^?1soil in the control plots. Our results suggest that fertilization in tropical forests may enhance long‐term C sequestration in the soils of tropical wet forests.