Stable soil nitrogen accumulation and flexible organic matter stoichiometry during primary floodplain succession

Large increases in nitrogen (N) inputs to terrestrial ecosystems typically have small effects on immediate N outputs because most N is sequestered in soil organic matter. We hypothesized that soil organic N storage and the asynchrony between N inputs and outputs result from rapid accumulation of N in stable soil organic pools. We used a successional sequence on floodplains of the Tanana River near Fairbanks, Alaska to assess rates of stable N accumulation in soils ranging from 1 to 500+ years old. One-year laboratory incubations with repeated leaching separated total soil N into labile (defined as inorganic N leached) and stable (defined as total minus labile N) pools. Stable N pools increased faster (2 g N m^–2 yr^–1) than labile N (0.4 g N m^–2 yr^–1) pools during the first 50 years of primary succession; labile N then plateaued while stable and total N continued to increase. Soil C pools showed similar trends, and stable N was correlated with stable C (r² = 0.95). From 84 to 95 % of soil N was stable during our incubations. Over successional time, the labile N pool declined as a proportion of total N, but remained large on an aerial basis (up to 38 g N m^–2). The stoichiometry of stable soil N changed over successional time; C:N ratios increased from 10 to 22 over 275 years (r² = 0.69). A laboratory ¹⁵N addition experiment showed that soils had the capacity to retain much more N than accumulated naturally during succession. Our results suggest that most soil N is retained in a stable organic pool that can accumulate rapidly but is not readily accessible to microbial mineralization. Because stable soil organic matter and total ecosystem organic matter have flexible stoichiometry, net ecosystem production may be a poor predictor of N retention on annual time scales.     

The nitrogen cycle in lodgepole pine forests,southeastern Wyoming

Storage and flux of nitrogen were studied in several contrasting lodgepole pine (Pinus contorta spp.latifolia) forests in southeastern Wyoming. The mineral soil contained most of the N in these ecosystems (range of 315–860 g · m^–2), with aboveground detritus (37.5–48.8g · m^–2) and living biomass (19.5–24.0 g · m^–2) storing much smaller amounts. About 60–70% of the total N in vegetation was aboveground, and N concentrations in plant tissues were unusually low (foliage = 0.7% N), as were N input via wet precipitation (0.25 g · m^–2 · yr^–1), and biological fixation of atmospheric N (<0.03 g · m^–2 · yr^–1, except locally in some stands at low elevations where symbiotic fixation by the leguminous herbLupinus argenteus probably exceeded 0.1 g · m^–2 · yr^–1).Because of low concentrations in litterfall and limited opportunity for leaching, N accumulated in decaying leaves for 6–7 yr following leaf fall. This process represented an annual flux of about 0.5g · m^–2 to the 01 horizon. Only 20% of this flux was provided by throughfall, with the remaining 0.4g · m^–2 · yr^–1 apparently added from layers below. Low mineralization and small amounts of N uptake from the 02 are likely because of minimal rooting in the forest floor (as defined herein) and negligible mineral N (< 0.05 mg · L^–1) in 02 leachate. A critical transport process was solubilization of organic N, mostly fulvic acids. Most of the organic N from the forest floor was retained within the major tree rooting zone (0–40 cm), and mineralization of soil organic N provided NH₄ for tree uptake. Nitrate was at trace levels in soil solutions, and a long lag in nitrification was always observed under disturbed conditions. Total root nitrogen uptake was calculated to be 1.25 gN · m^–2 · yr^–1 with estimated root turnover of 0.37-gN · m^–2 · yr^–1, and the soil horizons appeared to be nearly in balance with respect to N. The high demand for mineralized N and the precipitation of fulvic acid in the mineral soil resulted in minimal deep leaching in most stands (< 0.02 g · m^–2 · yr^–1). These forests provide an extreme example of nitrogen behavior in dry, infertile forests.     

Nitrogen mineralization,nitrification and denitrification in upland and wetland ecosystems

Summary Nitrogen mineralization, nitrification, denitrification, and microbial biomass were evaluated in four representative ecosystems in east-central Minnesota. The study ecosystems included: old field, swamp forest, savanna, and upland pin oak forest. Due to a high regional water table and permeable soils, the upland and wetland ecosystems were separated by relatively short distances (2 to 5 m). Two randomly selected sites within each ecosystem were sampled for an entire growing season. Soil samples were collected at 5-week intervals to determine rates of N cycling processes and changes in microbial biomass. Mean daily N mineralization rates during five-week in situ soil incubations were significantly different among sampling dates and ecosystems. The highest annual rates were measured in the upland pin oak ecosystem (8.6 g N m^–2 yr^–1), and the lowest rates in the swamp forest (1.5 g N m^–2 yr^–1); nitrification followed an identical pattern. Denitrification was relatively high in the swamp forest during early spring (8040 g N₂O–N m^–2 d^–1) and late autumn (2525 g N₂O–N m^–2 d^–1); nitrification occurred at rates sufficient to sustain these losses. In the well-drained uplands, rates of denitrification were generally lower and equivalent to rates of atmospheric N inputs. Microbial C and N were consistently higher in the swamp forest than in the other ecosystems; both were positively correlated with average daily rates of N mineralization. In the subtle landscape of east-central Minnesota, rates of N cycling can differ by an order of magnitude across relatively short distances.     

Benithic-pelagic coupling of nutrient metabolism along an estuarine eutrophication gradient

The shallow, brackish (11–18% salinity) Roskilde Fjord represents a eutrophication gradient with annual averages of chlorophyll, ranging from 3 to 25 mg chl a m^–3. Nutrient loadings in 1985 were 11.3–62.4 g N m^–2 yr^–1 and 0.4–7.3 g P m^–2 yr^–1. A simple one-layer advection-diffusion model was used to calculate mass balances for 7 boxes in the fjord. Net loss rates varied from –32.2 to 17.9 g P m^–2 yr^–1 and from –3.3 to 66.8 g N m^–2, corresponding to 74% of the external P-loading and 88% of the external N-loading to the entire estuary.Gross sedimentation rates measured by sediment traps were between 7 and 52 g p m^–2 yr^–1 and 50 and 426 g N M^–2 yr^–1, respectively. Exchangeable sediment phosphorus varied in annual average between 2.0 and 4.8 g P m^–2 and exchangeable sediment nitrogen varied from 1.9 to 33.1 g N m–1. Amplitudes in the exchangeable pools followed sedimentation peaks with delays corresponding to settling rates of 0.3 m d^–1. Short term nutrient exchange experiments performed in the laboratory with simultaneous measurements of sediment oxygen uptake showed a release pattern following the oxygen uptake, the changes in the exchangeable pools and the sedimentation peaks.The close benthic-pelagic coupling also exists for the denitrification with maxima during spring of 5 to 20 mmol N m^–2 d–1. Denitrification during the nitrogen-limited summer period suggests dependence on nitrification. Comparisons with denitrification from other shallow estuaries indicate a maximum for denitrification in estuaries of about 250 µmol N m^–2 h^–2 achieved at loading rates of about 25–125 g N m^–2 yr^–1.     

The soil fauna of a beech forest on limestone: trophic structure and energy budget

Summary The soil fauna of a mull beech forest on lime-stone in southern Lower Saxony (West Germany) was sampled quantitatively. Biomass estimates, trophic characteristics, and measurement and calculation of the energetic parameters of the constituent animal populations were used to construct an energy budget of the total heterotrophic subsystem of the forest. Mean annual zoomass amounted to about 15 g d wt m^–2; earthworms (about 10 g d wt m^–2) and other groups of the macrofauna were dominant. Protozoa constituted about 1.5 g d wt m^–2. Relative distribution of zoomass among the trophic categories was 50% macrosaprophages, 30% microsaprophages, 12% microphytophages, and 4% zoophages. Total annual consumption rate of the saprophagous and microphytophagous soil fauna (6328 and 4096 kJ m^–2 yr^–1, respectively) was of the same order of magnitude as annual litter fall (canopy leaves 6124 kJ m^–2 yr^–1, flowers and fruits 944 kJ m^–2 yr^–1, herbs 1839 kJ m^–2 yr^–1, fine woody material 870 kJ m^–2 yr^–1, tree roots 3404 kJ m^–2 yr^–1, without coarse woody litter). Primary decomposers (macrosaprophages) were the key group for litter comminution and translocation onto and into the soil, thus contributing to the high decomposition rate (k=0.8) for leaf litter. Consumption rates of the other trophic groups were (values as kJ m^–2 yr^–1): bacteriophages 2954, micromycophages 416, zoophages 153. Grazing pressure of macrophytophages (including rhizophages) was low. Faeces input from the canopy layer was not significant. Grazing pressure on soil microflora almost equalled microbial biomass; hence, a large fraction of microbial production is channelled into the animal component. Predator pressure on soil animals is high, as a comparison between consumption rates by zoophages and production by potential prey — mainly microsaprophages, microphytophages and zoophages — demonstrated. Soil animals contributed only about 11% to heterotrophic respiration. However, there is evidence that animals are important driving variables for matter and energy transfer: key processes are the transformation of dead organic material and grazing on the microflora. It is hypothesized that the soil macrosaprophages are donor-limited.     

Chitin production by arthropods in the hydrosphere

Chitin is widely distributed in nature and its annual production is thought to be huge. However, the chitin production has been rarely estimated in aquatic ecosystems, despite the growing economic interest in this polymer. Arthropods are one of the main chitin producers in the hydrosphere and a correct evaluation of the chitin production by these organisms in the different marine and freshwater ecosystems is of prime interest to understand their importance in the biogeochemical cycles of carbon and nitrogen. Such evaluation is also worth considering to achieve a rational exploitation of crustaceans which are currently the major source of chitin for the industry. Annual chitin production of crustaceans and insects in aquatic ecosystems was estimated on the basis of annual tissue production estimates and body chitin content measurements. About 800 annual tissue production estimates were collected from the literature. Estimates mainly concerned continental fresh waters and neritic ecosystems. Data were almost inexistent for athalassohaline and oceanic ecosystems. On the whole, 60% of the production estimates fell between 0.1 and 10.0 g dry weight m^–2 yr^–1. Published chitin levels in crustaceans and insects ranged from 3 to 16% of the whole body dry weight. Data were, however, lacking for some major groups such as trichopterans or amphipods. Aquatic insects and crustaceans were therefore collected and assayed for chitin using a highly specific enzymatic method. The chitin content of the collected insects (Coleoptera, Diptera, Ephemeroptera, Odonata, Plecoptera, Trichoptera) varied from 3 to 10% of the whole body dry weight; that of the collected crustaceans (Amphipoda, Branchiopoda, Copepoda) from 2.5 to 8.5% of the whole body dry weight. Total annual chitin production by arthropods had been estimated to 28 × 10⁶ T chitin yr^–1 for the freshwater ecosystems, to 6 × 10⁶ T chitin yr^–1 for athalassohaline ecosystems and to 1328 × 10⁶ T chitin yr^–1 for marine ecosystems. The importance of the chitin production corresponding to the formation of exuviae and peritrophic membranes in arthropods and the chitin production by non-arthropod organisms in the chitin budget of aquatic ecosystems was highlighted and discussed.     

Nitrogen mineralization in heathland ecosystems dominated by different plant species

Net N mineralization rates were measured in heathlands still dominated by ericaceous dwarf shrubs (Calluna vulgaris or Erica tetralix) and in heathlands that have become dominated by grasses (Molinia caerulea or Deschampsia flexuosa). Net N mineralization was measuredin situ by sequential soil incubations during the year. In the wet area (gravimetric soil moisture content 74–130%), the net N mineralization rates were 4.4 g N m^–2 yr^–1 in the Erica soil and 7.8 g N m^–2 yr^–1 in the Molinia soil. The net nitrification rate was negligibly slow in either soil. In the dry area (gravimetric soil moisture content 7–38%), net N mineralization rates were 6.2 g N M-2 yr^–1 in the Calluna soil, 10.9 g N m^–2 yr^–1 in the Molinia soil and 12.6 g N m^–2 yr^–1 in the Deschampsia soil. The Calluna soil was consistently drier throughout the year, which may partly explain its slower mineralization rate. Net nitrification was 0.3 g N m^–2 yr^–1 in the Calluna soil, 3.6 g N m^–2 yr^–1 in the Molinia soil and 5.4 g N m^–2 yr^–1 in the Deschampsia soil. The net nitrification rate increased proportionally with the net N mineralization rate suggesting ammonium availability may control nitrification rates in these soils. In the dry area, the faster net N mineralization rates in sites dominated by grasses than in the site dominated by Calluna may be explained by the greater amounts of organic N in the soil of sites dominated by grasses. In both areas, however, the net amount of N mineralized per gram total soil N was greater in sites dominated by Molinia or Deschampsia than in sites dominated by Calluna or Erica. This suggests that in heathlands invaded by grasses the quality of the soil organic matter may be increased resulting in more rapid rates of soil N cycling.     

Ecosystem respiration and organic carbon processing in a large,tidally influenced river: the Hudson River

We estimated whole-ecosystem rates of respiration over a 40-km stretch of the tidally influenced freshwater Hudson River every 2 to 3 weeks from May through November. We measured in situ concentrations of oxygen over depth at dusk and dawn at 10 stations spaced over this interval. The use of multiple stations allowed for the consideration of the influence of tidal advection of water masses. Respiration was estimated from the decrease in oxygen overnight with a correction for diffusive exchange of oxygen with the atmosphere. We estimated this flux of oxygen to or from the atmosphere using the measured oxygen gradient and a transfer velocity model which is a function of wind velocity.Integration of the data for the period of May through November yields an estimate of whole-ecosystem respiration of 591 g C m^–2 (S.E. = 66). That the standard error of this estimate is relatively low (11% of the estimate) indicates that the use of multiple stations adequately deals with error introduced through the advection of water between stations. The logarithm of average daily respiration rate was correlated with average daily temperature (p = 0.007;r ² = 0.62). We used this temperature-respiration relationship to derive an estimate of the annual respiration rate of 755 g C m^–2 yr^–1 (S.E. = 72). This estimate is moderately sensitive to the estimated flux of oxygen between the atmosphere and water; using the lower and upper 95% confidence limits of our model relating the transfer velocity of oxygen to wind speed gives a range of annual respiration estimates from 665 g C m^–2 yr^–1 to 984 g C m^–2 yr^–1.The river is strongly heterotrophic, with most respiration driven by allochthonous inputs of organic matter from terrestrial ecosystems. The majority of the allochthonous inputs to the river (over 60%) are apparently metabolized within the river. Any change in allochthonous inputs due to changes in land use or climate patterns would be expected to alter the oxygen dynamics and energy flow within this tidally influenced river.     

Impact of long-term alternations of discharge and spate on the chironomid community in the lowland Widawka River (Central Poland)

Spatial and temporal distribution, abundance and production of macroinvertebrate communities were estimated over two years in a fifth-order section of the Widawka River. Discharge of this river has been increased artificially by coal mine water inputs. Additionally, during the second year, one of the highest discharges of the current 20-year period was recorded. Chironomidae were co-dominant in macrobenthos, both in a straight reach (WIA) and in a meandering site (WIB). More mosaic habitats resulted in higher densities of midges, reaching 6215 ind.m^–2 in year 1 and 1141 ind.m^–2 in year 2 at WIA, while at WIB 896 densities were ind.m^–2 and 257 ind.m^–2, respectively. Flooding affected the distribution and abundance of the chironomid assemblages. Recolonization by psammophilous Polypedilum began after the various microhabitats were buried with sand. Chironomid production was estimated on a species-specific basis for the dominant taxa. In year 1 (mean annual water temperature 10.0° C) chironomid production was 12.4 g dry wt m^–2 yr^–1 1 at WIA and 1.9 g dry wt m^–2 yr^–1 at WIB. These values sharply decreased in year 2 (mean annual water temperature 9.8° C) reaching 1.9 g dry wt m^–2 yr^–1 at WIA and 0.4 g dry wt m^–2 yr^–1 at WIB, as effects of the high spate.     

The effect of sewage-enriched river Ganga water on the biomass production of theAzolla-Anabaena complex

The biomass formation ofAzolla was greatly enhanced by water of the River Ganga and by prevailing environmental conditions. It increased gradually from January to April (first maximum 2.409 g.m^–2.day^–1), declined during June (1.185 g.m^–2.day^–1), and reached a second its maximum during September (2.629 g.m^–2.day^–1). The biomass formation was related to the nutrient availability in the medium in a particular season (measured were: nitrate-N, available phosphorus, total suspended solids, and conductivity). The average annual production of 6.73 ton.ha^–1.yr^–1 is equivalent to the average production of 0.025 ton.ha^–1.yr^–1 phosphorus, 0.252 ton.ha^–1.yr^–1 nitrogen, and 1.57 ton.ha^–1.yr^–1 crude protein.     

Climate‐sensitive ecosystem carbon dynamics along the soil chronosequence of the Damma glacier forefield,Switzerland

We performed a detailed study on the carbon build‐up over the 140‐year‐long chronosequence of the Damma glacier forefield, Switzerland, to gain insights into the organic carbon dynamics during the initial stage of soil formation and ecosystem development. We determined soil carbon and nitrogen contents and their stable isotopic compositions, as well as molecular‐level composition of the bulk soils, and recalcitrance parameters of carbon in different fractions. The chronosequence was divided into three age groups, separated by small end moraines that resulted from two glacier re‐advances. The net ecosystem carbon balance (NECB) showed an exponential increase over the last decades, with mean annual values that range from 100 g C m^?2 yr^?1 in the youngest part to over 300 g C m^?2 yr^?1 in a 60–80 years old part. However, over the entire 140‐year chronosequence, the NECB is only 20 g C m^?2 yr^?1, similar to results of other glacier forefield studies. The difference between the short‐ and long‐term NECB appears to be caused by reductions in ecosystem carbon (EC) accumulation during periods with a colder climate. We propose that two complementary mechanisms have been responsible: 1) Reductions in net primary productivity down to 50% below the long‐term mean, which we estimated using reconstructed effective temperature sums. 2) Disturbance of sites near the terminus of the re‐advanced glacier front. Stabilization of soil organic matter appeared to play only a minor role in the coarse‐grained forefield. We conclude that the forefield ecosystem, especially primary productivity, reacts rapidly to climate changes. The EC gained at warm periods is easily lost again in a cooling climate. Our conclusions may also be valid for other high mountain ecosystems and possibly arctic ecosystems.     

Climatic variability,hydrologic anomaly,and methane emission can turn productive freshwater marshes into net carbon sources

Freshwater marshes are well‐known for their ecological functions in carbon sequestration, but complete carbon budgets that include both methane (CH₄) and lateral carbon fluxes for these ecosystems are rarely available. To the best of our knowledge, this is the first full carbon balance for a freshwater marsh where vertical gaseous [carbon dioxide (CO₂) and CH₄] and lateral hydrologic fluxes (dissolved and particulate organic carbon) have been simultaneously measured for multiple years (2011–2013). Carbon accumulation in the sediments suggested that the marsh was a long‐term carbon sink and accumulated ~96.9 ± 10.3 (±95% CI) g C m^?2 yr^?1 during the last ~50 years. However, abnormal climate conditions in the last 3 years turned the marsh to a source of carbon (42.7 ± 23.4 g C m^?2 yr^?1). Gross ecosystem production and ecosystem respiration were the two largest fluxes in the annual carbon budget. Yet, these two fluxes compensated each other to a large extent and led to the marsh being a CO₂ sink in 2011 (?78.8 ± 33.6 g C m^?2 yr^?1), near CO₂‐neutral in 2012 (29.7 ± 37.2 g C m^?2 yr^?1), and a CO₂ source in 2013 (92.9 ± 28.0 g C m^?2 yr^?1). The CH₄ emission was consistently high with a three‐year average of 50.8 ± 1.0 g C m^?2 yr^?1. Considerable hydrologic carbon flowed laterally both into and out of the marsh (108.3 ± 5.4 and 86.2 ± 10.5 g C m^?2 yr^?1, respectively). In total, hydrologic carbon fluxes contributed ~23 ± 13 g C m^?2 yr^?1 to the three‐year carbon budget. Our findings highlight the importance of lateral hydrologic inflows/outflows in wetland carbon budgets, especially in those characterized by a flow‐through hydrologic regime. In addition, different carbon fluxes responded unequally to climate variability/anomalies and, thus, the total carbon budgets may vary drastically among years.     

Carbon cycling and budget in a forested basin of southwestern Hokkaido,northern Japan

Quantification of annual carbon sequestration is very important in order to assess the function of forest ecosystems in combatting global climate change and the ecosystem responses to those changes. Annual cycling and budget of carbon in a forested basin was investigated to quantify the carbon sequestration of a cool-temperate deciduous forest ecosystem in the Horonai stream basin, Tomakomai Experimental Forest, northern Japan. Net ecosystem exchange, soil respiration, biomass increment, litterfall, soil-solution chemistry, and stream export were observed in the basin from 1999–2001 as a part of IGBP-TEMA project. We found that 258 g C m^–2 year^–1 was sequestered annually as net ecosystem exchange (NEE) in the forested basin. Discharge of carbon to the stream was 4 g C m^–2 year^–1 (about 2% of NEE) and consisted mainly of dissolved inorganic carbon (DIC). About 43% of net ecosystem productivity (NEP) was retained in the vegetation, while about 57% of NEP was sequestered in soil, suggesting that the movement of sequestered carbon from aboveground to belowground vegetation was an important process for net carbon accumulation in soil. The derived organic carbon from aboveground vegetation that moved to the soil mainly accumulated in the solid phase of the soil, with the result that the export of dissolved organic carbon to the stream was smaller than that of dissolved inorganic carbon. Our results indicated that the aboveground and belowground interaction of carbon fluxes was an important process for determining the rate and retention time of the carbon sequestration in a cool-temperate deciduous forest ecosystem in the southwestern part of Hokkaido, northern Japan.     

Phosphorus and nitrogen retention in five Precambrian shield wetlands

Phosphorus and nitrogen mass balances of five wetlands (two beaver ponds, two conifer-Sphagnum swamps and one sedge fen) situated in three catchments in central Ontario, Canada, were measured. Monthly and annual input-output budgets of total phosphorus (TP), total nitrogen (TN), total organic nitrogen (TON), total inorganic nitrogen (TIN), ammonium ion (NH₄ ⁺ -N), nitrate (NO ₃ ^– -N) and dissolved organic carbon (DOC) were estimated for the five wetlands during the 1982–83 and 1983–84 water years. Except for the deepest beaver pond (3.2 m) which had annual TP retention of –44% (–0.030 ± 0.015 g m^–2 yr^–1), the wetlands retained < 0.001 to 0.015 g M^–2 yr^–1 ; however, this wasless than 20% of the inputs and the estimated budget uncertainties were equal to or greater than the retention rates. Annual TN retentions ranged from –0.44 to 0.56 g m^–2 yr^–1 (–12 to 4%) but were not significantly different from zero. The wetlands transformed nitrogen by retaining TIN (16 to 80% RT) and exporting an equivalent amount as TON (–7 to 102% RT). The beaver ponds, however, retained NO ₃ ^– while NH ₄ ⁺ was passed through or the outputs exceeded the inputs. In contrast, the conifer swamps retained both NH ₄ ⁺ and NO ₃ ^– . DOC fluxes into and out of the beaver ponds were equal (–18 and 4% RT) but output from the conifer swamps exceeded input by > 90%. Marked seasonal trends in nutrient retention were observed. Nutrient retention coincided with low stream flow, increased evapotranspiration and biotic uptake during the summer. Net nutrient export occurred during the winter and spring when stream flows were highest and biotic uptake was low.     

Cation distribution,cycling, and removal from mineral soil in Douglas-fir and red alder forests

Overstory species influence the distribution and dynamics of nutrients in forest ecosystems. Ecosystem-level estimates of Ca, Mg, and K pools and cycles in 50-year old Douglas-fir and red alder stands were used to determine the effect of overstory composition on net cation removal from the mineral soil, i.e. cation export from the soil in excess of additions. Net cation removal from Douglas-fir soil was 8 kg Ca ha^–1 yr^–1, 1 kg Mg ha^–1 yr^–1, and 0.3 kg K ha^–1 yr^–1. Annual cation export from soil by uptake and accumulation in live woody tissue and O horizon was of similar magnitude to leaching in soil solution. Atmospheric deposition partially off-set export by adding cations equivalent to 28–88% of cation export. Net cation removal from red alder soil was 58 kg Ca ha^–1 yr^–1, 9 kg Mg ha^–1 yr^–1, and 11 kg K ha^–1 yr^–1. Annual cation accumulation in live woody tissue and O horizon was three times greater than in Douglas-fir, while cation leaching in soil solution was five to eight times greater. The lack of excessive depletion of exchangeable cations in the red alder soil suggests that mineral weathering, rather than exchangeable cations, was the source of most of the removed cations. Nitric acid generated during nitrification in red alder soil led to high rates of weathering and NO₃-driven cation leaching.     

Retention of nutrients in river basins

In Denmark, as in many other European countries, the diffuse losses of nitrogen (N) and phosphorus (P) from the rural landscape are the major causes of surface water eutrophication and groundwater pollution. The export of total N and total P from the Gjern river basin amounted to 18.2 kg ha^–1 and 0.63 kg P ha^–1 during June 1994 to May 1995. Diffuse losses of N and P from agricultural areas were the main nutrient source in the river basin contributing 76% and 51%, respectively, of the total export.Investigations of nutrient cycling in the Gjern river basin have revealed the importance of permanent nutrient sinks (denitrification and overbank sedimentation) and temporary nutrient storage in watercourses. Temporary retention of N and P in the watercourses thus amounted to 7.2–16.1 g N m^–2 yr^–1 and 3.7–8.3 g P m^–2 yr–1 during low-flow periods. Deposition of P on temporarily flooded riparian areas amounted from 0.16 to 6.50 g P m^–2 during single irrigation and overbank flood events, whereas denitrification of nitrate amounted on average to 7.96 kg N yr^–1 per running metre watercourse in a minerotrophic fen and 1.53 kg N yr^–1 per linear metre watercourse in a wet meadow. On average, annual retention of N and P in 18 Danish shallow lakes amounted to 32.5 g N m^–2 yr^–1 and 0.30 g P m^–2 yr^–1, respectively, during the period 1989–1995.The results indicate that permanent nutrient sinks and temporary nutrient storage in river systems represent an important component of river basin nutrient budgets. Model estimates of the natural retention potential of the Gjern river basin revealed an increase from 38.8 to 81.4 tonnes yr^–1 and that P-retention increased from –0.80 to 0.90 tonnes yr^–1 following restoration of the water courses, riparian areas and a shallow lake. Catchment management measures such as nature restoration at the river basin scale can thus help to combat diffuse nutrient pollution.     

Distribution and production ofChironomus salinarius (Diptera: Chironomidae) in a shallow coastal lagoon in the Bay of Cádiz

The abundance, generation time and production ofChironomus salinarius larvae in a lagoon fish-pond system in the Bay of Cádiz were studied by taking monthly samples at 3 sites during 1991 and 1992. Numerical abundance and biomass of larvae showed considerable spatial, seasonal and interannual variation (ANCOVAs,P<0.001). The maximum mean annual density was 7048 larvae m^–2, and corresponded to a biomass of 3.08 g dry weight (DW) m^–2. It was recorded at the site with the lowest rate of water renewal. Seasonal patterns were similar at all sites, with main annual peaks of abundance and biomass in autumn-early winter. Chironomid density was positively related to the biomass of benthic macroalgae (P<0.001). The population studied was multivoltine with a probable average of five generations per year, with overlapping cohorts and a predominance of third- and fourth-instar larvae. Estimates of annual production ranged between 72.2 g DW m^–2 yr^–1 at the site with the lowest rate of water renewal in 1991 and 0.1 g DW m^–2 yr^–1 at the site with the highest rate of water renewal in 1992. Mean annual production and the production/biomass ratio for the system was estimated to be 16.8 g DW m^–2 yr^–1 and 12.7, respectively. Possible factors leading to the observed density fluctuations are discussed, as well as possible sources of error in production estimates.     

Forest floor mass,litterfall and nutrient return in Central Himalayan high altitude forests

Dynamics of forest floor biomass, pattern of litter fall and nutrient return in three central Himalayan high elevation forests are described. Fresh and partially decomposed litter layer occur throughout the year. In maple and birch the highest leaf litter value was found in October and in low-rhododendron in August. The relative contribution of partially and more decomposed litter to the total forest floor remains greatest the year round. The total calculated input of litter was 627.7 g m^-2 yr^-1 for maple, 477.87 g m^-2 yr^-1 for birch and 345.9 g m^-2 yr^-1 for low-rhododendron forests. 49–61% of the forest floor was replaced per year with a subsequent turnover time of 1.6–2.0 yr. The annual nutrient return through litter fall amounted to (kg ha^-1 yr^-1) 25.5–56.1 N, 2.0–5.4 P and 9.9–23.3 K. The tree litter showed an annual replacement of 26–54% for different nutrients and it decreased towards higher elevation. The nutrient use efficiency in terms of litter produced per unit of nutrient was higher in present study compared to certain mid- and high-elevation forests of the central Himalaya.     

A historic perspective on Wadden Sea eutrophication

In this paper, a reconstruction of the pre-industrial trophic status of the Wadden Sea is presented. A conceptual model is outlined that links the organic matter and nutrient dynamics in the Wadden Sea with riverine nutrient input. Fundamental processes in this model are: a nutrient-limited offshore primary production and the subsequent import of primary produced organic matter from the North Sea into the Wadden Sea. Two approaches have been followed to estimate the production and remineralisation levels under pre-industrial conditions. The first approach is based on present-day relationships between the seasonal cycle of NH₄ and NO₂ in the western Dutch Wadden Sea and suggests, on average, sixfold lower production and remineralisation rates under pre-industrial conditions (range: four to eight times). The second approach is based on present carbon budgets extrapolated to pre-industrial budgets on the basis of present relationships between winter nutrient concentrations, annual primary production and annual organic matter turnover rates, and suggests a fivefold lower organic matter turnover under pre-industrial conditions (annual primary production: ~55 g C m^–2 year^–1, annual remineralisation: ~77 g C m^–2 year^–1). Better pre-industrial light conditions in the Wadden Sea may have allowed a more efficient use of nutrients, a higher annual primary production of about 86 g C m^–2 year^–1 and annual remineralisation rates of about 108 g C m^–2 year^–1.     

Sensitivity of Temperate Desert Steppe Carbon Exchange to Seasonal Droughts and Precipitation Variations in Inner Mongolia,China

Arid grassland ecosystems have significant interannual variation in carbon exchange; however, it is unclear how environmental factors influence carbon exchange in different hydrological years. In this study, the eddy covariance technique was used to investigate the seasonal and interannual variability of CO₂ flux over a temperate desert steppe in Inner Mongolia, China from 2008 to 2010. The amounts and times of precipitation varied significantly throughout the study period. The precipitation in 2009 (186.4 mm) was close to the long-term average (183.9±47.6 mm), while the precipitation in 2008 (136.3 mm) and 2010 (141.3 mm) was approximately a quarter below the long-term average. The temperate desert steppe showed carbon neutrality for atmospheric CO₂ throughout the study period, with a net ecosystem carbon dioxide exchange (NEE) of −7.2, −22.9, and 26.0 g C m⁻² yr⁻¹ in 2008, 2009, and 2010, not significantly different from zero. The ecosystem gained more carbon in 2009 compared to other two relatively dry years, while there was significant difference in carbon uptake between 2008 and 2010, although both years recorded similar annual precipitation. The results suggest that summer precipitation is a key factor determining annual NEE. The apparent quantum yield and saturation value of NEE (NEE_sat) and the temperature sensitivity coefficient of ecosystem respiration (R_eco) exhibited significant variations. The values of NEE_sat were −2.6, −2.9, and −1.4 µmol CO₂ m⁻² s⁻¹ in 2008, 2009, and 2010, respectively. Drought suppressed both the gross primary production (GPP) and R_eco, and the drought sensitivity of GPP was greater than that of R_eco. The soil water content sensitivity of GPP was high during the dry year of 2008 with limited soil moisture availability. Our results suggest the carbon balance of this temperate desert steppe was not only sensitive to total annual precipitation, but also to its seasonal distribution.