Changes in topsoil carbon stock in the Tibetan grasslands between the 1980s and 2004

Climate warming is likely inducing carbon loss from soils of northern ecosystems, but little evidence comes from large-scale observations. Here we used data from a repeated soil survey and remote sensing vegetation index to explore changes in soil organic carbon (SOC) stock on the Tibetan Plateau during the past two decades. Our results showed that SOC stock in the top 30 cm depth in alpine grasslands on the plateau amounted to 4.4 Pg C (1 Pg=10¹⁵ g), with an overall average of 3.9 kg C m⁻². SOC changes during 1980s–2004 were estimated at −0.6 g C m⁻² yr⁻¹, ranging from −36.5 to 35.8 g C m⁻² yr⁻¹ at 95% confidence, indicating that SOC stock in the Tibetan alpine grasslands remained relatively stable over the sampling periods. Our findings are nonconsistent with previous reports of loss of soil C in grassland ecosystems due to the accelerated decomposition with warming. In the case of the alpine grasslands on the Tibetan Plateau studied here, we speculate that increased rates of decomposition as soils warmed during the last two decades may have been compensated by increased soil C inputs due to increased grass productivity. These results suggest that soil C stock in terrestrial ecosystems may respond differently to climate change depending on ecosystem type, regional climate pattern, and intensity of human disturbance.     

Change in CO2 balance under a series of forestry activities in a cool-temperate mixed forest with dense undergrowth

To evaluate the effects on CO₂ exchange of clearcutting a mixed forest and replacing it with a plantation, 4.5 years of continuous eddy covariance measurements of CO₂ fluxes and soil respiration measurements were conducted in a conifer-broadleaf mixed forest in Hokkaido, Japan. The mixed forest was a weak carbon sink (net ecosystem exchange, −44 g C m⁻² yr⁻¹), and it became a large carbon source (569 g C m⁻² yr⁻¹) after clearcutting. However, the large emission in the harvest year rapidly decreased in the following 2 years (495 and 153 g C m⁻² yr⁻¹, respectively) as the gross primary production (GPP) increased, while the total ecosystem respiration (RE) remained relatively stable. The rapid increase in GPP was attributed to an increase in biomass and photosynthetic activity of Sasa dwarf bamboo, an understory species. Soil respiration increased in the 3 years following clearcutting, in the first year mainly owing to the change in the gap ratio of the forest, and in the following years because of increased root respiration by the bamboo. The ratio of soil respiration to RE increased from 44% in the forest to nearly 100% after clearcutting, and aboveground parts of the vegetation contributed little to the RE although the respiration chamber measurements showed heterogeneous soil condition after clearcutting.     

Amazonian peatlands: an ignored C sink and potential source

In tropical lowlands, peatlands are commonly reported from Southeast Asia, and especially Indonesian tropical peatlands are known as considerable C sinks and sources. In contrast, Amazonia has been clearly understudied in this context. In this study, based on field observations from 17 wetland sites in Peruvian lowland Amazonia, we report 0–5.9 m thick peat deposits from 16 sites. Only one of the studied sites did not contain any kind of peat deposit (considering pure peat and clayey peat). Historic yearly peat and C accumulation rates, based on radiocarbon dating of peat samples from five sites, varied from 0.94 ± 0.99 to 4.88 ± 1.65 mm, and from 26 ± 3 to 195 ± 70 g C m⁻², respectively. The long-term apparent peat and C accumulation rates varied from 1.69 ± 0.03 to 2.56 ± 0.12 mm yr⁻¹, and from 39 ± 10 to 85 ± 30 g C m⁻² yr⁻¹, respectively. These accumulation rates are comparable to those determined in the Indonesian tropical peatlands. Under altered conditions, Indonesian peatlands can release globally relevant amounts of C to the atmosphere. Considering the estimated total area of Amazonian peatlands (150 000 km²) close to that of the Indonesian ones (200 728 km²) as well as several factors threatening the Amazonian peatlands, we suggest that the total C stocks and fluxes associated with Amazonian peatlands may be of global significance.     

Near-zero recent carbon accumulation in a bog with high nitrogen deposition in SW Sweden

We present data on the accumulation of carbon and nitrogen into an open oceanic ombrotrophic bog, SW Sweden, with high levels of anthropogenic nitrogen deposition. The aim was to investigate if this peatland currently acts as a sink for atmospheric carbon. Peat cores were sampled from the top peat layer in five different vegetation types. Small pines were used to date the cores. The cores bulk density and carbon and nitrogen content were determined. A vegetation-classified satellite image was used to estimate the areal extent of the vegetation types and to scale up these results to bog level. The rate of current carbon input into the upper oxic acrotelm was 290 g m⁻² yr⁻¹, and there were no significant differences in accumulation rates among the vegetation types. This organic matter input to the acrotelm was almost completely decomposed before it was deposited for storage in the deeper peat layers (the catotelm) and only a small fraction (≪1%) or 0.012 g m⁻² yr⁻¹ of the carbon would be left, assuming a residence time of 100 years in the acrotelm. Nitrogen accumulation rates differed between the vegetation classes, and the average input via primary production varied from 5.33 to 16.8 g m⁻² yr⁻¹. Current nitrogen input rates into the catotelm are much lower, 0–0.059 g m⁻² yr⁻¹, with the highest accumulation rates in lawn-dominated communities. We suggest that one of the main causes of the low carbon input rates is the high level of nitrogen deposition, which enhances decomposition and changes the vegetation from peat-forming Sphagnum -dominance to dominance by dwarf shrubs and graminoids.     

Simulated chronic nitrogen deposition increases carbon storage in Northern Temperate forests

High levels of atmospheric nitrogen (N) deposition in Europe and North America were maintained throughout the 1990s, and global N deposition is expected to increase by a factor of 2.5 over the next century. Available soil N limits primary production in many terrestrial ecosystems, and some computer simulation models have predicted that increasing atmospheric N deposition may result in greater terrestrial carbon (C) storage in woody biomass. However, empirical evidence demonstrating widespread increases in woody biomass C storage due to atmospheric N deposition is uncommon. Increased C storage in soil organic matter due to chronic N inputs has rarely been reported and is often not considered in computer simulation models of N deposition effects. Since 1994, we have experimentally simulated chronic N deposition by adding 3 g N m⁻² yr⁻¹ to four different northern hardwood forests, which span a 500 km geographic gradient in Michigan. Each year we measured tree growth. In 2004, we also examined soil C content to a depth of 70 cm. When we compared the control treatment with the NO₃⁻ deposition treatment after a decade of experimentation, ecosystem C storage had significantly increased in both woody biomass (500 g C m⁻²) and surface soil (0–10 cm) organic matter (690 g C m⁻²). The increase in surface soil C storage was apparently driven by altered rates of organic matter decomposition, rather than an increase in detrital inputs to soil. Our results, for study locations stretching across hundreds of kilometers, support the hypothesis that chronic N deposition may increase C storage in northern forests, potentially contributing to a sink for anthropogenic CO₂ in the northern Hemisphere.     

Changes in organic carbon distribution with depth in agricultural soils in northern Belgium, 1960–2006

In most studies concerning the carbon (C) exchange between soil and atmosphere only the topsoil (0–0.3 m) is taken into account. However, it has been shown that important amounts of stable soil organic carbon (SOC) are also stored at greater depth. Here, we developed a quantitative model to estimate the evolution of the distribution of SOC with depth between 1960 (database 'Aardewerk') and 2006 in northern Belgium. This temporal analysis was conducted under different land use, texture and drainage conditions. The results indicate that intensified land management practices seriously affect the SOC status of the soil. The increase in plough depth and a change in crop rotation result in a significant decrease of C near the surface for dry silt loam cropland soils, (i.e. 1.02 ± 0.23 kg C m⁻² in the top 0.3 m between 1960 and 2006). In wet to extremely wet grasslands, topsoil SOC decreased significantly, indicating a negative influence of intensive soil drainage on SOC stock. This resulted in a decline of SOC between 1960 and 2006 in the top 1 m, ranging from 3.99 ± 2.57 kg C m⁻² in extremely wet silt loam soils to 2.04 ± 2.08 kg C m⁻² in wet sandy soils. A slight increase of SOC stock is observed under dry to moderately wet grasslands at greater depths corresponding to increased livestock densities in the region. The increase of SOC in the top 1 m under grassland ranges from 0.65 ± 1.39 kg C m⁻² in well drained silt loam soils to 2.59 ± 6.49 kg C m⁻² in moderately drained silt loam soils over entire period.     

Population dynamics and production of Jesogammarus annandalei, an endemic amphipod, in Lake Biwa, Japan

SUMMARY 1. Population dynamics and production of Jesogammarus annandalei , an endemic amphipod in Lake Biwa, were examined from April 1997 to June 1998. The life cycle of this species was 1 year with the new generation beginning in early autumn. They preferred low temperature (<12 °C) and their spatial distribution varied seasonally and accordingly.
2. In deep water, the abundance of J. annandalei ranged from 200 to 63 000 m⁻² and decreased towards summer and the biomass (0.01∼3.6 g C m⁻²) was on average comparable that of zooplankton. The density was much higher than that recorded by a study conducted 35 years ago.
3. Individual growth rate of this amphipod was high in winter and spring but decreased in summer. Annual production of J. annandalei (6.2 g C m⁻² year⁻¹) was only 2% of primary production but was at the higher end of the range reported for amphipods in oligo- and mesotrophic lakes.
4. These results are consistent with the view that Lake Biwa is becoming more eutrophic, with a consequent decrease in the abundance of predatory fish in the profundal zone.     

Temporal dynamics in epipelic, pelagic and epiphytic algal production in a clear and a turbid shallow lake

SUMMARY 1. Pelagic and epipelic microalgal production were measured over a year in a pre-defined area (depth 0.5 m) in each of two lakes, one turbid and one with clear water. Further estimates of epiphytic production within reed stands were obtained by measuring production of periphyton developed on artificial substrata.
2. Total annual production of phytoplankton and epipelon was 34% greater in the turbid lake (190 g C m⁻² year⁻¹) than in the clearwater lake (141 g C m⁻² year⁻¹). However, the ratio of total production to mean water column TP concentration was two fold greater in the clearwater lake.
3. Phytoplankton accounted for the majority of the annual production (96%) in the turbid lake, while epipelic microalgal production dominated (77%) in the clear lake. The relative contribution of epipelic algae varied over the year, however, and in the turbid lake was higher in winter (11–25%), when the water was relatively clear, than during summer (0.7–1.7%), when the water was more turbid. In the clearwater lake, the relative contribution of epipelon was high both in winter, when the water was most clear, and in mid-summer, when phytoplankton production was constrained either by nutrients or grazing.
4. Compared with pelagic and epipelic primary production, epiphytic production within a reed stand was low and did not vary significantly between the lakes.
5. The study supports the theory of a competitive and compensatory trade-off between primary producers in lakes with contrasting nutrient concentrations, resulting in relatively small differences in overall production between clear and turbid lakes when integrating over the season and over different habitats.     

Climate-induced changes in high elevation stream nitrate dynamics

Mountain terrestrial and aquatic ecosystems are responsive to external drivers of change, especially climate change and atmospheric deposition of nitrogen (N). We explored the consequences of a temperature-warming trend on stream nitrate in an alpine and subalpine watershed in the Colorado Front Range that has long been the recipient of elevated atmospheric N deposition. Mean annual stream nitrate concentrations since 2000 are higher by 50% than an earlier monitoring period of 1991–1999. Mean annual N export increased by 28% from 2.03 kg N ha⁻¹ yr⁻¹ before 2000 to 2.84 kg N ha⁻¹ yr⁻¹ in Loch Vale watershed since 2000. The substantial increase in N export comes as a surprise, since mean wet atmospheric N deposition from 1991 to 2006 (3.06 kg N ha⁻¹ yr⁻¹) did not increase. There has been a period of below average precipitation from 2000 to 2006 and a steady increase in summer and fall temperatures of 0.12 °C yr⁻¹ in both seasons since 1991. Nitrate concentrations, as well as the weathering products calcium and sulfate, were higher for the period 2000–2006 in rock glacier meltwater at the top of the watershed above the influence of alpine and subalpine vegetation and soils. We conclude the observed recent N increases in Loch Vale are the result of warmer summer and fall mean temperatures that are melting ice in glaciers and rock glaciers. This, in turn, has exposed sediments from which N produced by nitrification can be flushed. We suggest a water quality threshold may have been crossed around 2000. The phenomenon observed in Loch Vale may be indicative of N release from ice features such as rock glaciers worldwide as mountain glaciers retreat.     

Nitrogen effects on net ecosystem carbon exchange in a temperate steppe

It has widely been documented that nitrogen (N) enrichment stimulates plant growth and net primary production. However, there is still dispute on how N addition affects net ecosystem CO₂ exchange (NEE), which represents the balance between ecosystem carbon (C) uptake and release. We conducted an experimental study to examine effects of N addition on NEE in a temperate steppe in northern China from 2005 to 2008. N was added at a rate of 10 g N m⁻² yr⁻¹ with NH₄NO₃ alone or in combination with phosphorous (P, 5 g P₂O₅ m⁻² yr⁻¹) in both clipped and unclipped plots. Over the 4 years, N addition significantly stimulated growing-season NEE, on average, by 27%. Neither the main effects of P addition or clipping nor their interactions with N addition were statistically significant on NEE in any of the 4 years. However, the magnitude of N stimulation on NEE declined over time. N addition significantly increased NEE by 60% in 2005 and 21% in 2006, but its effect was not significant in 2007 and 2008. N-induced shift in species composition was primarily responsible for the declined N stimulation over time. The gradually increasing coverage of the upper canopy species ( Stipa krylovii ) and standing litter accumulation induced light limitation on the lower canopy species ( Artemisia frigida ). Thus, N-induced shifts in plant species composition strongly regulated the direct effects of N addition on C sequestration in the temperate steppe.     

Storms can cause Europe-wide reduction in forest carbon sink

Disturbance of ecosystems is a major factor in regional carbon budgets, and it is believed to be partly responsible for the large inter-annual variability of the terrestrial part of the carbon balance. Forest fires have so far been considered as the most important disturbance but also other forms of disturbance such as insect outbreaks or wind-throw might contribute significantly to the largely unexplained inter-annual variability, at least in specific regions. The effect of wind-throw has not yet been estimated because of lack of data on how carbon fluxes are affected. The Gudrun storm, which hit Sweden in January 2005, resulted in ca. 66 million m³ of wind-thrown stem wood on an area of ca. 272 000 ha. Using a model (BIOME-BGC) calibrated to CO₂ flux measurements at two sites, the annual net ecosystem productivity during the first year after the storm was estimated to be in the range −897 to −1259 g C m⁻² yr⁻¹. This is a much higher loss compared with harvested (clear-cut) forests in Europe, which ranged between ca. −420 and −100 g m⁻² yr⁻¹. The reduction in the carbon sink scaled to the whole wind-thrown area was estimated at ca. 3 million tons C during the first year. By historical data on wind-throw in Europe combined with modelling, we estimated that the large Lothar storm in 1999 reduced the European carbon balance by ca. 16 million tons C, this is ca. 30% of the net biome production in Europe. We conclude that the impact of increased forest damage by more frequent storms in future climate change scenarios must be considered and that intermittent large wind-throw events may explain a part of the large inter-annual variability in the terrestrial carbon sink.     

Electrical signalling and cytokinins mediate effects of light and root cutting on ion uptake in intact plants

Nutrient acquisition in the mature root zone is under systemic control by the shoot and the root tip. In maize, exposure of the shoot to light induces short-term (within 1–2 min) effects on net K⁺ and H⁺ transport at the root surface. H⁺ efflux decreased (from −18 to −12 nmol m⁻² s⁻¹) and K⁺ uptake (∼2 nmol m⁻² s⁻¹) reverted to efflux (∼−3 nmol m⁻² s⁻¹). Xylem probing revealed that the trans-root (electrical) potential drop between xylem vessels and an external electrode responded within seconds to a stepwise increase in light intensity; xylem pressure started to decrease after a ∼3 min delay, favouring electrical as opposed to hydraulic signalling. Cutting of maize and barley roots at the base reduced H⁺ efflux and stopped K⁺ influx in low-salt medium; xylem pressure rapidly increased to atmospheric levels. With 100 m m NaCl added to the bath, the pressure jump upon cutting was more dramatic, but fluxes remained unaffected, providing further evidence against hydraulic regulation of ion uptake. Following excision of the apical part of barley roots, influx changed to large efflux (−50 nmol m⁻² s⁻¹). Kinetin (2–4  µ m ), a synthetic cytokinin, reversed this effect. Regulation of ion transport by root-tip-synthesized cytokinins is discussed.     

Carbon stored in human settlements: the conterminous United States

Urban areas are home to more than half of the world's people, responsible for >70% of anthropogenic release of carbon dioxide and 76% of wood used for industrial purposes. By 2050 the proportion of the urban population is expected to increase to 70% worldwide. Despite fast rates of change and potential value for mitigation of carbon dioxide emissions, the organic carbon storage in human settlements has not been well quantified. Here, we show that human settlements can store as much carbon per unit area (23–42 kg C m⁻² urban areas and 7–16 kg C m⁻²exurban areas) as tropical forests, which have the highest carbon density of natural ecosystems (4–25 kg C m⁻²). By the year 2000 carbon storage attributed to human settlements of the conterminous United States was 18 Pg of carbon or 10% of its total land carbon storage. Sixty-four percent of this carbon was attributed to soil, 20% to vegetation, 11% to landfills, and 5% to buildings. To offset rising urban emissions of carbon, regional and national governments should consider how to protect or even to increase carbon storage of human-dominated landscapes. Rigorous studies addressing carbon budgets of human settlements and vulnerability of their carbon storage are needed.     

Does nitrogen deposition affect soil microfungal diversity and soil N and P dynamics in a high Arctic ecosystem?

In a high Arctic polar semidesert ecosystem (ambient N deposition c. 0.1 g N m⁻² a⁻¹), the effects of N enrichment on the diversity of soil microfungi and on N content and availability in organic and mineral soils were determined. Three N (total: 0, 0.5, 5 g N m⁻² a⁻¹) and two P (total 0, 1 g m⁻² a⁻¹) treatments were applied, since P may limit response to N in this ecosystem. Organic and mineral soils were sampled in June and August in the second year of treatment for microfungi, pH, moisture content, and total N and P. In the third year, soils were resampled for extractable and total N and P. The fungi isolated were typical of high pH soils in the High Arctic and Antarctic. The species richness and diversity of soil microfungi were very low, with ranges as follows: Shannon diversity, 0.56–1.5; richness, 2–6; evenness, 0.79–0.9. There was no significant effect of treatment on the frequency of occurrence of different taxa of soil microfungi. Time of sampling also had no significant impact on fungal assemblages, although different, more diverse communities were isolated from organic, rather than mineral, soils. Nitrate-N in organic soil decreased significantly when P was added alone, but not when P and N were added together. Addition of 0.5 g N m⁻² a⁻¹, a rate deposition already occurring in Greenland and Iceland, appeared to exceed N demand even when P limitation was relieved. There was no apparent soil acidification as a result of the N treatments.     

Phytoplankton and pelagic primary productivity in Øvre Heimdalsvatn

A phytoplankton investigation was carried out in the subalpine, low-productive Norwegian lake Øvre Heimdalsvatn in 1969–70 and 1972. This paper describes the temporal and spatial distribution of the standing stock of phytoplankton, and phytoplankton primary productivity. The annual average primary productivity in 1972 was 4.0–4.9 mg C m⁻³ d⁻¹; the annual average standing stock varied from 120 mg m⁻³ (freshweight) in 1969–70, to 250 mg m⁻³ in 1972. Phytoplankton species composition and size distribution is discussed. Throughout the year the phytoplankton is dominated by small (ultraplankton) species; μ-algae (< 5 μm) showed cell concentrations up to 15 mill. cells 1⁻¹. The dominating group was chrysophytes; cryptophytes, dinoflagellates or green algae were at times abundant. A phytoplankton monthly budget and a diagram showing annual average carbon flow through the standing stock of phytoplankton are presented; the phytoplankton dynamics in Øvre Heimdalsvatn is compared to that of other low-productive lakes.     

Artificial drainage and associated carbon fluxes (CO2/CH4) in a tundra ecosystem

Ecosystem flux measurements using the eddy covariance (EC) technique were undertaken in 4 subsequent years during summer for a total of 562 days in an arctic wet tundra ecosystem, located near Cherskii, Far-Eastern Federal District, Russia. Methane (CH₄) emissions were measured using permanent chambers. The experimental field is characterized by late thawing of permafrost soils in June and periodic spring floods. A stagnant water table below the grass canopy is fed by melting of the active layer of permafrost and by flood water. Following 3 years of EC measurements, the site was drained by building a 3 m wide drainage channel surrounding the EC tower to examine possible future effects of global change on the tundra tussock ecosystem. Cumulative summertime net carbon fluxes before experimental alteration were estimated to be about +15 g C m⁻² (i.e. an ecosystem C loss) and +8 g C m⁻² after draining the study site. When taking CH₄ as another important greenhouse gas into account and considering the global warming potential (GWP) of CH₄ vs. CO₂, the ecosystem had a positive GWP during all summers. However CH₄ emissions after drainage decreased significantly and therefore the carbon related greenhouse gas flux was much smaller than beforehand (475 ± 253 g C-CO₂-e m⁻² before drainage in 2003 vs. 23 ± 26 g C-CO₂-e m⁻² after drainage in 2005).     

The effect of an emergent macrophyte (Typha angustifolia) on sediment resuspension in a shallow north temperate lake

1. The effects of emergent macrophytes on water turbidity and sediment resuspension in the shallow Kirkkojärvi basin of Lake Hiidenvesi were studied with sediment traps, and concomitant sediment and water samples. The study was conducted during May–August in three different zones of a stand of the emergent Typha angustifolia .
2. Within the stand (5 m from the edge), both the concentration of suspended solids and the rate of sediment resuspension were significantly lower than at the edge and outside the stand (5 m from the edge). The differences between the zones increased towards the end of summer together with the growing stem density. During the study period (82 days), 2210 g dw m⁻² of sediment was resuspended in the outer zone. At the edge and in the inner zone, the corresponding numbers were 1414 and 858 g dw m⁻², respectively.
3. With the resuspended sediment, 39.4 mg P  m⁻² day⁻¹ was brought to the water column outside the stand, 22.4 mg P  m⁻² day⁻¹ at the edge and 13.4 mg P  m⁻² day⁻¹ within the stand.
4. In early summer, the concentration of suspended solids had a highly significant positive effect on soluble reactive phosphorus (SRP) concentration in the water, whereas in late summer no effect was found. During the study period, phosphorus retention by emergent macrophyte stands corresponded to 3–5% of the present annual external phosphorus loading of the Kirkkojärvi basin.     

High organic loading influences the physical characteristics of aerobic sludge granules

Aims:  The effect of high organic loading rate (OLR) on the physical characteristics of aerobic granules was studied.
Methods and Results:  Two column-type sequential aerobic sludge blanket reactors were fed with either glucose or acetate as the main carbon source, and the OLR was gradually raised from 6 to 9, 12 and 15 kg chemical oxygen demand (COD) m⁻³ d⁻¹. Glucose-fed granules could sustain the maximum OLR tested. At a low OLR, these granules exhibited a loose fluffy morphology dominated by filamentous bacteria. At higher OLRs, these granules became irregularly shaped, with folds, crevices and depressions. In contrast, acetate-fed granules had a compact spherical morphology at OLRs of 6 and 9 kg COD m⁻³ d⁻¹, with better settling and strength characteristics than glucose-fed granules at similar OLRs. However, acetate-fed granules could not sustain high OLRs and disintegrated when the OLR reached 9 kg COD m⁻³ d⁻¹.
Conclusions:  The compact regular microstructure of the acetate-fed granules appeared to limit mass transfer of nutrients at an OLR of 9 kg COD m⁻³ d⁻¹. The looser filamentous microstructure of the glucose-fed granules and the subsequent irregular morphology delayed the onset of diffusion limitation and allowed significantly higher OLRs to be attained.
Significance and Impact of the Study:  High organic loading rates are possible with aerobic granules. This research would be helpful in the development of aerobic granule-based systems for high-strength wastewaters.     

Abrupt change in primary productivity in a littoral zone of Lake Biwa with the development of a filamentous green-algal community

1. Some characteristics of the photosynthesis and primary production of benthic and planktonic algal communities were investigated in a littoral zone covered with gravel in the north basin of Lake Biwa, paying special attention to the recent development of filamentous green algae (FGA) in the benthic algal community.
2. P_max (maximum gross photosynthesis rate) values of the benthic algal community (0.1–1.2 mg C mg chl. a ⁻¹ h⁻¹) obtained from photosynthesis–irradiance (P–I) curves were lower than those of the planktonic algal community (2.4–11.5 mg C mg chl. a ⁻¹ h⁻¹). This is apparently a result of the high degree of self shading in the benthic algal community and its low turnover as compared with that of the planktonic algal community.
3. Relatively high I_k values (150–200 μmol photon m⁻² s⁻¹) were observed in the benthic algal community only in June–July when a FGA, Spirogyra sp., was abundant. This reflected a photosynthetic characteristic of the Spirogyra itself, in which photosynthesis was saturated at high light intensity.
4. The FGA community established in the layer between planktonic and sessile (benthic algae except for FGA) algal communities. It brought about extraordinarily high organic matter production in the littoral zone at the expense of production in the sessile algal community.     

Spatial and temporal patterns of microcrustacean assemblage structure and secondary production in a wetland ecosystem

1. In contrast to extensive studies of zooplankton in lakes, the role of microcrustaceans in wetlands is not well studied. In this study, spatial and temporal patterns of microcrustacean assemblage structure and secondary production were quantified over a 2-year period in a southeastern U.S.A. wetland.
2. Thirty-two species, including 19 cladocerans, 10 copepods and three ostracods, generated different temporal patterns of density and production between vegetated ( Nymphaea ) and non-vegetated (open-water) zones reflecting species-specific differences in life histories.
3. Summer assemblages were dominated by small, planktonic filter-feeders, typified by high annual production/biomass ( P / B ) and daily production. In contrast, winter assemblages were dominated by larger, epibenthic detritivores with low P / B and high biomass. Seasonal shifts in the relative importance of planktonic species in the warmer months to benthic and epiphytic species in the cooler months suggest that energy flow pathways through microcrustaceans may vary seasonally.
4. Total annual production was higher during both years in the Nymphaea zone (13.0 g and 13.6 g DM m⁻² year⁻¹) than the open-water (8.2 and 6.3 g DM m⁻² year⁻¹), and was similar between years for the entire wetland pond (12.3 and 12.2 g DM m⁻² year⁻¹).
5. Although wetland ecosystems have been the subject of considerable ecological research in the past 20 years, our study is one of the few to demonstrate a highly diverse and relatively productive microcrustacean assemblage. Such comprehensive production studies can be used to quantify the ecological importance of microcrustaceans in freshwater wetland ecosystems.