Effects of spring flood and water level draw-down on methane dynamics in the littoral zone of boreal lakes

1. The annual dynamics of methane (CH₄) in a temporarily flooded meadow, mire bank, lacustrine sedge fen, temporarily and continuously inundated sedge ( Carex sp.) and reed ( Phragmites australis ) marshes were studied from June to November in the humic mesoeutrophic Lake Mekrijärvi and in eutrophicated parts of the mesotrophic Lake Heposelkä in the southern part of East Finland. The effects of water level and temperature on littoral CH₄ fluxes were determined. Vegetation zonation along the moisture gradient, and associated CH₄ fluxes, were evaluated.
2. The CH₄ flux increased along the moisture gradient from –0.2 to 14.2 mg CH₄ m^–2 h^–1, and was highest in the permanently inundated marshes. The duration of anoxia in the sediment caused differences in the CH₄ flux. Estimated emissions for the period 1 June – 30 September in continuously inundated sparse reed and sedge marshes, drying sedge marsh, and lacustrine sedge fen were 13, 11 and 6 g CH₄ m^–2, respectively.
3. In continuously inundated vegetation, the fluxes were highest in late July/early August. The seasonal CH₄ flux pattern suggested that the fluxes were regulated by the supply of organic matter during the course of the summer and the water level. In the temporarily flooded zone, the seasonal CH₄ flux dynamics was greatly affected by changes in the lake water level, the fluxes being highest during the spring flood in early June.     

Variation in tissue element concentration in mire plants over a range of sites

Elements in surface peat and ground water were estimated on seven mire sites in central and north Sweden. The variation in tissue clement concentration was studied in eleven plants occurring on the studied sites. The elements estimated were: N, P, K. Ca, Mg, Fe, Mn, Zn, Al, Cu, Mo, Na, S, B, and Si. The species studied were; Carex diandra, Carex limosa, Carex rostrata, Epilobium palustre, Menyanthes trifoliate, Pedicularis palustris, Rumex acetosa, Saxifraga hirculus, Scirpus hudsonianus, Stellaria crassifolia and Tofteldia pusilla . In surface peat over the range of sites, the variation in total concentrations of nitrogen and phosphorus was six and tenfold, respectively. The variation in total concentration of potassium was four-fold. Extractable concentrations of these three elements showed an even greater variation between sites. Over the range of sites, the variation in tissue element concentration was larger in roots than in above-ground parls. In above-ground parts, floral parts varied less in tissue element concentration than did stems and leaves. The largest variation in tissue element concentration was found in Carex rostrata and Menyanthes trifoliata . The smallest variation was found in Saxifraga hirculus . Tissues of the rich fen species Saxifraga hirculus and Stellaria crassifolia had the lowest concentrations of heavy metals. The correlation between substrate element concentration and tissue concentration differed both between tissues and species. Concentrations of N, P and K in tissues of the Carex species were highly correlated with the concentrations of these elements in the substrate, while N, P and K concentrations in tissues of the herbaceous species, especially Saxifraga hirculus . correlated less well.     

In search of a hydrological explanation for vegetation changes along a fen gradient in the Biebrza Upper Basin (Poland)

The understanding of succession from rich fen to poorer fen types requires knowledge of changes in hydrology, water composition, peat chemistry and peat accumulation in the successional process. Water flow patterns, water levels and water chemistry, mineralisation rates and nutrient concentrations in above-ground vegetation were studied along a extreme-rich fen-moderate-rich fen gradient at Biebrza (Poland). The extreme-rich fen was a temporary groundwater discharge area, while in the moderate-rich fen groundwater flows laterally towards the river. The moderate-rich fen has a rainwater lens in spring and significant lower concentrations of calcium and higher concentrations of phosphate in the surface water. Mineralisation rates for N, P and K were higher in the moderate-rich fen. Phosphorus concentrations in plant material of the moderate-rich fen were higher than in the extreme-rich fen, but concentrations of N and K in plant material did not differ between both fen types. Water level dynamics and macro-remains of superficial peat deposits were similar in both fen types.We concluded that the differences observed in the moderate-rich and the extreme-rich fens were caused by subtile differences in the proportion of water sources at the peat surface (rainwater and calcareous groundwater, respectively). Development of an extreme-rich fen into a moderate-rich fen was ascribed to recent changes in river hydrology possibly associated with a change in management practices. The observed differences in P-availability between the fen types did not result in significantly different biomass. Moreover, biomass production in both fen types was primarily N-limited although P-availability was restricted too in the extreme-rich fen. Aulacomnium palustre, the dominant moss in the moderate-rich fen, might be favoured in competition because of its broad nutrient tolerance and its quick establishment after disturbance. It might outcompete low productive rich fen species which were shown to be N-limited in both fens. We present a conceptual model of successional pathways of rich fen vegetation in the Biebrza region.     

Hydro-ecological analysis of the Biebrza mire (Poland)

Vegetation composition and structure of 58 sites along gradients in the valley mire of Biebrza, Poland, are related to physical and chemical variables of groundwater and peat. The three most prominent hydrochemical processes in the valley are (a) dissolution of calcite; (b) dissolution of iron, manganese and aluminium; and (c) enrichment with nitrogen and potassium. Major factors determining these processes are vertical flow of the groundwater and river flooding.Within the rheophilous zone of the mire, calcium-richness of the shallow groundwater and base-saturation of the peat are caused by upward seepage of groundwater originating from adjacent higher grounds. This groundwater movement keeps the larger part of the mire saturated with calcium.Good correlations exist between hydrochemistry and vegetation patterns. Groundwater-fed sites support a characteristic rich fen vegetation (Caricetum limoso-diandrae) with a low biomass production. The flood-plain vegetation consists of highly-productive communities of Glycerietum maximae and Caricetum elatae. In a belt in the Upper Basin where neither flooding nor upward seepage occurs, succession, probably caused by intensified drainage, leads to a dwarf-shrub vegetation (Betuletum humilis; poor fen).     

Methanogen communities along a primary succession transect of mire ecosystems

Peat accumulating mires are important sources of the greenhouse gas methane. Methane emissions and methanogenic Archaea communities have been shown to differ between fens and bogs, implying that mire succession includes an ecological succession in methanogen communities. We investigated methane production and the methanogen communities along a chronosequence of mires (ca. 100-2,500 years), which consisted of five sites (1-5) located on the land-uplift coast of the Gulf of Bothnia. Methane production was measured in a laboratory incubation experiment. Methanogen communities were determined by amplification of a methyl coenzyme M-reductase (mcr) gene marker and analyzed by terminal-restriction fragment length polymorphism. The terminal-restriction fragment length polymorphism fingerprinting resulted in 15 terminal restriction fragments. The ordination configuration of the terminal restriction fragments data, using nonmetric multidimensional scaling, showed a clear gradient in the methanogen community structure along the mire chronosequence. In addition, fingerprint patterns of samples from the water table level and 40 cm below differed from one another in the bog site (site 5). Methane production was negligible in the three youngest fen sites (sites 1-3) and showed the highest rates in the oligotrophic fen site (site 4). Successful PCR amplification using mcr gene primers revealed the presence of a methanogen community in all five sites along the study transect.     

Classification and ordination of the mire vegetation of Stormyra near Tynset, S Norway

The vegetation of Stormyra, a flat fen at Tynset, in Hedmark province, Southern Norway is classified by cluster analyses into seven different plant communities, all of a nonfixed hierarchical, low rank as follows:
(1) Scorpidium scorpioides comm., (2) Campylium stellatum - Drepanocladus revolvers comm., (3) Carex rostrata - Calliergon giganteum comm., (4) Sphagnum papillosum - S. subfulvum comm., (5) Homalothecium nitens - Sphagnum warnstorfii comm., (6) Pinus sylvestris - Sphagnum angustifolium comm. and (7) Empetrum hermaphroditum - Sphagnum fuscum comm.
A DCA-ordination showed that the samples were distributed mainly along a hummock - mud-bottom vegetational gradient, reflecting the local groundwater level. The range of the plant communities in terms of the traditional poor - rich and mire expanse - mire margin vegetational gradients is also dealt with. The vegetation and its relationship to such environmental factors as the ground-water level, and its seasonal fluctuations, the supply or absence of running surface water, the pH and electrical conductivity, are also discussed.     

Base mineral inflow in a remnant cool-temperate mire ecosystem

Kiushitou (42°28N, 141°9E) is a lowland mire located in a residential area of northern Japan. We examined the 2-D distribution of hydrochemical variables and their seasonal changes in relation to plant communities in an attempt to conserve the Sphagnum fen (Sphagnum subfulvum). This mire is gently sloping and the upper area consists of alder and ash forests, while the lower area is covered with fen communities. The grassy fen, Moliniopsis japonica, occurs throughout the lower area, whereas the Sphagnum fen is restricted to the southwest part of the mire. anova and canonical correspondence analysis revealed that the occurrence of Sphagnum fen is negatively correlated with Mg, Ca and electrical conductivity (EC). These variables indicated that water in the upper forest area contained a high concentration of minerals from the neighboring residential area. Seasonal changes in EC values revealed that the watercourse from a spring point in an upper corner to the lowest drainage ditch was divided into two, southwest and northeast, courses. Because the northeast-course spring water joins mineral-rich water flowing from a point at the upper margin, the northeast part of the lower area contains considerable Ca and Mg. In contrast, the mineral-poor spring water flowing into the southwest part of the mire ensures the survival of the Sphagnum fen. Thus, when we stop the supply of mineral-rich water from the upper margin the area of the Sphagnum fen will expand into the northeast part of the mire. Two-dimensional details of the hydrochemical regime clarify the impact of mineral inflow and the expansion mechanisms of these minerals.     

Yield of wheat across a subambient carbon dioxide gradient

Yields and yield components of two cultivars of day-neutral spring wheat ( Triticum aestivum L.) were assessed along a gradient of daytime carbon dioxide (CO₂) concentrations from about 200 to near 350 μmol CO₂ (mol air)^–1 in a 38 m-long controlled environment chamber. The range in CO₂ concentration studied approximates that of Earth's atmosphere since the last ice age. This 75% rise in CO₂ concentration increased grain yields more than 200% under well-watered conditions and by 80–150% when wheat was grown without additions of water during the last half of the 100-day growing season. The 27% increase in CO₂ from the pre-industrial level of 150 years ago (275 μmol mol^–1) to near the current concentration (350 μmol mol^–1) increased grain yields of 'Yaqui 54' and 'Seri M82' spring wheats by 55% and 53%, respectively, under well-watered conditions. Yield increased because of greater numbers of grains per spike, rather than heavier grains or numbers of spikes per plant. Water use increased little with CO₂ concentration, resulting in improved water use efficiency as CO₂ rose. Data suggest that rising CO₂ concentration contributed to the substantial increase in average wheat yields in the U.S. during recent decades.     

Water chemistry of mires in relation to the poor-rich vegetation gradient and contrasting geochemical zones of the north-eastern fennoscandian Shield

The relationship of the poor-rich vegetation gradient with water chemistry and geochemical bedrock zones was studied with a data set from 36 pristine mire sites in the middle boreal zone, Finland. The bipartition of bedrock zones into a granitoid zone and three other, more alkaline bedrock zones was clearly connected to the poor-rich gradient in the study area. Mires in the granitoid bedrock zone were predominantly bogs and poor fens, while most of the rich fen sites were located in other bedrock zones. In fens with strongly minerogenous hydrology, the geochemically distinct bedrock zones showed significant differences in concentrations of calcium, manganese, phosphorus, as well as in the total equivalent charge of base cations and the ratio of calcium and magnesium. In light of the indicator species baseda priori poor-rich classification anda posteriori correlations with vegetation ordinations and a poor-rich indicator cover index, pH showed the strongest relationship of the water chemical variables with the poor-rich gradient. High calcium concentrations only separated extremely rich fens from other categories. The poor-rich gradient of mires is considered as a primarily pH-related vegetation continuum. A more general use of pH-classes along with hydrological and vegetation based classification is suggested for mire classification instead of the “poor-rich” terminology.     

Element concentrations in mosses and surface waters of western Canadian mires relative to precipitation chemistry and hydrology

Concentrations of N, P, S, Na, K, Mg, Ca, Mn, Fe, Cu, Cd, Zn, Pb, Al, and AIA (acid insoluble ash) m mosses (three Sphagnum species and Tomenthypnum nitens, all hummock species) from a variety of mires, both ombrotrophic and minerotrophic, in the coastal western and central parts of Canada are considered in relation to surface water pH and concentrations of Na⁺, K⁺, Mg²⁺, Ca²⁺, Cl⁻, and SO₄^2- Distinct west-east concentration gradients were present for most elements in both mosses and water, but there were correlations between surface water and moss concentrations only for Ca and Mg
On ombrotrophic sites and sites characterized by poor fen vegetation, wet deposition is the main source of elements in the surface water On rich fen sites, additional Ca and Mg from surrounding soils change the elemental proportions We conclude that hydrochemically the limit between poor and rich fen sites is more decisive than between bog and fen The increase in Ca may give brown mosses a competitive advantage over Sphagnum
Moss concentrations of Na and Mg are the only ones decreasing inland The constancy or inland increase of moss elemental concentrations may depend on either an increasing atmospheric supply (e g Pb), differences in moss growth rates (especially N, P, and K) or site conditions related to the water regime (e g Fe and Al)     

Plasma steroid concentrations in relation to the reproductive cycle of cultured male Rhamdia quelen

Seepage water through a lake bottom and the layering of tight and high-permeable layers in the aquifer produce groundwater under greater pressure than the water column of the lake. This results in under-water springs and seepages. When spawning substrata were available, there was a spatial correlation between the degree of groundwater influx and the redd density of brown trout Salmo trutta . In high density spawning areas with > 100 observed redds ha⁻¹, the mean seepage water influx during spring was c . 1200 ml m⁻² min⁻¹ compared to a mean flux of 113 ml m⁻² min⁻¹ in low density spawning areas with 5–10 redds ha⁻¹.     

The trophic basis of production of the macroinvertebrate community of a southeastern U.S.A. blackwater stream

The trophic basis of production of the macroinvertebrate communities at three sites on a second-order, low gradient blackwater stream in southeastern U.S.A. was determined. The sampling sites were located above, within and below a low-flow swamp system. From 47–64% of macroinvertebrate production was supported by FPOM at the three sites, with dependence on FPOM being greatest at the swamp site. Algae (filamentous species and diatoms) supported 15–31% of production, indicating that algae can be of considerable importance even in fully canopied headwater streams. The production of some collector-gatherers including Stenonema modestum (55%), Hexagenia munda (58%) and Baetis spp. (78%), was supported predominantly by algae. Algae also supported 61–79% of Hydropsychidae production and 68% of Simuliidae production. Animal material supported 16–26% of macroinvertebrate production at the three sites. CPOM was of minor direct importance to the macroinvertebrate community of this headwater stream, supporting only 1–3% of macroinvertebrate production. Shredders ingested only 1–3 g m⁻² y⁻¹ of CPOM, or about 1% of the annual direct leaf fall to this stream. Assuming a 10% assimilation efficiency for CPOM, shredders produced <3 g m⁻² y⁻¹ of FPOM through CPOM processing, this being approximately 2 orders of magnitude less than reported for high gradient headwater streams. These results indicate that low-order coastal plain streams vary somewhat from the River Continuum Concept in that they exhibit little utilization of and dependence on CPOM as a direct energy source. Only the smallest first-order streams and especially the extensive floodplains may be the functional headwaters of these stream systems.     

Composition and structure of a species rich Amazonian rain forest obtained by two different sample methods

Inventory data for trees ≥ 10 cm DBH from a hectare plot are compared to data obtained by the Point-Centered Quarter Method along a line transect from the same locality in Anangu, Amazonian Ecuador. The one-hectare quadrat plot of 100 × 100 m had 734 individuals, 153 species, 46 families, a total basal area of 22.2 m², and an estimated above ground tree volume of 240.5 m³. The line transect had a calculated density of 728 individuals per hectare, which included 239 species, 51 families, a total basal area of 34.1 m², and an estimated above ground tree volume of 409.6 m³. Of the 20 species with the highest IVI, only four were shared by the two samples. The most important species were Quararibea ochrocalyx on the hectare plot and Iriartea deltoidea on the line transect, constituting 26.6 and 13.3% of the individuals, respectively. The five families with the highest FIV on the hectare plot (Bombacaceae, Arecaceae, Moraceae, Caesalpinaceae, and Lauraceae) and on the line transect (Arecaceae, Moraceae, Meliaceae, Mimosaceae and Caesalpinaceae) constitute 40.4% and 35.4% of the Family Importance Values of the samples, respectively. The Point-Centered Quarter Method used along a line transect reflects maximum diversity and provides average values of density and tree size in the area. The quadrat plot reflects the local structure and composition of the forest within the plot.     

Multiple gradients in mire vegetation: a comparison of a Swedish and an Italian bog

The major environmental gradients underlying plant species distribution were outlined in two climatically and bio-geographically contrasting mires: a Swedish bog in the boreo-nemoral zone, and an Italian bog in the south-eastern Alps. Data on mire morphology, surface hydrology, floristic composition, peat chemistry and pore-water chemistry were collected along transects from the mire margin (i.e., the outer portion of the mire in contact with the surrounding mineral soil) towards the mire expanse (i.e., the inner portion of the mire). The delimitation and the extent of the minerotrophic mire margin were related to the steepness of the lateral mire slope which, in turns, controls the direction of surface water flow. The mineral soil water limit was mirrored in geochemical variables such as pH, alkalinity, Ca²⁺, Mg²⁺, Al³⁺, Mn²⁺, and SiO₂ concentrations in pore-water, as well as Ca, Al, Fe, N and P contents in surface peat. Depending on regional requirements of plant species, different species were useful as fen limit indicators at the two sites. The main environmental factors affecting distribution of habitat types and plant species in the two mires were the acidity-alkalinity gradient, and the gradient in depth to the water table. The mire margin – mire expanse gradient corresponds to a complex gradient mainly reflected in a differentiation of vegetation structure in relation to the aeration of the peat substrate.     

Hydrological landscape settings of base‐rich fen mires and fen meadows: an overview

Question: Why do similar fen meadow communities occur in different landscapes? How does the hydrological system sustain base‐rich fen mires and fen meadows? Location: Interdunal wetlands and heathland pools in The Netherlands, percolation mires in Germany, Poland, and Siberia, and calcareous spring fens in the High Tatra, Slovakia. Methods: This review presents an overview of the hydrological conditions of fen mires and fen meadows that are highly valued in nature conservation due to their high biodiversity and the occurrence of many Red List species. Fen types covered in this review include: (1) small hydrological systems in young calcareous dune areas, and (2) small hydrological systems in decalcified old cover sand areas in The Netherlands; (3) large hydrological systems in river valleys in Central‐Europe and western‐Siberia, and (4) large hydrological systems of small calcareous spring fens with active precipitation of travertine in mountain areas of Slovakia. Results: Different landscape types can sustain similar nutrient poor and base‐rich habitats required by endangered fen meadow species. The hydrological systems of these landscapes are very different in size, but their ground water flow pattern is remarkably similar. Paleoecological research showed that travertine forming fen vegetation types persisted in German lowland percolation mires from 6000 to 3000 BP. Similar vegetation types can still be found in small mountain mires in the Slovak Republic. Small pools in such mires form a cascade of surface water bodies that stimulate travertine formation in various ways. Travertine deposition prevents acidification of the mire and sustains populations of basiphilous species that elsewhere in Europe are highly endangered. Conclusion: Very different hydrological landscape settings can maintain a regular flow of groundwater through the top soil generating similar base‐rich site conditions. This is why some fen species occur in very different landscape types, ranging from mineral interdunal wetlands to mountain mires.     

Establishment and persistence of common ragweed (Ambrosia artemisiifolia L.) in disturbed soil as a function of an urban–rural macro-environment

No data are available on whether rising carbon dioxide concentration [CO₂] or increased air temperature can alter the establishment and persistence of common ragweed ( Ambrosia artemisiifolia L.) within a plant community following soil disturbance. To determine ragweed longevity, we exposed disturbed soil with a common seed bank population to an in situ temperature and [CO₂] gradient along an urban–rural transect beginning in early 2002. No other consistent differences in meteorological variables (e.g. wind speed, humidity, PAR, tropospheric ozone) as a function of urbanization were documented over the course of the study (2002–2005). Above-ground measurements of biomass over this period demonstrated that ragweed along the transect responded to urban induced increases in [CO₂]/temperature with peak biomass being observed at this location by the end of 2003. However, by the Fall of 2004, and continuing through 2005, urban ragweed populations had dwindled to a few plants. The temporal decline in ragweed populations was not associated with increased disease, herbivory or auto-allelopathy, but was part of a demographic reduction in the total number of annual plant species observed for the urban location. In a separate experiment, we showed that such a demographic shift is consistent with CO₂/temperature induced increases in biomass and litter accumulation, with a subsequent reduction in germination/survival of annual plant species. Overall, these data indicate that [CO₂]/temperature differences associated with urbanization may increase initial ragweed productivity and pollen production, but suggest that long-term, multi-year persistence of ragweed in the urban macro-environment may be dependent on other factors.     

Life-cycle, drift and production of Gammarus pulex L. (Amphipoda) in a Danish spring

SUMMARY. Gammarus pulex L. was studied at three sites in a Danish spring. Breeding took place during the summer and therefore cohorts could be followed. Sexual maturity was not reached until the next summer and maximum age appeared to be two years.
Changes in population size differed between stations and was tentatively related to differences in drift and numbers of predators. Generally the drift was non-selective between cohorts and within cohorts.
Yearly production estimated from production curves was 2.6 g dry weight m⁻² (the young cohort) and 1.2 g dry weight (the old cohort) at Station 1. The respective P / ratios were 3.35 and 1.22. Production of the youngest cohort at the three stations in the period September-May was 0.9–1.3 g m⁻² and P / ratios were 2.14–2.31. Total yearly production at the three stations was estimated at 4–7 g dry weight m⁻².     

Wind as a factor influencing flower-visiting by Hadena bicruris (Noctuidae) and Deilephila elpenor (Sphingidae)

Abstract. 1. In a low-speed wind tunnel, male as well as female moths of Hadena bicruris responded to floral odours with positive anemotaxis. Hitherto, such orientation has only been demonstrated for male moths in response to pheromones.
2. H.bicruris had a maximum flight speed of 4.5–5.4ms^-1 and stopped its flower visiting at a wind speed of 2.5–2.8 m s^-1.
3. Deilephila elpenor had a similar maximum flight speed (4.5— 5.1 ms^-1), but it continued visiting flowers up to wind speeds of 3.0–5.0 ms^-1.
4. Apart from mechanical resistance during flight and flower visits, wind might have adverse effects on the energy budget and on evaporative water loss.     

Predation and potential transfer of pollen in a population of Saxifraga hirculus

Body surface pollen load and gut pollen (pollen diet load) of flower visitors in a Danish population of Saxifraga hirculus were analyzed. The four most frequent visitors, Eurimyia lineata (Syrphidae), Neoascia tenur (Syrphidae), Asindulum nigrum (Mycetophilidae), and Zygaena trifolii (Zygaenidae) had total surface loads (and S. hirculus loads) of 1031 (771), 137 (106), 108 (78), and 318 (27) pollen grains, respectively. Other syrphid visitors had on average a load of 407 (192) pollen grains. Males of E. lineata, N. tenur , and A. nigrum carried more surface pollen than the females; females of E. lineata had a lot of pollen from Ranunculus acris and S. hirculus and males had a lot from S. hirculus . Females of N. tenur carried pollen mostly from S. hirculus, Lotus uliginosus and Galium uliginosum ; males carried pollen mostly from S. hirculus . The surface pollen load of A. nigrum was dominated by S. hirculus pollen, whereas that of Z. trifolii was dominated by Cirsium palustre and Lychnis flos-cuculi pollen. The amount and kind of pollen varied enormously both intraspecifically and interspecifically. Eurimyia lineata, N. tenur and other syrphids had total pollen diet loads (and S. hirculus pollen diet loads) of approximately 14900 (657), 3739 (2327) and 17784 (13040). The average syrphid contained pollen from five plant species, only one to two, however, were quantitatively important. The average diet load in females was nine times that in males in both E. lineata and N. tenur. Eurimyia lineata ate a lot of R. acris; N. tenur ate a lot of S. hirculus and G. uliginosum . Generally, no correlation was observed between size or season and the composition of surface load and diet load. Eurimyia lineata made most of the pollen visits. Thus, the seed set from this Danish population of S. hirculus was dependent on a single syrphid species.     

Biomass and oxygen dynamics of the epiphyte community in a Danish lowland stream

SUMMARY. 1. We examined the abundance and oxygen metabolism of epiphytic organisms on the dominant macrophyte, Potamogeton pectinatus , in headwaters of the eutrophic River Suså. Microbenthic algae were abundant in the stream during spring and macrophytes during summer.
2. The low macrophyte biomass in spring supported a dense epiphyte cover whereas the high macrophyte biomass during summer had a thin epiphyte cover of 10–100-fold lower abundance per unit area of macrophyte surface. The epiphyte community was dominated by microalgae in spring and by heterotrophs, probably bacteria, during summer. This seasonal shift was shown by pronounced reductions of the chlorophyll a content (from 2–3% to 0.1–0.7% of organic DW), the gross photosynthetic rate (from 20–85 to 3–15 mg O₂, g-¹ organic DW h⁻¹) and the ratio of gross photosynthesis to dark respiration in the epiphyte community (from 5–18 to 1). The reduced contributions of epiphytic microalgae correlated with reduced light availability during summer.
3. Both the density and the photosynthetic activity of epiphytic algae were low on a stream area basis relative to those of microbenthic algae and macrophytes. Rapid variations in water velocity and extensive light attenuation in water and macrophyte stands probably constrained the development of epiphytic algae. The epiphyte community was more important in overall stream respiration, contributing c. 10% to total summer respiration and c. 20% to summer respiration within the predominantly heterotrophic microbial communities on sediments and macrophyte surfaces.