Carbon flow in the littoral food web of an oligotrophic lake

Benthic food web dynamics and carbon flow were examined in the littoral zone of Lake Coleridge, a large deep oligotrophic lake, using radioactive and stable isotope techniques in conjunction with analyses of stomach contents of the fauna. We specifically address two hypotheses: (1) that macrophytes only contribute to the carbon flow to higher trophic levels when they have decayed; and (2) that epiphytic algae is the major source of carbon for macroinvertebrates, and thus fish, with only minor contributions from phytoplankton or terrestrial sources. Epiphytic diatoms were a major component of the stomach contents of the gastropod snail Potamopyrgus antipodarum, and of chironomids. Animal remains were also common in the diet of some chironomids, while amorphous organic matter predominated in the stomachs of oligochaetes. A variety of epiphytic algal taxa was found in trichopteran larvae. Feeding rate of P. antipodarum measured with radioactive tracers increased by 10× on decayed macrophytes (Elodea) compared with live material, while feeding rates on characean algae increased by a factor of 3 when decayed material was presented. However, assimilation rates were less than 20% on decayed material compared with 48–52% on live material. Potential carbon sources were easily distinguished based on their ¹³C values, although isotopic ratios showed significant variation among sites. Epiphytic algae showed less variation among sites than macrophytes and were depleted by 4–5 compared with macrophytes. Detrital material, organic matter in the sediments and plankton were significantly depleted in ¹³C relative to macrophytes and slightly depleted relative to epiphytic algae. Most macroinvertebrate taxa showed a similar pattern among sites to macrophytes and epiphytic algae. P. antipodarum and chironomids were slightly enriched compared with epiphytic algae. Ratios for the common bully (Gobiomorphus cotidianus) were generally consistent with a diet dominated by chironomids, while there was some evidence for terrestrial inputs for koaro (Galaxias brevipinnis) and juvenile brown trout. Epiphytic algae appear to underpin much of the production in the littoral zone of this oligotrophic lake, with trichopteran and chironomid larvae mediating carbon flows from algae to fish. Macrophytes do not make a major contribution directly to carbon flow to higher trophic levels even when decayed. The lack of a direct link between macrophytes and higher trophic levels is due to the faunal composition, including a lack of large herbivores.     

A Century of Change in Macrophyte Abundance and Composition in Response to Agricultural Eutrophication

Clear Lake, Iowa, USA is a shallow, agriculturally eutrophic lake that has changed drastically over the past century. Eight macrophyte surveys since 1896 were pooled and examined to characterize long-term impacts of eutrophication on macrophyte community composition and relative abundance. Surveys in 1981 and 2000 revealed few submergent and floating-leaved species and a dominance in emergent species (Scirpus, Typha). Over the past century, however, species richness has declined from a high of 30 species in 1951 to 12 found today, while the community composition has shifted from submergent-(99%) to emergent-dominated floras (84%). Potamogeton praelongus was the first emergent species to disappear but was followed by several other clear water Potamogeton species. Several floating leaved and emergent genera increased in relative abundance with eutrophication, notably Nuphar, Nymphaea, Phragmites, Polygonum, Sagittaria, Scirpus, and Typha. P. pectinatus was present over the entire century due to its tolerance of eutrophic conditions. Macrophyte growth was generally light-limited, with 93% of the variance in relative abundance of submergent species explained by changes in water transparency. Clear Lake exhibits signs of alternative stable states, oscillating between clear and turbid water, coupled with high and low submerged species relative abundance. The maximum macrophyte richness occurred as the lake oscillated between submergent- and emergent-dominated states. Changes in the water level have also impacted macrophyte growth since the area of the lake occupied by emergent macrophytes was negatively correlated with water level. Strongest correlations indicated that macrophytes respond to water level variations with a 2-year time-lag.     

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.     

Long‐term dynamics of submerged macrophytes and algae in a small and shallow,eutrophic lake: implications for the stability of macrophyte‐dominance

1. Submerged macrophyte and phytoplankton components of eutrophic, shallow lakes have frequently undergone dynamic changes in composition and abundance with important consequences for lake functioning and stability. However, because of a paucity of long‐term survey data, we know little regarding the nature, direction and sequencing of such changes over decadal–centennial or longer timescales. 2. To circumvent this problem, we analysed multiple (n = 5) chronologically correlated sediment cores for plant macro‐remains and a single core for pollen and diatoms from one small, shallow, English lake (Felbrigg Hall Lake, Norfolk, U.K.), documenting 250 years of change to macrophyte and algal communities. 3. All five cores showed broadly similar stratigraphic changes in macrophyte remains with three distinct phases of macrophyte development: Myriophyllum–Chara–Potamogeton (c. pre‐1900), to Ceratophyllum–Chara–Potamogeton (c. 1900–1960) and finally to Zannichellia–Potamogeton (c. post‐1960). Macrophyte species richness declined from at least 10 species pre‐1900 to just four species at the present day. Additionally, in the final Zannichellia–Potamogeton phase, a directional shift between epi‐benthic and phytoplankton‐based primary production was indicated by the diatom data. 4. Based on macrophyte–seasonality relationships established for the region, concomitant with the final shift to Zannichellia–Potamogeton, we infer a reduction in the seasonal duration of plant dominance (plant‐covered period). Furthermore, we hypothesise that this change in species composition resulted in a situation whereby macrophyte populations were seasonally ‘sandwiched’ between two phytoplankton peaks in spring and late summer as observed in the contemporary lake. 5. We suggest that eutrophication‐induced reductions in macrophyte species richness, especially if the number of plant‐seasonal strategies is reduced, may constrict the plant growing season. In turn, this may render a shallow lake increasingly vulnerable to seasonal invasions of phytoplankton resulting in further species losses in the plant community. Thus, as part of a slow (over perhaps 10–100s of years) and self‐perpetuating process, macrophytes may be gradually pushed out by phytoplankton without the need for a perturbation as required in the alternative stable states model of plant loss.     

Hydrologic and anthropogenic influences on aquatic macrophyte development in a large,shallow lake in China

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THE INFLUENCE OF SUBMERGED MACROPHYTES ON SEDIMENTARY DIATOM ASSEMBLAGES1

Submerged macrophytes are a central component of lake ecosystems; however, little is known regarding their long‐term response to environmental change. We have examined the potential of diatoms as indicators of past macrophyte biomass. We first sampled periphyton to determine whether habitat was a predictor of diatom assemblage. We then sampled 41 lakes in Quebec, Canada, to evaluate whether whole‐lake submerged macrophyte biomass (BiomEpiV) influenced surface sediment diatom assemblages. A multivariate regression tree (MRT) was used to construct a semiquantitative model to reconstruct past macrophyte biomass. We determined that periphytic diatom assemblages on macrophytes were significantly different from those on wood and rocks (ANOSIM R = 0.63, P < 0.01). A redundancy analysis (RDA) of the 41‐lake data set identified BiomEpiV as a significant (P < 0.05) variable in structuring sedimentary diatom assemblages. The MRT analysis classified the lakes into three groups. These groups were (A) high‐macrophyte, nutrient‐limited lakes (BiomEpiV ≥525 μg · L^?1; total phosphorus [TP] <35 μg · L^?1; 23 lakes); (B) low‐macrophyte, nutrient‐limited lakes (BiomEpiV <525 μg · L^?1; TP <35 μg · L^?1; 12 lakes); and (C) eutrophic lakes (TP ≥35 μg · L^?1; six lakes). A semiquantitative model correctly predicted the MRT group of the lake 71% of the time (P < 0.001). These results suggest that submerged macrophytes have a significant influence on diatom community structure and that sedimentary diatom assemblages can be used to infer past macrophyte abundance.     

Factors affecting the structure of epiphytic gastropod communities in the St. Lawrence River (Quebec,Canada)

A study of epiphytic gastropods associated with two submerged macrophytes (Myriophyllum spicatum and Vallisneria americana) was conducted at a soft and at a hard water site in the St. Lawrence River during two non-consecutive years in order to compare effects of macrophyte species and biomass, site and year in gastropod community structure. There was no effect of macrophyte species on total gastropod abundance nor on gastropod diversity, and few gastropod species showed a marked preference for either macrophyte species. Inter-site and inter-annual variations in diversity, total gastropod abundance and gastropod community structure were greater than variations among macrophytes. However, analysis of variations of epiphytic gastropod communities in one of the two sites shows that abiotic factors are important in explaining epiphytic gastropod distribution. Our results and results of other studies on gastropod population dynamics in the St. Lawrence River indicate that intra- and interspecific competition between gastropods is important on macrophytes and that they must be carefully considered in order to understand epiphytic community structure and population dynamics.     

The Seasonal Succession of Epiphytic Communities within an Equisetum fluviatile L. Stand in Lake Pääjärvi,Southern Finland

The seasonal succession of epiphytic communities on Equisetum fluviatile was controlled both directly and indirectly by the macrophyte. Decaying macrophytic material supported rich algal growth and biomass accumulation in spring and early summer. Emergence of the macrophytes severely reduced underwater light availability, and the epiphytic algal biomass declined rapidly as a result of both lower photosynthetic activity of the epiphytic algae, and more intensive grazing by invertebrate herbivores. Epiphytic N: P ratios were lower than those in the water around suggesting that either the water was not the only source of phosphorus for the epiphytic algae or the algae took phosphorus up selectively from the water. Low epiphytic C: N ratios suggested a high potential nutritional value for herbivores.     

Herbivory of invertebrates on submerged macrophytes from Danish freshwaters

1. Invertebrate herbivory on submerged freshwater macrophytes, measured as per cent leaf area lost, was determined for sixteen species and forty-two populations of macrophytes during peak summer biomass in Danish streams and lakes. 2. All seventeen Potamogeton populations and seventeen of the remaining twenty-five non-Potamogeton populations were grazed. Species of Potamogeton were significantly more heavily grazed (mean 4.2%) than non-Potamogeton species (mean 0.8%). Herbivory losses were not significantly different between stream (mean 2.4%) and lake populations (mean 1.9%). Wide ranges in herbivory loss were observed between species from the same locality and within species from different localities. The location of main damage to either old or young leaves was not species specific but varied among localities. Additional data for four macrophyte populations showed that herbivory loss had a strong seasonal variation (e.g. 1.0–26.3% for Potamogeton perfoliatus), with maximum losses during May-June. 3. Although the mean defoliation percentages were low during the period of maximum macrophyte biomass, they were not systematically lower than encountered for terrestrial plants.     

Factors affecting the contribution by epiphytic algae to the primary productivity of an oligotrophic freshwater lake

A diatom-dominated population of epiphytic algae was studied in an oligotrophic lake to determine the factors which limit epiphyte growth and to measure their contribution to primary productivity. Algae were collected from plants growing at four sites in Lake George, N.Y., during the spring, summer, and fall of 1974. Samples were taken from 3 m, corresponding to the depth at which macrophytes were most productive. Algae exhibited an optimum temperature for H¹⁴CO₃^- uptake at 30 C, although the summer littoral lake temperature ranged from 18 to 25 C. Light saturation occurred at an intensity of 8,608 lux, approximating the environmental intensity at the depth from which algae were taken. Epiphytes exhibited their maximum photosynthetic capacity of 0.6 mg of carbon fixed/m² of macrophyte surface area per h in the early afternoon in mid-August. They assimilated approximately 5% as much inorganic carbon as the macrophytes from which they were taken. Epiphyte population densities followed the seasonal growth patterns of the macrophytes, with maximal leaf colonization remaining essentially constant relative to the leaf position on the plant. There was little change in density between sampling sites at any given time. Productivities of epiphytes from bottom leaves were 10-fold greater than those of epiphytes from top leaves. Addition of PO₄^-3, NO₃^-, NH₃, Si, and SO₄^-2 had no stimulatory effect on photosynthesis. Addition of HCO₃^- stimulated photosynthesis greater than 30%, suggesting that carbon may be a limiting nutrient for epiphytic algae in Lake George.     

Periphyton in Lake Vechten,with emphasis on biomass and production of epiphytic algae

Summary Characteristics of the periphyton associated with the submerged macrophytesCeratophyllum, Elodea andMyriophyllum were similar. During the whole growing season the periphyton dry weight often exceeded that of the macrophyte. The periphyton contained much detritus and carbonates, and the epiphytic algae were accompanied by large numbers of bacteria, protozoans and rotifers. Attached algae were scarce, but motile diatoms were abundant throughout the year. Thus the periphyton showed the features of a benthic community. Consequently, the role of the macrophytes would be rather the extension of the bottom area instead of actively promoting the growth of microorganisms by providing minerals or excreting organic substances.During the peak of the littoral development the epiphytic algae expressed a high photosynthetic capacity, while at the same time the periphyton dark respiration was intense. After measurements of the depth distribution of biomass and light in aCeratophyllum stand, it was concluded that the contribution of the epiphytic algae to the production in the lake's littoral zone was substantial. Furthermore, on a daily basis, the total association of macrophytes and periphyton would be oxygen consuming due to heterotrophic activity, notably mineralization of the detritus.     

Die überwinterungsknospen des Krausen Laichkrauts (Potamogeton crispus) als Nahrung für den H?ckerschwan (Cygnus olor) auf dem Federsee

Summary In the highly eutrophicated, shallow Lake Federsee no submersed macrophytes grew, until 1982 a waste water pipeline around the lake was put in operation. In 1983Potamogeton crispus began to colonize the lake and its abundance increased from year to year. The growth cycle, however, remained very short and the macrophyte always died off between the end of June and the middle of July, after having developed large numbers of resting buds. Simultaneous with the increasing canopies ofPotamogeton a rise of Mute Swans in summer and autumn was observed, and because of the absence ofPotamogeton and other new food plants it was suggested that at this time the bird mainly feeds on the resting buds ofPotamogeton. This was confirmed by investigation of excrements, which sometimes consisted mainly of remnants of resting buds. Therefore it is supposed that Mute Swans essentially affect the recolonization of the lake byPotamogeton crispus and by this way also slow down the reestablishment of a diverse flora and fauna.     

Whole-lake food-web manipulation as a means to study community interactions in a small ecosystem

Whole-lake food-web manipulation was carried out in the hypertrophic Lake Zwemlust (The Netherlands), with the aim of studying the effects on the lake's trophic status and to gain an insight into complex interactions among lake communities. Before manipulation this small (1.5 ha) and shallow (1.5 m) lake was characterized byMicrocystis blooms in summer and high chlorophyll-a concentrations were common (ca. 250 μg 1⁻¹). In March 1987 the planktivorous and benthivorous fish species in the lake were completely removed (ca. 1000 kg ha⁻¹), a new simple fish community (pike and rudd) was introduced and artificial refuges were created. The effects of this manipulation on the light climate, nutrient concentrations, phytoplankton, zooplankton, fish, macrophytes, and macrofauna were monitored during 1987, 1988 and 1989. Community interactions were investigated in phytoplankton bioassays and zooplankton grazing experiments. After the manipulation, despite the still high P and N loads to the lake (ca. 2.2 g P m⁻² y⁻¹ andca. 5.3 g N m⁻² y⁻¹), the phytoplankton density was low (Chl-a<5μg l⁻¹), due to control by large-sized zooplankton in spring and N-limitation in summer and autumn. A marked increase in the abundance of macrophytes and filamentous green algae in 1988 and 1989, as well as N loss due to denitrification, contributed to the N limitation of the phytoplankton. Before manipulation no submerged macro-vegetation was present but in 1988, the second year after manipulation, about 50% of the lake bottom was covered by macrophytes increasing to 80% in 1989. This led to substantial accumulation of both N and P, namely 76% and 73% respectively of the total nutrients in the lake in particulate matter. Undesirable features of the increase in macrophytes were: 1) direct nuisance to swimmers; and, 2) the large scale development of snails, especiallyL. peregra, which may harbour the parasite causing ‘swimmers' itch’. But harvesting of only about 3% of the total macrophyte biomass from the swimmers' area, twice a year, reduced the nuisance for swimmers without adversely affecting the water clarity.     

Effect of macrophyte community composition and nutrient enrichment on plant biomass and algal blooms

Submerged freshwater macrophytes decline with increasing eutrophication. This has consequences for ecosystem processes in shallow lakes and ponds as macrophytes can reduce algal blooms under eutrophic conditions. We hypothesize that the productivity of submerged vegetation, biomass change under eutrophication and the suppression of algal blooms may be affected by macrophyte community composition. To test our hypothesis, we established three macrophyte community types in 36 fishless experimental ponds: one dominated by the oligotrophic species Chara globularis, one dominated by the eutrophic species Potamogeton pectinatus and a diverse vegetation which became co-dominated by Elodea nuttallii and C. globularis, and we fertilized half of the ponds.The macrophyte communities produced different amounts of biomass and they responded differently to fertilization. The community dominated by Potamogeton produced the lowest overall biomass, but was not affected by nutrient addition. The communities dominated by Chara and co-dominated by Elodea and Chara produced more than four-fold the amount of biomass produced in Potamogeton communities under oligotrophic conditions, but were strongly negatively affected by nutrient addition.Phytoplankton abundance did not differ significantly among the plant community types, but showed large variation within community types. There was a significant negative relationship between spring macrophyte biomass and the probability of summer algal blooms. The occurrence of algal blooms coincided with low daphnid densities and high pH (>10).We conclude that the macrophyte community composition, characterized by the dominant species, strongly affected the amount of biomass production as well as the short-term response of the vegetation to nutrient enrichment. Macrophyte community composition had no direct effect on algal blooms, but can affect the occurrence of algal blooms indirectly as these occurred only in ponds with low (<100 g/m² DW) spring macrophyte biomass.     

Submerged Macrophyte-bed Effects on Water-Column Phosphorus,Chlorophyll <Emphasis Type="Italic">a</Emphasis>, and Bacterial Production

Submerged macrophytes are a major component of freshwater ecosystems, yet their net effect on water column phosphorus (P), algae, and bacterioplankton is not well understood. A 4-month mass-balance study during the summer quantified the net effect of a large (5.5 ha) undisturbed macrophyte bed on these water-column properties. The bed is located in a slow-flowing (0.05–0.1 cm s^–1) channel between two lakes, allowing for the quantification of inputs and outputs. The P budget for the study period showed that, despite considerable short-term variation, the macrophyte bed was a negligible net sink for P (0.06 mg m^–2 day^–1, range from –0.76 to +0.79 mg m^–2 day^–1), demonstrating that loading and uptake processes in the weedbed roughly balance over the summer. Chlorophyll a was disproportionately retained relative to particulate organic carbon (POC), indicating that the algal component of the POC was preferentially trapped. However, the principal contribution of the weedbed to the open water was a consistent positive influence on bacterioplankton production over the summer. Conservative extrapolations based on measured August specific exports (m^–2 day^–1) of P and bacterial production exiting the weedbed applied to five regional lakes varying in lake morphometry and macrophyte cover suggest that even in the most macrophyte dominated of lakes (66% cover), P loading from submerged weedbeds never exceeds 1% day^–1 of standing epilimnetic P levels, whereas subsidization of bacterioplankton production can reach upward of 20% day^–1. The presence of submerged macrophytes therefore differentially modifies algae and bacteria in the water column, while modestly altering P dynamics over the summer.     

Herbivory in omnivorous fishes: effect of plant secondary metabolites and prey stoichiometry

1. Many animals that consume freshwater macrophytes are omnivorous (i.e., they include both plant and animal matter in their diet). For invertebrate omnivorous consumers, selection of macrophyte species depends partly on the presence of secondary metabolites in plants, plant carbon/nutrient balances and/or physical structure of plants. However, little is known about the mechanisms influencing consumption of macrophytes in aquatic vertebrates. 2. For two fish species, the omnivorous rudd (Scardinius erythrophthalmus) and herbivorous grass carp (Ctenopharyngodon idella), feeding preferences were determined in three choice experiments. We tested (i) whether the presence of secondary metabolites and macrophyte stoichiometry affects macrophyte species selection by fish, (ii) the importance of macrophyte stoichiometry by manipulating the macrophytes experimentally and (iii) the rate of herbivory when the most palatable macrophyte is offered simultaneously with a common animal prey. 3. In a choice experiment with five species of submerged macrophytes (Callitriche sp., Chara globularis, Elodea nuttallii, Myriophyllum spicatum and Potamogeton pectinatus), Myriophyllum was clearly consumed least by both fishes, which strongly correlated with the highest phenolic concentration of this macrophyte. Additionally, a significant negative relationship was found between consumption and C : N ratio of the five macrophytes. The two most consumed macrophytes also had the lowest dry matter concentration (DMC). 4. In a second choice experiment, the C : N ratio of the least (Myriophyllum) and most (Potamogeton) palatable plants was manipulated by growing the macrophytes under fertilised and unfertilised conditions and subsequently feeding them to rudd. The avoidance of consumption of the chemically defended Myriophyllum by rudd was partly alleviated by the lowered C : N ratio. 5. The third choice experiment showed that both fishes preferred animal prey (the amphipod Gammarus pulex) over the most palatable macrophyte (Potamogeton) when offered simultaneously. The C : N ratio of the amphipods was about half that of the lowest C : N ratio measured in the macrophytes. Consumption by the fishes could not clearly be related to C : P or N : P ratios of prey items in any of the experiments. 6. We conclude that omnivorous fish avoid macrophytes that are chemically defended. However, when these defences are only minor, stoichiometry (C : N ratio) in combination with DMC may be a determining factor for consumption by vertebrate facultative herbivores.     

On body size and habitat selection in rotifers in a macrophye-dominated lake Budzyńskie,Poland

Lake Budzyńskie is shallow, freshwater lake with a well-developed and differentiated macrophytic vegetation. Zooplankton samples were collected from five stations: two of them in submerged macrophytes (Chara and Myriophyllum), one in the zone of floating leaves (Potamogeton), a rush station (Typha) and one in the open water surrounding the vegetation beds. The mean Rotifera densities differed significantly between the lake parts. Furthermore, different habitats were characterised by differences in body size with the exception of the middle body size group (Keratella cochlearis, Polyarthra vulgaris and Trichocerca similis), which was dominated by limnetic representatives. However, in all the other size-dependent groups both stands of submerged macrophytes were characterised by much higher densities than other zones. Additionally, body size within the examined habitats significantly differed. Thus, the size structure of Rotifera communities was directly related to morphological and spatial structures of the substrata. Two groups of habitats were distinguished: the first one consisting of open water and two vegetated zones of less complicated structure (Potamogeton and Typha), and the second of more complex submerged macrophyte species (Chara and Myriophyllum). The differentiation of the architecture of macrophytes affected the nutritional conditions and refuge effectiveness of these habitats.     

FIELD MEASUREMENTS OF INORGANIC NITROGEN UPTAKE BY EPIFLORA COMPONENTS OF THE SEAGRASS POSIDONIA OCEANICA (MONOCOTYLEDONS,POSIDONIACEAE)1

Crustose corallines, crustose and erect brown algae, and sessile animals are major components of the epiphytic community of the Mediterranean seagrass Posidonia oceanica (L.) Delile. Production, biomass, and specific composition of this epiphyte–seagrass association are impacted by anthropogenic increase of nutrient load in this oligotrophic area. In this context, nitrogen uptake by P. oceanica and its epiflora was measured using the isotope ¹⁵N at a 10 m depth in the Revellata Bay (Corsica, Mediterranean Sea). Epiflora components showed various seasonal patterns of biomass and abundance. The epiphytic brown algae appeared at the end of spring, later than the crustose corallines, and after the nitrate peak in the bay. Because of their later development in the season, epiphytic brown algae mostly rely on ammonium for their N needs. We hypothesize that the temporal succession of epiphytic organisms plays a crucial role in the N dynamics of this community under natural conditions. The epiphytic brown algae, which have a growth rate one order of magnitude greater than that of crustose corallines, showed lower N‐uptake rates. The greater N‐uptake rates of crustose corallines probably reflect the greater N requirements (i.e., lower C/N ratios) of red algae. We determined that the epiflora incorporated ammonium and nitrate more rapidly than their host. Nevertheless, when biomass was taken into account, P. oceanica was the most important contributor to N uptake from the water column by benthic macrophytes in this seagrass bed.     

Removal of settled sediments and periphyton from macrophytes by grazing invertebrates in the littoral zone of a large oligotrophic lake

• 1 The resistance and resilience of littoral zone communities to sedimentation will depend both on the extent to which sediment deposition affects productivity, and on interactions within the communities. A series of hypotheses were set up and tested to examine interactions and feedback mechanisms among deposited sediments, periphyton, macrophytes and grazers in a large oligotrophic lake subject to fluctuating sediment loadings.
• 2 Although sediments incorporated into periphyton reduced light availability to macrophytes, periphytic algae were generally the dominant light absorbing component under natural conditions. When grazers were absent, both sediments incorporated in the periphyton and periphytic algal densities increased, and both were then important in reducing light available to macrophytes.
• 3 Grazing rate and assimilation efficiency for the dominant grazer, the prosobranch gastropod Potamopyrgus antipodarum, increased with increasing sediment content under natural lake conditions to reach a maximum at 10 mg sediment cm^?2.
• 4 An increase in sediment incorporation into periphyton films resulted in an increased grazing rate and hence grooming of sediments from macrophytes.
• 5 Grazing invertebrates can play a major role in maintenance of littoral communities by continuously grooming macrophyte hosts of periphytic algae and settled sediments.

     

A simple tide simulator for maintenance of marine (intertidal) algal and lichen cultures

Strumpshaw Broad is a medieval man-made lake, excavated about 500 years ago in valley peat deposits. Evidence from diatom fossils, and other remains (filamentous algae, Chrysophycean cysts, sponge spicules, snail shells) indicates a series of changes since the eighteenth century probably related to development of the city of Norwich and sophistications of its sewerage system. Progressive nutrient-enrichment led to replacement around 1912 of low-growing macrophytes (Chara) and sparse plankton populations by tall-growing vascular aquatic macrophytes, with increased epiphytic diatom numbers. A blanketing phase of filamentous algae (probably Vaucheria) was associated with the transition between 1795 and 1903. Around 1950, after a phase of increasing plankton populations and declining epiphyte and macrophyte numbers, the macrophyte-epiphyte complex disappeared altogether, to be replaced by dense plankton populations for a few years. These changes and the progressive decrease in volume of the lake owing to increased build up of sediment throughout the period covered by the core, led to alterations in hydrology which now prevent even plankton development. All data are expressed on absolute bases (amount sedimented per unit area per year) and the necessity for this, as opposed to relative measures (percentage representation, concentration per unit weight) in valid interpretation of palaeolimnological data is emphasized.