Regulation of submersed macrophyte biomass in a temperate lowland river: Interactions between shading by bank vegetation, epiphyton and water turbidity

Growth of aquatic vegetation is often controlled by light supply, which is potentially decreased by bank vegetation, water turbidity and epiphytic biofilm. To understand the relative importance of these shading factors and the interactions between them we analysed the seasonal course of macrophyte biomass, shading by bank vegetation, turbidity of the water column and epiphytic light absorption in shaded and sunny sections of a temperate eutrophic lowland river. At a shaded site, bank vegetation decreased the light supply by 79%, 0.5 m water column by 45% and 2-week-old epiphyton by 28% during the vegetation period. Growth of submersed macrophytes, but not of epiphyton, was light-limited in the shaded sections. We found a saturation-type correlation between light supply and macrophyte biomass. Therefore, the additional light absorption of the water column or epiphyton only shortened the period of optimum light supply at the sunny site, but was crucial for macrophyte development at the shaded site. Light absorption of phytoplankton was most important in spring and that of epiphyton in late summer. Submersed macrophytes effectively retained particles and thus improved light supply of downstream stands, but this positive feedback effect was only relevant for shaded sections in summer.     

An experimental study of the influence of periphytic algae on invertebrate abundance in a Hong Kong stream

1. Small cages (294cm²) containing unglazed clay quarry tiles were used to investigate the influence of periphytic algae on macroinvertebrate abundance in a Hong Kong stream. Algal biomass was manipulated by shading cages with plastic sheets. Individual cages were assigned to one of three treatment groups: unshaded, shaded and deeply shaded. Invertebrate densities and algal biomass within cages were monitored after 23, 37 and 65 days. 2. Multiple-regression analysis revealed that algal biomass, invertebrate morphospecies richness and total abundance declined with greater shading intensity. The responses of individual invertebrate taxa varied: some (especially Trichoptera) were unaffected by shading, whereas grazers (Baetidae, Psephenidae and Elmidae) declined as shading increased. 3. Significant regressions of the densities of individual taxa upon algal and detrital standing stocks in cages had positive slopes, but algal biomass increased during the study while detrital standing stocks declined. Abundance of invertebrates declined or remained rather stable over time. Density increases resulting from a positive association with algae were apparently offset by declines in abundance correlated with reductions in detritus. 4. Declines in algal biomass were associated with greater shading to which animals may respond directly. To uncouple the link between scarcity of algae and reduction of light intensity, the plastic covers on two groups of cages (deeply shaded and unshaded) which had been placed in the stream for 28 days were reversed so that cages which had been shaded became unshaded and vice versa. The cages were recovered on day 33, Only Coleoptera demonstrated a positive association with atgae inside cages; no relationship between population densities and algal biomass or light intensity was apparent for other taxa. However, the design may have been confounded by deposition of sediment in the cages (due to declining stream discharge) which reduced population densities of colonizers. 5. This study documents changes in invertebrate abundance and morphospecies richness in response periphyton and detritus standing stocks within patches. Summation of such responses may account for observed variations in benthic communities among Hong Kong streams which differ in the extent of shading by riparian vegetation.     

The relative importance of shading and nutrients on algal production in subtropical streams

1. We used artificial substrata in forested and open streams in South-East Queensland, Australia, to determine the relative importance of shading from riparian vegetation and of nutrients on periphyton growth, and whether nitrogen and/or phosphorus limited algal productivity.
2. Nutrient-diffusing substrata consisting of agar enriched with N, P and N + P, and controls without nutrients, were deployed in duplicate at 15 sites in headwater streams with riparian canopy cover ranging from 0 to 88%.
3. Shading was the over-riding factor controlling periphyton biomass accrual on the artificial substrata, with nutrients playing a relatively minor role. Microscopic examination of periphyton scrapings taken after 7 weeks revealed that diatoms dominated on the artificial substrata in shaded streams, whereas filamentous green algae dominated the algal assemblage in the more open canopy streams.
4. Whilst nutrients had little effect on the accrual of algal biomass compared with riparian shading, there was evidence that nitrogen, and not phosphorus, stimulated periphyton production in streams with sufficient light.     

Experimental shading alters leaf litter breakdown in streams of contrasting riparian canopy cover

1. Headwater stream ecosystems are primarily heterotrophic, with allochthonous organic matter being the dominant energy. However, sunlight indirectly influences ecosystem structure and functioning, affecting microbial and invertebrate consumers and, ultimately, leaf litter breakdown. We tested the effects of artificial shading on litter breakdown rates in an open‐canopy stream (high ambient light) and a closed‐canopy stream (low ambient light). We further examined the responses of invertebrate shredders and aquatic hyphomycetes to shading to disentangle the underlying effects of light availability on litter breakdown. 2. Litter breakdown was substantially slower for both fast‐decomposing (alder, Alnus glutinosa) and slow‐decomposing (beech, Fagus sylvatica) leaf litters in artificially shaded stream reaches relative to control (no artificial shading) reaches, regardless of stream type (open or closed canopy). 3. Shredder densities were higher on A. glutinosa than on F. sylvatica litter, and shading had a greater effect on reducing shredder densities associated with A. glutinosa than those associated with F. sylvatica litter in both stream types. Fungal biomass was also negatively affected by shading. Results suggest that the effects of light availability on litter breakdown rates are mediated by resource quality and consumer density. 4. Results from feeding experiments, where A. glutinosa litter incubated under ambient light or artificial shade was offered to the shredder Gammarus fossarum, suggest that experimental shading and riparian canopy openness influenced litter palatability interactively. Rates of litter consumption by G. fossarum were decreased by experimental shading in the open‐canopy stream only. 5. The results suggest that even small variations in light availability in streams can mediate substantial within‐stream heterogeneity in litter breakdown. This study provides further evidence that changes in riparian vegetation, and thus light availability, influence organic matter processing in heterotrophic stream ecosystems through multiple trophic levels.     

NUTRIENT AND LIGHT LIMITATION OF ALGAE IN TWO NORTHERN CALIFORNIA STREAMS1

Nutrient-diffusing subsrates were used to investigate nutrient limitation of attached algal assemblages in a shaded stream and an unshaded stream in northern California. Water from both streams contained low levels of nitrogen (< 14 μg.L^?1) and very low N:P ratios (< 2). After 31 days of colonization and growth, attached algal biomass on nitrate-diffusing substrates was significantly greater than on control substrates in the unshaded stream. Nitrate-diffusing substrates also supported larger numbers of grazing insects in the unshaded stream. The prostrate diatoms Achnanthes lanceolata Bréb. and Coconeis placentula Ehr. displayed the most consistent positive responses to nitrate enrichment. Nutrient enrichment did not increase the accrual of algal biomass in the shaded stream, but algal biomass was significantly greater at sites located under openings in the tree canopy, implicating light as a limiting factor in this stream. Several Navicula and Nitzschia species, and one unidentified Gomphonema species, were positively associated with higher light levels in the shaded stream. Shade appears to be the primary factor limiting algal growth in small northern California streams, but when its effect is reduced by logging, the inherently low levels of nitrogen in these streams can become limiting.     

Experimental increases and reductions of light to streams: effects on periphyton and macroinvertebrate assemblages in a coniferous forest landscape

Increased light reaching streams as a result of riparian vegetation management is often thought to be responsible for enhanced algal productivity. However, concomitant changes in nutrients and other physical processes confound that interpretation. We manipulated light in two separate experiments to test the role of light as a controlling factor for periphyton productivity and biomass, and to observe invertebrate responses in small streams in central British Columbia, Canada. We did this by adding artificial light to reaches of three forested streams, and in a second experiment we used shadecloth to cover reaches of two streams flowing through clearcuts. Periphyton growth, productivity and composition, and macroinvertebrate benthic densities were contrasted with control reaches within the same streams. Gross primary production (GPP) was increased at least 31% by light addition to forested streams. Periphyton biomass was higher under light additions, but only significantly so in one of the streams. In one stream grazers increased along with the periphyton response, whilst in the other two lit streams invertebrates, including grazers, decreased with increased light. The shading significantly reduced GPP to about 11% of that in clearcut sections, but failed to produce any significant responses in either periphyton standing stock or invertebrates in the clearcut streams. Measures of algal production and biomass responded as predicted; however, invertebrate responses to increased and decreased light were idiosyncratic amongst streams, perhaps indicating lagged responses and limitation by other resources.     

Regulation of algal biomass in a small lowland stream: field experiments on the role of invertebrate grazing, phosphorus and irradiance

1. Field experiments were undertaken in a small Danish lowland stream to study the role of invertebrate grazing, phosphorus concentration and irradiance in the regulation of benthic algal biomass on stones. 2. Algal biomass was regulated by invertebrate grazing. The gastropod Ancylus fluviatilis prevented algal biomass build up in early spring at a density of about 900 ind. m^–2, and reduced algal biomass to very low levels during the algal growth period at a density of about 6000 ind. m^–2. Grazing pressure therefore might regulate the magnitude of peak algal biomass in Gelbæk stream, a finding in agreement with earlier field observations. 3. As phosphorus enrichment to around 152 μg l^–1 during the period when irradiance was not limiting did not result in any further increase in algal biomass, it can be concluded that algal growth in spring was not controlled by phosphorus limitation. 4. Algal biomass development differed significantly in a shaded and in a non-shaded reach of the stream. Only very low algal biomass accumulation was evident in the shaded reach, while in the non-shaded reach an algal biomass peak of about 800 mg chlorophyll m^–2 was evident after 6 weeks of colonization. The shaded reach represented the light condition in Gelbæk stream after leaf proliferation of the overhead canopy and high bank vegetation. The findings therefore suggest that irradiance available to the algal community after shading from riparian vegetation prevents further algal biomass increase and hence determines the timing of peak algal biomass in the stream. 5. The irradiance experiment also suggests that if not regulated by invertebrate grazing or shading by an overhead canopy in the summer, then the magnitude of peak biomass in Gelbæk stream will be regulated by self-shading in the algal community. 6. These field experiments support theories, derived from laboratory experiments and field studies, that regulation of algal biomass is a complex interaction of top-down and bottom-up mechanisms.     

Evidence for the use of non-detrital dissolved organic matter by microheterotrophs on plant detritus in a woodland stream

SUMMARY 1. Recent studies provide evidence for the use of exudates from living plants by epilithic microheterotrophs in streams. This study investigated the possible use of such non-detrital sources of dissolved organic matter (DOM) by streatn microheterotrophs colonizing leaf litter. Biomass of bacteria and of fungi accumulating in situ on autumn-shed leaves in flow-through troughs from which light was excluded was compared to that accumulating on leaves in troughs open to natural illumination.
2. In experiments repeated at different times of year and in different stream sections, greater biomass of microheterotrophs consistently accumulated on the leaf detritus in troughs open to natural illumination. Differences in water temperature or in grazing of leaf surfaces by macroinvertebrates could not account for these consistent differences. Further, greater microheterotroph biomass accumulated on light- and dark-incubated leaves in a stream section relatively open to sunlight, compared to corresponding leaves in a section heavily shaded by canopy and understorey vegetation.
3. These and other results suggest that, to some yet undetermined extent, detritus-associated microheterotrophs use non-detrital DOM. This conclusion is consistent with a priori predictions based on consideration of microbial energetics involved in the use of detrital versus non-detrital DOM.
4. Studies of trophic pathways in streams and other aquatic habitats have failed to assess some potentially important sources of non-detrital DOM. The ability of available techniques to assess the relative roles of detrital and non-detrital sources of DOM is evaluated, and alternative approaches to this problem are suggested.     

Herbs and grasses as an allochthonous resource in open-canopy headwater streams

1. The organic matter dynamics of streams dominated by herbs and grass on their banks are poorly understood, despite the fact that such streams are common worldwide. Further, herbs and grasses can provide large quantities of detritus to stream food webs, and particularly small streams can be heavily shaded by overhanging vegetation, perhaps limiting in‐stream primary production. 2. We quantified the standing crop of edge vegetation and associated macroinvertebrate communities along three headwater streams with herbaceous and grass riparian vegetation on agricultural land in the Piedmont of Maryland, U.S.A., measured the decomposition of four common species of herbs and grasses using experimental leaf packs, and removed edge vegetation experimentally to determine the effect of shading on benthic algal production. 3. Large standing crops of plant material (average range: 68–276 g ash‐free dry mass per m⁻²), composed largely of monocotyledons, were found at all three study streams. These values are similar to those for coarse particulate organic matter in deciduous forested streams in the eastern U.S.A. In addition, diverse assemblages of shredding macroinvertebrates were observed at all three study sites. 4. Decomposition of the herbs was faster than that of the grasses, and both decomposed faster than most deciduous tree leaf litter. The decomposition rates of the herbs and grasses were significantly related to leaf quality as measured by leaf nitrogen content. Macroinvertebrate shredders colonized all experimental leaf packs, and the colonization of the herbs was faster than that of the grasses. 5. The accrual of chlorophyll‐a after the removal of shading vegetation was faster than that measured prior to removal as well as that in an unmanipulated control reach. 6. Given that the standing crop of organic matter in streams with herbs and grass along their banks was similar to that in forested streams, that the organic matter was rich in nitrogen and used by detritivores, and riparian shading limited algal growth, we suggest that herbaceous and grass plant material may be an important allochthonous food resource in such systems.     

Controlled release experiments to determine the effects of shade and plants on nutrient retention in a lowland stream

Understanding nutrient uptake and retention in streams remains an important challenge for lotic scientists. In this study a series of pulse and continuous releases of dissolved nutrients were made to shaded and unshaded (reference) reaches of a small lowland stream to determine whether suppression of macrophyte growth by riparian shade impaired nutrient retention. The nutrients were dissolved reactive phosphorus (DRP), total ammoniacal nitrogen (NH₄–N) and nitrate nitrogen (NO₃–N). Nutrient reductions ranged from 100% of DRP when stream water was anoxic, to 5–10% for NH₄–N and NO₃–N in the reference reach. Nutrient removals were affected by travel times in each reach. Percentage removals of NH₄–N (46 ± 10) and NO₃–N (52 ± 14) were higher in the shaded reach than in the swifter moving reference reach (15 ± 8 and 16 ± 10, respectively). DRP (%) removals were 75± 7 and 57 ± 12 for the shaded and reference reaches, respectively. The presence of emergent marginal macrophytes (Persicaria hydropiper) increased stream velocity in the reference reach by reducing the effective channel cross-section area. Shading reduced plant biomass, increased the channel cross-section and lowered velocity in the experimental reach, effecting dramatic reductions in nutrient concentrations over short distances. The opposite effect is more typical for larger, swifter streams having dense stands of submerged macrophytes, where lowering channel plant biomass will cause increased velocities and lower relative nutrient losses. Riparian shade does not necessarily impair nutrient uptake from small streams. Where invasive marginal species such as P. hydropiper dominate headwater streams shade may be beneficial to the protection of downstream waters from eutrophication. Where reduction of nutrient fluxes from small streams is a key objective for protection of downstream waters, active management of streams should seek to increase travel times, allowing greater potential for nutrient uptake. This will need to be weighed against the need for effective drainage in pastoral areas where reduced travel times are usually sought.     

Macroinvertebrate Responses to Algal and Bacterial Manipulations in Streams

Our study was designed to assess the relative importance of algae and bacteria as sources of energy for stream macroinvertebrates. In one experiment, we manipulated algae by artificially shading six sections in each of two streams, one stream with an open canopy (clear-cut drainage basin) and the other with a closed canopy (forested drainage basin); both streams were in Hubbard Brook Experimental Forest, New Hampshire, USA. Chlorophyll a concentrations were reduced from 0.2 to 0.05 μg/cm² in artificially shaded sections of both streams. However, macroinvertebrates showed no response to these algal manipulations in either the clear-cut or forested stream. Nutrient concentrations (N and P) were low and limiting to primary production in both the clear-cut and forested streams. Additionally, both streams had relatively low macroinvertebrate densities suggesting bottom-up controls were important in macroinvertebrate abundance. However, the forested stream did have higher macroinvertebrate densities presumably because of higher inputs of coarse particulate organic matter from the riparian vegetation. In a second experiment, in Augusta Creek, Michigan, we manipulated both algae and bacteria. To reduce algae, we artificially shaded experimental stream channels so that chlorophyll a was reduced from natural levels of 3.0–5.6 to 0.4–0.7 μg/cm². Half of the shaded channels had dissolved organic carbon (DOC – sucrose) dripped into them to raise DOC levels by 2–3 mg/l and thus stimulate bacterial abundance. Open channels, with higher algal abundance, had higher densities of Ephemerella, but only in November when nymphs were larger. Channels with increased DOC had higher bacterial abundances, higher densities of Chironomidae and lower densities of Heptageniidae. Several other macroinvertebrate taxa that were at relatively low abundance in our samples showed no significant response to these manipulations. Our results suggest that early instar Ephemerella may not rely as heavily on algae as later instars. Also, certain taxa were able to use the heterotrophic microbial community, especially chironomids which increased in numbers when bacterial density increased; thus, the bacterial carbon source may be more important to some stream macroinvertebrates than previous studies have suggested.     

Effect of Permanent Shading on Adult Size and Biomass of the Caddisfly <Emphasis Type="Italic">Lype phaeopa</Emphasis> (Stephens, 1836) (Psychomyiidae)

To investigate the effect of shading on the development of aquatic insect larvae, estimated by the size and biomass of freshly emerged adults, a laboratory experiment was conducted in which larvae of the psychomyiid caddisfly Lype phaeopa were reared in glass aquaria covered by gauze pervious to light and compared to aquaria which were covered by a black textile distinctly reducing the light irradiation. Seven submerged branches colonized by larvae of L. phaeopa were collected from a small lowland stream, cut in two equally sized halves and one of the halves was reared under natural light and the other one under shaded condition. The hypothesis of the study was that larvae reared under natural light conditions can develop faster resulting in earlier emergence and larger adults compared to larvae reared under a shaded conditions. Females were always significantly larger and heavier than simultaneously emergent males (paired t-test, p < 0.001) independent of the treatment. The experiment revealed no effects of the shading on the adult size and biomass of emergent adults, neither for males (t-test, p = 0.197) nor for females (p = 0.303). Adult biomass was influenced by seasonal temperature (smaller adults during the warm summer months). Females’ biomass was also different between the branches independent from the treatment (one way ANOVA, p = 0.002) whereas males showed only a trend (p = 0.055). The shading displayed a significant effect on the emergence date of males with earlier emergence in the light treatment compared to the dark treatment (t-test, p = 0.046), but this could be not ascertained for females (p = 0.828). The experiment indicated that shading has no significant effect on the adult biomass of L.␣phaeopa, but it could delay the time of emergence in males. Main driving factors of larval development and subsequent adult biomass in L. phaeopa were temperature and habitat quality.     

Microhabitat availability in Welsh moorland and forest streams as a determinant of macroinvertebrate distribution

SUMMARY. 1 Eighteen streams in mid-Wales were sampled for macro-invertebrates in both riffles and margins in April 1985–87. Stream macro-flora, substrata and marginal habitats were surveyed in May 1988.
2. TWINSPAN classification of the macroinvertebrate data indicated three major stream groups. One was distinguished by circumneutral pH and had a flora and fauna typical of such conditions. The other two groups consisted of acidic streams with moorland and conifer afforested catchments respectively. The forest streams were the more acidic but the two groups also differed significantly in the composition of their marginal habitats.
3 The acidic moorland streams had more vegetation ('soft' features) in the margins and supported several invertebrate taxa which were relative more abundant there than in the riffles. These taxa may be excluded from forest streams because the margins are 'hard' due to greater erosiveness and shading.
4. In view of the increasing cover by conifer afforestation in Britain, it is clearly necessary to elucidate all its effects on stream ecosystems, which include changes to the physical environment.     

Plant distribution and abundance in relation to physical conditions and location within Danish stream systems

The distribution of obligate submerged plants, amphibious plants and terrestrial plants in streams was examined in relation to water depth, substrate and distance to the banks using univariate and mulitivariate analyses. The analyses were based on recordings in more than 40000 quadrats (25×25 cm) in 208 unshaded sites in the predominantly small and shallow (<1.6 m) Danish streams. Also, the distribution of plant abundance and richness from the source to the outlet of the stream system was determined.The submerged plants in Danish streams include 87 terrestrial, 22 amphibious and 30 obligate submerged taxa. The distribution of plant types was mainly related to water depth and distance to the bank among the physical conditions, while the type of bottom substrate had no significant influence. Terrestrial plants and amphibious plants (excluding Sparganium emersum) dominated in shallow water near the bank, but declined rapidly with increasing depth and distance to the bank, reflecting the importance of dispersal by ingrowth from populations on the banks to the water among these plants. Accordingly, these two plant groups constituted a higher proportion of total plant abundance in small streams than large streams. S. emersum dominated on great depth and distance to the banks, probably reflecting the lengthwise dispersal of this species from upstream to downstream parts of the stream system, the tolerance of the species to weed cutting and the adaptation to grow at low light intensities. Obligate submerged plants dominated at intermediate depths and at all distances from the bank except from 0 to 50 cm. This distribution reflects the ability of these species to disperse lengthwise in streams and live permanently submerged.The species number of all species and obligate submerged plants was lower in the smallest stream sites compared to larger downstream sites, while there was no difference for terrestrial and amphibious plants. The downstream increase of submerged species can be explained by the increase of habitat area and the dispersal of plants with the current, implying that the species pool accumulates with distance from the source. This result is at variance with a maximum richness of submerged plant species predicted for intermediate-sized streams according to the River Continuum Concept developed for large North American streams having forested shallow upstream sections and unshaded, deep downstream sections both unsuitable to submerged plant growth. The results for Danish streams imply that both the longitudinal connection through the flowing water and the transversal connection to the local species on the banks are important for plant distribution in the streams.     

Responses of invertebrates and algae of a boreal coniferous forest stream to experimental manipulation of leaf litter inputs and shading

Forest harvesting alters leaf litter inputs and shading of small streams. Most of the previous studies of harvesting effects are limited to coastal or deciduous forests, so here we consider a sub-boreal forest stream. To test the hypothesis that changes in light and litter inputs would affect the benthic community in these streams, we experimentally manipulated these variables in stream mesocosms. We used a 2 × 2 factorial design with light (shaded or full light) and leaf litter inputs (equivalent to a forested stream or one quarter that rate) as factors. The high leaf litter treatment resulted in differences in macroinvertebrate community composition and higher densities of two shredders, Limnephilus sp. and Podmosta sp., suggesting food limitation. Algal filaments were longer in the high light treatments indicating a change in periphyton composition. There were no significant differences in chlorophyll a or ash-free dry mass, suggesting that light was not limiting to periphyton. The community structure clearly shifted in response to both resources, although primarily to detrital inputs. These results provide evidence that changes to shading and leaf inputs to small streams can affect the benthos and may limit secondary production.     

Watershed Land Use and Seasonal Variation Constrain the Influence of Riparian Canopy Cover on Stream Ecosystem Metabolism

Ecosystem metabolism is an important determinant of trophic structure, nutrient cycling, and other critical ecosystem processes in streams. Whereas watershed- and local-scale controls on stream metabolism have been independently investigated, little is known about how controls exerted at different scales interact to determine stream metabolic rates, particularly in urban streams and across seasons. To address this knowledge gap, we measured ecosystem metabolism in four urban and four reference streams in northern Kentucky, USA, with paired closed and open riparian canopies, during each of the four seasons. Gross primary production (GPP), ecosystem respiration, and net ecosystem production (NEP) were all best predicted by models with season as a main effect, but interactions between season, canopy, and watershed varied for each response. Urban streams exhibited higher GPP during most seasons, likely due to elevated nutrient loads. Open canopy reaches in both urban and forested streams, supported higher rates of GPP than the closed canopy which reaches during the summer and fall, when the overhead vegetation shaded the closed reaches. The effect of canopy cover on GPP was similar among urban and forested streams. The combination of watershed and local-scale controls resulted in urban streams that alternated between net heterotrophy (NEP <0) and net autotrophy (NEP >0) at the reach-scale during seasons with dense canopy cover. This finding has management relevance because net production can lead to accumulation of algal biomass and associated issues like nighttime hypoxia. Our study suggests that although watershed urbanization fundamentally alters ecosystem function, the preservation and restoration of canopied riparian zones can provide an important management tool at the local scale, with the strongest impacts on stream metabolism during summer.     

Shade avoidance in Trifolium repens: costs and benefits of plasticity in petiole length and leaf size

We tested whether the degree of shade-induced plasticity in petiole length and leaf area is related to the mean trait value expressed under high-light conditions, and to what extent trait values expressed under high-light and shaded conditions affect plant performance. Thirty-four Trifolium repens genotypes were used with a wide range of petiole lengths and leaf areas. Plants were subjected to a high-light environment and two shading regimes: homogeneous shading and a vertical light gradient. Absolute petiole elongation in response to both shading treatments and absolute leaf area expansion in response to homogeneous shading were independent of the trait values expressed in high light. Consequently, relative plasticity was higher for genotypes with lower high-light trait values. Plasticity was associated with enhanced plant performance in a vertical light gradient but not in homogeneously shaded conditions. We also found costs associated with the ability to express plasticity. Our results suggest that selection can act separately on trait values expressed under high-light conditions and on the degree of plasticity.     

Historical changes in species composition and richness accompanying perturbation and eutrophication of Danish lowland streams over 100 years

1. European lowland streams have experienced increased perturbation and eutrophication during the past 100 years. We use archive information from 27 Danish stream sites around 1896 and new data from 208 stream sites in 1996 to evaluate the accompanying changes in stream vegetation. Among the stream sites, 13 were both studied in 1896 and 1996. 2. The species richness of submerged plants has declined profoundly over the 100-year period, particularly among the large group of Potamogeton species. This is evident both from the direct comparison of the 13 stream reaches included in both studies, and from the general comparison of all stream reaches included in the two studies. 3. The Potamogeton vegetation has become less diverse, and is now dominated by species resistant to frequent disturbance through a high dispersal capacity. Potamogeton species, adapted to eutrophic conditions, have also increased relative to species more typical of oligotrophic conditions over this period. The dominant submerged species in the contemporary stream vegetation generally show a high capacity for dispersal and regrowth of detached shoots. 4. The decline of species richness in Danish streams can be partly explained by a decline in the species richness in lakes in the stream system. The rich vegetation observed downstream of lakes in the past has mostly disappeared due to loss of the vegetation in the now eutrophic lakes and increasing turbidity downstream. 5. The overall decline in richness, and the directional change in dominance patterns among stream species, can thus be explained by the loss of suitable habitats and the strong anthropogenic impacts, which have driven several European aquatic species close to extinction.     

Phytoplankton and Epiphyte Development and Their Shading Effect on Submerged Macrophytes in Lakes of Different Nutrient Status

The annual mean light intensity at the depth limit of the Littorella vegetation was 24–33% of the subsurface light intensity, despite large variations in each attenuation component (lake water, phytoplankton, and epiphytes). In oligotrophic, silicate-poor lakes, the light attenuation above the submerged vegetation was dominated by the water itself, which accounted for 65–72% of the total attenuation. Phytoplankton and epiphytes were equal in importance to each other. In oligotrophic, silicate-rich lakes and lakes receiving a nitrogen supply above background level, the epiphytes were more abundant, accounting for about 50% of the light attenuation. In one lake with a high nutrient supply, the epiphytes were responsible for 86% of the light attenuation. A new method of measuring the effect of shading by the epiphytic community on submerged macrophytes is presented. The light attenuation caused by the phytoplankton and the epiphytes was investigated and related to the depth distribution of the submerged angiosperm, Littorella uniflora. It is shown that the biomass of the epiphytes increased more than the biomass of the phytoplankton in response to an external or internal nutrient loading. Shading by epiphytes is of decisive importance for the depth distribution of Littorella at increasing nutrient supply.