Relative importance of periphyton and phytoplankton in turbid and clear vegetated shallow lakes from the Pampa Plain (Argentina): a comparative experimental study

We analyzed experimentally the relative contribution of phytoplankton and periphyton in two shallow lakes from the Pampa Plain (Argentina) that represent opposite scenarios according to the alternative states hypothesis for shallow lakes: a clear lake with submerged macrophytes, and a turbid lake with high phytoplankton biomass. To study the temporal changes of both microalgal communities under such contrasting conditions, we placed enclosures in the littoral zone of each lake, including natural phytoplankton and artificial substrata, half previously colonized by periphyton until a mature stage and half clean to analyze periphyton colonization. In the clear vegetated shallow lake, periphyton chlorophyll a concentrations were 3–6 times higher than those of the phytoplankton community. In contrast, phytoplankton chlorophyll a concentrations were 76–1,325 times higher than those of periphyton in the turbid lake. Here, under light limitation conditions, the colonization of the periphyton was significantly lower than in the clear lake. Our results indicate that in turbid shallow lakes, the light limitation caused by phytoplankton determines a low periphyton biomass dominated by heterotrophic components. In clear vegetated shallow lakes, where nitrogen limitation probably occurs, periphyton may develop higher biomass, most likely due to their higher efficiency in nutrient recycling.     

Community–habitat relationships in coastal streams in Big Sur, California, USA: travertine influences macroinvertebrate abundance and community structure

Travertine deposition occurs in streams worldwide but its effects on stream communities are poorly understood. I sampled benthic macroinvertebrates, periphyton, and reach-scale environmental variables in coastal streams in Big Sur, central California, USA, to determine the specific effects of travertine that occurred at some sites as well as to provide a broader assessment of community–habitat relationships. Total density and biomass of macroinvertebrates varied 6- and 9-fold across sites, respectively, and chlorophyll a concentrations varied 10-fold, but invertebrate and periphyton abundances were not correlated. Baetis tricaudatus (Ephemeroptera), Simuliidae (Diptera), and Chironomidae (Diptera) dominated macroinvertebrate communities across all sites, although differences in the relative abundances of these and other taxa produced moderate variation in community structure among sites (Bray-Curtis similarity coefficients of 47–84). Variation in community structure was related to a number of habitat features, notably travertine but also including variables reflecting channel morphology, flow, substrate size, and riparian tree type. Median density and biomass of macroinvertebrates were more than twice as high at sites without travertine than sites with travertine. Taxa richness also was higher at sites without travertine, and community structure differed moderately between sites with and without travertine, although there were no particular assemblages associated with either group. Non-metric multidimensional scaling (MDS) and cluster analysis of similarities in community structure appeared to separate sites with either travertine or high fines from sites without those conditions. These results demonstrate that travertine can have strong effects on stream communities, and additional studies are needed to identify the full range of effects on ecosystems and to evaluate the potential consequences of travertine for conservation efforts such as biomonitoring programs and threatened species management. Handling editor: R. Bailey     

Seasonal shifts in the importance of bottom–up and top–down factors on stream periphyton community structure

We examined the importance of temporal variability in top–down and bottom–up effects on the accumulation of stream periphyton, which are complex associations of autotrophic and heterotrophic microorganisms. Periphyton contributes to primary production and nutrient cycling and serves as a food resource for herbivores (grazers). Periphyton growth is often limited by the availability of nitrogen and phosphorus, and biomass can be controlled by grazers. In this study we experimentally manipulated nutrients and grazers simultaneously to determine the relative contribution of bottom–up and top–down controls on periphyton over time. We used nutrient diffusing substrates to regulate nutrient concentrations and an underwater electric field to exclude grazing insects in three sequential 16–17 day experiments from August to October in montane Colorado, USA. We measured algal biomass, periphyton organic mass, and algal community composition in each experiment and determined densities of streambed insect species, including grazers. Phosphorus was the primary limiting nutrient for algal biomass, but it did not influence periphyton organic mass across all experiments. Effects of nutrient additions on algal biomass and community composition decreased between August and October. Grazed substrates supported reduced periphyton biomass only in the first experiment, corresponding to high benthic abundances of a dominant mayfly grazer (Rhithrogena spp.). Grazed substrates in the first experiment also showed altered algal community composition with reduced diatom relative abundances, presumably in response to selective grazing. We showed that top–down grazing effects were strongest in late summer when grazers were abundant. The effects of phosphorus additions on algal biomass likely decreased over time because temperature became more limiting to growth than nutrients, and because reduced current velocity decreased nutrient uptake rates. These results suggest that investigators should proceed with caution when extending findings based on short‐term experiments. Furthermore, these results support the need for additional seasonal‐scale field research in stream ecology.     

空间层次及人工基质对着生藻类建群特性的影响

为全面了解着生藻类在建群中群落变化的生态学特性,揭示着生藻类的建群规律,在以丝状藻类为优势藻的生态塘中,采用花岗岩和瓷砖为附着材料,设置水体底部和中部为附着位点,进行频次为10d的采样分析。结果表明,生态塘中共检出8门73属117种着生藻类,其中以硅藻、蓝藻、绿藻为优势类群。同时不同人工基质和不同空间层次条件下着生藻类的建群特征较一致,早期以单细胞硅藻如舟形藻(Navicula sp.)、脆杆藻(Fragilaria sp.)、曲壳藻(Achnanthes sp.)等为优势,后期以丝状藻类如鞘丝藻(Lyngbya sp.)、颤藻(Oscillatoria sp.)、伪鱼腥藻(Pseudanabaena sp.)等为优势;研究结果发现不同人工基质(花岗岩和瓷砖)对着生藻类的种类组成、细胞密度、生物量和藻类多样性无显著影响,花岗岩和瓷砖上附着的着生藻类具有较高的相似性;但不同的空间层次对着生藻类建群特征影响明显,水体底部具有更多的硅藻种类数,中部具有更多的绿藻,随着建群时间的发展,蓝藻比例不断增加;就生物量而言,底部的着生藻类叶绿素a显著高于水体中部,但两者的细胞密度无显著性差异;随着建群过程的发展,水体底部的着生藻类生物量达峰值所需的时间比中部更长。通过相关性分析,生态塘中着生藻类的生长主要受总磷的影响。     

Experimental Analysis of Grazing by the Mayfly Meridialaris chiloeensis on Different Successional Stages of Stream Periphyton

In this study we determined grazing effects of the South Andean endemic mayfly Meridialaris chiloeensis on periphyton at different stages of successional development. Grazing effects were studied through a two‐factor experimental design (colonization stages X grazer density) in a stream‐side channel in spring and winter. Our results showed an absence of proportionality between grazer density and periphyton decline in response to grazers at low and intermediate levels of periphytic biomass; however, when periphyton biomass was high a direct inverse relationship was observed between post‐grazing biomass and grazer density. The relationship between periphytic algae (chlorophyll a concentration) and periphyton (total periphytic ash‐free dry mass) (C/OM index) was used as an estimation of the autotrophic fraction in the total periphyton matrix. Grazing did not alter the C/OM index indicating that both autotrophic and heterotrophic fractions of the periphyton components were reduced in the same proportion. Ordination of samples using the relative abundance of diatom species showed that herbivore effect was less evident at intermediate and late stage of colonization than at early one. These results support the statement that the outcome of the herbivore‐periphyton interaction may depend on the successional stage of the periphyton community. In spring Fragilaria pinnata relative abundance, on the basis of cell counts, was reduced by grazing and Nitzschia palea was enhanced. In the winter experiment, grazing decreased Achnanthes minutissima relative abundance. (© 2004 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Invertebrate communities associated with macrophytes in Lake Ladoga: effects of environmental factors

Littoral invertebrate communities (meio- and macrobenthos and zooplankton) were studied in seven types of macrophyte associations commonly encountered in Lake Ladoga: in reed (Phragmites) beds on sand, soft and hard bottoms, in associations with the prevalence of Potamogeton spp., Carex spp., and Equisetum fluviatile, and in diverse vegetation stands with e.g. Polygonum amphibium, Cicuta virosa, Typha latifolia and Eleocharis acicularis. Some of the studied habitats were affected by sewage pollution, others were in comparatively undisturbed areas. Statistically significant differences between invertebrate communities in the different macrophyte associations were found. In stepwise multiple linear regression analysis the following factors were identified as determinants of abundance of aquatic invertebrates in macrophyte associations: shoot density, plant dry weight biomass, periphyton biomass, periphyton chlorophyll a, periphyton primary production, and concentrations of Sr, Mg, Ca, P, Mn, Zn, Pb and Cu. Pollution was shown to have a minor effect on the composition of littoral invertebrate communities. It is not possible to determine one single principle factor responsible for the structure and density of invertebrates in macrophyte communities.     

Influence of substrate type on periphyton biomass and nutrient state at contrasting high nutrient levels in a subtropical shallow lake

How substrate affects periphyton biomass and nutrient state at different, but high, nutrient levels was tested in three large enclosures in a hypereutrophic subtropical shallow lake. We compared periphytic characteristics (1) on three hard substrates (stone, bamboo, and wood) incubated for 2 weeks and 1 year, respectively, to investigate the existence of the influences of substrate type at hypereutrophic levels, and (2) on artificial plants with contrasting (parvopotamid-like and myriophyllid-like) soft substrate morphology. In general, periphytic biomass and nutrient state were sensitive to variations in nutrient level, incubation time, hard substrate type (except 2-week incubated) and substrate morphology, but to a varying extent. The periphyton nutrient content increased with increasing nutrient levels on most substrates. Long-time incubated substrates supported more periphytic biomass, had a higher nutrient content and autotrophic proportion, while the effect of nutrient level on nutrient content in the periphyton was independent of incubation time. The effects of hard substrate type on periphyton characteristics were much weaker than those of nutrient level. By contrast, the effects of soft substrate morphology on periphyton biomass and carbon: nutrient ratios surpassed those of nutrient level. Chlorophyll a, dry mass, and ash free dry mass were much higher on parvopotamid than on myriophyllid substrates. Our results show that periphyton biomass and nutrient state are influenced by both substrate and nutrient level even in hypereutrophic lakes, which might have cascading effects on the benthic food web.     

DISTURBANCE OF STREAM PERIPHYTON BY PERTURBATIONS IN SHEAR STRESS: TIME TO STRUCTURAL FAILURE AND DIFFERENCES IN COMMUNITY RESISTANCE1

The resistance of stream periphyton to structural disturbance by increases in shear stress (simulating a spate) was investigated in a laboratory flow tank. We monitored loss of biomass from a filamentous community (dominated by Melosira varians) under four different levels of shear stress. In each case, any loss that was going to occur did so within 10 min for this community. In a second experiment, we tested the resistance of four different communities (two dominated by nonfilamentous diatoms and two dominated by filamentous green algae/diatoms) to increases in shear stress. Nine different levels of shear stress were used, ranging from 1- to 70-fold higher than the conditions to which the communities were acclimated. All communities were 14 days old, but some differences in initial biomass occurred that influenced the degree of resistance independently of species composition. Overall, the nonfilamentous diatom communities were the most resistant, and the filamentous communities were the least resistant. The kinetics of the sloughing process varied among community types, with a community dominated by Melosira varians/Gom-phonema parvulum losing 50% of its biomass with only a 3-fold increase in shear stress. In contrast, a community dominated by the nonfilamentous diatoms Fragilaria vaucheriae/Cymbella minuta lost <50% of its biomass after a 70-fold increase in shear stress. Shear stresses required for 50% loss of biomass for the different communities were as follows: 3.6 Newtons.m^?2 for the Melosira varians/Gomphonema parvulum community, 10.0 N.m^?2 for the Spirogyra sp./Gomphoneis her-culeana/Ulothrix zonata community, 50.6 N.m^?2 for the Fragilaria construens/Cymbella minuta/Ach-nanthes minutissima community, and >90.0 N.m^?2for the Fragilaria vaucheriae/Cymbella minuta community. These results show that spates without bedload movement can potentially have widely differing disturbance effects on periphyton loss among streams depending on the initial taxonomic composition of resident communities. These results have important implications for stream ecosystem analysis and modeling.     

Cadmium and lead toxicity on tropical freshwater periphyton communities under laboratory-based mesocosm experiments

Periphyton constitutes an important community that is useful for assessment of ecological conditions in lotic systems. The objective of this study was to assess the effects of different mixtures of Cd and Pb on periphyton growth as well as Cd and Pb mixtures toxicity to diatom assemblages in laboratory mesocosm experiments. A natural periphyton community sampled from the Monjolinho River (South of Brazil) was inoculated into five experimental systems containing clean glass substrates for periphyton colonization. The communities were exposed to mixtures of dissolved Cd and Pb concentrations of 0.01 and 0.1 mg l⁻¹ Cd and 0.033 and 0.1 mg l⁻¹ Pb. Periphyton ash-free dry weight, growth rate, diatom cell density and diatom community composition were analyzed on samples collected after 1, 2 and 3 weeks of colonization. High Cd concentration (0.1 mg l⁻¹) has negative effects on periphyton growth while high concentration of Pb (0.1 mg l⁻¹) decreased the toxic effects of Cd on periphyton growth. Shifts in species composition (development of more resistant species like Achnanthidium minutissimum and reduction of sensitive ones like Cymbopleura naviculiformis, Fragilaria capucina, Navicula cryptocephala, Encyonema silesiacum, Eunotia bilunaris, and Gomphonema parvulum), decreases in species diversity of diatom communities with increasing Cd and Pb concentrations and exposure duration have been demonstrated in this study making diatom communities appropriate monitors of metal mixtures in aquatic systems.     

Effect of periphyton biomass on hydraulic characteristics and nutrient cycling in streams

The effect of periphyton biomass on hydraulic characteristics and nutrient cycling was studied in laboratory streams with and without snail herbivores. Hydraulic characteristics, such as average water velocity, dispersion coefficients, and relative volume of transient storage zones (zones of stationary water), were quantified by performing short-term injections of a conservative tracer and fitting an advection-dispersion model to the conservative tracer concentration profile downstream from the injection site. Nutrient cycling was quantified by measuring two indices: (1) uptake rate of phosphorus from stream water normalized to gross primary production (GPP), a surrogate measure of total P demand, and (2) turnover rate of phosphorus in the periphyton matrix. These measures indicate the importance of internal cycling (within the periphyton matrix) in meeting the P demands of periphyton. Dense growths of filamentous diatoms and blue-green algae accumulated in the streams with no snails (high-biomass streams), whereas the periphyton communities in streams with snails consisted almost entirely of a thin layer of basal cells of Stigeoclonium sp. (low-biomass streams). Dispersion coefficients were significantly greater and transient storage zones were significantly larger in the high-biomass streams compared to the low-biomass streams. Rates of GPP-normalized P uptake from water and rates of P turnover in periphyton were significantly lower in high biomass than in low biomass periphyton communities, suggesting that a greater fraction of the P demand was met by recycling in the high biomass communities. Increases in streamwater P concentration significantly increased GPP-normalized P uptake in high biomass communities, suggesting diffusion limitation of nutrient transfer from stream water to algal cells in these communities. Our results demonstrate that accumulations of periphyton biomass can alter the hydraulic characteristics of streams, particularly by increasing transient storage zones, and can increase internal nutrient cycling. They suggest a close coupling of hydraulic characteristics and nutrient cycling processes in stream ecosystems.     

THE RELATIONSHIP OF LIPIDS WITH LIGHT AND CHLOROPHYLL MEASUREMENTS IN FRESHWATER ALGAE AND PERIPHYTON1

Lipid content and lipid class composition were determined in stream periphyton and the filamentous green algae Cladophora sp. and Spirogyra sp, Sterols and phospholipids were compared to chlorophyll a (chl a) as predictors of biomass for stream periphyton and algae. Chlorophyll a, phospholipids, and sterols were each highly correlated with ash-free dry mass (AFDM) (r² > 0.98). Stream periphyton exposed naturally to high light (HL) and low light (LL) had chl a concentrations (μg chl a-mg^?1AFDM) of 7.9± 0.7 and 12.4 ± 2.9, respectively, while the sterol concentrations of these HL and LL stream periphyton (1.6 ± 0.4) were not significantly different (P > 0.05). Periphyton exposed to an irradiance of 300 μmol photons·m^?2s^?1 in the laboratory for 60 h had 5.6 ± 0.55 μg chl a·mg^?1 AFDM, but the same periphyton exposed to 2% incident light for the same amount of time had 11.0 ± 0.56 μg chl a·mg^?1 AFDM. Sterol concentrations in these periphyton communities remained unchanged (1.5 ± 0.3 μg·mg^?1AFDM), Similar results (i.e. changes in chl a but stability of sterol concentrations in response to irradiance changes) were also found for Cladophora and Spirogyra in laboratory experiments. Sterols can be quantified rapidly from a few milligrams of algae and appear to be a useful predictor of eukaryote biomass, whereas cellular levels of chl a vary substantially with light conditions. Phospholipids (or phospholipid fatty acids) are considered to be a reliable measure of viable microbial biomass. Nevertheless, phospholipid content varied substantially and unpredictably among algae and periphyton under different light regimes. Irradiance also had a significant effect on storage lipids: HL Cladophora and HL periphyton had 2 × and 5 × greater concentrations of triacylglycerols, respectively, compared to their LL forms. HL and LL algae also differed in the concentration of several major fatty acids. These light-induced changes in algal lipids and fatty acids have important implications for grazers.     

Periphyton community responses to elevated metal concentrations in a Rocky Mountain stream

The effects of elevated metals on stream periphyton in the Eagle River, a mining impacted river in central Colorado, were assessed in 1991 and 1992 using assemblage information (taxa richness, community similarity) and non-taxonomic measures (biomass, chlorophyll a, autotrophic index). The number of periphyton genera collected ranged from 2 at a site adjacent to abandoned mining operations to 21 at a downstream site, but was not significantly correlated with dissolved metals concentrations. Fragilaria and Achnanthes were the dominant genera at all sites, with Fragilaria dominating the less impacted sites and Achnanthesdominating at the more impacted sites. Taxonomic similarity was greatest among those sites receiving the greatest inputs of metals from mining operations, where the coefficient of similarity ranged from 0.87 to 0.99. Cluster analyses revealed significant differences among sites adjacent to the mine and either the upstream or downstream sites. Chlorophyll a content of periphyton and the autotrophic index in both years showed significant downstream decreases associated with increasing dissolved metals concentrations. Overall, the periphyton community data were able to separate metal contaminated sites from reference or less impacted sites, and responded in predictable ways to increasing metal concentrations of Eagle River water.     

Periphyton responds differentially to nutrients recycled in dissolved or faecal pellet form by the snail grazer Theodoxus fluviatilis

1. We aimed to separate the effects of grazers on periphyton via grazing from that of nutrient recycling from their faecal pellets. 2. We set up three different experimental treatments (snails/no snails/faecal pellets) and sampled over 16 days. The ‘snail’ treatment contained a low density (snail biomass c. 14 g^?2) of the gastropod grazer Theodoxus fluviatilis and the ‘faecal pellet’ treatment received the same amount of faecal pellets as were produced in the ‘snail’ treatment. Whereas the ‘faecal pellet’ treatment provided extra nutrients to periphyton from the faeces, the ‘snail’ treatment provided nutrients in the form of both faeces and in excreta. There was also direct grazing on periphyton in the ‘snail’ treatment. The ‘no snail’ was not grazed and received no nutrients in faeces or excreta. 3. We measured periphyton C, N and P content, chlorophyll‐a (chl‐a), primary production, bacterial biomass, bacterial production and bacterial respiratory activity. In the water column we measured dissolved inorganic N and soluble reactive P. 4. Snails increased the amount of dissolved inorganic N in the water. On day 16, the periphyton N : P ratio in the ‘faecal pellet’ treatment was lower, and periphyton P content was higher, than in the other two treatments. N : P ratios decreased over time in the ‘faecal pellet’ treatment. Primary and bacterial production were positively correlated in all treatments. 5. Algal chl‐a and primary production of periphyton per unit area and periphyton chl‐a : C ratios increased over the 16 day in the ‘snail’ treatment, and thus excretion of dissolved N by snails had a stronger positive effect on the periphyton community than N and P in faecal pellets. 6. Our data show that excretion and egestion can have different effects on periphyton, probably because of the higher proportion of dissolved N in excreta and the higher proportion of P recycled in faecal pellets. The relative effect of nutrients recycled in egesta or in excretions, probably depends on the form of nutrient limitation of the periphyton. Further, the different components of the periphyton matrix could react differently to the different forms of nutrient recycling. 7. We conclude that direct grazing effects are less important than nutrient effects when nutrients are limiting and grazing pressure is low. Further, the spatial separation of different grazing effects can lead to differences in periphyton production and nutrient stoichiometry. This might be an explanation for the patchiness of periphyton in nature.     

Indirect effects of a parasite on a benthic community: an experiment with trematodes, snails and periphyton

1. Few studies have directly addressed the role played by parasites in the structure and function of ecosystems. Parasites influence the behaviour, reproduction and overall fitness of their hosts, but have been usually overlooked in community and ecosystem‐level studies. We investigated the effects of trematode parasites on snail–periphyton interactions. 2. Physa  acuta (Gastropoda: Pulmonata) snails infected with the trematode Posthodiplostomum minimum (often >30% of within‐shell biomass) grazed more rapidly than uninfected snails. Trematode effects on snail grazing indirectly affected the standing stock and community structure of periphyton. Populations of snails with 50% infected individuals reduced algal biomass by 20% more than populations with lesser (10% or 0%) infection rates. 3. The alga Cladophora glomerata dominated periphyton communities grazed by snail populations with 50% infection rates, whereas diatoms and blue–green algal taxa dominated when grazed by snail populations with lower infection rates. 4. Thus, trematodes indirectly affected periphyton communities by altering host snail behaviour, a trait‐mediated indirect effect. These results indicate that trematodes can indirectly influence benthic community structure beyond simple population fitness, with possible related effects on ecosystem function.     

Phosphate limitation influences the sensitivity to copper in periphytic algae

1. To assess the influence of nutrient limitation on copper toxicity, periphitic communities from an oligotrophic stream were exposed to copper for six to 12 days with and without the supply of nutrients (mainly P). 2. In contrast to the hypothesis that nutrient cycling in mature biofilms would protect them from Cu toxicity, low and high biomass biofilms did not differ in their physiological response to copper after 6 days of exposure. 3. A clear influence of P‐limitation on copper toxicity was observed. Periphytic communities that were previously fertilised for 18 days were three times more tolerant than control communities indicating that P‐limitation enhanced Cu toxicity and tolerance induction were probably related to the higher P‐availability. In addition, a compensation of Cu toxicity after P‐addition was observed in the long‐term (after 12 days). 4. We conclude that periphyton from oligotrophic streams is more sensitive to copper than periphyton from fertilised streams, and that therefore a higher effect of chronic copper exposure is expected to occur in oligotrophic P‐limited fluvial systems. 5. Extrapolation of our results to the area of study (Catalonia, north‐east of Spain) indicates that while the levels of Cu commonly found in the zone may negatively affect the periphyton from oligotrophic streams, because of the interaction between Cu and P, they are not able to control the growth of nuisance algae which is common under high nutrient conditions.     

Nutrient limitation of algal periphyton in streams along an acid mine drainage gradient

Metal oxyhydroxide precipitates that form from acid mine drainage (AMD) may indirectly limit periphyton by sorbing nutrients, particularly P. We examined effects of nutrient addition on periphytic algal biomass (chl a), community structure, and carbon and nitrogen content along an AMD gradient. Nutrient diffusing substrata with treatments of +P, +NP and control were placed at seven stream sites. Conductivity and SO₄ concentration ranged over an order of magnitude among sites and were used to define the AMD gradient, as they best indicate mine discharge sources of metals that create oxyhydroxide precipitates. Aqueous total phosphorous (TP) ranged from 2 to 23 μg · L^?1 and significantly decreased with increasing SO₄. Mean chl a concentrations at sites ranged from 0.2 to 8.1 μg · cm^?2. Across all sites, algal biomass was significantly higher on +NP than control treatments (Co), and significantly increased with +NP. The degree of nutrient limitation was determined by the increase in chl a concentration on +NP relative to Co (response ratio), which ranged from 0.6 to 9.7. Response to nutrient addition significantly declined with increasing aqueous TP, and significantly increased with increasing SO₄. Thus, nutrient limitation of algal biomass increased with AMD impact, indicating metal oxyhydroxides associated with AMD likely decreased P availability. Algal species composition was significantly affected by site but not nutrient treatment. Percent carbon content of periphyton on the Co significantly increased with AMD impact and corresponded to an increase in the relative abundance of Chlorophytes. Changes in periphyton biomass and cellular nutrient content associated with nutrient limitation in AMD streams may affect higher trophic levels.     

Controls on periphyton biomass in heterotrophic streams

1. Headwater streams of the Hubbard Brook Experimental Forest (HBEF) are typically characterised by a periphyton assemblage of low biomass and diversity. However, periphyton blooms have been observed following catchment deforestation experiments and occasionally during the annual spring thaw before canopy leaf‐out. 2. There is pronounced seasonal variation in both nutrient and light availability in HBEF streams. Stream water nitrogen (N) concentrations and light levels are higher before canopy leaf‐out and after leaf senescence and are lower during the growing season. Periphyton accrual rates also change seasonally; they are highest in spring prior to leaf‐out and significantly lower during summer and in autumn. 3. Periphyton biomass rarely responded positively to in‐situ experimental enrichment with nitrogen or phosphorus. In the summer, nutrient enrichment overall had no effect on periphyton biomass, while outside the growing season N enrichment had inhibitory effects on periphyton. 4. Despite these experimental results, surveys of ambient chlorophyll a concentrations in streams across the HBEF demonstrated no relationship between streamwater dissolved inorganic N or P concentrations and benthic chlorophyll a. 5. Our results suggest that HBEF periphyton communities are not closely regulated by nutrient availability, even during periods of high light availability. The inhibitory effects of nutrient enrichment outside the growing season are interesting, but further research is necessary to elucidate the mechanisms driving these responses.     

Changes in periphyton abundance and community structure with the dispersal of a caddisfly grazer, <Emphasis Type="Italic">Micrasema quadriloba</Emphasis>

We examined the larval population densities and biomass of a caddisfly grazer, Micrasema quadriloba, and the abundance and community structures of periphyton at a segment scale (7.4 km with four study sites), along a second-to fourth-order Japanese mountain stream throughout the grazer’s life cycle. In the uppermost riffle of the study segment (site 1), periphyton abundance was kept at low levels when the larvae occurred. The larval distribution spread downstream as larvae developed from first instars in May to fifth instars in January. We performed multiple regression analyses to test the effects of environmental variables and larval biomass on periphyton abundance in both the riffle of site 1 and the study segment; the results revealed that the larval biomass was significantly negatively correlated with periphyton abundance similarly in both the riffle and the study segment. In addition, both the correlation and community analyses showed that the larval biomass was significantly negatively correlated to the relative abundance of large and/or filamentous microalgae, which appeared in the uppermost layer of the periphyton mat, and that larval biomass was significantly positively correlated to the relative abundance of small diatoms, which strongly adhered to the substrate. Thus, the present study implied that the grazing of M. quadriloba larvae would regulate the abundance of periphyton in a riffle and also regulate the abundance and community structure of periphyton at the segment scale with the expansion of their longitudinal distribution.     

Interactions among invaders: community and ecosystem effects of multiple invasive species in an experimental aquatic system

With ecosystems increasingly supporting multiple invasive species, interactions among invaders could magnify or ameliorate the undesired consequences for native communities and ecosystems. We evaluated the individual and combined effects of rusty crayfish (Orconectes rusticus) and Chinese mystery snails [Bellamya (=Cipangopaludina) chinensis] on native snail communities (Physa, Helisoma and Lymnaea sp.) and ecosystem attributes (algal chlorophyll a and nutrient concentrations). Both invaders are widespread in the USA and commonly co-occur within northern temperate lakes, underscoring the importance of understanding their singular and joint effects. An outdoor mesocosm experiment revealed that while the two invaders had only weakly negative effects upon one another, both negatively affected the abundance and biomass of native snails, and their combined presence drove one native species to extinction and reduced a second by >95%. Owing to its larger size and thicker shell, adult Bellamya were protected from crayfish attack relative to native species (especially Physa and Lymnaea), suggesting the co-occurrence of these invaders in nature could have elevated consequences for native communities. The per capita impacts of Orconectes (a snail predator) on native snails were substantially greater than those of Bellamya (a snail competitor). Crayfish predation also had a cascading effect by reducing native snail biomass, leading to increased periphyton growth. Bellamya, in contrast, reduced periphyton biomass, likely causing a reduction in growth by native lymnaeid snails. Bellamya also increased water column N:P ratio, possibly because of a low P excretion rate relative to native snail species. Together, these findings highlight the importance of understanding interactions among invasive species, which can have significant community- and ecosystem-level effects.