Current velocity influences the facilitation and removal of algae by stream grazers

The modification of flows in lotic ecosystems can have dramatic effects on abiotic and biotic processes and change the structure of basal trophic levels. In high-gradient streams, most of the biota are benthic, and decreased flow may homogenize and reduce benthic current velocity, potentially changing stream ecosystem function. Grazing by macroinvertebrates is an important component of stream function because grazers regulate energy flow from primary producers to higher trophic levels. We conducted an experiment to examine how macroinvertebrate grazers facilitated or removed algal biomass across a gradient of benthic current velocity (0–40 cm s^?1). We chose three grazers (Drunella coloradensis, Cinygmula spp., and Epeorus deceptivus) from a montane stream and conducted our experiment using 24 artificial stream channels that had three treatments: no grazers (control), single-grazer, and combined-grazer treatments. In the absence of grazers, algal biomass increased with benthic current velocity. Grazer treatments differed from the control in that more algal biomass was removed at higher velocities, whereas algal accrual was largely facilitated at low velocities. The transition from facilitation to removal ranged from 4.5 to 5.9 cm s^?1 for individual grazer treatments and occurred at 11.7 cm s^?1 for the combined-grazer treatment. Our data suggest that velocity plays a significant role in the facilitation and removal of algae by macroinvertebrate grazers. Additionally, the patterns revealed here could have general implications for algal accrual in systems where flow is reduced.     

Relationship between littoral grazers and metaphytic algae in five softwater lakes

The zoobenthos communities within the littoral zones of five low alkalinity, Ontario lakes were examined during summer 1987. Detached algal clouds or metaphyton, dominated by algae from the family Zygnemataceae, accumulated over the summer in three of these lakes. Previous researchers have hypothesized that diminished grazing pressure due to extirpation of acid sensitive herbivores might enhance the development of metaphytic algae at pHs less than 6. No overt consistent differences existed among the abundances of small (cladocerans, cyclopoids, ostracods, oligochaetes, dipterans) or medium (amphipods, gastropods, emphemeropterans, trichopterans) grazers in study lakes with or without metaphyton. Notable differences did exist, however, with respect to what we term large grazers. Crayfish, algivorous fish (mostly cyprinids), and to a lesser extent tadpoles, were absent or rare in the three lakes with metaphytic algae but abundant in those without. If a reduction in grazing pressure can be implicated as a factor in enhanced metaphyton development, our correlative results suggest that it is the large grazers, particularly crayfish, that are the most likely candidates.     

Striking convergence in the mouthpart evolution of stream-living algae grazers

This scanning-electron microscopic study demonstrates the convergent evolution of the mouthparts of various herbivorous stream animals (insects from different orders, an isopod, snails, fish, and a tadpole) feeding on epilithic algal pastures. This food source is rich but is often difficult to harvest. Nevertheless, a large number of species can live on it because they have evolved highly specialized mouthparts. There are four functional problems that an algae grazer has to solve: the algae must be removed from the stone, they have to be collected and crushed, and a current shield is needed to prevent the water flow sweeping away the food. Among the 30 algae grazers examined in this study, a limited number of morphological solutions have been found for each of these adaptational problems. There are multiple evolutionary pathways for mouthpart adaptation and even closely related species have often evolved different types of tools for the same function. This refects the existence of a certain amount of evolutionary scope. Such freedom of evolution is present, however, only at the beginning of the adaptiogenesis of an algae grazer. Once one of the evolutionary pathways is taken, further improvement of the mouthparts is possible only by the refinement of the ‘chosen’ type of tools. The consequence of this is that a large number of astonishing convergences have occurred in algae grazers that have independently trodden the same evolutionary pathway.     

Effects of freshwater eutrophication on species and functional beta diversity of periphytic algae

Studies about beta diversity and environmental heterogeneity have shown that the strength of the environmental filtering effect may decrease with the increasing scale. These empirical results have related eutrophic aquatic environments to higher values of beta diversity, but never to dissimilarity of species and functional traits of periphytic algae. We tested the hypotheses that periphytic algae have higher dissimilarity of both species and functional traits in eutrophic environments, and that these dissimilarities are related to environmental dissimilarity. To this end, we used richness, density, and four functional traits of periphytic algae and local limnological data from wetlands in the Brazilian savanna (Cerrado). We analyzed the beta diversity and the relationship of species and functional dissimilarities with the environmental dissimilarity and geographic distances. Our hypothesis was confirmed for functional traits dissimilarity and for the importance of the environmental dissimilarity for both species and functional beta diversity. The cultural eutrophication led to a functional homogenization in urban wetlands, which indicates the establishment of species with similar ecological requirements, and consequently, similar ‘roles’ in the ecosystem, and also that sensitive species may have been replaced by tolerant species, leading to declining biodiversity.

     

Predatory fish impact on competition between stream insect grazers: a consideration of behaviorally- and density-mediated effects on an apparent coexistence pattern

A manipulative field experiment and theoretical analyses of a simple competition model were used to show how exploitative competition between a caddisfly (Glossosoma sp.) and three mayfly grazers (Ameletus sp., Baetisthermicus and Cinygmula sp.) was mediated by a predatory fish, freshwater sculpin (Cottus nozawae). The field experiment followed a two‐factorial design, with Glossosoma densities (natural vs reduced) and sculpin presence (present vs absent) as treatments. Diet analysis revealed that all four prey species were eaten under the natural condition and the sculpin preferred mayfly grazers to Glossosoma. Our experiment showed that although mayfly densities in the presence of either sculpin or Glossosoma were lower than in the no‐sculpin plus reduced‐Glossosoma treatment, no difference in mayfly densities was found between the following three treatments: sculpin plus natural‐Glossosoma, no‐sculpin plus natural‐Glossosoma, and sculpin plus reduced‐Glossosoma. These results indicated that fish predator produced no effects additive to the competitive operation on the mayfly density, and also that competitive operation of Glossosoma on the mayfly densities produced no additional effects to fish predators. In addition, although the competitive effect of the mayflies on Glossosoma could not be manipulated in isolation, the density of Glossosoma in the presence of sculpin was greater than that in the absence of sculpin in the natural‐Glossosoma treatment. Thus, the densities of these competing grazers either stayed the same or increased in the presence of the fish predator relative to predator‐free treatments. A theoretical model, developed to explain the experimental results here, predicted that the densities of the two competing prey under predation pressure could be simultaneously greater than those under predator absent conditions when the behaviorally‐mediated effects of the predator were strongly operative. Although we were unable to distinguish experimentally the two different effects of predator on the prey competition, the behaviorally‐ and density‐mediated effects, the reality of the behaviorally‐mediated effects in the experiment was discussed.     

The influence of crevice size on the protection of epilithic algae from grazers

1. This study investigated how the size of crevices might affect their effectiveness as refuges for diatom‐dominated algal assemblages from the grazing minnow Campostoma anomalum. 2. Crevice size was manipulated by making fired‐clay substrates, using moulds to produce eight substrates with pits from 1.17 to 22.0 mm diameter. Non‐pitted clay‐stones and limestone were also tested. Cages were used to control the access of Campostoma to arrays of the 10 different algal‐colonised substrates. The grazing treatments were: open and grazed, caged and ungrazed, and a grazed cage control. The experiment was replicated in eight large outdoor tanks. After 3 weeks, substrates were brushed and chlorophyll a concentrations of the removed algae and the algae remaining in pits were measured. 3. The experiment was field‐validated by exposing arrays of substrates to grazing Campostoma in five pools of a limestone stream. 4. The clay‐stone and limestone substrates accrued similar algal biomass and assemblages. 5. Smaller crevices provided more protection against grazing than larger crevices. Specifically, pits with openings smaller than 2 mm protected the enclosed algal assemblages in both the tank and field experiments. Larger pits provided less protection and pits over 7 mm in diameter were heavily grazed and may even be preferentially grazed by Campostoma. 6. None of the tested pit sizes were protective against larval chironomid grazers in the tank experiment, demonstrating that differences in the grazer size influence the effectiveness of crevice refuges.     

Effects of removal of riparian vegetation on algae and heterotrophs in a Mediterranean stream

The effect of removal of a riparian strip on aquaticautotrophic (algae) and heterotrophic (bacteria,macroinvertebrates) organisms was monitored in aMediterranean stream during the canopy growing period.Community composition, biomass and metabolicactivities were compared with those recorded during apre-riparian removal period and in a forested stretchdownstream. Higher irradiance was associated with Cladophora increase in the logged section. Algalbiomass increased up to ten times, and productivitywas up to four times higher than in the pre-removalperiod and the forested section. Bacterialcommunities showed higher ectoenzymatic activities(-glucosidase, -xylosidase) in thelogged section than in forested conditions. Moreoverthe coincidence between the maxima of-glucosidase and chlorophyll-a suggeststhat bacterial activity was enhanced by the higheravailability of high-quality algal material. Themacroinvertebrate community had higher density andbiomass in the logged section than in the forestedsection and in the pre-removal period. Scrapers andfilterers become dominant after riparian removal,while shredders, predators and collectors did not showsignificant changes either between sites or periods.Responses of environmental variables and bioticcommunities indicate that the changes occurring in thestream due to riparian removal could be consideredbottom-up controlled, as increased illumination wasthe main mechanism responsible.     

Nutrients, algae and grazers in some British streams of contrasting pH

1. The relationship between algal biomass accumulation, invertebrate colonization, and stream-water pH was investigated in seven streams in three regions of England and Wales. Possible nutrient limitation of algal production at all sites was examined with diffusion substrata. 2. Periphyton assemblages on experimental substrata after 30 days were dominated by diatoms, notably Eunotia spp., at all sites. Algal pigment concentration (chlorophyll a and phaeopigments) was not correlated with stream-water pH, and mean concentrations on control (unenriched) substrata ranged from 0.08 to 1.94 μg cm^?2. 3. The growth response of periphyton to nutrient additions was site specific. Algal production was stimulated by nutrient additions at sites in the English Lake District and Llyn Brianne (south-west Wales), but not in the Ashdown Forest (southern England). 4. Larval Chironomidae were the main invertebrates retrieved from substrata at all sites. Within all three regions, larval abundance was positively related to algal pigment concentration (biomass). Abundance of the stonefly Nemurella pictetii was also positively correlated with algal biomass at the one site where it occurred. 5. Our results indicate that epilithic algal production in small, oligotrophic streams is unlikely to be determined primarily by pH. Neither do they support the view that an absence of grazers from acid streams is necessarily due to an inadequate food supply.     

Group size of feeding stream case-bearing caddisfly grazers and resource abundance

Several herbivorous insects are known to benefit from feeding in groups; however, little is known about (1) the resource conditions under which herbivorous insects perform group feeding and (2) the optimum population size to get any benefits by group feeding, for example, in terms of growth performance. To test the hypotheses that the benefits from group feeding change with resource level and population size, we performed field investigations and an enclosure experiment using the grazer caddisfly larva Micrasema quadriloba. The field investigations revealed aggregated distributions of larvae (indicator of aggregation, I_δ=4.1±1.55, aggregated density: 12.7±5.3 individuals per 3.1×3.1 cm² (mean±1 SD)) when periphyton was abundant on stream cobbles and random distributions (I_δ=1.0±0.11) when periphyton was scarce. In the enclosure experiment, the relative growth rate (RGR) of the larvae at each population size showed different tendencies at high and low periphyton abundance levels; RGR with abundant periphyton had a convex curve with a peak at intermediate population size, whereas RGR with scarce periphyton decreased linearly with increasing population size. The benefits from group feeding thus changed with resource level; larvae obtained high growth performance by group feeding behavior only when the resource was sufficiently abundant. The present study revealed not only that the optimum group size of larvae increased their growth performance, but also that this optimum group size occurred frequently in the field. We also discuss the mechanisms and benefits of group feeding by case-bearing caddisfly grazers.     

Effects of microspatial complexity on a herbivore—epilithon interaction in an Australian upland stream

1. Increases in spatial heterogeneity may provide more food, create new resources or interfere with the foraging activities of a herbivore. 2. Three colonization experiments were performed in an upland stream to investigate the effects of one type of spatial heterogeneity (grooves on the substratum) on the relationship between an abundant herbivore, Agapetus monticolus (Banks, Trichoptera), and the epilithon on which it feeds. 3. The results indicate that grooves do not provide any different resources or interfere with the foraging activities of A. monticolus, but may increase food abundance, although the effect of grooves on food abundance appears to vary temporally. 4. Temporal variation in the effects of microspatial complexity raises the possibility that its importance to the community may be over- or underestimated unless experiments are designed to incorporate this temporal variation.     

Epilithic biofilm metabolism during the high water flow period in an Andean neotropical stream

Tropical rivers show high discharge during the rainy period, and this might limit the epilithic biofilm biomass. While high discharge may limit algal biomass, high light and nutrient availability common in disturbed watersheds may promote high productivity, and even autotrophy. Metabolism may be affected by discharge peaks, but well-adapted algal communities should support quick recovery. These assumptions were analyzed through the patterns of oxygen production and consumption in the Rio Tota, an Andean stream in the north of the Colombian Andes with relevant farming activities. Metabolism was measured for 26 consecutive days with in situ recirculating incubation chambers. The results stressed that these are net autotrophic systems during the high water flow period and that this occurs with algal communities of low biomass and low diversity. The environmental conditions of high light, high but non-catastrophic flow, and moderate temperature favor the algal community performance. However, the stream metabolism was sensitive to the day-by-day variations of environmental conditions, in particular to the above-average hydrological variations.     

Control of periphyton standing crop in an Atlantic Forest stream: the relative roles of nutrients,grazers and predators

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Allelopathic interaction between a reef coelenterate and benthic algae

Summary We did in situ experiments to study the influence of a water-borne substance, exuded by the reef anthozoan Condylactis gigantea, on filamentous algae. Algal biomass accumulation on experimental glass slides was significantly inhibited by the exudate. Algal spores and diatoms were present in equal numbers on experimental and control slides, but the exudate inhibited germination of spores and was toxic to developing green, red and brown algae. The algal population on the control slides consisted of early colonizers as well as later successional species and reflected the natural population. The described allelochemic mechanism will be important in interference competition between the anemone and algae. Settling of larvae in supposedly space-limited systems such as reefs is facilitated when algaecovered surfaces are open to settlement. Present address: Netherlands Institute for Sea Research (NIOZ) P.O. Box 59, Texel, The Netherlands     

Interactions between a stream caddisfly and the algae on its case: factors affecting algal quantity

1. Field experiments were conducted to examine interactions between larvae of the caddisfly Gumaga nigricula and the algae (primarily diatoms) located on their sand grain cases. Floating experimental enclosures were placed in sunlit pools of a California stream. Treatments applied to occupied cases, or to those from which caddisflies had been removed, included variations in grazer density, addition of nutrients, and modification of illumination using shade cloth. 2. Neither faecal-enriched agar in empty cases nor injections of faecal material into plugged cases affected case chlorophyll a concentration; injected nutrients (N and P), however, did increase chlorophyll a concentration. Faecal material may be nutrient poor, or nutrients may not be readily released. In addition, the relatively impermeable case wall may reduce the likelihood that nutrients from caddisflies reach the algae on the outer case surface. 3. Although a Gumaga larva may graze algae from its own case, grazing by conspecifics and the caddisfly Helicopsyche borealis causes a greater reduction in the chlorophyll a content of cases. 4. Chlorophyll a content of empty cases was decreased by experimental reduction of light. Because of shading associated with burrowing, chlorophyll a content of cases occupied by Gumaga was low and was unaffected by experimental light reduction. 5. Removal of Gumaga from its case results in rapid accrual of algae. Thus, the net effect of larval presence is to diminish the algal content of its case.     

Contrasting patterns in local and zonal family richness of stream invertebrates along an Andean altitudinal gradient

1. Describing and understanding patterns in biological diversity along major geographical gradients is an important topic in ecology. Samples collected from a large number of physically and chemically comparable stream sites along a 4000 m gradient of altitude in the Andes of Ecuador served to characterise patterns of family richness of aquatic macroinvertebrates at the scale of the stream site (local) and at that of discrete altitudinal zones. 2. Both mean local and zonal family richness decreased by about 50% from sea level to 4000 m a.s.l. Local richness declined linearly, while zonal richness remained constant from sea level up to a threshold altitude of about 1800 m, whereafter it decreased. 3. From sea level to 1800 m few families were lost from zonal richness and few were gained. From 1800 to 3800 m the decrease in the number of families was accounted for by a loss of families present in lowland streams, with few new families gained. Hence, there was relatively little turnover of families along the entire gradient. 4. The diverging pattern of local and zonal richness was caused by sporadically occurring families inflating zonal richness at mid‐altitudes. If the sporadic families were represented by the same species found commonly in the lowlands, then the mid‐altitudinal zonal richness would be maintained by a ‘rescue effect’. More probably, however, the sporadically occurring families found at mid‐altitudes are each represented by new species replacing each other along the gradient, the families progressively diminishing in species richness and occurrence as the overall temperature tolerance of the family is approached. 5. This study demonstrates that spatial scale affects altitudinal patterns in the taxonomic richness of stream invertebrates. It also showed that family‐level identification can facilitate interpretation of sources and sinks of biodiversity along geographic gradients.     

Effects of nutrients and grazers on periphyton phosphorus in lake enclosures

SUMMARY. 1. Periphyton. measured as particulate phosphorus (PP) and expressed as periphyton PP, growing on vertically oriented substrata (polyvinyl impregnated nylon) under different nutrient loadings, light intensities (exposures), and grazer communities was examined in eight large enclosures (750 m3) where nutrients (N and P) and planktivorous fish (1+yellow perch) were added in a 2x2 factorial design.
2. During the first 3 weeks of the experiment (25 June to 15 July), there was a significantly higher accumulation of phosphorus into periphyton (periphyton PP) with fertilization, but fish addition had no effect. During the fourth to seventh weeks (16 July to 12 August), addition of fish was associated with lower abundance of amphipods and chironomids and higher concentration of periphyton PP. In the enclosures without fish, these invertebrates were over 25 times more abundant, and periphyton PP decreased substantially compared to the June-July period. Fertilization increased periphyton PP only at high exposures in the enclosures with fish.
3. Exposure had a significant effect on periphyton PP. In the enclosures with fish, high abundance of nanoplankton reduced water transparency, and periphyton PP was lower in the deeper waters which may have been due to limitation by low light. Lower periphyton PP was also observed at the surface on sunny sides of enclosures without fish, and therefore with high water transparency. This pattern may have been due to inhibitory effects of high light intensity.
4. Periphyton communities in the enclosures with fish had higher uptake rates for planktonic phosphorus, and lower rates of phosphorus release, suggesting that periphyton with high phosphorus demand may have high internal cycling of assimilated phosphorus.     

Trematodes and nematodes parasitizing the benthic insect community of an Andean Patagonian stream,with emphasis on plagiorchiid metacercariae

In freshwater systems, parasitological studies have mainly been carried out on vertebrates and molluscs, but little is known about parasites of aquatic insects. We describe the trematodes and nematodes parasitizing the benthic insects of an Andean Patagonian stream and the presence of parasites in the terrestrial adult stages. Members of 3 of 20 insect taxa were found to be parasitized by larval nematodes, and members of six taxa harbored metacercariae of digeneans. In benthic samples, chironomids, simuliids (Order Diptera), and baetids (Order Ephemeroptera) harbored mermithid larvae (Nematoda). The stonefly Antarctoperla michaelseni (Order Plecoptera), the caddisfly Smicridea annulicornis (Order Trichoptera), a watersnipe fly (Order Diptera: Athericidae), and three species of leptophlebiid mayflies (Order Ephemeroptera) were parasitized by encysted plagiorchiid metacercariae (Order Plagiorchiida). Most metacercariae were found in the three species of mayflies with prevalences ranging 15–63% and mean intensities ranging 1.2–4.9. Prevalence declined from summer to early winter, probably because of the emergence of infected nymphs and the recruitment of uninfected new cohorts. The imagos had live metacercariae with higher prevalences and intensities of infection than nymphs. We suggest that these plagiorchiids have an allogenic life cycle, involving a terrestrial definitive host.     

Harmful algae on tropical coral reefs: Bottom-up eutrophication and top-down herbivory

A conceptual paradigm, the “Relative Dominance Model”, provides the perspective to assess the interactive external forcing-mechanisms controlling phase shifts among the dominant benthic functional groups on tropical coral reefs [i.e., microalgal turfs and frondose macroalgae (often harmful) versus reef-building corals and calcareous coralline algae (mostly beneficial due to accretion of calcareous reef framework)]. Manipulative experiments, analyses of existing communities and bioassays tested hypotheses that the relative dominances of these functional groups are mediated by two principal controlling factors: nutrients (i.e., bottom-up control) and herbivory (i.e., top-down control). The results show that reduced nutrients alone do not preclude fleshy algal growth when herbivory is low, and high herbivory alone does not prevent fleshy algal growth when nutrients are elevated. However, reduced nutrients in combination with high herbivory virtually eliminate all forms of fleshy micro- and macro-algae. The findings reveal considerable complexity in that increases in bottom-up nutrient controls and their interactions stimulate harmful fleshy algal blooms (that can alter the abundance patterns among functional groups, even under intense herbivory); conversely, elevated nutrients inhibit the growth of ecologically beneficial reef-building corals. The results show even further complexity in that nutrients also act directly as either limiting factors (e.g., physiological stresses) or as stimulatory mechanisms (e.g., growth enhancing factors), as well as functioning indirectly by influencing competitive outcomes. Herbivory directly reduces fleshy-algal biomass, which indirectly (via competitive release) favors the expansion of grazer-resistant reef-building corals and coralline algae. Because of the sensitive nature of direct/indirect and stimulating/limiting interacting factors, coral reefs are particularly vulnerable to anthropogenic reversal effects that decrease top-down controls and, concomitantly, increase bottom-up controls, dramatically altering ecosystem resiliencies.     

维生素B12在细菌与真核藻类相互作用中的功能研究进展

自然环境中细菌与藻类发生着直接或间接的相互作用,这些相互作用对双方的生理功能、群落组成、逆境抵御等方面具有重要影响。维生素B12 (VB12)是一种结构复杂、功能重要的维生素,是许多真核藻类所必需的生长因子。由于VB12在自然界中只能由产VB12的细菌合成,产VB12细菌对真核藻类的生理功能具有重要影响。本文将主要介绍近年来在细菌-真核藻类相互作用中VB12所参与的藻类生长调节、抗逆反应与基因表达调控等方面的研究进展,并讨论在细菌-真核藻类分子相互作用这一研究模式中应注重的发展方向。     

The effects of disturbance on a herbivore-epilithon interaction in an upland stream

Two experiments were performed in an upland stream to determine the effects of the frequency of physical disturbance on the relationship between an abundant glossosomatid trichopteran (Agapetus monticolus; Banks) and the epilithon upon which it feeds. Artificial cobbles with an established epilithic community were tumbled either every 1, 2 or 4 weeks. The first experiment failed to detect any significant effects of rock tumbling on the abundance of A. monticolus or the epilithon: a result due to several spates. The first experiment did reveal that disturbances may disrupt the ability of A. monticolus to locate patches of abundant food. The second experiment found that although the abundance of A. monticolus was not affected by the disturbances, periphyton abundance was significantly reduced. Increasing the frequency of disturbance did not magnify this effect. Comparisons of these results with other studies of disturbance in streams indicate that the effects of disturbance on herbivory may be highly variable. A variety of factors, such as the relative resistances of the herbivores and the epilithon, need to be examined before the effects of disturbances on lotic herbivorous interactions can be completely understood.