Resolving resource partitioning in parrotfishes (Scarini) using microhistology of feeding substrata

Parrotfishes (Scarini) are considered key agents in coral reef health and recovery, but the drivers of parrotfish–coral dynamics remain contentious. The prevailing view of parrotfishes as ecosystem engineers is based on the perceived removal of algal turf, macroalgae and sediment, but these are effects of feeding, not causes. The recent proposal that most parrotfishes are ‘microphages’ that target microscopic photoautotrophs (particularly cyanobacteria) identifies the need to resolve dietary targets at a microscopic scale. Here, we investigate parrotfish dietary targets by posing the following two questions: (1) are microscopic photoautotrophs the most consistent and dominant elements of the prey community, and (2) do the prey community and substratum taphonomy vary between parrotfish species? In order to identify and quantify dietary targets, five parrotfish species were followed until focused feeding was observed at Lizard Island on the Great Barrier Reef, Australia. Feeding sites were photographed in situ and extracted as substratum bite cores. Cores were analysed microscopically to identify and quantify all epilithic photoautotrophs. Endolithic photoautotrophs accessible to excavating parrotfish were also investigated by vacuum-embedding cores with epoxy resin followed by decalcification to expose endolith microborings. The dominant functional groups of epilithic biota on the cores were tufted cyanobacteria, turfing algae and crustose coralline algae (CCA). The only consistent feature across all cores was the high density of filamentous cyanobacteria, supporting the view that these parrotfishes target microphotoautotrophs. Macroalgae was absent or a minor component on cores, supporting the hypothesis that parrotfishes avoid larger algae. The microchlorophyte Ostreobium was the dominant photoautotrophic euendolith (true borer) in the cores of the excavating parrotfish Chlorurus microrhinos. Significant differences in CCA coverage, turf height and substrate taphonomy were found among the five parrotfish species, suggesting that interspecific resource partitioning is based on successional stage of feeding substrata.

     

Allelochemicals produced by Caribbean macroalgae and cyanobacteria have species-specific effects on reef coral microorganisms

Coral populations have precipitously declined on Caribbean reefs while algal abundance has increased, leading to enhanced competitive damage to corals, which likely is mediated by the potent allelochemicals produced by both macroalgae and benthic cyanobacteria. Allelochemicals may affect the composition and abundance of coral-associated microorganisms that control host responses and adaptations to environmental change, including susceptibility to bacterial diseases. Here, we demonstrate that extracts of six Caribbean macroalgae and two benthic cyanobacteria have both inhibitory and stimulatory effects on bacterial taxa cultured from the surfaces of Caribbean corals, macroalgae, and corals exposed to macroalgal extracts. The growth of 54 bacterial isolates was monitored in the presence of lipophilic and hydrophilic crude extracts derived from Caribbean macroalgae and cyanobacteria using 96-well plate bioassays. All 54 bacterial cultures were identified by ribotyping. Lipophilic extracts from two species of Dictyota brown algae inhibited >50% of the reef coral bacteria assayed, and hydrophilic compounds from Dictyota menstrualis particularly inhibited Vibrio bacteria, a genus associated with several coral diseases. In contrast, both lipo- and hydrophilic extracts from 2 species of Lyngbya cyanobacteria strongly stimulated bacterial growth. The brown alga Lobophora variegata produced hydrophilic compounds with broad-spectrum antibacterial effects, which inhibited 93% of the bacterial cultures. Furthermore, bacteria cultured from different locations (corals vs. macroalgae vs. coral surfaces exposed to macroalgal extracts) responded differently to algal extracts. These results reveal that extracts from macroalgae and cyanobacteria have species-specific effects on the composition of coral-microbial assemblages, which in turn may increase coral host susceptibility to disease and result in coral mortality.     

Parrotfish abundance and selective corallivory on a Belizean coral reef

Parrotfish are important members of coral reef communities because they consume macroalgae that would otherwise outcompete reef-building corals for space. However, some Caribbean parrotfish species also feed directly on live corals, and thus have the potential to negatively impact coral fitness and survival. This study investigates selective grazing by parrotfish on particular coral species, differences in grazing incidence among reef habitats and intraspecific discrimination among colonies of several coral species. We also investigate spatial and temporal patterns of parrotfish species abundance across habitats on the Belize barrier reef, and examine correlations between parrotfish abundance and grazing intensity across reef habitats. We found that members of the Montastraea annularis species complex, major builders of Caribbean reefs, were preferred targets of parrotfish grazing across all reef habitats, while M. cavernosa, Agaricia agaricites, Diploria strigosa, Porites astreoides and Porites porites were not preferred; Siderastrea siderea was preferentially grazed only in the spur and groove habitats. Parrotfish grazing preferences varied across habitats; M. annularis was grazed most often in shallow habitats, whereas M. franksi was consumed more at depth. Although it was not possible to directly observe parrotfish grazing on corals, we did find a positive correlation between Sparisoma aurofrenatum abundance and M. franksi grazing incidence across habitats. Finally, when we compared our results to parrotfish abundances measured by a previous study, we found that Sparisoma viride and Sp. aurofrenatum, two species known to be corallivorous, had increased abundances between 1982 and 2004. In light of escalating threats on Caribbean reef corals, it would be important for future studies to evaluate the impact of parrotfish corallivory on coral survival.     

IMPACT OF THE INVASIVE SEAWEED SARGASSUM MUTICUM (PHAEOPHYTA) ON AN INTERTIDAL MACROALGAL ASSEMBLAGE1

The impact of the invasive seaweed Sargassum muticum (Yendo) Fensholt on a low intertidal macroalgal assemblage was assessed at a semiexposed rocky shore in northern Spain between 2002 and 2004. Sargassum muticum plants were removed from the mature macroalgal assemblage and from those occurring along the successional process of the assemblage. Biomass, richness, diversity, and percentage cover of macroalgae in experimental plots were compared with unmanipulated controls. The effect of S. muticum removal on the macroalgal assemblage more than 2 years after the beginning of the experiment was negligible. Moreover, no differences between treatments were detected in the general patterns of succession. Only significant differences in S. muticum abundance were detected between treatments at the end of the experiment. We suggest that the low abundance of S. muticum at this intertidal level and its pseudoperennial life cycle may limit competition with native macroalgae. However, long‐term removal experiments may be a more indicator of the impact of S. muticum at the upper limit of its vertical distribution.     

Application of diet theory reveals context-dependent foraging preferences in an herbivorous coral reef fish

Dietary preferences of grazers can drive spatial variability in top-down control of autotroph communities, because diet composition may depend on the relative availability of autotroph species. On Caribbean coral reefs, parrotfish grazing is important in limiting macroalgae, but parrotfish dietary preferences are poorly understood. We applied diet-switching analysis to quantify the foraging preferences of the redband parrotfish (Sparisoma aurofrenatum). At 12 Caribbean reefs, we observed 293 redband parrotfish in 5-min feeding bouts and quantified relative benthic algal cover using quadrats. The primary diet items were macroalgal turfs, Halimeda spp., and foliose macroalgae (primarily Dictyota spp. and Lobophora spp.). When each resource was evaluated independently, there were only weak relationships between resource cover and foraging effort (number of bites taken). Electivity for each resource also showed no pattern, varying from positive (preference for the resource) to negative (avoidance) across sites. However, a diet-switching analysis consisting of pairwise comparisons of relative cover and relative foraging effort revealed clearer patterns: parrotfish (a) preferred Halimeda and macroalgal turfs equally, and those two resources were highly substitutable; (b) preferred Halimeda to foliose macroalgae, but those two resources were complementary; and (c) also preferred turf to foliose macroalgae, and those resources were also complementary. Thus parrotfish grazing rates depend on relative, not absolute, abundance of macroalgal types, due to differences in substitutability among resources. Application of similar analyses may help predict potential changes in foraging effort of benthic grazers over spatial gradients that could inform expectations for reef recovery following the protection of herbivore populations.     

Impact of Herbivore Identity on Algal Succession and Coral Growth on a Caribbean Reef

Background

Herbivory is an important top-down force on coral reefs that regulates macroalgal abundance, mediates competitive interactions between macroalgae and corals, and provides resilience following disturbances such as hurricanes and coral bleaching. However, reductions in herbivore diversity and abundance via disease or over-fishing may harm corals directly and may indirectly increase coral susceptibility to other disturbances.

Methodology and Principal Findings

In two experiments over two years, we enclosed equivalent densities and masses of either single-species or mixed-species of herbivorous fishes in replicate, 4 m² cages at a depth of 17 m on a reef in the Florida Keys, USA to evaluate the effects of herbivore identity and species richness on colonization and development of macroalgal communities and the cascading effects of algae on coral growth. In Year 1, we used the redband parrotfish (Sparisoma aurofrenatum) and the ocean surgeonfish (Acanthurus bahianus); in Year 2, we used the redband parrotfish and the princess parrotfish (Scarus taeniopterus). On new substrates, rapid grazing by ocean surgeonfish and princess parrotfish kept communities in an early successional stage dominated by short, filamentous algae and crustose coralline algae that did not suppress coral growth. In contrast, feeding by redband parrotfish allowed an accumulation of tall filaments and later successional macroalgae that suppressed coral growth. These patterns contrast with patterns from established communities not undergoing primary succession; on established substrates redband parrotfish significantly reduced upright macroalgal cover while ocean surgeonfish and princess parrotfish allowed significant increases in late successional macroalgae.

Significance

This study further highlights the importance of biodiversity in affecting ecosystem function in that different species of herbivorous fishes had very different impacts on reef communities depending on the developmental stage of the community. The species-specific effects of herbivorous fishes suggest that a species-rich herbivore fauna can be critical in providing the resilience that reefs need for recovery from common disturbances such as coral bleaching and storm damage.     

Context-dependent corallivory by parrotfishes in a Caribbean reef ecosystem

This study assesses the patterns of corallivory by parrotfishes across reefs of the Florida Keys, USA. These reefs represent a relatively unique combination within the wider Caribbean of low coral cover and high parrotfish abundance suggesting that predation pressure could be intense. Surveys across eight shallow forereefs documented the abundance of corals, corallivorous parrotfishes, and predation scars on corals. The corals Porites porites and Porites astreoides were preyed on most frequently with the rates of predation an order of magnitude greater than has been documented for other areas of the Caribbean. In fact, parrotfish bite density on these preferred corals was up to 34 times greater than reported for corals on other reefs worldwide. On reefs where coral cover was low and corals such as Montastraea faveolata, often preferred prey for parrotfishes, were rare, predation rates on P. porites and P. astreoides, and other less common corals, intensified further. The intensity of parrotfish predation increased significantly as coral cover decreased. However, parrotfish abundance showed only a marginal positive relationship with predation pressure on corals, likely because corallivorous parrotfish were abundant across all reefs. Parrotfishes often have significant positive impacts on coral cover by facilitating coral recruitment, survival, and growth via their grazing of algae. However, abundant corallivorous parrotfishes combined with low coral cover may result in higher predation on corals and intensify the negative impact that parrotfishes have on remaining corals.     

Chronic spearfishing may indirectly affect reef health through reductions in parrotfish bite rates

The grazing behaviour of two Caribbean parrotfish, a fished species, the stoplight parrotfish Sparisoma viride and a non-fished species, the striped parrotfish Scarus iseri, were studied in the presence (fished site) and absence (marine reserve) of chronic spearfishing activity. Diurnal feeding periodicity did not differ between the sites in either species: roving individuals had significantly higher bite rates in the afternoon, while territorial individuals foraged consistently throughout the day. Mean bite rate varied between sites in both species. Abundance, biomass and bite rates of S. viride were all significantly higher within the reserve, except for roving S. viride which had a higher mean bite rate in the afternoon outside the reserve compared with within it, attributable to maximisation of feeding in the afternoon when fishing risk was lower. Scarus iseri mean abundance and bite rate were greater outside the reserve, potentially because reduction in large territorial herbivores allowed S. iseri to feed more rapidly. By reducing the grazing potential of the remaining S. viride individuals the effect of fishing is greater than would be predicted from biomass changes alone. Less grazing by S. viride would not be compensated for by the increase in grazing by S. iseri because the latter feeds on different algae. Spearfishing of key parrotfish species reduces grazing potential directly by extraction and indirectly by changing behaviour.     

Parrotfish predation on massive Porites on the Great Barrier Reef

Parrotfish grazing scars on coral colonies were quantified across four reef zones at Lizard Island, Northern Great Barrier Reef (GBR). The abundance of parrotfish grazing scars was highest on reef flat and crest, with massive Porites spp. colonies having more parrotfish grazing scars than all other coral species combined. Massive Porites was the only coral type positively selected for grazing by parrotfishes in all four reef zones. The density of parrotfish grazing scars on massive Porites spp., and the rate of new scar formation, was highest on the reef crest and flat, reflecting the lower massive Porites cover and higher parrotfish abundance in these habitats. Overall, it appears that parrotfish predation pressure on corals could affect the abundance of preferred coral species, especially massive Porites spp, across the reef gradient. Parrotfish predation on corals may have a more important role on the GBR reefs than previously thought.     

New incubation device forin situ measurement of acetylene-reducing activity in ricefields

A device forin situ estimation of biological nitrogen fixation in shallow-water ricefields was developed using the acetylene-reducing assay. The device consists of a rigid transparent bottomless plastic bottle provided with an agitation system. Laboratory experiments using flooded pots inoculated withAnabaena UAM 202 indicated that agitation significantly reduced the time needed to detect the production of ethylene by eliminating the slow diffusion of acetylene and ethylene in water. A direct relationship between the abundance of cyanobacteria and the rate of acetylene reduction was observed in laboratory and field experiments. A negative correlation between the amount of combined nitrogen and the abundance of cyanobacteria was observed in the field.     

Bait worm packaging as a potential vector of invasive species

Invasive species have become an increasingly greater concern for the ecological health of coastal ecosystems, yet vectors of these introductions often are unclear. This project evaluated the potential for the brown seaweed Ascophyllum nodosum ecad scorpiodes (Hauck) Reinke, packaged with bait worms (Nereis virens) harvested from the coast of Maine (USA), as a vector of invasive marine fauna and flora. Often, the seaweed and contents of the bait boxes are discarded into the water by recreational fishermen after using the bait worms, and any included non-native species may then be introduced. Bait boxes were purchased from several commercial vendors in Connecticut and New York over a two-year period. Subsamples of the seaweed were placed in laboratory culture and the growth of associated macro- and microalgae was monitored. Marine invertebrate species present in the samples were also identified and quantified. Results indicated 13 species of macroalgae and 23 species of invertebrates were associated with baitboxes. Among the highly diverse microbial assemblage detected, two species of potentially toxic marine microalgae, Alexandrium fundyense Balech and Pseudonitzschia multiseries (Hasle) Hasle, were found both prior to and after incubation at various temperatures, indicating these harmful algae are brought to and can survive in receiving waters. These findings highlight the need to consider alternative choices of bait box packaging materials or appropriate disposal methods of the seaweed in order to minimize the transport of species which are not native to the receiving coastal waters.     

Feeding by coral reef mesograzers: algae or cyanobacteria?

Marine studies on herbivory have addressed the role of algae as food and shelter for small consumers, but the potential of benthic cyanobacteria to play similar roles is largely unknown. Here, feeding preferences were measured for eight invertebrate consumers from Guam, offered four common macroalgae and two cyanobacteria. The survivorship of another consumer raised on either macroalgae or cyanobacteria was also assessed. From the choices offered, the sacoglossans Elysia rufescens and E. ornata consumed the green macroalga Bryopsis pennata. The crab Menaethius monoceros preferred the red alga Acanthophora spicifera. The amphipods Parhyale hawaiensis and Cymadusa imbroglio consumed macroalgae and cyanobacteria in equivalent amounts, with C. imbroglio showing less selectivity among diets. In contrast to these patterns, in these assays the gastropods Stylocheilus striatus, Haminoea cymbalum, H. ovalis, and Haminoea sp. fed exclusively, or survived only, on cyanobacteria. Preferences for different cyanobacteria varied. Field surveys of cyanobacteria-associated species yielded 34 different invertebrate taxa and suggested different degrees of specificity in these associations. Tropical mesograzers exploit considerably different food resources, with some species adapted to consume cyanobacterial mats. Benthic cyanobacteria may play important roles as food and shelter for marine consumers and may indirectly influence local biodiversity through their associated fauna.     

Mesoherbivores reduce net growth and induce chemical resistance in natural seaweed populations

Herbivory on marine macroalgae (seaweeds) in temperate areas is often dominated by relatively small gastropods and crustaceans (mesoherbivores). The effects of these herbivores on the performance of adult seaweeds have so far been almost exclusively investigated under artificial laboratory conditions. Furthermore, several recent laboratory studies with mesoherbivores indicate that inducible chemical resistance may be as common in seaweeds as in vascular plants. However, in order to further explore and test the possible ecological significance of induced chemical resistance in temperate seaweeds, data are needed that address this issue in natural populations. We investigated the effect of grazing by littorinid herbivorous snails (Littorina spp.) on the individual net growth of the brown seaweed Ascophyllum nodosum in natural field populations. Furthermore, the capacity for induced resistance in the seaweeds was assessed by removing herbivores and assaying for relaxation of defences. We found that ambient densities of gastropod herbivores significantly reduced net growth by 45% in natural field populations of A. nodosum. Seaweeds previously exposed to grazing in the field were less consumed by gastropod herbivores in feeding bioassays. Furthermore, the concentration of phlorotannins (polyphenolics), which have been shown to deter gastropod herbivores, was higher in the seaweeds that were exposed to gastropod herbivores in the field. This field study corroborates earlier laboratory experiments and demonstrates that it is important to make sure that the lack of experimental field data on marine mesoherbivory does not lead to rash conclusions about the lack of significant effects of these herbivores on seaweed performance. The results strongly suggest that gastropods exert a significant selection pressure on the evolution of defensive traits in the seaweeds, and that brown seaweeds can respond to attacks by natural densities of these herbivores through increased chemical resistance to further grazing.     

Biological and chemical factors driving the temporal distribution of cyanobacteria and heterotrophic bacteria in a eutrophic lake (West Lake,China)

Physico-chemical parameters, hydrological conditions, and microbial interactions can affect the growth and persistence of cyanobacteria, but the interacting effects among these bloom-forming factors are still poorly known. This hampers our capacity to predict the occurrence of cyanobacterial bloom accurately. Here, we studied the relationship between temperature, N and P cycles, and the microbial community abundance and diversity at 0.5 m under the surface of West Lake (China) from January 21 to November 20, 2015, in order to better understand the key factors regulating temporal changes in the cyanobacterial community. Using high throughput sequencing of the 16S rRNA gene V3-V4 region, we studied the diversity and abundance of bacteria. In parallel, we measured physico-chemical parameters and followed the abundance of key genes involved in N fixation, denitrification, and nutrient uptake. Multivariate analyses suggest that P concentration and water temperature are the key factors controlling the outbreak of summer cyanobacterial bloom. RT-qPCR analyses of the bacterial community and measurements of the copy number of denitrification-related gene (nirK, nosZ, nirS) show that denitrification potential and denitrifying bacteria relative abundance (Pseudomonas and Bacillus) increased in concert with diazotrophic cyanobacterial genera (Anabaena, Nostoc, Aphanizomenon flos-aquae) and the common bloom-forming non-diazotrophic cyanobacterium genus Microcystis. The present study brings new insights on the complex interplay between physico-chemical parameters, heterotrophic bacterial community composition, nitrogen cycle, and cyanobacteria dominance in a eutrophic lake.

     

Effects of fish predation and seaweed competition on the survival and growth of corals

On Caribbean coral reefs, high rates of grazing by herbivorous fishes are thought to benefit corals because fishes consume competing seaweeds. We conducted field experiments in the Florida Keys, USA, to examine the effects of grazing fishes on coral/seaweed competition. Initially, fragments of Porites divaracata from an inshore habitat were transplanted into full-cage, half-cage, and no-cage treatments on a fore-reef. Within 48 h, 56% of the unprotected corals in half-cage and no-cage treatments (62 of 111) were completely consumed. Stoplight parrotfish (Sparisoma viride) were the major coral predators, with redband parrotfish (S. aurofrenatum) also commonly attacking this coral. Next, we transplanted fragments of P. porites collected from the fore-reef habitat where our caging experiments were being conducted into the three cage treatments, half in the presence of transplanted seaweeds, and half onto initially clean substrates. The corals were allowed to grow in these conditions, with concurrent development of competing seaweeds, for 14 weeks. Although seaweed cover and biomass were both significantly greater in the full-cage treatment, coral growth did not differ significantly between cage treatments even though corals placed with pre-planted seaweeds grew significantly less than corals placed on initially clean substrate. This surprising result occurred because parrotfishes not only grazed algae from accessible treatments, but also fed directly on our coral transplants. Parrotfish feeding scars were significantly more abundant on P. porites from the half and no-cage treatments than on corals in the full cages. On this Florida reef, direct fish predation on some coral species (P. divaracata) can exclude them from fore-reef areas, as has previously been shown for certain seaweeds and sponges. For other corals that live on the fore-reef (P. porites), the benefits of fishes removing seaweeds can be counterbalanced by the detrimental effects of fishes directly consuming corals. Received: 31 May 1997 / Accepted: 2 September 1997     

Can tropical freshwater zooplankton graze efficiently on cyanobacteria?

Zooplankton may at times graze cyanobacteria. However, their top-down effects are considered to be low, particularly in tropical regions dominated by small-size grazers that may be unable to consume efficiently filamentous or colonial species. Recently, cyanobacteria blooms were reported in the Senegal River hydrosystem. We conducted feeding experiments to assess the ability of copepods (Pseudodiaptomus hessei and Mesocyclops ogunnus), cladocerans (Moina micrura and Ceriodaphnia cornuta), and rotifers (Brachionus angularis, B. falcatus, and Keratella sp.) to control different cyanobacteria (Cylindrospermopsis raciborskii, Anabaena solitaria, A. flos-aquae, and Microcystis aeruginosa). None of the zooplankton species ingested M. aeruginosa. Mesocyclops ogunnus did not consume any of the cyanobacteria. Both cladocerans consumed the smallest filaments of cyanobacteria, whereas all the rotifers and P. hessei consumed a broader food-size spectrum. The functional feeding responses suggest that the concentration and size of the filaments are not the sole criteria for food consumption. The high zooplankton community grazing rates, estimated by applying the clearance rates measured in the laboratory to the in situ zooplankton abundance, indicate that grazing by zooplankton potentially constitutes an important controlling factor for the filamentous cyanobacteria in the tropics.     

The effect of epibionts on the susceptibility of the red seaweed Cryptonemia seminervis to herbivory and fouling

Epibiosis or fouling on living organisms can have direct and indirect detrimental effects, in particular on photosynthetic organisms such as seaweeds. It thus seems reasonable to hypothesize that macroalgae have been selected for the presence or induction of antifouling (AF) defences. The red seaweed Cryptonemia seminervis is usually found in nature with an elevated cover of epibionts. To assess the effect of epibiosis on the susceptibility of this seaweed to herbivory and fouling, the abundance of fouling was evaluated and compared to herbivore consumption (by amphipods and sea urchins) of fouled (bryozoan and sponge) and non-fouled C. seminervis. Attachment of the mussel Perna perna to surfaces treated with extracts from seaweeds with and without epibionts was also assessed. Epibiosis corresponded to ca. 51% of the blade surface of C. seminervis, sometimes covering as much as 90% and up to 51% of the thallus weight, encompassing mainly the bryozoan Membranipora membranacea and an unidentified sponge. Algae colonized by M. membranacea were preferred compared to algae devoid of epibionts, a ‘shared doom’ effect, either by the amphipod Elasmopus brasiliensis or by the urchin Lytechinus variegatus (p < 0.01). Sponge epibiosis also increased consumption by both herbivores (p < 0.001), suggesting that epibionts may act as lures to herbivores, attracting consumers that otherwise would not feed significantly on the seaweed. Foods containing extracts from fouled C. seminervis were preferred by urchins over the alga devoid of epibionts. However, extracts from fouled alga inhibited mussel attachment when compared to epibiont-free alga. Differences might be a direct detrimental effect of the presence of epibionts. On the other hand, epibiosis may induce the production of AF defences in C. seminervis.     

7. Tundra stream macroalgae of North America: composition, distribution and physiological adaptations

Eighty-three infrageneric taxa of stream macroalgae have been reported from tundra regions of North America, composed of 32 cyanobacteria, 35 Chlorophyta, 10 Chrysophyta and 6 Rhodophyta. There are few if any endemics represented in this flora. The most widespread species are the cyanobacteria Rivularia minutula, Nostoc commune and Tolypothrix tenuis as well as asexual populations of the chlorophyte genus Zygnema. The relative contribution of cyanobacteria compared to that of the Chlorophyta increases from the low to high arctic. Number of species per segment ranges from 0 to 7, with a mean of 2.8, and varies little between the low and high arctic. The percentage of stream bottom covered by macroalgae ranges from 0 to ca. 75%; mean cover values for low and high arctic streams are ca. 12 and 8%, respectively. Tundra macroalgae tend to be more abundant and diverse in less rigorously flowing stream sections. Most species tolerate prolonged freezing by forming resistant vegetative cells with thick walls, plentiful reserves and low molecular weight solutes to lower the freezing point. Many tundra stream macroalgae also produce ‘sunscreen’ pigments to reduce exposure to damaging radiation in the blue and ultraviolet regions. Nutrients tend to be low and phosphorus is often limiting in these systems. Arctic streams appear to differ from those of Antarctica in having potential grazers of macroalgae, such as the chironomid Diamesa, the mayfly Baetis and the caddisfly Brachycentrus.     

Increasing dominance of small zooplankton with toxic cyanobacteria

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Impacts of differential consumption by the grazing fish, Plecoglossus altivelis, on the benthic algal composition in the Chikuma River, Japan

Grazing effects of ayu, Plecoglossus altivelis Temminck et Schegel, on the benthic algal assemblages were investigated in the Chikuma River, Japan. Comparison of the algal composition on boulders with and without intensively grazed patches indicated that fish grazing decreased the abundance of diatoms and prostrate filamentous cyanobacteria and caused upright filamentous cyanobacteria to predominate. Differential consumption by ayu was estimated by comparing the relative abundance of algae in the stomach contents of ayu and that in the algal assemblages within the grazed patches. The results showed that ayu consumed the prostrate filamentous cyanobacteria proportionally to their abundance, whereas they ingested diatoms and the upright filamentous cyanobacteria in a larger and lower quantity, respectively, than that expected from their abundance. Differential consumption would involve the change in the algal composition toward the predominance of upright filamentous cyanobacteria under fish grazing conditions.