The role of light availability and herbivory on algal responses to nutrient enrichment in a riparian wetland,Alaska

We investigated how the relative availability of solar radiation in the presence or absence of grazing alters the ability of benthic algae to respond to nutrient enrichment in an Alaskan marsh. We used a factorial mesocosm experiment that included nutrient enrichment (enriched or control), grazing (grazed or ungrazed), and light (unshaded or shaded) to simulate shading by macrophytes early and late in the growing season, respectively. We found stronger effects of grazers and nutrients compared to light on benthic algal biomass and taxonomic composition. Algal biomass increased in nutrient‐enriched treatments and was reduced by grazing. Shading did not have an effect on algal biomass or taxonomic composition, but the concentration of chl a per algal biovolume increased with shading, demonstrating the ability of algae to compensate for changes in light availability. Algal taxonomic composition was more affected by grazer presence than nutrients or light. Grazer‐resistant taxa (basal filaments of Stigeoclonium) were replaced by diatoms (Nitzschia) and filamentous green algae (Ulothrix) when herbivores were removed. The interacting and opposing influences of nutrients and grazing indicate that the algal community is under dual control from the bottom‐up (nutrient limitation) and from the top‐down (consumption by herbivores), although grazers had a stronger influence on algal biomass and taxonomic composition than nutrient enrichment. Our results suggest that low light availability will not inhibit the algal response to elevated nutrient concentrations expected with ongoing climate change, but grazers rapidly consume algae following enrichment, masking the effects of elevated nutrients on algal production.     

Interactions among irradiance,nutrients, and herbivores constrain a stream algal community

Using stream-side, flow-through channels, I tested for the effects of nutrients (NU) (nitrogen plus phosphorus), irradiance (L), and snail grazing (G) on a benthic algal community in a small, forested stream. Grazed communities were-dominated by a chlorophyte (basal cells ofStigeoclonium) and a cyanophyte (Chamaesiphon investiens), whereas ungrazed communities were comprised almost entirely of diatoms, regardless of nutrient and light levels. Snails maintained low algal biomass in all grazed treatments, presumably by consuming increased algal production in treatments to which L and NU were increased. When nutrients were increased, cellular nutrient content increased under ambient conditions (shaded, grazed) and biomass and productivity increased when snails were removed and light was increased. Together, nutrients and light had positive effects and grazing had negative effects on biomass (chlorophylla, AFDM, algal biovolume) and chlorophyll-and areal-specific productivity in ANOVAs. However, in most cases, only means from treatments in which all three factors were manipulated (ungrazed, +NU&L treatments) were significantly different from controls; effects of single factors were generally undetectable. These results indicate that all three factors simultaneously limited algal biomass and productivity in this stream during the summer months. Additionally, the effects of these factors in combination were in some cases different from the effects of single factors. For example, light had slight negative effects on some biomass parameters when added at ambient snail densities and nutrient concentrations, but had strong positive effects in conjunction with nutrient addition and snail removal. This study demonstrates that algal biomass and productivity can be under multiple constraints by irradiance, nutrients, and herbivores and indicates the need to employ multifactor experiments to test for such interactive effects.     

Gastropod grazers and nutrients,but not light,interact in determining periphytic algal diversity

The potential interactions of grazing, nutrients and light in influencing autotroph species diversity have not previously been considered. Earlier studies have shown that grazing and nutrients interact in determining autotroph species diversity, since grazing decreases species diversity when nutrients (i.e. N or P) limit autotroph growth, but increases it when nutrients are replete. We hypothesized that increased light intensities would intensify the interactions between grazing and nutrients on algal species diversity, resulting in even stronger reductions in algal species diversity through grazing under nutrient–poor conditions, and to even stronger increases of algal species diversity through grazing under nutrient-rich conditions. We studied the effects of grazing (absent, present), nutrients (ambient, N + P enriched) and light (low light, high light) on benthic algal diversity and periphyton C:nutrient ratios (which can indicate algal nutrient limitation) in a factorial laboratory experiment, using the gastropod grazer Viviparus viviparus. Grazing decreased algal biomass and algal diversity, but increased C:P and N:P ratios of periphyton. Grazing also affected periphyton species composition, by decreasing the proportion of Spirogyra sp. and increasing the proportion of species in the Chaetophorales. Grazing effects on diversity as well as on periphyton N:P ratios were weakened when nutrients were added (interaction between grazing and nutrients). Chlorophyll a (Chl a) per area increased with nutrient addition and decreased with high light intensities. Light did not increase the strength of the interaction between grazing and nutrients on periphytic algal diversity. This study shows that nutrient addition substantially reduced the negative effects of grazing on periphytic algal diversity, whereas light did not interact with grazing or nutrient enrichment in determining periphytic algal diversity.     

Grazer control of nutrient availability in the periphyton

Summary Benthic algal assemblages are regulated by both abiotic (e.g., nutrient) and biotic (e.g., grazing) constraint. The objective of this study was to determine how changes in these two factors affected the structure of an algal assemblage in an ephemeral stream. Coverslips were incubated for 21 days in enclosures containing one of three nutrient environments (ambient, phosphorus-enriched, or phosphorus and nitrogen enriched) and one of four densities of the snail Gonibasis (0, 40, 80, or 120 snails/m²) and examined directly to enumerate the algal assemblage. The effect of grazing on algal biomass was dependent on the nutrient environment. An overstory of diatoms was susceptible to removal by grazing and was not strongly affected by nutrient enrichment. An understory of Stigeoclonium was more resistant to grazing and responded strongly to nutrient enrichment only in the presence of grazers. Snail grazers may mediate nutrient availability to the understory indirectly by removing overlying cells or by direct excretion of nutrients. Multiple interactions occur between benthic herbivores and algae, and, as shown here, some of them are positive and involve modifications of the nutrient environment.     

Independent and interactive effects of nutrients and grazers on benthic algal community structure

Algal responses to nutrients, grazing by Helicopsyche borealis, and concurrent grazing by Helicopsyche and Baetis tricaudatus were examined in recirculating stream chambers. Alagl communities, dominated by Achnanthes minutissima, Cocconeis placentula, and Synedra ulna, were primarily phosphorus-limited. Algal populations responded after only 6 days of nutrient enrichment. Initially, both the adnate diatom Cocconeis and erect diatom Synedra showed positive response to nutrient enrichment. Accumulation of algal biomass between day 3 and 6 in the P enriched treatment was resulted primarily from the growth of Synedra, an overstory rosette-like diatom colony. Such a shift in dominant growth from adnate to erect diatoms is a general phenomenon in periphyton succession in the absence of disturbance. Algal species showed differential responses to an increase of Helicopsyche densities. The accrual rate of Achnanthes continuously decreased with increasing grazer densities. The accrual rates of both Cocconeis and Synedra declined but reached plateaus between medium and high grazing densities. Baetis effectively and exclusively depressed Synedra and had no significant impact on Cocconeis. After concurrent grazing, algal communities were mainly dominated by Cocconeis (approximately 80% of total algal biovolume). The grazer' s mouth structures, grazing efficiencies, and mobility may account for the differential effects of concurrent grazing on algal communities. Significant interactive effects of P and grazing by Helicopsyche indicated that both nutrient addition and grazing may exert significant impact on algal communities. However, grazing may have a much stronger effect on algae than nutrients. Our results indicate that enhancement of algal biomass by P was dampened by grazing activities and that P had no effect on algal biomass in the presence of grazers.     

Impact of Urban Pollution from the Hanoi Area on Benthic Diatom Communities Collected from the Red, Nhue and Tolich Rivers (Vietnam)

The effects of urban pollution from Hanoi city on the benthic diatom communities of the Nhue–Tolich river system were studied during the 2003 dry season. Benthic diatoms were allowed to grow on glass slides suspended in the water flow for 4 weeks. To reveal the relationship between water quality and diatom communities, Canonical correspondence analysis (CCA) was used on data concerning relative abundances of diatom species and environmental variables. Two diatom indices, IPS and DAIpo, were applied to evaluate water quality in the three rivers. A total of 291 diatom taxa were found in the Red, Nhue and Tolich Rivers. These were mainly cosmopolitan taxa, with some tropical, subtropical and endemic taxa. The most abundant taxa at the Red site were Aulacoseira granulata, Achnanthidium minutissimum, Encyonema minutum, Navicula recens and other halophilous taxa such as Nitzschia kurzii, Seminavis strigosa, Entomoneis paludosa, Bacillaria paradoxa. Diatom assemblages at the Tolich site consisted mainly of Nitzschia umbonata, Nitzschia palea and Eolimna minima. Diatom density ranged from 660 to 30,000 cells/cm². Environmental variables and diatom assemblage composition at all sites were significantly correlated. Two diatom indices gave similar results and indicate the Tolich River with the lowest values as a highly polluted site.     

Diatoms as indicators: The influences of experimental nitrogen enrichment on diatom assemblages in sub-Arctic streams

Indices using diatoms are widely used to assess water quality, but are usually constructed from field correlations and not tested through rigorous experimentation. We tested experimentally the performance of the Sørensen and the Shannon indices, and the trophic diatom index (TDI). Nitrogen was naturally limiting in the eight remote sub-Arctic streams used and we measured the effects of experimental nitrogen enrichment on diatom assemblages. Diatom densities increased significantly in the enriched reaches but there was no significant difference in invertebrate density between control and treatment reaches. Grazing effects were thus controlled for. Diversity within streams (Shannon index) was significantly reduced by nutrient addition but the Sørensen index did not change. The trophic diatom index (TDI), which is presumed to reflect nutrient concentration, was not influenced by nutrient addition and generally the values were low in both control and treatment reaches. Densities of the diatom genera Achnanthes and Gomphonema increased significantly with enrichment while those of Nitzschia and Fragilaria decreased significantly. Less abundant diatom species, which collectively constituted around 40% in relative abundance in the control reaches, were around 15–18% in treatment reaches. Growth forms were altered by the nutrients. Diatoms attached by mucilage pads were more abundant in treated reaches compared with control reaches. Motile diatoms became scarcer. The size of diatom species was unaffected by nutrient enrichment. This study showed that it is important to test experimentally indices that are developed for particular habitats before using them elsewhere.     

Ingestion and digestion of epilithic algae by larval insects in a heavily grazed montane stream

1. Epilithic algae grown on elevated or non-elevated ceramic tiles were exposed (to produce assemblages with different grazing histories) in a heavily grazed, montane stream in New Mexico, U.S.A. to Ameletus nymphs (Ephemeroptera) and Ecclisomyia larvae (Trichoptera) and the algal composition in insect faeces was compared to that on the tiles. Differences in grazing and digestion efficiency between grazers were then assessed and also differences in susceptibility to ingestion and digestibility among common algae. 2. Ordination of tile and faecal samples, using the relative abundance of common algae, revealed that: (i) algal assemblages on elevated vs. non-elevated tiles differed only slightly; (ii) the taxonomic composition of algae in faeces of both caddis and mayflies differed substantially from that on the tiles, indicating low grazing efficiency for some algal taxa; and (iii) the algal composition of faeces produced by caddis larvae and mayflies was similar, indicating little difference in grazing efficiency between them. However, some algal taxa were more susceptible to ingestion by caddisfly larvae when occurring on elevated tiles than on non-elevated tiles, suggesting that previous exposure to caddis grazing influenced assemblage attributes. 3. Although Ameletus and Ecclisomyia differed little in grazing efficiency, the percentage of diatoms that were dead after passage through the gut was greatest in the mayfly treatment, suggesting that mayflies digested diatoms more efficiently than the caddis. Analyses of differences in the condition of chloroplasts within diatoms in tile and faecal samples showed that losses of ’live‘ diatom cells (i.e. those containing full chloroplasts) during gut passage through mayflies equalled the increase, in faeces, of ’dead‘ (empty frustules) cells of all common diatoms. In contrast, some diatoms were digested inefficiently by caddis larvae. 4. Algae on elevated tiles contained a higher proportion of dead diatoms than those on non-elevated tiles, possibly because mayflies visited raised tiles more often and, consequently, ingested and defaecated cells at a higher rate in the absence of caddis larvae. Moreover, diatom taxa differed in the percentage of cells that were dead within tile assemblages, with populations of typically grazer-resistant taxa (e.g. Achnanthidium minutissimum, Planothidium lanceolatum and Cocconeis placentula var. euglypta) containing significantly more dead cells than grazer-susceptible taxa [e.g. small, chain-forming Fragilaria (= Staurosirella)]. This result suggests that a trade-off exists between ingestion vs. digestion resistance of microalgae. Both the ingestion and digestion efficiency of algivorous macroinvertebrates could influence the structure and function of algal assemblages. In heavily grazed systems, where algal cells are probably processed through grazer guts repeatedly, differential resistance to digestion among algae may be particularly important.     

Effects of consumers and enrichment on abundance and diversity of benthic algae in a rocky intertidal community

Human alteration of nutrient cycling and the densities of important consumers have intensified the importance of understanding how nutrients and consumers influence the structure of ecological systems. We examined the effects of both grazing and nutrient enrichment on algal abundance and diversity in a high-intertidal limpet-macroalgal community on the South Island of New Zealand, a relatively nutrient-poor environment. We used a fully factorial design with three levels each of grazing (manipulations of limpet and snail densities) and nutrients (nutrient-diffusers attached to the rock). Top-down control by grazers appears to be the driving organizing mechanism for algal communities in this system, with strong negative effects of grazing on algal diversity and abundance across all levels of nutrient enrichment. However, in contrast to the conclusions drawn from the analysis of the whole algal community, there was an interactive effect of grazing and enrichment on foliose algae, an important component of the algal system. When herbivory was reduced to very low levels, enrichment generated increases in the abundance and biomass of foliose algae. As expected, top-down control was the primary determinant of algal community structure in this system, controlling abundance and diversity of macrophytes on the upper shore. Contrary to expectations, however, increased nutrients had no community-wide effects, although foliose algal abundance increases were greatest with high nutrients and reduced grazing. It seems likely that most of the corticated algal species have limited capacity to respond to nutrient pulses in this nutrient-poor environment.     

Assessment of ecological status in U.K. rivers using diatoms

1. The European Union's Water Framework Directive requires all water bodies to achieve ‘good ecological status’ by 2015 and this paper describes a rationale for defining ‘good ecological status’ based on diatoms, a significant component of the biological quality element ‘macrophyte and phytobenthos’. 2. A database of benthic diatom samples collected over the past 20 years was assembled. New sampling, specifically for this project, was undertaken during 2004 to supplement these data. In total 1051 samples were included in the database with matching environmental data. 3. ‘Reference sites’, relatively unimpacted by human activity, were selected from this database by a series of screening steps and these sites were used to develop a site‐specific reference typology. 4. Environmental variables not related to the pressure gradient were used to predict the ‘expected’ Trophic Diatom Index (TDI) values at each site. Site‐specific TDI predictions were used to generate ecological quality ratios (EQRs) ranging from ≥1, where the diatom assemblage showed no impact, to (theoretically) 0, when the diatom assemblage was indicative of major anthropogenic activities. 5. The boundary between ‘high’ and ‘good’ status was defined as the 25th percentile of EQRs of all reference sites. The boundary between ‘good’ and ‘moderate’ status was set at the point at which nutrient‐sensitive and nutrient‐tolerant taxa were present in equal relative abundance. An ecological rationale for this threshold is outlined in the paper.     

Comparing Effects of Nutrients on Algal Biomass in Streams in Two Regions with Different Disturbance Regimes and with Applications for Developing Nutrient Criteria

Responses of stream algal biomass to nutrient enrichment were studied in two regions where differences in hydrologic variability cause great differences in herbivory. Around northwestern Kentucky (KY) hydrologic variability constrains invertebrate biomass and their effects on algae, but hydrologic stability in Michigan (MI) streams permits accrual of high herbivore densities and herbivory of benthic algae. Multiple indicators of algal biomass and nutrient availability were measured in 104 streams with repeated sampling at each site over a 2−month period. Many measures of algal biomass and nutrient availability were positively correlated in both regions, however the amount of variation explained varied with measures of biomass and nutrient concentration and with region. Indicators of diatom biomass were higher in KY than MI, but were not related to nutrient concentrations in either region. Chl a and % area of substratum covered by Cladophora were positively correlated to nutrient concentrations in both regions. Cladophora responded significantly more to nutrients in MI than KY. Total phosphorus (TP) and total nitrogen (TN) explained similar amounts of variation in algal biomass, and not significantly more variation in biomass than dissolved nutrient concentrations. Low N:P ratios in the benthic algae indicated N as well as P may be limiting their accrual. Most observed responses in benthic algal biomass occurred in nutrient concentrations between 10 and 30 μg TP l⁻¹ and between 400 and 1000 μg TN l⁻¹.     

GRACILARIA EDULIS (RHODOPHYTA) AS A BIOLOGICAL INDICATOR OF PULSED NUTRIENTS IN OLIGOTROPHIC WATERS

The response of the marine macroalga Gracilaria edulis (Gmelin) Silva to nutrient pulses of varying magnitude was investigated to test its applicability as a marine bioindicator at two oligotrophic locations. After exposure to nutrient pulses, algal amino acid, tissue nitrogen, and chlorophyll a content were assessed relative to algae incubated under control conditions (no nutrient enrichment). The smallest nutrient pulse involved a nutrient enrichment experiment conducted within a coral atoll, whereas two larger pulses resulted from sewage discharge to a tropical coastal bay. After exposure to the smallest nutrient pulse (10 × ambient), only changes in macroalgal amino acid concentration and composition were detected (mainly as increases in citrulline). At 100 × ambient concentrations, increases in tissue % nitrogen of the macroalgae were detected, in addition to responses in amino acids. Macroalgae exposed to the highest nutrient pulse (1000 × ambient) responded with increased chlorophyll a , tissue nitrogen, and amino acids within the three day incubation period. In contrast to these algal responses, analytical water sampling techniques failed to detect elevated nutrients when nutrient pulses were not occurring. The responses of this algal bioindicator to variable nutrient pulses may provide a useful tool for investigating the source and geographical extent of nutrients entering oligotrophic coastal waters.     

Interactive effects of grazing and enrichment on diversity; conceptual implications of a rocky intertidal experiment

Understanding the ways in which consumers and productivity act and interact to yield differences in diversity is of primary conceptual and pragmatic importance in a world in which humans are simultaneously changing ecological communities and substantially altering the availability of nutrients. Here, I used macroalgal communities on rocky reefs to examine the effects of both limpet grazing and nutrient enrichment on algal diversity throughout almost two years of succession. The experimental design included three levels each of grazing and nutrients, with unglazed terracotta pots attached to the rock as replicate plots in a high intertidal limpet-macroalgal community. Grazing effects varied by year. During the first year, grazing effects were context-dependent with limpets resulting in lower species richness, especially at the highest level of limpet density. However, at this highest level of limpet density, high enrichment counteracted the negative effect of limpets such that diversity was similar to that in treatments with lower limpet densities. In the second year, grazing generally decreased richness, regardless of enrichment. The results of this experiment are partially consistent with the grazer-reversal hypothesis – grazing decreased richness in low nutrient conditions and this effect was neutralized (rather than reversed) under high enrichment. Inconsistencies with model predictions may be explained by the apparent unresponsiveness of algal productivity to experimental enrichment, the unique substrate-scraping feeding mechanisms of limpets, and potentially limited propagule supply.     

Changes in diatom-dominated biofilms during simulated improvements in water quality: implications for diatom-based monitoring in rivers

Although benthic diatoms are used to assess river water quality, there are few data on the rate at which diatom assemblages react to changes in water quality. The aim of this study was to assess the reaction time of diatoms and to discuss the changes occurring during water quality improvement on the basis of their autecological characteristics. In order to simulate this improvement, diatom-dominated biofilms grown on artificial sandstone substrata were transferred from several polluted rivers to an unpolluted river. They were sampled three times: before transfer and 1 and 2 months after transfer. The ecology and growth-forms of the taxa explained most of the changes in species composition observed during the experiment. Adnate diatoms gradually replaced motile and stalked taxa. Gomphonema parvulum, a stalked diatom positioned vertically in the biofilm, is adapted for light and space competition in high-density algal biofilms. When transferred to an unpolluted site, this growth-form is less competitive and does not tolerate the high grazing pressure. Fistulifera saprophila is a single celled motile diatom, living in organic matrices. When the artificial substrata were transferred to the unpolluted site, this particular ecological niche disappeared quickly. On the other hand, Achnanthidium minutissimum, which is considered to be cosmopolitan and an early colonizer, increased during the first month of transfer and then decreased. It was gradually replaced by A. biasolettianum, which was the taxon best suited to this pristine stream. The changes observed differed between treatments depending on the species composition and architecture of the biofilms. In particular, biofilms dominated by stalked and motile diatoms were more quickly modified than those dominated by small motile diatoms. The diatom index reflects these changes, and its values showed that about 60 days following a water quality improvement were necessary for transferred diatom assemblages to reach diatom index values similar as those at the unpolluted river.     

Top-down and bottom-up control in an eelgrass–epiphyte system

Nutrient supply and the presence of grazers can control primary producers in aquatic ecosystems, but the relative importance of bottom-up and top-down effects remains inconclusive. We conducted a mesocosm experiment and a field study to investigate the independent and interactive effects of nutrient enrichment and grazing on primary producers in an eelgrass bed Zostera marina . Nutrient treatments consisted of ambient or enriched (2× and 4× ambient) concentrations of inorganic nitrogen and phosphate. Grazer treatments consisted of presence or absence of field densities of the common isopod Idotea baltica . We found strong and interacting effects of nutrients and grazing on epiphytes. Epiphyte biomass and productivity were enhanced by nutrient enrichment and decreased in the presence of grazers. The absolute amount of epiphyte biomass consumed by grazers increased under high nutrient supply, and thus, nutrient effects were stronger in the absence of grazing. The effects of grazers and fertilisation on epiphyte composition were antagonistic: chain-forming diatoms and filamentous algae profited from nutrient enrichment, but their proportions were reduced by grazing. Eelgrass growth was positively affected by grazing and by nutrient enrichment at moderate nutrient concentrations. High nutrient supply reduced eelgrass productivity compared to moderate nutrient conditions. The monthly measured field data showed a nitrogen limitation for epiphytes and eelgrass in summer, which may explain the positive effect of nutrient enrichment on both primary producers. Generally, the field data suggested the possibility of seasonally varying importance of bottom-up and top-down control on primary producers in this eelgrass system.     

Effects of grazer immigration and nutrient enrichment on an open algae-grazer system

After disturbance, recovery dynamics of local populations depend on arrival rates of immigrants and local growth conditions. We studied the effects of herbivore immigration rates and nutrient enrichment on the dynamics of grazing insect larvae, benthic microalgae, and filamentous macroalgae recovering from low local densities in an open stream system. The two types of algae approximate a trade‐off between capabilities for growing at low resource levels and resisting herbivory. Many microalgae achieve relatively high growth rates at low nutrient levels but are vulnerable to grazers, whereas many macroalgae require high nutrient levels for growth but become increasingly defended with filament growth. We hypothesized that macroalgae should benefit more strongly than microalgae from increasing nutrient levels and decreasing grazer immigration rates, because both conditions increase macroalgal chances to grow into a size refuge from herbivory. We created a gradient of nutrient concentrations and manipulated drift immigration rates of macroinvertebrates. Macro‐ and microalgal biomass and the relative contribution of macroalgae to total algal biomass increased with increasing nutrient enrichment and decreased with increasing grazer immigration. Grazer densities responded positively to nutrient enrichment. The densities of large baetids responded positively to higher immigration rates of large baetids, whereas small baetids and chironomid larvae showed the opposite response. Per capita emigration of small baetids decreased with increasing algal biomass. The data suggest that large baetids negatively affected algal biomass and that small baetid and chironomid densities tracked resource levels set by nutrient enrichment and large baetids. Our experiments highlight the prospects of integrating disturbance with nutrient supply, immigration rates and local trophic interactions (determining recovery trajectories) into conceptual models of open system dynamics. We suggest that recovery trajectories towards micro‐ or macroalgal dominated states may depend on the spatial scale of disturbance relative to the movement ranges of migrating grazers and to nutrient supply.     

The effects of nutrient enrichment on a freshwater meiofaunal assemblage

1. The effects of eutrophication on phytoplankton, zooplankton and fish in lakes are well known. By contrast, little is known about the response of the zoobenthos to nutrient enrichment, while smaller organisms, such as the meiofauna, have for the most part been neglected. 2. In a long‐term (16 months) microcosm experiment, we assessed the effects of five levels of nutrients [total phosphorus (TP), 7–250 μg L^?1; nitrate, 2–8 mg L^?1] on a freshwater meiofaunal assemblage and on nematode diversity in particular. 3. Within the first 8 months, meiofaunal succession was only weakly affected, whereas, during the last 4 months, nutrient addition influenced most of the main taxa, with a concomitant change in the assemblage structure. 4. The density of the numerically dominant nematodes decreased upon nutrient enrichment, whereas ostracods became more numerous. Other taxa, including copepods, reached a maximum at intermediate nutrient levels or, in case of oligochaetes, were almost unaffected by nutrient enrichment. However, the changes in the density of the main taxa were usually insufficient to alter their biomass. Consequently, meiofaunal biomass was remarkably unresponsive to nutrient addition, while meiofaunal density displayed a unimodal relationship, with a peak at a TP concentration of 30 μg L^?1. In addition, nematode species richness decreased significantly with increasing nutrient concentrations. 5. We hypothesise that the response of meiofaunal taxa to nutrients is attributable to the development of primary producers, which shifted with enrichment from low densities of edible diatoms and unicellular green algae to large standing stocks of inedible forms, such as Lemna minor and Cladophora spp.     

Nutrient loading and grazing by the minnow Phoxinus erythrogaster shift periphyton abundance and stoichiometry in mesocosms

1. Anthropogenic activities in prairie streams are increasing nutrient inputs and altering stream communities. Understanding the role of large consumers such as fish in regulating periphyton structure and nutritional content is necessary to predict how changing diversity will interact with nutrient enrichment to regulate stream nutrient processing and retention. 2. We characterised the importance of grazing fish on stream nutrient storage and cycling following a simulated flood under different nutrient regimes by crossing six nutrient concentrations with six densities of a grazing minnow (southern redbelly dace, Phoxinus erythrogaster) in large outdoor mesocosms. We measured the biomass and stoichiometry of overstory and understory periphyton layers, the stoichiometry of fish tissue and excretion, and compared fish diet composition with available algal assemblages in pools and riffles to evaluate whether fish were selectively foraging within or among habitats. 3. Model selection indicated nutrient loading and fish density were important to algal composition and periphyton carbon (C): nitrogen (N). Nutrient loading increased algal biomass, favoured diatom growth over green algae and decreased periphyton C : N. Increasing grazer density did not affect biomass and reduced the C : N of overstory, but not understory periphyton. Algal composition of dace diet was correlated with available algae, but there were proportionately more diatoms present in dace guts. We found no correlation between fish egestion/excretion nutrient ratios and nutrient loading or fish density despite varying N content of periphyton. 4. Large grazers and nutrient availability can have a spatially distinct influence at a microhabitat scale on the nutrient status of primary producers in streams.     

The effects of top–down versus bottom–up control on benthic coral reef community structure

While climate change and associated increases in sea surface temperature and ocean acidification, are among the most important global stressors to coral reefs, overfishing and nutrient pollution are among the most significant local threats. Here we examined the independent and interactive effects of reduced grazing pressure and nutrient enrichment using settlement tiles on a coral-dominated reef via long-term manipulative experimentation. We found that unique assemblages developed in each treatment combination confirming that both nutrients and herbivores are important drivers of reef community structure. When herbivores were removed, fleshy algae dominated, while crustose coralline algae (CCA) and coral were more abundant when herbivores were present. The effects of fertilization varied depending on herbivore treatment; without herbivores fleshy algae increased in abundance and with herbivores, CCA increased. Coral recruits only persisted in treatments exposed to grazers. Herbivore removal resulted in rapid changes in community structure while there was a lag in response to fertilization. Lastly, re-exposure of communities to natural herbivore populations caused reversals in benthic community trajectories but the effects of fertilization remained for at least 2 months. These results suggest that increasing herbivore populations on degraded reefs may be an effective strategy for restoring ecosystem structure and function and in reversing coral–algal phase-shifts but that this strategy may be most effective in the absence of other confounding disturbances such as nutrient pollution.     

The effects of nutrient enrichment and herbivore abundance on the ability of turf algae to overgrow coral in the Caribbean

Turf algae are multispecies communities of small marine macrophytes that are becoming a dominant component of coral reef communities around the world. To assess the impact of turf algae on corals, we investigated the effects of increased nutrients (eutrophication) on the interaction between the Caribbean coral Montastraea annularis and turf algae at their growth boundary. We also assessed whether herbivores are capable of reducing the abundance of turf algae at coral-algae boundaries. We found that turf algae cause visible (overgrowth) and invisible negative effects (reduced fitness) on neighbouring corals. Corals can overgrow neighbouring turf algae very slowly (at a rate of 0.12 mm 3 wk⁻¹) at ambient nutrient concentrations, but turf algae overgrew corals (at a rate of 0.34 mm 3 wk⁻¹) when nutrients were experimentally increased. Exclusion of herbivores had no measurable effect on the rate turf algae overgrew corals. We also used PAM fluorometry (a common approach for measuring of a colony''s “fitness”) to detect the effects of turf algae on the photophysiology of neighboring corals. Turf algae always reduced the effective photochemical efficiency of neighbouring corals, regardless of nutrient and/or herbivore conditions. The findings that herbivores are not capable of controlling the abundance of turf algae and that nutrient enrichment gives turf algae an overall competitive advantage over corals together have serious implications for the health of Caribbean coral reef systems. At ambient nutrient levels, traditional conservation measures aimed at reversing coral-to-algae phase shifts by reducing algal abundance (i.e., increasing herbivore populations by establishing Marine Protected Areas or tightening fishing regulations) will not necessarily reduce the negative impact of turf algae on local coral communities. Because turf algae have become the most abundant benthic group on Curaçao (and likely elsewhere in the Caribbean), new conservation strategies are required to mitigate their negative impact on coral communities.