The Response of Experimental Rocky Shore Communities to Nutrient Additions

The aim of this study was to determine whether the experimental nutrient enrichment of littoral rocky shore communities would be followed by a predicted accumulation of fast-growing opportunistic algae and a subsequent loss of perennial benthic vegetation. Inorganic nitrogen (N) and potassium (P) was added to eight concrete mesocosms inhabited by established littoral communities dominated by fucoids. The response to nutrient enrichment was followed for almost 2 1/2 years. Fast-growing opportunistic algae (periphyton and ephemeral green algae) grew significantly faster in response to nutrient enrichment, but the growth of red filamentous algae and large perennial brown algae was unaffected. However, these changes were not followed by comparable changes in the biomass and composition of the macroalgae. The biomass of opportunistic algae was stimulated only marginally by the nutrient enrichment, and perennial brown algae (fucoids) remained dominant in the mesocosm regardless of nutrient treatment level. Established rocky shore communities thus seem able to resist the effects of heavy nutrient loading. We found that the combined effects of the heavy competition for space and light imposed by canopy-forming algae, preferential grazing on opportunistic algae by herbivores, and physical disturbance, succeeded by a marked export of detached opportunistic algae, prevented the fast-growing algae from becoming dominant. However, recruitment studies showed that the opportunistic algae would become dominant when free space was available under conditions of high nutrient loading and low grazing pressure. These results show that established communities of perennial algae and associated fauna in rocky shore environments can prevent or delay the accumulation of bloom-forming opportunistic algae and that the replacement of long-lived macroalgae by opportunistic species at high nutrient loading may be a slow process. Nutrient enrichment may not, in itself, be enough to stimulate structural changes in rocky shore communities.     

Stressor intensity determines antagonistic interactions between species invasion and multiple stressor effects on ecosystem functioning

Biological invasions, nutrient enrichment and ocean warming are known to threaten biodiversity and ecosystem functioning. The independent effects of these ecological stressors are well studied, however, we lack understanding of their cumulative effects, which may be additive, antagonistic or synergistic. For example, the impacts of biological invasions are often determined by environmental context, which suggests that the effects of invasive species may vary with other stressors such as pollution or climate change. This study examined the effects of an invasive seaweed (Sargassum muticum) on the structure and functioning of a synthetic macroalgal assemblage and tested explicitly whether these effects varied with nutrient enrichment and ocean warming. Overall, the presence of S. muticum increased assemblage productivity rates and warming altered algal assemblage structure, which was characterised by a decrease in kelp and an increase in ephemeral green algae. The effects of S. muticum on total algal biomass accumulation, however, varied with nutrient enrichment and warming, producing antagonistic cumulative effects on total algal biomass accumulation. These findings show that the nature of stressor interactions may vary with stressor intensity and among response variables, which leads to less predictable consequences for the structure and functioning of communities.     

Responses to nutrient enrichment,wave action and disturbance in rocky shore communities

A high degree of resistance against nutrient enrichment has previously been demonstrated for macroalgal-dominated rocky shore communities in the presence of moderate to large amounts of macroinvertebrate grazers. To experimentally examine, under controlled conditions, the possible roles for this resistance of two other factors, i.e. disturbance (presence/absence of the macroalgal canopy itself) and wave action, the canopy algae and associated algal and animal assemblages were removed by scraping from approximately one third of the area of eight littoral mesocosms, subjected to two different wave action regimes. After this, excessive nutrients were added to four mesocosms with the factor nutrients fully crossed with the factor wave action with two replicate mesocosm basins of each nutrient/wave treatment combination. Disturbance was added to the design as a within-basin factor thus making up a split-plot experiment. The abundance of grazers was allowed to vary freely and under the influence of the treatments. After 11 summer weeks, there were significant differences in community structure between nutrient enrichment levels for both algal and animal assemblages when examined by multivariate statistical techniques. Univariate analyses confirmed a significantly stimulated colonisation by green algae, mainly Ulva lactuca, in both disturbed (scraped) and undisturbed areas of nutrient-enriched mesocosms. In un-enriched mesocosms, the green algae were absent from undisturbed areas and rare in disturbed areas, where mainly brown Ectocarpus spp. and red algae had settled. Among the macrofauna, the total abundance of grazers was stimulated in nutrient-enriched mesocosms with individuals of the amphipod genus Gammarus and the isopod genus Jaera being especially numerous. With regard to wave action, no significant differences occurred in community structure, although there were indications of significant nutrient × wave effects for both the amount of exported red algae and the amount of accumulated brown algae. The study shows that eutrophication-related community shifts on rocky shores may occur very rapidly, regardless of the level of wave-energetic stress and the abundance of grazers, if the nutrient concentrations are high and the colonisation and growth of opportunistic algae are facilitated by disturbance such as (naturally or anthropogenically driven) canopy gap forming processes.     

Responses to nutrient enrichment,wave action and disturbance in rocky shore communities

A high degree of resistance against nutrient enrichment has previously been demonstrated for macroalgal-dominated rocky shore communities in the presence of moderate to large amounts of macroinvertebrate grazers. To experimentally examine, under controlled conditions, the possible roles for this resistance of two other factors, i.e. disturbance (presence/absence of the macroalgal canopy itself) and wave action, the canopy algae and associated algal and animal assemblages were removed by scraping from approximately one third of the area of eight littoral mesocosms, subjected to two different wave action regimes. After this, excessive nutrients were added to four mesocosms with the factor nutrients fully crossed with the factor wave action with two replicate mesocosm basins of each nutrient/wave treatment combination. Disturbance was added to the design as a within-basin factor thus making up a split-plot experiment. The abundance of grazers was allowed to vary freely and under the influence of the treatments. After 11 summer weeks, there were significant differences in community structure between nutrient enrichment levels for both algal and animal assemblages when examined by multivariate statistical techniques. Univariate analyses confirmed a significantly stimulated colonisation by green algae, mainly Ulva lactuca, in both disturbed (scraped) and undisturbed areas of nutrient-enriched mesocosms. In un-enriched mesocosms, the green algae were absent from undisturbed areas and rare in disturbed areas, where mainly brown Ectocarpus spp. and red algae had settled. Among the macrofauna, the total abundance of grazers was stimulated in nutrient-enriched mesocosms with individuals of the amphipod genus Gammarus and the isopod genus Jaera being especially numerous. With regard to wave action, no significant differences occurred in community structure, although there were indications of significant nutrient × wave effects for both the amount of exported red algae and the amount of accumulated brown algae. The study shows that eutrophication-related community shifts on rocky shores may occur very rapidly, regardless of the level of wave-energetic stress and the abundance of grazers, if the nutrient concentrations are high and the colonisation and growth of opportunistic algae are facilitated by disturbance such as (naturally or anthropogenically driven) canopy gap forming processes.     

Identifying the interacting roles of stressors in driving the global loss of canopy‐forming to mat‐forming algae in marine ecosystems

Identifying the type and strength of interactions between local anthropogenic and other stressors can help to set achievable management targets for degraded marine ecosystems and support their resilience by identifying local actions. We undertook a meta‐analysis, using data from 118 studies to test the hypothesis that ongoing global declines in the dominant habitat along temperate rocky coastlines, forests of canopy‐forming algae and/or their replacement by mat‐forming algae are driven by the nonadditive interactions between local anthropogenic stressors that can be addressed through management actions (fishing, heavy metal pollution, nutrient enrichment and high sediment loads) and other stressors (presence of competitors or grazers, removal of canopy algae, limiting or excessive light, low or high salinity, increasing temperature, high wave exposure and high UV or CO₂), not as easily amenable to management actions. In general, the cumulative effects of local anthropogenic and other stressors had negative effects on the growth and survival of canopy‐forming algae. Conversely, the growth or survival of mat‐forming algae was either unaffected or significantly enhanced by the same pairs of stressors. Contrary to our predictions, the majority of interactions between stressors were additive. There were however synergistic interactions between nutrient enrichment and heavy metals, the presence of competitors, low light and increasing temperature, leading to amplified negative effects on canopy‐forming algae. There were also synergistic interactions between nutrient enrichment and increasing CO₂ and temperature leading to amplified positive effects on mat‐forming algae. Our review of the current literature shows that management of nutrient levels, rather than fishing, heavy metal pollution or high sediment loads, would provide the greatest opportunity for preventing the shift from canopy to mat‐forming algae, particularly in enclosed bays or estuaries because of the higher prevalence of synergistic interactions between nutrient enrichment with other local and global stressors, and as such it should be prioritized.     

Nutrient Addition to Experimental Rocky Shore Communities Revisited: Delayed Responses, Rapid Recovery

Coastal eutrophication may alter the dominance patterns of marine macroalgae, with potential consequences for the associated fauna and the entire ecosystem. Benthic macroalgae and animals in control and nutrient-enriched mesocosms were monitored to investigate eutrophication-induced changes in rocky shore communities. During a 3-year project, nutrient addition had only minor effects on the community structure, such as increased cover and biomass of green Ulva spp. and increased abundance of certain animal species at high nutrient levels. This study is a 4-year extension of a previously reported project, with 2 extra years of effect studies (altogether 5 years) and a subsequent 2 years for recovery. During the 4th year of nutrient enrichment, the cover of Fucus vesiculosus and Fucus serratus started to decline. In the 5th year, these canopy species crashed and there was an evident take-over by green algae at high nutrient addition levels. The previously observed abundance stimulation for fauna disappeared later in the time series, probably due to the loss of the macroalgal canopy. After less than 2 years on regular seawater, the algal and animal communities had returned to within the range of normal variability. The results indicate that established rocky shore communities of perennial algae with associated fauna are able to persist for several years, even at very high nutrient levels, but that community shifts may suddenly occur if eutrophication continues. They also indicate that rocky shore communities have the ability to return rapidly to natural undisturbed conditions after the termination of nutrient enhancement.     

Understory Plant Community Composition Is Associated with Fine-Scale Above- and Below-Ground Resource Heterogeneity in Mature Lodgepole Pine (Pinus contorta) Forests

Understory plant communities play critical ecological roles in forest ecosystems. Both above- and below-ground ecosystem properties and processes influence these communities but relatively little is known about such effects at fine (i.e., one to several meters within-stand) scales, particularly for forests in which the canopy is dominated by a single species. An improved understanding of these effects is critical for understanding how understory biodiversity is regulated in such forests and for anticipating impacts of changing disturbance regimes. Our primary objective was to examine the patterns of fine-scale variation in understory plant communities and their relationships to above- and below-ground resource and environmental heterogeneity within mature lodgepole pine forests. We assessed composition and diversity of understory vegetation in relation to heterogeneity of both the above-ground (canopy tree density, canopy and tall shrub basal area and cover, downed wood biomass, litter cover) and below-ground (soil nutrient availability, decomposition, forest floor thickness, pH, and phospholipid fatty acids (PLFAs) and multiple carbon-source substrate-induced respiration (MSIR) of the forest floor microbial community) environment. There was notable variation in fine-scale plant community composition; cluster and indicator species analyses of the 24 most commonly occurring understory species distinguished four assemblages, one for which a pioneer forb species had the highest cover levels, and three others that were characterized by different bryophyte species having the highest cover. Constrained ordination (distance-based redundancy analysis) showed that two above-ground (mean tree diameter, litter cover) and eight below-ground (forest floor pH, plant available boron, microbial community composition and function as indicated by MSIR and PLFAs) properties were associated with variation in understory plant community composition. These results provide novel insights into the important ecological associations between understory plant community composition and heterogeneity in ecosystem properties and processes within forests dominated by a single canopy species.     

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 grazing on the distribution and composition of low-shore algal communities on the central coast of Portugal and on the southern coast of Britain

A red algal turf is often found just below the barnacle/limpet zone of many European shores, especially on steep shores of moderate exposure. The hypothesis that grazing by limpets determines the upper limit of distribution of this red algal turf was tested on moderately exposed shores in Portugal and Britain. We also aimed to assess whether the grazing effect is modified at various spatial scales. Grazers were excluded by fences, with half-fenced and unfenced controls. Exclusion plots were rapidly colonised by green ephemeral algae in the months immediately after the beginning of the experiment (summer); these algae were later replaced by perennial algae. The percentage cover of turf-forming macroalgae showed a significant increase at both locations. The upper limit of distribution extended more than 50 cm on most of the shores studied. In contrast, control and half-fenced plots remained devoid of algae. After 2 years, ungrazed plots were mainly colonised by a red algal turf (e.g. Caulacanthus ustulatus, Gelidium spp., Laurencia pinnatifida) in Portugal, while canopy cover (Fucus serratus and Himanthalia elongata) dominated in Britain in marked contrast to the grazed plots. Physical factors acting at both local and geographical scales may explain these differences. However, although physical factors probably have an important influence on the identity, size and abundance of sublittoral fringe macroalgae, grazers play a major role in directly setting their upper limits. The effect of grazing by limpets was not consistent for all of the morphological algal groups and spatial scales considered in the present study. The effect of grazing on the cover of turf algae varied between Portugal and Britain (location scale), while effects on ephemeral and canopy algal cover varied at the shore scale within location.     

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.     

The effect of grazing and nutrient supply on periphyton associated bacteria

The effects of nutrient additions and grazing by macro-invertebrates on periphyton-associated algae and bacteria were studied by performing an enclosure experiment on three occasions from early spring to summer at mesotrophic Lake Erken and V?dd?, at the Swedish Baltic coast. There were significant interactions between nutrient additions and grazing on bacterial biomass and specific activity in Lake Erken. Thus, the importance of either bottom-up or top-down effects could not be singled out. Bacterial biomass increased with enrichment only in the absence of grazers. Grazer presence tended to increase bacterial biomass in ambient nutrient conditions, but to decrease bacterial biomass under enrichment. For specific activity the positive response to enrichment was restricted to grazer presence. Hence, grazing by macro-invertebrates may have an indirect positive effect on bacterial activity by enhancing nutrient conditions through their feeding activities and/or fecal pellets production. In addition, we found a significant relationship between bacterial production and chlorophyll a at both sites. This relationship weakened in the presence of macro-invertebrates. Thus, the importance of internal nutrient regeneration by bacteria and algae decreased, possibly due to increased nutrient availability, in the presence of macro-invertebrate grazers.     

Environmental context determines multi‐trophic effects of consumer species loss

Loss of biodiversity and nutrient enrichment are two of the main human impacts on ecosystems globally, yet we understand very little about the interactive effects of multiple stressors on natural communities and how this relates to biodiversity and ecosystem functioning. Advancing our understanding requires the following: (1) incorporation of processes occurring within and among trophic levels in natural ecosystems and (2) tests of context‐dependency of species loss effects. We examined the effects of loss of a key predator and two groups of its prey on algal assemblages at both ambient and enriched nutrient conditions in a marine benthic system and tested for interactions between the loss of functional diversity and nutrient enrichment on ecosystem functioning. We found that enrichment interacted with food web structure to alter the effects of species loss in natural communities. At ambient conditions, the loss of primary consumers led to an increase in biomass of algae, whereas predator loss caused a reduction in algal biomass (i.e. a trophic cascade). However, contrary to expectations, we found that nutrient enrichment negated the cascading effect of predators on algae. Moreover, algal assemblage structure varied in distinct ways in response to mussel loss, grazer loss, predator loss and with nutrient enrichment, with compensatory shifts in algal abundance driven by variation in responses of different algal species to different environmental conditions and the presence of different consumers. We identified and characterized several context‐dependent mechanisms driving direct and indirect effects of consumers. Our findings highlight the need to consider environmental context when examining potential species redundancies in particular with regard to changing environmental conditions. Furthermore, non‐trophic interactions based on empirical evidence must be incorporated into food web‐based ecological models to improve understanding of community responses to global change.     

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.     

The underappreciated role of life history in mediating the functional consequences of biodiversity change

Biodiversity is changing on both global and local scales, motivating research to understand the consequences of these changes for how communities and ecosystems function. Here, we explore the role of life history strategies in mediating biodiversity and ecosystem functioning. In particular, we evaluate how the composition, biomass (% cover), and richness of perennial (persistence ≥ 1 year) and ephemeral (persistence < 1 year) species change along a gradient of increasing seaweed species richness on a rocky shoreline. We show that the majority of biomass is comprised of perennial species, especially where overall richness is low, whereas the majority of species are ephemeral, especially where overall richness is high. We then present and discuss the results of an 18‐month field manipulation quantifying the factorial effects of tidal elevation, wave exposure, herbivore removals, thermal and desiccation stress amelioration, and nutrient additions on perennial versus ephemeral species. In particular, the diversity of ephemeral species was strongly affected, relative to perennial species, by tidal elevation, wave exposure, and herbivory; herbivores reduced diversity of ephemeral species relative to perennials. Relative to perennial cover, ephemeral cover was greater higher on the shore, in more wave‐exposed habitats, and where herbivores were removed, plots were unscreened, and/or nutrients were added. Thus, perennials and ephemerals responded differently to environmental context and experimental manipulation. We compared nitrate uptake and photosynthesis rates of ephemeral and perennial species and found that maximum nitrate uptake and photosynthesis rates of ephemerals were twice as high as those of perennials. These results highlight the disproportionate roles that ephemeral species play in mediating ecosystem‐level processes. In combination with our comparisons of the diversity and cover of perennial and ephemeral species along a biodiversity gradient, these results demonstrate the utility of incorporating life history traits into our efforts to understand the functional consequences of biodiversity change.     

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.     

Algal propagule banks modify competition, consumer and resource control on Baltic rocky shores

We hypothesized that supply from macroalgal propagule banks may influence the relative abundance of annual and perennial algae and that this may alter the effects of grazers and nutrients on species composition. In a factorial field experiment in the Baltic Sea littoral system we tested the effects of manipulating propagule banks, the abundance of crustacean and gastropod grazers, and nutrient supply on recruitment and growth of macroalgae over a year. Moreover, we determined seasonal patterns of macroalgal propagule dispersal at the experimental site and quantified algal abundance and recruitment at 25 locations throughout the Baltic Sea. Experimental manipulations had minor effects on adults of the dominating perennial alga, Fucus vesiculosus. Instead, we found that species composition was determined by processes operating at early life stages. Propagule supply from a propagule bank strongly favored the fast-growing annual alga Enteromorpha spp. which then blocked settlement and recruitment of Fucus. Grazers reduced the abundance of annual algae and indirectly favored Fucus recruitment. There was an apparent trade-off between gains from the propagule bank and losses to herbivory in five of seven colonizing species. Nutrient enrichment overrode grazer control of annual algae and accelerated the decline of Fucus only when annual algae had already achieved high densities through the propagule bank. Corroborating the experimental findings, field surveys across the Baltic showed that Fucus recruit densities can be predicted from the cover of annual algae during the period of Fucus reproduction and settlement. Recruitment inhibition by annual algae, which is driven by the abundance of annuals in the propagule bank, increasing nutrient levels, and declining consumer control, is suggested as a mechanistic explanation of the current decline of perennial algae in the Baltic Sea.     

A few is enough: a low cover of a non-native seaweed reduces the resilience of Mediterranean macroalgal stands to disturbances of varying extent

There is increasing evidence that the severity of the ecological impact of non-native species does not necessarily scale linearly with their abundance in the introduced range. Nonetheless, the potential of low abundance invaders to alter the resilience of native communities to disturbance has been poorly explored. On Mediterranean rocky reefs, we tested the hypothesis that (1) a pulse disturbance opening gaps within canopy stands formed by the fucoid seaweed Cystoseira brachycarpa facilitates the establishment of the non-native seaweed Caulerpa cylindracea and that (2), once established, the seaweed can reduce the recovery of macroalgal canopies. In July 2011, C. cylindracea was experimentally transplanted in small and large plots that were either cleared of the canopy or left untouched. After 45 months, the cover of C. cylindracea was greatest in small canopy-removal plots, without, however, achieving values exceeding ~10%. Nonetheless, such a low abundance of C. cylindracea caused a threefold reduction in canopy recovery. The establishment of C. cylindracea in canopy-removal plots did not alter the structure of the understory assemblages or the cover of turf-forming, erect and encrusting algae and sessile invertebrates. Our results suggest that some non-native species may be stronger competitors than natives, despite their low abundance. This property has important implications for control programs since not achieving the total eradication of the targeted invader would make little progress towards the mitigation of its impacts. Finally, our results show that non-native species acting as passengers of change can ultimately promote the persistence of alternative degraded states.     

Nutrient Enrichment and Food Web Composition Affect Ecosystem Metabolism in an Experimental Seagrass Habitat

Background

Food web composition and resource levels can influence ecosystem properties such as productivity and elemental cycles. In particular, herbivores occupy a central place in food webs as the species richness and composition of this trophic level may simultaneously influence the transmission of resource and predator effects to higher and lower trophic levels, respectively. Yet, these interactions are poorly understood.

Methodology/Principal Findings

Using an experimental seagrass mesocosm system, we factorially manipulated water column nutrient concentrations, food chain length, and diversity of crustacean grazers to address two questions: (1) Does food web composition modulate the effects of nutrient enrichment on plant and grazer biomasses and stoichiometry? (2) Do ecosystem fluxes of dissolved oxygen and nutrients more closely reflect above-ground biomass and community structure or sediment processes? Nutrient enrichment and grazer presence generally had strong effects on biomass accumulation, stoichiometry, and ecosystem fluxes, whereas predator effects were weaker or absent. Nutrient enrichment had little effect on producer biomass or net ecosystem production but strongly increased seagrass nutrient content, ecosystem flux rates, and grazer secondary production, suggesting that enhanced production was efficiently transferred from producers to herbivores. Gross ecosystem production (oxygen evolution) correlated positively with above-ground plant biomass, whereas inorganic nutrient fluxes were unrelated to plant or grazer biomasses, suggesting dominance by sediment microbial processes. Finally, grazer richness significantly stabilized ecosystem processes, as predators decreased ecosystem production and respiration only in the zero- and one- species grazer treatments.

Conclusions/Significance

Overall, our results indicate that consumer presence and species composition strongly influence ecosystem responses to nutrient enrichment, and that increasing herbivore diversity can stabilize ecosystem flux rates in the face of perturbations.     

Effects of herbivore exclusion and nutrient enrichment on coral reef macroalgae and cyanobacteria

Although phase shifts on coral reefs from coral-dominated to algal-dominated communities have been attributed to the effects of increased nutrient availability due to eutrophication and reduced herbivore abundance due to overfishing and disease, these factors have rarely been manipulated simultaneously. In addition, few studies have considered the effects of these factors on benthic, filamentous cyanobacteria (blue-green algae) as well as macroalgae. We used a combination of herbivore-exclusion cages and nutrient enrichment to manipulate herbivore abundance and nutrient availability, and measured the impacts of these treatments on macroalgal and cyanobacterial community structure. In the absence of cages, surface cover of the cyanobacterium Tolypothrix sp. decreased, while surface cover of the cyanobacteria Oscillatoria spp. increased. Cyanobacterial cover decreased in partial cages, and Tolypothrix sp. cover decreased further in full cages. Lower cyanobacterial cover and biomass were correlated with higher macroalgal cover and biomass. Dictyota bartayresiana dominated the partial cages, while Padina tenuis and Tolypiocladia glomerulata recruited into the full cages. Palatability assays demonstrated that herbivore-exclusion shifted macroalgal species composition from relatively unpalatable to relatively palatable species. Nutrient enrichment interacted with herbivore exclusion to increase the change in cover of D. bartayresiana in the uncaged and fully caged plots, but did not affect the final biomass of D. bartayresiana among treatments. Nutrient enrichment did not significantly affect the cover or biomass of any other taxa. These results stress the critical role of herbivory in determining coral reef community structure and suggest that the relative palatabilities of dominant algae, as well as algal growth responses to nutrient enrichment, will determine the potential for phase shifts to algal-dominated communities.     

The effects of grazing by gastropods and physical factors on the upper limits of distribution of intertidal macroalgae

Summary The cover of foliose algae is sparse to non-existent above a low-level algal zone on many shores in N.S.W., except in rock-pools. Above this algal zone, encrusting algae, mostly Hildenbrandia prototypus, occupy most of the primary substratum on sheltered shores. Experimental manipulations at midtidal levels were used to test hypotheses about the effects of grazing by molluses and of physical factors during low tide on this pattern of algal community structure.Fences and cages were used to exclude grazers: molluscs grazed under roofs and in open areas. Cages and roofs provided shade, and decreased the harshness of the environment during low tide: fences and open areas had the normal environmental regime.In the absence of grazers, rapid colonization of Ulva and slower colonization by other foliose algae occurred in all experimental areas. The rate of colonization by Ulva sporelings was initially retarded on existing encrusting algae, but after a few months, cover of Ulva equalled that on cleared rock.Most species of algae only grew to maturity inside cages, and remained as a turf of sporelings inside fences. No foliose algae grew to a visible size in open, grazed areas. Grazing thus prevents the establishment of foliose algae above their normal upper limit on the shore, but the effects of physical factors during low tide prevent the growth of algae which become established when grazers are removed. Physical factors thus limit the abundance of foliose algae at mid-tidal levels.The recolonization of cleared areas by Hildenbrandia was not affected by the presence of a turf of sporelings, nor by the shade cast by roofs, but was retarded in cages where mature algae formed a canopy. Even under such a canopy, Hildenbrandia eventually covered as much primary substratum as in open, grazed areas. This encrusting alga is able to escape from the effects of grazing by having a tough thallus, and by its vegetative growth which allows individual plants to cover a lot of substratum, and by the tendency for new individuals to start growing from small cracks and pits in the rock, which are apparently inaccessible to the grazers.Mature foliose algae are removed from the substratum by waves, and many individual plants died during periods of hot weather. Sporelings in a turf were eliminated, after experimental fences were removed, by the combined effects of macroalgal grazers, which invaded the areas, and microalgal grarers which ate the turt from the edges inwards.The results obtained here are discussed with respect to other studies on limits to distribution of intertidal macroalgae, and the role of grazing in the diversity and structure of intertidal algal communities. Some problems of these experimental treatments are also discussed.