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.     

Herbivore abundance and grazing intensity on a Caribbean coral reef

Herbivory is a primary factor in determining the structure of coral reef communities. Spatial variation among reef habitats in the intensity of herbivory has been documented, but underlying variation in species composition and abundance within the herbivore guild has received little attention. The distribution and relative abundances of herbivorous fishes and sea urchins across several habitats were studied on the Belizean barrier reef off the Caribbean coast of Central America. Marked variation in total herbivore density as well as major changes in the composition of the herbivore guild were found across reef habitats. Acanthurids (surgeonfishes) predominated in shallow areas (< 5 m) while scarids (parrotfishes) were dominant in deeper habitats. Significant differences among habitats in an experimental assay of grazing intensity were strongly correlated with herbivore abundance. The spatial distribution of herbivorous fishes across reef habitats does not appear to be simply explained by differences in reef topography, but may depend on complex interactions among proximity to nearby shelter, predator abundance, density of territorial competitors, and local availability of food resources.     

Decadal-scale changes in abundance of non-scleractinian invertebrates on a Caribbean coral reef

The benthic community structure of the shallow reefs of St. John, US Virgin Islands, was studied from 1992 to 2007 to test the hypotheses that the abundances of non-scleractinian invertebrates have changed, and further, that the changes are associated with variation in the percentage cover of scleractinian coral and macroalgae. The study utilized photoquadrats (0.25 m²) from fringing reefs (7-9 m depth) characterized by igneous boulders and low (< 5%) coral cover, and results from six sites were pooled to describe these reefs as a single habitat. Photoquadrats were analyzed biennially for the abundance of invertebrates, particularly those belonging to four well represented classes (Anthozoa, Demospongiae, Echinoidea, and Polychaeta), as well as the cover of scleractinians and macroalgae. Overall, the combined (multivariate) abundance of 30 invertebrate taxa changed over time, the combined (multivariate) abundance of the four invertebrate classes changed over time, and the individual (univariate) abundance of anthozoans, sponges, and echinoids changed over time. Throughout the study, coral cover remained < 5%, and while it varied significantly, it did not display a consistent trajectory of change; in contrast, the cover of macroalgae increased throughout the study. While it is unsurprising that the abundances of invertebrates changed over 15 y, notably they varied even though coral cover remained stable, in only a few cases were they related positively to macroalgal cover, and in most cases it was members of the suspension feeding guild that became more abundant. These outcomes suggest that: (1) benthic invertebrates on the shallow reefs of St. John may be more strongly influenced by regional (e.g., larva supply) than local (e.g., coral cover) conditions, (2) Caribbean reefs have changed more since the early 1990s than can be inferred from variation in cover of coral and macroalgae, and (3) suspension feeding invertebrates have become more common on shallow fringing reefs in at least one location.     

Multitrophic interactions mediate the effects of climate change on herbivore abundance

Climate change can influence the abundance of insect herbivores through direct and indirect mechanisms. In this study, we evaluated multitrophic drivers of herbivore abundance for an aphid species (Aphis helianthi) in a subalpine food web consisting of a host plant (Ligusticum porteri), mutualist ants and predatory lygus bugs (Lygus spp.). We used a model-selection approach to determine which climate and host plant cues best predict year-to-year variation in insect phenology and abundance observed over 6 years. We complemented this observational study with experiments that determined how elevated temperature interacts with (1) host plant phenology and (2) the ant-aphid mutualism to determine aphid abundance. We found date of snowmelt to be the best predictor of yearly abundance of aphid and lygus bug abundance but the direction of this effect differed. Aphids achieved lower abundances in early snowmelt years likely due to increased abundance of lygus bug predators in these years. Elevating temperature of L. porteri flowering stalks reduced their quality as hosts for aphid populations. However, warming aphid colonies on host plants of similar quality increased population growth rates. Importantly, this effect was apparent even in the absence of ants. While we observed fewer ants tending colonies at elevated temperatures, these colonies also had reduced numbers of lygus bug predators. This suggests that mutualism with ants becomes less significant as temperature increases, which contrasts other ant-hemipteran systems. Our observational and experimental results show the importance of multitrophic species interactions for predicting the effect of climate change on the abundances of herbivores.     

Indirect effects of algae on coral: algae-mediated, microbe-induced coral mortality

Declines in coral cover are generally associated with increases in the abundance of fleshy algae. In many cases, it remains unclear whether algae are responsible, directly or indirectly, for coral death or whether they simply settle on dead coral surfaces. Here, we show that algae can indirectly cause coral mortality by enhancing microbial activity via the release of dissolved compounds. When coral and algae were placed in chambers together but separated by a 0.02  μ m filter, corals suffered 100% mortality. With the addition of the broad-spectrum antibiotic ampicillin, mortality was completely prevented. Physiological measurements showed complementary patterns of increasing coral stress with proximity to algae. Our results suggest that as human impacts increase and algae become more abundant on reefs a positive feedback loop may be created whereby compounds released by algae enhance microbial activity on live coral surfaces causing mortality of corals and further algal growth.     

Effect of nutrient enrichment on growth and photosynthesis of the zooxanthellate coral Stylophora pistillata

The effect of prolonged (9 week) nutrient enrichment on the growth and photosynthetic rates of the zooxanthellate coral Stylophora pistillata was investigated. The main questions were: (1) what is the exposure time needed to induce measurable change in growth rate? (2) which are the concentrations of nitrogen and phosphorus required to cause changes in these rates? (3) what is the recovery potential of the corals after the nutrient stress? For this purpose, three tanks (N, P, NP) were enriched with ammonium (N), phosphorus (P) or both nutrients (NP), respectively. A fourth tank (C) served as a control. The growth of 40 nubbins (10 in each tank) was monitored during four periods: period 1 (nutrient-poor conditions), period 2 (10?μm NH₄ and/or 2?μm PO₄ enrichment), period 3 (20?μm NH₄ and/or 2?μm PO₄) and period 4 (nutrient-poor conditions). Period 4 was performed to study the recovery potential of corals after a nutrient stress. During period 1, growth rates remained constant in all tanks. In the P tank, growth rates declined during the two enrichment periods, with a total decrease of 60% by the end of period 3. In the N tank, growth rates remained nearly constant during period 2 but decreased in period 3 (60% decrease). In the NP tank, 50% and 25% decreases were observed during periods 2 and 3. At the end of the recovery period, a regain in growth rate was observed in the N and NP tanks (35 and 30% increase, respectively, compared with the rates measured at the end of period 3) and growth rates returned to 60% of the initial rates. By contrast, in the P tank, there was no regain in growth and a further decrease of 5% was observed. Rates of photosynthesis were often higher during the enriched than the nutrient-poor period (up to 150% increase). Corals with the highest percent increases in maximal gross photosynthetic rate (P ^g _max ) had the smallest decreases in growth rate due to nutrient enrichment. In conclusion, high ammonium (20?μm) and relatively low phosphorus concentrations (2?μm) are required to induce a significant decrease in coral growth rate. The largest reduction was observed with both ammonium and phosphorus enrichment. The decrease in growth rate was rapid following nutrient enrichment, since a 10% decrease or more could be observed after the first week of treatment.     

The influence of salt and nitrogen on herbivore abundance: direct and indirect effects

We report on the influence of experimentally increased interstitial salinity and plant nitrogen on the abundance of the delphacid planthopper, Prokelisia marginata (Van Duzee) (Homoptera: Delphacidae), which feeds on salt marsh cordgrass, Spartina alterniflora. We also report the effects of these treatments on parasitism of P. marginata eggs by the fairyfly parasitoid, Anagrus sophiae (Hymenoptera: Mymaridae). Soil salinity was significantly elevated following the addition of salt pellets broadcast over the ground and plant foliar nitrogen was significantly increased after the addition of fertilizer. The addition of fertilizer increased P. marginata densities on Spartina but addition of salt did not. Neither treatment significantly affected levels of egg parasitism by A. sophiae. In this system direct effects of plants on their herbivores via changes in plant chemistry appear more important than indirect effects of plants on herbivores via their natural enemies. Received: 1 August 1997 / Accepted: 29 September 1997     

Contrasting responses of phytoplankton and benthic algae to recent nutrient enrichment in Arctic tundra ponds

相似文献   

Chronic nutrient enrichment increases prevalence and severity of coral disease and bleaching

Nutrient loading is one of the strongest drivers of marine habitat degradation. Yet, the link between nutrients and disease epizootics in marine organisms is often tenuous and supported only by correlative data. Here, we present experimental evidence that chronic nutrient exposure leads to increases in both disease prevalence and severity and coral bleaching in scleractinian corals, the major habitat‐forming organisms in tropical reefs. Over 3 years, from June 2009 to June 2012, we continuously exposed areas of a coral reef to elevated levels of nitrogen and phosphorus. At the termination of the enrichment, we surveyed over 1200 scleractinian corals for signs of disease or bleaching. Siderastrea siderea corals within enrichment plots had a twofold increase in both the prevalence and severity of disease compared with corals in unenriched control plots. In addition, elevated nutrient loading increased coral bleaching; Agaricia spp. of corals exposed to nutrients suffered a 3.5‐fold increase in bleaching frequency relative to control corals, providing empirical support for a hypothesized link between nutrient loading and bleaching‐induced coral declines. However, 1 year later, after nutrient enrichment had been terminated for 10 months, there were no differences in coral disease or coral bleaching prevalence between the previously enriched and control treatments. Given that our experimental enrichments were well within the ranges of ambient nutrient concentrations found on many degraded reefs worldwide, these data provide strong empirical support to the idea that coastal nutrient loading is one of the major factors contributing to the increasing levels of both coral disease and coral bleaching. Yet, these data also suggest that simple improvements to water quality may be an effective way to mitigate some coral disease epizootics and the corresponding loss of coral cover in the future.     

The impact of CO2 emission scenarios and nutrient enrichment on a common coral reef macroalga is modified by temporal effects

Future coral reefs are expected to be subject to higher pCO₂ and temperature due to anthropogenic greenhouse gas emissions. Such global stressors are often paired with local stressors thereby potentially modifying the response of organisms. Benthic macroalgae are strong competitors to corals and are assumed to do well under future conditions. The present study aimed to assess the impact of past and future CO₂ emission scenarios as well as nutrient enrichment on the growth, productivity, pigment, and tissue nutrient content of the common tropical brown alga Chnoospora implexa. Two experiments were conducted to assess the differential impacts of the manipulated conditions in winter and spring. Chnoospora implexa's growth rate averaged over winter and spring declined with increasing pCO₂ and temperature. Furthermore, nutrient enrichment did not affect growth. Highest growth was observed under spring pre‐industrial (PI) conditions, while slightly reduced growth was observed under winter A1FI (“business‐as‐usual”) scenarios. Productivity was not a good proxy for growth, as net O₂ flux increased under A1FI conditions. Nutrient enrichment, whilst not affecting growth, led to luxury nutrient uptake that was greater in winter than in spring. The findings suggest that in contrast with previous work, C. implexa is not likely to show enhanced growth under future conditions in isolation or in conjunction with nutrient enrichment. Instead, the results suggest that greatest growth rates for this species appear to be a feature of the PI past, with A1FI winter conditions leading to potential decreases in the abundance of this species from present day levels.     

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.     

Changes in the abundance of blue-green algae related to nutrient loadings in the nearshore of Lake Michigan

Nutrient loadings to the nearshore of southeastern Lake Michigan have undergone a remarkable reduction. This reduction can affect the nutrient supply and result in biological changes. Changes in phytoplankton community, particularly the blue-green algae, can be related to nutrient changes. After thermal stratification, sudden increases in the blue-green algae population were significantly correlated to soluble reactive phosphorus concentrations. Phosphorus-stimulated low dissolved silica and phosphorus limitations after stratification appear to be primary factors contributing to the success of these algae.     

Effects of whole-stream nutrient enrichment on the concentration and abundance of aquatic hyphomycete conidia in transport

The concentrations and relative abundances of aquatic hyphomycete conidia in water were followed during a three-year study in two headwater streams at Coweeta Hydrologic Laboratory, North Carolina, using the membrane-filtration technique. After a one-year pretreatment period, one of the streams was enriched continuously with inorganic nutrients (N+P) for two years while the other stream served as the reference. This ecosystem-level nutrient manipulation resulted in concentrations of aquatic hyphomycete conidia in the water of the treated stream that were 4.5-6.9 times higher than the concentrations observed during the pretreatment period and in the reference stream. Nutrient enrichment led to an increase in the number of fungal species detected on each sampling date. Changes in dominance patterns and relative abundances of individual species also were detected after treatment. Nutrient addition stimulates the reproductive activity of aquatic hyphomycetes, their colonization success and fungal-mediated leaf-litter decomposition. Such changes in the activity of the fungal community might affect higher trophic levels in lotic ecosystems.     

Long-term effects of imidacloprid on the abundance of surface- and soil-active nontarget fauna in turf

• 1 Outbreaks of root‐feeding scarab larvae in turfgrass are widely managed through preventive applications of imidacloprid. Long‐residual activity and application before feasible scouting probably lead to its overuse and overexposure.
• 2 Recent investigations revealed a selective impact of imidacloprid (not trichlorfon or halofenozide) on certain nontarget turf arthropods, motivating the present study on the persistence of abundance effects over 6 years of annual applications.
• 3 Arthropods were sampled monthly (July to October) in replicated plots using soil core heat extraction and pitfall traps to quantify soil‐ and surface‐active arthropods. Captures were identified to class, order or family. The most represented taxa were analysed to test for cumulative effects and their change over season and year.
• 4 Imidacloprid had no impact on pitfall captures, although the abundance of Hemiptera, Thysanoptera, Coleoptera and Collembola was suppressed in soil core captures. Among beetles, impact was expressed in adults (not larvae), and in Carabidae and Staphylinidae (not Chrysomelidae or Curculionidae). Among springtails, impact was expressed in Entomobryomorpha (not Poduromorpha or Symphypleona). Impact did not diminish with year but there was variable recovery between applications.
• 5 There may therefore be a diverging response of soil‐ and surface‐active fauna to the nontarget impacts of imidacloprid. The suppression of predaceous (not phytophagous) beetles indicates an indirect effect mediated through declines in prey populations.
• 6 The magnitude of abundance effects confirms that the balance between target and nontarget impact should be explicitly examined. Implications are discussed with respect to functional relevance for nutrient cycling and the natural regulation of pests.

     

Response of cyanobacteria and phytoplankton abundance to warming,extreme rainfall events and nutrient enrichment

Cyanobacterial blooms are an increasing threat to water quality and global water security caused by the nutrient enrichment of freshwaters. There is also a broad consensus that blooms are increasing with global warming, but the impacts of other concomitant environmental changes, such as an increase in extreme rainfall events, may affect this response. One of the potential effects of high rainfall events on phytoplankton communities is greater loss of biomass through hydraulic flushing. Here we used a shallow lake mesocosm experiment to test the combined effects of: warming (ambient vs. +4°C increase), high rainfall (flushing) events (no events vs. seasonal events) and nutrient loading (eutrophic vs. hypertrophic) on total phytoplankton chlorophyll‐a and cyanobacterial abundance and composition. Our hypotheses were that: (a) total phytoplankton and cyanobacterial abundance would be higher in heated mesocosms; (b) the stimulatory effects of warming on cyanobacterial abundance would be enhanced in higher nutrient mesocosms, resulting in a synergistic interaction; (c) the recovery of biomass from flushing induced losses would be quicker in heated and nutrient‐enriched treatments, and during the growing season. The results supported the first and, in part, the third hypotheses: total phytoplankton and cyanobacterial abundance increased in heated mesocosms with an increase in common bloom‐forming taxa—Microcystis spp. and Dolichospermum spp. Recovery from flushing was slowest in the winter, but unaffected by warming or higher nutrient loading. Contrary to the second hypothesis, an antagonistic interaction between warming and nutrient enrichment was detected for both cyanobacteria and chlorophyll‐a demonstrating that ecological surprises can occur, dependent on the environmental context. While this study highlights the clear need to mitigate against global warming, oversimplification of global change effects on cyanobacteria should be avoided; stressor gradients and seasonal effects should be considered as important factors shaping the response.     

Biogeographic consequences of nutrient enrichment for plant–herbivore interactions in coastal wetlands

A major challenge in ecology is to understand broadscale trends in the impact of environmental change. We provide the first integrative analysis of the effects of eutrophication on plants, herbivores, and their interactions in coastal wetlands across latitudes. We show that fertilisation strongly increases herbivory in salt marshes, but not in mangroves, and that this effect increases with increasing latitude in salt marshes. We further show that stronger nutrient effects on plant nitrogen concentration at higher latitudes is the mechanism likely underlying this pattern. This biogeographic variation in nutrient effects on plant–herbivore interactions has consequences for vegetation, with those at higher latitudes being more vulnerable to consumer pressure fuelled by eutrophication. Our work provides a novel, mechanistic understanding of how eutrophication affects plant–herbivore systems predictably across broad latitudinal gradients, and highlights the power of incorporating biogeography into understanding large‐scale variability in the impacts of environmental change.     

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.     

Macroalgal blooms on southeast Florida coral reefs: I. Nutrient stoichiometry of the invasive green alga Codium isthmocladum in the wider Caribbean indicates nutrient enrichment

Invasive blooms of the siphonaceous green algae Codium spp. have been considered a symptom of coastal eutrophication but, to date, only limited biochemical evidence supports a linkage to land-based nutrient pollution. Beginning in the summer of 1990, spectacular blooms of unattached Codium isthmocladum developed on deep coral reef habitats in southern Palm Beach County and northern Broward County, and in subsequent years, attached populations formed on reefs in northern Palm Beach County.To better understand the nutrition of these HABs, we collected C. isthmocladum and other reef macroalgae from various locations in southeast Florida as well as the wider Caribbean region for tissue C:N:P analysis in order to gauge variability in the type and degree of N- and/or P-limited growth. Widespread nutrient enrichment in floridian C. isthmocladum populations was evidenced by significantly higher tissue P (0.06% versus 0.04% of dry weight) and lower C:N (12 versus 19), C:P (425 versus 980), and N:P (35 versus 50) ratios compared to more nutrient-depleted Caribbean populations. To determine nutrient availability on southeast Florida's reefs, we sampled near-bottom waters at a variety of locations for DIN (NH₄⁺ + NO₃⁻ + NO₂⁻) and SRP analysis. In general, concentrations of NH₄⁺, NO₃⁻and SRP were all high on southeast Florida's reefs compared to values reported for Caribbean coral reefs. Although summertime upwelling provides episodic NO₃⁻ and SRP enrichment to reefs in southeast Florida, these transient nutrient pulses have not historically supported C. isthmocladum blooms.We suggest that the widespread P enrichment of C. isthmocladum tissue and water column DIN:SRP ratios <16:1 in southeast Florida drive this system toward N limitation where low level NH₄⁺ enrichment becomes of paramount importance. Hence, the recent C. isthmocladum blooms appear to be supported by increasing land-based nutrient pollution, particularly, sewage that is enriched in NH₄⁺ and SRP at a low N:P ratio (<10:1) critical to sustaining balanced growth during bloom formation.