Effect of grazing by isopods and amphipods on growth of Ulva spp. (Chlorophyta)

Eutrophication of shallow coastal waters often leads to blooms of macroalgae. Grazing by crustaceans, such as amphipods and isopods, can reduce macroalgal biomass accumulation. At the same time, growth of the macroalgae can be stimulated by epiphyte removal. The role of grazing by isopods and amphipods on Ulva spp. biomass development was investigated in the Veerse Meer, a brackish lagoon situated in the southwest Netherlands. Exclusion of grazing in the field did not stimulate Ulva spp. growth. In fact, growth rates were higher in exclosures that allowed grazers to enter. Edibility tests identified the amphipod Gammarus locusta, and the isopods Idotea chelipes and Sphaeroma hookeri as potential grazers on Ulva spp. However, when epiphytic diatoms were present on the Ulva spp. thalli, Gammarus and Sphaeroma grazed on ephiphytes and not on Ulva tissue. Only Idotea continued to graze on Ulva spp. A laboratory growth experiment revealed a positive effect of Gammarus presence on Ulva spp. growth, probably caused by preferential removal of epiphytic diatoms from the Ulva spp. thalli. The growth stimulation by epiphyte removing grazers such as Gammarus may explain the higher growth rates in the presence of grazers observed in the field. When determining the potential role of invertebrate grazers in controlling macroalgal biomass accumulation, it is important to include an assessment of the epiphyte abundance on the macroalgae, as preferential removal of epiphytes may stimulate growth and thus have the opposite effect.     

Nutrient and iron limitation to <Emphasis Type="Italic">Ulva</Emphasis> blooms in a eutrophic coastal lagoon (Sacca di Goro,Italy)

Growth patterns and bloom formation of the green seaweed Ulva rigida were analysed in the eutrophic Sacca di Goro lagoon (Po River Delta, Italy). Variations of standing biomasses and elemental composition of Ulva were analysed through an annual cycle with respect to nitrogen, phosphorus and iron. Growth rates, nutrient and iron uptake and nitrate storage by macroalgal thalli were also assessed with field experiments during the formation of a spring bloom. The control of Ulva growth and the bloom formation depended on multiple factors, especially on nitrogen availability and iron deficiency. In the nitrate rich waters of the Sacca di Goro lagoon, nitrate accumulation in Ulva thalli was inversely related with Fe uptake, indicating an influence of Fe limitation on N acquisition. Since length and magnitude of nitrate luxury uptake are inversely related to the size of the intracellular nitrate pools, in nitrate rich waters the fast growing Ulva may face risk of N-limitation not only when exposed to low N concentrations or at high biomass levels, but also when exposed to pulsed dissolved nitrate concentrations at low iron availability. The potential Fe limitation could be affected by processes controlled by geochemical reactions and by macroalgal growth and decomposition. Both Fe oxidation during the active macroalgal growth and the formation of insoluble FeS and FeS₂ during bloom collapse can result in a drastic decrease of soluble iron. Thus, a potential limitation of Fe to macroalgae can occur, determining positive feedbacks and potentially controlling the extent of bloom development and persistence.     

Growth of the seaweed Ulva rigida C. Agardh in relation to biomass densities, internal nutrient pools and external nutrient supply in the Sacca di Goro lagoon (Northern Italy)

Growth of the seaweed Ulva rigida C. Agardh was investigated in relation to biomass densities, internal nutrient pools and external nutrient supply. Research was carried out from 23 March to 5 July 1994 in the Sacca di Goro (Po Delta, Northern Italy), whose south-eastern part was covered by extensive mats of Ulva rigida. Two types of field experiments were conducted by incubating Ulva thalli inside large cages. In the first experiment, beginning on 23 March, 100 g of wet thalli were placed into the cages, allowed to grow for two weeks, then collected and replaced. This procedure was repeated 8 times over the study period. In the second experiment, Ulva thalli were left inside the cages and collected at selected time intervals (14, 27, 41, 64 and 76 days) in order to simulate the effects of increased density on growth and nutrient storage.We recorded specific growth rates (NGR) ranging from 0.025 to 0.081 d^–1 for a period up to two months in the repeated short-term experiments performed at relatively low initial algal densities (300–500 g AFDW m^–3). These NGR resulted significantly related to dissolved inorganic nitrogen (DIN) in the water column. Tissue concentrations of total Kjeldahl nitrogen (TN) were almost constant, while extractable nitrate decreased in a similar manner to DIN in the water column. Total phosphorus showed considerable variation, probably linked to pulsed freshwater inflow.In the long-term incubation experiment, NGR of Ulva was inversely related to density. Internal concentrations of both total P and TN reached maximum values after one month; thereafter P concentration remained almost constant, while TN decreased below 2% w/w (by dry weight). The TN decrease was also accompanied by an abrupt decrease in nitrate tissue concentration. The biomass incubated over the two month period suffered a progressive N limitation as shown by a decreasing NY ratio (49.4 to 14.6). The reciprocal control of Ulva against biogeochemical environment and viceversa is a key factor in explaining both resource competition and successional stages in primary producer communities dominated by Ulva. However, when the biomass exceeds a critical threshold level, approximately 1 kg AFDW m^–3, the macroalgal community switches from active production to rapid decomposition, probably as a result of selfshading, biomass density and development of anaerobic conditions within the macroalgal beds.     

Spatial and temporal variability in macroalgal blooms in a eutrophied coastal estuary

All three macroalgal clades (Chlorophyta, Rhodophyta, and Phaeophyceae) contain bloom-forming species. Macroalgal blooms occur worldwide and have negative consequences for coastal habitats and economies. Narragansett Bay (NB), Rhode Island, USA, is a medium sized estuary that is heavily influenced by anthropogenic activities and has been plagued by macroalgal blooms for over a century. Over the past decade, significant investment has upgraded wastewater treatment from secondary treatment to water-quality based limits (i.e. tertiary treatment) in an effort to control coastal eutrophication in this system. The goal of this study was to improve the understanding of multi-year macroalgal bloom dynamics through intensive aerial and ground surveys conducted monthly to bi-monthly during low tides in May–October 2006–2013 in NB. Aerial surveys provided a rapid characterization of macroalgal densities across a large area, while ground surveys provided high resolution measurements of macroalgal identity, percent cover, and biomass.Macroalgal blooms in NB are dominated by Ulva and Gracilaria spp. regardless of year or month, although all three clades of macroalgae were documented. Chlorophyta cover and nutrient concentrations were highest in the middle and upper bay. Rhodophyta cover was highest in the middle and lower bay, while drifting Phaeophyceae cover was patchy. Macroalgal blooms of >1000 g fresh mass (gfm)/m² (max = 3510 gfm/m²) in the intertidal zone and >3000 gfm/m³ (max = 8555 gfm/m³) in the subtidal zone were observed within a heavily impacted embayment (Greenwich Bay). Macroalgal percent cover (intertidal), biomass (subtidal), and diversity varied significantly between year, month-group, site, and even within sites, with the highest species diversity at sites outside of Greenwich Bay. Total intertidal macroalgal percent cover, as well as subtidal Ulva biomass, were positively correlated with temperature. Dissolved inorganic nitrogen concentrations were correlated with the total biomass of macroalgae and the subtidal biomass of Gracilaria spp. but not the biomass of Ulva spp. Despite seasonal reductions in the nutrient output of wastewater treatment facilities emptying into upper Narragansett Bay in recent years, macroalgal blooms still persist. Continued long-term monitoring of water quality, macroalgal blooms, and ecological indicators is essential to understand the changes in macroalgal bloom dynamics that occur after nutrient reductions from management efforts.     

INVERTEBRATE‐MEDIATED NUTRIENT LOADING INCREASES GROWTH OF AN INTERTIDAL MACROALGA1

Even in nitrogen‐replete ecosystems, microhabitats exist where local‐scale nutrient limitation occurs. For example, coastal waters of the northeastern Pacific Ocean are characterized by high nitrate concentrations associated with upwelling. However, macroalgae living in high‐zone tide pools on adjacent rocky shores are isolated from this upwelled nitrate for extended periods of time, leading to nutrient limitation. When high‐intertidal pools are isolated during low tide, invertebrate‐excreted ammonium accumulates, providing a potential nitrogen source for macroalgae. I quantified the influence of mussels (Mytilus californianus Conrad) on ammonium accumulation rates in tide pools. I then evaluated the effects of ammonium loading by mussels on nitrogen assimilation and growth rates of Odonthalia floccosa (Esp.) Falkenb., a common red algal inhabitant of pools on northeastern Pacific rocky shores. Odonthalia was grown in artificial tide pool mesocosms in the presence and absence of mussels. Mesocosms were subjected to a simulated tidal cycle mimicking emersion and immersion patterns of high‐intertidal pools on the central Oregon coast. In the presence of mussels, ammonium accumulated more quickly in the mesocosms, resulting in increased rates of nitrogen assimilation into algal tissues. These increased nitrogen assimilation rates were primarily associated with higher growth rates. In mesocosms containing mussels, Odonthalia individuals added 41% more biomass than in mesocosms without mussels. This direct positive effect of mussels on macroalgal biomass represents an often overlooked interaction between macroalgae and invertebrates. In nutrient‐limited microhabitats, such as high‐intertidal pools, invertebrate‐excreted ammonium is likely an important local‐scale contributor to macroalgal productivity.     

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.     

Influence of outbreak of macroalgal blooms on phosphorus release from the sediments in Swan Lake Wetland,China

Macroalgal blooms have occurred worldwide frequently in coastal areas in recent decades, which dramatically modify phosphorus (P) cycle in water column and the sediments. Rongcheng Swan Lake Wetland, a coastal wetland in China, is suffering from extensive macroalgal blooms. In order to verify the influence of macroalgal growth on sediment P release, the sediments and filamentous Chaetomorpha spp. were incubated in the laboratory to investigate the changes of water quality parameters, P levels in overlying water, and sediments during the growth period. In addition, algal biomass and tissue P concentration were determined. In general, Chaetomorpha biomasses were much higher in high P treatments than in low P treatments. Compared with algae+low P water treatment, the addition of sediments increased the algal growth rate and P accumulation amount. During the algal growth, water pH increased greatly, which showed significant correlation with algal biomass in treatments with high P (P < 0.05). P fractions in the sediments showed that Fe/Al–P and organic P concentrations declined during the algal growth, and great changes were observed in algae+low P water+sediment treatment for both. As a whole, the sediments can supply P for Chaetomorpha growth when water P level was low, and the probable mechanism was the release of Fe/Al–P at high pH condition induced by intensive Chaetomorpha blooms.     

Regulation of spatial and seasonal variation of macroalgal biomass in a brackish, eutrophic lake

Processes leading to biomass variation of Ulva were investigated at two contrasting sites in the eutrophic Veerse Meer (The Netherlands). Ulva species dominated at the Middelplaten site, while at the Kwistenburg site a mixture of Ulva spp. and Chaetomorpha linum dominated. Total summer macroalgal biomass was higher at Middelplaten than at Kwistenburg (282 and 79 g DW m^–2, respectively). Growth rates of Ulva spp. were high at both sites in May 1992 (cage mean 0.28–0.30 day^–1), but quickly dropped to lower values (0.05–0.10 day^–1). In May, growth rates were significantly highest at Kwistenburg, while during the rest of the season growth rates were similar for both sites. Temperature, pH, dissolved oxygen, salinity, light attenuation, phytoplankton and nutrient concentrations did not differ between sites. The relation between variation in Ulva spp. growth rates and environmental parameters was analysed using stepwise multiple regression, showing that light and temperature were the main variables regulating Ulva spp. growth rates. As Ulva growth rates were similar for both sites but biomass was much lower at Kwistenburg it was concluded that a large amount of produced biomass was lost at Kwistenburg. Although the exact reason for this remains unclear, it seems most likely that transport of macroalgae by wind and waves is a very important factor. This study shows the importance of simultaneously measuring growth rates and biomass at a high temporal resolution to reveal the mechanisms responsible for spatial variation in macroalgal biomass in shallow coastal areas. Electronic Publication     

Spatiotemporal variations of inorganic nutrients along the Jiangsu coast,China, and the occurrence of macroalgal blooms (green tides) in the southern Yellow Sea

Large macroalgal blooms (i.e. green tides of Ulva prolifera) occurred in the southern Yellow Sea, China, yearly from 2007 to 2016. They were among the largest of such outbreaks around the world, and these blooms likely originated along the coast of the Jiangsu Province, China. Understanding the roles of nutrients in the onset of these macroalgal blooms is needed to identify their origin. This study analyzes the spatiotemporal variations in dissolved inorganic nitrogen and phosphorus (DIN and PO₄–P) and the N/P ratio along the Jiangsu coast from 1996 to 2014 during late-March to April, the months which corresponds to the pre-bloom period of green tides since 2007. A zone of high DIN and PO₄–P concentrations has developed along the Jiangsu coast, between the cities of Sheyang and Nantong, since 1996. There was an 18-year trend of increasing DIN concentrations during the pre-bloom period as well as a positive correlation between the U. prolifera biomass and DIN concentrations. Nutrient inputs from rivers and mariculture in the Jiangsu Province may have provided nitrogen that contributed the magnitude of macroalgal blooms that subsequently spread into the southern Yellow Sea.     

珠江口及毗邻海域营养盐对浮游植物生长的影响

基于2006年7月(夏季),10月(秋季)和2007年3月(春季)的现场调查数据,对珠江口及毗邻海域中的营养盐和叶绿素a等环境生态因子的时空分布特性进行了对比分析,研究了氮磷比与叶绿素a含量和种群多样性之间的联系,探讨了该海域营养盐对于浮游植物生长的影响。结果表明:(1)研究海域营养盐表现出较强的季节和空间差异性,总氮(TN)和总磷(TP)浓度均值春季(1.545 mg/L、0.056 mg/L)和夏季(1.570 mg/L、0.058 mg/L)均大于秋季(1.442 mg/L、0.034 mg/L),且春夏季浓度空间差异更明显。(2)调查期间海域营养盐含量超标现象突出,夏季尤为明显。无机氮(DIN)总体均值0.99 mg/L,超四类海水标准限值1倍,活性磷酸盐(PO4-P)总体均值0.021 mg/L,DIN∶PO4-P平均值为130;叶绿素a浓度与营养盐、p H、温度有较显著的相关性。(3)叶绿素a浓度较高的站位,具有较高的DIN∶PO4-P值,但浮游植物多样性指数偏低,优势种明显,主要为中肋骨条藻。氮磷比的改变会影响不同生长特性的浮游植物间的竞争和种群结构的改变;今后海洋污染治理中,在控制氮、磷污染时要注意氮磷比的改变可能造成的浮游生态影响。     

Modelling the distribution and growth of ‘problem’ green seaweed in the Medway estuary,UK

The results of a modelling study to investigate the mechanisms controlling macroalgal growth within the Medway estuary, UK, are presented. Intertidal zone bathymetry, tidal dynamics and turbidity control the time available for nutrient uptake and photosynthesis, and were used as a basis for predicting areas where macroalgae will grow. Tidal bed stress was also considered as a controlling factor for the presence of the less robust green macroalgae species. Two approaches to predicting macroalgal distributions were applied: (1) a simple ‘suitability index’ method based on tidal flooding and drying, taking account of the conflict between time available for nutrient uptake and for photosynthesis; and (2) a biological macroalgal growth model that includes a detailed treatment of nutrient uptake and plant growth. The former approach assigns a value between zero and one for the suitability of a location for macroalgal occurrence, while the latter predicts the full macroalgal growth dynamics over an annual cycle. Tidal bed stress was included in both approaches as an independent modifier of macroalgal occurrence/growth. Results were compared with aerial survey maps of observed vegetation cover and time series of measured biomass density. Both approaches gave good predictions of non-species-specific vegetation cover in the intertidal zone of the Medway. Tidal bed stress was found to be a strong predictor of the specific occurrence of Enteromorpha spp. and Ulva spp., with these species favouring areas of low tidal energy. It was concluded that light and a lack of suitable regions with low tidal bed stress, rather than nutrients, were the main factors limiting excessive growth of Enteromorpha spp. and Ulva spp. in the estuary. Although this study was focussed on the Medway, the results are likely to be applicable to a broad range of relatively turbid, meso- and macro-tidal estuaries. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Guest editors: J. H. Andersen & D. J. Conley Eutrophication in Coastal Ecosystems: Selected papers from the Second International Symposium on Research and Management of Eutrophication in Coastal Ecosystems, 20–23 June 2006, Nyborg, Denmark     

The relative importance of sediment and water column supplies of nutrients to the growth and tissue nutrient content of the green macroalga Enteromorpha intestinalis along an estuarine resource gradient

Large blooms of opportunistic green macroalgae such as Enteromorpha intestinalis are of ecological concern in estuaries worldwide. Macroalgae derive their nutrients from the water column but estuarine sediments may also be an important nutrient source. We hypothesized that the importance of these nutrient sources to E. intestinalis varies along a nutrient-resource gradient within an estuary. We tested this in experimental units constructed with water and sediments collected from 3 sites in Upper Newport Bay estuary, California, US, that varied greatly in water column nutrient concentrations. For each site there were three treatments: sediments + water; sediments + water + Enteromorpha intestinalis (algae); inert sand + water + algae. Water in units was exchanged weekly simulating low turnover characteristic of poorly flushed estuaries. The importance of the water column versus sediments as a source of nutrients to E. intestinalis varied with the magnitude of the different sources. When initial water column levels of dissolved inorganic nitrogen (DIN) and soluble reactive phosphorus (SRP) were low, estuarine sediments increased E. intestinalis growth and tissue nutrient content. In experimental units from sites where initial water column DIN was high, there was no effect of estuarine sediments on E. intestinalis growth or tissue N content. Salinity, however, was low in these units and may have inhibited growth. E. intestinalis growth and tissue P content were highest in units from the site with highest initial sediment nutrient content. Water column DIN was depleted each week of the experiment. Thus, the water column was a primary source of nutrients to the algae when water column nutrient supply was high, and the sediments supplemented nutrient supply to the algae when water column nutrient sources were low. Depletion of water column DIN in sediment + water units indicated that the sediments may have acted as a nutrient sink in the absence of macroalgae. Our data provide direct experimental evidence that macroalgae utilize and ecologically benefit from nutrients stored in estuarine sediments.     

Effects of herbivores, nutrient enrichment, and their interactions on macroalgal proliferation and coral growth

We conducted a 20-week manipulative field experiment on shallow forereefs of the Florida Keys to assess the separate and interactive effects of herbivory and nutrient enrichment on the development of macroalgal communities and the fitness of the corals Porites porites and Siderastrea siderea. Excluding large herbivorous fishes produced macrophyte blooms both with and without nutrient enrichment. In contrast, there were no direct effects of nutrient enrichment. There were, however, small, but significant, interactive effects of herbivory and enrichment on macroalgal cover. Following nutrient enrichment, total macroalgae and the common seaweeds Dictyota spp. were suppressed in the presence, but not in the absence, of large herbivorous fishes—suggesting that fishes were selectively feeding on nutrient-enriched macrophytes. Access by large herbivores prevented algal overgrowth of corals, but these large fishes also directly grazed both corals. Excluding fishes did not alter survivorship of either coral species, but did decrease parrotfish grazing scars on both corals and increased the net growth of P. porites. Nutrient additions had no direct effects on the survivorship of corals, but there was a trend (P = 0.097) for nutrients to stimulate the growth of P. porites. The preponderance of experiments available to date indicates that loss of key herbivores is a major factor driving macroalgal blooms on coral reefs; anthropogenic nutrient pollution generally plays a more minor role.     

Tracing the origin of green macroalgal blooms based on the large scale spatio-temporal distribution of Ulva microscopic propagules and settled mature Ulva vegetative thalli in coastal regions of the Yellow Sea,China

From 2008 to 2016, massive floating green macroalgal blooms occurred annually during the summer months in the Yellow Sea. The original source of these blooms was traced based on the spatio-temporal distribution and species composition of Ulva microscopic propagules and settled Ulva vegetative thalli monthly from December 2012 to May 2013 in the Yellow Sea. High quantities of Ulva microscopic propagules in both the water column and sediments were found in the Pyropia aquaculture area along the Jiangsu coast before a green macroalgal bloom appeared in the Yellow Sea. The abundance of Ulva microscopic propagules was significantly lower in outer areas compared to in Pyropia aquaculture areas. A molecular phylogenetic analysis suggested that Ulva prolifera microscopic propagules were the dominant microscopic propagules present during the study period. The extremely low biomass of settled Ulva vegetative thalli along the coast indicated that somatic cells of settled Ulva vegetative thalli did not provide a propagule bank for the green macroalgal blooms in the Yellow Sea. The results of this study provide further supporting evidence that the floating green macroalgal blooms originate from green macroalgae attached to Pyropia aquaculture rafts along the Jiangsu coastline of the southern Yellow Sea.     

Comparison of population growth and photosynthetic apparatus changes in response to different nutrient status in a diatom and a coccolithophore

In many marine ecosystems, diatoms dominate in nutrient‐rich coastal waters while coccolithiophores are found offshore in areas where nutrients may be limiting. In lab‐controlled batch cultures, mixed‐species competition between the diatom Phaeodactylum tricornutum and the coccolithophore Emiliana huxleyi and the response of each species were examined under nitrate (N) and phosphate (P) starvation. Based on the logistic growth model and the Lotka–Volterra competition model, E. huxleyi showed higher competitive abilities than P. tricornutum under N and P starvation. For both species, cell growth was more inhibited by P starvation, while photosynthetic functions (chl a fluorescence parameters) and cellular constituents (pigments) were impaired by N starvation. The decline of photosynthetic functions occurred later in E. huxleyi (day 12) than in P. tricornutum (day 9); this time difference was associated with greater damage of the photosynthetic apparatus in P. tricornutum compared with E. huxleyi. Xanthophyll cycle pigment accumulation and the transformation from diadinoxanthin to diatoxanthin was more active in E. huxleyi than P. tricornutum, under similar N and P starvation. We concluded that E. huxleyi and P. tricornutum have different mechanisms to allocate resources and energy under nutrient starvation. It appears that E. huxleyi has a more economic strategy to adapt to nutrient depleted environments than P. tricornutum. These findings provided additional evidence explaining how N versus P limitation differentially support diatom and coccolithophore blooms in natural environments.     

Trophic cascades in a temperate seagrass community

We assessed the relative importance of bottom–up and top–down processes in structuring an eelgrass community in Sweden, a system impacted both by eutrophication and overfishing. Using artificial seagrass as substrate, we manipulated nutrient levels and predator abundance in a full‐factorial cage‐experiment. The results revealed a seagrass community dominated by strong top–down processes controlling the aggregate biomass of mesograzers and macroalgae. In the absence of predators the large amphipod Gammarus locusta became very abundant resulting in a leaf community with low biomass of algae and smaller mobile fauna. One enclosed gobid fish predator reduced the abundance of adult G. locusta by >90%, causing a three to six times increase in the biomass of algae, smaller mesograzers and meiofauna. Numerous small predators in uncaged habitats reduced the biomass of G. locusta and other mesograzers by >95% in comparison to the fish treatment, further increasing the biomass of epiphytic algae and meiofauna. Although water column nutrient enrichment caused a temporal bloom of the filamentous macroalgae Ulva spp., no significant nutrient‐effects were found on the algal community at the end of the experiment. The only lasting nutrient‐effect was a significant increase in the biomass of G. locusta, but only in the absence of ambient predators. These results demonstrate that mesograzers can respond to enhanced food supply, increase their biomass and control the algal growth when predation rates are low. However, in the assessed system, high predation rates appear to make mesograzers functionally extinct, causing a community‐wide trophic cascade that promotes the growth of ephemeral algae. This top–down effect could penetrate down, despite a complex food‐web because the interaction strength in the community was strongly skewed towards two functionally dominant algal and grazer species that were vulnerable to consumption. These results indicate that overexploitation of gadoid fish may be linked to increased macroalgal blooms and loss of eelgrass in the area through a trophic cascade affecting the abundance of mesograzers.     

Modeling the influence of nutrients, turbulence and grazing on Pfiesteria population dynamics

A semi-idealized marine ecosystem model, designed as a heuristic tool for exploring the population dynamics of non-inducible versus toxic forms of Pfiesteria is described. The model is based on empirical evidence suggesting that these differing functional types of Pfiesteria also differ substantially in terms of what they eat and how they utilize it to optimize their growth. Non-inducible strains are similar to other mixotrophic dinoflagellates, whereas toxic strains may consume organic matter and detritus, produce toxins and attack fish. In our model formulation we represent these differences in a simplified way: the non-inducible strain is kleptochloroplastidic and it can take up DIN, but it cannot utilize DON, whereas the toxic strain is heterotrophic, it cannot utilize DIN, but it can utilize DON directly. These differences give rise to very different impacts on prey and nutrient concentrations in our model. Under high DIN/DON ratio conditions, the non-inducible cells grew much faster and were therefore more likely to bloom, but this advantage is substantially mitigated when the DIN/DON ratio is low. A turbulence parameterization was also incorporated into our model. The effect of this was to reduce the grazing rate of Pfiesteria when turbulence levels are high. According to our model, increased turbulence is more detrimental to the toxic functional type because it grows more slowly. The further imposition of microzooplankton grazing in the model showed that top-down control effects can be very significant, which is consistent with both laboratory and field studies and the general idea that plankton blooms can only happen in the absence of substantial grazing control. In general, our model results suggest that non-toxic blooms are more likely to occur in more turbulent inorganic-nutrient rich conditions, which are often found in more open coastal and estuarine waters that are subject to high inorganic loading. In contrast, toxic blooms are more likely to occur in calm, organic-nutrient rich conditions, which are often found in shallow, protected tributaries that are subject to high organic nutrient loading. Our model results also support the idea that the absence of strong grazing pressure is a prerequisite to bloom formation for both non-inducible and toxic strains of Pfiesteria. These results are generally consistent with observed patterns of toxic Pfiesteria blooms in Chesapeake Bay, the Neuse River of North Carolina and many other coastal and estuarine environments.     

Patterns of Distribution and Environmental Correlates of Macroalgal Assemblages and Sediment Chlorophyll A in Oregon Tidal Wetlands

Algae have important functional roles in estuarine wetlands. We quantified differences in macroalgal abundance, composition and diversity, and sediment chl a and pheophytin a (pheo a) among three National Wetlands Inventory (NWI) emergent marsh classes in four Oregon estuaries spanning a range of riverine to marine dominance. We also assessed the strength of macroalgal‐vascular plant associations and the degree to which environmental variables correlated with algal community metrics in marsh and woody wetlands. The frequency of occurrence of most macroalgal genera, total benthic macroalgal cover, macroalgal diversity, and sediment chl a content were several times higher in low emergent marsh than in high marsh or palustrine tidal marsh. Conversely, pheo a: chl a ratios were highest in high and palustrine marsh. Attached macroalgae (Fucus and Vaucheria) were strongly associated with plants common at lower tidal elevations such as Sarcocornia perennis and Jaumea carnosa; Ulva (an unattached alga) was not strongly associated with any common low marsh plants. In structural equation models, intertidal elevation was the most influential predictor of macroalgal cover and richness and chl a; light availability and soil salinity played secondary roles. Although common taxa such as Ulva spp. occurred across a broad range of salinities, wetlands with oligohaline soils (salinity < 5) had the lowest macroalgal diversity and lower sediment chl a. These types of baseline data on algal distributions are critical for evaluating the structural and functional impacts of future changes to coastal estuaries including sea‐level rise (SLR), altered salinity dynamics, and habitat modification.     

THE EFFECT OF NITROGEN SOURCE ON THE GROWTH AND TOXICITY OF THREE POTENTIALLY HARMFUL DINOFLAGELLATES

Increases in population and agriculture in coastal areas can result in increased nutrient inputs and alterations in the ratios of organic to inorganic nutrients in coastal waters. Such changes in coastal nutrient regimes can affect phytoplankton community structure by creating conditions favorable for growth and dominance of algae that were not dominant before. The effect that changes in ratios and concentrations of nutrients have on toxicity of harmful algal species is not well known. There seems to be a relationship; however, between nutrient stress and toxin production among harmful phytoplankton producing low‐N toxins, e.g. Diarrhetic Shellfish Poisoning (DSP) toxins. Even less is known about the relationship between organic nutrient uptake and toxin production. Benthic species and species in coastal areas are probably exposed to greater fluxes of dissolved organic nitrogen (DON). In this study, benthic and planktonic species of Prorocentrum were grown on L1 media with the sole N‐source varying among treatments as nitrate, ammonium, urea, L‐glutamic acid, and high molecular weight natural DON. An ELISA specific to the DSP toxins, okadaic acid and 35‐methylokadaic acid, was used to determine toxin production by each species when grown on the different N sources. Preliminary results indicate that some organic forms of N support growth as well as inorganic forms for Prorocentrum minimum, P. mexicanum, and P. hoffmannianum.     

Management of natural <Emphasis Type="Italic">Ulva</Emphasis> spp. blooms in San Quintin Bay,Baja California: Is it justified?

According to Zertuche-González et al. (2009), Ulva spp. blooms, favored by oyster cultivation, are likely displacing subtidal meadows of Zostera marina in San Quintin Bay, Baja California. The authors propose a partial removal of the seaweed to reduce the risk of eutrophication and eelgrass displacement in the bay. We warn about the removal of Ulva spp. biomass by raising six arguments that emphasize the necessity of a historical and ecosystem-based management for San Quintin Bay. First, processes other than competitive exclusion by Ulva spp. blooms more likely explain changes of Z. marina subtidal meadows in the past decade. Second, there is no consistent evidence that oyster cultivation is promoting blooms of Ulva spp. and the loss of eelgrass. Third, the removal of Ulva spp. biomass is based on experiences of heavily anthropogenically eutrophic systems, while San Quintin Bay is not. Fourth, the proposed course of actions to restore eelgrass meadows ignores general historical baselines of estuarine and coastal systems by confusing what it means to be “pristine.” Fifth, despite the important experimental evidence indicating strong top–down control in temperate seagrass meadows, Zertuche-González et al. (2009) underestimated the capacity of consumers in structuring dynamics of vegetated soft-bottom communities in San Quintin Bay. Sixth, Ulva expansa may exert positive effects on seagrass ecosystem properties and functions. Instead, we propose protection against the propagation of unsustainable practices in the bay, and the reintroduction of large consumers that are now absent in this ecosystem. An ecosystem-based analysis of the role of Ulva spp. on eelgrass dynamics is needed.