IMPACTS OF MESOGRAZERS ON EPIPHYTE AND ENDOPHYTE GROWTH ASSOCIATED WITH CHEMICALLY DEFENDED MACROALGE FROM THE WESTERN ANTARCTIC PENINSULA: A MESOCOSM EXPERIMENT1

It has been hypothesized that the extensive mesograzer community along the western Antarctic Peninsula regulates epiphytic algae as well as emergent filaments from endophytic species. Should grazing limit growth of fouling or potentially pathogenic microphytes, then Antarctic macrophytes may actually benefit from the remarkably high densities of mesograzer amphipods that occur in these waters. Although initially counterintuitive, the negative impacts of epi/endophyte fouling may outweigh stresses caused by limited amphipod grazing on chemically defended macrophytes by reducing stress from endo/epiphyte biomass. If so, then alleviating mesograzing stress should result in significant increases in endo/epiphytic biomass. To test this hypothesis, a mesocosm experiment was conducted. Individuals representing four common species of Antarctic macroalgae were placed in flow‐through seawater mesocosms. Amphipods were added to five mesocosms at simulated natural densities, while the other five remained herbivore free. At the end of 7 weeks, endo/epiphytic growth on individual macrophytes was quantified. Most species of macroalgae demonstrated noticeably higher instances of endophyte coverage, epiphytic diversity, and diatom colonization in consumer‐free mesocosms than in the presence of amphipods. These data suggest that macroalgae along the western Antarctic Peninsula rely on grazers to control populations of potentially harmful epiphytes. We hypothesize that the chemically defended macroalgal flora lives in mutualism with high densities of mesograzers, providing amphipods with shelter from predation while continually being cleaned of potentially harmful endo/epiphytes.     

Relative effects of nutrient enrichment and grazing on epiphyte-macrophyte (Zostera marina L.) dynamics

The independent and interactive effects of nutrient concentration and epiphyte grazers on epiphyte biomass and macrophyte growth and production were examined in Zostera marina L. (eelgrass) microcosms. Experiments were conducted during early summer, late summer, fall, and spring in a greenhouse on the York River estuary of Chesapeake Bay. Nutrient treatments consisted of ambient or enriched (3× ambient) concentrations of inorganic nitrogen (ammonium nitrate) and phosphate. Grazer treatments consisted of the presence or absence of field densities of isopods, amphipods, and gastropods. epiphyte biomass increased with both grazer removal and nutrient enrichment during summer and spring experiments. The effect of grazers was stronger than that of nutrients. There was little epiphyte response to treatment during the fall, a result possibly of high ambient nutrient concentrations and low grazing pressure. Under low grazer densities of early summer, macrophyte production (g m^–2 d^–1) was reduced by grazer removal and nutrient enrichment independently. Under high grazer densities of late summer, macrophyte production was reduced by enrichment only with grazers absent. During spring and fall there were no macrophyte responses to treatment. The relative influence of epiphytes on macrophyte production may have been related to seasonally changing water temperature and macrophyte requirements for light and inorganic carbon.     

Nutrient impacts on epifaunal density and species composition in a subtropical seagrass bed

The capacity of epifauna to control algal proliferation following nutrient input depends on responses of both grazers and upper trophic level consumers to enrichment. We examined the responses of Thalassia testudinum (turtle grass) epifaunal assemblages to nutrient enrichment at two sites in Florida Bay with varying levels of phosphorus limitation. We compared epifaunal density, biomass, and species diversity in 2 m² plots that had either ambient nutrient concentrations or had been enriched with nitrogen and phosphorus for 6 months. At the severely P-limited site, total epifaunal density and biomass were two times higher in enriched than in unenriched plots. Caridean shrimp, grazing isopods, and gammarid amphipods accounted for much of the increase in density; brachyuran crabs, primary predatory fish, and detritivorous sea cucumbers accounted for most of the increase in biomass. At the less P-limited site, total epifaunal density and biomass were not affected by nutrient addition, although there were more caridean shrimp and higher brachyuran crab and pink shrimp biomass in enriched plots. At both sites, some variation in epifaunal density and biomass was explained by features of the macrophyte canopy, such as T. testudinum and Halodule wrightii percent cover, suggesting that enrichment may change the refuge value of the macrophyte canopy for epifauna. Additional variation in epifaunal density and biomass was explained by epiphyte pigment concentrations, suggesting that enrichment may change the microalgal food resources that support grazing epifauna. Increased epifaunal density in enriched plots suggests that grazers may be able to control epiphytic algal proliferation following moderate nutrient input to Florida Bay. Electronic supplementary material Electronic supplementary material is available for this article at and accessible for authorised users.     

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.     

Extracts of North Sea macroalgae reveal specific activity patterns against attachment and proliferation of benthic diatoms: a laboratory study

A variety of macroalgae (Ceramium rubrum, Corallina officinalis, Palmaria palmata, Mastocarpus stellatus, Fucus vesiculosus, Cladophora rupestris, Ulva sp.) were investigated by scanning electron microscopy to visualize epiphytic colonizers. The macroalgae differed in terms of their epiphytic coverage of bacteria, fungi and diatoms. Macroalgae, largely devoid of epiphytic diatoms, were hypothesized to employ effective antifouling means to reduce epiphytic coverage, whilst heavily fouled macroalgae were proposed to lack antifouling strategies. To test these hypotheses from an allelochemical perspective with regard to fouling diatoms, dichloromethane-methanol (1:1) crude extracts of macroalgae were concentrated in dimethylsulfoxide and investigated in diatom attachment and proliferation assays using four benthic diatoms (Nitzschia sp., Navicula phyllepta, Navicula arenaria and Amphora sp.). Algal extracts exhibited a distinct pattern of activity against the test diatoms, suggesting a targeted and selective effect of macroalgal metabolites on individual fouling diatoms. The main outcome of this study was that visual inspection and quantitative categorization of epiphytic colonizers on macroalgal thalli could not be used to predict reliably whether macroalgae employed a chemical defense mechanism.     

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.     

EPIPHYTE-GRAZER INTERACTIONS ON ZOSTERA MARINA (ANTHOPHYTA: MONOCOTYLEDONES): EFFECTS OF DENSITY ON COMMUNITY FUNCTION1

Interactions between algal epiphytes and their grazers can have a significant impact on the structure and function of eelgrass (Zostera marina L.) meadows. In Puget Sound, the herbivorous gastropod Lacuna variegata Carpenter and its congeners appear to remove large quantities of the epiphytic community from eelgrass blades. When snails at typical field densities were used in microcosms, Lacuna significantly reduced epiphytic biomass and areal productivity. Biomass-specific productivity of the epiphytic community showed an increasing trend with increasing snail density. Epiphytic productivity increased nonlinearly with increasing epiphytic biomass. The commonly used logistic population growth formula adequately described this relationship. Grazing rate also increased nonlinearly with increasing epiphytic biomass. The Hailing equation adequately described the relationship between grazing rate and epiphytic biomass. The proportion of the epiphytic biomass found on the oldest blade of an eelgrass shoot was related linearly to epiphytic biomass, suggesting that a constant fraction of the epiphytic community is lost regardless of epiphytic density. Lacuna clearly removed large quantities of epiphytic material from eelgrass blades, significantly altering community function. Modified Lotka-Volterra equations, incorporating the logistic growth form and Hailing grazing equation, should prove useful in modeling the epiphyte–grazer interaction.     

Amphipods on seaweeds: partners or pests?

Summary Herbivorous marine amphipods have been implicated as important grazers on filamentous and ephemeral algae, and thus as beneficial to macrophytes in reducing overgrowth by epiphytic competitors. In North Carolina, USA, amphipods comprise 97% of all macroscopic animals inhabiting the abundant brown seaweed Sargassum filipendula, and peak in abundance between late winter and early summer. I used outdoor tank experiments to test the species-specific impact of common phytal amphipods on the growth of Sargassum and its epiphytes. The results show that seaweed-associated amphipods are a trophically diverse group that could either increase or decrease host fitness depending on their feeding preferences. The amphipods Ampithoe marcuzii, Caprella penantis, and Jassa falcata each significantly reduced growth of epiphytes on Sargassum plants relative to amphipod-free controls, while Ericthonius brasiliensis had no significant effect on Sargassum or its epiphytes. However, amphipod grazing was not necessarily beneficial to Sargassum. A. marcuzii consumed Sargassum in one outdoor tank experiment, reducing its mass by 11%, while Sargassum plants without amphipods grew by 81%. Epiphytes (mostly diatoms and the filamentous brown alga Ectocarpus siliculosus) and detritus remained abundant on these plants suggesting that A. marcuzii preferred the host to its epiphytes. Similarly, when given simultaneous access to Sargassum and to several common foliose and filamentous epiphytes in the lab, A. marcuzii ate Sargassum almost exclusively. The other three amphipods ate no macroalgae. In contrast to A. marcuzii, C. penantis consistently reduced epiphytes with no negative effect on Sargassum. Thus the species composition of the amphipod fauna can determine whether these animals increase or decrease seaweed fitness.     

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     

Relationships between macroalgal biomass and microbiological quality of water in a phytotreatment pond

The microbiological quality of wastewater in phytotreatment ponds with foliose macroalgae can be influenced by biofilm formation on thallus surface. This hypothesis was tested with an in situ experiment which was carried out in a pond with Ulva spp. receiving wastewater from a land based fish farm at Piombino (Italy). The total bacterial load (TBL) was determined in the inflowing and outflowing waters and a multifactorial design was employed to investigate the effect of different macroalgal biomass. Microbiological analysis revealed a high TBL in the water column (18.4 ± 7.4 × 10⁸ cells ml⁻¹). TBL of inlet water was significantly correlated with quantity and quality of particulate organic matter (POM) of inflowing water, whereas no correlation was found between TBL and POM in the outlet water. A significant decrease in the POM concentration was detected within macroalgal ponds, due to the mechanical action of thalli which favoured POM sedimentation. Nevertheless, great TBLs were found in the outlet water. These findings suggest that TBL probably depended upon macroalgae. Indeed high bacterial density was found on macroalgal thallus surface (~10⁸ cells cm²). Furthermore, high plate counts of faecal bacteria (faecal enterococci) were determined on thallus surface (~ 40 CFU cm²) and outlet water (11 886 ± 3984 CFU 100 ml⁻¹) supporting the evidence that macroalgae negatively affect the microbiological quality of treated water. Bacterial activities in terms of exoenzymatic rates and secondary production were two folds higher in the water within macroalgal beds, than in the open water. These preliminary results suggest that high macroalgal biomass represents a ‚hot spot’ of bacterial density and activity that may affect microbiological quality of the treated water. Bacterial control of inlet water and management of macroalgal biomass through periodic removal are essential for a more efficient treatment of wastewater in phytotreatment ponds.     

Invasion of a Habitat-Forming Seaweed: Effects on Associated Biota

Fucus evanescens is a brown alga of arctic origin that has invaded European coasts. The epiphytic community of F. evanescens in southern Sweden was compared with that of the native Fucus vesiculosus, to examine to what extent an invading seaweed can modify local biodiversity. F. evanescens was much less fouled than F. vesiculosus, supporting both less biomass and fewer species of epiphytes. Multivariate analysis of the most common epiphyte taxa showed that the epiphytic community composition of F. evanescens was not entirely separated from that of F. vesiculosus, but host species contributed significantly to explain the variation in community composition. The biomass of free-living invertebrates was also lower on F. evanescens, although the pattern differed between taxonomic groups. While the biomass of amphipods was lower on F. evanescens, there was no significant difference in biomass of isopods or gastropods between the Fucus species. The good correlation between biomass of epiphytes and free-living animals suggests that the epiphytes play an important role in providing a suitable habitat for many species of free-living epifauna. The study shows that the invasion of F. evanescens affects the environmental conditions for many species associated with the Fucus community but that the direct effect on biodiversity is probably low.     

CONSUMPTION OF ULVA LACTUCA (CHLOROPHYTA) BY THE OMNIVOROUS MUD SNAIL ILYANASSA OBSOLETA (SAY)

Marine invertebrate grazing on temperate macroalgae may exert a significant “top-down” control on macroalgal biomass. We conducted two laboratory experiments to test (1) if consumption by the omnivorous mud snail Ilyanassa obsoleta (Say) on the macroalga Ulva lactuca Linnaeus was a function of food quality (nitrogen content) and (2) if grazing on benthic macroalgae occurred at significant rates in the presence of alternative food sources in the sediment (detritus, larvae, benthic microalgae). Grazing rates were higher for N-enriched macroalgae; however, all snails lost weight when grazing on macroalgae alone, indicating that U. lactuca was a poor food source. The presence of sediment from two sites, a sandy lagoon and an adjacent organic-rich muddy tidal creek, did not affect consumption of macroalgae in microcosm experiments, and the grazing snails were capable of significantly reducing macroalgal biomass associated with both sediment types. Grazing rates by this omnivore were as high as 10.83 mg wet weight·individuals ^{− 1}·d ^{− 1} and were similar to those recorded for herbivorous species. In situ loss rates calculated from average grazing rates per individual and snail abundances (up to 3.5 g dry weight·m ^{− 2}·d ^{− 1}) also were comparable with those calculated for herbivorous species. This level of grazing could remove up to 88% of new macroalgal growth at the lagoon site where the N supply was relatively low but had a much smaller effect (18% of new growth) at the high-nutrient creek site. Snails facilitated macroalgal growth at both sites by increasing tissue N content by 40%–80%. Consumption and digestion of macroalgae aided in the recycling of nutrients temporarily bound in the algae and resulted in enrichment of surficial sediments. Increased N sequestration in the sediments also was associated with an interruption of snail burrowing behavior due to persistent anoxia in sediments rich in decaying algal material. Our data suggest that in shallow lagoons where mud snails and benthic macroalgae coexist, grazing may influence N retention in macroalgal biomass.     

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.     

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     

Effect of nutrient availability, grazer assemblage and seagrass source population on the interaction between Thalassia testudinum (turtle grass) and its algal epiphytes

Seagrass leaves are often densely covered by epiphytic algae which can suppress seagrass productivity and has been implicated in declines of seagrass meadows worldwide. The net effect of epiphytes on seagrass growth and morphology depends on the independent and interactive effects of a variety of factors, including nutrient availability and the intensity of grazing on epiphytes. Here I report the results of a mesocosm experiment designed to test the effects of nutrient addition and within-functional group variation (grazer species composition and the source population of seagrass) on the strength of the interactions among grazers, epiphytes, and turtle grass (Thalassia testudinum). Turtle grass ramets from two sites in the northern Gulf of Mexico were cleared of epiphytes and transplanted into common-garden mesocosms. Replicate ramets were grown in a split-split plot design with two levels of dissolved nutrients and four different grazer species combinations (Tozeuma carolinense alone, Pagurus maclaughlinae alone, both species together, and no grazers present). As expected, grazers had a significant negative effect on epiphyte biomass/leaf area and a significant positive effect on turtle grass growth in the mesocosms. The two species were more similar in their direct effects on epiphyte biomass than in their indirect effects on turtle grass growth; this may reflect differences in epiphyte community composition under different grazer treatments. The effect of nutrient addition on turtle grass growth depended critically on the intensity of grazing: in the presence of grazers, turtle grass tended to produce a greater biomass of new leaf tissue in the tanks with nutrients added than in the control tanks. However, when grazers were absent, the direction of the effect was reversed, and plants with nutrients added grew less than the control plants. The two source populations of turtle grass differed significantly in epiphyte biomass/leaf area accrued in the mesocosms as well as in the strength of the effect of grazers on turtle grass growth. This suggests that population differentiation in seagrass interactions with epiphytes, as well as spatial and temporal variation in resources and grazer community composition, can greatly effect the role of epiphytes in limiting seagrass productivity.     

The role of macroalgae in epiphytism of the toxic dinoflagellate Gambierdiscus toxicus (Dinophyceae)

Populations of the toxic, epiphytic dinoflagellate Gambierdiscus toxicus Adachi et Fukuyo are asSociated closely with Jania sp. on Hitiaa and Papara fringing reefs in Tahiti. Small populations were also observed to be asSociated with Amphiroa sp. and Halimeda opuntia (L.) Lamouroux. The cells attached themselves to the thallus by means of a short thread. When the thalli were irradiated, the cells began to detach from them and swim around the branches. The swimming cells stopped and attached to substrata when a disturbance occurred. The attached cells began to swim within a short time under light conditions when the thallus of Jania sp. were placed near the attached cells. Amphiroa sp. and H. opuntia also induced this re-commencement of swimming of the attached cells. These observations suggest that G. toxicus usually swims around macroalgal thalli on coral reefs. When sudden disturbance or strong water motion occurs, they attach to the surface of macroalgae and are not dispersed. Soon after water motion becomes slow, the cells begin to swim into the water around the thalli. The epiphytism of G. toxicus is different from epiphytic pennate diatoms, most of which adhere to the thallus all the time. The population of G. toxicus is maintained as an asSociation to a limited number of species of macroalgae which support the re-commencement of swimming after disturbance.     

Palatability of the Antarctic rhodophyte Palmaria decipiens (Reinsch) RW Ricker and its endo/epiphyte Elachista antarctica Skottsberg to sympatric amphipods

Although many macroalgae that occur throughout Western Antarctic Peninsular waters are known to produce defensive secondary metabolites that deter grazing, the rhodophyte, Palmaria decipiens is palatable to several sympatric meso- and macro-grazers. It has been hypothesized that high levels of mesoherbivory by amphipods may account for the conspicuous lack of filamentous epiphytes emerging from the thalli of marcophytes in this region. Nonetheless, Elachista antarctica is a filamentous phaeophyte found growing within, and emerging from the thallus of the rhodophyte P. decipiens. It is surprising that E. antarctica occurs exclusively in association with a palatable species of macroalgae considering the standing biomass of other chemically defended unpalatable species is very high. We tested the hypothesis that E. antarctica grows on P. decipiens due to the host's overwhelming palatability compared to that of the epiphyte. That is, the hypothesis that mesograzers prefer the host over the epiphyte, grazing around emerging filaments. Choice and no choice feeding assays with live tissues of E. antarctica and P. decipiens were conducted in three different trials with four sympatric amphipod species (Prostebbingia gracilis, Gondogeneia antarctica, Oradarea bidentata, and Paraphimedia integricauda) commonly found in association with P. decipiens. G. antarctica consumed both species but ate P. decipiens at a faster rate than the epiphyte in two of three trials. P. gracilis, O. bidentata, and P. integricauda fed on the epiphyte, E. antarctica at faster rates than upon P. decipiens. Aggressive grazing of the epiphyte by this suite of amphipods indicates that differences in palatability and differences in grazing pressure on host and epiphyte do not explain the exclusive epiphytism of E. antarctica on P. decipiens.     

Effects of macrophyte growth forms on invertebrate communities in saline lakes of the Wyoming High Plains

In saline lakes, areal cover and both species and structural diversity of macrophytes often decline as salinity increases. To assess effects of the loss of certain macrophyte growth forms, we characterized benthic and epiphytic invertebrates in three growth forms (thin-stemmed emergents, erect aquatics, and low macroalgae) in oligosaline lakes (0.8–4.2 mS cm⁻¹) of the Wyoming High Plains, USA. We also measured the biomass and taxonomic composition of epiphytic and benthic invertebrates in two erect aquatics with very similar structure that are found in both oligosaline (Potamogeton pectinatus) and mesosaline (9.3–23.5 mS cm⁻¹) (Ruppia maritima) lakes. Although total biomass of epiphytic invertebrates varied among oligosaline lakes, the relative distribution of biomass among growth forms was similar. For epiphytic invertebrates, biomass per unit area of lake was lowest in emergents and equivalent in erect aquatics and low macroalgae; biomass per unit volume of macrophyte habitat was greatest in low macroalgae. For benthic invertebrates, biomass was less beneath low macroalgae than other growth forms. Taxonomic composition did not differ appreciably between growth forms for either benthic or epiphytic invertebrates, except that epiphytic gastropods were more abundant in erect aquatics. Total biomass of epiphytic and benthic invertebrates for the same growth form (erect aquatic) did not differ between oligosaline (Potamogeton pectinatus) and mesosaline (Ruppia maritima) lakes, but taxonomic composition did change. In the oligosaline to mesosaline range, direct toxic effects of salinity appeared important for some major taxa such as gastropods and amphipods. However, indirect effects of salinity, such as loss of macrophyte cover and typically higher nutrient levels at greater salinities, probably have larger impacts on total invertebrate biomass lake-wide.     

Are epiphytes a significant component of intertidal Zostera noltii beds?

The role of epiphytes in an intertidal Zostera noltii seagrass bed in Marennes-Oléron Bay was assessed in comparison with the other main benthic primary producers (Z. noltii, microphytobenthos) at two bathymetric levels and on a seasonal basis. Assemblage and biomass of epiphytes were studied using scanning electron microscopy (SEM). Z. noltii and its detrital matter followed a typical seasonal pattern: microphytobenthos was present in large quantities throughout the year representing 21% of the total biomass while detrital matter, above-ground parts and below-ground parts accounted for 65, 9 and 5%, respectively. Only two species of epiphytic diatoms, Cocconeis scutellum and Cocconeis placentula, were observed on seagrass leaves. Epiphyte biomass was very low, representing on average less than 0.001% of that of microphytobenthos or leaves. This low epiphyte biomass is linked with the absence of macroalgae and also with the low biovolume of Cocconeis, which formed a monolayer of cells on leaves. This can be explained by the severe conditions of the intertidal and the high leaf turn-over of Z. noltii leaves.     

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.