Crustacean epifauna of seagrass and macroalgae in Apalachee Bay,Florida, USA

Epifaunal crustaceans on turtlegrass (Thalassia testudinum) and five dominant macroalgae (Anadyomene stellata, Digenia simplex, Halimeda incrassata, Laurencia poitei and Penicillus lamourouxii) were quantitatively sampled bimonthly over a one-year period from September 1979 to September 1980 in a subtropical seagrass meadow in Apalachee Bay, Florida (northeastern Gulf of Mexico). These plant species exhibited a wide range of morphologies, with surface area-to-biomass ratios differing by over 2.5 times. A similar suite of crustaceans occurred on all macrophytes despite differences in shape or architecture among plant species. Relative abundances of many crustaceans, however, varied among plant hosts. Similarity analysis indicated that the epifaunal associates of T. testudinum were distinct from those of the macroalgae. Species richness was generally higher on turtlegrass than on any of the macroalgae. Abundances of total crustaceans per plant biomass or per plant surface area, on the other hand, were greater on all macroalgal species compared to the seagrass. Abundances (per plant biomass or plant surface area) of 14 of the 16 numerically dominant epifaunal species differed significantly among macrophytes. Twelve of the 16 species had greater abundance on one or more macroalgae, while only two species were more abundant on T. testudinum. Almost half of the dominant species had greatest abundances on the branching red alga L. poitei. Although abundances per plant biomass and plant surface area were greater on macroalgae relative to turtlegrass, densities (individuals per meter square of bottom) of animals associated with T. testudinum were significantly greater than those associated with macroalgae, primarily because of the greater abundance of turtlegrass in the grass bed. Both surface area-to-biomass ratios and degree of branching were poorly correlated with epifaunal abundance and number of species. Neither structural feature is an adequate predictor of faunal abundance and species richness among plant species, especially when macrophytes with very different morphologies are compared.     

Effects of O2, pH and DIC on photosynthetic net-O2 evolution by marine macroalgae

Net photosynthetic O₂ evolution by five marine macroalgae:Ulva lactuca L.,Enteromorpha sp.,Ceramium strictum Harvey,Fucus serratus L., andF. vesiculosus L., collected from Danish waters in the summer of 1983 was followed at increasing O₂ and with pH either fixed close to pH 7, 8 or 9, or drifting upwards during photosynthesis in a closed chamber to determine the effects of changing O₂, pH and DIC (dissolved inorganic carbon) on photosynthesis. Increasing O₂, increasing pH and decreasing DIC together limited O₂ evolution. Raising the O₂ concentration with pH and DIC held constant resulted in less inhibition of net-O₂ evolution than when all three factors acted together. The O₂ inhibition of photosynthesis was similar to the reported O₂ inhibition of ribulose 1,5-bisphosphate carboxylase isolated from lower and higher plants. Net-O₂ evolution as a function of the molar ratio of O₂ to HCO^– ₃ + CO₂ in solution provided a general, linear relationship (r ² = 0.72 to 0.84), predicting inhibition of photosynthesis based on O₂ pH and DIC changing together. Slopes of this relationship, representing competition between O₂ and carbon based on external concentrations, were similar for the five taxonomically different algae, suggesting that similar processes act to reduce net-O₂ evolution.     

Changes in pH as a measure of photosynthesis by marine macroalgae

An automatically operated method for high precision measurements of steady-state photosynthesis by macroalgae was developed. Changes in pH and oxygen content of seawater passing the algae in a flowthrough system, could be measured with extremely high accuracy over very long periods of time. The method is especially suitable for measurements on flowthrough systems with high rates of water exchanges (i.e. short retention time), and can be used to study exchange processes for marine plants, animals and small ecosystems. Since the same measuring unit is used for several flowthrough chambers, the method is very suitable for comparisons between different species, or between differently pretreated specimens of the same species (e.g. in toxicological studies). The method was used to study the ratio: [oxygen production] to [CO₂+H⁺ uptake] at different light intensities for several macroalgae belonging to different systematic groups and from different habitats. At lower photosynthetic rates this ratio was similar for all of the algae studied (1.17±0.02). For brown algae of the fucacean family, the ratio increased by 0.08 units at higher photosynthetic rates. This increase was thought to be related to the crassulacean acid metabolism (CAM)-like strategies connected to these algae. For all other algae studied, the ratio remained constant or decreased slightly (at most by 0.04 units) at higher photosynthetic rates. The relations between the abovementioned ratio and the photosynthetic quotient are discussed on a theoretical basis.     

Modeling the role of macroalgae in a shallow sub-estuary of Narragansett Bay, RI (USA)

Proliferation of macroalgal mats is a frequent consequence of nutrient-driven eutrophication in shallow, photic coastal marine ecosystems. These macroalgae have the potential to significantly modify water quality, plankton productivity, nutrient cycling, and dissolved oxygen dynamics. We developed a model for Ulva lactuca and Gracilaria tikvahiae in Greenwich Bay, RI (USA), a shallow sub-estuary of Narragansett Bay, as part of a larger estuarine ecosystem model. The model predicts the biomass of both species in units of carbon, nitrogen, and phosphorus as a function of primary production, respiration, grazing, decay, and physical exchange, with particular attention to the effects of biomass layering on light attenuation and suppression of metabolic rates. The model successfully reproduced the magnitude and seasonal cycle of area-weighted and peak biomass in Greenwich Bay along with tissue C:N ratios, and highlighted the importance of grazing and inclusion of self-limitation primarily in the form of self-shading to overcome an order of magnitude difference in rates of production and respiration. Inclusion of luxury nutrient uptake demonstrated the importance of internal nutrient storage in fueling production when nutrients are limiting. Macroalgae were predicted to contribute a small fraction of total system primary production and their removal had little effect on predicted water quality. Despite a lack of data for calibration and a fair amount of sensitivity to individual parameter values, which highlights the need for further autecological studies to constrain formulations, the model successfully predicted macroalgal biomass dynamics and their role in ecosystem functioning. Our formulations should be exportable to other temperate systems where macroalgae occur in abundance.     

Settlement of larval blacklip abalone, Haliotis rubra, in response to green and red macroalgae

Surfaces from the habitat of adult Haliotis rubra were tested as inducers of larval settlement to determine the cues that larvae may respond to in the field. Settlement was high on the green algal species Ulva australis and Ulva compressa (Chlorophyta), the articulated coralline algae Amphiroa anceps and Corallina officinalis, and encrusting coralline algae (Rhodophyta). Biofilmed abiotic surfaces such as rocks, sand and shells did not induce settlement. Ulvella lens was also included as a control. Treatment of U. australis, A. anceps and C. officinalis with antibiotics to reduce bacterial films on the surface did not reduce the settlement response of H. rubra larvae. Similarly, treatment of these species and encrusting coralline algae with germanium dioxide to reduce diatom growth did not significantly reduce larval settlement. These results suggest that macroalgae, particularly green algal species, may play an important role in the recruitment of H. rubra larvae in the field and can be used to induce larval settlement in hatchery culture.     

Allowing macroalgae growth forms to emerge: Use of an agent-based model to understand the growth and spread of macroalgae in Florida coral reefs,with emphasis on Halimeda tuna

The growth patterns of macroalgae in three-dimensional space can provide important information regarding the environments in which they live, and insights into changes that may occur when those environments change due to anthropogenic and/or natural causes. To decipher these patterns and their attendant mechanisms and influencing factors, a spatially explicit model has been developed. The model SPREAD (SPatially-explicit Reef Algae Dynamics), which incorporates the key morphogenetic characteristics of clonality and morphological plasticity, is used to investigate the influences of light, temperature, nutrients and disturbance on the growth and spatial occupancy of dominant macroalgae in the Florida Reef Tract. The model species, Halimeda and Dictyota spp., are modular organisms, with an “individual” being made up of repeating structures. These species can also propagate asexually through clonal fragmentation. These traits lead to potentially indefinite growth and plastic morphology that can respond to environmental conditions in various ways. The growth of an individual is modeled as the iteration of discrete macroalgal modules whose dynamics are affected by the light, temperature, and nutrient regimes. Fragmentation is included as a source of asexual reproduction and/or mortality. Model outputs are the same metrics that are obtained in the field, thus allowing for easy comparison. The performance of SPREAD was tested through sensitivity analysis and comparison with independent field data from four study sites in the Florida Reef Tract. Halimeda tuna was selected for initial model comparisons because the relatively untangled growth form permits detailed characterization in the field. Differences in the growth patterns of H. tuna were observed among these reefs. SPREAD was able to closely reproduce these variations, and indicate the potential importance of light and nutrient variations in producing these patterns.     

UV-radiation can affect depth-zonation of Antarctic macroalgae

Due to depletion of stratospheric ozone over polar regions of the Northern and Southern Hemispheres UV-B-radiation has increased at the surface of the earth. Measurements of variable chlorophyll fluorescence were conducted to document UV-induced photoinhibition of photosystem II in cultivated macroalgae with different depth distributions in Antarctica. The reactions during artificial UV-exposure were observed on a short time scale (hours) and in light–dark cycles over several days. The nine species of investigated macroalgae show great differences in UV-tolerance of the photosynthetic process. Photosynthesis of the studied green algae was inhibited to a minor degree, while the brown algae showed an intermediate inhibition of photosynthesis. The response of the studied red algae varied with species. The differences in the degree of inhibition and recovery of photosynthetic efficiency and capacity indicate that UV-radiation is one important factor affecting the vertical distribution of macroalgae in nature. Received: 11 December 1997 / Accepted: 27 March 1998     

Interactive effects of nitrogen and dissolved inorganic carbon on photosynthesis,growth, and ammonium uptake of the macroalgae Cladophora vagabunda and Gracilaria tikvahiae

Dissolved inorganic carbon (DIC) is rarely considered limiting for macroalgae, but some research suggests that under conditions of N sufficiency, photosynthetic capacity is enhanced with DIC enrichment. During spring (April–May) and summer (July–August) 1993, we investigated the interactive effects of nitrogen (N) and DIC on photosynthetic capacity, growth, and nutrient uptake rates of the macroalgae, Cladophora vagabunda (L.) van den Hoek and Gracilaria tikvahiae (McLachlan), dominant species in a temperate eutrophic estuary (Cape Cod, Massachusetts, USA). Water-column CO₂ concentrations showed significant diurnal fluctuations, ranging from a morning CO₂ peak (21 M) to an afternoon low (13 M) during summer, probably associated with metabolic activities in a thick algal mat. Results from instantaneous photosynthesis measurements and microcosm experiments indicate that DIC limits photosynthetic capacity and growth rates of C. vagabunda during summer, perhaps related to tissue N sufficiency and low water-column CO₂ concentrations. For example, this species showed enhanced growth (F=8.69, P<0.02) under DIC but not N enrichment. G. tikvahiae showed marginal DIC enhancement of maximum photosynthetic rate, while growth was significantly stimulated by addition of N. Reduced thallus N of this species during the summer further identifies N as the primary factor limiting growth. In addition, G. tikvahiae has the ability to use DIC in its several forms, while C. vagabunda primarily uses dissolved CO₂. DIC enrichment resulted in a depression of NH₄ ⁺ uptake rates for both species, particularly during summer at saturating (60 M) ammonium levels, suggesting competition between NH₄ ⁺ uptake and DIC acquisition under conditions of N sufficiency. Dominance of C. vagabunda and G. tikvahiae in areas undergoing eutrophication has been attributed to their successful procurement and storage of N as well as to high growth rates. The present study revealed that under conditions of N sufficiency during summer, DIC may control rates of production of these opportunistic macroalgae.     

Carbon acquisition strategies for marine macroalgae

A model system was developed to analyse differences in carbon acquisition strategies among macroalgae. During photosynthesis in a limited volume of seawater the capability of the algae to assimilate inorganic carbon as well as to change the alkalinity of the seawater was analysed. These properties were then related to the status of the carbonate equilibrium system of the seawater. The experimental system was assumed to simulate the conditions in the boundary layer during periods of low water exchange or high intensity irradiations. Fundamental differences were found between different algal classes, suggesting that capabilities to adapt to specific environmental conditions may be connected with dissimilarities in carbon acquisition strategies. In general, green algae were able to reach the highest pH (10.8 at 5°C), and thus to achieve the highest reduction in the level of inorganic carbon via a simple HCO₃ ^–/OH^– ion exchange process. For brown algae, pH increases due to carbon uptake never exceeded pH 9.7 (9.5 in a saltwater scale). In spite of this, members of the Fucaceae (littoral brown algae) were able to extract almost all of the dissolved inorganic carbon (DIC). This was achieved through a gradual decrease in the alkalinity of the enclosed water, so that the carbon assimilation could continue without any concomitant increase in pH. For red algae, the specific response was an increase in the level of inorganic carbon. Thus, for this algal class, no specific strategy for handling a shortage of inorganic carbon was documented. Within each algal class, differences in pH and DIC compensation points could be related to differences in the depths at which the algal species occurred. This paper also introduces a low cost and convenient method of analysing DIC in seawater.     

Biomass, diversity and production of rocky shore macroalgae at two nutrient enrichment and wave action levels

The littoral zone of temperate rocky shores is normally dominated by perennial macroalgae (e.g. Fucus, Ascophyllum, Laminaria), but nutrient enrichment and/or permanently decreased wave action may lead to structural community changes from dominance of perennials to increased amounts of annual opportunistic species (mainly green algae). Macroalgal biomass, diversity and production as well as relationships between the two latter were studied using Solbergstrand’s rocky shore mesocosms in SE Norway in connection with a long-term experimental manipulation of nutrient addition and wave action (high and low levels of both factors applied in a crossed way to eight outdoor basins). After more than 2 years of experimental treatment, the total standing stock of macroalgae was larger in low nutrient than in high nutrient treatments as well as in high wave compared to low wave treatments (in autumn only). For macroalgal functional groups, bushy and filamentous brown and filamentous red algae were generally favoured by low nutrient concentrations, while annual filamentous and sheet-like green algae were stimulated by the nutrient enrichment. There was only one significant interaction between nutrient enrichment and wave action (for brown filamentous algae in autumn) and also only one significant main effect of the wave treatment (for bushy brown algae in autumn). Surprisingly, the high nutrient treatments supported a higher diversity of macroalgae, whereas the low nutrient treatments generally showed higher production rates. Moreover, significantly negative correlations were found between macroalgal diversity and primary productivity in both summer and autumn. This study shows that it is the biological components of the communities subjected to external forcing (nutrient addition or decreased wave action) that regulate production and this contradicts the common misperception that resource production in natural systems simply can be fast-forwarded by fertilization. The negative relationships between diversity and productivity, although a consequence of unexpected results for diversity and production, are also novel and hint towards species identities having more important functional consequences than general species dominance patterns and the amount of species per se. These results also emphasise the context dependency of findings within the field of biodiversity and ecosystem functioning.     

Long-term effects of ultraviolet radiation on growth and photosynthetic performance of polar and cold-temperate macroalgae

Long-term effects of artificial ultraviolet radiation (UV) and natural solar radiation on growth and photosynthetic activity, as measured by chlorophyll fluorescence, were investigated in 13 different polar and cold-temperate macroalgal species. Isolates of five different species from the Arctic and Antarctic were exposed to different light treatments of photosynthetically active radiation (PAR), PAR+UVA and PAR+UVA+UVB. Eight different species collected on the island of Helgoland, North Sea, Germany, were studied in the laboratory and under natural solar radiation conditions. Increase in fresh weight and changes in photosynthetic performance were monitored over a period of 3-4 weeks. The sublittoral polar species, particularly the Antarctic red algal species Gymnogongrus antarcticus and G. turquetii, the Arctic cold-temperate brown alga Alaria esculenta and, very drastically, the Arctic-endemic Laminaria solidungula, exhibited strong inhibiting effects of artificial UVB radiation on growth. In the cold-temperate sublittoral growth of the red algae Phycodrys rubens and, to a lesser extend, Membranoptera alata was substantially inhibited by UV radiation. In contrast, eulittoral species, e.g. Fucus serratus, did not show any differences in growth with respect to artificial irradiation conditions, with or without UV radiation. In the laboratory, some individuals of the green alga Codium fragile exhibited strong morphological changes of the whole thallus, particularly under UVB exposure. In the experimental outdoor set up, growth of most of the algal species was already inhibited by the full solar UV waveband, but, apart from Polyides rotundus, no additional UVB effect could be detected. Changes of in vivo fluorescence were not always consistent with the measurable changes in growth rate, indicating that physiological processes leading to an inhibition of growth may act independently of changes in photosynthetic activity. For the polar species, a general correlation between the natural vertical distribution in the field and the individual sensitivity towards UV radiation was indicated, while for eulittoral species from Helgoland no clear relationship was found. The obtained results show that measuring growth is a good ecological parameter to monitor long-term effects of UV radiation on single macroalgal species and the possible resulting changes of whole algal communities in coastal ecosystems.     

Stressed, but not defenceless: no obvious influence of irradiation levels on antifeeding and antifouling defences of tropical macroalgae

The production of defence metabolites is assumed to be costly in metabolic terms. If this holds true, low-light stress should reduce the ability of seaweeds to defend themselves chemically against herbivory and fouling. We investigated the effect of energy limitation on the defensive status of seaweeds by assessing their attractiveness to mesograzers and their activity against a bivalve macrofouler in comparison with non-stressed conspecifics. The macroalgae Codium decorticatum (Woodw.) M. Howe, Osmundaria obtusiloba (C. Agardh) R. E. Norris, Pterocladiella capillacea (S. G. Gmel.) Santel. and Hommer., Sargassum vulgare C. Agardh and Stypopodium zonale (Lamour.) Papenf. collected at the southeastern Brazilian coast were exposed to six levels of irradiation (between 1 and 180 μmol photons m⁻² s⁻¹) for 10–14 days. After this period, algae from all treatment levels were: (a) processed as artificial food and offered to an amphipod community dominated by Elasmopus brasiliensis Dana and (b) extracted to test for differences in settlement rates of the fouling mussel Perna perna L. on filter paper loaded with the crude extracts. Generally, photosynthesis rates and growth were reduced under low light conditions. Attractiveness to herbivores and macrofoulers, however, was insensitive to energy limitation. We discuss possible explanations for the observed absence of a relationship between light availability and algal defence including the change in nutritional value of the algal tissue, the allocation of resources towards defence instead of growth and the absence of costs for defence.     

Modelling the dynamics of coral reef macroalgae using a Bayesian belief network approach

Macroalgae are a major benthic component of coral reefs and their dynamics influence the resilience of coral reefs to disturbance. However, the relative importance of physical and ecological processes in driving macroalgal dynamics is poorly understood. Here we develop a Bayesian belief network (BBN) model to integrate many of these processes and predict the growth of coral reef macroalgae. Bayesian belief networks use probabilistic relationships rather than deterministic rules to quantify the cause and effect assumptions. The model was developed using both new empirical data and quantified relationships elicited from previous studies. We demonstrate the efficacy of the BBN to predict the dynamics of a common Caribbean macroalgal genus Dictyota. Predictions of the model have an average accuracy of 55% (implying that 55% of the predicted categories of Dictyota cover were assigned to the correct class). Sensitivity analysis suggested that macroalgal dynamics were primarily driven by top–down processes of grazing rather than bottom–up nutrification. BBNs provide a useful framework for modelling complex systems, identifying gaps in our scientific understanding and communicating the complexities of the associated uncertainties in an explicit manner to stakeholders. We anticipate that accuracies will improve as new data are added to the model.     

Light requirements for growth of six shade-acclimated Mediterranean macroalgae

Six mediterranean macroalgae were cultivated for more than 2 yr under shade culture conditions, after which light requirements for growth were investigated at 16±2°C. The saturation light levels for growth in the logarithmic phase were related to the bathymetric distribution of the algae on the shore. The eulittoral to supralittoral red alga Bangia atropurpurea was saturated at a photon fluence rate of 71 mol photons m^-2 s^-1, the upper sublittoral to eulittoral brown algae Scytosiphon lomentaria, Colpomenia peregrina and Kuckuckia spinosa and the sublittoral brown alga Stictyosiphon soriferus at 39 to 71 mol photons m^-2 s^-1, and the deep-water alga Choristocarpus tenellus at 19 mol photons m^-2 s^-1. The minimum light requirements for growth of B. atropurpurea and C. tenellus were determined by observing length increase for 56 d under limiting light conditions. The compensation and minimum irradiances required for growth of B. atropurpurea were 0.5 and 1 mol photons m^-2 s^-1 respectively. The corresponding values for C. tenellus were 0.15 to 0.28 and 0.5 mol photons m^-2 s^-1 respectively. C. tenellus was the siowest-growing species tested at saturating light conditions, but it grew faster than B. atropurpurea at 1 mol photons m^-2 s^-1. Both B. atropurpurea and C. tenellus were able to survive 56 d in darkness, but only the latter grew under darkness in the first 14 d.     

Assimilation efficiency of a temperate-zone intertidal fish (Cebidichthys violaceus) fed diets of macroalgae

We studied assimilation efficiencies of the temperate-zone intertidal fish Cebidichthys violaceus (Girard, 1854) fed in the laboratory on each of the following species of macroalgae: Spongomorpha coalita (Chlorophyta), Ulva lobata (Chlorophyta), Iridaea flaccida (Rhodophyta) and Porphyra perforata (Rhodophyta). Together, these 4 algae make up over 75% of the natural summer diet of C. violaceus. Assimilation efficiency was calculated by proximate organic analysis of food and feces; the amount of ash in food and feces was used as a standard. Depending on the algal species, the fish assimilated 43 to 81% of the protein, 21 to 44% of the lipid, 45 to 62% of the carbohydrate and 31 to 52% of all three classes of organic material combined. These data are the first results showing that a temperate-zone marine fish can assimilate macroalgal constituents. Protein, carbohydrate and total organic material were absorbed more efficiently from rhodophytes than from chlorophytes. Conversely, lipid was absorbed more efficiently from chlorophytes than from rhodophytes. These results are compared with previous work showing that C. violaceus in nature eats more chlorophytes than rhodophytes, but in laboratory preference tests prefers rhodophytes to chlorophytes.     

Method for determination of turgor pressure in macroalgae,with particular reference to the Phaeophyta

A procedure is described for the determination of the internal osmotic pressure and turgor pressure of marine macroalgae, for use in the laboratory and on the shore. A volume-related parameter (either thallus fresh weight, or area) is measured before and after transfer of plant material to a range of hyperosmotic solutions. Plotting the final fresh weight/area as a percentage of the initial value gives a biphasic curve, with an initial component of negative slope due to the change in thallus volume in less extreme hyperosmotic solutions, where the non-rigid thallus contracts in response to decreasing cell turgor pressure. The second component has a shallower slope and represents plasmolysis in more extreme hyperosmotic solutions, i.e., where turgor pressure is reduced to zero and the protoplast shrinks away from the cell wall; the extraprotoplast space created by plasmolysis will be filled with the external solution and thus no further changes in weight occur. These two components intersect at the lowest osmotic pressure at which cell turgor is zero. By correcting for any effects of the cell wall on thallus volume, the relationship can be used to calculate internal osmotic pressure and hence turgor pressure, assuming that the remaining change in thallus volume of the initial component is due entirely to variation in the intraprotoplast volume (approximately equivalent to the intraprotoplast water content, determined by subtraction of the extraprotoplast water and dry weight from the thallus fresh weight). Using this procedure, the turgor pressures of Fucus spiralis L., Ectocarpus siliculosus, (Dillw.) Lyngb. and Laminaria digitata (Huds.) Lamour. (from Fife Ness, Scotland, May–August 1987) in a seawater-based medium were 0.82, 0.58 and 1.34 Osmol kg^–1, respectively. The turgor pressure of F. spiralis on the shore at Fife Ness (June 1987) was 0.74 Osmol kg^–1.     

Effects of detrital non-native and native macroalgae on the nitrogen and carbon cycling in intertidal sediments

The decay of non-native and native seaweed mixing may modify sediment biogeochemistry and organic matter transfers within benthic food webs according to their composition and biomass. The non-native species Sargassum muticum was deliberately added to the sediment of an intertidal sandflat at different biomass and mixed to the native species Ulva sp. and Fucus vesiculosus. The sediment porewater was then ¹³C and ¹⁵N enriched to test whether both detrital diversity and biomass influenced the transfer of porewater carbon and nitrogen to the sediment and to the macrofauna consumers. More ¹⁵N-nitrogen was mobilized to sediments and macrofauna when the 3-species detrital mixing was buried, probably because this mixing provided species-specific compounds such as polyphenols due to the presence of S. muticum and F. vesiculosus, as well as large amounts of nitrogen due to the presence of Ulva. Our study revealed the importance of detrital diversity and non-native seaweeds for the nitrogen cycling in the benthic food web.     

Interactions between wave action and grazing control the distribution of intertidal macroalgae

Canopy-forming macroalgae are key species on temperate rocky shores. However, there is a lack of understanding of how the relative balance of physical and biological factors controls the establishment and persistence of intertidal macroalgae. Here we present an integrated study of the relative importance of wave-induced forces and grazing for the recruitment and survival of the canopy-forming intertidal macroalgae Fucus vesiculosus and F. spiralis. A set of overtopped breakwaters provided a nearly unconfounded gradient in wave exposure between seaward and landward sides. A biomechanical analysis was performed based on empirical measurements of maximum drag forces in breaking waves, a model of long-term maximum wave height, and the breaking stress of Fucus spp. The estimated maximum flow speed (7-8 m/s) on the seaward side of the breakwaters was predicted to completely dislodge or prune Fucus spp. larger than approximately 10 cm, while dislodgment was highly unlikely on the landward side for all sizes. Experimental transplantation of Fucus spp. supported the biomechanical analysis but also suggested that mechanical abrasion may further limit survival in wave-exposed locations. Experimental removal of the limpet Patella vulgata, which was the principal grazer at this site, resulted in recruitment of Fucus spp. on the seaward side. We present a model of limpet grazing that indicates that limpet densities >5-20 individuals/m2 provide a proximate mechanism preventing establishment of Fucus spp., whereas wave action >2 m/s reduces persistence through dislodgment and battering. In a conceptual model we further propose that recruitment and survival of juvenile Fucus spp. are controlled indirectly by wave exposure through higher limpet densities at exposed locations. This model predicts that climate change, and in particular an increased frequency of storm events in the northeast Atlantic, will restrict fucoids to more sheltered locations.