Wave-exposed Shores Research Articles

Loss, resilience and recovery of kelp forests in a region of rapid ocean warming.

Changes in kelp abundances on regional scales have been highly variable over the past half-century owing to strong effects of local and regional drivers. Here, we assess patterns and dominant environmental variables causing spatial and interspecific variability in kelp persistence and resilience to change in Nova Scotia over the past 40 years. We conducted a survey of macrophyte abundance at 251 sites spanning the Atlantic coast of Nova Scotia from 2019 to 2022. We use this dataset to describe spatial variability in kelp species abundances, compare species occurrences to surveys conducted in 1982 and assess changes in kelp abundance over the past 22 years. We then relate spatial and temporal patterns in abundance and resilience to environmental metrics. Our results show losses of sea urchins and the cold-tolerant kelp species Alaria esculenta, Saccorhiza dermatodea and Agarum clathratum in Nova Scotia since 1982 in favour of the more warm-tolerant kelps Saccharina latissima and Laminaria digitata. Kelp abundances have increased slightly since 2000, and Saccharina latissima and L. digitata are widely abundant in the region today. The highest kelp cover occurs on wave-exposed shores and at sites where temperatures have remained below thresholds for growth (21 °C) and mortality (23 °C). Moreover, kelp has recovered from turf dominance following losses at some sites during a warm period from 2010 to 2012. Our results indicate that dramatic changes in kelp community composition and a loss of sea urchin herbivory as a dominant driver of change in the system have occurred in Nova Scotia over the past 40 years. However, a broad-scale shift to turf-dominance has not occurred, as predicted, and our results suggest that resilience and persistence are still a feature of kelp forests in the region despite rapid warming over the past several decades.

Effects of temperature and pH on the growth, calcification, and biomechanics of two species of articulated coralline algae

Ocean warming and acidification are predicted to impact the physiology of marine organisms, especially marine calcifiers that must deposit calcium carbonate and resist dissolution. Of particular concern are articulated coralline algae, which must maintain both calcified segments (intergenicula) and uncalcified joints (genicula) in order to thrive along wave-swept rocky coastlines. We examined the effect of pH and temperature, both individually and in combination, on the growth, calcification, and biomechanical properties of 2 species of articulated coralline algae, Corallina vancouveriensis and Calliarthron tuberculosum, common on wave-exposed shores in the NE Pacific. Increased temperature and reduced pH were found to reduce growth rates in both species (30-89% lower) but had little influence on the amount of intergenicular calcium carbonate or on the genicular biomechanical properties of these species. Results suggest that although growth rates may decline, these 2 coralline species will maintain the integrity of their tissues and continue to persist under future climate stress.

Comparing the genetic diversity and population structure of sister marine snails having contrasting habitat specificity

To grasp the processes of spatial genetic structuring in open and connectable marine environments is the principal study goal in molecular biological studies. Comparative seascape genetics using multiple species are a powerful approach to understand the physical geographic and oceanographic effects on genetic variation. Besides, species-specific ecological traits such as dispersal abilities and habitat specificity are important factors for spatial genetic structuring. We focused on the sister marine snail species Tegula kusairo and T. xanthostigma around the Japanese mainland, which have contrasting habitat specificities for wave strength. Tegula kusairo only inhabits sheltered coastal environments, while T. xanthostigma is found mainly on wave-exposed rocky shores facing the open sea. We estimated their genetic diversity indices and levels of population differentiation based on mtDNA. We found that the genetic diversity of T. kusairo was lower than that of T. xanthostigma, while their level of population genetic differentiation was higher than that of T. xanthostigma. Namely, the species specific to weak wave environments had a higher level of population genetic differentiation than the species specific to strong wave action. Ecological traits linked not only to dispersal abilities but also to habitat specificity can influence genetic variation in a pair of closely related sister species distributed in the same seascape.

Influence of environmental variables over multiple spatial scales on the population structure of a key marine invertebrate

Quantifying scale-dependent patterns and linking ecological to environmental variation is required to understand mechanisms regulating biodiversity. We conducted a large-scale survey in rocky shores along the SE Brazilian coast to examine spatial variability in body size and density of an intertidal barnacle (Chthamalus bisinuatus) and its relationships with benthic and oceanographic predictors. Both the size and density of barnacles showed most variation at the smallest spatial scales. On average, barnacle body size was larger on shores located in areas characterised by higher chlorophyll levels, colder waters, low wave action and low influence of freshwater. Barnacles were more abundant at wave-exposed shores. We identified critical scales of spatial variation of an important species and linked population patterns to essential environmental predictors. Our results show that populations of this barnacle are coupled to scale-dependent oceanographic variation. This study offers insights into the mechanisms regulating coastal populations along a little studied coastline.

Facilitation of an invader by a native habitat-former increases along interacting gradients of environmental stress.

Native habitat-forming species can facilitate invasion by reducing environmental stress or consumer pressure. However, the intensity of one stressor along a local gradient may differ when expanding the scale of observation to encompass major variations in background environmental conditions. In this study, we determined how facilitation of the invasive porcelain crab, Petrolisthes elongatus, by the native tube-forming serpulid, Galeolaria caespitosa, varied with environmental gradients at local (tidal height) and larger (wave exposure) spatial scales. G.caespitosa constructs a complex calcareous matrix on the underside of intertidal boulders and we predicted that its positive effects on P.elongatus density would increase in intensity with shore height and be stronger at wave-sheltered than wave-exposed locations. To test these predictions, we conducted two experiments. First, we determined the effects of serpulid presence (boulders with live or dead serpulid matrix vs. bare boulders) at six shore heights that covered the intertidal distribution of P.elongatus. Second, we determined the effects of serpulid presence (present vs. absent), shore height (high vs. low) and wave exposure (sheltered vs. exposed) on crabs across six locations within the invaded range in northern Tasmania, Australia. In Experiment 1, the presence of serpulids (either dead or alive) enhanced P.elongatus densities at all shore heights, with facilitation intensity (as determined by a relative interaction index; RII) tending to increase with shore height. In Experiment 2, serpulids facilitated P.elongatus across shore heights and wave exposures, although crab densities were lower at high shore levels of wave-sheltered locations. However, the intensity of crab facilitation by serpulids was greater on wave-sheltered than on wave-exposed shores, but only at the high shore level. This study demonstrates that local effects of native habitat-formers on invasive species are dependent on prevailing environmental conditions at larger spatial scales and that, under more stressful conditions, invaders become increasingly reliant on positive interactions with native habitat-formers. Increased strength of local-scale facilitation by native species, dampening broader scale variations in environmental stressors, could enhance the ability of invasive species to establish self-sustaining populations in the invaded range.

Beyond tides: surge-dominated submersion regimes on rocky shores of central Chile

By definition, intertidal organisms are exposed to fluctuations between submersion and exposure to air due to the variation in the sea level caused, primarily, by tides and waves. Such submersion regimes have great consequences on the evolution, physiology, behavior, and ecology of intertidal marine organisms, but, paradoxically, our quantitative knowledge about submersion regimes is extremely limited, especially in wave-exposed shores. We here quantify submersion regimes during summer (February 2012) in a wave-exposed rocky shore of central Chile (Las Cruces, 33°30′S, 71°38′W) using a remote sensing method that permits high-frequency measurements. We provide metrics of submersion for the high (barnacle), mid (mussel) and low (upper kelp) intertidal zones and developed statistical models to hindcast submersion regimes from tide and wave data. Results revealed that the submersion consists solely of numerous but brief submersion events produced by waves. The three intertidal zones therefore experience intermittent submersion most of the time, and submersion events are so brief that, in these three zones, emersion predominates over submersion. In this region, submersion cannot be determined from tidal charts, because most of the intertidal zone lies above the highest tidal level, but still the tidal cycle strongly modulates submersion regimes, by altering wave swash into the shore. Combined statistical models including tides and waves are necessary to reproduce the most biologically relevant aspects of submersion. We discuss the consequences of these findings for our understanding of adaptation to intertidal life, and the design of studies on responses to current environmental variability and future climate change.

Recreational boating degrades vegetation important for fish recruitment

Recreational boating increases globally and associated moorings are often placed in vegetated habitats important for fish recruitment. Meanwhile, assessments of the effects of boating on vegetation, and potential effects on associated fish assemblages are rare. Here, we analysed (i) the effect of small-boat marinas on vegetation structure, and (ii) juvenile fish abundance in relation to vegetation cover in shallow wave-sheltered coastal inlets. We found marinas to have lower vegetation cover and height, and a different species composition, compared to control inlets. This effect became stronger with increasing berth density. Moreover, there was a clear positive relationship between vegetation cover and fish abundance. We conclude that recreational boating and related moorings are associated with reduced cover of aquatic vegetation constituting important habitats for juvenile fish. We therefore recommend that coastal constructions and associated boating should be allocated to more disturbance tolerant environments (e.g. naturally wave-exposed shores), thereby minimizing negative environmental impacts.

Threats to large brown algal forests in temperate seas: the overlooked role of native herbivorous fish

Canopy-forming algae are declining globally due to multiple disturbances. This decline has recently been on the increase due to the spread of some tropical herbivorous fishes. This new phenomenon has drawn attention to the effects of fish herbivory in temperate areas, which have been assumed to be negligible compared to that of invertebrates, such as sea urchins. In this study, the impact of a Mediterranean native herbivorous fish (Sarpa salpa, salema) was assessed on the canopy-forming seaweed Cystoseira amentacea var. stricta. Cystoseira amentacea forms belts in the infralittoral fringe of wave-exposed shores, which has so far been considered a refuge from fish herbivory. To test the effects of salema feeding on natural C. amentacea belts, an innovative herbivore deterrent device was conceived. Salema had a significant effect on C. amentacea by decreasing algal size, biomass and fertility, by up to 97%. The results suggest that the contribution of salema feeding to the loss of Cystoseira forests in the Mediterranean may have been overlooked. In addition, the analysis of temporal and spatial patterns of salema landings in the Mediterranean Sea suggests that salema abundance may have increased recently. Thus, along with invertebrate herbivory and anthropogenic stressors, fish herbivory may also represent a potential threat to algal forests in temperate areas.

Morphological and physiological differences between two morphotypes of Durvillaea antarctica (Phaeophyceae) from the sub-Antarctic ecoregion of Magallanes, Chile

The coastal marine ecosystems of the sub-Antarctic ecoregion of Magallanes in southern Chile are strongly characterized by environmental heterogeneity with a unique and high biodiversity of benthic macroalgae. The kelp Durvillaea antarctica (Phaeophyceae) is abundant in the intertidal in this region. Two morphotypes, one with elongated-cylindrical (“morphotype EC”) and one with laminar (“morphotype L”) blades, have been collected from a wave-protected site at the Strait of Magellan and the wave-exposed shores of the isolated Otway Sound. Morphometric and physiological (by chlorophyll fluorescence) analyses showed significant differences between the two morphotypes regarding the morphology, the maximum electron transport rates (ETRmax), and the chlorophyll c concentration. These morphological and physiological differences can be linked to their hydrodynamic environment. The presence of two different species in the region of Magallanes based on the measured parameters is discussed. This study may be useful for future genetic and biotechnological investigations and draws attention to D. antarctica as a species of commercial value among the marine resources of Chile.

First visual record of rare purple dogwhelks ( Nucella lapillus) on the Atlantic coast of Nova Scotia, Canada.

The dogwhelk Nucella lapillus is a rocky intertidal gastropod of the North Atlantic coast. Individual shell color varies. Common colors range between white and brown, with darker dogwhelks being more affected by heat stress than lighter-colored conspecifics. Other reported shell colors are black, mauve, pink, yellow, and orange from European coasts, red and grey from the Bay of Fundy coast of New Brunswick and Nova Scotia (Canada), and purple, black, gray, yellow, and orange from the coasts of Maine and Massachusetts (USA), with purple being considered as a rare color. On the Atlantic coast of Nova Scotia, dogwhelks are active from April until November, but information on dogwhelk shell color is missing for this coast. On 16 June 2016, we found two purple dogwhelks in the mid-to-high intertidal zone of a moderately wave-exposed rocky shore near DuncansCove, on the Atlantic coast of Nova Scotia while collecting dogwhelks (n= 1000) for manipulative field experiments. All other dogwhelks collected on that day were of common white and brown colors. During earlier dogwhelk collections in Atlantic Nova Scotia (between 2011-2013) and field surveys in Duncans Cove (between 2014-2016), we did not find any purple dogwhelks, indicating the rareness of this color in that region. Interestingly, the purple dogwhelks were detected on a relatively cool day (12.3 ± 0.4 °C, mean ± se, n= 96 temperature measurements) compared to the intertidal temperatures of all other survey days (≥ 18.2 ± 0.5 °C), suggesting that purple dogwhelks may find it less thermally stressful to venture out of crevices and macroalgal cover under relatively cool temperatures. Our observations provide the first visual record of rare purple dogwhelks on the Atlantic coast of Nova Scotia, Canada.

A 20-year dataset of species replacement patterns in the middle-intertidal zone of Tatoosh Island, Washington, USA

This data set documents changes in the sessile species occupying several rock benches on wave-exposed shores of Tatoosh Island, Washington, USA from 1993-2012. Plots and transects were located within the middle intertidal zone dominated by the mussel Mytilus californianus. Data were taken in 14 60 x 60 cm quadrats positioned at two corners with permanent marking screws. A 10 x 10 grid defined by the intersection of equally-spaced monofilament lines yielded a set of 100 fixed points per quadrat per census. Plots were generally located initially in sites that had undergone natural wave disturbance in the past 0-3 years, to better document transient successional dynamics, and most plots have experienced 1-2 disturbance/succession cycles over the course of the data collection. A further set of points was monitored on 11 permanent transects 9.1 m long with 30 initially randomly placed points, which cover a broader span of shoreline than the quadrats. The data collection was implemented to parameterize Markov Chain models and use these to make predictions about the effects of local species extinction that could subsequently be tested in an experimentally tractable ecosystem. To date, the data have been used in 1) a parameterization and analysis of a basic multi-species Markov Chain model, 2) a spatially-explicit cellular automata, 3) a reformulation, parameterization and experimental test of the Neutral Theory of Biodiversity, 4) a comparative analysis of Markov Chain models across different marine habitats, 5) development of an approach to link global change to multi-species interactions using an environment-dependent (ocean acidification) series of Markov Chain models, 6) analysis of changes in system dynamics following experimental species extinction, and 7) parameterization of population dynamic models of mussels revealing density-linked stochastic patterns. The Markov Chain and Neutral models have subsequently been tested in independent experiments. These data may be of further use in analyzing detailed patterns of species transitions, as well as more standard analyses of spatial and temporal patterns of species abundance and richness. This article is protected by copyright. All rights reserved.

Attachment strength of the herbivorous rockweed isopod, Idotea wosnesenskii (Isopoda, Crustaceae, Arthropoda), depends on properties of its seaweed host

Wave-induced forces can affect the distribution of intertidal organisms, changing the structure of communities along wave-swept shores. The risk of dislodgement for an intertidal animal depends on the forces experienced, relative to the force that animal is able to apply to resist dislodgement (hereby termed attachment strength). Isopods in the genus Idotea (Isopoda, Crustaceae, Arthropoda) are well adapted to wave-swept environments. They possess hooked claws on their walking appendages (peripods) that allow them to attach to seaweeds. In this study, we sought to compare the attachment strength of this isopod from various macroalgae. We used previously reported hydrodynamic data (drag coefficient) for Idotea wosnesenskii to predict the required velocity for dislodgement. Using this information, we were able to predict whether differences in attachment strength could limit substratum choice, and whether wave velocities could be mediating algal–invertebrate interactions. We found that attachment strength was dependent on substratum, and that seaweed breaking stress (MPa) and thickness (m) act as significant predictors of isopod attachment strength. Furthermore, we determined that differences between substrata were relevant to velocities commonly experienced in wave-swept environments. I. wosnesenskii are likely limited to stronger and thicker seaweeds in areas with greater wave intensity. Here, we identify a novel ecological implication of seaweed material properties that could be impacting organismal interactions along wave-exposed shores.

Forecasting invasions: resource use by mussels informs invasion patterns along the South African coast

Invasive species are often more able to rapidly and efficiently utilise resources than natives, and comparing per capita resource use at different resource densities among invaders and trophically analogous natives could allow for reliable predictions of invasiveness. In South Africa, invasion by the Mediterranean mussel Mytilus galloprovincialis has transformed wave-exposed shores, negatively affecting native mussel species. Currently, South Africa is experiencing a second mussel invasion with the recent detection of the South American Semimytilus algosus. We tested per capita uptake of an algal resource by invading M. galloprovincialis, S. algosus, and the native Aulacomya atra at different algal concentrations and temperatures, representing the west and south coasts of South Africa, to examine whether their per capita resource use could be a predictor of their spread and subsequent invasiveness. Regardless of temperature, M. galloprovincialis was the most efficient consumer, significantly reducing algal cells compared to the other species when the resource was presented in both low and high starting densities. Furthermore, these findings aligned with a greater biomass of M. galloprovincialis on the shore in comparison with the other species. Resource use by the new invader S. algosus was dependent on the density of resource and, although this species was efficient at low algal concentrations at cooler temperatures, this pattern broke down at higher algal densities. This was once more reflected in lower biomass in surveys of this species along the cool west coast. We therefore forecast that S. algosus will be become established along the south coast; however, we also predict that M. galloprovincialis will maintain dominance on these shores.

Effects of wave action and grazers on frond perforation of the green alga, Ulva australis

2 of Ulva frond, hole areas ranged from 0.37 to 5.94 cm 2 , and between 4.9 and 36.2 holes were observed. Fourteen residential animal species were observed at the three evaluated sites, 75.0 (Haseom) to 408.7 individuals 100 g -1 Ulva (Pohang) per site. The dominant residential species at each site differed with Amphithoe sp. at Haseom, Monodonta spp. at Pohang, and Pagurus sp. at Woedo. The growth (frond area, wet weight) and hole number of Ulva fronds, and the number of residential animals were significantly greater in samples collected from the sheltered shore than the wave-exposed shore of Seongsan. The present results showed U. australis grew well at sheltered shores and had more holes on the fronds due to abundance of residen- tial animals. The dominant residential animals (crabs, gammaridea, and snails) were similar in the Ulva populations of sheltered and wave-exposed shores, but greater species diversity was observed at the exposed shore (18 species ver. 11 species). In conclusion, U. australis is a keystone species providing habitat to various invertebrates and frond holes are positively correlated to the number of residential animals.

Three new macroalgae from the Three Kings Islands New Zealand including the first southern Pacific Ocean record of the Furcellariaceae (Rhodophyta)

:The Three Kings Islands are an uninhabited archipelago 57 km north of the North Island of New Zealand. Three new species of red algae were described from the Three Kings Islands: Perplexiramosus clintonii gen. et sp. nov., the first record of a member of the Furcellariaceae (Gigartinales) in the South Pacific Ocean, found growing exclusively on Sargassum johnsonii, and two species found on the steep, wave-exposed intertidal shores. Pyropia francisii sp. nov. (Bangiaceae) had a rosette growth form and a very distinctive arrangement of the reproductive regions on the blade. Gigartina ewenii sp. nov. (Gigartinaceae) grew in clumps and had channelled thalli that fork near the apex, with cystocarps and tetrasporangial sori on upper dichotomies. These algae have not been collected in any other areas and are believed to be endemic. This study adds to our knowledge of the seaweed flora of New Zealand and to our understanding of diversity on this special island group.

Does wave exposure determine the interactive effects of losing key grazers and ecosystem engineers?

The consequences of biodiversity loss in the face of environmental change remain difficult to predict, given the complexity of interactions among species and the context-dependency of their functional roles within ecosystems. Predictions may be enhanced by studies testing how the interactive effects of species loss from different functional groups vary with important environmental drivers. On rocky shores, limpets and barnacles are recognised as key grazers and ecosystem engineers, respectively. Despite the large body of research examining the combined effects of limpet and barnacle removal, it is unclear how their relative importance varies according to wave exposure, which is a dominant force structuring intertidal communities. We tested the responses of algal communities to the removal of limpets and barnacles on three sheltered and three wave-exposed rocky shores on the north coast of Ireland. Limpet removal resulted in a relative increase in microalgal biomass on a single sheltered shore only, but led to the enhanced accumulation of ephemeral macroalgae on two sheltered shores and one exposed shore. On average, independently of wave exposure or shore, ephemeral macroalgae increased in response to limpet removal, but only when barnacles were removed. On two sheltered shores and one exposed shore, however, barnacles facilitated the establishment of fucoid macroalgae following limpet removal. Therefore, at the scale of this study, variability among individual shores was more important than wave exposure per se in determining the effect of limpet removal and its interaction with that of barnacles. Overall, these findings demonstrate that the interactive effects of losing key species from different functional groups may not vary predictably according to dominant environmental factors.

Deja vu? A second mytilid mussel, Semimytilus algosus, invades South Africa's west coast

A second marine mussel is shown to have invaded South Africa's west coast. Molecular techniques, based on intraspecific gene sequence divergences, prove its identity as Semimytilus algosus, a member of the family Mytilidae, native to Chile. The identity of an older introduced population found in Namibia is also confirmed. The present geographic range of S. algosus in South Africa extends some 500 km, from Groenriviersmond in the north to Bloubergstrand in the south. Together with Mytilus galloprovincialis, another previously established invasive mussel, S. algosus has become a dominant intertidal organism on wave-exposed rocky shores across this region. Both species are now much more abundant intertidally than either of the indigenous mussels Aulacomya ater and Choromytilus meridionalis, which have become largely confined to sublittoral and sand-inundated habitats respectively. The two invasive mussels display strong spatial segregation, with M. galloprovincialis dominating the midshore and S. algosus blanketing the lower shore. Through a combination of its small size and high abundance, S. algosus is likely to greatly increase food availability for a range of intertidal predators, many of which cannot consume mussels above a threshold size.

Biogeographic variation in behavioral and morphological responses to predation risk

The expression of prey antipredator defenses is often related to ambient consumer pressure, and prey express greater defenses under intense consumer pressure. Predation is generally greater at lower latitudes, and antipredator defenses often display a biogeographic pattern. Predation pressure may also vary significantly between habitats within latitudes, making biogeographic patterns difficult to distinguish. Furthermore, invasive predators may also influence the expression of prey defenses in ecological time. The purpose of this study was to determine how these factors influence the strength of antipredator responses. To assess patterns in prey antipredator defenses based upon geographic range (north vs. south), habitat type (wave-protected vs. wave-exposed shores), and invasive predators, we examined how native rock (Cancer irroratus) and invasive green (Carcinus maenas) crab predators influence the behavioral and morphological defenses of dogwhelk (Nucella lapillus) prey from habitats that differ in wave exposure across an ~230 km range within the Gulf of Maine. The expression of behavioral and morphological antipredatory responses varied according to wave exposure, geographic location, and predator species. Dogwhelks from areas with an established history with green crabs exhibited the largest behavioral and morphological antipredator responses to green crabs. Dogwhelk behavioral responses to rock crabs did not vary between habitats or geographic regions, although morphological responses were greater further south where predation pressure was greatest. These findings suggest that dogwhelk responses to invasive and native predators vary according to geographic location and habitat, and are strongly affected by ambient predation pressure due to the invasion history of an exotic predator.

Distribution and reproductive biology of the little cuttlefish Sepiola atlantica (Cephalopoda: Sepiolidae) around Anglesey, North Wales

Sepiola atlantica were captured in seine nets at twelve locations around the coast of Anglesey, UK. Animals were most abundant on the very sheltered shore of Y Foryd Bay and the wave-exposed shore of Traeth Penrhos, and these two locations were further sampled seasonally to examine the seasonal occurrence, population structure and reproductive biology of S. atlantica. Sepiola atlantica (5–28 mm dorsal mantle length (DML)) migrated inshore seasonally and first appeared in July where they attained peak abundances between July and August at seawater temperatures of 17°C. Numbers declined between September and October as temperatures fell below 15–16°C, and in late October, they migrated offshore. Male S. atlantica was significantly smaller and matured at a smaller DML than females. The number of spermatangia on the bursa copulatrix of female S. atlantica varied seasonally attaining maximum numbers in October with a mean of 22 spermatangia on the bursa copulatrix of Y Foryd Bay females. At Y Foryd Bay and Traeth Penrhos, the number of spermatophores in male and potential fecundity in female S. atlantica ranged between 1 and 147 and 25 and 141, respectively. The spermatophoric complex and gonadosomatic index showed a high degree of variability in individuals of similar wet body weight with the female gonad constituting a far greater percentage of the total wet body weight than the male gonad. It is concluded that S. atlantica of all sizes and maturity stages congregate in the shallow waters around Anglesey between July and October when environmental conditions are favourable for enhanced growth and maturation and where the high numbers of animals enhance opportunities for mating.

Interactive effects of losing key grazers and ecosystem engineers vary with environmental context

Loss of biodiversity may cause significant changes to ecosystem structure and functioning. Evidence from long-term in situ removal experiments is rare but important in determining the effects of biodiversity loss against a background of environmental variation. Limpets and mussels are thought to be important in controlling community structure on wave-exposed shores in the UK: limpets as key grazers, mussels as ecosystem engineers. A long-term factorial removal experiment revealed interactive effects that varied between 2 shores in SW England. At one site (Harlyn), removing limpets caused a significant shift in community structure, but where limpets were lost, the presence or absence of mussels made little difference. Where limpets were present, however, the removal of mussels changed the structure and variability of the community. At the other site (Polzeath), the loss of mussels caused significant changes in community structure, and limpets played a less important role. At Harlyn, fucoid algae were abundant throughout the year. There were fewer algae at Polzeath, and cover was dominated by the summer bloom of ephemerals. At Harlyn, the limpets played a major role in controlling algae, but their effects were mediated by the presence of mussels. Other grazers were not able to fulfil their role. At Polzeath, mussels were far more important, and ephemeral algae grew on them regardless of the presence or loss of limpets. These findings emphasise the need to assess spatial and temporal variation in the effects of biodiversity loss and the importance of interactive effects of loss of multiple species from different functional groups.