Differential recruitment of introduced Pacific oysters and native mussels at the North Sea coast: coexistence possible?

Pacific oysters (Crassostrea gigas Thunberg 1793) have been introduced into the Wadden Sea (North Sea), where they settle on native mussel beds (Mytilus edulis L.), which represent the only extensive insular hard substrata in this soft-sediment environment. As abundances of C. gigas rose, some mussel beds became increasingly overgrown with oysters, whereas others did not. Field experiments revealed that recruitment of C. gigas was higher in the lower intertidal than in the upper subtidal zone, that it was higher on conspecifics than on mussels, and that it was not affected by barnacle epigrowth except when settling on mussels. Mussel recruitment is known from inter- and subtidal zones. It occurred equally on oyster and mussel shells but showed a clear preference for barnacle epigrowth over clean shells. Assuming that settlement and recruitment are key processes for species abundances on the North Sea coast, it is predicted that the positive feedback in oyster settlement will lead to rapid reef formation of this invader at the expense of mussel beds. Mussels, however, may escape competitive exclusion by settling between or on the larger oysters especially when barnacles are abundant. Experimental patches with mussels were more often covered by fucoid algae (Fucus vesiculosus forma mytili Nienburg) than patches with oysters, and oyster recruitment was poor underneath such algal canopies. Thus, fucoids may provide the native mussels with a refuge from the invading oysters and the two bivalves may coexist, provided food is not limiting.     

Marine invasive aliens on South African shores: implications for community structure and tropillc functioning

Up to 12 marine mollusc and three crab species are thought to have been deliberately or accidentally introduced to South Africa. Of these, only the Mediterranean mussel Mytilus galloprovincialis and the European shore crab Carcinus meanas have become invasive. M. galloprovincialis, probably introduced in the late 1970s, is already the dominant mussel throughout the Cape west coast. As a result, indigenous intertidal mussels Aulacomya ater have been largely displaced, mussel standing stock has increased and the upshore limit of mussel beds has become considerably elevated. Implications include competitive exclusion of large limpets but enhanced recruitment of juvenile limpets, increased habitat availability for mussel infauna, reduction of habitat for algal infauna and enhanced food availability for predators, particularly terrestrial species. C. maenas was first recorded from Table Bay Docks in 1983 and currently ranges from Camps Bay to Saldanha Bay. Although it is a voracious predator, it appears unable to colonize wave-swept shores, so is unlikely to displace indigenous crabs, or to severely impact prey species on the open coast. Valuable conservation areas and mariculture sites in sheltered lagoonal areas are, however, threatened by it.     

Comparing quality of estuarine and nearshore intertidal habitats for Carcinus maenas

Estuarine and nearshore marine areas are vital habitats for several fish and benthic invertebrates. The shore crab Carcinus maenas (Crustacea: Brachyura: Portunidae) inhabits a variety of coastal, estuarine and lagoon habitats. At low tide, habitat structural complexity may be most important for crabs in the intertidal, providing refuge from predation and desiccation. The quality of different vegetated and nonvegetated estuarine and rocky shore habitats in SW Portugal and SW England was evaluated for intertidal C. maenas populations. We estimated population density, size–structure, and potential growth (RNA/DNA ratios) to investigate habitat quality. Vegetated estuarine habitats supported higher crab densities, than nonvegetated estuarine and rocky shore habitats. Investigation of population size–structure revealed that all habitats seem important recruitment and nursery areas although estuarine habitats in SW Portugal appeared to support higher densities of new recruits than equivalent habitats in SW England. Significant variation was found in RNA/DNA ratios among habitats. Ratios were highest in the rocky shore suggesting a high quality habitat where growth potential is high. We speculate that competition from other top-predators (Pachygrapsus spp.) rather than low habitat quality may limit the occurrence of C. maenas in intertidal rocky shore habitats in SW Portugal. In estuarine environments RNA/DNA ratios were significantly higher in the vegetated than in the nonvegetated estuarine habitats in SW Portugal but not in SW England, suggesting geographic differences in the extent to which highly structure habitats represent high quality. Our results challenge the current paradigm that structured habitats are necessarily those of higher quality for C. maenas.     

Adult-larval interactions: Predictions,mussels and cocoons

The purpose of this investigation was to test two hypotheses regarding adultlarval interactions in dense assemblages of suspension feeders. Benthic communities outside and inside a dense population of the mussel Mytilus edulis were sampled at a New England soft-bottom intertidal flat. The hypothesis that recruitment of M. edulis is inhibited by the presence of densely packed adults was not supported. Mytilus edulis population size-class structure revealed no dominance of large individuals, indicating that older cohorts were not able to prevent subsequent mussel recruitment. The hypothesis that M. edulis has a negative effect on the recruitment of other infaunal species was not supported. Infaunal density was significantly higher inside the mussel bed than outside. Oligochaete worms were the most common organisms at the study site and had higher absolute and relative abundances inside the mussel bed. The production of cocoons and dispersal of non-larval benthic offspring may confer upon the oligochaetes relative immunity from M. edulis ingestion and suffocation in mussel feces and pseudofeces. These results may be extended to other infaunal species, including densely packed suspension feeders themselves, which form cocoons, fragment asexually, brood, or disperse as juveniles and adults.     

Small is profitable: No support for the optimal foraging theory in sea stars Asterias rubens foraging on the blue edible mussel Mytilus edulis

Doubt has been shed recently on the most popular optimal foraging theory stating that predators should maximize prey profitability, i.e., select that prey item that contains the highest energy content per handling time. We hypothesized that sea stars do not forage on blue mussels according to the classical optimal foraging theory but are actively avoiding damage that may be caused by e.g. capture of foraging on too-strong mussel shells, hence the sea stars will have a stronger preference for mussels that are smaller than the most profitable ones. Here we present experimental evidence of the sea star Asterias rubens as a predator that indeed chooses much smaller blue mussels Mytilus edulis to forage on than the most profitable ones. Hence this study does not support the optimal foraging theory. There may be other constraints involved in foraging than just optimizing energy intake, for example predators may also be concerned with preventing potential loss or damage of their foraging instruments.     

Predators on rocky shores in the northern Atlantic: Can the results of local experiments be generalized on a geographical scale?

Experimental manipulations of invertebrate prey and predators on rocky shores have been done by many authors. In the northern Atlantic the predators involved are usually the green crab Carcinus maenas and/or the dogwhelk Nucella lapillus, and the prey species studied are acorn barnacles (balanid Cirripedia), mussels (Mytilus spp.) and winkles (Littorina spp.). Usually the predators are found to have a regulating “top–down” effect on the prey species. In Iceland the acorn barnacle Semibalanus balanoides, the blue mussel Mytilus edulis and the flat periwinkle Littorina obtusata (including to some extent Littorina mariae) are found on rocky shores all around Iceland in what would seem to be near-optimal physical conditions. The predators Carcinus maenas and Nucella lapillus, on the other hand, are relatively southern species that do not thrive on the colder coasts of Iceland. Thus general surveys of different coasts of Iceland would seem to offer opportunities to see whether the results of local experiments can be discerned on a geographical scale (hundreds of km). The roughly 4900 km of the rocky coastline in Iceland was in this study subdivided into four regions, I–IV, according to the commonness or presence of the two predators. With the hope of reducing compounding factors the surveys were confined to sheltered or semi-sheltered fucoid shores, which were further divided into Ascophyllum (more sheltered) and Fucus vesiculosus (less sheltered) shores. Estuaries or other low-salinity environments were avoided. The study was based on 761 stations distributed around the rocky coastline on these two types of shores. The results for barnacles and mussels, being generally more abundant in regions were predators were scarce or absent, and being less common in Ascophyllum than F. vesiculosus shores in contrast to the predatory dogwelks, were in large measure in accord with predictions from experiments indicating “top–down” regulation. The results for the periwinkles, however, suggested that “bottom–up” regulation might be of greater importance, as their density was closely linked to the biomass of macroalgae, while the abundance of predators appeared to have little effect. The periwinkle, although certainly eaten, may not be a preferred prey of the two predators in question in Iceland. This paper forms a link between experimental and observational study approaches.     

Fate and effects of a heavy fuel oil spill on a Georgia salt marsh

Addition of a heavy oil to a Spartina salt marsh in the autumn resulted in high concentrations of polycyclic aromatic hydrocarbons in sediment and benthic animals. The highest concentrations of phenanthrene, chrysene and fluoranthene in the sediment were 112, 105 and 75 ng/g sediment, respectively. These concentrations rapidly decreased during the 20 week period following the spill. The times for these hydrocarbons to decrease to 50% of their highest values, i.e. half-life, were approximately 100, 70 and 30 days in sediment, mussels and oysters, respectively. Benthic macrofauna species showed three responses to oil addition which included no change, an increase, or a decrease in the population. No changes were noted in populations of fiddler crabs (Uca pugnax), oysters (Crassostrea virginica), and mussels (Modiolus demissus). Mud snails (Nassarius obsoleta) increased in density after the spill due to immigration of adult snails from untreated areas to scavenge on animals killed by the oil. Many of the adult periwinkles (Littorina irrorata) were killed by the oil. In the spring, juvenile periwinkles recolonised to oiled areas as a result of larvae settling.     

Biotic consequences of a shift in invertebrate ecosystem engineers: Invasion of New Zealand rocky shores by a zone‐forming ascidian

The pervasive effects of invasive ecosystem engineers, that is those species that modify their environment, are well documented, but rarely have the broader impacts of one foundation invertebrate species being replaced by another been examined. In New Zealand, green‐lipped mussels, Perna canaliculus, commonly dominate wave‐exposed rocky shores. The recent appearance of an invasive ecosystem engineer, the ascidian Pyura doppelgangera, at the very northern tip of New Zealand now threatens to exclude these bivalves from this habitat. Here, we report major shifts in assemblages associated with the invader and chronicle its continued spread. We examined epibiota associated with clumps of mussels and clumps of Pyura from two rocky shore habitats—pools and emergent substrata at two locations. We detected some differences in species richness in biota associated with the two foundation species, but faunal abundance only differed between the locations. These minor changes were dwarfed by the shift in species composition within clumps of each foundation species. Molluscs, particularly gastropods, and crustaceans dominated the assemblage within mussels. In contrast, tubicolous polychaetes dominated the fauna associated with the ascidian. Sessile epifauna, notably barnacles and calcareous tube‐dwelling polychaetes, were common on mussels, but never encountered on the ascidian. Multivariate analysis revealed marked dissimilarity (>80%) between the characteristic mussel and ascidian faunas with virtually no overlap. This biotic shift overshadowed any differences between habitats and locations. The broader implications of these faunal shifts for local and regional patterns of biodiversity, as well as ecosystem function, remain unclear, but deserve further attention.     

Negative effects of blue mussel (Mytilus edulis) presence in eelgrass (Zostera marina) beds in Flensborg fjord,Denmark

The effect of blue mussel (Mytilus edulis) presence in eelgrass (Zostera marina) beds was studied from June 2004 to July 2005 in Flensborg fjord, Denmark. The field experiments were conducted at two stations, one with only Z. marina (Eelgrass station) present and one where M. edulis were present in the Z. marina beds (Mixed station). Zostera marina parameters were measured (growth of leaves, shoot density, leaf length, and nutrient content) in combination with epiphyte cover and sediment parameters (sulphate reduction rates, sediment nutrient fluxes, organic content, C, N and P content) to examine possible positive and negative effects of the mussels on eelgrass performance. The fluxes of ammonium from the sediments were stimulated at all sampling dates at the Mixed station, and possibly stimulated epiphyte growth at this station. Further ¹⁵N signals in epiphytes from the Mixed station suggested that excretion products from the mussels were important nitrogen sources at this station. Sulphate reduction rates were enhanced at the Mixed station and also sediment sulphide concentrations increased under mussel influence, which may have resulted in sulphide toxicity and decreased growth of Z. marina at this station. The study indicates that for Z. marina beds in Flensborg Fjord the effects of M. edulis in seagrass beds are primarily negative, and raises the question whether this leads to negative effects on the stability and expansion of Z. marina beds.     

Enhancement of polycyclic aromatic hydrocarbons in estuarine invertebrates by surface runoff at a decommissioned military fuel depot

Accumulation of polycyclic aromatic hydrocarbons (PAHs) was determined in blue mussels (Mytilus spp.) and shore crabs (Hemigrapsus sp.) at a recently closed military fuel depot in central San Francisco Bay, California. In April 1996, during a period of above average precipitation, specimens were collected at the depot, near the depot, and at sites 10 and 20 km south of the depot. Four weeks after the rains ended, blue mussels were again collected at the depot, and at two additional sites in the central Bay region. In April, total PAHs in mussels from the depot were significantly higher only than that in mussels collected 20 km from the depot; however, seven specific, substituted PAHs were higher at the depot than at all other sites. In June, only two of the 38 PAHs common in mussels in April were detected at the depot; these concentrations were comparable to ambient concentrations in mussels at the Bay. It seemed that bioavailability of PAHs at the depot was enhanced by rainfall, probably due to the mobilization of PAHs via groundwater into the Bay. Concentrations in mussels from chronically contaminated sites were about five times higher than mussels collected from the depot. Low PAH concentrations were detected in shore crabs near the depot, and the highest levels were not associated with the depot. Observed PAH concentrations are discussed in relation to upper trophic organisms.     

Further refutation of the primary-secondary settlement hypothesis for the brown mussel Perna perna

The primary-secondary settlement hypothesis, that mussels first settle in algae and then move to mussel beds, was rejected as the only mode of recruitment for the brown mussel Perna perna in a previous study at one location over one year on the south coast of South Africa. We investigated recruitment of P. perna into mussel beds and algal turfs over a five-year period (1995–2000) at 3–4 sites at each of four locations on the South and East coasts. Attachment to the two substrata was significantly temporally synchronous at two of the four locations. The disparity among locations could be due to differences in the algal species present or to differences in the importance of turf algae as a facilitator of recruitment. At Zululand, on the East Coast, attachment to mussel beds and algae was measured monthly and found not to be temporally synchronous. At this location, primary settlement into algae was significantly correlated with an increase in the density of juveniles in the mussel beds 2–3 months later, indicating that turf algae can act as irregular but important facultative facilitators of mussel recruitment, supplementing direct input into the mussel beds. However, as amounts of settlers and recruits in mussel beds exceeded those found in algal turfs most years and places, the primary-secondary settlement hypothesis is inapplicable to P. perna under most circumstances.     

The interactive roles of predation and tidal elevation in structuring populations of the edible cockle,Cerastoderma edule

The size and age structure of the Cerastoderma edule population at Traeth Melynog, North Wales, varies dramatically with tidal level. In areas low on the shore up to 96% of cockle spat fail to survive their first summer, but mortality rate subsequently declines and remains at a low level. By contrast cockles high on the shore suffer moderate mortality during their first year (47%), but increasing rates thereafter. High-shore populations consequently consist mainly of smaller (younger) individuals and low-shore ones of a transient spatfall, plus a few larger and older individuals. The potential role of shore crabs, Carcinus maenas, and oystercatchers, Haematopus ostralegus in the determination of these patterns is assessed. Shore crabs move up into the intertidal to feed with each flood tide from about April to December. They selectively consume cockles < 15 mm in length, taking an estimated 236 × 10³ cockles, or 2432 g dry flesh year⁻¹ per linear meter of shoreline, mostly from lower shore levels. Oystercatchers are present only during winter and preferentially select large cockles of at least 20 mm length. They are estimated to remove 9 × 10³ cockles, or 1204 g dry flesh year⁻¹ per linear meter of shoreline, most of this from mid- and high-shore levels. These results indicate that shore crabs are far more important predators than previously suspected, taking 25 × the numbers and 2 × the biomass cosumed by oystercatchers. Predation also appears to be the key factor controlling the structure of the C. edule population. Crabs consume almost all the cockles settling low on the shore during their first summer, but avoid older individuals, which subsequently survive and grow well under low levels of oystercatcher predation. On the high shore, crabs are unimportant and the cockles survive well as they slowly grow into the size range attractive to oystercatchers. Thereafter they suffer increasingly severe winter mortality and are soon eliminated.     

The effect of shore location on biomarker expression in wild Mytilus spp. and its comparison with long line cultivated mussels

Biomarkers are a common tool in the assessment of potential effects of contaminants in aquatic organisms. In order to identify the effects of anthropogenic pollution it is essential to identify background levels and to know the range of natural variability in the biomarker response. In this study, we examined various biomarkers of stress (glutathione S-transferase and metallothionein), damage (lipid peroxidation (LPO) and DNA damage (DNA)) and reproduction (vitellin-like proteins) in marine mussels (Mytilus spp.) from four locations along a vertical transect from high to low shore and compared them with cultivated long line mussels. High shore and cultivated mussels showed significantly higher LPO and DNA damage expression than the low shore mussels indicating a level of oxidative stress resulting from mussel location. Significant effects in physiological endpoints were also found. This study highlights the need to consider the diversity of natural environmental stress factors when using biomarkers in environmental assessment.     

Long-term changes (1979–1994) in two coastal benthic communities (English Channel): analysis of structural developments

The long-term variations of two coastal benthic communities (Eastern English Channel, France) were studied between 1979 and 1994 by means of factor correspondence analysis (FCA), trophic structure and rank-frequency diagrams (RFD). FCA identified periods of relative homogeneity and periods of strong variation in the structure of the two communities. The first community, established on heterogeneous sand, showed a multi-annual cycle linked to grain-size variations in the sediment. As passive supplies of organic matter were not important and suspension feeders could hardly live in these conditions, the community remained poor and stable and was dominated by deposit-feeders and carnivores. The other one, a rich and diverse mussel (Mytilus edulis) bed, showed great variations in the abundance of the main species which generated the enrichment of the community by its suspension-feeding behaviour (biodeposition). Beyond these variations, the community remained stable until 1990 as no continuous trend or permanent change could be seen. Since 1990, no recruitment of M. edulis occurred and the relative dominance of suspension feeders, as well as the number of species, strongly decreased. During the following years, several species of tunicates (sessile suspension feeders with gregarious recruitment) recruited and partly took the ecological niche of M. edulis but they did not induce an increase of biodeposition and the previous richness was not restored. After a strong and rapid shift in the structure, a new equilibrium has been established within the community.     

A chemical investigation of the transport and fate of petroleum hydrocarbons in littoral and benthic environments: The TSESIS oil spill

The fate of saturated and aromatic hydrocarbons discharged into the coastal Baltic Sea environment from the TSESIS oil spill has been studied in the acute and postacute (one year) phases of the spill. Periodic samples of Mytilus edulis (mussels) from eight littoral zone stations and Macoma balthica from nine soft bottom stations were obtained as well as sediment trap samples and surface sediment samples. Glass capillary gas chromatography and gas chromatographic mass spectrometry were used as the analytical tools to determine saturated and aromatic hydrocarbon composition and concentrations in these samples.Sediment trap samples indicated that sizable quantities of chemically and microbially weathered oil were sedimented, and available for benthic uptake shortly after the spill. After initial uptake of sedimented oil (500 to 1000 μ/g dry weight), Macoma populations appear to have begun slow depuration through the first winter after the spill, but TSESIS oil was again introduced to the benthic stations studied during the following summer. Mytilus populations in the region were severely impacted by the oil. Initial depuration of spilled oil during the first month was rapid and nearly complete at all but the most heavily impacted stations one year after the spill. The post-spill depuration of assimilated hydrocarbons was characterised by a relative retention of alkylated dibenzothiophenes and alkylated phenanthrenes compared to their unsubstituted parent compounds and compared with the entire homologous naphthalene series.These data suggest that petroleum hydrocarbons from the TSESIS spill have become a chronic source of degraded saturated and aromatic hydrocarbons to the soft bottom benthic communities. Petroleum hydrocarbons in the benthic environment from this spill appear to reside in the difficulty sampled and mobile flucculent layer at the sediment/water interface and may affect epifaunal communities for an extended period of time.     

Different crab species influence feeding of the snail Nucella lapillus through trait‐mediated indirect interactions

Indirect interactions are among the many important factors that influence the community structure of the rocky intertidal zone. Trait‐mediated indirect interactions, in which the presence of a predator or competitor can influence the relationship between two other species, have emerged as vital for understanding community dynamics. This study examined the effect of different crab species on the feeding habits of an intertidal snail, Nucella lapillus. Crab species were defined as being sympatric predatory (Carcinus maenas and Cancer irroratus), sympatric non‐predatory (Uca pugnax and Pagurus longicarpus), or allopatric predatory (Mithrax sculptus and Percnon gibbesi). Nucella lapillus were potentially exposed to risk cues from each of the crab species. Crabs were kept in perforated boxes, which allowed any chemical cues to be emitted but prevented direct contact. Nucella lapillus had significantly lower feeding rates in the presence of sympatric predatory crab species than N. lapillus exposed to either sympatric non‐predatory crabs or allopatric predatory crabs. There was no difference in feeding rate between N. lapillus exposed to the sympatric non‐predatory crabs and to the allopatric predatory crabs. Nucella lapillus in the presence of sympatric predatory crabs had a feeding rate of only 0.07 barnacles per snail per day, whereas N. lapillus housed with non‐predatory crabs and allopatric predatory crabs had rates of 0.11 and 0.12, respectively, suggesting that N. lapillus alter their behavior in response to chemical risk cues from local predators. These results suggest that the ability to detect and respond to risk cues is a selectively evolved trait.     

Accumulation of Zn,Cu and Cd by crabs and barnacles

The crab Carcinus maenas (L.) and the barnacle Elminius modestus Darwin were exposed to a range of dissolved concentrations of Zn, Cu and Cd for 21 days in artificial seawater. Accumulation of Zn and Cu by crabs has been interpreted in terms of the presence of a regulation mechanism to maintain constant body concentrations (83·2 ± 19·4 μg Zn g^?1 dry wt.; 39·8 ± 9·8 μg Cu g^?1 dry wt.) under varying external dissolved metal levels, until a threshold dissolved metal concentration (c. 400 μg Zn l^?1; c. 170 μg Cu l^?1) beyond which net accumulation of metal begins. Cadium appears to be accumulated by C. maenas at all exposures with no evidence for regulation of body cadmium concentrations. Exposure of E. modestus to Zn, Cu or Cd caused net accumulation of the respective metal in the bodies of the barnacles, with no evidence for regulation of body metal concentrations.     

Teneurs en benzo 3,4 pyrène chez Mytilus edulis L. de l'estuaire et du golfe du Saint-Laurent

Blue mussels (Mytilus edulis L.) taken from 27 stations along the estuary and gulf of St. Lawrence were analyzed for their content in 3,4-benzopyrene. Most of the concentrations are below the detection limit for the analytical method, i.e. 0·15 μg kg^?1 (dry weight) and probably reflect the natural baseline level. However, mussels from two stations located at the mouth of Saguenay Fjord exhibit very high contents: 24 and 28·5μg kg^?1 (dry weight). A mussel transplantation experiment has shown that the entire Fjord is contaminated by anthropogenic 3,4-benzopyrene.     

The effect of life-history variation on the population size structure of a rocky intertidal snail (Littorina sitkana)

On wave-sheltered shores of the northeastern Pacific, the population size structure of Littorina sitkana varies with intertidal height, as larger snails are mostly found only in the upper intertidal. This pattern has been attributed to high predation rates by crabs (and perhaps fish) on large snails inhabiting low-intertidal areas; i.e., large snails are presumed to be rare there simply because predators kill them. In this study we investigate the hypothesis that predation contributes to the shore-level size gradient displayed by L. sitkana by selecting for (or inducing) earlier sexual maturation and reduced somatic growth in low-shore snails relative to high-shore individuals.In the first part of our study, we carried out laboratory dissections, field experiments (mark-release-recapture and caging), and field surveys on a wave-protected shore in Bamfield Inlet, Barkley Sound (British Columbia, Canada). The principal results were: (1) adult survivorship was greater at higher, than at lower, intertidal level, (2) snails displayed a preference for their shore level of origin, (3) immature adults from the high intertidal displayed greater rates of somatic growth relative to immature adults from the low intertidal, and (4) low-shore snails matured at a smaller size than high-shore individuals. In the second part of the study, a large-scale survey showed intra-specific variation in size at sexual maturity (point 4 above) to be relatively consistent over time (winter of 1999 and 2001 for snails from our main study site) and space (13 different sites in winter 2001), although the magnitude of these differences varied greatly from shore to shore.Our results indicate that L. sitkana individuals inhabiting upper and lower parts of their intertidal range allocate resources differently to somatic and gonadal growth, an intra-specific difference that is best interpreted as a response to spatial and size-dependent variation in predation pressure. Taken together, results of this and other recent studies indicate that phenotypic responses to contrasting selection pressures operating in upper- and lower-intertidal areas contribute to the intertidal size gradient of L. sitkana. We believe that greater consideration of evolutionary processes in ecological studies will lead to a more complete understanding of the mechanisms responsible for structuring marine coastal communities.     

Effect of an exotic prey on the feeding pattern of a predatory snail

The mussel Brachidontes pharaonis, which invaded the Mediterranean from the Red Sea about 120 years ago, has recently become abundant in many midlittoral and some infralittoral rocky habitats along the Israeli rocky shore. We investigated the influence of B. pharaonis, as a novel prey, on the foraging patterns of the large whelk Stramonita haemastoma in the field, and examined food preferences in the laboratory. S. haemastoma has shifted from indigenous species to feeding on the novel mussel when abundant. The whelk prefers to prey upon the invasive mussel over all indigenous species offered (e.g. barnacles and mussels), probably due to its larger size. In the midlittoral zone, the foraging activity of S. haemastoma is considerably low even where refuges are readily available (incisioned-rocks) and food density is high (mainly B. pharaonis). Higher proportions of whelks are actively foraging in the infralittoral zone but usually on smaller prey, mostly barnacles. We suggest that this differential foraging activity in the two zones is related to the degree of exposure to wave action. The midlittoral is inherently more exposed to wave action than the infralittoral, where sea conditions are more benign and the whelks may enjoy longer activity periods.