Sea otter foraging on deep-burrowing bivalves in a California coastal lagoon

Sea otter, Enhydra lutris, predation had no detectable effect on abundance and size distribution of deep-burrowing bivalve prey in the Elkhorn Slough, California, USA. Up to 23 otters were present for 6 mo of the study period (March 1984 through April 1985). This is in contrast to previous studies of sea otter predation, especially on the shallow-burrowing Pismo clam Tivela stultorum, which can be found along the wave-exposed coast near the slough. The deep-burrowing clams Tresus nuttallii and Saxidomus nuttalli made up 61% of the prey taken in the slough, and are more difficult for otters to excavate than Pismo clams. The occurrence of foraging otters was highest in an area where the two bivalve prey were extremely abundant (18 individuals m^–2). However, the otters did not selectively prey on the largest clams available within the study sight, but foraged preferentially in a patch of smaller individuals where bivalve burrow depth was restricted by the presence of a dense clay layer. This foraging strategy maximized the amount of prey biomass obtained per unit volume of sediment excavated. Our findings suggest that in soft-sediment habitats deep-burrowing bivalves may be more resistant to otter predation than shallower burrowers.     

Adaptations of subtropical Venus clams to predation and desiccation: endurance of<Emphasis Type="Italic"> Gafrarium tumidum</Emphasis> and avoidance of<Emphasis Type="Italic"> Ruditapes variegatus</Emphasis>

Intertidal endobenthic bivalves are often dislodged from sediments by hydrodynamic forces. As a result, they encounter the dangers of predation and desiccation, which are generally harsh near the sediment surface. To cope with such dangers, the bivalves possibly possess: (1) a strong body to endure predation and desiccation stress, (2) quick mobility to avoid the stresses, or (3) a high growth rate for attaining a size refuge. The present study examined which of these modes are adopted by the subtropical cobbled-shore Venus clams Gafrarium tumidum (Röding, 1798) and Ruditapes variegatus (Sowerby, 1852), revealing the following interspecific differences. (1) G. tumidum survived better than R. variegatus did in harsh experimental conditions, namely: the experimental cages exposed to predation and desiccation on a cobbled shore; a laboratory aquarium with a predatory crab Scylla serrata; and ovens with high temperatures (27°C and 34°C). (2) R. variegatus was more mobile than G. tumidum was, digging into the sediment on a cobbled shore more rapidly at both high and low tides. (3) The two species with shell lengths 20–30 mm showed similar growth rates (median: –0.2 to 44.5 m day^–1) in seasonal mark–recapture surveys over 2 years. Overall, to cope with the dangers of predation and desiccation G. tumidum appears to have a strong body, while R. variegatus displays rapid mobility, and neither species seems to attain a size refuge through rapid growth. Such species-specific modes are discussed in relation to the interspecific differences found in shell morphology.Communicated by T. Ikeda, Hakodate     

Further insight on carapid—holothuroid relationships

Carapidae (or pearlfish) are eel-like fishes that live inside different invertebrates, such as holothurians, sea stars, or bivalves. Those of the genus Carapus are commensal and use their host as a shelter, while Encheliophis species are parasitic and eat the hosts gonads. In areas where they live in sympatry, C. boraborensis, C. homei, C. mourlani and E. gracilis are able to inhabit the same host species. Infestation is considered as monospecific when several conspecifics are observed in the same host. However, many aspects of this particular relation remain obscure, e.g. communication between carapids and the defence systems of the different protagonists (carapids and hosts). Experiments have been conducted in the field and laboratory to investigate several aspects of the carapids relationships with their hosts. Sampling carried out in the Bay of Opunohu (Moorea, French Polynesia) determined the sex ratio of C. boraborensis (3:1) and C. homei (1:1) and their distribution rate within different Echinodermata. Our study showed that neither species was capable of determining whether a heterospecific already occupied a sea cucumber or not. They were, however, able to locate the sea cucumbers cloaca, due to the excurrent resulting from respiration. The sea cucumbers defence system (Cuverian tubules) minimises predator attacks, but is not effective against carapid intrusion. The Carapidae defence system is twofold. Due to a passive system related to the sea cucumbers low cloacal position, the Cuverian tubules are not expelled when fish enter the cloaca. Moreover, carapids resist sea cucumber toxins better than other reef fish. Their increased resistance might be related to their gills rather than to their mucus coating; however, the latter may assist the fish in resisting the sticky substances emitted by the Cuverian tubules.Communicated by S.A. Poulet, Roscoff     

Diet and food preferences of the adult horseshoe crab Limulus polyphemus in Delaware Bay,New Jersey,USA

Adult horseshoe crabs Limulus polyphemus (L.) feed on a wide variety of infaunal and epifaunal invertebrates during their spring spawning migration in Delaware Bay, New Jersey, USA. Comparison of the gut contents with estimates of available prey showed that the most abundant potential prey item, the bivalve Gemma gemma, was avoided. The thinner shelled but comparatively scarce clam Mulinia lateralis was a preferred prey item. In the laboratory, crabs fed on G. gemma when it was the only available item but not when M. lateralis or soft-shell clams, Mya arenaria, were offered in conjunction. Large M. lateralis (>10mm) were preferred to small M. lateralis; there was no discrimination between M. lateralis and M. arenaria of the same size. Male and female horseshoe crabs had similar gut contents and laboratory feeding preferences, despite the fact that females are larger than males. Crabs spawning later in the summer contained more food than did crabs collected at the peak of spawning activity.     

Anatomical and experimental evidence for particulate feeding in<Emphasis Type="Italic"> Lucinoma aequizonata</Emphasis> and<Emphasis Type="Italic"> Parvilucina tenuisculpta</Emphasis> (Bivalvia: Lucinidae) from the Santa Barbara Basin

Previous nutritional models for adults of the lucinid bivalve Lucinoma aequizonata contend that symbiotic chemoautotrophic bacteria provide most of the organic carbon for the host. The existence of this symbiosis, coupled with the hosts distinctive anatomical features, shaped the impression that particulate feeding was not a significant part of L. aequizonata nutrition. Here, we use several techniques to show that particulate feeding is a consistent and important part of the L. aequizonata nutritional strategy. Histological and scanning electron microscopy observations reveal that the gills of L. aequizonata, like those of the lucinid Parvilucina tenuisculpta, have functional mucociliary epithelia, able to transport captured particles to the mouth. Observations of gut content and radiolabeled feeding experiments indicate that L. aequizonata does ingest and assimilate carbon from particulate organic matter. Furthermore, molecular identification of a broad spectrum of organisms in the guts of native adult specimens demonstrates that L. aequizonata is non-selective when ingesting organic material, and has a mixotrophic diet.Electronic Supplementary Material Supplementary material is available in the online version of this article at Communicated by P.W. Sammarco, Chauvin     

Stomach contents and size-frequency distributions of two coexisting sea star species,Astropecten aranciacus and A. bispinosus,with reference to competition

Stomach contents of the sea star species Astropecten aranciacus (L.) and A. bispinosus (Otto) consisted of roughly the same sort of prey, mostly bivalves and irregular sea urchins. However, the two predators ingested prey of different size classes. The stomach contents of A. bispinosus reflected the size distribution of the benthic macrofauna, while larger prey was more frequent in the stomach contents of A. aranciacus. As the density of prey in the investigated area is low, A. aranciacus and A. bispinosus are likely to compete for food. Possible mechanisms reducing the effect of competition are discussed.     

Dietary constraints on reproductive periodicity in two sympatric deep-sea astropectinid seastars

Examination of the diet of two sympatric species of seastar, Bathybiaster vexillifer and Plutonaster bifrons from 2200 m depth in the Rockall Trough, NE Atlantic Ocean (57°18'N; 10°28'W), suggested that diet may determine the different reproductive patterns found between these two species. In the non-seasonally breeding B. vexillifier, the diet showed a high Shannon-Wiener prey diversity index, the dominant prey being the irregular echinoid Hemiaster expergitus together with a variety of prosobranch gastropods and protobranch bivalves. By contrast, the prey diversity in the seasonally breeding P. bifrons was significantly lower than that of B. vexillifer. In addition, organic carbon content in the sediment residue in the stomachs of P. bifrons displayed a seasonal cycle, while no such seasonality was detected in B. vexillifer. The stomachs of P. bifrons also contained a higher proportion of scavenged material, including the seasonally available remains of mesopelagic blue whiting. These data, together with Bathysnap (time-lapse camera) observations of feeding behaviour in both species, suggest that B. vexillifer is a predator feeding deep in the sediment, whereas P. bifrons feeds close to the sediment surface where it is affected by the seasonal availability of phytodetritus and fish carcasses.     

Diel vertical migration of the marine copepod<Emphasis Type="Italic"> Calanopia americana</Emphasis>. I. Twilight DVM and its relationship to the diel light cycle

Marine copepods commonly exhibit vertical movements in the water column over the diel cycle, termed diel vertical migration (DVM), with the most common pattern being an ascent in the water column to minimum depth around sunset and descent to maximum depth around sunrise. The present study characterized the DVM pattern of the pontellid copepod Calanopia americana Dahl in the Newport River estuary (North Carolina, USA, in July 2003). The estuary is shallow and well-mixed, and the study site (34°43N; 76°40W), 1.5 km inside the estuary entrance, is unusual in lying within a gyre where tidal currents are always in the seaward direction. Changes in C. americana vertical abundance were related to spectrally relevant changes in light throughout the diel cycle. Simultaneous measurements of light and zooplankton abundance near the surface (0.5 m depth) and near the bottom (0.5 m above bottom) were made over one 4-h period and two 3-day periods during different phases of the tide. These observations suggest that C. americana undertook twilight DVM in the Newport River estuary; an ascent to the surface occurred at sunset, followed by a descent to near the bottom around midnight, with a second ascent to the surface and then descent to near bottom at sunrise. DVM in C. americana was independent of the tidal cycle, with the initial ascent in the water column at sunset possibly associated with relative rates of irradiance change. Copepod vertical movements were consistent with a night-active endogenous rhythm, and appeared independent of the abundance of predatory chaetognaths, Sagitta spp. In DVM studies with migrators like C. americana that are broadly sensitive to visible wavelengths of light, measuring photosynthetically active radiation may be a reasonable alternative to measuring light in a spectrally relevant photometric unit.Electronic Supplementary Material Supplementary material is available in the online version of this article at http://dx.doi.org/10.1007/s00227-005-1569-x.Communicated by J.P. Grassle, New Brunswick     

The types of escape reactions elicited in the scallopPecten maximus by selected sea-star species

Pecten maximus (L.), when stimulated by contact with various sea-star species, displays several fairly distinct types of responses. These range from valve closure, through jumping to the well-known swimming escape reaction. These responses are described in detail. The most violent responses are evoked by contact with the predatory sea starsAsterias rubens,Astropecten irregularis, andMarthasterias glacialis, whereas non-predatory sea stars may evoke minor reactions or none at all. It is suggested that, in this instance, the adaptive value of the various types of responses lies not only in enabling the scallop to escape from predators, but also in preventing needless and even hazardous movement away from the refuge of its prepared depression in the substratum when confronted by the less harmful species of sea stars. Crude chemical extracts of all sea stars always evoked escape reactions, but only those ofAsterias rubens andAstropecten irregularis caused the full swimming response.     

Ontogeny of diet changes in a tropical benthic carnivorous fish, Parupeneus barberinus (Mullidae): relationship between foraging behaviour, habitat use, jaw size, and prey selection

Parupeneus barberinus forages on benthic invertebrates using a wide range of foraging modes, including vigorous digging in the substratum, resulting in considerable disturbance to the benthos. Polychaetes were the most important prey item for all size classes, but fishes less than 120 mm total length consumed more small ostracods and nematodes than did larger fishes. Fishes greater than 120 mm total length consumed mostly bivalves, and fishes over 240 mm total length consumed mostly bivalves and crabs. A morphological examination of the feeding apparatus suggested that the size of important prey items consumed was determined by gape height and jaw width. Prey available to different size classes of fishes was determined by combining information on microhabitat use, foraging behaviours, and prey volumes in the substratum. Small fishes spent more time foraging on the reef flat and slope, compared with larger fishes that foraged mostly on the reef edge and base. In addition smaller fishes foraged mostly in the upper 2 cm of sediment, whereas larger fishes often foraged to depths of 10 cm. Selection ratios showed that different size classes of fishes selectively extracted different prey items from the substratum. Small fishes showed a preference for ostracods whereas large fishes selected for bivalves and crabs. Although polychaetes were the dominant prey item for all size classes, they were consistently selected against.     

The “piece-by-piece predation” of<Emphasis Type="Italic"> Eteone longa</Emphasis> on<Emphasis Type="Italic"> Scolelepis squamata</Emphasis> (Polychaetes )—traces on the sediment documenting chase,defence and mutilation

On many sea shores of the Niedersachsen coast, the polychaete Scolelepis squamata is the dominant animal species living in the sediment of exposed beaches.The population of the predatory species Eteone longa, with a main distribution in more sheltered intertidal and subtidal habitats, has a certain overlap with S. squamata. In these restricted areas, S. squamata suffers from a permanent pursuit by E. longa during low tide. Field and laboratory studies have revealed that this predator-prey relationship follows a distinct behavioural pattern and is an exploitation of surviving animals: the predator does not ingest the whole prey individual but only feeds on regenerable parts of the body. Both species draw characteristic tracks on the sediment surface that illustrate the phases of the chase, the attempts at defence and the mutilation of the prey species.Communicated by O. Kinne, Oldendorf/Luhe     

Feeding mechanism of Pelagia noctiluca (Scyphozoa: Semaeostomeae); laboratory and open sea observations

The behaviour of the oral arms of Pelagia noctiluca (Forsskål) has been investigated in open sea predation and in predation induced both in the laboratory and the natural environment. Specimens were first studied in the field and then collected from coastal aggregations in the Gulf of Trieste, North Adriatic Sea, in December 1985 (Lat. 13°40 E, Long. 45°42 N) and in June 1986 (Lat. 13°39 E, Long. 45°43 N). The results of laboratory experiments and in situ observations, recorded on videotapes and photographs, show that the marginal tentacles are utilized to: (1) paralyze the prey; (2) contract and bend inward towards the nearest oral arm. Occasionally the tentacle does not contract and the prey is released; thus prey selection may occur. The oral arms are therefore involved in the: (1) transport of prey from the tentacle to the gastric cavity; (2) catching of motionless prey; (3) anchoring the medusa to the substratum. A similar feeding pattern can explain the survival of several specimens of P. noctiluca near the bottom during the winter of 1985/1986 in the Gulf of Trieste (North Adriatic Sea) in spite of the severe climatic conditions.     

Measurement of filtration rates by infaunal bivalves in a recirculating flume

A flume system and protocol for measuring the filtration rate of infaunal bivalves is described. Assemblages of multi-sized clams, at natural densities and in normal filter-feeding positions, removed phytoplankton suspended in a unidirectional flow of water. The free-stream velocity and friction velocity of the flow, and bottom roughness height were similar to those in natural estuarine waters. Continuous variations in phytoplankton (Chroomonas salinay) cell density were used to measure the filtration rate of the suspension-feeding clam Potamocorbula amurensis for periods of 2 to 28 h. Filtration rates of P. amurensis varied from 100 to 580 liters (gd)^-1 over a free-stream velocity range of 9 to 25 cm s^-1. Phytoplankton loss rates were usually constant throughout the experiments. Our results suggest that suspension-feeding by infaunal bivalves is sensitive to flow velocity.     

Feeding ecology of three species of Astropecten (Asteroidea) coexisting on shallow sandy bottoms of the northwestern Mediterranean Sea

Predation and competition are important biotic interactions influencing populations and communities in marine soft sediments. Sea stars are ubiquitous predators with diverse diets that play functionally important roles in the benthos. In this study, we examined the diet and the ecological roles of three sympatric species of the genus Astropecten (A. aranciacus, A. irregularis pentacanthus and A. platyacanthus). The study was performed between March 2010 and February 2011 on the Maresme coast (northwestern Mediterranean Sea). Results showed that their main diet consisted on gastropods and bivalves, such as Glycymeris glycymeris, Callista chione, Gibbula guttadauri and Cyclope neritea. Food competition between species was avoided by partition of prey resources. Intraspecific differences in the dietary compositions between seasons were found, but not between size classes. Ontogenetic patterns of prey size consumption were recognized in the three species. A large diet overlap was detected between A. aranciacus and A. platyacanthus in winter, due to changes in prey availability. Nevertheless, the analysis of the infaunal community composition and stomach contents indicated that food selection was not associated with prey availability.     

Surface sequestration of chemical feeding deterrents in the Antarctic sponge<Emphasis Type="Italic"> Latrunculia apicalis</Emphasis> as an optimal defense against sea star spongivory

Latrunculia apicalis is a spherically shaped demosponge that previous investigations have shown is rarely preyed upon by sea stars which are the dominant spongivores in antarctic benthic communities. Prior studies have also demonstrated that L. apicalis produces organic compounds that elicit a tube foot retraction response in the keystone spongivorous sea star Perknaster fuscus that can be used as a reliable assay for feeding deterrence. L. apicalis is known to contain discorhabdin alkaloids which serve, among other roles, as the source of its green coloration. To assess the defensive nature of the discorhabdin alkaloids toward P. fuscus, we have determined discorhabdin G concentrations in discrete sponge layers and evaluated those concentrations in the P. fuscus bioassay. In discorhabdin G-bearing sponges, we found a gradient of discorhabdin G that falls off rapidly toward the center of the sponge. On average, 52% of total discorhabdin G in a given sponge was found within 2 mm of the sponge surface. Tube foot retraction responses to extracts from the surface tissues (0–2 mm depth) of L. apicalis were compared to those of an inner layer (8–10 mm depth) and to a sample comprised of the same inner layer spiked with discorhabdin G at a concentration equivalent to that of the surface tissues. Tube foot retraction response times to extracts of the surface layers and the spiked inner layers were not statistically different, but were significantly greater than responses to the unaltered inner layer and controls. These results support the predictions of the optimal defense theory as L. apicalis sequesters its defensive chemistry (discorhabdin G) in its most vulnerable surface tissues, where the likelihood of predation from sea stars is highest. As antarctic sponges are generally preyed upon by extraoral feeding sea stars rather than deeper biting predators such as fish, surface sequestration may be uniquely adaptive in sessile macroinvertebrates occurring in antarctic marine benthic environments.Communicated by P.W. Sammarco, Chauvin     

Spatial and temporal variation in the diet of a high trophic level predator,the Australian fur seal (<Emphasis Type="Italic">Arctocephalus pusillus doriferus</Emphasis>)

This study quantifies the manner in which Australian fur seals, Arctocephalus pusillus doriferus, use their prey in a spatial and temporal context. We analysed 977 scat and 66 regurgitate samples collected from Tasmanian breeding colonies and haul-outs between 1994 and 2000. Diagnostic prey remains identified in the scats represented 35 fish taxa and 8 cephalopod taxa. The main taxa identified in scats, where frequency of occurrence was 10%, were leatherjacket species (family Monocanthidae), redbait (Emmelichthys nitidus), barracouta (Thyrsites atun), jack mackerel (Trachurus declivis) and red cod (Pseudophysis bachus). Regurgitates were dominated by cephalopods, primarily Goulds squid (Nototodarus gouldi), Octopus maorum, O. berrima/pallidus and Sepia apama. Discriminant function analyses indicated that there were generally no significant differences in the composition of the diet between colonies within a year, suggesting that prey distribution is fairly uniform throughout Bass Strait at those time scales. The diet at breeding colonies, however, exhibited significant inter- and intra-annual variation, determined by the presence of several key taxa, such as barracouta and a species of scorpionfish (family Scorpaenidae). The diet composition also varied regionally, between Bass Strait and southern Tasmania in spring 1999 and autumn 2000, with redbait, barracouta and a species of scorpionfish identified as the main taxa contributing to this difference. Redbait occurred in the diet only in southern Tasmania, whereas barracouta and scorpionfish occurred only in Bass Strait.Electronic Supplementary Material Supplementary material is available in the online version of this article at http://dx.doi.org/10.1007/s00227-003-1219-0.Communicated by G.F. Humphrey, Sydney     

Chemosensory responses and foraging behavior of the seastar<Emphasis Type="Italic"> Pycnopodia helianthoides</Emphasis>

Chemical cues released by damaged or dead organisms can affect how and where benthic organisms feed. These cues may cause predators to act as opportunistic scavengers in lieu of their normal predatory role. A scavenger, as defined in this study, is an organism that consumes damaged and/or dead organisms. In-situ experiments were performed to determine how the seastar Pycnopodia helianthoides (Brandt) reacts in the presence of chemical cues from one of its prey species, the butter clam Saxidomus giganteus (Deshayes), using both intact and damaged individuals. The results of these experiments suggest that P. helianthoides use their chemosensory abilities to locate damaged/dead prey. The role of current in propagating chemical cues was paramount in this foraging activity. P. helianthoides chose damaged prey over live prey even when live prey was encountered en route to the damaged individual. This study suggests that chemical cues emitted from damaged or dead individuals may cause significant changes in foraging tactics of key predators, thus altering food-web dynamics.Communicated by J.P. Grassle, New Brunswick     

Comparative allometries of gut-passage time,gut content and metabolic faecal loss inMytilus edulis andCerastoderma edule

Allometric relations were determined in bivalves collected from approximate mid-tide levels in Biscay, Spain, during March 1987. Species compared included the epifaunal suspension-feederMytilus edulis L. (9 to 1 108 mg dry soft-tissue weight) from a rocky-shore population at Meinakotz Beach, and the infaunal deposit-feederCerastoderma edule (L.) (1 to 192 mg dry soft-tissue weight) from the mudflats in Mundaka Ría. Relative to M. edulis, and compared per unit dry tissue weight,C. edule had similar palp areas but smaller gill areas. In addition, to help maximize absorption from organically-poor deposits,C. edule ingested three to four times as much food per hour, but had gut contents that were five to six times greater, so that gut-passage times available for the extraction of nutrients were 2.5 times longer. Metabolic faecal losses, which were comprised of endogenous materials lost from the bivalve into the gut, were two to three times greater inC. edule, but similar to those ofM. edulis when expressed per unit mass ingested by each species. These losses were very substantial, being equivalent to as much as 15% of the ingested mass, and represent a significant indirect cost that is presumably incurred largely by the intracellular digestion characteristic of bivalves. Weight exponents indicated that such metabolic investment represented an unchanging proportion of the total costs of growth. They also showed that age-related constraints on total production did not stem from decreasing gut content. Rather, associated exponents identified limitations to the production in each species as being linked with marked reductions of gutpassage time and ingestion rate, and indicated that these limitations do not derive from corresponding decreases in gill or palp areas.     

Biochemical composition,energy content and chemical antifeedant and antifoulant defenses of the colonial Antarctic ascidian<Emphasis Type="Italic"> Distaplia cylindrica</Emphasis>

The colonial ascidian Distaplia cylindrica occurs both as scattered individual colonies or in gardens of colonies in fine-grained soft substrata below 20 m depths off Anvers Island along the Antarctic Peninsula. Individual colonies, shaped as tall rod-like cylinders and anchored in the sediments by a bulbous base, may measure up to 7 m in height. D. cylindrica represent a considerable source of materials and energy for prospective predators, as well as potential surface area for fouling organisms. Nonetheless, qualitative in situ observations provided no evidence of predation by sympatric predators such as abundant sea stars, nor obvious biofouling of colony surfaces. Mean energy content of whole-colony tissue of D. cylindrica was relatively high for an ascidian (14.7 kJ g^–1 dry wt), with most of this energy attributable to protein (12.7 kJ g^–1 dry wt). The sympatric omnivorous sea star Odontaster validus consistently rejected pieces of D. cylindrica colonies in laboratory feeding assays, while readily ingesting similarly sized alginate food pellets. Feeding deterrence was determined to be attributable to defensive chemistry, as colonies of D. cylindrica are nutritionally attractive and lack physical protection (conspicuous skeletal elements or a tough outer tunic), and O. validus display significant feeding-deterrent responses to alginate food pellets containing tissue-level concentrations of organic extracts. In addition, high acidity measured on outer colony surfaces (pH 1.5) as well as homogenized whole-colony tissues (pH 2.5) are indicative of surface sequestration of inorganic acids. Agar food pellets prepared at tissue levels of acidity resulted in significant feeding deterrence in sea stars. Thus, both inorganic acids and secondary metabolites contribute to chemical feeding defenses. D. cylindrica also possesses potent antifoulant secondary metabolites. Tissue-level concentrations of hydrophilic and lipophilic extracts caused significant mortality in a sympatric pennate diatom. Chemical feeding deterrents and antifoulants are likely to contribute to the abundance of D. cylindrica and, in turn, play a role in regulating energy transfer and community structure in benthic marine environments surrounding Antarctica.Communicated by P.W. Sammarco, Chauvin     

Induced defensive responses by the bivalve Mytilus edulis to predators with different attack modes

Predators such as crabs, whelks, and sea stars attack their bivalve prey in different ways, and predator-induced defenses are an important means of protection. The degree to which induced defenses are specific to different predators, however, remains largely unknown. In laboratory experiments (June to August 1998), we raised mussels (Mytilus edulis L.) in the presence of a drilling predator [the whelk Nucella lapillus (L.)] or a crushing predator [the crab Carcinus maenas (L.)] to determine whether induced changes in prey shell thickness, size, or shape occurred and whether changes were predator-specific. Over a 2 month period, juvenile mussels were exposed to waterborne cues from actively feeding crabs or whelks. Mussels produced thicker shell lips in response to both predators relative to control mussels raised in their absence, and the difference was significantly greater in response to whelks than to crabs. Mussels exposed to whelks showed significantly smaller increases in shell length and width and total wet weight than did mussels exposed to crabs. Thus, there may be a trade-off between shell thickness and linear shell growth and a potential delay in attaining a size refuge from predation. Received: 4 August 1999 / Accepted: 31 January 2000