PREDATION ON THE BIVALVE CHOROMYTILUS MERIDIONALIS (KR.) BY THE GASTROPOD NATICA (TECTONATICA) TECTA ANTON

The effects of a population of the boring gastropod Natica tectaon the bivalve Choromytilus meridionalis were investigated atBailey's Cottage, False Bay, South Africa. In July 1979 theN. tecta density on the mussel bed averaged 69 m^–2 andthe population consisted mainly of reproductively mature individualsbetween 20–33 mm shell width. Laboratory experiments on N. tecta showed that prey size selectionis an increasing function of predator size. The prey size rangetaken by large N. tecta is also greater than that taken by smallindividuals. The position of the borehole on the mussel shellis a function of the way in which the shell is held by the footduring the boring process. Consumption rates measured in thelaboratory showed an increase from approximately 1 kJ per weekin 18 mm N. tecta to 4.5 kJ per week in 28 mm individuals. Populationconsumption in the field was calculated as 663 kJ m^–2month^–1. It was estimated that at this rate the standingcrop of mussels in the pool would be eliminated within 10 months.Field measurements showed significant depletion after 6 months. New spat settlement of mussels occur every 4–6 years.The growth curve shows that after one year the population meansize exceeds 30 mm shell length, which is beyond the prey selectionsize range of small N. tecta. It was concluded that at the timeof a new mussel settlement a niche is provided for the simultaneoussettlement and growth of juvenile N. tecta in high densities.However, within one year the increase in prey size, togetherwith depletion due to over-exploitation, limits population growthand density in N. tecta. (Received 14 March 1980;     

Optimal foraging versus shared doom effects: interactive influence of mussel size and epibiosis on predator preference

In the western Baltic Sea, the highly competitive blue mussel Mytilus edulis tends to monopolize shallow water hard substrata. In many habitats, mussel dominance is mainly controlled by the generalist predator Carcinus maenas. These predator-prey interactions seem to be affected by mussel size (relative to crab size) and mussel epibionts.There is a clear relationship between prey size and predator size as suggested by the optimal foraging theory: Each crab size class preferentially preys on a certain mussel size class. Preferred prey size increases with crab size.Epibionts on Mytilus, however, influence this simple pattern of feeding preferences by crabs. When offered similarly sized mussels, crabs prefer Balanus-fouled mussels over clean mussels. There is, however, a hierarchy of factors: the influence of attractive epibiotic barnacles is weaker than the factor ‘mussel size’. Testing small mussels against large mussels, presence or absence of epibiotic barnacles does not significantly alter preferences caused by mussel size. Balanus enhanced crab predation on mussels in two ways: Additional food gain and, probably more important, improvement in handling of the prey. The latter effect is illustrated by the fact that artificial barnacle mimics increased crab predation on mussels to the same extent as do live barnacles.We conclude that crab predation preferences follows the optimal foraging model when prey belong to different size classes, whereas within size classes crab preferences is controlled by epibionts.     

Predation on exotic zebra mussels by native fishes: effects on predator and prey

SUMMARY 1. Exotic zebra mussels, Dreissena polymorpha, occur in southern U.S. waterways in high densities, but little is known about the interaction between native fish predators and zebra mussels. Previous studies have suggested that exotic zebra mussels are low profitability prey items and native vertebrate predators are unlikely to reduce zebra mussel densities. We tested these hypotheses by observing prey use of fishes, determining energy content of primary prey species of fishes, and conducting predator exclusion experiments in Lake Dardanelle, Arkansas. 2. Zebra mussels were the primary prey eaten by 52.9% of blue catfish, Ictalurus furcatus; 48.2% of freshwater drum, Aplodinotus grunniens; and 100% of adult redear sunfish, Lepomis microlophus. Blue catfish showed distinct seasonal prey shifts, feeding on zebra mussels in summer and shad, Dorosoma spp., during winter. Energy content (joules g⁻¹) of blue catfish prey (threadfin shad, Dorosoma petenense; gizzard shad, D. cepedianum; zebra mussels; and asiatic clams, Corbicula fluminea) showed a significant species by season interaction, but shad were always significantly greater in energy content than bivalves examined as either ash-free dry mass or whole organism dry mass. Fish predators significantly reduced densities of large zebra mussels (>5 mm length) colonising clay tiles in the summers of 1997 and 1998, but predation effects on small zebra mussels (≤5 mm length) were less clear. 3. Freshwater drum and redear sunfish process bivalve prey by crushing shells and obtain low amounts of higher-energy food (only the flesh), whereas blue catfish lack a shell-crushing apparatus and ingest large amounts of low-energy food per unit time (bivalves with their shells). Blue catfish appeared to select the abundant zebra mussel over the more energetically rich shad during summer, then shifted to shad during winter when shad experienced temperature-dependent stress and mortality. Native fish predators can suppress adult zebra mussel colonisation, but are ultimately unlikely to limit population density because of zebra mussel reproductive potential.     

Prey selection and the functional response of sea stars (Asterias vulgaris Verrill) and rock crabs (Cancer irroratus Say) preying on juvenile sea scallops (Placopecten magellanicus (Gmelin)) and blue mussels (Mytilus edulis Linnaeus)

Predators in nature include an array of prey types in their diet, and often select certain types over others. We examined (i) prey selection by sea stars (Asterias vulgaris) and rock crabs (Cancer irroratus) when offered two prey types, juvenile sea scallops (Placopecten magellanicus) and blue mussels (Mytilus edulis), and (ii) the effect of prey density on predation, prey selection, and component behaviours. We quantified predation rates, behavioural components (proportion of time spent searching for prey, encounter probabilities) and various prey characteristics (shell strength, energy content per prey, handling time per prey) to identify mechanisms underlying predation patterns and to assess the contribution of active and passive prey selection to observed selection of prey. Sea stars strongly selected mussels over scallops, resulting from both active and passive selection. Active selection was associated with the probability of attack upon encounter; it was higher on mussels than on scallops. The probability of capture upon attack, associated with passive selection, was higher for mussels than for scallops, since mussels can not swim to escape predators. Sea stars consumed few scallops when mussels were present, and so did not have a functional response on scallops (the target prey). Rock crabs exhibited prey switching: they selected mussels when scallop density was very low, did not select a certain prey type when scallop density was intermediate, and selected scallops when scallop density was high relative to mussel density. The interplay between encounter rate (associated with passive selection) and probability of consumption upon capture (associated with both active and passive selection) explained observed selection by crabs. Scallops were encountered by crabs relatively more often and/or mussels less often than expected from random movements of animals at all scallop densities. However, the probability of consumption varied with scallop density: it was lower for scallops than mussels at low and intermediate scallop densities, but tended to be higher for scallops than mussels at high scallop densities. When mussels were absent, crabs did not have a functional response on scallops, but rather were at the plateau of the response. When mussels were present with scallops at relatively low density, crabs exhibited a type II functional response on scallops. Our results have implications for the provision of protective refuges for species of interest (i.e., scallops) released onto the sea bed, such as in population enhancement operations and bottom aquaculture.     

Predation on invasive zebra mussel, Dreissena polymorpha, by pumpkinseed sunfish, rusty crayfish, and round goby

The enemy release hypothesis states that invasive species are successful in their new environment because native species are not adapted to utilize the invasive. If true for predators, native predators should have lower feeding rates on the invasive species than a predator from the native range of the invasive species. We tested this hypothesis for zebra mussel (Dreissena polymorpha) by comparing handling time and predation rate on zebra mussels in the laboratory by two North American species (pumpkinseed, Lepomis gibbosus, and rusty crayfish, Orconectes rusticus) and one predator with a long evolutionary history with zebra mussels (round goby, Neogobius melanostomus). Handling time per mussel (7 mm shell length) ranged from 25 to >70 s for the three predator species. Feeding rates on attached zebra mussels were higher for round goby than the two native predators. Medium and large gobies consumed 50–67 zebra mussels attached to stones in 24 h, whereas pumpkinseed and rusty crayfish consumed <11. This supports the hypothesis that the rapid spread of zebra mussels in North America was facilitated by low predation rates from the existing native predators. At these predation rates and realistic goby abundance estimates, round goby could affect zebra mussel abundance in some lakes.     

Predator (<Emphasis Type="Italic">Carcinus maenas</Emphasis>) nonconsumptive limitation of prey (<Emphasis Type="Italic">Nucella lapillus</Emphasis>) feeding depends on prey density and predator cue type

Predators often have nonconsumptive effects (NCEs) on prey. For example, upon detection of predator cues, prey can reduce feeding activities to hamper being detected by predators. Previous research showed that waterborne chemical cues from green crabs (Carcinus maenas, predator) limit the dogwhelk (Nucella lapillus, prey) consumption of barnacles regardless of dogwhelk density, even though individual predation risk for dogwhelks decreases with conspecific density. Such NCEs might disappear with dogwhelk density if dogwhelks feed on mussels, as mussel stands constitute better antipredator refuges than barnacle stands. Through a laboratory experiment, we effectively found that crab chemical cues limit the per-capita consumption of mussels by dogwhelks at low dogwhelk density but not at high density. The combination of tactile and chemical cues from crabs, however, limited the dogwhelk consumption of mussels at both dogwhelk densities. The occurrence of such NCEs at both dogwhelk densities could have resulted from tactile cues indicating a stronger predation risk than chemical cues alone. Overall, the present study reinforces the notions that prey evaluate conspecific density when assessing predation risk and that predator cue type affects their perception of risk.     

Prey selection within a size-class of mussels, Mytilus edulis, by oystercatchers, Haematopus ostralegus

By comparing the shells of those mussels Mytilus edulis that had been opened by oystercatchers Haematopus ostralegus with mussels of similar size that had not been opened, it was shown that oystercatchers that break into their prey by hammering a hole in the shell selected between prey within a size-class. Ventral hammerers selected mussels that were relatively thin on the ventral surface, were brown in colour and carried few barnacles. Dorsal hammerers selected eroded mussels with thin dorsal shells. Stabbing oystercatchers did not select for thin-shelled prey. In conjunction with the great individual variation in flesh content between mussels of the same length, prey size in this case can be only a poor predictor of prey profitability.     

Drilling under threat: An experimental assessment of the drilling behavior of Nucella lamellosa in the presence of a predator

Using the drilling muricid, Nucella lamellosa (Gmelin 1791) and its prey, the mussel Mytilus trossulus (Gould 1850), the impact of a secondary predator, the crab Cancer gracilis, on drilling was investigated experimentally. The frequency of incomplete holes was compared under two conditions: (1) when the gastropod's natural predator, C. gracilis, was present and (2) when it was absent. The results indicate that the presence of a secondary predator can affect drilling activity, leading to a significant increase in the frequency of incomplete drill holes. The introduction of a secondary predator can also decrease the overall drilling frequency. The size distributions of completely and incompletely drilled mussels suggest that in the presence of the secondary predator the decision by the gastropod to either abandon or continue drilling its prey might be influenced by how much time it has already invested into drilling or the size of the prey item. These results are important for the ecological and evolutionary implications of incomplete drill holes frequencies, especially with regard for their use as proxies for evolutionary prey improvement.     

Adjustment of the activity of α-amylase extracted from the style of the black mussel Choromytilus meridionalis (Krauss) in response to thermal acclimation

The mussel Choromytilus meridionalis (Krauss) is a common inhabitant of the intertidal zone on the west coast of the Cape Peninsula, South Africa, and experiences temperatures of between 8°C when immersed by the tide and at least 25°C on exposure to air. The activity of α-amylase extracted from the crystalline style of freshly-collected mussels has a low temperature coefficient of ≈ 1.12 over much of the temperature range experienced in the natural environment. Warm acclimation results in an increase in the α-amylase activity, despite the fact that individual rate: temperature curves for extracts from mussels acclimated to 8, 15 and 22°C have rather low temperature coefficients of 1.14–1.17 between 10 and 20°C. The increase of activity of the α-amylase following warm acclimation may form an integral part of the improved filtration, ingestion and assimilation which is necessary to offset increased metabolic losses during the warm conditions of the summer months.     

Sub-optimal mussel selection by tufted ducks Aythya fuligula: Test of a hypothesis

A series of tests with four tufted ducks Aythya fuligula, feeding on freshwater mussels Dreissena polymorpha was performed to test the hypothesis that tufted ducks select sub-optimal mussel sizes to avoid the risk of taking items that are too large to be swallowed, or at least, have highly variable profitabilities. The results indicate that the largest mussel size class available influenced the overall pattern of selection and that an increasing risk of taking unprofitable mussels caused a decrease in mean mussel size eaten. Some of the recorded differences between the birds tested could be explained by differences in the shape of the profitability curves for each individual.     

Optimal foraging and risk of claw damage: How flexible are shore crabs in their prey size selectivity?

Understanding the behavioural mechanisms that underlie prey size preference of predators is an essential component of unravelling the processes that govern predator-prey dynamics. In marine systems, despite being able to consume larger and more profitable prey, many molluscivorous predators show a preference for smaller, less profitable prey, most likely to minimize the risk of damaging feeding extremities. Here we assessed the flexibility of this prey size preference. We observed that shore crabs (Carcinus maenas) that were food deprived, and which were offered mussels (Mytilus edulis) of different sizes in dichotomous preference tests, preferred smaller, less profitable mussels. The same result was observed for crabs foraging with a conspecific competitor. Only crabs that were conditioned to feed on the larger, most profitable mussels shifted their prey size preference and ranked the most profitable mussels as highest. Although shore crabs showed flexibility in prey size preference, through which they would be able to cope with environmental variability, our results in general emphasize preference for smaller prey. We discuss the possibility that crabs maximize their long-term feeding rate, in which case it can be optimal to select these smaller mussels.     

GROWTH AND MORTALITY OF THE MUSSELS CHOROMYTILUS MERIDIONALIS (KRAUSS) AND AULACOMYA ATER(MOLINA) AS INDICATORS OF BIOTIC CONDITIONS

Two mussels, Choromytilus meridionalis (Krauss) and Aulacomyaater, (Molina) were used as indicators of biotic factors thatapparently result in radically different sublittoral communitiesat two closely situated islands on the South African west coast.Specimens of both species were attached to submerged plates,which were either unprotected or protected from predation bycages. C. meridionalis grew faster and tolerated silting andsmothering better than A. ater. At Malgas Island adults of C.meridionalis are completely absent and those of A. ater scarcedue to intense predation by rock-lobsters. When protected againstpredation, growth of both species was significantly faster andsurvivership higher at Malgas than at Marcus Island, where C.meridionalis is abundant but A. ater absent. When the musselswere unprotected at Malgas, they were eaten in a few hours.At Marcus Island where rock-lobsters are virtually absent, cagedmussels grow faster than unprotected individuals and survivedlonger. Drilling whelks (mostly Nucella squamosa (Lamarck))were major predators at Marcus. Relative intensity of rock-lobsterpredation and competition between the mussels appear to controlthe structure and diversity of the sublittoral communities atthese sites, rather than physical factors. Growth rate, shellthickness and tolerance of smothering by other species determinewhich species of mussel predominates. (Received 5 April 1988; accepted 11 July 1988)     

Zebra mussels affect benthic predator foraging success and habitat choice on soft sediments

The introduction of zebra mussels (Dreissena spp.) to North America has resulted in dramatic changes to the complexity of benthic habitats. Changes in habitat complexity may have profound effects on predator-prey interactions in aquatic communities. Increased habitat complexity may affect prey and predator dynamics by reducing encounter rates and foraging success. Zebra mussels form thick contiguous colonies on both hard and soft substrates. While the colonization of substrata by zebra mussels has generally resulted in an increase in both the abundance and diversity of benthic invertebrate communities, it is not well known how these changes affect the foraging efficiencies of predators that prey on benthic invertebrates. We examined the effect of zebra mussels on the foraging success of four benthic predators with diverse prey-detection modalities that commonly forage in soft substrates: slimy sculpin (Cottus cognatus), brown bullhead (Ameirus nebulosus), log perch (Percina caprodes), and crayfish (Orconectes propinquus). We conducted laboratory experiments to assess the impact of zebra mussels on the foraging success of predators using a variety of prey species. We also examined habitat use by each predator over different time periods. Zebra mussel colonization of soft sediments significantly reduced the foraging efficiencies of all predators. However, the effect was dependent upon prey type. All four predators spent more time in zebra mussel habitat than in either gravel or bare sand. The overall effect of zebra mussels on benthic-feeding fishes is likely to involve a trade-off between the advantages of increased density of some prey types balanced against the reduction in foraging success resulting from potential refugia offered in the complex habitat created by zebra mussels.     

Mussel beds in deeper water provide an unusual situation for competitive interactions between the seastars Leptasterias polaris and Asterias vulgaris

The common seastars Leptasterias polaris and Asterias vulgaris show competitive interactions in shallow subtidal communities in the northern Gulf of St. Lawrence, particularly during summer when aggregations of the two seastars forage on mussel beds at 1-2 m in depth. We examined interactions between the two seastars in a different situation, in a mussel bed at 6 m in depth (a rare situation in this region). In the deeper mussel bed, seastars were three times more abundant than in the shallower beds, and the mussels were larger. The deeper bed disappeared rapidly due to the intense predation. Although decreased prey abundance should have favored interference interactions, we did not detect either partitioning of mussels by size or avoidance of A. vulgaris by L. polaris as previously reported when mussels are in short supply in shallower water. The lack of an avoidance behavior by L. polaris, together with the higher proportion of L. polaris than A. vulgaris that were feeding, suggests that in this situation, the dominance of A. vulgaris (observed in shallower water) is attenuated, or that L. polaris may dominate.     

Plastic Responses of a Sessile Prey to Multiple Predators: A Field and Experimental Study

Background

Theory predicts that prey facing a combination of predators with different feeding modes have two options: to express a response against the feeding mode of the most dangerous predator, or to express an intermediate response. Intermediate phenotypes protect equally well against several feeding modes, rather than providing specific protection against a single predator. Anti-predator traits that protect against a common feeding mode displayed by all predators should be expressed regardless of predator combination, as there is no need for trade-offs.

Principal Findings

We studied phenotypic anti-predator responses of zebra mussels to predation threat from a handling-time-limited (crayfish) and a gape-size-limited (roach) predator. Both predators dislodge mussels from the substrate but diverge in their further feeding modes. Mussels increased expression of a non-specific defense trait (attachment strength) against all combinations of predators relative to a control. In response to roach alone, mussels showed a tendency to develop a weaker and more elongated shell. In response to crayfish, mussels developed a harder and rounder shell. When exposed to either a combination of predators or no predator, mussels developed an intermediate phenotype. Mussel growth rate was positively correlated with an elongated weaker shell and negatively correlated with a round strong shell, indicating a trade-off between anti-predator responses. Field observations of prey phenotypes revealed the presence of both anti-predator phenotypes and the trade-off with growth, but intra-specific population density and bottom substrate had a greater influence than predator density.

Conclusions

Our results show that two different predators can exert both functionally equivalent and inverse selection pressures on a single prey. Our field study suggests that abiotic factors and prey population density should be considered when attempting to explain phenotypic diversity in the wild.     

Type and nature of cues used by Nucella lapillus to evaluate predation risk

The ability of prey to detect and adequately respond to predation risk influences immediate survival and overall fitness. Chemical cues are commonly used by prey to evaluate risk, and the purpose of this study was to elicit the nature of cues used by prey hunted by generalist predators. Nucella lapillus are common, predatory, intertidal snails that evaluate predatory risk using chemical cues. Using Nucella and a suite of its potential predators as a model system, we explored how (1) predator type, (2) predator diet, and (3) injured conspecifics and heterospecifics influence Nucella behavior. Using laboratory flumes, we determined that Nucella responded only to the invasive green crab (Carcinus maenas), the predator it most frequently encounters. Nucella did not respond to rock crabs (Cancer irroratus) or Jonah crabs (Cancer borealis), which are sympatric predators but do not frequently encounter Nucella because these crabs are primarily subtidal. Predator diet did not affect Nucella responses to risk, although starved predator response was not significantly different from controls. Since green crabs are generalist predators, diet cues do not reflect predation risk, and thus altering behavior as a function of predator diet would not likely benefit Nucella. Nucella did, however, react to injured conspecifics, a strategy that may allow them to recognize threats when predators are difficult to detect. Nucella did not react to injured heterospecifics including mussels (Mytilus edulis) and herbivorous snails Littorina littorea, suggesting that they are responding to chemical cues unique to their species. The nature of cues used by Nucella allows them to minimize costs associated with predator avoidance.     

Stereotypic and size-selective predation in Polinices pulchellus (Gastropoda: Naticidae) Risso 1826

Polinices pulchellus were size-selective in their choice of Cerastoderma edule. Large predators (12-15.9 mm shell length) selected both larger and a wider size range of cockles than smaller individuals (4-11.9 mm shell length). Considerable overlap occurred in the sizes of cockles frequently drilled by different size classes of snails, indicating that certain sizes of cockles may be most profitable to a wide range of predator sizes. Consumption rates were highest during July and August and were closely related to seawater temperature. Inner and outer drill hole diameters were both correlated with predator size, and the morphology of the drill hole was geometrically similar across a range of predator sizes. Polinices pulchellus showed no preference for either the left or right valve and drilled most cockles in the centre of the shell valve. The relationship between the distance of the drill hole from the umbo and prey size was unaffected by predator size, such that predators of different sizes were not found to drill cockles in different positions. When disturbed during drilling, incomplete drill holes were abandoned and, when drilling resumed, it occurred in new locations on the surface of the shell valve. The findings of this study highlight the stereotyped nature of drilling behaviour seen in the family Naticidae.     

Feeding physiology of the carnivorous gastropod Thais clavigera (Kuster): do they eat “soup”?

The ability to feed on suspended and dissolved organic nutrients may have been retained in predatory gastropods during evolution. The carnivorous muricid neogastropod Thais clavigera feeds on prey by boring through their shells, followed by extracellular digestion and suction of the nutrient-rich fluid of the prey's body tissues. This study reports on the effect of feeding on suspended and soluble organic nutrients (SSONs) on the survival, growth, and various physiological activities including scope for growth and glycogen stores of T. clavigera. Juvenile T. clavigera of similar shell length (23.8±1.7 mm) were either starved, fed with mussel Septifer virgatus, fed with SSONs from homogenized mussel flesh (S. virgatus), or fed with both mussels and SSONs, and kept in artificial seawater (salinity: 30‰) for 50 days. Ingestion of SSONs by the animals was significant. Feeding with the “soup” (i.e., SSONs) reduced tissue wastage and improved condition index of the snails. T. clavigera fed in this manner were intermediates between the starved and the mussel-fed groups in terms of mortality, growth, food consumption, respiration, scope for growth, and glycogen content measurements. Furthermore, T. clavigera fed with both mussels and SSONs exhibited an identical energy requirement and similar values of various physiological measurements as that of those fed solely on mussel flesh. Feeding of SSONs contributed >10% of the overall energy requirement when both SSONs and mussel prey were available. The results indicate that energy from suspended and dissolved organic nutrients can contribute to the maximization of energy input in T. clavigera, which may favor better survivorship and thus lifetime fitness.     

Predator size interacts with habitat structure to determine the allometric scaling of the functional response

While both predator body size and prey refuge provided by habitat structure have been established as major factors influencing the functional response (per capita consumption rate as a function of prey density), potential interactions between these factors have rarely been explored. Using a crab predator (Panopeus herbstii) – mussel prey (Brachidontes exustus) system, we examined the allometric scaling of the functional response in oyster (Crassostrea virginica) reef habitat, where crevices within oyster clusters provide mussels refuge from predation. A field survey of mussel distribution showed that mussels attach closer to the cluster periphery at high mussel density, indicating the potential for saturation of the refuge. In functional response experiments, the consumption rate of large crabs was depressed at low prey density relative to small crabs, while at high prey density the reverse was true. Specifically, the attack rate coefficient and handling time both decreased non‐linearly with crab size. An additional manipulation revealed that at low prey densities, the ability of large crabs to maneuver their claws and bodies to extract mussels from crevices was inhibited relative to small crabs by the structured habitat, reducing their attack rate. At high prey densities, crevices were saturated, forcing mussels to the edge of clusters where crabs were only limited by handling time. Our study illuminates a potentially general mechanism where the quality of the prey refuge provided by habitat structure is dependent on the relative size of the predator. Thus anthropogenic influences that alter the natural crab size distribution or degrade reef habitat structure could threaten the long‐term stability of the crab –mussel interaction in reefs.     

Clumping behavior as a strategy against drilling predation: Implications for the fossil record

Drilling gastropod predators are of particular interest to paleontologists, because predatory drill-holes in marine invertebrates serve as one of the rare sources of data for the study of ancient predator-prey interactions. Modern laboratory studies are an important part of predation research providing valuable ecological insight and constraining fossil evidence and interpretations. Previous studies have shown that mussels use clumping behavior against durophagous predation [Okamura, B., 1986. Group living and the effects of spatial position in aggregations of Mylitus edulis. Oecologia 69, 341-347.; Lin, J., 1991. Predator-prey interactions between blue crabs and ribbed mussels living in clumps. Estuar. Coast. Shelf Sci. 32, 61-69.], but its role against drilling predation had not been explored. In this study, we explore the effect of clumping on predator success (drill-hole frequency) and prey handling (drill-hole position) using the mussel, Mytilus trossulus, as prey and the gastropod, Nucella lamellosa, as drilling predator. We assigned mussels to two groups: in one, mussels were allowed to clump together with their byssal threads, and in the other, they were kept separate. We observed a significant decrease in the drilling frequency within the group containing clumped mussels, confirming that clumping acts as a successful anti-predatory strategy against drilling predators. The use of clumping as an effective strategy against multiple types of predators may relax the trade-offs associated with aggregated lifestyles [Bertness, M.D., Grosholz, E., 1985. Population dynamics of the ribbed mussel, Geukensia demissa: the costs and benefits of an aggregated distribution. Oecologia 67, 192-204.]. The increased benefit and unchanged metabolic cost of clumped living alters estimates of individual fitness with evolutionarily significant implications (e.g., eliminating the need to invoke group or species selection to explain the adaptive benefit of an aggregated lifestyle). In spite of potential differences in prey handling and grappling due to clumping, mean drill-hole placement and variation in drill-hole placement showed no significant differences between the two groups. These observations suggest that comparison of predation intensities across clumping and non-clumping taxa must consider the anti-predatory effect of this behavior.