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.     

Effects of juvenile non-indigenous Carcinus maenas on the growth and condition of juvenile Cancer irroratus

The Atlantic rock crab, Cancer irroratus, is a commercially fished species and a critical prey item for the American lobster, Homarus americanus, in Atlantic Canada. The recent invasion of European green crab, Carcinus maenas, may have significant effects on the growth and condition of native C. irroratus, because both species overlap spatially and temporally and have similar habitat and dietary requirements. To examine such potential effects, we measured the growth of juvenile C. irroratus in the presence of juvenile C. maenas over a period of 4 months (growing season), under the following species combinations: (1) one C. irroratus (10-25 mm CW); (2) two C. irroratus (10-25 mm CW); (3) one C. irroratus (10-25 mm CW) and one C. maenas (10-15 mm CW). Morphological measurements included pre- and post-molt carapace width, chela height, abdomen width (mm), weight (g), and estimates of molt increment (%) and intermolt duration (days). Analysis of the hepatopancreas for % lipid content at the end of the experiment provided an estimate of physiological condition. The effect of the presence of C. maenas on the growth of C. irroratus shifted from negative to positive, when C. irroratus reached CW of 19-22 mm and gained a presumably significant size advantage over C. maenas. The positive effect resulted from increased energy intake through crab consumption. In the absence of crab consumption, the presence of a second crab (conspecific or C. maenas) had no effect on growth. C. irroratus consumed crabs more frequently when the second individual was a green crab than a conspecific. Consumption of C. maenas had a pronounced effect on the growth rate of C. irroratus, resulting in shorter intermolt periods and larger percent molt increments than in the presence of a conspecific. Therefore, the presence of juvenile C. maenas does not appear to have a prolonged negative effect on the growth of C. irroratus; rather, it may provide an additional food item as rock crabs grow, as long as encounters between the two species occur at high enough rates.     

Effects of substrate on interactions between juvenile sea scallops (Placopecten magellanicus Gmelin) and predatory sea stars (Asterias vulgaris Verrill) and rock crabs (Cancer irroratus Say)

We investigated the effect of substrate (glass bottom, sand, granule, pebble) on predation of juvenile sea scallops (Placopecten magellanicus) by sea stars (Asterias vulgaris) and rock crabs (Cancer irroratus) at two prey sizes (11-15 mm and 24-28 mm shell height), and two prey densities (10 and 30 scallops per aquarium) in laboratory experiments. Specifically, we quantified predation rate and underlying behaviours (proportion of time a predator spent searching for and handling prey, encounter rate between predators and prey, and various outcomes of encounters). We detected a significant gradual effect of particle size of natural substrates on sea star predation: specifically, predation rate on and encounter rate with small scallops tended to decrease with increasing particle size (being highest for sand, intermediate for granule, and lowest for pebble). Substrate type did not significantly affect predation rates or behaviours of sea stars preying on large scallops or of rock crabs preying on either scallop size classes. Other factors, such as prey size and density, were important in the scallop-sea star and scallop-rock crab systems. For example, predation rate by sea stars and crabs and certain sea star behaviours (e.g. probability of consuming scallops upon capture) were significantly higher with small scallops than with large scallops. As well, in interactions between small scallops and sea stars, predation rate and encounter rate increased with prey density, and the proportion of time sea stars spent searching was higher at low prey density than high prey density. Thus, substrate type may be a minor factor determining predation risk of seeded scallops during enhancement operations; prey size and prey density may play a more important role. However, substrate type still needs to be considered when choosing a site for scallop enhancement, as it may affect other scallop behaviours (such as movement).     

Subtidal-intertidal trophic links: American lobsters [Homarus americanus (Milne-Edwards)] forage in the intertidal zone on nocturnal high tides

Homarus americanus (Milne-Edwards), the American lobster, is a predator in New England subtidal communities, feeding on ecologically important grazers (sea urchins), mesopredators (crabs), and basal species (mussels). In this study, we provide the first report of adult American lobsters foraging in rocky intertidal habitats during nocturnal high tides. Censuses by SCUBA divers in the low intertidal (Chondrus crispus Stackhouse) zone showed mean densities of 2.2 lobsters/20 m² on nocturnal high tides, with contrasting low densities of 0.18/20 m² during diurnal high tides. Nocturnal high-tide intertidal densities were 62% of those reported in a previous study of lobsters in nearby subtidal rocky areas (Novak, 2004). The average carapace length of lobsters in the intertidal at night was > 50 mm. These lobsters were actively foraging in the intertidal with collected individuals having a mean stomach fullness of 67%. Prey found in the stomach contents primarily consisted of crabs, mussels and snails. Field experiments showed that lobsters rarely fed on medium to large size individuals of the common intertidal snail, Littorina littorea (L.). In contrast, experiments with local crab species demonstrated that lobsters actively and readily prey on Cancer irroratus (Say) and Carcinus maenas (L.), but were significantly less likely to consume Cancer borealis (Stimpson). The abundance of Carcinus maenas and blue mussels (Mytilus edulis L.) in the intertidal zone may explain the upshore movement of lobsters. Since nocturnal migration of Homarus americanus into the intertidal zone has not been documented before, our understanding of the dynamics of New England intertidal communities needs to be expanded to include this predator.     

Predatory behavior and preference of a successful invader, the mud crab Dyspanopeus sayi (Panopeidae), on its bivalve prey

Predator-prey relationships between the panopeid crab, Dyspanopeus sayi, and the mytilid, Musculista senhousia, were investigated. Through laboratory experiments, prey-handling behavior, prey size selection, predator foraging behavior and preferences for two types of prey (M. senhousia and the Manila clam Ruditapes philippinarum) were assessed. Handling time differed significantly with respect to the three prey sizes offered (small: 15.0-20.0 mm shell length, SL; medium: 20.1-25.0 mm SL; and large: 25.1-30.0 mm SL); mud crabs were more efficient in predating medium-small than large prey. Although differences in prey profitability were not evident, D. sayi exhibited a marked reluctance to feed on larger-sized prey whilst smaller, more easily predated mussels were available. Size selection may be the result of a mechanical process in which encountered prey are attacked but rejected if they remain unbroken after a certain number of opening attempts. D. sayi exhibited inverse density-dependent foraging. A significant higher mortality of prey was evident at low prey density. Thus, at low predator density, the D. sayi-M. senhousia interaction was a destabilizing type II functional response. Interference responses affected the magnitude of predation intensity by D. sayi on M. senhousia, since as the density of foraging crabs increased, their foraging success fell. At high density (4 crabs tank⁻¹), crabs engaged in a high amount of agonistic activity when encountering a conspecific specimen, greatly diminished prey mortality. Finally, presenting two types of prey, Manila clam juveniles were poorly predated by mud crabs, which focused their predation mostly on M. senhousia. It is hypothesized that, when more accessible prey is available, mud crabs will have a minimal predatory impact on commercial R. philippinarum juvenile stocks.     

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.     

Disentangling interference competition from exploitative competition in a crab–bivalve system using a novel experimental approach

In predator–prey relationships such as those between crabs and their bivalve prey, interference competition is a topic of intense investigation as it can have profound consequences on the dynamics of both predator and prey populations. However in laboratory experiments – also those on crab–bivalve systems – workers never adequately disentangled interference competition from exploitative competition, as prey depletion was never compensated. Hitherto, experimental studies on crab–bivalve systems lack direct behavioural observations and have provided only indirect and thus inconclusive evidence of interference competition. We studied interference competition in adult male shore crabs Carcinus maenas that foraged on blue mussels Mytilus edulis. We developed a novel type of experimental tank to replenish each consumed mussel, and thus to keep prey levels constant. We conducted two experiments in which we varied number of crabs (1, 2, 4) and number of mussels (first experiment: 4, 8, 16, 32; second experiment: 8, 32, 128) and directly observed the foraging behaviour of crabs (foraging area=0.25 m²). In the first experiment, feeding rates decreased with increasing crab density only at mussel density 16 because both search time and time spent in agonistic interactions increased. At other mussel densities, variation in crab density did not affect feeding rates, possibly because of low statistical power and the narrow range of mussel densities offered. In the second experiment feeding rates decreased with increasing crab density because crabs spent more time in agonistic interactions and handling their prey. Feeding rates increased with increasing mussel density. Overall, crabs spent on average 14–18% of their foraging time in agonistic behaviours, while on three out of 64 occasions feeding rates decreased because mussels were stolen (kleptoparasitism). Concluding, we have shown that interference competition occurs in absence of prey depletion, while conducting direct behavioural observations aid to identify the behavioural processes that underlie interference competition.     

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.     

The Effects of a Competitor on the Foraging Behaviour of the Shore Crab Carcinus maenas

Optimal Diet Theory suggests that individuals make foraging decisions that maximise net energy intake. Many studies provide qualitative support for this, but factors such as digestive constraints, learning, predation-risk and competition can influence foraging behaviour and lead to departures from quantitative predictions. We examined the effects of intraspecific competition within a classic model of optimal diet – the common shore crab, Carcinus maenas, feeding on the mussel, Mytilus edulis. Unexpectedly, we found that breaking time (Tb), eating time (Te), and handling time (Th) all decreased significantly in the presence of a conspecific. Reduced handling time in the presence of a competitor resulted in an increased rate of energy intake, raising the question of why crabs do not always feed in such a way. We suggest that the costs of decreased shell breaking time may be increased risk of claw damage and that crabs may be trading-off the potential loss of food to a competitor with the potential to damage their claw whilst breaking the shell more rapidly. It is well documented that prey-size selection by crabs is influenced by both the risk of claw damage and competition. However, our results are the first to demonstrate similar effects on prey handling times. We suggest that crabs maximise their long-term rate of energy intake at a scale far greater than individual foraging events and that in order to minimise claw damage, they typically break shells at a rate below their maximum. In the presence of a competitor, crabs appear to become more risk-prone and handle their food more rapidly, minimising the risk of kleptoparasitism.     

Predation by gulls on crabs in rocky intertidal and shallow subtidal zones of the Gulf of Maine

Most organisms in intertidal areas are marine in origin; many have distributions that extend into the subtidal zone. Terrestrial predators such as mammals and birds may exploit these animals during low tide and can have considerable effects on intertidal food webs. Several studies have shown that avian predators are capable of reducing densities of sessile and slow-moving intertidal invertebrates but very few studies have considered avian predation on mobile invertebrate predators such as crabs. In this study, we investigated predation by Great Black-backed Gulls (Larus marinus Linnaeus) on three species of crabs (Cancer borealis Stimpson, Cancer irroratus Say, and Carcinus maenas Linnaeus). The study was at Appledore Island, ME (a gull breeding island) and 8 other sites throughout the Gulf of Maine, including breeding islands and mainland sites. On Appledore Island, intertidal and subtidal zones provided over one-third of prey remains found at gull nests, and crabs were a substantial proportion (∼ 30% to 40%) of the total remains. Similarly, collections of prey remains from intertidal areas indicated that crabs were by far the most common marine prey. C. borealis was eaten far more often and C. irroratus and C. maenas less often than expected at each site. Comparing numbers of carapaces to densities of crabs in low intertidal and shallow subtidal zones at each site, we estimated that gulls remove between 15% and 64% of C. borealis during diurnal low tides. The proportion of C. borealis eaten by gulls was independent of proximity to a gull colony. Approximately 97% of the outer coast of Maine is within 20 km of a breeding island. Thus, a lot of gull predation on crabs may occur throughout the Gulf of Maine during summer months. Crabs are important predators of other invertebrates; if predation by gulls reduces the number of crabs in intertidal and shallow subtidal areas, gulls may have important indirect effects on intertidal food webs.     

Position of horseshoe crabs in estuarine food webs: N and C stable isotopic study of foraging ranges and diet composition

To discern the position of horseshoe crabs as a potentially important predator in estuarine food webs, we determined where they foraged and what they ate. We used N and C stable isotopes to link adult horseshoe crabs to their oraging locations and potential food sources in Pleasant Bay, Cape Cod. The δ¹⁵N in tissues of horseshoe crabs and their potential foods suggest crabs were loyal to local foraging sites and did not forage substantially in subestuaries receiving >110 kg N ha⁻¹ year⁻¹. Among locations where crabs foraged, δ¹³C values in potential foods showed that food webs in subestuaries subject to higher N loads were supported by algal producers, while food webs in subestuaries with lower N loads were also supported by Spartina. δ¹³C values in horseshoe crab tissue did not change with load, suggesting they ate a mixed diet, regardless of N load. N and C isotopes in horseshoe crab feces were similar to signatures of estimated diet, suggesting low assimilation efficiency, perhaps due to ingestion of low quality organic matter. Although horseshoe crabs were relatively opportunistic in foraging habits, conservation or culture of horseshoe crabs may require habitats with higher water quality, ample particulate organic matter, and supporting a variety of prey.     

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.     

Exploring the decline of oyster beds in Atlantic Canada shorelines: potential effects of crab predation on American oysters (<Emphasis Type="Italic">Crassostrea virginica</Emphasis>)

Atlantic Canada’s American oyster (Crassostrea virginica) beds, while economically and ecologically important, have been in decline over the past few decades. Predation by crabs, in particular by the European green crab (Carcinus maenas), has been proposed as one of the potential causes of such decline. Hence, this study examined oyster mortality levels in multiple beds across Prince Edward Island (PEI) and then experimentally assessed the contribution of green crab predation to oyster mortality. Results from surveys conducted in 10 estuaries across PEI in 2014 indicate that the probability of mortality for small oysters was significantly higher when green crabs were present then in areas without green crabs. This probability of mortality was significantly less when there was the presence of alternative prey like natural mussel beds (Mytilus edulis). The odds of oyster mortality were also higher when beds had rock crabs (Cancer irroratus) compared to beds with no rock crabs. Given the potential importance of green crab predation, its influence was assessed in 2015 using two field experiments with tethered oysters. Our results indicate that odds of small oyster mortality occurring were much higher in green crab inclusion cages than in the open environment and the exclusion cages. These results reaffirm that oysters up to ~40 mm SL are vulnerable to predation, and at least some of the mortality affecting these oysters can be causally attributed to green crab predation. Green crab predation rates upon small oysters are relevant given the economic benefits and ecosystem services provided by these bivalves. They highlight the need for the industry to consider mitigation measures and potentially adapt their oyster growing strategies.     

Effects of chemical context on shell investigation behavior in hermit crabs

Specific chemicals in the environment evoke significant changes in the behavior of many aquatic organisms. We studied in the laboratory whether satiated individuals of the hermit crab, Pagurus longicarpus Say 1817, adjust their investigatory behavior towards an empty, optimal gastropod shell according to differences of chemical context. We also explored to what extent shell investigation by a crab in the same hunger state was affected by occupying an inadequately sized shell. Our results confirmed in part previous findings that crabs can discriminate the odor of freshly dead snails from the odor of freshly dead conspecifics. In the presence of the former odor, crabs inhabiting shells of inadequate size were more responsive and active than those in better-fitting shells. To the contrary, regardless of the quality of the inhabited shell, P. longicarpus remained practically motionless when presented with the odor of freshly dead conspecifics, possibly because the risks of incurring in predators would outweigh the benefits of acquiring a new shell. Unexpectedly, we found that crabs in both types of shell quality exhibited nearly the same behavior in control water, while crabs in adequate shells were more responsive in the presence of food odor. Individuals appeared insensitive to the odor of live snails; indeed, only one hermit crab species has been seen removing living snails from their shells. An intriguing result was that water conditioned by the odors of live conspecifics exerted a strong effect on all the individuals by inducing an intense shell investigation. Our study underlines the central role exerted by chemical detection in hermit crabs' behavior and demonstrates the existence of a complex interplay among chemical context, the physiological state of the animal, and the ecological pressures of the habitat.     

Impact scenario for the invasive red king crab <Emphasis Type="Italic">Paralithodes camtschaticus</Emphasis> (Tilesius, 1815) (Reptantia,Lithodidae) on Norwegian,native, epibenthic prey

Large invasive predators like the king crab, Paralithodes camtschaticus, deserve particular attention due to their potential for catastrophic ecological impact on recipient communities. Conspicuous, epibenthic prey species, such as the slow growing commercial scallop Chlamys islandica, are particularly exposed to the risk of local extinction. A research program integrating experiments and field monitoring is attempting to predict and track the impact of invasive king crab on scallop beds and associated fauna along the north Norwegian coast. The claw gape of the crab shows no limitations in handling the flat-bodied scallop. However, the potential impact of the crab on scallop may depend on the availability of other calcified prey associated with scallop beds, such as the sea star, sea urchin, and blue mussel, all species recorded in the diet of P. camtschaticus. To address this issue, a laboratory experiment on foraging behaviour of P. camtschaticus was conducted. The experimental results show that all size classes of red king crab prefer scallops, but small juveniles and medium sized crabs demonstrate active selection for starfish (Asterias rubens) that equals or surpasses the electivity of the large crab. The selection of sea urchin (Strongylocentrotus droebachiensis) and blue mussel (Mytilus edulis) is slightly positive or neutral for the three crab size classes. These results suggest that scallop beds with a rich associated fauna are less vulnerable to red king crabs predation and possibly more resilient than beds with few associated species. Also, crab size distribution is likely relevant for invasion impact, with increasing abundance of small and medium sized crabs being detrimental for alternative calcified prey associated with scallop beds. Successive stages of crab invasion will see an acceleration of scallop mortality rates associated with (i) decreasing availability of alternative prey, due to protracted predation pressure intensified by recruitment of juvenile crabs, and (ii) increased number of large crabs. Estimates of crab density and intake rates suggest that the accelerated loss rates will eventually endanger scallop beds persistence.     

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.     

Selective crab predation on native and introduced bivalves in British Columbia

Experiments were conducted to determine whether locally abundant crab species prefer co-occurring littleneck clams, Protothaca staminea (Conrad, 1837) and Tapes philippinarum (A. Adams and Reeve, 1850), relative to a recently introduced species, the varnish clam, Nuttallia obscurata, (Reeve, 1857). Prey preference, handling time, pick-up success, profitability and consumption rates were investigated for two crab species, Dungeness crab, Cancer magister (Dana, 1852) and red rock crab, Cancer productus (Randall, 1839) crabs. Both crab species preferred varnish clams over the native species. This may be attributable to the lower handling time, higher pick-up success and increased profitability of consuming varnish clams. Handling time appeared to be a factor not only in species preference, but also in the degree of preference, with shorter handling times corresponding to stronger preference values. Both native and introduced bivalves burrow into the substratum, with the varnish clam burrowing deepest. When feeding on clams in limited substratum both crab species preferred the varnish clam. In the unlimited substratum trials Dungeness crabs preferred varnish clams (although to a lesser degree) while red rock crabs preferred littleneck clams. This was likely due to the significantly deeper burial of the varnish clam, making it less accessible. Although the morphology (i.e. thin shell, compressed shape) of the invader increases its vulnerability to predation, burial depth provides a predation refuge. These results demonstrate how interactions between native predators and the physical characteristics and behaviour of the invader can be instrumental in influencing the success of an invasive species.     

Biogeographic variation in behavioral and morphological responses to predation risk

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

Predation by blue crabs, Callinectes sapidus, on rapa whelks, Rapana venosa: possible natural controls for an invasive species?

Blue crabs Callinectes sapidus are voracious predators in Chesapeake Bay and other estuarine habitats. The rapa whelk Rapana venosa is native to Asian waters but was discovered in Chesapeake Bay in 1998. This predatory gastropod grows to large terminal sizes (in excess of 150 mm shell length (SL)) and has a thick shell that may contribute to an ontogenetic predation refuge. However, juvenile rapa whelks in Chesapeake Bay may be vulnerable to predation by the blue crab given probable habitat overlap, relative lack of whelk shell architectural defenses, and the relatively large size of potential crab predators. Feeding experiments using three size classes of blue crab predators in relation to a size range of rapa whelks of two different ages (Age 1 and Age 2) were conducted. Blue crabs of all sizes tested consumed Age 1 rapa whelks; 58% of all Age 1 whelks offered were eaten. Age 2 rapa whelks were consumed by medium (67% of whelks offered were eaten) and large (70% of whelks offered were eaten) blue crabs but not by small crabs. The attack methods of medium and large crabs changed with whelk age and related shell weight. Age 1 whelks were typically crushed by blue crabs while Age 2 whelk shells were chipped or left intact by predators removing prey. Rapa whelks less than approximately 35 mm SL are vulnerable to predation by all sizes of blue crabs tested. Rapa whelk critical size may be greater than 55 mm SL in the presence of large blue crabs indicating that a size refugia from crab predation may not be achieved by rapa whelks in Chesapeake Bay until at least Age 2 or Age 3. Predation by blue crabs on young rapa whelks may offer a natural control strategy for rapa whelks in Chesapeake Bay and other estuarine habitats along the North American Atlantic coast.     

The invasive green crab and Japanese shore crab: behavioral interactions with a native crab species,the blue crab

Blue crabs, Callinectes sapidus (Rathbun), are an ecologically and commercially important species along the East coast of North America. Over the past century and a half, blue crabs have been exposed to an expanding set of exotic species, a few of which are potential competitors. To test for interactions with invasive crabs, juvenile C. sapidus males were placed in competition experiments for a food item with two common non-indigenous crabs, the green crab Carcinus maenas (L.) and the Japanese shore crab, Hemigrapsus sanguineus (De Haan). Agonistic interactions were evaluated when they occurred. In addition, each species’ potential to resist predators was examined by testing carapace strength. Results showed that C. maenas was a superior competitor to both C. sapidus and H. sanguineus for obtaining food, while the latter two species were evenly matched against each other. Regarding agonism, C. sapidus, was the “loser” a disproportionate number of times. C. sapidus carapaces also had a significantly lower breaking strength. These experiments suggest that both as a competitor, and as potential prey, juvenile blue crabs have some disadvantages compared with these common sympatric exotic crab species, and in areas where these exotics are common, juvenile native blue crabs may be forced to expend more energy in conflict that could be spent foraging, and may be forced away from prime food items toward less optimum prey.