Can postlarval bivalves select sediment type during settlement? A field test with Macoma balthica (L.) and Cerastoderma edule (L.)

Many soft-bottom benthic invertebrates display sediment-associated patterns of dispersion. Habitat selection experiments have shown that the larvae of some species can choose to settle in favourable habitats, and this process could establish patterns observed in the field. However, many soft-bottom infauna, including the bivalves Cerastoderma edule and Macoma balthica, show postlarval relocation. Such movements could obviate patterns established at initial settlement, and active habitat selection in these postlarval stages may be responsible for adult distributions. We investigated sediment selection in postlarval stages of these bivalves on an intertidal mudflat using a field survey and experiment. Our 1225 m² grid survey showed a strong spatial correlation between the smallest individuals of both species, which showed no correlation with sediment characteristics. Larger size groups showed increasingly strong associations with sediment type: C. edule with sandy sediments and M. balthica with muddy. In our sediment choice experiment, the numbers of bivalves recruited into muddy, sandy and muddy depression (to detect small-scale hydrodynamic effects) treatments after 2 days in the field were recorded. There were no significant treatment effects in either C. edule or M. balthica. All bivalves were too large to be initial settlers; they were thus relocating individuals. Neither the survey nor the experiment provided any evidence that relocating individuals of these two species could choose the sediment in which they alight. The sediment-related patterns observed in adults of these species must therefore be due to other mechanisms, possibly postsettlement predation by Carcinus maenas and/or an ability to leave less-preferred sediments in a series of migrations until the preferred habitat is found.     

Behavioral plasticity of the soft-shell clam, Mya arenaria (L.), in the presence of predators increases survival in the field

Soft-shell clams, Mya arenaria, are sessile, suspension-feeding bivalves that are preyed upon by the exotic green crab, Carcinus maenas. Clams evade crab consumers by burrowing deeper into the sediment after perceiving a threat from a nearby predator. The purpose of this study was to determine the types of signals that M. arenaria use to detect predators and the types of behaviors clams use to avoid being eaten. In a field study, clams increased their burial depth in the presence of green crab predators consuming conspecifics that were caged nearby, and also increased burial depth after artificial tactile stimulation in the laboratory assay. These results indicate that clams can use chemical and mechanical cues to detect potential predatory threats. We performed a field study to examine the difference in survivability of clams that had burrowed deeper into the sediment in response to predators vs. control clams that were burrowed less deeply. Significantly higher survival rates were observed in clams that had initially burrowed more deeply, suggesting that increasing burial depth is a valid predator avoidance strategy. Some bivalves also alter their pumping rates in the presence of predators, making them less apparent and providing more structural defense by covering soft tissue, and we measured pumping time of soft-shell clams in the presence and absence of predators, when burrowing was not an option for escape. Soft-shell clams did not alter their pumping time in the presence of green crab predators, possibly because they employ a burrowing method called “hydraulic” or “jet-propelled” burrowing, where it is necessary for the clam to pump in order to burrow. Chemical signals and tactile cues instigated behavioral changes in M. arenaria, and this change in behavior (increasing burial depth) increased clam survival in the field.     

Evaluation of the sample size used for the rapid bioassessment of rivers using macroinvertebrates

Experiments were designed to investigate selective predation by medium (40–55 mm carapace width: CW) and large (55–70 mm CW) Carcinus maenas when feeding on four bivalves of contrasting shell morphology. Size-selection was examined by presenting individual crabs with a wide size range of Mytilus edulis, Ostrea edulis, Crassostrea gigas and Cerastoderma edule. Medium-sized crabs preferred mussels 5–15 mm shell length (maximum shell dimension: SL) and cockles 5–10 mm SL, whereas large crabs preferred mussels 15–25 mm and cockles 10–20 mm SL. Crabs generally showed no preference for any particular size of either oyster species. Species-selection was examined by presenting individual crabs with paired combinations of the four bivalves in various proportions. When offered mussels and oysters simultaneously, both size categories of crabs consistently selected mussels, and food choice was independent of prey relative abundance. By contrast, C. maenas selected mussels and cockles as expected by the frequency in which each size category of crab encountered the preferred size ranges of prey. Crab preference clearly paralleled the rank order of prey profitability, which in turn was mainly determined by prey biomass, suggesting that active selection takes place at some point of the predation cycle. Experiments with epoxy resin models showed that initial reluctance of crabs to attack oysters was not associated with the ultimate energy reward. Moreover, they suggest that foraging decisions are partly based on evaluations of overall prey shape and volume, and that the minimum dimension of the shell constitutes an important feature which crabs recognise and associate with prey value.     

Activity patterns, feeding and burrowing behaviour of the crab Ucides cordatus (Ucididae) in a high intertidal mangrove forest in North Brazil

Activity patterns, feeding and burrowing behaviour of the economically important semi-terrestrial mangrove crab Ucides cordatus (Ucididae, L. 1763) was studied in a high intertidal Rhizophora mangle forest stand in Bragança, North Brazil. Video observations in the rainy and dry season were conducted over 24 h cycles at different lunar phases to investigate the behaviour of these litter-feeding crabs outside their burrows. During the rainy season, crabs stayed inside their burrows for 79% and 92% of the time during day and night, respectively. Time spent for feeding, burrowing and other activities outside their burrows was significantly longer during the day with 9.9% (night: 1.7%) and at waning and waxing moon with 9% (full and new moon: 0.9%). At neap tides (no tidal inundation) foraging and feeding activities outside burrows were clearly light-dependent, increasing at dawn and decreasing at dusk. Highest activities during daytime relate to the visual localisation of food. During the dry season, crabs spent less time inside burrows at neap tides than during the rainy season (80% and 91%, respectively). However, time spent for feeding activities was similar during both seasons. During almost all observation periods crabs collected leaf litter, but rarely fed on it outside burrows. At neap tides nearly all available litter was collected, suggesting that the U. cordatus population is litter-limited during these times. At spring tides (regular tidal inundation) the surface activity of U. cordatus was tide-dependent. Crabs closed their burrow entrances 2-3 h before flooding and re-emerged as soon as the tide retreated. During the day, burrow maintenance was the second most frequent behaviour after feeding. Agonistic interactions were regularly observed and were mainly related to burrow defence. The mean foraging radius of the crabs was only 19 cm (max: 1 m) underneath high Rhizophora mangle trees where crab densities were high. The results point to a high competition for burrows and show that U. cordatus is territorial. It is concluded that several exogenous factors, in particular light, leaf litter availability, flooding of burrows and the presence of conspecifics are important in controlling the crabs' activity patterns.     

Effect of dietary polyunsaturated fatty acids on reproductive output and larval growth of bivalves

The pre-spawning condition of adult bivalves is influenced by quantity and quality of available food. For bivalves, the essential polyunsaturated fatty acids (PUFAs) eicosapentaenoic acid (EPA) 20:5(n-3) and docosahexaenoic acid (DHA) 22:6(n-3) are presumed to determine the nutritional value of algae. Macoma balthica kept on a broodstock diet supplemented with PUFAs spawned a larger number of eggs (average 22220) per female and larger sized eggs (106.7 μm) compared to adults kept on a diet without PUFA supplementation (962.5 eggs with average size 99.8 μm).Larvae of M. balthica from the same parental pool however did not profit from a diet where a part was replaced with PUFA spheres. Instead, larvae reared on Isochrysis sp. showed lower mortality and higher growth rates than larvae fed on the same algae supplemented with lipid spheres. Crassostrea gigas larvae showed no clear response to a PUFA supplemented diet.     

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.     

Physiological responses of three crustacean species to infection by the dinoflagellate-like protist Hematodinium (Alveolata: Syndinea)

This is the first study comparing physiological responses of three decapod species to infection by parasites of the genus Hematodinium, which belongs to the dinoflagellate-like Syndinea. Responses varied profoundly between the crabs Carcinus maenas and Cancer pagurus (Brachyura), but also differed to those of hermit crabs, Pagurus bernhardus (Anomura). Osmoregulatory capacity was reduced significantly in Hematodinium-infected C. maenas, haemolymph pH increased in parasitised C. pagurus and P. bernhardus, and L-lactate concentration decreased in infected P. bernhardus. Changes to tissues and exoskeletons were observed in C. pagurus, but not in C. maenas and P. bernhardus.     

Biodiversity of benthic invertebrates and organic matter processing in shallow marine sediments: an experimental study

The main objective of this study was to measure the impact of benthic invertebrate diversity on processes occurring at the water-sediment interface. We analyzed the effects of interactions between three shallow water species (Cerastoderma edule, Corophium volutator, and Nereis diversicolor). The impacts of different species richness treatments were measured on sediment reworking, bacterial characteristics, and biogeochemical processes (bromide fluxes, O₂ uptake, nutrient fluxes, and porewater chemistry) in sediment cores. The results showed that the three species exhibited different bioturbation activities in the experimental system: C. edule acted as a biodiffusor, mixing particles in the top 2 cm of the sediments; C. volutator produced and irrigated U-shaped tubes in the top 2 cm of the sediments; and N. diversicolor produced and irrigated burrow galleries in the whole sediment cores. C. edule had minor effects on biogeochemical processes, whereas the other species, through their irrigation of the burrows, increased the solute exchange between the water column and the sediment two-fold. These impacts on sediment structure and solute transport increased the O₂ consumption and the release of nutrients from sediments. As N. diversicolor burrowed deeper in the sediment than C. volutator, it irrigated a greater volume of sediments, with great impact on the sediment cores.Most treatments with a mixture of species indicated that observed values were often lower than predicted values from the addition of the individual effects of each species, demonstrating a negative interaction among species. This type of negative interaction measured between species on ecosystem processes certainly resulted from an overlap of bioturbation activities among the three species which lived and foraged in the same habitat (water-sediment interface). All treatments with N. diversicolor (in isolation and in mixture) produced similar effect on sediment reworking, water fluxes, nutrient releases, porewater chemistry, and bacterial characteristics. Whichever species associated with N. diversicolor, the bioturbation activities of the worm hid the effect of the other species. The results suggest that, in the presence of several species that use and modify the same sediment space, impact of invertebrates on ecosystem processes was essentially due to the most efficient bioturbator of the community (N. diversicolor). In consequence, the functional traits (mode of bioturbation, depth of burrowing, feeding behaviour) of an individual species in a community could be more important than species richness for some ecosystem processes.     

Metabolic responses of sand fiddler crabs, Uca pugilator, in northwest Florida to seasonal temperature change

Metabolic responses of sand fiddler crab, Uca pugilator, populations in northwest Florida are greatly influenced by seasonal temperature fluctuations. Crabs acclimated at 20 °C and immediately transferred to either 14 or 26 °C produced an acute metabolic response with respective temperature quotient (Q₁₀) values of 3.46 and 3.91. Crabs acclimated at 10 and 20 °C exhibited a Q₁₀ of 2.62 indicating a partial compensation response. A brumation (reverse) response (Q₁₀ value of 20.11) was observed for acclimated crabs between 5 and 10 °C. Brumation is advantageous during winter when food supplies are scarce and crabs must survive extensive periods of inactivity.     

The functional group approach to bioturbation: The effects of biodiffusers and gallery-diffusers of the Macoma balthica community on sediment oxygen uptake

The Macoma balthica community, which is widely distributed in intertidal soft sediments bordering the north Atlantic, is dominated by two functional groups with different sediment mixing modes: the biodiffusers M. balthica and Mya arenaria and the gallery-diffuser Nereis virens. To compare the effects of these two groups on sediment oxygen uptake rates, we used experimental microcosms with identical biovolumes to measure the influence of each species on oxygen uptake. The two biodiffusers had similar effects on oxygen uptake in spite of different space occupation and different feeding, ventilation and burrowing modes. Biodiffusers and gallery-diffusers had different effects on oxygen uptake. Periodic ventilation by the gallery-diffusers stimulated the oxygen uptake by the sediment more than the steady activities of the biodiffusers. Temporal variation in oxygen fluxes in bioturbated microcosms was linked to construction and maintenance of biogenic structures. The results confirm that the functional group approach to bioturbation is a useful tool for quantifying the effects of intertidal benthic communities on benthic fluxes.     

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.     

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.     

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.     

Immunological detection of winter flounder (Pseudopleuronectes americanus) eggs and juveniles in the stomach contents of crustacean predators

Predation on the early life history of fish is an important factor regulating year-class strength. Verifying predation events, however, is difficult when analyses rely on visually identifying the remnants of partially digested fish in the stomachs of suspected predators. The objective of this study was to assess the utility of using immunological assays to detect the presence of winter flounder eggs and juveniles (Pseudopleuronectes americanus) in the gut contents of sand shrimp (Crangon septemspinosa) and green crab (Carcinus maenas). After defining assay capabilities, the stomach contents of field-collected shrimp and crabs were examined to determine if these predator-prey relationships occur under natural conditions. Winter flounder-specific antisera developed and used in this study successfully identified homologous antigens (egg or juvenile flounder extracts) without appreciably cross-reacting with antigenic material from predators or nontarget prey. Moreover, antisera detected flounder eggs 10.8-16.4 h after initial feeding by various sized shrimp, and identified juvenile flounder 9.4 and 7.8 h after initial ingestion by shrimp and crabs, respectively. Immuonological dietary analysis of decapod crustaceans collected from Niantic River, Connecticut, revealed that C. septemspinosa and C. maenas are potentially important predators on the early life stages of winter flounder. The temporal trends and magnitude of flounder predator-induced mortality was affected primarily by the spatial and temporal overlap between predator and prey (egg mortality), and the size-dependent relationships underlying crustacean and flatfish predator-prey interactions (juvenile mortality).     

You are what you eat: diet-induced chemical crypsis in a coral-feeding reef fish

The vast majority of research into the mechanisms of camouflage has focused on forms that confound visual perception. However, many organisms primarily interact with their surroundings using chemosensory systems and may have evolved mechanisms to ‘blend in’ with chemical components of their habitat. One potential mechanism is ‘chemical crypsis'' via the sequestration of dietary elements, causing a consumer''s odour to chemically match that of its prey. Here, we test the potential for chemical crypsis in the coral-feeding filefish, Oxymonacanthus longirostris, by examining olfactory discrimination in obligate coral-dwelling crabs and a predatory cod. The crabs, which inhabit the corals consumed by O. longirostris, were used as a bioassay to determine the effect of coral diet on fish odour. Crabs preferred the odour of filefish fed their preferred coral over the odour of filefish fed a non-preferred coral, suggesting coral-specific dietary elements that influence odour are sequestered. Crabs also exhibited a similar preference for the odour of filefish fed their preferred coral and odour directly from that coral, suggesting a close chemical match. In behavioural trials, predatory cod were less attracted to filefish odour when presented alongside the coral it had been fed on, suggesting diet can reduce detectability. This is, we believe, the first evidence of diet-induced chemical crypsis in a vertebrate.     

Parasite infection and sand coarseness increase sand crab (Emerita analoga) burrowing time

Parasites with indirect life cycles require trophic transmission from intermediate hosts to definitive (vertebrate) hosts. Transmission may be facilitated if parasite infection alters the behavior of intermediate hosts such that they are more vulnerable to predation. Vulnerability to predation may also be influenced by abiotic factors; however, rarely are the effects of parasites and abiotic factors examined simultaneously. The swash zone of sandy beaches is a particularly harsh environment. Sand crabs (Emerita analoga) burrow rapidly in the swash zone to avoid predators and dislodgment. We examined prevalence and abundance of the acanthocephalan parasite Profilicollis altmani in sand crabs, and investigated the synergistic effects of sand grain size (an important abiotic factor), parasite infection, body size and reproductive condition on burrowing speed in females, from three California sites. More heavily parasitized crabs burrowed more slowly, making them potentially more vulnerable to predation by marine bird definitive hosts. Ovigerous females harbored more parasites than non-ovigerous females, but burrowed more quickly. All crabs burrowed slowest in the coarsest sand, and burrowing times increased with repeated testing, suggesting that it is energetically costly. Abiotic and biotic factors influence burrowing, and behavioral variation across sites may reflect the response to natural variation in these factors.     

Brown shrimp (Crangon crangon, L.) functional response to density of different sized juvenile bivalves Macoma balthica (L.)

Variability in infaunal bivalve abundance in the Wadden Sea is largely determined by recruitment variability. Post-settlement, but pre-recruitment bivalve mortality is high and related to the occurrence of their most abundant predator, the brown shrimp Crangon crangon. To investigate if the mortality patterns of newly settled bivalves can be explained by the foraging behavior of brown shrimp, we carried out experiments on shrimp functional response to three size classes of juveniles of the Baltic Tellin Macoma balthica. The functional response curves for all three prey sizes (0.62 mm, 0.73 mm, and 0.85 mm) were the hyperbolic Holling's type II. The attack rate was highest for the smallest prey size (a = 0.31, medium and large prey a = 0.22); the handling time was longest for the largest prey size (T_h = 29 s, small and medium prey T_h = 15 s). Thus, a large body size is advantageous for the bivalves over the whole density range. Knowledge of individual foraging behavior is needed to model predation mortality of bivalves. The consumption rates in the experiment were theoretically high enough to account for M. balthica mortality in the field.     

Biotic resistance experienced by an invasive crustacean in a temperate estuary

Communities high in species diversity tend to be more successful in resisting invaders than those low in species diversity. It has been proposed that the biotic resistance offered by native predators, competitors and disease organisms plays a role. In Yaquina Bay, Oregon, we observed very little overlap in the distribution of the invasive European green crab, Carcinus maenas, and the larger red rock crab, Cancer productus. C. productus dominates the more saline, cooler lower estuary and C. maenas, the less saline, warmer upper estuary. Because caged C. maenas survive well in the lower estuary, we decided to test the hypothesis that C. productus prey on C. maenas and thus contribute to their exclusion from the more physically benign lower estuary. A laboratory species interaction experiment was designed to determine whether C. productus preys on smaller C. maenas at a higher rate than on smaller crabs of their own species. Crabs of both species were collected and sorted by weight into three size classes: small, medium and large. Small and medium crabs of both species were paired with C. maenas and C. productus of various sizes. When conspecifics were paired, mortality was less than 14%, even in the presence of larger crabs. Smaller C. productus survived well in the presence of larger C. maenas, but the reverse was not true. When small C. maenas (60–67 mm carapace width) were matched with medium and large C. productus, their mortality increased to 52% and 76%, respectively. A less dramatic pattern was observed for medium C. maenas (73–80 mm) in the presence of medium and large C. productus. Thus on the West Coast of North America, the more aggressive red rock crab, C. productus, has the potential to reduce the abundance of C. maenas in the more saline and cooler lower estuaries.     

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.