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.     

Learning in an invasive and a native predatory crab

The invasion of the green crab Carcinus maenas in the northeastern U.S. and its competition with the native blue crab Callinectes sapidus and other native crustaceans has been well-documented and researched. Various reasons for the invader’s success against native crabs have been examined (juvenile predation, food source flexibility, etc.), but another possibility is a difference in the learning ability of invasive versus native crab species. In this study, the learning ability of C. maenas and C. sapidus was tested by their increased speed in locating hidden food over successive days. The data suggest that C. maenas possesses a learning ability significantly greater than that of C. sapidus, which may partially contribute to its success.     

A comparison of the predatory impacts of an invasive and native crab species using a functional response approach

The European green crab (Carcinus maenas) is invasive on the West coast of North America, but the ecological consequences of this invasion remain poorly understood. Comparative functional response analysis has arisen as a method of elucidating ecological consequences of invasive species by comparing the impact of these species to native analogues. Through comparative functional response experiments of green crabs and native red rock crabs (Cancer productus) we found that green crab predation increased asymptotically (Type II functional response) when fed increasing densities of Pacific oysters (Magallana gigas), while red rock crab predation displayed a sigmoidal (Type III) response. At high oyster densities red rock crabs consume more Pacific oysters than green crabs do, due to their reduced handling time, though green crabs consume more Pacific oysters relative to their size than red rock crabs. However, compared to red rock crabs, green crabs consume more oysters at low prey densities, which implies that they have a larger, potentially destabilizing impact on low densities of Pacific oysters. As green crabs continue to spread across the West coast of North America, Pacific oysters will face increased predation pressure. Our results show the advantage of using functional response analysis to compare density dependent predation between an invasive species and a native species to predict the ecological consequences of invasions.

     

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.     

Cold tolerance of the invasive Carcinus maenas in the east Pacific: molecular mechanisms and implications for range expansion in a changing climate

Physiological studies have long been utilized to understand the role of environmental temperature in the distribution of native organisms within marine communities. For the invasive crab Carcinus maenas, temperature has been implicated as the main predictor of establishment success across temperate regions. Therefore, we determined whether the lower temperature tolerances of this non-native crab would restrict it from spreading farther poleward from a relatively new recipient environment. Cold tolerance capacity was determined in the laboratory by holding crabs sampled from Vancouver Island, British Columbia (BC)—near the present northern limit for the northeast Pacific metapopulation to an overwintering thermal profile generated from Sitka, Alaska, USA. These crabs were physiologically capable of overwintering north of their present range boundary. The cellular response to cold stress was investigated using two functional categories of the cellular stress response. We measured cyclin D1, a cell-cycle regulator, and Hsp70, a protein chaperone, after laboratory acclimation and acute cold stress on two populations of C. maenas from the west coast of North America that have disparate thermal histories (crabs sampled from CA or BC). We found site-specific differential expression of cyclin D1 after cold acclimation and cold shock, perhaps affecting invasion capacity in this species. Determining what physiological mechanisms are in place with respect to thermal tolerance and preference can give insight into what makes an invasive organism successful and aid in predicting probable distribution of such species within a new environment.     

Does consumer injury modify invasion impact?

Predicting the impacts of an invasive species solely by its abundance is common, yet it ignores other potentially important moderating factors. One such factor is injury. Severe injury can lead to mortality, which can directly reduce the abundance of the invader. However, more moderate, sublethal injury can also temper the impact of invasive species. Therefore, to predict impacts, it may be useful to examine not only abundance, but also moderating factors (e.g., injury) and predictors of these factors (e.g., density, size). We documented the density, injury (i.e., limb loss), and size of two conspicuous invaders, the European green crab (Carcinus maenas) and the Asian shore crab (Hemigrapsus sanguineus), at thirty sites from Shinnecock County Park, New York to Lubec, Maine. In addition, we used a field experiment to determine how injury influenced the consumption rate of mussels (Mytilus edulis) by each crab species. 31.6% of all C. maenas (1,493/4,721) and 30.7% of H. sanguineus (2,003/6,523) were missing appendage(s). Of the crabs injured, 38.4% (573/1,493) and 30.5% (611/2,003) were missing cheliped(s) for C. maenas and H. sanguineus, respectively. In our experiments, cheliped loss reduced consumption of both species on M. edulis. Injured C. maenas consumed 21% fewer mussels than uninjured crabs. Injury completely eliminated mussel consumption by H. sanguineus. Previous studies have highlighted the detrimental impacts of these two invaders on native bivalve prey. While the loss of a single cheliped can greatly reduce or even eliminate the ability of C. maenas and H. sanguineus to consume M. edulis, our results suggest that injury has a relatively minor role in reducing overall population-level impacts on prey such as mussels. However, injury on an individual-level can play a role in moderating the consumptive impacts of these invaders.     

Marine invasive species: validation of citizen science and implications for national monitoring networks

Approximately 1,000 volunteers assessed the presence of invasive (Carcinus maenas and Hemigrapsus sanguineus) and native crabs within the intertidal zone of seven coastal states of the US, from New Jersey to Maine. Identification of crab species and determination of the gender of the observed crabs was documented at all 52 sites across a 725-km coastal transect. Using quantitative measures of accuracy of data collected by citizen scientists, a significant predictor of a volunteer’s ability was determined and eligibility criteria were set. Students in grade three and seven had the ability to differentiate between species of crabs with over 80% and 95% accuracy, respectively. Determination of gender of the crabs was more challenging and accuracy exceeded 80% for seventh grade students, while 95% accuracy was found for students with at least 2 years of university education. We used the data collected by citizen scientists to create a large-scale standardized database of the distribution and abundance of the native and invasive crabs. Hemigrapsus sanguineus dominated the rocky intertidal zone from Sandy Hook, New Jersey to Boston Harbor, Massachusetts while C. maenas dominated the northern extent of the sampled coastline. A citizen scientist of this monitoring network detected a range expansion of H. sanguineus. We identified obstacles to creating a national monitoring network and proposed recommendations that addressed these issues.     

Chemically induced predator avoidance behaviour in the burrowing bivalve Macoma balthica

The responses of the burrowing bivalves Macoma balthica and Cerastoderma edule to chemical cues emitted by feeding shore crabs Carcinus maenas were investigated. M. balthica held in the laboratory and exposed to chemical signals in effluent water discharging from tanks containing C. maenas fed 20 M. balthica day^− 1 reacted by increasing their burial depths from approximately 30 mm to depths of > 60 mm, over a period of several days. When the signal was removed the bivalves gradually returned to their original depth over 5 days. C. edule similarly exposed to effluent from crabs feeding on conspecifics showed no response. In an attempt to identify the signal inducing this burrowing response, M. balthica were exposed to a variety of chemical signals. Crabs fed M. balthica elicited the strongest response, followed by crabs fed C. edule. There were also small responses to effluent from crabs fed on fish, crabs previously fed on M. balthica and to crab faeces, but no responses to starved crabs, crushed M. balthica, or controls. We conclude that increased burrowing depth of M. balthica is induced by some as yet unidentified chemical cue produced by feeding crabs and is strongest when the crabs were fed on M. balthica. Unexpectedly, neither the presence of crabs themselves, nor of damaged conspecifics, was effective in eliciting a burrowing response. The mortality rates of M. balthica and C. edule selected by crabs when burrowed at normal depths and after exposure to effluent from feeding crabs were different. Crabs selected 1.5 times more C. edule than M. balthica when both species were burrowed at their normal depths, but 15 times more after the tanks had been exposed to effluent from feeding crabs for 5 days. The burrowing response of M. balthica thus appears to reduce mortality significantly by displacing predation pressure on to the more accessible C. edule.     

Scaling the consequences of interactions between invaders from the individual to the population level

The impact of human‐induced stressors, such as invasive species, is often measured at the organismal level, but is much less commonly scaled up to the population level. Interactions with invasive species represent an increasingly common source of stressor in many habitats. However, due to the increasing abundance of invasive species around the globe, invasive species now commonly cause stresses not only for native species in invaded areas, but also for other invasive species. I examine the European green crab Carcinus maenas, an invasive species along the northeast coast of North America, which is known to be negatively impacted in this invaded region by interactions with the invasive Asian shore crab Hemigrapsus sanguineus. Asian shore crabs are known to negatively impact green crabs via two mechanisms: by directly preying on green crab juveniles and by indirectly reducing green crab fecundity via interference (and potentially exploitative) competition that alters green crab diets. I used life‐table analyses to scale these two mechanistic stressors up to the population level in order to examine their relative impacts on green crab populations. I demonstrate that lost fecundity has larger impacts on per capita population growth rates, but that both predation and lost fecundity are capable of reducing population growth sufficiently to produce the declines in green crab populations that have been observed in areas where these two species overlap. By scaling up the impacts of one invader on a second invader, I have demonstrated that multiple documented interactions between these species are capable of having population‐level impacts and that both may be contributing to the decline of European green crabs in their invaded range on the east coast of North America.     

Spontaneous alternation in marine crabs: Invasive versus native species

Organisms ranging from paramecia to humans tend to explore places that have been least recently explored, which is referred to as spontaneous alternation. Although organisms rely on different sources of information in alternating between places, the emergent behavioral pattern is likely advantageous during exploration and foraging. Under this rationale, continuous spontaneous alternation performance of the invasive green crab, Carcinus maenas was assessed and compared with the native blue crab, Callinectes sapidus in a plus-maze submerged in seawater. For the first time spontaneous alternation behavior was demonstrated in Crustacea (i.e., C. maenas) and significant interspecific differences in alternation performance were observed between the invasive versus the native species. Carcinus maneas exhibited a pronouncedly higher spontaneous alternation performance than C. sapidus. Carcinus maneas on average alternated at levels higher than chance, which was not the case for C. sapidus. These observations point to an additional behavioral mechanism that might result in the competitive success of green crabs over blue crabs in areas where they co-occur. Most of the subjects exhibited asymptotic alternation performance from the onset; there was no improvement in their performance over the course of the experimental session. This finding implies the innate nature of this behavioral policy.     

Native species vulnerability to introduced predators: testing an inducible defense and a refuge from predation

To manage the impacts of biological invasions, it is important to determine the mechanisms responsible for the effects invasive species have on native populations. When predation by an invader is the mechanism causing declines in a native population, protecting the native species will involve elucidating the factors that affect native vulnerability. To examine those factors, this study measured how a native species responded to an introduced predator, and whether the native response could result in a refuge from predation. Predation by the green crab, Carcinus maenas, has contributed to the decline in numbers of native soft-shell clams, Mya arenaria, and efforts to eradicate crabs have proven futile. We tested how crab foraging affected clam burrowing, and how depth in the sediment affected clam survival. Clams responded to crab foraging by burrowing deeper in the sediment. Clams at shallow depths were more vulnerable to predation by crabs. Results suggest soft-shell clam burrowing is an inducible defense in response to green crab predation because burrowing deeper results in a potential refuge from predation by crabs. For restoring the native clam populations, tents could exclude crabs and protect clams, but when tents must be removed, exposing the clams to cues from foraging crabs should induce the clams to burrow deeper and decrease vulnerability. In general, by exposing potential native prey to cues from introduced predators, we can test how the natives respond, identify whether the response results in a potential refuge, and evaluate the risks to native species survival in invaded communities.     

A developmental model for predicting handedness frequencies in crabs

Brachyuran crabs develop handedness in at least two different ways. Some crabs, such as Cancer productus, become heterochelous through use-induced differences in claw growth. Other crab species, for instance Carcinus maenas, appear to have a genetic predisposition towards right handedness. In this latter case, however, handedness reversal may take place following autotomy of the major claw. Thus, in C. maenas, and other species with this developmental strategy, younger cohorts are strongly biased towards right-handed individuals and the frequency of left-handedness increases with each subsequent moult. In the absence of differential mortality the ratio of left-handed to right-handed crabs in a given population should be predictable if the frequencies of right and left claw loss are known for different stages in the life history. Here, we develop a simple mathematical model for predicting handedness in crabs under the Carcinus-model of claw ontogeny and apply it to two species with very different ecologies and life histories; the green crab (Carcinus maenas (L.)) and the Trinidadian mountain crab (Eudaniela garmani (Rathburn)). The predicted and observed handedness frequencies were in complete concordance for the early intermoults of both species but significantly deviated in mature C. maenas where left-handed individuals were under-represented in the population. These results are discussed in the context of the evolution and functional significance of claw autotomy and handedness in crabs.     

Comparative brain architecture of the European shore crab <Emphasis Type="Italic">Carcinus maenas</Emphasis> (Brachyura) and the common hermit crab <Emphasis Type="Italic">Pagurus bernhardus</Emphasis> (Anomura) with notes on other marine hermit crabs

The European shore crab Carcinus maenas and the common hermit crab Pagurus bernhardus are members of the sister taxa Brachyura and Anomura (together forming the taxon Meiura) respectively. Both species share similar coastal marine habitats and thus are confronted with similar environmental conditions. This study sets out to explore variations of general brain architecture of species that live in seemingly similar habitats but belong to different major malacostracan taxa and to understand possible differences of sensory systems and related brain compartments. We examined the brains of Carcinus maenas, Pagurus bernhardus, and three other hermit crab species with immunohistochemistry against tyrosinated tubulin, f-actin, synaptic proteins, RF-amides and allatostatin. Our comparison showed that their optic neuropils within the eyestalks display strong resemblance in gross morphology as well as in detailed organization, suggesting a rather similar potential of processing visual input. Besides the well-developed visual system, the olfactory neuropils are distinct components in the brain of both C. maenas and P. bernhardus as well as the other hermit crabs, suggesting that close integration of olfactory and visual information may be useful in turbid marine environments with low visibility, as is typical for many habitats such as, e.g., the Baltic and the North Sea. Comparing the shape of the olfactory glomeruli in the anomurans showed some variations, ranging from a wedge shape to an elongate morphology. Furthermore, the tritocerebrum and the organization of the second antennae associated with the tritocerebrum seem to differ markedly in C. maenas and P. bernhardus, indicating better mechanosensory abilities in the latter close to those of other Decapoda with long second antennae, such as Astacida, Homarida, or Achelata. This aspect may also represent an adaptation to the “hermit lifestyle” in which competition for shells is a major aspect of their life history. The shore crab C. maenas, on the other hand seems to rely much less on mechanosensory information mediated by the second antennae but in water, the visual and the olfactory senses seem to be the most important modalities.     

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.     

Thermogeography predicts the potential global range of the invasive European green crab (Carcinus maenas)

Aim The highly adaptable estuarine crab (Carcinus maenas) has successfully invaded five temperate geographic regions outside of its native Europe. Here, we determine which environmental factors predict the current distribution of C. maenas and what the potential geographic range of this species might be. We also investigated whether the invasion potential of C. maenas differs with respect to the origin of a native subpopulation. Location Models were developed using global observation records of C. maenas. Methods Boosted regression trees were used to model observations from the (1) native, (2) invasive, (3) southern European, (4) northern European and (5) the combined native and invasive geographic ranges of C. maenas. Results Most established invasions were predicted mainly based on temperature. Interestingly, the environment encountered by established invasions failed to predict the majority of northern European populations; suggesting that invasion potential may differ between distinct native populations. Supporting this suggestion, a model of northern European populations, distinguished from southern European populations based on genetic structure, only predicted established invasions south of Nova Scotia. By contrast, a model of southern European populations predicted most established invasions. Main conclusions These results suggest that invasion potential depends on the European origin of an invasive population and that most invasions have arisen from southern Europe. Finally, a model based on combined native and invasive ranges of C. maenas identified potential geographic range extension along many currently invaded coastlines and the potential invasion of countries like Chile, China, Russia, Namibia and New Zealand.     

Host specificity of <Emphasis Type="Italic">Sacculina carcini</Emphasis>, a potential biological control agent of the introduced European green crab Carcinus maenas in California

The European green crab, Carcinus maenas, is an introduced marine predator established on the west coast of North America. We conducted laboratory experiments on the host specificity of a natural enemy of the green crab, the parasitic barnacle Sacculina carcini, to provide information on the safety of its use as a possible biological control agent. Four species of non-target, native California crabs (Hemigrapsus oregonensis, H. nudus, Pachygrapsus crassipes and Cancer magister) were exposed to infective larvae of S. carcini. Settlement by S. carcini on the four native species ranged from 33 to 53%, compared to 79% for green crabs. Overall, cyprid larvae tended to settle in higher numbers on individual green crabs than on either C. magister or H. oregonensis. However, for C. magister this difference was significant for soft-shelled, but not hard-shelled individuals. Up to 29% of the native crabs arrested early infections by melanizing the rootlets of the parasite. Most native and green crabs settled on by S. carcini became infected, especially when settled on by >3 cyprids. Infected green crabs died at more than twice the rate of uninfected green crabs. In contrast to green crabs, all infected native crabs died without producing an externa (reproductive sac). At high settlement intensities, infected native crabs frequently exhibited neurological symptoms (twitching, loss of movement) before death. These results indicate that use of S. carcini as a biological control agent could result in the death of native crabs. The magnitude of this effect would be proportional to the density of infected green crabs in the environment and the probability that cyprids would contact native crabs in the wild. Potential benefits of biological control should be assessed in relation to these potential non-target effects.     

Short‐term effect of hypoxia on ammonia excretion and respiration rates in the crab carcinus maenas

The metabolic response of the crab Carcinus maenas to short‐term hypoxia (60% and 35% saturated seawater) was studied at 17.5°C in fed, 3 day‐unfed and 6 day‐unfed crabs.

Ammonia excretion rate decreased under hypoxia: a 40% and 45% decrease in the normoxic rate was observed in fed crabs at 35% saturation and in 3 day‐unfed crabs at both hypoxic levels respectively. In the 6 day‐unfed crabs, the effect of hypoxia was concealed by the effect of starvation.

Oxygen consumption rate was directly related to the external O₂ tension irrespective of the crab's nutritional state. Stressed crabs behaved as a whole, as oxygen‐conformers.

A strong relationship was observed between ammonia excretion and oxygen consumption rates in fed crabs under hypoxia but not in starved crabs.     

Effects of acclimation and acute temperature change on specific dynamic action and gastric processing in the green shore crab, Carcinus maenas

The effects of temperature acclimation and acute temperature change were investigated in postprandial green shore crabs, Carcinus maenas. Oxygen uptake, gut contractions and transit rates and digestive efficiencies were measured for crabs acclimated to either 10 °C or 20 °C and subsequently exposed to treatment temperatures of 5, 15, or 25 °C. Temperature acclimation resulted in a partial metabolic compensation in unfed crabs, with higher oxygen uptake rates measured for the 10 °C acclimated group exposed to acute test temperatures. The Q₁₀ values were higher than normal, probably because the acute temperature change prevented crabs from fully adjusting to the new temperature. Both the acclimation and treatment temperature altered the characteristics of the specific dynamic action (SDA). The duration of the response was longer for 20 °C acclimated crabs and was inversely related to the treatment temperature. The scope (peak oxygen consumption) was also higher for 20 °C acclimated crabs with a trend towards an inverse relationship with treatment temperature. Since the overall SDA (energy expenditure) is a function of both duration and scope, it was also higher for 20 °C acclimated crabs, with the highest value measured at the treatment temperature of 15 °C. The decline in total SDA after acute exposure to 5 and 25 °C suggests that both cold stress and limitations to oxygen supply at the temperature extremes could be affecting the SDA response. The contractions of the pyloric sac of the foregut region function to propel digesta through the gut, and contraction rates increased with increasing treatment temperature. This translated into faster transit rates with increasing treatment temperatures. Although pyloric sac contractions were higher for 20 °C acclimated crabs, temperature acclimation had no effect on transit rates. This suggests that a threshold level in pyloric sac contraction rates needs to be reached before it manifests itself on transit rates. Although there was a correlation between faster transit times and the shorter duration of the SDA response with increasing treatment temperature, transit rates do not make a good proxy for calculating the SDA characteristics. The digestive efficiency showed a trend towards a decreasing efficiency with increasing treatment temperature; the slower transit rates at the lower treatment temperatures allowing for more efficient nutrient absorption. Even though metabolic rates of 10 °C acclimated crabs were higher, there was no effect of acclimation temperature on digestive efficiency. This probably occurred because intracellular enzymes and digestive enzymes are modulated through different control pathways. These results give an insight into the metabolic and digestive physiology of Carcinus maenas as it makes feeding excursions between the subtidal and intertidal zones.     

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.     

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.