Predatory blue crabs induce byssal thread production in hooked mussels

Abstract. Blue crabs (Callinectes sapidus) prey on hooked mussels (Ischadium recurvum) growing epizoically on oyster clumps in estuaries along the Louisiana coast. In prey size‐selection experiments, blue crabs preferred small mussels (<30‐mm shell length) to larger mussels, possibly because handling time increased with mussel size. When crabs were given a choice of solitary mussels versus mussels in clumps on oysters in the laboratory, mortality was lower by 86% in clumped mussels. However, no size selection by crabs occurred with mussels in clumps, likely because smaller mussels escaped predation in crevices between larger mussels or oysters. When individuals of two size classes of mussels were exposed to water containing the scent of crabs and of mussels consumed by blue crabs, an increase in byssal thread production was induced in all mussels, but byssal thread production rate was higher for small mussels than for large mussels. We conclude that increased predation risk for small mussels has resulted in higher size‐specific production of byssal threads, and that predator‐induced production of byssal threads, which may increase clumping behavior, may reduce their risk of mortality to predatory blue crabs.     

Predation by xanthid crabs on early post-settlement gastropods: the role of prey size, prey density, and habitat complexity

Small predators in marine benthic communities create a hazardous environment for newly settled invertebrates, especially for the smallest individuals. To explore the effects of predation on a newly settled gastropod, queen conch (Strombus gigas Linnaeus), by a xanthid crab (Micropanope sp.), prey size, prey density, and habitat complexity were manipulated in five laboratory experiments. All crabs >3.1 mm CW killed all conch <2 mm SL when individual crabs (<14 mm carapace width (CW)) were offered individual conch that were 2–35 days old after metamorphosis (1.2–8.8 mm shell length (SL)). Only 10% of the crabs >5.0 mm CW, however, killed conch that were >5.0 mm SL, suggesting that conch may reach a size refuge from xanthid crabs at 5 mm SL. Furthermore, when given a choice, crabs (4.8 mm CW) preferred smaller conch (2.0 mm SL) to larger (3.7 mm SL), suggesting that 1 week of additional growth in shell length is advantageous to survivorship. Proportional mortality decreased as conch density increased when crabs were offered conch at seven different densities (two to 96 individuals). Crabs proved to be effective predators regardless of the amount of seagrass structure provided in a microcosm experiment, and could consume two conch in 10 s. The high densities of xanthid crabs that occur in the wild, their effectiveness as predators, and their large appetites point to the important role that small predators may potentially play in structuring the population dynamics of their small prey immediately after settlement.     

Broken pieces: can variable ecological interactions be deduced from the remains of crab attacks on bivalve shells?

Shell fragmentation patterns that result from attacks by durophagous predators on hard‐shelled marine invertebrates are a rich source of indirect evidence that have proved useful in interpreting predation pressure in the fossil record and recent ecology. The behaviour and effectiveness of predators are known to be variable with respect to prey size. It is less well understood if variable predator–prey interactions are reflected in shell fragmentation patterns. Therefore, we conducted experimental trials to test the behavioural response of a living crab, Carcinus maenas, during successful predatory attacks on the blue mussel Mytilus edulis on two prey size categories. Further, we examined resultant shell fragments to determine whether specific attack behaviours by C. maenas could be successfully deduced from remaining mussel shells. In contrast to previous studies, we observed no significant differences in attack behaviour by the predators attributable to prey size. In most experimental predation events, crabs employed an ad hoc combination of five mechanisms of predation previously described for this species. We identified seven categories of shell breakage in predated mussels, but none of these were unambiguously correlated with specific attack behaviour. Combined attack behaviours may produce shell breakage patterns that have previously been assumed to be attributable to a single behaviour. While specific patterns of shell breakage are clearly attributable to durophagy, the results of this study provide important insights into the limitations of indirect evidence to interpret ecological interactions.     

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.     

The mechanics of predation by the shore crab,Carcinus maenas (L.), on the edible mussel,Mytilus edulis L.

Summary Mechanical aspects of predation by the shore crab, Carcinus maenas, on the edible mussel, Mytilus edulis, were examined. The shore crabs from the population studied utilized five distinct, largely size-related, mussel-opening techniques. Crushing the mussel umbone appeared the most successful opening method for medium-sized prey. Small mussels were crushed outright and large mussels could be opened by a slow, uneconomical, boring technique. The strengths of mussels, from an exposed shore, were tested under compression in four separate planes to determine the loads a crab would need to apply to crush the shells outright and the mechanical properties of mussels. Little inter-plane variability in compressive strength was observed, although intra-plane variability appeared high. The compressive strengths of mussels from a sheltered shore were found to be significantly higher than those from the exposed shore in the plane tested. A strain gauge was embedded in a mussel shell enabling the pattern and magnitude of forces produced by crab chelae in opening a mussel to be studied. The crab's chelae did not appear overwhelmingly strong when compared directly to the compressive strength of the crab's preferred mussel sizes. It is, therefore, postulated that crabs usually seek out and exploit weak spots in the umbone of mussels by trial and error, eventually breaking through the shell by a cumulative process of extending minute fractures in the shell substructure.     

The effects of prey size, predator size, and sediment composition on the rate of predation of the blue crab, Callinectes Sapidus Rathbun, on the hard clam, MercenariaMercenaria (Linné)

Sediment preferences of blue crabs, Callinectes sapidus Rathbun, and predation rates on various size classes of the hard clam, Mercenaria mercenaria (Linné), in a variety of sediment types were studied in the laboratory. Blue crabs of all size classes exhibited a preference for sand, mud, and sand/mud rather than crushed oyster shell or granite gravel. Clams were more vulnerable to predation by crabs in sand and sand/mud than in crushed oyster shell or granite gravel. When crabs were given a choice of clam sizes based on carapace width (CW), small crabs (<75 mm CW) consumed 5- and 10-mm shell length (SL) clams. Medium crabs (75–125 mm CW) preferentially consumed 10-mm SL clams. Large crabs (> 125 mm CW) consumed 10- and 25-mm SL clams equally. Blue crabs did not eat clams that were >40-mm SL.     

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.     

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

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

An invasive crab alters interaction webs in a marine community

Over the last decade, the non-native, filter-feeding crab Petrolisthes armatus invaded oyster reefs of the South Atlantic Bight at densities of thousands m⁻². Mesocosm and field experiments demonstrated that P. armatus at ∼10–75% of mean summer densities: (1) suppressed growth of small oysters, biomass of benthic microalgae, and recruitment of native mud crabs, (2) enhanced oyster, mussel, and total bivalve recruitment, macroalgal cover, and survivorship of predatory oyster drills, but (3) did not affect native taxonomic richness. Laboratory feeding assays, field tethering experiments, and population changes in field and mesocosm experiments suggest that P. armatus is a preferred prey for native mud crabs and other consumers, thus relieving predation on native species and enhancing recruitment or survival of bivalves and oyster drills. In contrast, the invasive crab can consume crustacean larvae and via this feeding may suppress recruitment of native mud crabs. Our findings should be conservative given the low densities of P. armatus seeded into experimental plots and our inability to run longer-term experiments due to controls rapidly being colonized by non-native crabs recruiting from the plankton. Invasive crabs commonly impact native communities via predation, but community impacts of this invasive crab may be as much due to its role as a preferred prey of native consumers as to its predation on native prey. Given that oysters are foundation species for shallow reefs in the South Atlantic Bight, the long-term effects of this invasion could be considerable.     

Differential vulnerability to predation in two sympatric whelks is mediated by juvenile traits

In many taxa, initial differences in offspring size play an important role in mediating subsequent performance; however, the consequences of interspecific variation in size for the performance of co‐occurring taxa have been rarely examined. We used the whelks Cominella virgata and C. maculosa, which co‐occur on rocky shores throughout their life cycles, to examine the vulnerability of early life‐stages to native predators under controlled laboratory conditions. Among all the predators evaluated (the cushion sea star Patiriella spp., the olive rockfish Acanthoclinus fuscus, the oyster borer snail Haustrum scobina, the smooth shore crab Cyclograpsus lavauxi, and the pebble crab Heterozius rotundifrons), hatchlings of both species (C. virgata: ~3 mm shell length [SL] and C. maculosa: ~1.5 mm SL) were especially vulnerable to the smooth shore crab Cy. lavauxi, the only potential predator in which mortality was greater than in the control treatment. Small shore crabs (~8 mm carapace width [CW]) were unable to eat hatchlings of either whelk species, whereas medium and large shore crabs (~12 and ~18 mm CW, respectively) consumed hatchlings of both prey species. Hatchlings of C. virgata were less vulnerable to predation by medium crabs than large ones, and those of C. maculosa were equally vulnerable to both sizes of crabs. In hatchlings of both prey species, shell length and shell thickness increased over time. Two months after hatching, only individuals of C. virgata had reached a size refuge from predation. Our results show that interspecific vulnerability to predators can be mitigated by larger sizes and thicker shells at hatching; nonetheless, our results also suggest that other species‐specific factors, such as juvenile growth rate, may also play key roles in determining the vulnerability of hatchling and juvenile snails to shell‐crushing predators.     

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.

     

Ontogenetic shift in susceptibility to predators in juvenile northern abalone, Haliotis kamtschatkana

Predation has been suggested as a major cause of juvenile mortality in benthic marine invertebrates. However, the extent to which juveniles are susceptible to predators is unknown for most species, and it remains unclear to what extent ontogenetic shifts in susceptibility to predators are common among marine invertebrates. This study examined the northern abalone Haliotis kamtschatkana, a species listed as threatened in British Columbia, Canada. Our goals were to characterize the diversity and abundance of species that prey on juvenile abalone and determine if abalone experience an ontogenetic shift in susceptibility to predators. Juvenile H. kamtschatkana were found to be susceptible to a broad variety of predators: 14 of the 37 potential predator species to which we offered juvenile abalone (≤ 28 mm shell length (SL)) consumed at least one juvenile abalone. Four of those species (three crabs and one seastar) consumed ≥ 10% of the juvenile abalone that were offered in the laboratory. These species were present at field sites where abalone are found, indicating that they have the potential to be significant predators of juvenile H. kamtschatkana in the wild. The most abundant predators were small crabs, especially Lophopanopeus bellus (black-clawed crabs) and Scyra acutifrons (sharp-nosed crabs). Juvenile H. kamtschatkana also experienced a pronounced ontogenetic shift in susceptibility to predators. The risk of predation for juvenile H. kamtschatkana decreased rapidly with increasing body size, especially over the 12–13 mm SL size range. Susceptibility remained low beyond 13 mm SL, indicating relatively low and unchanging levels of predation risk once the individual reaches this size. Although abalone are susceptible to several species during the first 1–2 years of life, predator effects on juvenile abalone abundance and microhabitat use may largely be attributable to the influence of only 1 or 2 predator species that can only kill abalone < 13 mm SL.     

Halotolerance of the oyster predator, Imogine mcgrathi, a stylochid flatworm from Port Stephens, New South Wales, Australia

The stylochid flatworm, Imogine mcgrathi was confirmed as a predator of the pteriid oyster Pinctada imbricata. Occurring at an average of 3.2 per oyster spat collector bag, the flatworms were found to consume oysters at a rate of 0.035–0.057 d^–1 in laboratory trials. Predation was affected by flatworm size with larger worms capable of consuming larger oysters and of consuming greater dry weights of oyster flesh. Irrespective of flatworm size, predation was generally confined to oysters less than 40 mm in shell height. Although all predation occurred at night, shading flatworms during the day did not significantly increase the rate of predation, but there were significant increases in the dry weight of oyster meat consumed. As a means of controlling flatworm infestations, salt, brine baths (250 g kg^–1) and freshwater baths were effective in killing I. mcgrathi. The ease of use of hyper- or hyposaline baths then encouraged assessments of I. mcgrathi halotolerance. The flatworms were exposed to solutions ranging in salinity from 0 to 250 g kg^–1for periods of from 5 min to 3 h. Despite showing both behavioural and physiological signs of stress, I. mcgrathi survived the maximum exposure time of 3 h at salinities in the range 7.5–60 g kg^–1, inclusive. Beyond this range, the duration of exposure tolerated by flatworms decreased until 0 and 250 g kg^–1, at which the flatworms no longer survived the minimum tested exposure of 5 min. Thus, despite the significant impact of other stylochids on commercial bivalves, at their current prevalence, I. mcgrathi can be controlled by exposing them to hyper- and hyposaline baths for the culture of P. imbricata in Port Stephens, NSW, Australia.     

Quantifying factors influencing American lobster (Homarus americanus) predation on the invasive green crab (Carcinus maenas) in Newfoundland,Canada

The European green crab (Carcinus maenas) is a newly invasive species in Newfoundland, where it has likely been present for ≤15 years. The green crab has been found in stomach contents of American lobster (Homarus americanus) in New England and Nova Scotia, Canada, but predation on this species has not yet been quantified in Newfoundland. We conducted feeding experiments to determine whether lobsters from Newfoundland were as likely as those from Nova Scotia (which have coexisted with green crabs for >60 years) to recognize and prey upon this new species. We also performed experiments to determine whether green crabs reach a size refuge from predation and whether factors including starvation, availability of alternate food sources, or habitat complexity would influence the probability of lobster attacking or feeding on green crabs. In our trials, lobster origin had no significant effect on crab predation; lobsters, irrespective of origin, were more likely to consume small (<40 mm carapace width [CW]) and medium (40–65 mm CW) crabs than larger (>65 mm CW) ones. Nevertheless, even small lobsters (73–76 mm carapace length, 300 g) were able to kill and consume the largest green crabs (78 mm CW, 100 g). Green crabs were less likely to be attacked or eaten when an alternative food source was present, suggesting that the lobsters were preying on the crabs, rather than simply killing them in a dispute over territory. The addition of a shelter provided a refuge for the green crabs; however, the crabs were only able to avoid being injured or eaten if this shelter was structurally complex. The green crab is slowly spreading westward around the island of Newfoundland, and so its long‐term effects, interactions with other organisms, and contribution to the diet of Newfoundland lobsters remain to be seen.     

Size-selective predation by roach (Rutilus rutilus) on zebra mussel (Dreissena polymorpha): field stuides

Summary Studies of predation by roach (Rutilus rutilus) on zebra mussel (Dreissena polymorpha) in a large, eutrophic lake showed that there was a clearly marked size threshold ( 160 mm SL) above which roach began to feed on mussels. Roach preying on various sizes of mussels selected them in proportions different from their abundance and accessibility in the habitat. The mean size of mussels ingested by roach of 220 mm and larger, which fed predominantly on Dreissena, closely followed the pattern expected for a constant ratio of mean prey size to mean predator mouth size = 0.59. To explain the size selection we applied an optimal foraging approach, based on the ability of different-sized fish to crush (cost) mussels of different sizes, and hence crushing resistance, and energy contents (benefit). We found that fish smaller than 160 mm, which showed no inclination to eat Dreissena, would only be able to take small mussels with a very high cost/benefit ratio. The real switch to Dreissena would be expected in fish of 230–240 mm that could take most of their prey from highly profitable, numerous, and easily accessible size classes while keeping the mean prey size at the optimal level relative to mean predator mouth size.     

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.     

Attack responses of the southern Australian whelk, Lepsiella vinosa (Lamarck, 1822) (Gastropoda: Muricidae), to novel bivalve prey: an experimental approach

In their native ranges, muricid gastropods feed on similar prey, often bivalves and barnacles, which they usually drill. Throughout its wide southern Australian distribution, the intertidal Lepsiella vinosa feeds on a range of prey from barnacles and littorinid gastropods in the southeast to mussels in the southwest. A number of muricids have been introduced throughout the world, either with oysters or in ship ballast water. In their new environments, they switch to native prey but their feeding responses to them have never been studied in the laboratory. The object of this study was to study the feeding of L. vinosa on a suite of non-native species. Australian Lepsiella vinosa was taken to Hong Kong, offered five different possible prey species and allowed to feed to satiation for many weeks. Replacement of consumed prey items by similar-sized conspecifics was undertaken until trends emerged. Lepsiella vinosa readily attacked thick- and thin-shelled bivalves from Hong Kong’s sandy beaches, Anomalocardia squamosa and Caecella chinensis, respectively, and from rocky shores, Septifer virgatus and Hormomya mutabilis, again respectively. It attacked them all, as it does its major prey item, Xenostrobus inconstans, in its native southwestern Australia, by either drilling or marginal probing with its proboscis. It also preferred intermediate sized prey (10–15 mm shell length), as with its natural prey. It quickly attacked the sandy shore species, and Hormomya mutabilis and Mytilopsis sallei, the latter two being closely similar in shell form and size to its natural prey X. inconstans. Hormomya mutabilis was the most favored prey, and was most similar in shell form and thickness to X. inconstans. This study therefore suggests that if introduced elsewhere, L. vinosa could radically affect intertidal community structure.     

THE INFESTATION OF MYTILUS EDULIS LINNAEUS BY POLYDORA CILIATA (JOHNSTON) IN THE CONWY ESTUARY, NORTH WALES

The incidence of the shell-boring parasite Polydora ciliatain Mytilus edulis from the Conwy estuary, North Wales is described.Amongst mussels >40 mm in shell length both the occurrenceand abundance of P. ciliata increased steeply with host size.Infestation was highest amongst mussels near the mouth of theestuary where around 60–65% of the population was parasitized;no systematic relationship between the incidence of P. ciliataand tidal elevation was observed. Mussel condition was negativelycorrelated with the degree of infestation. Moderately and heavilyinfested mussels, however, never accounted for more than ca.10% of this population and the overall detrimental effects ofP. ciliata on growth and production is thus probably quite small.Whilst heavily infested mussels were evidently more vulnerableto predation, the shore crab, Carcinus maenas, selected smaller,non-infested mussels whenever these were available. No obviousrelationships were found between the presence of P. ciliataand the incidence of either pearls or pea crabs. (Received 14 January 1991; accepted 2 April 1991)     

Prey state shapes the effects of temporal variation in predation risk

The ecological impacts of predation risk are influenced by how prey allocate foraging effort across periods of safety and danger. Foraging decisions depend on current danger, but also on the larger temporal, spatial or energetic context in which prey manage their risks of predation and starvation. Using a rocky intertidal food chain, we examined the responses of starved and fed prey (Nucella lapillus dogwhelks) to different temporal patterns of risk from predatory crabs (Carcinus maenas). Prey foraging activity declined during periods of danger, but as dangerous periods became longer, prey state altered the magnitude of risk effects on prey foraging and growth, with likely consequences for community structure (trait-mediated indirect effects on basal resources, Mytilus edulis mussels), prey fitness and trophic energy transfer. Because risk is inherently variable over time and space, our results suggest that non-consumptive predator effects may be most pronounced in productive systems where prey can build energy reserves during periods of safety and then burn these reserves as ‘trophic heat’ during extended periods of danger. Understanding the interaction between behavioural (energy gain) and physiological (energy use) responses to risk may illuminate the context dependency of trait-mediated trophic cascades and help explain variation in food chain length.     

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.