Post-settlement effects of habitat type and predator size on cannibalism of glaucothoe and juveniles of red king crab Paralithodes camtschaticus

Postlarval (glaucothoe) and juvenile (first crab stage, C1) red king crab Paralithodes camtschaticus actively select structurally complex substrata for settlement. Such habitats may provide them with shelter from predation during critical early stages. We tested this hypothesis by placing glaucothoe and juvenile crab in aquaria with or without natural or artificial habitats, and with or without predators (1-3-year-old red king crab) of two different sizes. Predators caused increased mortality of glaucothoe, but predator size, habitat presence and habitat type had no effect on survival. Predators caused significant mortality of C1 crabs in the absence of habitat, and mortality was inversely related to predator size. Density of glaucothoe on habitats was similar with or without predators, but density of C1 crab on habitats was higher than that of glaucothoe, and increased in the presence of large predators. Active selection for complex substrata by settling glaucothoe does not reduce cannibalism, but may pre-position them for improved survival after metamorphosis. In contrast, juvenile crabs modify their behavior to achieve higher densities in refuge habitats, which tends to dampen the effect of predation. These survival strategies may have evolved to compensate for the greater risk of predation in open habitats.     

The competitive and predatory impacts of the nonindigenous crab Carcinus maenas (L.) on early benthic phase Dungeness crab Cancer magister Dana

We evaluate the potential competitive and predatory impacts of nonindigenous European green crab Carcinus maenas on native Dungeness crab Cancer magister in the northeast Pacific. The coastal estuaries of Washington State, USA, provide appropriate habitat for recently introduced green crab, yet these areas are important nursery grounds for Dungeness crab and contribute greatly to the coastal crab fishery. Juvenile Dungeness crabs are dependent on limited intertidal epibenthic shell for refuge habitat during early benthic life and experience increased mortality on open sand and mud as a result of predation by fish and birds. Early juveniles throughout the subtidal are similarly at risk due to predation by fish and especially adult conspecifics. Laboratory experiments and infrared video observations revealed that juvenile green crab displace Dungeness crab of equal size from shelters during one-on-one competition. Green crab also consistently win nocturnal foraging trials in which the species compete for fresh, damaged clams. Field and laboratory enclosure experiments show that juvenile Dungeness crab emigrate from oyster shell habitat as a result of competition and predation by adult green crab. Depending on the extent to which the two species overlap, interactions with the dominant nonindigenous species could have a negative influence on juvenile Dungeness crab survival and could conceivably impact recruitment to the fishery. However, current evidence indicates that the distribution of green crab in Washington State is far removed from nursery areas of Dungeness crab.     

Molecular keys unlock the mysteries of variable survival responses of blue crabs to hypoxia

Hypoxia is a major stressor in coastal ecosystems, yet generalizing its impacts on fish and shellfish populations across hypoxic events is difficult due to variability among individuals in their history of exposure to hypoxia and related abiotic variables, and subsequent behavioral and survival responses. Although aquatic animals have diverse physiological responses to cope with hypoxia, we know little about how inter-individual variation in physiological state affects survival and behavioral decisions under hypoxic conditions. Laboratory experiments coupled with molecular techniques determined how extrinsic factors (e.g., water body and temperature) and respiratory physiology (hemocyanin concentration and structure) affected survival and behavior of adult blue crabs (Callinectes sapidus) exposed to different levels of hypoxia over a 30-h time period. Nearly 100% of crabs survived the 1.3 mg dissolved oxygen (DO) l^?1 treatment (18.4% air saturation), suggesting that adult blue crabs are tolerant of severe hypoxia. Probability of survival decreased with increasing hypoxic exposure time, lower DO, and increasing temperature. Individual-level differences in survival correlated with water body and crab size. Crabs collected from the oligo/mesohaline and hypoxic Neuse River Estuary (NRE), North Carolina, USA survived hypoxic exposures longer than crabs from the euhaline and normoxic Bogue and Back Sounds, North Carolina. Furthermore, small NRE crabs survived longer than large NRE crabs. Hemocyanin (Hcy) concentration did not explain these individual-level differences, however, hypoxia-tolerant crabs had Hcy structures indicative of a high-O₂-affinity form of Hcy, suggesting Hcy “quality” (i.e., structure) may be more important for hypoxia survival than Hcy “quantity” (i.e., concentration). The geographic differences in survival we observed also highlight the importance of carefully selecting experimental animals when planning to extrapolate results to the population level.     

Predation risk predicts use of a novel habitat

Predator–prey interactions are often highly co‐evolved, with selection over time for prey with morphological and behavioral traits that minimize predation risk. Consequently, in many environments prey choose among potential habitats according to their refuge value. It is unclear, however, when presented with new habitats, if prey are able to evaluate the predation risk of these relative to familiar habitats and utilize these in accordance with their value. We tested whether, along the east coast of the USA, native mud crabs Panopeus herbstii utilize the non‐native alga Gracilaria vermiculophylla according to its relative refuge value. Experiments examining predation by blue crabs Callinectes sapidus on mud crabs revealed that the non‐native alga had an intermediate refuge value relative to native oysters, which were the most protective, and unvegetated sediment, which was the least. In subsequent choice experiments, mud crabs selected oysters over alga over unvegetated sediment, in accordance with habitat refuge values. Further, in field experiments, the use of Gracilaria by mud crabs was inversely related to the proximity of the alga to the preferred habitat type, oysters, and was reduced by the presence of a blue crab predator. Consequently, mud crabs are utilizing the non‐native alga Gracilaria in accordance with its intermediate refuge value. The relative refuge value of non‐native vs native habitat‐forming species may provide a baseline expectation against which to measure the speed of learning and opportunism in the response of native prey to novel protective habitats.     

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

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

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.     

The relative importance of habitat-specific settlement, predation and juvenile dispersal for distribution and abundance of young juvenile shore crabs Carcinus maenas L.

Young juveniles of many motile benthic species are concentrated in structurally complex habitats, but the proximate causes of this distribution are usually not clear. In the present study, I assessed three potentially important processes affecting distribution and abundance of early benthic stages in the shore crab (Carcinus maenas): (1) selection of habitat by megalopae (postlarvae); (2) habitat-specific predation; and (3) post-settlement movements by juveniles. These processes were assessed concurrently over 3-9 days at two spatial scales: at the scale of square meters using cage techniques within nursery areas, and at the scale of hectares using isolated populations of juvenile shore crabs in small nursery areas as mesocosms. The results were compared to habitat-specific distribution in the field.Shore crab megalopae and first instar juveniles (settlers) were distributed non-randomly among micro-habitats in the assessed nursery areas, with great densities in both mussel beds, eelgrass and filamentous algal patches (on average 114-232 settlers m⁻²), and significantly smaller densities on open sand habitats at all times (on average 4 settlers m⁻²). The same habitat-specific settlement pattern was found in cages where predators were excluded, suggesting that active habitat selection at settlement was responsible for the initial distribution. Older juveniles (second to ninth instar crabs) were also sparse on sand, but in contrast to settlers, were concentrated in mussel beds, which showed significantly greater densities than eelgrass and algal habitats. The cage experiment demonstrated a dynamic distribution of juvenile crabs. Young juveniles constantly migrated over open sand habitats (20 m or further) and colonized the experimental plots in a habitat-specific pattern that reflected the distribution in the field. This pattern was also found for very small crabs colonizing predator-exclusion cages, suggesting that selection of habitat by migrating juveniles caused the ontogenetic change in habitat use. Although post-settlement movements were great within nursery areas, juvenile dispersal at a regional scale appeared to be small, and the recruitment of juvenile shore crabs to the shallow bays occurred mainly through pelagic megalopae.Conservative estimates at the scale of whole nursery areas, based on migration trap data and field samples, indicated great mortality of settlers and early benthic stages of shore crabs. Results from the cage experiment suggest that predation by crabs and shrimp were responsible for the high settlement mortality. Both enclosed cannibalistic juvenile crabs and local predators on uncaged habitat plots caused significant losses of settlers in all habitats (on average 22% and 64% 3 day⁻¹, respectively). The effect of predators was highly variable between trials, but differed little between habitat types, and predation had no detectable proximate effect on juvenile distribution, despite the great losses. Small settlement densities on sand habitats in combination with a refuge at low prey numbers, and an aggregation of cannibalistic juvenile crabs in nursery habitats appear to decrease the effect of habitat-specific predation rates on the distribution of juvenile shore crabs. This study demonstrates that active habitat selection at settlement followed by a dynamic redistribution of young juveniles can be the proximate processes responsible for habitat-specific distribution of epibenthic juveniles, and indicate that predation represents a major evolutionary process reinforcing this behavior.     

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.     

Diel variation in predator abundance, predation risk and prey distribution in shallow-water estuarine habitats

Predation by visual predators is often affected by light conditions and may therefore exhibit strong diel variation. The dominant predators on grass shrimp, Palaemonetes pugio, are finfish predators that are thought to locate their prey by visual cues. We examined the response of grass shrimp to diel variation in predation risk in the nearshore shallow waters of the Chesapeake Bay. We used diel shoreline seines to assess the relative abundance of predators. We assessed the relative risk of predation with shrimp tethered at refuge (30 cm) and nonrefuge (60 cm) depths. To measure grass shrimp response to predation risk, we used dipnets to monitor habitat use. Four predominantly visual predators dominated the shoreline seine catches, Fundulus heteroclitus, Micropogonias undulatus, Morone americana and Morone saxatilis. Total predator abundance had a diel component, with dramatic nighttime decreases in total abundance, whereas guild composition and relative abundance remained unchanged. Relative predation risk for tethered shrimp exhibited significant time by habitat interaction. During the day, depth negatively affected survivorship of tethered shrimp while at night overall survivorship increased and there was no effect of depth. Shrimp habitats use reflected diel predation risks. Abundances in the near shore were highest during the day with decreased abundances at night. Together, the seine and tethering data highlight the importance for a refuge (e.g., shallow water) from predation during the daytime and a relaxation of predation pressure at night.     

Effects of seagrass habitat fragmentation on juvenile blue crab survival and abundance

Seagrasses form temporally dynamic, fragmented subtidal landscapes in which both large- and small-scale habitat structure may influence faunal survival and abundance. We compared the relative influences of seagrass (Zostera marina L.) habitat fragmentation (patch size and isolation) and structural complexity (shoot density) on juvenile blue crab (Callinectes sapidus Rathbun) survival and density in a Chesapeake Bay seagrass meadow. We tethered crabs to measure relative survival, suction sampled for crabs to measure density, and took seagrass cores to measure shoot density in patches spanning six orders of magnitude (ca. 0.25-30,000 m²) both before (June) and after (September) seasonally predictable decreases in seagrass structural complexity and increases in seagrass fragmentation. We also determined if juvenile blue crab density and seagrass shoot density varied between the edge and the interior of patches. In June, juvenile blue crab survival was not linearly related to seagrass patch size or to shoot density, but was significantly lower in patches separated by large expanses of unvegetated sediment (isolated patches) than in patches separated by <1 m of unvegetated sediment (connected patches). In September, crab survival was inversely correlated with seagrass shoot density. This inverse correlation was likely due to density-dependent predation by juvenile conspecifics (i.e. cannibalism); juvenile blue crab density increased with seagrass shoot density, was inversely correlated with crab survival, and was greater in September than in June. Shoot density effects on predator behavior and on conspecific density also likely caused crab survival to be lower in isolated patches than in connected patches in June. Isolated patches were either large (patch area >3000 m²) or very small (<1 m²). Large isolated patches had the lowest shoot densities, which may have allowed predators to easily find tethered crabs. Very small isolated patches had the highest shoot densities and consequently a high abundance of predators (=juvenile conspecifics). Though shoot density did not differ between the edge and the interior of patches, crabs were more abundant in the interior of patches than at the edge. These results indicate that seagrass fragmentation does not have an overriding influence on juvenile blue crab survival and density, and that crab cannibalism and seasonal changes in landscape structure may influence relationships between crab survival and seagrass habitat structure. Habitat fragmentation, structural complexity, faunal density, and time all must be incorporated into future studies on faunal survival in seagrass landscapes.     

Effects of periodic hypoxia on mortality, feeding and predation in an estuarine epifaunal community

The York River Estuary, a tributary of the Chesapeake Bay, USA, experiences periodic low oxygen stress (hypoxia), yet epifaunal species form dense communities there. We studied hypoxia tolerance of common epifaunal species in the York River by exposing sessile and mobile epifauna to high and low oxygen concentrations in laboratory aquaria. Mortality in hypoxia varied among species, ranging from 0% to 100%, with trends of decreased tolerance by mobile species relative to sessile species. While most species tested experienced some mortality after being exposed to hypoxia (at 1 mg O(2)/l or 0.5 mg O(2)/l) for 5 days, many species had a median lethal time (LT(50)) in hypoxia greater than 1 week (3 of 6 species at 1 mg O(2)/l and 6 of 14 species at 0.5 mg O(2)/l), the maximum duration of typical hypoxic episodes in the York River, suggesting that hypoxia may cause little mortality for some species in this system. However, hypoxia had sub-lethal effects on behavior in all species tested. Epifaunal animals responded to hypoxia with behaviors that moved them higher in the water column or by entering resting states until hypoxia passed. Feeding and predation by a variety of taxa (the hydroid Obelia bicuspidata, the mud crab Neopanope sayi, juvenile blue crabs Callinectes sapidus, the flatworm Stylochus ellipticus, and the nudibranch Doridella leucolena) decreased during hypoxia, despite varying mortality responses to low oxygen stress, suggesting that short hypoxic episodes may create predation refuges for prey species. At least one highly tolerant species (O. bicuspidata) showed substantially decreased growth in hypoxia. Although relatively high tolerance of hypoxia by many estuarine epifaunal species limits serious disturbance during brief hypoxic episodes, hypoxia's greatest impact on York River epifaunal communities might be through its indirect effects on behavior and predation.     

Phenotypic plasticity in oysters (Crassostrea virginica) mediated by chemical signals from predators and injured prey

Prey organisms reduce predation risk by altering their behavior, morphology, or life history. Avoiding or deterring predators often incurs costs, such as reductions in growth or fecundity. Prey minimize costs by limiting predator avoidance or deterrence to situations that pose significant risk of injury or death, requiring them to gather information regarding the relative threat potential predators pose. Chemical cues are often used for risk evaluation, and we investigated morphological responses of oysters (Crassostrea virginica) to chemical cues from injured conspecifics, from heterospecifics, and from predatory blue crabs (Callinectes sapidus) reared on different diets. Previous studies found newly settled oysters reacted to crab predators by growing heavier, stronger shells, but that adult oysters did not. We exposed oysters at two size classes (newly settled oyster spat and juveniles ~2.0 cm) to predation risk cue treatments including predator or injured prey exudates and to seawater controls. Since both of the size classes tested can be eaten by blue crabs, we hypothesized that both would react to crab exudates by producing heavier, stronger shells. Oyster spat grew heavier shells that required significantly more force to break, an effective measure against predatory crabs, when exposed to chemical exudates from blue crabs as compared to controls. When exposed to chemical cues from injured conspecifics or from injured clams (Mercenaria mercenaria), a sympatric bivalve, shell mass and force were intermediate between predator treatments and controls, indicating that oysters react to injured prey cues but not as strongly as to cues released by predators. Juvenile oysters of ~ 2.0 cm did not significantly alter their shell morphology in any of the treatments. Thus, newly settled oysters can differentiate between predatory threats and adjust their responses accordingly, with the strongest responses being to exudates released by predators, but oysters of 2.0 cm and larger do not react morphologically to predatory threats.     

Predator biomass determines the magnitude of non-consumptive effects (NCEs) in both laboratory and field environments

Predator body size often indicates predation risk, but its significance in non-consumptive effects (NCEs) and predator risk assessment has been largely understudied. Although studies often recognize that predator body size can cause differing cascading effects, few directly examine prey foraging behavior in response to individual predator sizes or investigate how predator size is discerned. These mechanisms are important since perception of the risk imposed by predators dictates behavioral responses to predators and subsequent NCEs. Here, we evaluate the role of predator body size and biomass on risk assessment and the magnitude of NCEs by investigating mud crab foraging behavior and oyster survival in response to differing biomasses of blue crab predators using both laboratory and field methods. Cues from high predator biomass treatments including large blue crab predators and multiple small blue crab predators decreased mud crab foraging and increased oyster survival, whereas mud crab foraging in response to a single small blue crab did not differ from controls. Mud crabs also increased refuge use in the presence of large and multiple small, but not single small, blue crab predators. Thus, both predator biomass and aggregation patterns may affect the expression of NCEs. Understanding the impact of predator biomass may therefore be necessary to successfully predict the role of NCEs in shaping community dynamics. Further, the results of our laboratory experiments were consistent with observed NCEs in the field, suggesting that data from mesocosm environments can provide insight into field situations where flow and turbulence levels are moderate.     

Diet selectivity of juvenile blue crabs (Callinectes sapidus) in Chesapeake Bay

Shallow coves in Chesapeake Bay have abundant food and serve as nursery grounds for juvenile blue crabs. In this study, we examined the relationships between the diet of very small (4-40?mm CW) juvenile blue crabs and the benthic infauna in shallow, unvegetated nursery coves. We compared infauna in benthic samples with gut contents of juvenile blue crabs from six shallow coves in each of two sub-estuaries (Rappahannock and York Rivers) in Chesapeake Bay, Virginia, USA. Benthic communities differed depending on river and location, with abundant clams in upriver regions and abundant polychaetes in downriver regions. Juvenile crabs, like adults, appeared to be opportunistic feeders, with gut contents including clams, amphipods, polychaetes, small crustaceans, plant matter, and detritus. There was a positive relationship between polychaetes in the benthic samples and in crab guts, suggesting that juvenile crabs are opportunistic feeders on polychaetes in the benthos. Moreover, Ivlev's electivity index and foraging ratio showed that clams and polychaetes were selectively eaten at all locations. Alternatively, crabs selectively rejected amphipods. Crab densities corresponded positively with polychaete densities, which suggests that there may be bottom-up control of crab distributions and that food resources are important in nursery habitats.     

Antipredator defence of the hermit crab Pagurus filholi induced by predatory crabs

Hermit crabs have two antipredator tactics: taking refuge in its shell and fleeing. We examined the following two hypotheses using the hermit crab Pagurus filholi : (1) hermit crabs change their preference for shell types that they take refuge in and/or change the timing of fleeing (i.e. the duration of refuge in the shell) when they perceive a predator threat; (2) the type of shell that a hermit crab occupies affects the fleeing tactic of the individual. Under the stimulus of a crushed conspecific, hermit crabs changed neither their preference for shell species nor their refuge duration. On the other hand, under the stimulus of the predatory crab Gaetice depressus , hermit crabs increased their preference for Batillaria cumingi shells, which provide superior protection against predators, and shortened their refuge duration in the shells even when they occupied those effective against predation. Refuge duration was longer in B. cumingi shells than in the more vulnerable shells of Homalopoma sangarense . These results suggest that both antipredator defences (changing shell and timing of fleeing) are induced by the stimulus of a predator, and the timing of fleeing is affected by the shell type occupied.     

Predator-induced defenses in a salt-marsh gastropod

Predator-induced defenses are among the most ecologically important forms of phenotypic plasticity. Although predation and induced defenses are well documented in rocky-intertidal systems, they have received less attention in soft-bottom communities. Shell-crushing predators are common in soft-bottom, vegetated habitats, which often exhibit substantial spatial heterogeneity in predation intensity. We examined variations in shell morphology of the salt-marsh periwinkle, Littoraria irrorata, among marsh microhabitats in the northern Gulf of Mexico that vary in their accessibility to predatory blue crabs, Callinectes sapidus. Littoraria from high-predation sites exhibited more extensively calcified apertural lips and narrower apertural openings relative to snails from low-predation sites. Thick apertural lips generally increased the handling time required by Callinectes to breach Littoraria shells in laboratory experiments, although the method of shell entry used by crabs was dependent on the crab:snail size ratio. Apertural-lip thickness was not related to past predation events in field-collected snails. Snails exposed to water treated with the effluent of Callinectes and crushed conspecifics produced significantly thicker apertural lips than controls, with a response time and morphological extent comparable to that of their rocky-shore counterparts. This study underscores the widespread occurrence of predator-induced plasticity in marine gastropods and emphasizes its role in soft-bottom, vegetated marine habitats, where shell-crushing predation can be as prevalent a selective force as in the rocky intertidal.     

Fight or flight: an investigation of aggressive behavior and predator avoidance in two populations of blue crabs (<Emphasis Type="Italic">Callinectes sapidus</Emphasis> Rathbun) in New Jersey

Recent literature has suggested aggression may be context dependent. The purpose of this investigation was to examine aggressive and predator avoidance behaviors in juvenile blue crabs of two populations. Furthermore, we wanted to determine whether aggression persisted into the adult stages. Juvenile blue crabs collected from an impacted estuary, the Hackensack Meadowlands (HM), were found to attack a threatening stimulus significantly more often (70%) than conspecifics from a less impacted estuary (Tuckerton—TK). TK juveniles responded significantly more often with a flight (~35%) or mixed response (~30%). Additionally, HM juveniles were significantly more successful than TK juveniles at avoiding an adult blue crab predator when sandy substrate was present in laboratory experiments. However, the video clarity made it impossible to determine which interactions were allowing survival. To determine if “aggression” exhibited by the HM juveniles was the reason for their enhanced survival, follow-up predator avoidance experiments were conducted without substrate and videotaped. The results of these experiments suggest that aggression per se is not the reason since aggressive juveniles were no more successful than non-aggressive individuals. The aggressive behavior exhibited by HM juveniles continues into the adult stages. This behavior may be important to recognize when estimating population size as well as local fishery efforts.     

Predation by Callinectes sapidus (Rathbun) within Spartina alterniflora (Loisel) marshes

This study examined predation by the blue crab, Callinectes sapidus Rathbun, within intertidal Spartina alterniflora (Loisel) marshes of Dauphin Island, Alabama. Species and size preferences displayed by the predator when foraging within the marsh were investigated using nektonic, epifaunal, and infaunal prey populations including Fundulus similis Baird and Girard, Littorina irrorata Say, and Geukensia demissa Dillwyn.

Short-term field experiments involving the use of predator inclusion cages, in which the relative abundances of all prey species and the density of macrophyte vegetation were manipulated, indicated that mean mortality differed significantly among species. Blue crabs exhibited a distinct species preference for Littorina, and to a lesser extent, for Fundulus. However, the predator rarely choses infaunal individuals. Within predator inclusion cages, size selection by the crabs among three size classes of each prey was evident for Littorina and Fundulus but not for Geukensia. Blue crabs tended to select intermediate-sized snails and large fish while not exhibiting a size preference for infaunal bivalves.

In the marsh, mean percentage of the Littorina population within the 14–18 mm size class exhibited an increased mortality as compared to two other size classes, which was negatively correlated with increasing tidal height. Such a relationship may have been due to a decreasing gradient of crab predation associated with increasing tidal height. Geukensia size class distributions showed little evidence of differences along the tidal height gradient. No data are available for Fundulus, a mobile species which would not experience such differential predation along a marsh gradient.

In comparing crab predation patterns among prey species, it is apparent that Callinectes utilizes prey species differentially. Such differential utilization may be based on optimization of energy yield and minimization of energy expenditure. Thus, the preference of blue crabs for nektonic and epifaunal prey is hypothesized to be the result of a smaller energy expediture as a result of the crab's visual evaluation of these prey. Infaunal prey species (e.g., Geukensia) require a greater energy investment because of the necessity of excavating the prey item. Such prey also allow little selection by size because of being cryptic.     

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

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

Crab shell-crushing predation and gastropod architectural defense

The shell-breaking behavior of the crabs Ozius verreauxii Saussure 1853 and Eriphia squamata, Stimpson 1859 from the Bay of Panama is described. The master claws of both these crabs are well designed for breaking shells. Small shells, relative to the size of a crab predator, are crushed by progressively breaking off larger segments of a shell's apex, while larger shells are peeled by inserting a large dactyl molar into the aperture of a shell and progressively chipping away the lip of the shell.

Heavy gastropod shells are shown to be less vulnerable to crab predators than lighter shells, and narrow shell apertures and axial shell sculpture are demonstrated to be architectural features that deter crab predation. The incidence of architectural features which deter crab predation appears to be higher for smaller gastropod species than for larger gastropods which are too large for most crab predators. Large fish predators prey upon both gastropods and shell-crushing crabs. To avoid fish predators, both these prey groups seek refuge under rocks when covered by the tide. Fish predation thus appears to enforce a close sympatry between smaller gastropods and their crab predators.