Crab-resistant features of shells of burrowing bivalves: Decreasing vulnerability by increasing handling time

The vulnerability of burrowing bivalves to shell-breaking predation by crabs was found to be influenced strongly by shell features: size, shell thickness, degree of inflation, and the presence or absence of a gape.The relationship between the critical size of a bivalve (maximum size of vulnerability) and crab size was determined for four different morphotypes of bivalves. For the three bivalves where a “size refuge” was present, critical size increased with crab size. Nevertheless, when offered a choice crabs preferred clams well below the critical size and ate them in the order predicted by the critical-size experiment.Examination of the mechanics of shell crushing revealed how these shell features decreased vulnerability. Larger crabs could efficiently handle larger clams because both chela strength and degree of chela gape increased with crab size. Strain gauges attached to crab chelae showed that thick-shelled clams resisted a greater total number of force pulses than did thin-shelled clams of the same body weight. This may be related to the ability of thick-shelled clams to withstand greater loads than thin-shelled clams when loaded only once. This suggests that the reason for the increased resistance to crabs is prolongation of the shell-breaking time. Even though a large thick-shelled, tightly-closing, clam could eventually be opened, it will probably be rejected in favour of prey with shorter handling times.     

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.     

Morphology and muscle stress of chelae of temperate and tropical stone crabs Menippe mercenaria

Chela morphology and muscle stress were compared between temperate and tropical populations of stone crabs Menippe mercenuriu (Say) to test whether environmental differences might result in greater crushing strength in the tropics. Such differences include increased crab diversity in the tropics (which might lead to greater fighting among congeners), increased prey exoskeleton calcification in the tropics, and year round chela use in the tropics as opposed to seasonal chela use in the temperate population.
No latitudinal differences were found in any aspect of chela morphology, including relative chela size, mechanical advantage, apodeme surface area, and angle of muscle fibre pinnation. Summer measurements of crusher chela muscle stress were also similar between the two populations.
The maximum muscle stress determined for M. mercenariu was 220 N ∼r n -∼, much higher than stress levels previously reported for crustaceans. Other researchers have typically measured forces either from autotomized chelae or by measuring forces required to open a closed chelae. I have determined muscle stress using a force transducer that measures active gripping strength in live crabs.     

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.     

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.     

The geography of crushing: Variation in claw performance of the invasive crab Carcinus maenas

The major claws of predatory, durophagous decapods are specialized structures that are routinely used to crush the armor of their prey. This task requires the generation of extremely strong forces, among the strongest forces measured for any animal in any activity. Laboratory studies have shown that claw strength in crabs can respond plastically to, and thereby potentially match, the strength of their prey's defensive armor. These results suggest that claw strength may be variable among natural populations of crabs. However, very few studies have investigated spatial variation in claw strength and related morphometric traits in crabs. Using three geographically separate populations of the invasive green crab in the Gulf of Maine, we demonstrate, for the first time, geographic variation in directly measured claw crushing forces in a brachyuran. Despite variation in mean claw strength however, the scaling of claw crushing force with claw size was consistent among populations. We found that measurements of crushing force were obtained with low error and were highly repeatable for individual crabs. We also show that claw mass, independent of a linear measure of claw size, and carapace color, which is an indicator of time spent in the intermoult, were important predictors of claw crushing force.     

Specialized shell-breaking crab claws in Cretaceous seas

Here we report on a large brachyuran crab species from the Late Cretaceous of Mexico that has claws indicative of highly specialized shell-breaking behaviour. This crab possessed dimorphic claws (the right larger than the left), armed with several broad teeth, including a curved tooth structure found at the base of the movable finger of the right claw. The curved tooth is similar to the one observed on claws of many living durophagous crabs that use it as a weapon to peel, crush or chip the edges of hard-shelled prey, particularly molluscs. These morphological traits suggest that specialized shell-breaking crab predators had evolved during the Cretaceous, which contradicts previous findings supporting an Early Cenozoic origin for specialized shell crushers within the brachyuran clade.     

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.     

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.     

Selective crab predation on native and introduced bivalves in British Columbia

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

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

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

The 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.     

Changes in the selection differential exerted on a marine snail during the ontogeny of a predatory shore crab

Empirical estimates of selection gradients caused by predators are common, yet no one has quantified how these estimates vary with predator ontogeny. We used logistic regression to investigate how selection on gastropod shell thickness changed with predator size. Only small and medium purple shore crabs (Hemigrapsus nudus) exerted a linear selection gradient for increased shell‐thickness within a single population of the intertidal snail (Littorina subrotundata). The shape of the fitness function for shell thickness was confirmed to be linear for small and medium crabs but was humped for large male crabs, suggesting no directional selection. A second experiment using two prey species to amplify shell thickness differences established that the selection differential on adult snails decreased linearly as crab size increased. We observed differences in size distribution and sex ratios among three natural shore crab populations that may cause spatial and temporal variation in predator‐mediated selection on local snail populations.     

Predation induced changes in behavior and growth rate in three populations of the intertidal snail, Littorina sitkana (Philippi)

We investigated the sublethal effects of a predatory crab, Cancer productus (Randall), on the behavior and growth of its snail prey, Littorina sitkana, by setting up controlled rearing and prey-size selection experiments. L. sitkana were collected from three sites on San Juan Island, WA, USA. These sites varied in snail size, abundance, and vertical distribution, and in the abundance of the crab predator C. productus. Snails from all three populations were raised for 34 days under the following treatments: no-crab control, a non-feeding C. productus encased in mesh box, and an encased C. productus feeding on L. sitkana. The non-feeding crab treatment did not affect snail foraging behavior or growth rate in comparison with the no-crab control. In contrast, the presence of a feeding crab elicited escape behavior in the snails, halted grazing, and consequently reduced growth rates. A population difference in escape behavior was observed: upward migration in snails from rocky shores and hiding in crevices in snails from a mud flat. It thus appears that chemicals leaching from crushed conspecific snails, rather than the presence of the crab predator, act as the “alarm substance” to which L. sitkana react. The magnitude of the growth depression in the presence of feeding crabs was 85%, with no difference among the three populations. Once the feeding crab stimulus was removed, snails in all populations resumed normal growth, suggesting that this response to feeding predators is reversible with changing environmental conditions. Laboratory experiments were set up to determine if all size classes of L. sitkana are equally susceptible to C. productus predation. C. productus consistently selected the largest of three size classes of L. sitkana. These results suggest that slow growth rate and small size in L. sitkana may actually be an adaptation for coexisting with high C. productus abundance, rather than simply a cost of escape behavior.     

What makes a good home for hermits? Assessing gastropod shell density and relative strength

The survival and reproductive success of hermit crabs is intrinsically linked to the quality of their domicile shells. Because damaged or eroded shells can result in greater predation, evaluating shell structure may aid our understanding of population dynamics. We assessed the structural attributes of Cerithium atratum shells through assessments of (a) density using a novel approach involving computed tomography and (b) tolerance to compressive force. Our goal was to investigate factors that may influence decision making in hermit crabs, specifically those that balance the degree of protection afforded by a shell (i.e. density and strength) with the energetic costs of carrying such resources. We compared the density and relative strength (i.e. using compression tests) of shells inhabited by live gastropods, hermit crabs (Pagurus criniticornis) and those found empty in the environment. Results failed to show any relationship between density and shell size, but there was a notable effect of shell density among treatment groups (gastropod/empty/hermit crab). There was also a predictable effect of shell size on maximum compressive force, which was consistent among occupants. Our results suggest that hermit crabs integrate multiple sources of information, selecting homes that while less dense (i.e. reducing the energy costs of carrying these resources), still offer sufficient resistance to compressive forces (e.g. such as those inflicted by shell-breaking predators). Lastly, we show that shell size generally reflects shell strength, thus explaining the motivation of hermit crabs to search for and indeed fight over the larger homes.     

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.     

Does the handedness of the pebble crab Eriphia smithii influence its attack success on two dextral snail species?

The predaceous crab Eriphia smithii (Xanthidae) has one larger claw with molar teeth on either the right or the left cheliped, which it uses to crush the shell of prey. Whether the handedness of crabs affected successful predation on two snail species, Nerita albicilla (Neritidae) and Planaxis sulcatus (Planaxidae) was experimentally investigated. The fate of snails of each species was analysed by multiple logistic regression with three explanatory variables: handedness, shell-size index and individuality of crabs. No effect of handedness was detected in attacks on N. albicilla , probably as a result of the spherical and more symmetrical shell morphology of this species. In contrast, right-handedness contributed to greater attack success on P. sulcatus , which has more conical shells. Further investigation of how snail shells were broken revealed that left-handed crabs had more difficulty breaking the aperture of larger P. sulcatus , which was thought to cause the difference in attack success between right- and left-handed crabs. The advantages conferred by handedness are discussed.     

Interference and exploitation components in interespecific competition between sympatric intertidal hermit crabs

This study was designed to evaluate the effect of interference and exploitation competition in shell partitioning between two hermit crab species (Pagurus criniticornis and Clibanarius antillensis). Field samples revealed that shells of the gastropod Cerithium atratum were the main resource used by both hermit crab species and that Pagurus used eroded or damaged shells in higher frequency than Clibanarius. The exploitative ability of each species was compared between species in the laboratory using dead gastropod (Cerithium) baits to simulate predation events and signalize newly available shells to hermit crabs. Pagurus reached the baits more rapidly than Clibanarius, but this higher exploitative ability did not explain shell utilization patterns in nature. Another experiment evaluated the dominance hierarchy between these two hermit crab species and revealed that Clibanarius was able to outcompete Pagurus for higher quality shells in agonistic encounters. This higher interference competitive ability of Clibanarius in relation to Pagurus may explain field observations. Nevertheless, Pagurus may be responsible to enhance shell availability to other hermit crab species that have lower ability to find and use newly available shells. Differently, the poorer condition of shells used by Pagurus, the higher ability of this species to attend gastropod predation events and its higher consumption rate by shell-breaking crabs (Menippe nodifrons) may increase its predation risks, thus revealing the disadvantages of such an exploitative competitive strategy for hermit crabs.     

Green crab (Carcinus maenas) foraging efficiency reduced by fast flows

Predators can strongly influence prey populations and the structure and function of ecosystems, but these effects can be modified by environmental stress. For example, fluid velocity and turbulence can alter the impact of predators by limiting their environmental range and altering their foraging ability. We investigated how hydrodynamics affected the foraging behavior of the green crab (Carcinus maenas), which is invading marine habitats throughout the world. High flow velocities are known to reduce green crab predation rates and our study sought to identify the mechanisms by which flow affects green crabs. We performed a series of experiments with green crabs to determine: 1) if their ability to find prey was altered by flow in the field, 2) how flow velocity influenced their foraging efficiency, and 3) how flow velocity affected their handling time of prey. In a field study, we caught significantly fewer crabs in baited traps at sites with fast versus slow flows even though crabs were more abundant in high flow areas. This finding suggests that higher velocity flows impair the ability of green crabs to locate prey. In laboratory flume assays, green crabs foraged less efficiently when flow velocity was increased. Moreover, green crabs required significantly more time to consume prey in high velocity flows. Our data indicate that flow can impose significant chemosensory and physical constraints on green crabs. Hence, hydrodynamics may strongly influence the role that green crabs and other predators play in rocky intertidal communities.     

Impact Scenario for an Introduced Decapod on Arctic Epibenthic Communities

The intentional introduction of a species for the enhancement of stock or establishment of new fisheries, often has unforeseen effects. The red king crabs, Paralithodes camtschaticus, which was introduced into the Barents Sea by Russian scientists, has established a self-sustaining population that has expanded into Norwegian waters. As top benthic predators, the introduced red king crabs may have possible effects upon native epifaunal scallop (Chlamys islandica) communities. These benthic communities may be a source of prey species in late spring, when the red king crabs feed most intensively. Foraging rates (consumption, killing or severely damaging) of red king crab on native prey organisms were measured by factorial manipulation of crab density (0.5, 1.5 and 3 per m ²), size classes (immature, small mature, and large mature crabs), and by evaluating prey consumption after 48 h, in order to extrapolate a scenario of the likely impacts. Foraging rates of the red king crab on scallops ranged between 150 and 335 g per m² within 48 h. These rates did not change when crab density was altered, though an increased amount of crushed scallops left uneaten at the tank floor, were correlated with high density of small mature crabs. Foraging rate changed significantly with crab size. Consequently, the susceptibility of native, shallow water epibenthic communities to red king crab predation in the early life history stages, and during the post-mating/molting spring period, must be considered significant when foraging rates are contrasted with natural scallop biomass between 400 and 1200 g scallops per m².