Predator (<Emphasis Type="Italic">Carcinus maenas</Emphasis>) nonconsumptive limitation of prey (<Emphasis Type="Italic">Nucella lapillus</Emphasis>) feeding depends on prey density and predator cue type

Predators often have nonconsumptive effects (NCEs) on prey. For example, upon detection of predator cues, prey can reduce feeding activities to hamper being detected by predators. Previous research showed that waterborne chemical cues from green crabs (Carcinus maenas, predator) limit the dogwhelk (Nucella lapillus, prey) consumption of barnacles regardless of dogwhelk density, even though individual predation risk for dogwhelks decreases with conspecific density. Such NCEs might disappear with dogwhelk density if dogwhelks feed on mussels, as mussel stands constitute better antipredator refuges than barnacle stands. Through a laboratory experiment, we effectively found that crab chemical cues limit the per-capita consumption of mussels by dogwhelks at low dogwhelk density but not at high density. The combination of tactile and chemical cues from crabs, however, limited the dogwhelk consumption of mussels at both dogwhelk densities. The occurrence of such NCEs at both dogwhelk densities could have resulted from tactile cues indicating a stronger predation risk than chemical cues alone. Overall, the present study reinforces the notions that prey evaluate conspecific density when assessing predation risk and that predator cue type affects their perception of risk.     

Adult Prey Neutralizes Predator Nonconsumptive Limitation of Prey Recruitment

Recent studies have shown that predator chemical cues can limit prey demographic rates such as recruitment. For instance, barnacle pelagic larvae reduce settlement where predatory dogwhelk cues are detected, thereby limiting benthic recruitment. However, adult barnacles attract conspecific larvae through chemical and visual cues, aiding larvae to find suitable habitat for development. Thus, we tested the hypothesis that the presence of adult barnacles (Semibalanus balanoides) can neutralize dogwhelk (Nucella lapillus) nonconsumptive effects on barnacle recruitment. We did a field experiment in Atlantic Canada during the 2012 and 2013 barnacle recruitment seasons (May–June). We manipulated the presence of dogwhelks (without allowing them to physically contact barnacles) and adult barnacles in cages established in rocky intertidal habitats. At the end of both recruitment seasons, we measured barnacle recruit density on tiles kept inside the cages. Without adult barnacles, the nearby presence of dogwhelks limited barnacle recruitment by 51%. However, the presence of adult barnacles increased barnacle recruitment by 44% and neutralized dogwhelk nonconsumptive effects on barnacle recruitment, as recruit density was unaffected by dogwhelk presence. For species from several invertebrate phyla, benthic adult organisms attract conspecific pelagic larvae. Thus, adult prey might commonly constitute a key factor preventing negative predator nonconsumptive effects on prey recruitment.     

Behavioural components of prey-selection by dogwhelks, Nucella lapillus (L.), feeding on barnacles, Semibalanusbalanoides (L.), in the laboratory

Adult dogwhelks, maintained on mussels for 60 days before experimentation to reduce prior effects of ingestive conditioning and handling skills appropriate to barnacles, and thence deemed “inexperienced”, preferred the largest barnacles presented to them. Juvenile and adult dogwhelks, maintained on barnacles and deemed experienced, preferred barnacles of intermediate sizes that were correlated with the sizes of the dogwhelks. “Inexperienced” dogwhelks penetrated barnacles significantly more often by drilling than by prising, whereas experienced dogwhelks did the reverse. A predominant tendency to prise open barnacles was developed by previously “inexperienced” dogwhelks after they had eaten six to eight consecutive prey. Larger dogwhelks prised open greater proportions of larger barnacles than did smaller dogwhelks. Experienced adult dogwhelks prised open all barnacles of ≈2-mm opercular diameter but only 20% of those ≈ 6 mm in diameter. The preferred barnacles, 4–5 mm in opercular diameter, were of a size that the experienced adult dogwhelks could prise open in 50% of attacks.Dogwhelks inspected barnacles by crawling over them for ≈20 min when the prey were subsequently rejected and ≈30 min when subsequently eaten. Penetration time and ingestion time were linear functions of barnacle diameter, total handling time ranging from 4–26 h for barnacles of 2.5–2.7 mm opercular diameter. Experienced dogwhelks handled barnacles faster than “inexperienced” dogwhelks, largely because they used the quicker method of penetration by prising whenever possible. The yield of flesh per unit handling time was an accelerating function of barnacle size and, with experience, could be increased by a factor of ≈1.2 for barnacles of 7 mm opercular diameter to ≈2.0 for barnacles of 2 mm opercular diameter. The preference of experienced dogwhelks for barnacles somewhat smaller than the largest available, may have reflected the greater frequency with which these prey could be penetrated by the quicker method of prising. Minimizing handling time could be important in nature if there is a risk to feeding dogwhelks of being displaced by competitors. In the laboratory, 12% of dogwhelks where thus displaced.Barnacles of ≈5 mm opercular diameter were estimated to be slightly more profitable than mussels of ≈17.5-mm shell length to a dogwhelk experienced with barnacles, but less profitable to a dogwhelk that has recently fed extensively on mussels. Dogwhelks, therefore, might switch between diets predominately of barnacles or of mussels if prey of comparable profitabilities change drastically in their relative abundance on the shore.     

Generalizing from experiments: is predation strong or weak in the New England rocky intertidal?

Summary Petraitis (1990) recently critized previous generalizations regarding the effects of predation in the New England rocky intertidal region (e.g., Menge 1976; Lubchenco and Menge 1978). Contrary to Lubchenco's and my conclusions, Petraitis concluded that (1) barnacles and not mussels are the favored prey of dogwhelks and (2) barnacles and not dogwhelks control mussel abundances in the mid and low rocky intertidal region. I provide evidence that these criticisms are unwarranted. First, Lubchenco and I never claimed that diet composition reflected prey preference. Moreover, predators can influence prey abundance without preferring the prey. Hence, claims regarding preferences have no bearing on our conclusions. Second, Petraitis' experiments do not invalidate Lubchenco's and my experimental results. Reanalyses of our experimental data support the earlier conclusion that at wave-sheltered sites, whelks reduce the abundance of mussels independently of barnacle abundances. Further, at all but one of Lubchenco's and my study sites, predator densities were higher than at Petraitis' site. Thus, the absence of a predator effect in Petraitis's study was most likely due to low predator density rather than a lack of generality of our earlier results. This reevaluation therefore suggests that within a broader conceptual framework, Petraitis' apparently divergent results are actually consistent with ours.     

Experimental determination of predation intensity in an intertidal predator guild: dominant versus subordinate prey

Theoretical and empirical ecologists have long acknowledged that information about the intensity or strength of the interaction between species is crucial for an understanding of community dynamics. In communities in which predation is an important structuring process, and some predator species are commercially exploited, quantitative estimates of predation by different predator species within a guild are necessary to make even the simplest recommendations about conservation and resource management. Here, we evaluated per capita and population level components of predation intensity of three intertidal predators that feed on monospecific stands of barnacles and mussels at wave exposed sites in the rocky intertidal zone of central Chile. These prey species represent the two most distinctive stages of the mid-intertidal seascape, with mussels being competitively dominant. Our results showed that the commercially exploited gastropod Concholepas concholepas and the sea star Heliaster helianthus have similarly large per capita and population effects on the competitively dominant mussel Perumytilus purpuratus . Their per capita (by average size individual) and population effects on mussels were more than two orders of magnitude larger than those of Acanthocyclus gayi crabs and likely even larger than the effect of other predator species in this system (other crabs, whelks, birds, fish). The overall pattern of predation on barnacles was similar to that on mussels, but some differences occurred in the way different components of predation intensity were distributed across predator species. Despite the roughly similar pattern of population predation intensity between prey species, the expected consequences for the prey population, and hence the rest of the community, were acutely different for mussels and barnacles.     

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.     

Response of predatory snails to a novel prey following the geographic range expansion of an intertidal barnacle

During a geographic range expansion, the success of a species colonizing a new region may be influenced by how resident predators respond to this novel prey. The volcano barnacle Tetraclita rubescens has recently expanded its range > 300 km northward along the coast of California, USA. In the historic portion of its range, Tetraclita is preyed upon by a southern dogwhelk, Nucella emarginata. In newly colonized northern habitats, Tetraclita has joined an assemblage of mid-intertidal barnacles preyed upon by two species of northern dogwhelks (N. ostrina and N. canaliculata). Using a combination of field enclosures and laboratory experiments, we tested the impacts of northern whelks on volcano barnacles relative to other barnacle species. In the field, juvenile Tetraclita were rarely consumed. In laboratory trials with larger barnacles, Tetraclita were nearly immune to drilling by northern whelks, perhaps due to superior morphological defenses. In all experiments, prior encounters with Tetraclita did not appear to increase the frequency of predation on this barnacle; northern whelk species rarely preyed on Tetraclita even when collected from a site where this barnacle was abundant. In contrast, the southern whelk N. emarginata readily preyed on Tetraclita in laboratory trials, perhaps reflecting a longer shared evolutionary history between these species. Overall, our results suggest that the success of a population established beyond a species' historical range boundary may be influenced by the functional traits of that species in relation to the composition and evolutionary history of the surrounding community.     

Enhanced condition in dogwhelks, Nucella lapillus (L.) living under mussel hummocks

Dogwhelks, Nucella lapillus eat both barnacles and mussels, Mytilus edulis. However, mussels are dangerous prey because they can snare and immobilize dogwhelks with their byssi. Despite this danger, adult dogwhelks are common inside mussel hummocks or stacks found on mature mussel beds. The whelks live in debris-filled chambers within the hummocks, unsnared by byssi. Evidence is presented to show that this is a highly favourable microhabitat for dogwhelks. Condition index (CI) measurements showed that mean CI for “hummock” dogwhelks was 20.99 in early spring, 29.65 in midsummer and 34.86 in early winter. In comparison, CI values for dogwhelks from nearby barnacle zones were 20.13, 25.03 and 24.66 respectively. The summer and winter CI values of “hummock” dogwhelks were enhanced to a highly significant extent. Data are also presented to show that shore elevation or dogwhelk shell height (over the size range of animals studied) had no confounding effect on this result. Predicted flesh and shell mass values for a standard 25 mm high whelk showed that occupation of hummocks was associated with a thin-walled shell morphology, and that the shell mass became proportionately lower in summer and early winter. A feeding experiment showed that CI rises slowly, even in dogwhelks fed to satiation on mussels. Dogwhelks were taken from the barnacle zone in March 1997 and either fed to satiation on Mytilus or starved, whilst being held in seawater at ambient seawater temperature. The CI of fed dogwhelks rose from 19.7 to 27.4 in 3 months, while that of starved Nucella fell from 19.7 to 16.3. In the meantime, the CI of dogwhelks in situ in the barnacle zone had risen from 19.7 to 23.8. Hence, 3 months' satiation ration had led to a nett 3.6 unit rise in CI; starvation to a nett 7.5 unit fall. All of the changes in CI were due to changes in flesh mass; shell mass was stable. Since “hummock” dogwhelks collected in early winter have a CI value about 10.2 units greater than barnacle zone dogwhelks, this indicates that “hummock” dogwhelks are long-term inhabitants of this microhabitat.     

Mussel and dogwhelk distribution along the north‐west Atlantic coast: testing predictions derived from the abundant‐centre model

Aim We performed the first test of predictions from the abundant‐centre model using north‐west Atlantic coastal organisms. We tested the hypotheses that the density of intertidal mussels (Mytilus edulis and M. trossulus) and dogwhelks (Nucella lapillus) and mussel age and size would peak at an intermediate location along their distribution range. We also assessed the latitudinal variation in critical aerial exposure time. Location North‐west Atlantic coast between Newfoundland (Canada) and New York (USA), covering 1800 km of shoreline. Methods Using a nested design, we measured mussel density, age and size and dogwhelk density in 60 wave‐exposed rocky intertidal sites spread evenly in six regions. Critical aerial exposure times were determined using online data. Results Mytilus edulis peaked in abundance in Maine and was much less abundant in the other regions. Mytilus trossulus peaked in abundance in southern Nova Scotia and Maine, was less abundant in the other regions to the north, and was absent in the southernmost region (New York). Both mussel species were least abundant in a northern region (Cape Breton), although not in the northernmost region (Newfoundland). Critical aerial exposure times were negatively correlated with overall mussel density. Mussel age and size were similar among regions. Dogwhelks peaked in abundance in Maine and were much less abundant in the other regions, being positively correlated with overall mussel density across regions. Main conclusions Density data for M. edulis and N. lapillus provide limited support for an abundant‐centre pattern, while M. trossulus shows a clear ramped‐south distribution. Critical aerial exposure times suggest that physiological stress during summer and winter low tides may be lowest in Maine and southern Nova Scotia, which might partially explain mussel predominance in those regions. Winter ice scour in Cape Breton may explain the abundance trough observed there. Mussel size and age may be more limited by wave exposure at our sites (as they all face open waters) than by regional differences in environmental stress. Dogwhelks, which prey on mussels, seem to respond positively to prey density at the regional scale. Our study supports the notion that, while the abundant‐centre model is a useful starting point for research, it often represents an oversimplification of reality.     

Survivorship of juvenile barnacles and mussels: spatial dependence and the origin of alternative communities

Mussels, barnacles, and rockweeds often form a distinct mosaic of patches on rocky intertidal shores, and it has been suggested that these communities may represent alternative community states. One way that alternative community states can arise is if early successional events are scale-dependent, but it is not known if juvenile survivorships of mussels and barnacles are, in fact, scale-dependent. Scale-dependence of barnacles (Semibalanus balanoides (L.)) and mussels (Mytilus edulis L.) was tested in the Gulf of Maine, USA. In winter 1997, clearings of 1, 2, 4 and 8 m in diameter and uncleared controls were made in stands of the rockweed Ascophyllum nodosum (L.) Le Jolis at 12 sites spread evenly across four bays on Swan's Island, ME. Summer and fall-winter survivorship of barnacles, which recruited in spring 1997, were estimated by tracking the 1997 cohort until late winter 1998. Survivorship of mussels was estimated from following the fate of transplanted juveniles over 4 days in late August 1997. Both barnacles and mussels showed better survival in 4 and 8 m clearings than in small clearings and controls. There was also significant variation in survivorship among sites. Densities of gastropods in the clearings did not reflect survivorship patterns of barnacles and mussels. Barnacle survivorship increased in fall and winter, and in large clearings was comparable to survivorship in barnacle-dominated habitats. Mussel survivorship was low in all clearing sizes suggesting that mussel beds develop slowly.     

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.     

Predation by gulls on crabs in rocky intertidal and shallow subtidal zones of the Gulf of Maine

Most organisms in intertidal areas are marine in origin; many have distributions that extend into the subtidal zone. Terrestrial predators such as mammals and birds may exploit these animals during low tide and can have considerable effects on intertidal food webs. Several studies have shown that avian predators are capable of reducing densities of sessile and slow-moving intertidal invertebrates but very few studies have considered avian predation on mobile invertebrate predators such as crabs. In this study, we investigated predation by Great Black-backed Gulls (Larus marinus Linnaeus) on three species of crabs (Cancer borealis Stimpson, Cancer irroratus Say, and Carcinus maenas Linnaeus). The study was at Appledore Island, ME (a gull breeding island) and 8 other sites throughout the Gulf of Maine, including breeding islands and mainland sites. On Appledore Island, intertidal and subtidal zones provided over one-third of prey remains found at gull nests, and crabs were a substantial proportion (∼ 30% to 40%) of the total remains. Similarly, collections of prey remains from intertidal areas indicated that crabs were by far the most common marine prey. C. borealis was eaten far more often and C. irroratus and C. maenas less often than expected at each site. Comparing numbers of carapaces to densities of crabs in low intertidal and shallow subtidal zones at each site, we estimated that gulls remove between 15% and 64% of C. borealis during diurnal low tides. The proportion of C. borealis eaten by gulls was independent of proximity to a gull colony. Approximately 97% of the outer coast of Maine is within 20 km of a breeding island. Thus, a lot of gull predation on crabs may occur throughout the Gulf of Maine during summer months. Crabs are important predators of other invertebrates; if predation by gulls reduces the number of crabs in intertidal and shallow subtidal areas, gulls may have important indirect effects on intertidal food webs.     

Effects of temperature,salinity and aerial exposure on predation and lysosomal stability of the dogwhelk Thais (Nucella) lapillus (L.)

The ingestion and absorption rate of standard length Thais lapillus (L.) stepwise-acclimated to constant temperature-salinity conditions and preying on Mytilus edulis (L.) varied directly with environmental salinity at 10, 15 and 20°C. Dogwhelk ingestion and absorption rates indicate that cold torpor existed at 5°C and heat stress was evident at 20°C. The feeding cycle duration was significantly longer for dogwhelks acclimated to 20%. S than in those acclimated to 30%. S at 10°C even though no significant difference existed between the two groups of snails in the drilling and ingestion or postfeeding phases of the cycle. Ingestive conditioning of dogwhelks to mussels occurred; the duration of the drilling and ingestion and total feeding cycle declined as a function of the number of mussels consumed by a snail. Dogwhelks of all sizes prey on a wide length range of mussels and there is also a high degree of variability in the ingestion rate of snails as a function of their size. A prominent feature of the lack of a relationship between dogwhelk ingestion rate and snail size was that the percentage of nonfeeding snails increased at low salinity and temperature extremes. Digestive-tubule cell lysosomal stability was tested as an index of digestive capability and animal condition; in stepwise-acclimated dogwhelks, it correlated well with their ingestion and absorption rates. The ingestion rate of dogwhelks acclimated to 30%. S and subjected to a 30?17.5?30%. S semidiurnal pattern of fluctuating salinity for 21 days was significantly lower than for snails maintained at 30%. S; however, snails acclimated to 17.5%. S and exposed to the same pattern of fluctuating salinity fed at a higher rate than snails maintained at 17.5%. S. Aerial exposure of snails maintained at 30%. S and 10°C water temperature resulted in an ingestion rate 2.1 times faster than for snails constantly submerged suggesting that tidal emersion is not always stressful to intertidal carnivores. The postfeeding phase of the feeding cycle was shortened in dogwhelks subjected to aerial exposure. Although significant variation occurred in digestive-tubule cell lysosomal stability during the first cycle of fluctuating salinity, the variability had declined significantly by Day 21. This observation suggests that digestive tubule lysosomal stability becomes adapted to a fluctuating osmotic environment, although the initial changes in lysosomal stability are probably related to intralysosomal protein catabolism and production of amino acids for intracellular osmoregulation. Variations in the osmotic environment of T. lapillus have resulted in unexpected outcomes with respect to their ingestion rate under conditions of fluctuating salinity and aerial exposure.     

Biogeographic variation in behavioral and morphological responses to predation risk

The expression of prey antipredator defenses is often related to ambient consumer pressure, and prey express greater defenses under intense consumer pressure. Predation is generally greater at lower latitudes, and antipredator defenses often display a biogeographic pattern. Predation pressure may also vary significantly between habitats within latitudes, making biogeographic patterns difficult to distinguish. Furthermore, invasive predators may also influence the expression of prey defenses in ecological time. The purpose of this study was to determine how these factors influence the strength of antipredator responses. To assess patterns in prey antipredator defenses based upon geographic range (north vs. south), habitat type (wave-protected vs. wave-exposed shores), and invasive predators, we examined how native rock (Cancer irroratus) and invasive green (Carcinus maenas) crab predators influence the behavioral and morphological defenses of dogwhelk (Nucella lapillus) prey from habitats that differ in wave exposure across an ~230 km range within the Gulf of Maine. The expression of behavioral and morphological antipredatory responses varied according to wave exposure, geographic location, and predator species. Dogwhelks from areas with an established history with green crabs exhibited the largest behavioral and morphological antipredator responses to green crabs. Dogwhelk behavioral responses to rock crabs did not vary between habitats or geographic regions, although morphological responses were greater further south where predation pressure was greatest. These findings suggest that dogwhelk responses to invasive and native predators vary according to geographic location and habitat, and are strongly affected by ambient predation pressure due to the invasion history of an exotic predator.     

Using patterns of variability to test for multiple community states on rocky intertidal shores

Predictions based on theory of multiple stable states suggest that larger perturbations should lead to more unpredictable patterns of succession. This prediction was tested in the Gulf of Maine using data from 60 intertidal plots of varying size that were experimentally cleared of the rockweed Ascophyllum nodosum and from 14 benchmark sites from throughout the Gulf. Rockweed was removed from the experimental clearings ranging from 1 to 8 m in diameter in 1996 and data collected in 2004 were used to test effects of clearing size and location on divergence and variability in species composition. Benchmark data were collected in 2005, and the 14 sites were from a dataset on 53 sites throughout the Gulf of Maine. The selected sites were randomly chosen from all sites with > 80% canopy cover by A. nodosum and were expected to be similar to uncleared control plots from the experiment. Experimental removal of A. nodosum resulted in clearings at 12 sites within 4 bays. Abundances of gastropods, barnacles, mussels, and fucoid algae and the percentage cover of barnacles, mussels, fucoid algae, bare space, and other species were sampled. CAP and PERMDISP analyses revealed significant differences in multivariate dispersion and variability with both clearing size and location. Variability generally increased with clearing size and location effects were related to the north-south positioning of the sites. Benchmark sites were similar to the experimental control plots but as variable as the largest clearings. Results suggest that succession in larger clearings has been more unpredictable than in small clearings. The pattern of variability in the experimental clearings is consistent with the predictions of multiple stable states. However, the large amount of variation among the benchmark sites was due to mussels and was unexpected. This unexpected variability underscores the importance of sampling benchmark sites as part of experiments.     

Factors organizing rocky intertidal communities of New England: Herbivory and predation in sheltered bays

In New England, U.S.A., shores exposed to severe wave action are dominated by the common blue mussel Mytilus edulis L. while moderately protected areas are covered with perennial algae. It is thought that algae are limited by mussels which are a superior competitor. Because the effectiveness of predators is inhibited by wave activity, it is assumed that the rate of predation, which varies across this environmental gradient, accounts for the observed distribution of mussels and algae.Shores along sheltered bays appear to be an exception to this pattern and this study addresses some of the possible causes. In New England bays, mussels and barnacles Semibalanus balanoides (L.) are the most common organisms on the solid surfaces in the lower intertidal zone. Perennial macroalgae, such as Chondrus crispas Stackhouse and Fucus vesiculosus L., are rare. The distribution and abundance of species differs from that on moderately protected shores and is similar to very exposed shores which are dominated by mussels and barnacles.Herbivory by the common periwinkle Littorina littorea (L.) limits the abundance of F. vesiculosus and indirectly affects the success of mussels. During 4 years of experimental manipulations, F. vesiculosus rarely recruited in the presence of periwinkles but dominated experimental surfaces if periwinkles were excluded. When experimental surfaces with F. vesiculosus, which had been protected from herbivory for > 1 year, were exposed to natural conditions, herbivores cleared most of the surfaces within several months. Recruitment by barnacles and mussels was higher when periwinkles were excluded. However, the effect of periwinkles on mussels was indirect; the snails reduced barnacle success and thus reduced mussel recruitment which was enhanced by the surface irregularities provided by barnacles.The occurrence of mussels in sheltered bays is not due to a lack of predators. Predators were commonly seen at all sites. Most mussels on experimental surfaces were removed <4 wk when surfaces were exposed to natural levels of predation. Experiments do not provide an explanation for the occurrence of mussels, although the enhancement of mussel recruitment by barnacles suggests that the availability of settlement sites may be important.     

Isolated and combined impacts of blue mussels (Mytilus edulis) and barnacles (Balanus improvisus) on structure and diversity of a fouling community

Effects of two presumably dominant competitors, the blue mussel Mytilus edulis and the barnacle Balanus improvisus on recruitment, population dynamics and community structure on hard substrata were experimentally investigated in the subtidal Kiel Fjord, Western Baltic. The hypothesis that blue mussels and/or barnacles are local dominants and strongly influence succession and community structure was tested by monitoring succession in the presence and absence of simulated predation on either or both species. Manipulations included blue mussel removal, barnacle removal, combined blue mussel and barnacle removal, as well as a control treatment for natural (non-manipulated) succession. In the second part of the experiment, recovery from the treatments was monitored over 1 year.During the manipulative phase of the experiment, blue mussels had a negative effect on recruitment of species, whereas barnacles had no significant effect. Even so, a negative synergistic effect of blue mussels and barnacles was detected. Calculation of species richness and diversity H′ (Shannon Index) showed a negative synergistic effect of blue mussels and barnacles on community structure. Additionally, diversity H′ was negatively affected by the dominant competitor M. edulis. These effects were also detectable in the ANOSIM-Analysis. The non-manipulative phase of the experiment brought about a drastic loss of diversity and species richness. Blue mussels dominated all four communities. Barnacles were the only other species still being able to coexist with mussels. Effects of simulated predation disappeared fast.Thus, in the absence of predation on blue mussels, M. edulis within a few months dominates available space, and diversity of the benthic community is low. In contrast, when mussel dominance is controlled by specific predators, more species may persist and diversity remains high.     

Prey selection and the impact of the starfish Marthasterias glacialis (L.) and other predators on the mussel Choromytilus Meridionalis (Krauss)

The diet of the starfish, Marthasterias glacialis (L.), consists of a variety of mollusc species, as well as ascidians and barnacles. Starfish densities are maximal where mussels, Choromytilus meridionalis (Krauss), are abundant and in such areas mussels form the bulk of the diet. Laboratory feeding experiments indicate that Marthasterias glacialis select mussels of particular sizes and that the length of prey taken is an increasing function of predator arm length. The time taken to consume each mussel is determined by the ratio of shell length to starfish size. The number of mussels consumed per day increases only slightly with starfish size, but because the prey taken increase in size, energy consumption is maintained at a relatively consistent 1% of predator body energy per day. Using prey selection and feeding rate data for different sized starfish, predictive three dimensional predation surfaces are developed for a natural starfish population feeding on either one or two cohort Choromytilus meridionalis populations. The models indicate that predatory effort should be concentrated on the smallest mussels when a single adult cohort is present, but on recruiting mussels just above the minimum prey size limit where two cohorts are present. Other major predators of mussels, the rock lobster, Jasus lalandii (Milne Edwards), and the whelk, Natica tecta Anton, appear to select similar size-ranges of prey to starfish, despite their differing body forms and feeding methods. Since the juveniles of all three predators can only take small mussels, predator recruitment may well depend upon the successful settlement of strong mussel cohorts. Evidence for such entrainment of predator cohorts to settlements of mussels is presented.     

Subtidal-intertidal trophic links: American lobsters [Homarus americanus (Milne-Edwards)] forage in the intertidal zone on nocturnal high tides

Homarus americanus (Milne-Edwards), the American lobster, is a predator in New England subtidal communities, feeding on ecologically important grazers (sea urchins), mesopredators (crabs), and basal species (mussels). In this study, we provide the first report of adult American lobsters foraging in rocky intertidal habitats during nocturnal high tides. Censuses by SCUBA divers in the low intertidal (Chondrus crispus Stackhouse) zone showed mean densities of 2.2 lobsters/20 m² on nocturnal high tides, with contrasting low densities of 0.18/20 m² during diurnal high tides. Nocturnal high-tide intertidal densities were 62% of those reported in a previous study of lobsters in nearby subtidal rocky areas (Novak, 2004). The average carapace length of lobsters in the intertidal at night was > 50 mm. These lobsters were actively foraging in the intertidal with collected individuals having a mean stomach fullness of 67%. Prey found in the stomach contents primarily consisted of crabs, mussels and snails. Field experiments showed that lobsters rarely fed on medium to large size individuals of the common intertidal snail, Littorina littorea (L.). In contrast, experiments with local crab species demonstrated that lobsters actively and readily prey on Cancer irroratus (Say) and Carcinus maenas (L.), but were significantly less likely to consume Cancer borealis (Stimpson). The abundance of Carcinus maenas and blue mussels (Mytilus edulis L.) in the intertidal zone may explain the upshore movement of lobsters. Since nocturnal migration of Homarus americanus into the intertidal zone has not been documented before, our understanding of the dynamics of New England intertidal communities needs to be expanded to include this predator.     

Increase in density of genetically diverse invasive Asian shore crab (<Emphasis Type="Italic">Hemigrapsus sanguineus</Emphasis>) populations in the Gulf of Maine

Hemigrapsus sanguineus, the Asian shore crab, has rapidly replaced Carcinus maenas, the green crab, as the most abundant crab on rocky shores in the northwest Atlantic since its introduction to the United States (USA) in 1988. The northern edge of this progressing invasion is the Gulf of Maine, where Asian shore crabs are only abundant in the south. We compared H. sanguineus population densities to those from published 2005 surveys and quantified genetic variation using the cytochrome c oxidase subunit I gene. We found that the range of H. sanguineus had extended northward since 2005, that population density had increased substantially (at least 10-fold at all sites), and that Asian shore crabs had become the dominant intertidal crab species in New Hampshire and southern Maine. Despite the significant increase in population density of H. sanguineus, populations only increased by a factor of 14 in Maine compared to 70 in southern New England, possibly due to cooler temperatures in the Gulf of Maine. Genetically, populations were predominantly composed of a single haplotype of Japanese, Korean, or Taiwanese origin, although an additional seven haplotypes were found. Six of these haplotypes were of Asian origin, while two are newly described. Large increases in population sizes of genetically diverse individuals in Maine will likely have a large ecological impact, causing a reduction in populations of mussels, barnacles, snails, and other crabs, similar to what has occurred at southern sites with large populations of this invasive crab species.