Short- and long-term dietary effects on disease and mortality in American lobster Homarus americanus

The American lobster Homarus americanus fishery is heavily dependent on the use of fish as bait to entice lobsters into traps. There is concern that this food supplementation is nutritionally insufficient for lobsters, but previous experiments reported conflicting results. We conducted a long-term feeding experiment in which 1 yr old American lobsters were fed one of 7 diets for a period of 352 d, a time that allowed the lobsters to molt thrice. The diets consisted of fresh frozen herring, a 'wild' diet (rock crab, mussel, and Spirulina algae), a formulated artificial diet for shrimp, paired combinations of these 3 diets or a diet formulated at the New England Aquarium (Artemia, fish and krill meal, Spirulina algae, soy lecithin, vitamins and minerals). The lobsters fed the diet of 100% fish had higher initial molting rates, but within the period of this experiment all either contracted shell disease or died. Mixed diets resulted in higher survival and a lower probability of mortality. This research demonstrated a critical time component to diet studies in lobsters. Short- and long-term impacts of diet differ. In the long term, continual high consumption rates of fish by the lobsters promote poor health in all lobsters, not just those of market size. The use of fish as bait may make lobsters more susceptible to the stress associated with environmental fluctuation, thereby leading to increased disease and mortality. This nutritional stress can be used to develop a laboratory model of shell disease in American lobsters.     

Use of Herring Bait to Farm Lobsters in the Gulf of Maine

Background

Ecologists, fisheries scientists, and coastal managers have all called for an ecosystem approach to fisheries management, yet many species such as the American lobster (Homarus americanus) are still largely managed individually. One hypothesis that has yet to be tested suggests that human augmentation of lobster diets via the use of Atlantic herring (Clupea harengus) as bait may contribute to recent increases in lobster landings. Currently 70% of Atlantic herring landings in the Gulf of Maine are used as bait to catch lobsters in traps throughout coastal New England.

Methodology/Principal Findings

We examined the effects of this herring bait on the diet composition and growth rate of lobsters at heavily baited vs. seasonally closed (i.e., bait free) sites in coastal Maine. Our results suggest that human use of herring bait may be subsidizing juvenile lobster diets, thereby enhancing lobster growth and the overall economic value and yield of one of the most valuable fisheries in the U.S.

Conclusions/Significance

Our study illustrates that shifting to an ecosystem approach to fisheries management should require consideration of cross-fishery interactions.     

Lesion bacterial communities in American lobsters with diet-induced shell disease

In southern New England, USA, shell disease affects the profitability of the American lobster Homarus americanus fishery. In laboratory trials using juvenile lobsters, exclusive feeding of herring Clupea harengus induces shell disease typified initially by small melanized spots that progress into distinct lesions. Amongst a cohabitated, but segregated, cohort of 11 juvenile lobsters fed exclusively herring, bacterial communities colonizing spots and lesions were investigated by denaturing gradient gel electrophoresis of 16S rDNA amplified using 1 group-specific and 2 universal primer sets. The Bacteroidetes and Proteobacteria predominated in both spots and lesions and included members of the orders Flavobacteriales (Bacteriodetes), Rhodobacterales, Rhodospirillales and Rhizobiales (Alphaproteobacteria), Xanthomonadales (Gammaproteobacteria) and unclassified Gammaproteobacteria. Bacterial communities in spot lesions displayed more diversity than communities with larger (older) lesions, indicating that the lesion communities stabilize over time. At least 8 bacterial types persisted as lesions developed from spots. Aquimarina 'homaria', a species commonly cultured from lesions present on wild lobsters with epizootic shell disease, was found ubiquitously in spots and lesions, as was the 'Candidatus Kopriimonas aquarianus', implicating putative roles of these species in diet-induced shell disease of captive lobsters.     

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

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

Laboratory comparison of American lobster,Homarus americanus,foraging responses to invasive green crab,Carcinus maenas,and two traditional finfish baits

The American Lobster (Homarus americanus) is trapped primarily with finfish baits that are becoming more costly and difficult to acquire as stocks dwindle. Thus, alternative bait options should be considered. We made video recordings of lobster foraging responses in the laboratory to compare the attractiveness of the invasive green crab (Carcinus maenas) bait, two traditional finfish baits and an inert control. Solitary lobsters approached and handled all three biological baits with similar latencies and spent similar time in contact with the biological baits. In contrast, lobsters took longer to approach and handle, and spent significantly less time in contact with control bait. As green crabs were statistically indistinguishable in attractiveness from traditional baits in our laboratory setting, we suggest field trials should test this alternate bait in circumstances more directly relevant to the fishery.     

Effect of natural and laboratory diet on O : N ratio in juvenile lobsters (Homarus americanus)

Oxygen consumption and ammonia excretion rates of juvenile lobsters spanning the size range from shelter-restricted to vagile (7-60 mm CL, carapace length) were measured. There are no significant size-dependent break points in the slope of oxygen consumption or nitrogen excretion rates versus weight in freshly caught wild or laboratory fed lobsters. This suggests that there are no inherent changes in energy metabolism in juvenile lobster in the 7-60 mm CL range (stage V to adolescent lobsters), despite major behavioral shifts related to foraging and shelter use. In addition, the O : N ratio of freshly caught wild lobsters (15.7+/-7.4, mean+/-S.D.) is not significantly different from that of lobsters fed brown algae (13.4+/-4.9) or a mixed carbohydrate/protein diet (13.7+/-6.8) in lab. The O : N ratio of wild lobsters is significantly higher than lobsters fed a high protein diet (7.5+/-2.5) in the lab. This suggests that carbohydrates, particularly algal-derived carbohydrates, make up a significant portion of the metabolic fuel of wild juvenile lobsters from settlement through adolescence.     

Transmission of Panulirus argus virus 1 (PaV1) and its effect on the survival of juvenile Caribbean spiny lobster

The Caribbean spiny lobster Panulirus argus, an important fisheries species, is host to Panulirus argus virus 1 (PaV1), a lethal, unclassified virus--the first found in any species of lobster--prevalent in juvenile lobsters. We describe a series of laboratory experiments aimed at assessing the likely modes of disease transmission, determining the survival of lobsters relative to each transmission pathway and identifying potential alternate hosts. Given evidence for lower prevalence of PaV1 in large lobsters, the effect of lobster size on susceptibility was also examined. Results demonstrated that PaV1 can be transmitted to juvenile lobsters via inoculation, ingestion of diseased tissue, contact with diseased lobsters and--among the smallest juveniles--through water over distances of a few meters. Contact and waterborne transmission, the most likely modes of transmission in the wild, were less efficient than inoculation or ingestion. Nevertheless, about half of the smallest lobsters in contact and waterborne trials contracted the disease and died within 3 mo. Other decapods that co-occur with P. argus (e.g. spotted lobster P. guttatus, stone crab Menippe mercenaria, channel crab Mithrax spinosissimus) did not acquire the disease after inoculation with PaV1-infected hemolymph. Our results confirmed that PaV1 is highly infectious and lethal to juvenile P. argus, particularly early benthic juveniles in the wild, and, hence, is a threat to mariculture.     

Growth, feeding frequency, protein turnover, and amino acid metabolism in European lobster Homarus gammarus L

The effect of feeding frequency on growth and protein metabolism in the European lobster, Homarus gammarus, was investigated. Fourth (IV) stage lobsters H. gammarus were fed individually a marine animal meal (herring/mussels meal) for 56 days. Feeding a daily ration equivalent to 10% of their body weight gave better growth than feeding daily rations of 5% and 20%. Protein synthesis rates were similar for the three food rations but protein growth rates were significantly lower and protein degradation rates highest in the 5% body weight per day ration group. The efficiency with which synthesised protein was retained as growth was found to be 38% in the in the 10% ratio group. Protein synthesis rates of lobsters were found to be lower than those for shrimps (Penaeus vannamei). The amino acid flux also suggests a lower protein conversion efficiency than shrimps P. vannamei. The results suggests that lobsters are slow, periodic feeders and that growth can be readily increased by manipulation of particular environmental factors such as feeding frequency.     

Conflict between two inshore fisheries: for whelk (Buccinum undatum) and brown crab (Cancer pagurus), in the southwest Irish Sea

Landings from the whelk fishery in the southwest Irish Sea rose from 63 t in 1990 to 6587 t in 1996 although landings declined to 2919 tonnes in 1998. Whelk are caught in pots baited with a combination of dogfish Scyliorhinus spp and brown crab Cancer pagurus. The input of bait crab is estimated at 7.2% of whelk landed. Most of the crab comes from southeast Ireland where some of it is a by-product of the claw fishery; some is targeted as bait crab, and this includes sub-size or recently moulted individuals. Fishing effort has been increasing on brown crab for human consumption also and there is concern about the sustainability of the crab fishery. Alternative bait trials examined a number of readily available fish species, some of them offals, whose attractiveness was evaluated by reference to crab and dogfish: whitefish (whiting and cod), pelagic species (mackerel and scad) and blue mussel. An artificial bait ration which included some crab meat, but 60% less than its natural equivalent, proved as effective as natural bait crab. A bait which consisted of a single species was less successful than two species fished together and some of the combinations (notably of whitefish) performed very effectively. Pelagic species performed poorly. The trade in brown crab for human consumption will always produce some form of discards which should be used; extending its use in an artificial bait is desirable.     

The shifting balance of littoral predator-prey interaction in regimes of hydrodynamic stress

Above lowshore levels of wave-beaten rocky shores, desiccation from tidal exposure and hydrodynamics stresses from wave action are thought to create refuges from predation, allowing concentrations of sedentary prey such as mussel beds. Underwater time-lapse photography on rocky shores in Southern California revealed that dense aggregations of spiny lobsters prey on mussels during nocturnal high tides. In contradiction of the refuge hypothesis, the densest aggregations occurred on midshore levels of the most wave-exposed site, a semi-protected site showed intermediate densities, and a protected site showed only sparse numbers of lobsters. On wave-beaten shores, the lobsters' high mobility and rapid prey handling allowed them to exploit intertidal prey in the brief period at extreme high tide, when both desiccation and hydrodynamic stresses were at a minimum. The spatial differences in lobster densities were, however, positively related to the recruitment rates of juvenile mussels, the preferred prey. A field experiment demonstrated that predation by lobsters within a mussel bed affects the age/size structure of the bed without changing primary percent coverage. Therefore, concentrations of adult prey on some wave-swept sites appear to result from elevated rates of prey recruitment that surpass rates of predation, rather than absolute refuges from predation.     

Isotopic discrimination between food and blood and feathers of captive penguins: implications for dietary studies in the wild

Using measurements of naturally occurring stable isotopes to reconstruct diets or source of feeding requires quantifying isotopic discrimination factors or the relationships between isotope ratios in food and in consumer tissues. Diet-tissue discrimination factors of carbon ((13)C/(12)C, or delta (13)C) and nitrogen ((15)N/(14)N, or delta (15)N) isotopes in whole blood and feathers, representing noninvasive sampling techniques, were examined using three species of captive penguins (king Aptenodytes patagonicus, gentoo Pygoscelis papua, and rockhopper Eudyptes chrysocome penguins) fed known diets. King and rockhopper penguins raised on a constant diet of herring and capelin, respectively, had tissues enriched in (15)N compared to fish, with discrimination factors being higher in feathers than in blood. These data, together with previous works, allowed us to calculate average discrimination factors for (15)N between whole lipid-free prey and blood and feathers of piscivorous birds; they amount to +2.7 per thousand and +4.2 per thousand, respectively. Both fish species were segregated by their delta (13)C and delta (15)N values, and importantly, lipid-free fish muscle tissue was consistently depleted in (13)C and enriched in (15)N compared to whole lipid-free fish. This finding has important implications because previous studies usually base dietary reconstructions on muscle of prey rather than on whole prey items consumed by the predator. We tested the effect of these differences using mass balance calculations to the quantification of food sources of gentoo penguins that had a mixed diet. Modeling indicated correct estimates when using the isotopic signature of whole fish (muscle) and the discrimination factors between whole fish (muscle) and penguin blood. Conversely, the use of isotopic signatures of muscle together with discrimination factors between whole fish and blood (or the reverse) leads to spurious estimates in food proportions. Consequently, great care must be taken in the choice of isotopic discrimination factors to apply to wild species for which no controlled experiments on captive individuals have been done. Finally, our results also indicate that there is no need to remove lipids before isotopic analysis of avian blood.     

Effects of food quality on tissue-specific isotope ratios in the mussel <Emphasis Type="Italic">Perna perna</Emphasis>

Investigations into trophic ecology and aquatic food web resolution are increasingly accomplished through stable isotope analysis. The incorporation of dietary and metabolic changes over time results in variations in isotope signatures and turnover rates of producers and consumers at tissue, individual, population and species levels. Consequently, the elucidation of trophic relationships in aquatic systems depends on establishing standard isotope values and tissue turnover rates for the level in question. This study investigated the effect of diet and food quality on isotopic signatures of four mussel tissues: adductor muscle, gonad, gill and mantle tissue from the brown mussel Perna perna. In the laboratory, mussels were fed one of the two isotopically distinct diets for 3 months. Although not all results were significant, overall δ¹³C ratios in adductor, mantle and gill tissues gradually approached food source signatures in both diets. PERMANOVA analyses revealed significant changes over time in tissue δ¹³C (mantle and gill) with both diets and in δ¹⁵N (all tissues) and C:N ratios (mantle and gill) for one diet only. The percentage of replaced carbon isotopes were calculated for the 3 month period and differed among tissues and between diets. The tissue with the highest and lowest amount of replaced isotopes over 81 days were mantle tissue on the kelp diet (33.89%) and adductor tissue on the fish food diet (4.14%), respectively. Percentages could not be calculated for any tissue in either diet for δ¹⁵N due to the lack of significant change in tissue nitrogen. Fractionation patterns in tissues for both diets can be linked to nutritional stress, suggesting that consumer isotopic signatures are strongly dependent on food quality, which can significantly affect the degree of isotopic enrichment within a trophic level.     

Testing baits prepared from by‐product of the shrimp and snow crab industry in the pot fishery for Gadus morhua (Linnaeus, 1758) and Pollachius virens (Linnaeus, 1758)

In a study between October and November 2015 in northern Norway, bait prepared from the by‐product of the shrimp and snow crab industry was employed in the pot fishery to capture cod and saithe. Frozen natural herring was used as control bait. Pots with the three bait types were placed 1,000 m apart in order to eliminate the risk of interaction between the different baits. In total 11 sampling periods were conducted. There were no significant differences in catch rates between the different baits tested on cod and saithe. The manufactured bait tested in this study showed a continuous release of odours over a 14‐day period. Results show that the water content of the bait increases rapidly in the first phase of the immersion (until day 7), and the corresponding loss of bait mass (dry weight) suggest some physical disintegration (erosion) of the bait until the water content stabilizes. This study illustrates that it is possible to replace natural herring with hydrolysates from by‐products from shrimp or snow crab industry in the Norwegian pot fishery of cod and saithe.     

INTERACTIONS BETWEEN THE INDIAN RIVER LAGOON BLUE CRAB FISHERY AND THE BOTTLENOSE DOLPHIN, TURSIOPS TRUNCATUS

Anecdotal reports from blue crab fishermen in the Indian River Lagoon (IRL), Florida suggested that bottlenose dolphins ( Tursiops truncatus ) followed their boats, and stole bait fish from crab pots soon after they were deployed. To investigate these reports, we made biweekly observations from IRL commercial crab boats from January 1998 to January 1999 (670 h). Only 2.8% of the 18,891 crab pots surveyed revealed evidence of dolphin/crab pot interaction. Dolphin interactions included: (1) begging at boats, (2) feeding on discarded bait fish, (3) engaging in crab pot tipping behavior, and (4) dolphin mortality from crab pot float line entanglement. Overall, 16.6% of the 1,296 dolphins sighted interacted with fishing boats. Seasonal trends were evident, with fishery interactions peaking in the summer. Crab pot interactions ranged from 0% to 36% of the traps checked daily. Different methods of securing the bait-well door and the role of trap locality were tested using a replicated experimental design. Results showed significant differences in successful bait removal by dolphins, among the degrees of door security ( P < 0.001) and between trap location ( P < 0.01). Thus, increased door security may help to reduce the negative impacts to the fishery and dolphins involved.     

Competition with stone crabs drives juvenile spiny lobster abundance and distribution

Interspecific competition is assumed to have a strong influence on the population dynamics of competing species, but is not easily demonstrated for mobile species in the wild. In the Florida Keys (USA), anecdotal observations have long pointed to an inverse relationship in abundance of two large decapod crustaceans found co-occurring in hard-bottom habitat, the stone crab Menippe mercenaria and the Caribbean spiny lobster Panulirus argus. We used them to explicitly test whether competition for a renewable resource (shelter) can drive the abundance and distribution of the inferior competitor. We first explored this relationship in shelter competition mesocosm experiments to determine the competitively dominant species. Results showed that stone crabs are clearly the dominant competitors regardless of the number of lobsters present, the presence of co-sheltering species such as the spider crab, Damithrax spinosissimus, or the order of introduction of competitors into the mesocosm. We also found that lobsters use chemical cues from stone crabs to detect and avoid them. We then tested the ramifications of this competitive dominance in the field by manipulating stone crab abundance and then tracking the abundance and distribution of spiny lobsters through time. Increased stone crab abundance immediately resulted in decreased lobster abundance and increased aggregation. The opposite occurred on sites where stone crabs were removed. When we stopped removing stone crabs from these sites, they soon returned and lobster abundance decreased. This study explicitly demonstrated that interspecific competition can drive population dynamics between these species, and ultimately, community composition in these shallow water habitats.     

Predation Risk within Fishing Gear and Implications for South Australian Rock Lobster Fisheries

Depredation of southern rock lobster (Jasus edwardsii) within fishing gear by the Maori octopus (Pinnoctopus cordiformis) has economic and ecological impacts on valuable fisheries in South Australia. In addition, depredation rates can be highly variable resulting in uncertainties for the fishery. We examined how in-pot lobster predation was influenced by factors such as lobster size and sex, season, fishing zone, and catch rate. Using mixed modelling techniques, we found that in-pot predation risk increased with lobster size and was higher for male lobsters. In addition, the effect of catch rate of lobsters on predation risk by octopus differed among fishing zones. There was both a seasonal and a spatial component to octopus predation, with an increased risk within discrete fishing grounds in South Australia at certain times of the year. Information about predation within lobster gear can assist fishery management decision-making, potentially leading to significant reduction in economic losses to the fishery.     

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.     

Characterization of Vibrio fluvialis-like strains implicated in limp lobster disease

Studies were undertaken to characterize and determine the pathogenic mechanisms involved in a newly described systemic disease in Homarus americanus (American lobster) caused by a Vibrio fluvialis-like microorganism. Nineteen isolates were obtained from eight of nine lobsters sampled. Biochemically, the isolates resembled V. fluvialis, and the isolates grew optimally at 20 degrees C; none could grow at temperatures above 23 degrees C. The type strain (1AMA) displayed a thermal reduction time (D value) of 5.77 min at 37 degrees C. All of the isolates required at least 1% NaCl for growth. Collectively, the data suggest that these isolates may embody a new biotype. Pulsed-field gel electrophoresis (PFGE) analysis of the isolates revealed five closely related subgroups. Some isolates produced a sheep hemagglutinin that was neither an outer membrane protein nor a metalloprotease. Several isolates possessed capsules. The isolates were highly susceptible to a variety of antibiotics tested. However, six isolates were resistant to erythromycin. Seventeen isolates harbored plasmids. Lobster challenge studies revealed that the 50% lethal dose of a plasmid-positive strain was 100-fold lower than that of a plasmid-negative strain, suggesting that the plasmid may enhance the pathogenicity of these microorganisms in lobsters. Microorganisms that were recovered from experimentally infected lobsters exhibited biochemical and PFGE profiles that were indistinguishable from those of the challenge strain. Tissue affinity studies demonstrated that the challenge microorganisms accumulated in heart and midgut tissues as well as in the hemolymph. Culture supernatants and polymyxin B lysates of the strains caused elongation of CHO cells in tissue culture, suggesting the presence of a hitherto unknown enterotoxin. Both plasmid-positive and plasmid-negative strains caused significant dose-related intestinal fluid accumulations in suckling mice. Absence of viable organisms in the intestinal contents of mice suggests that these microorganisms cause diarrhea in mice by intoxication rather than by an infectious process. Further, these results support the thermal reduction data at 37 degrees C and suggest that the mechanism(s) that led to fluid accumulation in mice differs from the disease process observed in lobsters by requiring neither the persistence of viable microorganisms nor the presence of plasmids. In summary, results of lobster studies satisfy Koch's postulates at the organismal and molecular levels; the findings support the hypothesis that these V. fluvialis-like organisms were responsible for the originally described systemic disease, which is now called limp lobster disease.     

Diel variation in feeding rate and prey composition of herring and mackerel in the southern Gulf of St Lawrence

Diel feeding patterns of herring Clupea harengus and mackerel Scomber scombrus in the southern Gulf of St Lawrence were examined based on samples obtained by midwater trawling between 19 and 26 June 2001. Within 3 h time periods, stomach contents tended to be more similar between fish from the same tow than between fish from different tows. Thus, in contrast to previous diet studies, which have used individual fish stomachs as independent observations, tow was used as the experimental unit in statistical analyses in this study. Diel patterns in stomach fullness were identified using generalized additive models. Two peaks in stomach fullness occurred for herring, one in the morning and the other in the evening. Mackerel showed an increase in feeding intensity throughout the day with a peak in mid‐afternoon. The diel changes in stomach contents suggested rapid gastric evacuation rates for both species, especially for herring. The estimate of the instantaneous evacuation rate for herring was twice that for mackerel. Calanus copepods (mainly C. hyperboreus ), fishes (mainly capelin Mallotus villosus ) and euphausiids were the main prey found in the stomachs of both species. Calanus copepods dominated the diet of herring regardless of time period. They also dominated the diet of mackerel during the late afternoon, evening and night while fishes and euphausiids were dominant during the morning and early afternoon. These diel patterns emphasize the need for sampling throughout the day and night in order to estimate ration and diet composition for bioenergetic and ecosystem models.     

A long-term mesocosm study on the settlement and survival of juvenile European lobster Homarus gammarus L. in four natural substrata

To date, the natural substratum preferences of early benthic phase (EBP) European lobsters (Homarus gammarus) remain largely unknown. This study utilised a large scale mesocosm experiment to determine if the animal favours cobble ground, similar to its American counterpart (Homarus americanus), or has other substratum preferences. Postlarvae were provided with the choice of settling on four natural substrata: sand, coralline algae, mussel shell and cobble. Over a nine month period, the number and size of juveniles on each substratum was recorded, with loss of chelipeds used as an indication of social interaction. After a 30 day period, a non-random distribution of lobsters was observed on the four substrata. Juveniles were more abundant in substrata which provided pre-existing shelter in the form of interstitial spaces, i.e. cobble and mussel shell, than in sand or coralline algae. The survival of individuals from postlarvae to 30 day old juveniles ranged from 5 to 14% with surviving benthic recruits showing a clear mode at 6-8 mm carapace length (CL) in size distribution. The density of lobsters per m(2) of cobble remained relatively constant (18/m(2)) throughout the study period while the density of juveniles on mussel shell decreased significantly (35 to 5/m(2)). The size distribution of lobsters on each substratum also varied with time. By the conclusion of the trial, lobsters found in mussel shell had a mode of 8-10 mm CL within a range of 6-14 mm CL while those in cobble had a mode of 10-12 mm CL within a range of 8-24 mm CL. Overall, the results underline the importance of shelter-providing habitat such as cobble or crevice-type substrata to EBP European lobsters. They also confirm that for a shelter-dwelling animal such as a lobster, the physical structure of the habitat is a key factor in determining both the size and number of its inhabitants.