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Multiple enveloped viruses with rod-shaped nucleocapsids have been described, infecting the epithelial cell nuclei within the hepatopancreas tubules of crustaceans. These bacilliform viruses share the ultrastructural characteristics of nudiviruses, a specific clade of viruses infecting arthropods. Using histology, electron microscopy and high throughput sequencing, we characterise two further bacilliform viruses from aquatic hosts, the brown shrimp (Crangon crangon) and the European shore crab (Carcinus maenas). We assembled the full double stranded, circular DNA genome sequences of these viruses (~113 and 132 kbp, respectively). Comparative genomics and phylogenetic analyses confirm that both belong within the family Nudiviridae but in separate clades representing nudiviruses found in freshwater and marine environments. We show that the three thymidine kinase (tk) genes present in all sequenced nudivirus genomes, thus far, were absent in the Crangon crangon nudivirus, suggesting there are twenty-eight core genes shared by all nudiviruses. Furthermore, the phylogenetic data no longer support the subdivision of the family Nudiviridae into four genera (Alphanudivirus to Deltanudivirus), as recently adopted by the International Committee on Taxonomy of Viruses (ICTV), but rather shows two main branches of the family that are further subdivided. Our data support a recent proposal to create two subfamilies within the family Nudiviridae, each subdivided into several genera.

Current concerns about climate change have led to intensive research attempting to understand how climate-driven stressors affect the performance of organisms, in particular the offspring of many invertebrates and fishes. Although stressors are likely to act on several stages of the life cycle, little is known about their action across life phases, for instance how multiple stressors experienced simultaneously in the maternal environment can modulate the responses to the same stressors operating in the offspring environment. Here, we study how performance of offspring of a marine invertebrate (shore crab Carcinus maenas ) changes in response to two stressors (temperature and salinity) experienced during embryogenesis in brooding mothers from different seasons. On average, offspring responses were antagonistic: high temperature mitigated the negative effects of low salinity on survival. However, the magnitude of the response was modulated by the temperature and salinity conditions experienced by egg-carrying mothers. Performance also varied among cohorts, perhaps reflecting genetic variation, and/or maternal conditions prior to embryogenesis. This study contributes towards the understanding of how anthropogenic modification of the maternal environment drives offspring performance in brooders.

This paper analyses the population dynamics, growth and feeding ecology of Dicentrarchus labrax in order to gain a better understanding of its present role in the western Dutch Wadden Sea ecosystem. Otolith analysis showed that the population is mostly comprised of individuals aged 3-5 years old and between 20 and 45cm in length. In autumn, 0-group juveniles are also an important part of the population. Both juveniles and adults use the area as a feeding ground exhibiting an opportunistic feeding strategy that relies on available prey, especially the brown shrimp Crangon crangon. Stomach content analysis and nitrogen stable isotope analysis showed an ontogenetic shift towards piscivory and a general decrease in the dominance of invertebrates with increasing size. Over the last 50 years, large between-year fluctuations in D. labrax abundance have been observed with an underlying increasing trend from about 1990 until 2007 followed by a subsequent decline. Spring abundance showed significant relationships with temperature and salinity while autumn abundance was only related to temperature. Spring and autumn D. labrax abundance were also strongly related to abundance of brown shrimp C. crangon prey. Long-term trends in temperature and salinity in the area suggest that environmental conditions for juvenile growth have become optimal, resulting in increased abundance since the mid-1980s. Continued monitoring of the dynamics of this species in the Dutch Wadden Sea is important to understand and anticipate the effects of climate change on the D. labrax population and its role in the local food web.

Volatile organic compounds (VOCs) characterize the spoilage of seafood packaged under modified atmospheres (MAs) and could thus be used for quality monitoring. However, the VOC profile typically contains numerous multicollinear compounds and depends on the product and storage conditions. Identification of potential spoilage indicators thus calls for multivariate statistics. The aim of the present study was to define suitable statistical methods for this purpose (exploratory analysis) and to consequently characterize the spoilage of brown shrimp (Crangon crangon) and Atlantic cod (Gadus morhua) stored under different conditions (selective analysis). Hierarchical cluster analysis (HCA), principal components analysis (PCA) and partial least squares regression analysis (PLS) were applied as exploratory techniques (brown shrimp, 4 °C, 50%CO2/50%N2) and PLS was further selected for spoilage marker identification. Evolution of acetic acid, 2,3-butanediol, isobutyl alcohol, 3-methyl-1-butanol, dimethyl sulfide, ethyl acetate and trimethylamine was frequently in correspondence with changes in the microbiological quality or sensory rejection. Analysis of these VOCs could thus enhance the detection of seafood spoilage and the development of intelligent packaging technologies. acceptedVersion

This paper analyses the population dynamics, growth and feeding ecology ofDicentrarchus labraxin order to gain a better understanding of its present role in the western Dutch Wadden Sea ecosystem. Otolith analysis showed that the population is mostly comprised of individuals aged 3–5 years old and between 20 and 45 cm in length. In autumn, 0-group juveniles are also an important part of the population. Both juveniles and adults use the area as a feeding ground exhibiting an opportunistic feeding strategy that relies on available prey, especially the brown shrimpCrangon crangon. Stomach content analysis and nitrogen stable isotope analysis showed an ontogenetic shift towards piscivory and a general decrease in the dominance of invertebrates with increasing size. Over the last 50 years, large between-year fluctuations inD. labraxabundance have been observed with an underlying increasing trend from about 1990 until 2007 followed by a subsequent decline. Spring abundance showed significant relationships with temperature and salinity while autumn abundance was only related to temperature. Spring and autumnD. labraxabundance were also strongly related to abundance of brown shrimpC. crangonprey. Long-term trends in temperature and salinity in the area suggest that environmental conditions for juvenile growth have become optimal, resulting in increased abundance since the mid-1980s. Continued monitoring of the dynamics of this species in the Dutch Wadden Sea is important to understand and anticipate the effects of climate change on theD. labraxpopulation and its role in the local food web.

Common seals Phoca vitulina give birth in the Wadden Sea area during the summer months. We provide evidence that the pupping season has advanced in date in this region. Analysis of stranding dates of recently-born, orphaned pups admitted for rehabilitation, revealed a shift of, on average, 0.88 days per year over the period 1974–2008, yielding a total advance of 26 days. Although the pupping season has become progressively earlier, there were no indications of any negative impact on the weight of the pups, nor was there an increase in the proportion of seals with a lanugo coat. These observations suggest that the most likely explanation for the change in phenology of the pupping season is a corresponding change in the timing of cessation of the period of delayed implantation. It is suggested that shifts in phenology could reflect an adaptive response of the animals to altered local circumstances. The latter may in turn be induced by larger scale phenomena such as food availability or climate change.

The bivalve Macoma balthica is a common species in the Wadden Sea and North Sea. Juveniles temporarily use nurseries in the high intertidal. To explain this nursery use, predation pressure was examined for both juvenile and adult Macoma at low and high tidal flats. The study was carried out in the eastern Dutch Wadden Sea. Shrimps Crangon crangon, adult crabs Carcinits maenas, gobies Pomatoschistus and juvenile flatfish were more abundant and larger on low than on high tidal flats, but 0-group Carcinus was more abundant on the high tidal flats. Crangon and 0-group Carcinits stomachs frequently contained Macoma remains. These predators selectively preyed on small 0-group Macoma, both in the field and in laboratory experiments. The effect of predation by epibenthic animals and birds, on the low and high tidal flats, was examined in exclosure experiments (2 mm mesh). There was no effect of epibenthos exclosure on adult Macoma. For 0-group Macoma, densities were higher in exclosures than in the controls where predators had normal access. The density reduction by epibenthic predators was much larger in the low than in the high intertidal. We found no effect of bird predation on densities of 0- and 1+ group Macoma. Thus, 0-group Macoma is under high predation pressure by epibenthos in the low intertidal, especially by shrimps, while they are relatively safe in the high intertidal. However, most of the shellfish outgrow their epibenthic predators during their first summer. Therefore, it becomes safe for the bivalves to redistribute to locations where epibenthic predators are abundant, during their first winter. On the other hand, it did not become clear from this study why many of the larger Macoma leave the high intertidal. Concluding, the nursery use of Macoma-spat in the high intertidal is probably, at least partly, an adaptation to avoid epibenthic predation. (C) 2002 Elsevier Science B.V. All rights reserved.

The bivalve Macoma balthica is a common species in the Wadden Sea and North Sea. Juveniles temporarily use nurseries in the high intertidal. To explain this nursery use, predation pressure was examined for both juvenile and adult Macoma at low and high tidal flats. The study was carried out in the eastern Dutch Wadden Sea. Shrimps Crangon crangon, adult crabs Carcinits maenas, gobies Pomatoschistus and juvenile flatfish were more abundant and larger on low than on high tidal flats, but 0-group Carcinus was more abundant on the high tidal flats. Crangon and 0-group Carcinits stomachs frequently contained Macoma remains. These predators selectively preyed on small 0-group Macoma, both in the field and in laboratory experiments. The effect of predation by epibenthic animals and birds, on the low and high tidal flats, was examined in exclosure experiments (2 mm mesh). There was no effect of epibenthos exclosure on adult Macoma. For 0-group Macoma, densities were higher in exclosures than in the controls where predators had normal access. The density reduction by epibenthic predators was much larger in the low than in the high intertidal. We found no effect of bird predation on densities of 0- and 1+ group Macoma. Thus, 0-group Macoma is under high predation pressure by epibenthos in the low intertidal, especially by shrimps, while they are relatively safe in the high intertidal. However, most of the shellfish outgrow their epibenthic predators during their first summer. Therefore, it becomes safe for the bivalves to redistribute to locations where epibenthic predators are abundant, during their first winter. On the other hand, it did not become clear from this study why many of the larger Macoma leave the high intertidal. Concluding, the nursery use of Macoma-spat in the high intertidal is probably, at least partly, an adaptation to avoid epibenthic predation. (C) 2002 Elsevier Science B.V. All rights reserved.

The population development of the 1998 year class of the bivalve Macoma balthicd was studied by repeated sampling of a tidal flat area in the eastern Dutch Wadden Sea from May 1998 to August 2000. The juveniles migrated twice, once in mid-1998 from their primary settlement locations in the low sandy intertidal to the nursery in the high intertidal (spring migration, 25 % of all juvenile Mbalthica relocated from low to high intertidal) and once in late 1998/early 1999 from the nursery to the low intertidal and the subtidal (winter migration, between 8 and 15 % of all 0-group (less than 1 yr old) M. balthica relocated from high to low intertidal). During winter, M. balthica migration was most intensive at the lowest temperatures, Relative to the abundance on the tidal flats, the 1+ group (older than 1 yr) M. balthica (i.e. from the second summer onwards) were rare in the tidal channels of the Wadden Sea and were only slightly more common in the North Sea adjacent to the tidal inlet. During both the spring and winter migration, many bivalves disappeared from the tidal flat population. This could partly be explained by normal mortality and by emigration to the subtidal channels and the North Sea. The remaining mortality was probably due to the risks inherent in migration: predation during pelagic floating or not reaching the right locations, The number of bivalves that disappeared was very high. In spring, slightly more disappeared than relocated. In winter, > 4 times more bivalves disappeared than relocated, Migration is obviously a very dangerous activity, therefore great advantages must be associated with the nursery use that makes such migrations necessary. An analysis of literature data on the density of M. balthica shows that in the eastern Dutch Wadden Sea, 90 % of the population lives in the intertidal, about 10% in the adjacent North Sea, and a negligible fraction in the subtidal channels of the Wadden Sea.

Beach nourishment is widely applied as a coastal protection measure because of its reduced ecological impact relative to hard coastal protection. In order to predict expected ecological impact on the sandy beach ecosystem, we developed a simulation model that integrates species envelope-based projections for the dominant macrobenthos species and mechanistic food web modules for higher trophic levels.Species envelopes are estimated by using Bayesian inference of species’ biomass relationships according to the three main determining abiotic variables, i.e. median grain size, total organic matter and the intertidal elevation, obtained from multiple independent sampling campaigns along the Belgian coast. Maximal possible abundances of higher trophic levels, being birds, shrimp and flatfish, are estimated based on their functional and energetic relationships with macrobenthos as an important food item.After validation, we demonstrate that sediment grain size is the most important factor determining beach-level diversity and production, with strong deterioration after nourishment with too coarse sediment (e.g. >> than 300 µm). Nourishment slope had a smaller impact on the species zonation patterns compared to the sediment grain size. Patterns for higher trophic levels do not follow the decreasing patterns in macrobenthos abundance and biomass.The advised gradient in sediment grain size for nourishment of fine-grained beaches is defined as 200 – 300 µm. This modelling approach shows that the assessment of ecosystem health needs to include the evaluation of different species richness and biomass variables. Focusing solely on for instance the potential abundance of species from higher trophic levels might lead to deceptive conclusions due to the dominance of opportunistic prey species.

Deposit-feeders are the dominant bioturbators of aquatic sediments, where they profoundly impact biogeochemical processes, but they are also vulnerable to both lethal and non-lethal predation by a large variety of predators. In this study, we performed a series of experiments to test the effects of predation avoidance on the feeding activity and sediment reworking intensity of the deposit-feeding bivalve Macoma balthica. Feeding activity at the sediment–water interface and sediment reworking intensity (vertical displacements of inert particle tracers) were monitored using image analysis techniques for treatments including and excluding the predatory shrimp, Crangon crangon. Detection of C. crangon by M. balthica resulted in an immediate retraction of the feeding siphon and a reduction in feeding activity. M. balthica also buried deeper into the sediment in the presence of C. crangon. This predator avoidance behaviour indirectly affected sediment reworking modes and rates, increasing the thickness of the bioturbated sediment layer as well as the non-local transport of sediment particles at depth. Conversely, feeding activity and sediment reworking processes remained unaffected when C. crangon was present, but isolated from the sediment, suggesting that predator perception in M. balthica is tactile (i.e. induced by direct encounter) rather than being chemosensory. Collectively, these results demonstrate that predatory avoidance behaviour by benthic infauna can significantly impact benthic bioturbation and the incorporation of organic matter into the benthic food web.

The bivalve Macoma balthica is a common species in the Wadden Sea and North Sea. Juveniles temporarily use nurseries in the high intertidal. To explain this nursery use, predation pressure was examined for both juvenile and adult Macoma at low and high tidal flats. The study was carried out in the eastern Dutch Wadden Sea. Shrimps Crangon crangon, adult crabs Carcinits maenas, gobies Pomatoschistus and juvenile flatfish were more abundant and larger on low than on high tidal flats, but 0-group Carcinus was more abundant on the high tidal flats. Crangon and 0-group Carcinits stomachs frequently contained Macoma remains. These predators selectively preyed on small 0-group Macoma, both in the field and in laboratory experiments. The effect of predation by epibenthic animals and birds, on the low and high tidal flats, was examined in exclosure experiments (2 mm mesh). There was no effect of epibenthos exclosure on adult Macoma. For 0-group Macoma, densities were higher in exclosures than in the controls where predators had normal access. The density reduction by epibenthic predators was much larger in the low than in the high intertidal. We found no effect of bird predation on densities of 0- and 1+ group Macoma. Thus, 0-group Macoma is under high predation pressure by epibenthos in the low intertidal, especially by shrimps, while they are relatively safe in the high intertidal. However, most of the shellfish outgrow their epibenthic predators during their first summer. Therefore, it becomes safe for the bivalves to redistribute to locations where epibenthic predators are abundant, during their first winter. On the other hand, it did not become clear from this study why many of the larger Macoma leave the high intertidal. Concluding, the nursery use of Macoma-spat in the high intertidal is probably, at least partly, an adaptation to avoid epibenthic predation. (C) 2002 Elsevier Science B.V. All rights reserved.