The first occurrence of the ctenophore <Emphasis Type="Italic">Mnemiopsis leidyi</Emphasis> in the North Sea

After the discovery of large densities of Mnemiopsis leidyi in the Baltic Sea near Kiel by Javidpour et al. (First record of Mnemiopsis leidyi A. Agassiz 1865 in the Baltic Sea, 2006) in October 2006, we investigated the gelatinous zooplankton in the North Sea near Helgoland and recorded Mnemiopsis leidyi for the first time in the North Sea, albeit in much lower densities than those recorded in the Baltic Sea.     

The diet of Common Guillemot Uria aalge chicks provides evidence of changing prey communities in the North Sea

Local differences in feeding conditions have been suggested as a cause of regional variation in seabird demography but multi‐colony comparisons of diet are rare. In UK waters the main fish eaten by seabirds during the breeding season belong to three families: Ammodytidae, Clupeidae and Gadidae. Climate change and fishing are affecting these fish stocks and so probably impact on predators such as seabirds. We used standardized observations of prey brought in for chicks to make the first integrated assessment of the diet of Common Guillemot Uria aalge chicks at a UK scale. Chick diet varied markedly among the 23 colonies sampled between 2006 and 2011. Sandeels (Ammodytidae), probably Lesser Sandeels Ammodytes marinus, were the commonest prey. Their contribution to the diet varied both latitudinally and among marine regions, with the proportion significantly higher for a given latitude on the west coast compared to the east. The non‐sandeel component of the diet showed latitudinal changes, with small clupeids, probably Sprats Sprattus sprattus, predominant at southern colonies whereas juvenile gadids were the main alternative to sandeels in the north. Comparison of our Guillemot chick diet with data collected 15–30 years earlier suggests that the proportion of sandeels in the diet has decreased at colonies bordering the North Sea. No significant change was apparent in Atlantic colonies but historical data were limited. The early years of our study coincided with a population explosion of Snake Pipefish Entelurus aequoreus in the Northeast Atlantic and North Sea. Pipefish were recorded in Guillemot chick diet at several northern and northwestern colonies in 2006 and 2007 but have been absent since 2009. Spatial and temporal variation in chick diet accorded broadly with patterns expected as a result of rising sea temperatures and impacts of fishing. Guillemot chick diet could potentially be a useful indicator of changes in the distribution and abundance of forage fish.     

The diet of whiting Merlangius merlangus in the western Baltic Sea

The diet of whiting Merlangius merlangus in the western Baltic Sea was investigated and compared to the diet in the southern North Sea. Clupeids were important prey in both areas, but especially in the western Baltic Sea where they constituted up to 90% of the diet of larger individuals. Gobies, brown shrimps and polychaetes were the main prey of juveniles in the western Baltic Sea, while a wider range of species were consumed in the North Sea. The shift to piscivory occurred at smaller sizes in the western Baltic Sea and the fish prey consumed was proportionately larger than in the southern North Sea. Estimates of prey abundance and food intake of M. merlangus are required to evaluate its predatory significance in the western Baltic Sea, but its diet suggests that it could be just as significant a fish predator here as in the southern North Sea.     

Rapid increase of the grey seal (<Emphasis Type="Italic">Halichoerus grypus</Emphasis>) breeding stock at Helgoland

Over the past 25 years, the grey seal (Halichoerus grypus) population in the North Sea increased on average at 5% per year, while founding a number of new colonies. Based on published data, this note compares recent trends in pup production at nine breeding sites, with focus on the German island of Helgoland. Since 1997, numbers born at Helgoland increased at 35% per year (95% CI: 29–42), reaching an all-time high of 53 in 2007. Yet, this was less than 1% of all grey seal offspring in the North Sea area. Some other small stocks had similarly high growth rates of 20–50%, which must be explained by immigration from the larger colonies, as sustained autochthonous increase cannot exceed 13% per year in the species. Further, the Helgoland breeding stock increased significantly faster than the neighbouring colonies at Amrum (13%) and in the Dutch Wadden Sea (20%), indicating higher offspring survival and/or higher relative influx of migrants. This is attributed to the fact that at Helgoland newborn seals are safe from floodwater. In the other two sites, in contrast, pups are born on sandbanks and are thus often hit by winterly storm tides, which poses a risk to their survival. Because of the favourable breeding conditions at Helgoland, this new colony is expected to soon become an important stronghold of the grey seal in the Dutch-German North Sea area.     

Linking sandeel consumption and the likelihood of starvation in harbour porpoises in the Scottish North Sea: could climate change mean more starving porpoises?

Sandeels are known to be negatively affected by climate change in a number of ways. This study investigated whether these changes are affecting the harbour porpoise (Phocoena phocoena), a species which consumes sandeels. Porpoise diet was examined in spring (March-May), a critical time of year for survival when sandeels are important prey, from 1993 to 2001 to provide baseline information on the proportion of sandeels consumed. When data from spring 2002 and 2003 were compared to these baseline data, the diet was found to be substantially different, with a significant and substantially smaller proportion of sandeels being consumed in March and May. There were also differences in the number of porpoises starving between the two time periods (33% in spring 2002 and 2003 died of starvation, but only 5% in the baseline period). This suggests that a lower proportion of sandeels in the diet of porpoises in spring increases the likelihood of starvation. Therefore, we suggest that the negative effects of climate change on sandeel availability may have serious negative effects on harbour porpoise populations in the North Sea by increasing the likelihood of starvation in spring.     

Seabirds and sandeels: the conflict between exploitation and conservation in the northern North Sea

Sandeels, especiallyAmmodytes marinus are a major component of the diet of many predatory fish, seabirds and seals. The industrial fishery for sandeels is now the largest of the North Sea fisheries. A sandeel fishery in the Shetland area began in 1974 but has recently declined. This change was accompanied by dramatic declines in the breeding success of certain seabirds, particularly Arctic terns and kittiwakes. Current information on seabirds and sandeels in the Shetland area is reviewed and areas where further research is needed, highlighted. The Shetland problem illustrates the difficulties of reconciling conservation and exploitation when fundamental ecological and behavioural knowledge is lacking, and also the need to obtain further information on the ecological impact of industrial fisheries.     

Does diet in Celtic Sea fishes reflect prey availability?

Feeding preferences of Celtic Sea fishes were investigated using a database of stomach content records, collected between 1977 and 1994. The diet of cod Gadus morhua, hake Merluccius merluccius, megrim Lepidorhombus whiffiagonis, whiting Merlangius merlangus and saithe Pollachius virens changed markedly as the animals grew larger, and although large predators generally chose larger bodied prey, the variability of prey sizes consumed also increased. Large predators continued to select small, low value, benthic prey (e.g. Callionymus spp. and Trisopterus spp.) which were easier to catch, rather than larger, more energy lucrative pelagic prey (e.g. mackerel Scomber scombrus), even though these pelagic prey‐fishes were nearly always available and were often very abundant. Stock estimates of the International Council for the Exploration of the Sea and U.K. groundfish survey catches were used as indices of prey abundance. Blue‐whiting Micromesistius poutassou and other small pelagic fishes (Argentina spp. and clupeoids) were identified as being particularly important, and were consumed by some predators more often than would be expected given the abundance of these prey in the environment. There was no evidence for density‐dependent feeding by predators on mackerel and only hake exhibited density‐dependent feeding on horse‐mackerel. Hake, cod and megrim consumed more blue‐whiting when this prey was at higher abundance in the environment. In choosing what prey to consume, predators must balance costs and benefits, considering the quality of prey and the energy expended during search, capture and handling.     

Predation of whiting and haddock on sandeel: aggregative response, competition and diel periodicity

The predation of haddock Melanogrammus aeglefinus and whiting Merlangius merlangus on sandeel Ammodytes marinus was investigated based on intensive sampling [performed with a bottom trawl (GOV) equipped with a small meshed codend cover; sediment samples were taken with a van Veen grab] in a restricted area of c . 15 × 20 nautical miles in the northern North Sea during a 5 day period in July 1996. The analysis of the spatial distribution of predators revealed a pronounced aggregation of whiting in the south‐west part of the area, where sandeel catches were also highest. This pattern was thought to be the result of an aggregative behaviour of whiting. The sandeel concentration most likely reflected a restricted patch of coarse sediment of the preferred grain size for sandeels to bury in. In haddock the aggregation was less obvious. Both predators fed almost exclusively on sandeel in the south‐west part of the area with haddock stomachs containing more sandeel than whiting stomachs. The stomach contents in both predators increased rapidly during the night, indicating that the predators were targeting burying sandeels. This would explain the competitive advantage of the benthivorous haddock.     

Consumption rates and prey preference of the invasive gastropod Rapana venosa in the Northern Adriatic Sea

The alien Asian gastropod Rapana venosa (Valenciennes 1846) was first recorded in 1973 along the Italian coast of the Northern Adriatic Sea. Recently, this predator of bivalves has been spreading all around the world oceans, probably helped by ship traffic and aquaculture trade. A caging experiment in natural environment was performed during the summer of 2002 in Cesenatico (Emilia-Romagna, Italy) in order to estimate consumption rates and prey preference of R. venosa. The prey items chosen were the Mediterranean mussel Mytilus galloprovincialis (Lamarck 1819), the introduced carpet clam Tapes philippinarum (Adams and Reeve 1850), both supporting the local fisheries, and the Indo-Pacific invasive clam Anadara (Scapharca) inaequivalvis (Bruguière 1789). Results showed an average consumption of about 1 bivalve prey per day (or 1.2 g wet weight per day). Predation was species and size selective towards small specimens of A. inaequivalvis; consumption of the two commercial species was lower. These results might reduce the concern about the economical impact on the local bivalve fishery due to the presence of the predatory gastropod. On the other hand, selective predation might probably alter local community structure, influencing competition amongst filter feeder/suspension feeder bivalve species and causing long-term ecological impact. The large availability of food resource and the habitat characteristics of the Emilia-Romagna littoral makes this area an important breeding ground for R. venosa in the Mediterranean Sea, thus worthy of consideration in order to understand the bioinvasion ecology of this species and to control its likely further dispersal.     

The diet of king eiders wintering in Nuuk,Southwest Greenland,with reference to sympatric wintering common eiders

Coastal and offshore waters of Southwest Greenland are internationally important wintering areas for king eiders (Somateria spectabilis) breeding in eastern Canadian Arctic and in northwestern Greenland. This paper presents the first assessment of their winter diet. Based on esophageal-proventricular samples from 26 females (13 juveniles and 13 older birds) and 15 males (11 juveniles and four older birds) collected in 2000–2002 (November–May) in coastal waters of Nuuk, we identified 28 prey species. The diet consisted of almost equal proportions (aggregate fresh mass) of polychaetes, echinoderms, crustaceans and molluscs. The dominant prey species were Pectinaria spp. (26.8%), Strongylocentrotus droebachiensis (18.4%), Mya eideri (11.2%) and Hyas araneus (9.7%). The polychaetes have previously been identified as important prey for eiders in Greenland, but apparently not outside Greenland. Compared with a diet study of common eiders Somateria mollissima from the same wintering area, the king eiders consumed significantly less bivalves and significantly more echinoderms. This difference corresponded with observations that common eiders were feeding in shallow waters, while king eiders were feeding in deeper waters farther from the shore. Benthic surveys are needed to confirm that diet corresponds with prey availability.     

Diet composition and food consumption rate of harbor porpoises (Phocoena phocoena) in the western Baltic Sea

Stomach content composition and prey‐specific consumption rates of juvenile and adult harbor porpoises (Phocoena phocoena) were estimated from a data set including 339 stomachs collected over a 32 yr period (1980–2011) in the western Baltic Sea. The stomach contents were mainly hard parts of fish prey and in particular otoliths. The bias originating from differential residence time of otoliths in the stomachs was addressed by use of a recently developed approach. Atlantic cod and herring were the main prey of adults, constituting on average 70% of the diet mass. Juvenile porpoises also frequently consumed gobies. Here, the mass contribution by gobies was on average 25%, which was as much as cod. Other species such as whiting, sprat, eelpout, and sandeels were of minor importance for both juveniles and adults. The diet composition differed between years, quarters, and porpoise acquisition method. Yearly consumption rates for porpoises in the western Baltic Sea were obtained in three scenarios on the daily energy requirements of a porpoise in combination with an estimate including the 95% CLs of the porpoise population size. Cod of age groups 1 and 2 and intermediate‐sized herring suffered the highest predation from porpoises.     

Diet of harbor porpoises along the Dutch coast: A combined stable isotope and stomach contents approach

High stranding frequency of porpoises, Phocoena phocoena, along the Dutch coast since 2006 has led to increased interest in the ecology of porpoises in the North Sea. Stranded porpoises were collected along the Dutch coast (2006–2008) and their diet was assessed through stomach content and stable isotope analysis (δ¹³C and δ¹⁵N) of porpoise muscle and prey. Stable isotope analysis (SIAR) was used to estimate the contribution of prey species to the porpoises' diet. This was compared to prey composition from stomach contents, to analyze differences between long‐ and short‐term diet. According to stomach contents, 90.5% of the diet consisted of gobies, whiting, lesser sandeel, herring, cod, and sprat. Stable isotope analysis revealed that 70‐83% of the diet consisted of poor cod, mackerel, greater sandeel, lesser sandeel, sprat, and gobies, highlighting a higher importance of pelagic, schooling species in the porpoises' diet compared to stomach contents. This could be due to prey distribution as well as differences in behavior of porpoises and prey between the coastal zone and offshore waters. This study supports the need for multi‐method approaches. Future ecological and fishery impact assessment studies and management decisions for porpoise conservation should acknowledge this difference between the long‐ and short‐term diet.     

Diet comparison between pelagic and demersal whiting in the North Sea

Pelagic North Sea whiting Merlangius merlangus preyed upon (pelagic) sprat and sandeel, while demersal whiting preyed upon (demersal) Norway pout and herring. Values of diet breadth were low for both feeding groups using Levin's index. Diet overlaps within layers were low ( D <0·25), while the between layer food overlap was moderate ( D =0·25–0·74) to high ( D >0·74) using Schoener's index. Selection of prey was density dependent. However, prey size also played an important role. The diet of whiting shifted from amphipods and mysids to fish with increasing predator length, and the length of prey consumed increased significantly with length of whiting. The fact that the stomach contents differed between the feeding groups demonstrates the need for a sampling design that includes both pelagic and demersal habitats when trying to quantify the diet of whiting.     

First report of the oriental shrimp <Emphasis Type="Italic">Palaemon macrodactylus</Emphasis> Rathbun, 1902 (Decapoda,Caridea, Palaemonidae) from German waters

The native East Asian shrimp Palaemon macrodactylus has become a common inhabitant of estuaries along the Pacific coast of North America. More recently (documented since 1999), the species has also been colonising European waters and has been reported from Spain, England, Belgium and the Netherlands. In this study, we present a chronology of the reported introductions of this species and provide the first detailed report of its occurrence in German waters. P. macrodactylus was found in the Geeste river mouth (Weser Estuary) as well as in Hooksiel, north of Wilhelmshaven between 2004 and 2005. We assume its presence in other estuarine habitats of the North Sea and predict its introduction into the Baltic Sea.     

VARIABILITY IN THE DIET OF HARBOR PORPOISES (PHOCOENA PHOCOENA) IN SCOTTISH WATERS 1992–2003

Interannual, seasonal, and regional variation in the diet of porpoises, Phocoena phocoena , in Scottish (UK) waters was studied using stomach contents of animals stranded between 1992 and 2003. Most samples came from the east coast (including many porpoises killed by bottlenose dolphins), with smaller numbers from the west coast and from Shetland. The most important prey types, in terms of contribution by number and mass, were whiting ( Merlangius merlangus ) and sand eels (Ammodytidae). Multivariate analysis confirmed the existence of regional, seasonal, and interannual variation in diet, as well as differences ( i. e. , biases) related to cause of death. These differences were further explored using univariate analyses. Sand eels were more important in the summer months (quarters 2 and 3) and on the east coast, whereas gadids were more important in winter and in the Shetland area. Some, but not all, observed trends in the numerical importance and size of prey taken were consistent with trends in abundance and size of fish taken during research trawl surveys. There was some evidence that porpoises <1 yr old took more gobies (Gobiidae) and shrimps than did older porpoises. Clupeids (herring Clupea harengus and sprat Sprattus sprattus ) formed a relatively small proportion of the diet, but their importance varied from year to year. Although possible methodological biases prevent firm conclusions, it appears that the importance of clupeids in porpoise diet may have decreased since the 1960s, mirroring the decline in North Sea herring abundance. The recovery of the North Sea herring stock in recent years is not as yet reflected in porpoise diet.     

Chick starvation in yellow‐eyed penguins: Evidence for poor diet quality and selective provisioning of chicks from conventional diet analysis and stable isotopes

The link between poor reproductive success and diet was investigated in yellow‐eyed penguins Megadyptes antipodes, by assessing diet at two localities separated by about 30 km: the north coast of Stewart Island where breeding success is low (0.38–0.67 chicks per pair in recent years), and Codfish Island where breeding success is higher (0.96–1.51 chicks per pair), and relating this to published data from South Island localities, where average breeding success was 1.1 chicks per pair. Diet composition, meal sizes and energetic content of meals and prey were determined from stomach contents, and stable isotope analyses of chick down, fledgling feathers and adult blood provided information on diet throughout the fledging period. The high proportion of stomachs that were empty or lacked diagnostic remains reduced sample size considerably, and variability between samples reduced the power to detect significant differences in meal size, proportions of empty stomachs and prey diversity of meals. Energetic content of Stewart Island meals was less than Codfish Island meals, and there was a non‐significant trend for smaller meal sizes and reduced prey diversity among Stewart Island samples. Both localities had lower prey diversity and smaller meals than South Island penguins. Blue cod Parapercis colias accounted for 99% of prey biomass in Stewart Island and 70% in Codfish Island stomach samples, where 27% of prey biomass was opalfish Hemerocoetes monopterygius. Isotopic mixing models carried out on larger sample sizes indicated that opalfish comprised a large proportion of the diet at both locations, with adults selectively provisioning chicks with opalfish while feeding mainly on blue cod themselves. We suggest the large blue cod consumed by Codfish Island and Stewart Island penguins, larger than those consumed by South Island penguins, is difficult to transfer to chicks by regurgitation. Oyster dredging around Stewart Island may have reduced the availability and abundance of alternative prey to Stewart Island penguins.     

Short-term variation of nutritive and metabolic parameters in <Emphasis Type="Italic">Temora longicornis</Emphasis> females (Crustacea,Copepoda) as a response to diet shift and starvation

Changes in fatty acid patterns, digestive and metabolic enzyme activities and egg production rates (EPR) were studied in the small calanoid copepod Temora longicornis. Female copepods were collected in spring 2005 off Helgoland (North Sea). In the laboratory one group of copepods was fed with the cryptophycean Rhodomonas baltica for a period of 3 days. Another group of copepods was maintained without food. According to the fatty acid patterns, animals from the field were feeding on a more detrital, animal-based and to a minor extent to a diatom-based diet. Under laboratory conditions, females rapidly accumulated fatty acids such as 18:4 (n-3), 18:3 (n-3) and 18:2 (n-6) which are specific of R. baltica. Diatom-specific fatty acids such as 16:1 (n-7) were strongly reduced. In fed animals the activities of digestive and metabolic enzymes remained constant and egg production rates were highest on day 2. Starving animals, in contrast, showed significantly reduced faecal pellet production and EPR. Proteolytic enzyme activity decreased rapidly within 24 h and remained at a low level until the end of the experiment. Citrate synthase decreased continuously as well. T. longicornis rapidly reacts to dietary changes and food depletion. It has limited energy stores and, thus, strongly depends on continuous food supply.     

Multicolony tracking reveals the winter distribution of a pelagic seabird on an ocean basin scale

Aim An understanding of the non‐breeding distribution and ecology of migratory species is necessary for successful conservation. Many seabirds spend the non‐breeding season far from land, and information on their distribution during this time is very limited. The black‐legged kittiwake, Rissa tridactyla, is a widespread and numerous seabird in the North Atlantic and Pacific, but breeding populations throughout the Atlantic range have declined recently. To help understand the reasons for the declines, we tracked adults from colonies throughout the Atlantic range over the non‐breeding season using light‐based geolocation. Location North Atlantic. Methods Geolocation data loggers were deployed on breeding kittiwakes from 19 colonies in 2008 and 2009 and retrieved in 2009 and 2010. Data from 236 loggers were processed and plotted using GIS. Size and composition of wintering populations were estimated using information on breeding population size. Results Most tracked birds spent the winter in the West Atlantic, between Newfoundland and the Mid‐Atlantic Ridge, including in offshore, deep‐water areas. Some birds (mainly local breeders) wintered in the North Sea and west of the British Isles. There was a large overlap in winter distributions of birds from different colonies, and colonies closer to each other showed larger overlap. We estimated that 80% of the 4.5 million adult kittiwakes in the Atlantic wintered west of the Mid‐Atlantic Ridge, with only birds from Ireland and western Britain staying mainly on the European side. Main conclusions The high degree of mixing in winter of kittiwakes breeding in various parts of the Atlantic range implies that the overall population could be sensitive to potentially deteriorating environmental conditions in the West Atlantic, e.g. owing to lack of food or pollution. Our approach to estimating the size and composition of wintering populations should contribute to improved management of birds faced with such challenges.     

Seabirds as monitor organisms of contaminants along the German North Sea coast

In the 1980s, significant regional, interspecific and annual differences in contamination with toxic chemicals were found in eggs of breeding birds along the German North Sea coast. In some regions residue levels approached a range endangering breeding success, especially in terns. In 1987, a three-year monitoring-program for common ternSterna hirundo and oystercatcherHaematopus ostralegus was started, in order to recognize such contamination trends. Eggs from 9–14 areas along the coast are now being analysed for chlororganic residues and mercury. First results on geographical and year-to-year variation are presented here, and the advantages of seabird eggs as spatial and temporal monitors of marine pollution are discussed. The suggestion is made to include a shore-breeding bird species in the European monitoring programs. Presented at the VI International Wadden Sea Symposium (Biologische Anstalt Helgoland, Wattenmeerstation Sylt, D-2282 List, FRG, 1–4 November 1988)     

Trophic relationships and seasonal utilization of salt-marsh creeks by zooplanktivorous fishes

Synopsis In a high salinity estuary at North Inlet, South Carolina, co-occurrence and possible competition among adults of four dominant zooplanktivorous fishes were minimized by seasonal adjustments in lateral and vertical distributions as well as in dietary preferences. In winter, Atlantic silversides, Menidia menidia, occupied the entire water column while other planktivores were rare or absent from the estuary, and they consumed large prey such as mysid shrimps and fish larvae. An immigration of bay anchovies, Anchoa mitchilli, in the spring resulted in a redistribution of species with Atlantic silversides shifting to the surface waters and bay anchovies dominating the lower half of the water column. Both fishes consumed mostly copepods in the spring, but each favored a different species. There was little similarity in the large prey items consumed by the two fishes. Striped anchovies, Anchoa hepsetus, arrived in mid-summer and were most abundant at the surface while bay anchovies continued to dominate the bottom waters. Atlantic silversides were rare in all summer collections. The diets of the two anchovies were similar, but vertical separation during the period of maximum zooplankton abundance probably minimized competition. Rough silversides, Membras martinica, which were obligate surface dwellers, shared the upper water column with striped anchovies, but the two species had very different diets during their period of co-occurrence. Although seasonal changes in fish diets reflected shifts in zooplankton composition and all fishes consumed a variety of prey types, preferences for some prey taxa and total avoidance of others were indicated. Electivity indices indicated an especially strong selection for fiddler crab megalopae by all fishes in the summer and fall. All fishes, except rough silversides, which fed almost exclusively on copepods and crab zoeae, consumed large prey items when they were available. Fine scale partitioning of the food resources was apparent in the selection of different copepod and insect species by the fishes. Spatial and temporal separation in the distribution and/or dietary preferences of the zooplanktivores fishes probably reduces the potential for resource competition. Given the high abundances and selectivity of the planktivores, significant impacts on some zooplankton populations probably result.