Description: We conducted a year-round mesozooplankton study in the Arctic Kongsfjord from August 1998 until July 1999 to investigate seasonal abundance and vertical as well as stage distributions of the prevalent taxa. It is the first investigation in Kongsfjord that covers the Arctic winter season and provides reasonable estimates also of small-sized copepod species. Abundant smaller copepods comprised Oithona similis, Pseudocalanus minutus, Microcalanus spp., Triconia borealis and Acartia longiremis. Among the larger copepods, Calanus finmarchicus, C. glacialis, C. hyperboreus and Metridia longa dominated. The thecosome pteropod Limacina helicina was also an important component. Abundance maxima occurred in November (988,669 ind. m−2) with one to two orders of magnitude higher numbers as compared to all other months (39,832–200,067 ind. m−2). The summers of 1998 and 1999 were characterized by intrusions of Atlantic water, but the community was not entirely dominated by advected boreal species. During winter, the majority of the mesozooplankton occurred below 100 m. Advection is the most likely reason for the accumulation of zooplankton at depth in winter, but local production may also contribute to high overwintering numbers. Much lower abundances of most species in spring suggest high winter mortality and emphasize the importance of sufficient reproductive success during the previous summer to ensure enough winter survivors as seed stock for the coming reproductive season. This study was conducted prior to the recent warming trend in the Arctic. Therefore, it provides valuable baseline data and allows comparing present and future states of the zooplankton community in Kongsfjord.

Description: Copepods comprise the majority of mesozooplankton communities in all marine regions. Their diversity is likely regulated by environmental parameters and species-specific ecophysiological aspects making the distributional pattern of certain species an environmental indicator to detect changes in the marine habitat. We studied copepod distribution and diversity at six stations along a meridional transect in the eastern Atlantic Ocean (25°N to 21°S) from surface to 2000 m depths. Community and trophic structures with special regard to calanoid copepods were analyzed. Below the euphotic zone, representatives of the family Spinocalanidae were particularly abundant and diverse but morphological species identification both of adults and juveniles was very challenging. To elucidate both diversity and species- and stage-specific vertical distribution of Spinocalanidae we therefore applied an integrated taxonomic approach combining morphology, DNA sequence analyses and proteomics using the matrix-assisted laser desorption/ionization time-of-flight mass spectrometry (MALDI-TOF MS). Each specimen was analysed by the three methodologies by portioning the whole organism. Species clusters were similar for DNA sequence analyses and MALDI-TOF MS and resulted in a higher amount of species clusters compared to the morphological identification. Since MALDI-TOF MS is a rather time- and cost-efficient technology, we were able to identify high individual numbers of the Spinocalanidae and received a quantitative, high species-resolution picture of adult and juvenile Spinocalanidae and their distributional patterns.

Description: Hyperiid amphipods are predatory pelagic crustaceans that are particularly prevalent in high-latitude oceans. Many species are likely to have co-evolved with soft-bodied zooplankton groups such as salps and medusae, using them as substrate, for food, shelter or reproduction. Compared to other pelagic groups, such as fish, euphausiids and soft-bodied zooplankton, hyperiid amphipods are poorly studied especially in terms of their distribution and ecology. Hyperiids of the genus Themisto, comprising seven distinct species, are key players in temperate and cold-water pelagic ecosystems where they reach enormous levels of biomass. In these areas, they are important components of marine food webs, and they are major prey for many commercially important fish and squid stocks. In northern parts of the Southern Ocean, Themisto are so prevalent that they are considered to take on the role that Antarctic krill play further south. Nevertheless, although they are around the same size as krill, and may also occur in swarms, their feeding behaviour and mode of reproduction are completely different, hence their respective impacts on ecosystem structure differ. Themisto are major predators of meso- and macrozooplankton in several major oceanic regions covering shelves to open ocean from the polar regions to the subtropics. Based on a combination of published and unpublished occurrence data, we plot out the distributions of the seven species of Themisto. Further, we consider the different predators that rely on Themisto for a large fraction of their diet, demonstrating their major importance for higher trophic levels such as fish, seabirds and mammals. For instance, T. gaudichaudii in the Southern Ocean comprises a major part of the diets of around 80 different species of squid, fish, seabirds and marine mammals, while T. libellula in the Bering Sea and Greenland waters is a main prey item for commercially exploited fish species. We also consider the ongoing and predicted range expansions of Themisto species in light of environmental changes. In northern high latitudes, sub-Arctic Themisto species are replacing truly Arctic, ice-bound, species. In the Southern Ocean, a range expansion of T. gaudichaudii is expected as water masses warm, impacting higher trophic levels and biogeochemical cycles. We identify the many knowlegde gaps that must be filled in order to evaluate, monitor and predict the ecological shifts that will result from the changing patterns of distribution and abundance of this important pelagic group.

Description: Environmental fluctuations can impose energetic constraints on organisms in terms of food shortage or compensation for metabolic stress. To better understand the biochemical strategies that support adaptive physiological processes in variable environments, we studied the lipid dynamics of the brown shrimp Crangon crangon and the pink shrimp Pandalus montagui by analysing their midgut glands during an annual cycle. Both species have an overlapping distribu- tion range in the southern North Sea, but differ in their habitat preferences, reproductive strate- gies, and life-history traits. C. crangon showed minor total lipid accumulation in their midgut glands, ranging between 14 and 17% of dry mass (DM), dominated by phospholipids. In contrast, P. montagui stored significantly larger amounts of total lipid (47−70% DM, mainly triacylglycer- ols) and showed a distinct seasonal cycle in lipid accumulation with a maximum in summer. Fatty acid trophic markers indicated a wide food spectrum for both species, with higher preferences of P. montagui for microalgae. In C. crangon, feeding preferences were less distinct due the low total lipid levels in the midgut gland. PCA based on fatty acid compositions of both species suggested that C. crangon has a broader dietary spectrum than P. montagui. C. crangon seems to have the capacity to use sufficient energy directly from ingested food to fuel all metabolic requirements, including multiple spawnings, without building up large lipid reserves in the midgut gland. P. montagui, in contrast, relies more on the energy storage function of the midgut gland to over- come food scarcity and to allocate lipids for reproduction.

Description: The Asian shore crab H. sanguineus first appeared at the French coast in the late 1990’s. It rapidly extended its range further north to the German Wadden Sea and recently to western Sweden. In the intertidal area, it co-occurs with the European green crab C. maenas. As the ecophysiology of H. sanguineus is virtually unknown, the physiological capacities of both species and their potential for intra-guild competition were investigated. The aim of this study was to specifically compare the energy deposition and dietary preferences of ovigerous females of both species. Females of H. sanguineus and C. maenas carrying immature or mature eggs were collected in April, June, August and October 2015 in an intertidal area of the Island of Helgoland, North Sea. Total lipid levels and fatty acid compositions were determined of both midgut glands and eggs. In H. sanguineus, total lipid levels of the midgut glands were clearly higher than those of C. maenas (40% vs. 10% dry mass, DM). Immature eggs were quite lipid-rich in both species with 30% and 25%DM, respectively, whereas in mature eggs, lipid levels decreased to ~15%DM each. A Principal Component Analysis of the fatty acid compositions of midgut glands and eggs revealed separate clusters for both species with C. maenas lipids more characterized by membrane fatty acids. In C. maenas fatty acids of midgut glands and eggs clustered together largely dominated by carnivory biomarkers. Fatty acids of midgut glands and all eggs of H. sanguineus formed separate clusters and trophic markers indicate a more herbivorous diet. Higher lipid levels and thus more pronounced energy deposition in H. sanguineus midgut glands indicate higher starvation tolerance for females, a potential competitive advantage over C. maenas. Direct food competition, however, seems negligible, as H. sanguineus prefers a more herbivorous diet than C. maenas. Deviating fatty acid compositions in H. sanguineus midgut glands and eggs suggest that this species may represent an income breeder, utilizing energy from both the midgut gland but also from dietary input. Most brachyuran crabs are capital breeders, which rely exclusively on internal reserves.

Description: Biological invasions can entail major threats to marine biodiversity. Non-indigenous species may induce changes in community structure and ecosystem functioning, thereby affecting ecosystem services and human economic interest and health. Decapod brachyuran crabs are among the most invasive marine animal taxa. The originally European species Carcinus maenas and the Asian shore crab Hemigrapsus sanguineus invaded different coastal areas around the world. While their invasion history and ecology has been thoroughly investigated, physiological properties of H. sanguineus are virtually unknown. The aim of this study was to compare the metabolic energy demand of both crab species and - based on these data - to assess their potential dietary impact on the ecosystem. Respiration measurements were conducted with a flow-through system covering a temperature range naturally experienced by these crabs (5, 10, 15 and 20 °C). Both species were analyzed on the island of Helgoland in April, June and August 2015. A general linear mixed-effects model (LMM) was applied to test for the effects of species, temperature, biomass and sex on respiration rates. Overall rates increased with temperature but decreased with the mass of the crabs. Respiration rates did not differ significantly between sexes in both species. From the full model, two separated LMMs were created for either species. They allowed establishing species-specific equations for the prediction of respiration rates y (nmol d-1 g-1) for a crab of any given mass xMass (g) at any given ambient temperature xTemp (°C): ln y = 10.39 + (-0.34 * ln xMass) + (0.06 * xTemp) for C. maenas and ln y = 10.42 + (-0.39 * ln xMass) + (0.08 * xTemp) for H. sanguineus. The mass-specific respiration rates of C. maenas and H. sanguineus were quite similar. By applying the diet-dependent respiratory quotient, oxygen uptake may be used to calculate carbon uptake and metabolic energy demand either for single crabs or for entire populations of a given area. On the population level, the metabolic energy demand and thus ecosystem impact of both species depend primarily on their abundance in the field and, less so on their dietary preferences.

Description: This study aimed at understanding how life-cycle strategies of the primarily herbivorous Pseudocalanus minutus and the omnivorous Oithona similis are reflected by their lipid carbon turnover capacities. The copepods were collected in Billefjorden, Svalbard, and fed with 13C labeled flagellates and diatoms during 3 weeks. Fatty acid (FA) and fatty alcohol compositions were determined by gas chromatography, 13C incorporation was monitored using isotope ratio mass spectrometry. Maximum lipid turnover occurred in P. minutus, which exchanged 54.4% of total lipid, whereas 9.4% were exchanged in O. similis. In P. minutus, the diatom markers 16:1(n-7), 16:2(n-4) and 16:3(n-4) were almost completely renewed from the diet within 21 days, while 15% of the flagellate markers 18:2(n-6), 18:3(n-3) and 18:4(n-3) were exchanged. In O. similis, 15% of both flagellate and diatom markers were renewed. P. minutus exhibited typical physiological adaptations of herbivorous copepod species, with a very high lipid turnover rate and the ability to integrate FAs more rapidly from diatoms than from flagellates. O. similis depended much less on lipid reserves and had a lower lipid turnover rate, but was able to ingest and/or assimilate lipids with the same intensity from various food sources, to sustain shorter periods of food shortage.

Description: The intertidal Asian shore crab H. sanguineus was initially found on the French coast in the late 1990’s. It rapidly extended its range further north and is now well established in the German Wadden Sea. Recently, it was also found in western Sweden. In its intertidal habitat, it co-occurs with the European green crab C. maenas. The ecophysiology of H. sanguineus is virtually unknown. In this study, the physiological capacities of both species and their potential for intra-guild competition were investigated. We specifically compared the energy deposition and dietary preferences of ovigerous females of both species. Females of both species carrying immature or mature eggs were collected in April, June, August and October 2015 in an intertidal site on the Island of Helgoland, North Sea, Germany. Total lipid levels of midgut glands and eggs were acquired via extraction. Subsequently, fatty acid compositions were determined through gas chromatography. Total lipid levels of H. sanguineus midgut glands were clearly higher than those of C. maenas (40% vs. 10% dry mass, DM). Immature eggs of both species were quite lipid-rich with 30% and 25%DM, respectively. In mature eggs, lipid levels decreased to ~15%DM each. A Principal Component Analysis of the fatty acid compositions of midgut glands and eggs revealed separate clusters for both species. Lipids of C. maenas were characterized more by membrane fatty acids. Fatty acids of C. maenas midgut glands and eggs clustered together. They were largely dominated by carnivory biomarkers. Contrastingly, fatty acids of midgut glands and all eggs of H. sanguineus formed separate clusters and trophic markers indicated a more herbivorous diet. Higher lipid levels and thus more pronounced energy deposition in H. sanguineus midgut glands indicate higher starvation tolerance for females, a potential competitive advantage over C. maenas. Direct food competition, however, seems negligible, as H. sanguineus prefers a more herbivorous diet than C. maenas. Deviating fatty acid compositions among H. sanguineus midgut glands and eggs suggests that this species may represent an income breeder, utilizing energy from both the midgut gland and dietary input. In contrast, most brachyuran crabs are capital breeders, relying exclusively on internal reserves.