Description: The Asian shore crab Hemigrapsus sanguineus (De Haan, 1853) has recently established populations in the North Sea and now occurs within the native ranges of the green crab Carcinus maenas (Linnaeus, 1758). To determine potential competitive effects and to assess the progress of the invasion, species-specific population characteristics (numerical abundances, biomasses, and size distributions) of the two species around the island of Helgoland (German Bight, southern North Sea) were compared for surveys conducted in 2009 and 2014. Sampling sites were chosen based on accessibility and differed in their topography and wave exposure, which allowed testing for the influence of these factors on the establishment success of H. sanguineus. The numerical abundance and biomass of H. sanguineus increased markedly and approached those of C. maenas in 2014. At a sheltered site, H. sanguineus even outnumbered C. maenas, whereas the converse was observed at a site exposed to strong winds and waves. Although such contrasting abundance patterns between the native and the introduced shore crab may be the result of direct interference, the dominance of H. sanguineus at the sheltered site may also be explained by enhanced larval settling rates caused by odors of conspecifics. The results suggest that the invasion of H. sanguineus has not yet reached its equilibrium, and population abundances in the North Sea are expected to further increase in the future.

Description: The copepods' impact on vertical carbon flux was assessed for stratified depth layers down to 2000 m at six stations along a transect between 24°N and 21°S in the eastern Atlantic Ocean in October/November 2012. Total copepod community consumption ranged from 202-604 mg C m⁻² day⁻¹, with highest ingestion rates in the tropical North Atlantic. Calanoids consumed 75-90% of the particulate organic carbon (POC) ingested by copepods, although the relative contribution of cyclopoids (mostly Oncaeidae) increased with depth. Net ingestion (=consumption - fecal pellet egestion) of POC varied from 106-379 mg C m⁻² day⁻¹ for calanoids and 37-51 mg C m⁻² day⁻¹ for cyclopoids, corresponding to 16-58% and 5-9%, respectively, of primary production (PP). In total, 9-33% and 2-5% of PP were respired as inorganic carbon by calanoids and cyclopoids, respectively. Copepod ingestion was highly variable between stations and depth layers, especially in the epi- and upper mesopelagic zone. Diel vertical migrants such as Pleuromamma enhanced the vertical flux to deeper layers, particularly in the region influenced by the Benguela Current. The impact of copepod communities on POC flux decreased below 1000 m and POC resources reaching the bathypelagic zone were far from being fully exploited by copepods. As key components, copepods are important mediators of carbon fluxes in the ocean. Their biomass, community composition and interactions strongly affect the magnitude of organic carbon recycled or exported to deeper layers. High variability, even at smaller vertical scales, emphasizes the complex dynamics of the biological carbon pump.

Description: Respiration rates and electron transport system (ETS) activities were measured in dominant copepod species from the northern Benguela upwelling system in January-February 2011 to assess the accuracy of the ETS assay in predicting in vivo respiration rates. Individual respiration rates varied from 0.06 to 1.60 µL O2/h/ind, while ETS activities converted to oxygen consumption ranged from 0.14 to 4.46 µL O2/h/ind. ETS activities were significantly correlated with respiration rates (r**2 = 0.79, p = 0.0001). R:ETS ratios were lowest in slow-moving Eucalanidae (0.11) and highest in diapausing Calanoides carinatus copepodids CV (0.76) while fast-moving copepods showed intermediate R:ETS (0.23-0.37). 82% of the variance of respiration rates could be explained by differences in dry mass, temperature and the activity level of different copepod species. Three regression equations were derived to calculate respiration rates for diapausing, slow- and fast-moving copepods, respectively, based on parameters such as body mass and temperature. Thus, knowledge about the activity level and behavioral characteristics of copepod species can significantly increase the predictive accuracy of metabolic models, which will help to better understand and quantify the impact of copepods on nutrient and carbon fluxes in marine ecosystems.

Description: Decapods were sampled with a 1 m**2 MOCNESS (mainly upper 1000 m) in the northern Benguela Current during three cruises in December 2009, September/October 2010 and February 2011. Although pelagic decapods are abundant members of the micronekton community, information about their ecophysiology is very limited. Species-specific regional distribution limits were detected for various decapod species (e.g. Plesionika carinata, Sergestes arcticus, Pasiphaea semispinosa). Significant diel vertical migration patterns were determined for three caridean and three penaeiodean species. Biomass was variable and ranged from 23 to 2770 mg dry mass m**-2 with highest values for P. semispinosa. Fatty acid and stable isotope analyses revealed that the examined decapod species are omnivorous tocarnivorous except for the herbivorous to omnivorous species P. carinata. Calanid copepods such as Calanoides carinatus were identified as an important prey item especially for caridean species. Community consumption rates of pelagic decapods derived from respiration rates ranged from 7 mg C m**-2 d**-1 (231S) to 420 mg C m**-2 d**-1 (191S, 171S). A potential active respiratory carbon flux was calculated for migrating pelagic decapods with 4.4 mg C m**- d**-1 for the upper 200 m and with 2.6 mg C m**-2 d**-1 from the base of the euphotic zone to a depth of 600 m. Overall, pelagic decapods apparently play a more prominent role in the northern Benguela Current ecosystem than previously assumed and may exert a substantial predation impact on calanid copepods (up to 13% d**-1 of standing stock).

Description: Temora longicornis, a dominant calanoid copepod species in the North Sea, is characterised by low lipid reserves and high biomass turnover rates. To survive and reproduce successfully, this species needs continuous food supply and thus requires a highly flexible digestive system to exploit various food sources. Information on the capacity of digestive enzymes is scarce and therefore the aim of our study was to investigate the enzymatic capability to respond to quickly changing nutritional conditions. We conducted two feeding experiments with female T. longicornis from the southern North Sea off Helgoland. In the first experiment in 2005, we tested how digestive enzyme activities and enzyme patterns as revealed by substrate SDS-PAGE (sodium dodecylsulfate polyacrylamide gel electrophoresis) responded to changes in food composition. Females were incubated for three days fed ad libitum with either the heterotrophic dinoflagellate Oxyrrhis marina or the diatom Thalassiosira weissflogii. At the beginning and at the end of the experiment, copepods were deep-frozen for analyses. The lipolytic enzyme activity did not change over the course of the experiment but the enzyme patterns did, indicating a distinct diet-induced response. In a second experiment in 2008, we therefore focused on the enzyme patterns, testing how fast changes occur and whether feeding on the same algal species leads to similar patterns. In this experiment, we kept the females for 4 days at surplus food while changing the algal food species daily. At day 1, copepods were offered O. marina. On day 2, females received the cryptophycean Rhodomonas baltica followed by T. weissflogii on day 3. On day 4 copepods were again fed with O. marina. Each day, copepods were frozen for analysis by means of substrate SDS-PAGE. This showed that within 24 h new digestive enzymes appeared on the electrophoresis gels while others disappeared with the introduction of a new food species, and that the patterns were similar on day 1 and 4, when females were fed with O. marina. In addition, we monitored the fatty acid compositions of the copepods, and this indicated that specific algal fatty acids were quickly incorporated. With such short time lags between substrate availability and enzyme response, T. longicornis can successfully exploit short-term food sources and is thus well adapted to changes in food availability, as they often occur in its natural environment due seasonal variations in phyto- and microzooplankton distribution.

Description: The majority of global ocean production and total export production is attributed to oligotrophic oceanic regions due to their vast regional expanse. However, energy transfers, food-web structures and trophic relationships in these areas remain largely unknown. Regional and vertical inter- and intra-specific differences in trophic interactions and dietary preferences of calanoid copepods were investigated in four different regions in the open eastern Atlantic Ocean (38°N to 21°S) in October/November 2012 using a combination of fatty acid (FA) and stable isotope (SI) analyses. Mean carnivory indices (CI) based on FA trophic markers generally agreed with trophic positions (TP) derived from d15N analysis. Most copepods were classified as omnivorous (CI ~0.5, TP 1.8 to ~2.5) or carnivorous (CI 〉=0.7, TP 〉=2.9). Herbivorous copepods showed typical CIs of 〈=0.3. Geographical differences in d15N values of epi- (200-0 m) to mesopelagic (1000-200 m) copepods reflected corresponding spatial differences in baseline d15N of particulate organic matter from the upper 100 m. In contrast, species restricted to lower meso- and bathypelagic (2000-1000 m) layers did not show this regional trend. FA compositions were species-specific without distinct intra-specific vertical or spatial variations. Differences were only observed in the southernmost region influenced by the highly productive Benguela Current. Apparently, food availability and dietary composition were widely homogeneous throughout the oligotrophic oceanic regions of the tropical and subtropical Atlantic. Four major species clusters were identified by principal component analysis based on FA compositions. Vertically migrating species clustered with epi- to mesopelagic, non-migrating species, of which only Neocalanus gracilis was moderately enriched in lipids with 16% of dry mass (DM) and stored wax esters (WE) with 37% of total lipid (TL). All other species of this cluster had low lipid contents (〈 10% DM) without WE. Of these, the tropical epipelagic Undinula vulgaris showed highest portions of bacterial markers. Rhincalanus cornutus, R. nasutus and Calanoides carinatus formed three separate clusters with species-specific lipid profiles, high lipid contents (〉=41% DM), mainly accumulated as WE (〉=79% TL). C. carinatus and R. nasutus were primarily herbivorous with almost no bacterial input. Despite deviating feeding strategies, R. nasutus clustered with deep-dwelling, carnivorous species, which had high amounts of lipids (〉=37% DM) and WE (〉=54% TL). Tropical and subtropical calanoid copepods exhibited a wide variety of life strategies, characterized by specialized feeding. This allows them, together with vertical habitat partitioning, to maintain high abundance and diversity in tropical oligotrophic open oceans, where they play an essential role in the energy flux and carbon cycling.

Description: The crabeater seal (Lobodon carcinophaga) is the most abundant Antarctic seal and inhabits the circumpolar pack ice zone of the Southern Ocean. Until now, information on important environmental factors affecting its distribution as well as on foraging behaviour is limited. In austral summer 1998, 12 crabeater seals of both sexes and different age classes were equipped with satellitelinked dive recorders at Drescher Inlet (72.85°S, 19.26°E), eastern Weddell Sea. To identify suitable habitat conditions within the Weddell Sea, a maximum entropy (Maxent) modelling approach was implemented. The model revealed that the eastern and southern Weddell Sea is especially suitable for crabeater seals. Distance to the continental shelf break and sea ice concentration were the two most important parameters in modelling species distribution throughout the study period. Model predictions demonstrated that crabeater seals showed a dynamic response to their seasonally changing environment emphasized by the favoured sea ice conditions. Crabeater seals utilized ice-free waters substantially, which is potentially explained by the comparatively low sea ice cover of the Weddell Sea during summer 1998. Diving behaviour was characterized by short (〉90 % = 0-4 min) and shallow (〉90 % = 0-51 m) dives. This pattern reflects the typical summer and autumn foraging behaviour of crabeater seals. Both the distribution and foraging behaviour corresponded well with the life history of the Antarctic krill (Euphausia superba), the preferred prey of crabeater seals. In general, predicted suitable habitat conditions were congruent with probable habitats of krill, which emphasizes the strong dependence on their primary prey.

Description: Respiration rates of 16 calanoid copepod species from the northern Benguela upwelling system were measured on board RRS Discovery in September/October 2010 to determine their energy requirements and assess their significance in the carbon cycle. Individual respiration rates were standardised to a mean copepod body mass and a temperature regime typical of the northern Benguela Current. These adjusted respiration rates revealed two different activity levels (active and resting) in copepodids C5 of Calanoides carinatus and females of Rhincalanus nasutus, which reduced their metabolism during dormancy by 82% and 62%, respectively. An allometric function (Imax) and an energy budget approach were performed to calculate ingestion rates. Imax generally overestimated the ingestion rates derived from the energy budget approach by 〉75%. We suggest that the energy budget approach is the more reliable approximation with a total calanoid copepod (mainly females) consumption of 78 mg C m-2 d-1 in neritic regions and 21 mg C m-2 d-1 in oceanic regions. The two primarily herbivorous copepods C. carinatus (neritic) and Nannocalanus minor (oceanic) contributed 83% and 5%, respectively, to total consumption by calanoid copepods. Locally, C. carinatus can remove up to 90% of the diatom biomass daily. In contrast, the maximum daily removal of dinoflagellate biomass by N. minor was 9%. These estimates imply that C. carinatus is an important primary consumers in the neritic province of the northern Benguela system, while N. minor has little grazing impact on phytoplankton populations further offshore. Data on energy requirements and total consumption rates of dominant calanoid copepods of this study are essential for the development of realistic carbon budgets and food-web models for the northern Benguela upwelling system.

Description: The region of the Filchner Outflow System (FOS) in the southeastern Weddell Sea is characterized by intensive and complex interactions of different water masses. Dense Ice Shelf Water (ISW) emerging from beneath the ice shelf cavities on the continental shelf, meets Modified Warm Deep Water (MWDW) originating from the Antarctic Circumpolar Current at the sill of the Filchner Trough. These hydrographic features convert the FOS into an oceanographic key region, which may also show enhanced biological productivity and corresponding aggregations of marine top predators. In this context, six adult Weddell seals (Leptonychotes weddellii) were instrumented with CTD-combined satellite relay data loggers in austral summer 2014. By means of these long-term data loggers we aimed at investigating the influence of environmental conditions on the seals' foraging behaviour throughout seasons, focussing on the local oceanographic features. Weddell seals performed pelagic and demersal dives, mainly on the continental shelf, where they presumably exploited the abundant bentho-pelagic fish fauna. Diurnal and seasonal variations in light availability affected foraging activities. MWDW was associated with increased foraging effort. However, we observed differences in movements and habitat use between two different groups of Weddell seals. Seals tagged in the pack ice of the FOS focussed their foraging activities to the western and, partly, eastern flank of the Filchner Trough, which coincides with inflow pathways of MWDW. In contrast, Weddell seals tagged on the coastal fast ice exhibited typical central-place foraging and utilized resources close to their colony. High foraging effort in MWDW and high utilization of areas associated with an inflow of MWDW raise questions on the underlying biological features. This emphasizes the importance of further interdisciplinary ecological investigations in the near future, as the FOS may soon be impacted by predicted climatic changes.