Description: Hyperiidean amphipods are a major prey for fish and seabirds. In the Southern Ocean, they are particularly abundant, with distributions ranging from the Polar Frontal Zone to Antarctic shelf waters. The species Hyperiella dilatata has previously been reported to show a peculiar anti-predatory behaviour: It captures chemically protected, gymnosome pteropods in the water column and carries them on its dorsum, like a backpack. We report this association at four oceanic sampling sites between latitudes 45° and 71° S. Molecular barcodes of both hosts and pteropods are provided and compared with those of other hyperiidean and pteropod specimens. Morphological identifications as well as molecular analyses show a so far undocumented association of Hyperiella antarctica with the pteropod Spongiobranchaea australis in the Polar Frontal Zone (Lazarev Sea). H. dilatata carried Clione limacina antarctica specimens in the Weddell Sea, as recorded previously for the Ross Sea. Lengths of the abducted pteropods varied between 1 and 5 mm, with the biggest pteropod measuring more than half the host’s size. One of the abducting amphipods was a female carrying eggs. The formation of such tandem is known to be very efficient as protection from visually hunting icefish in the crystal-clear coastal waters around the Antarctic continent; however, in the open ocean, this behaviour was so far undocumented. Here, we develop hypotheses on its origin and function.

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: Climate change proceeding at unprecedented pace is currently redistributing life on Earth. Warming of the upper ocean and the atmosphere have altered sea ice extent and seasonal dynamics in the Arctic, and similar changes are observed in the Atlantic sector of the Southern Ocean and West Antarctic. This affects stocks of major pelagic species such as krill. A so far neglected key player, the predatory amphipod genus Themisto, covers a distribution from temperate to polar regions where it regionally represents the dominant food source for higher trophic levels. Poleward range expansions of several Themisto species have been already been documented or predicted as warming continues. In the Atlantic sector of the Southern Ocean, changes in seasonal sea ice extension are causing strong annual variations of krill stocks, whilst salps are on the rise and shifting their distribution poleward. A third key player, Themisto gaudichaudii, is now also anticipated to extend its range southward. In the Arctic, the ice-dependent Arctic T. libellula and the sub-Arctic boreal T. abyssorum co-exist. Considering the ongoing Atlantification of the Arctic, a range expansion of T. abyssorum concomitant with a retraction of T. libellula’s range are very likely to occur. Due to major knowledge gaps in the ecology, biology and genetic connectivity of Themisto species, the likelihood of range shifts and their consequences for food web structures and biogeochemical cycles remain largely unexplored. We conducted a comparative study of Themisto populations to investigate their distribution, connectivity and trophic ecology. Using population genetic methods, the species status and phylogeography of T. gaudichaudii, were characterized. A genetic homogeneity and high degree of phenotypic plasticity related to feeding morphology were revealed, indicating the potential of T. gaudichaudii lineages to thrive in regions further south. In order to predict the consequences of an increased predation pressure of T. gaudichaudii which may influence the standing stock of Antarctic krill and salps, feeding experiments and molecular diet analyses were carried out. Similar analyses were applied on Themisto species from the northern hemisphere, in order to characterize their genetic connectivity within and across different water masses (Atlantic vs. polar) in Fram Strait and East Greenland and compare with the Antarctic populations.

Description: The brown shrimp Crangon crangon is a key component of the North Atlantic coastal food web and an important target species for the fishery economy. As the brown shrimp contains large amounts of protein and essential fatty acids, its consumption makes it a beneficial choice for humans. Commercially harvested crustaceans like C. crangon are frequently affected by bacterial shell disease, with necrotizing erosions and ulcerations of the cuticle. To determine whether shell disease influences the nutritional value of C. crangon, total protein and lipid contents, as well as fatty acid compositions of muscle tissue and hepatopancreas, together with the hepatosomatic index, were examined in healthy and affected individuals. The biochemical composition of the tissues did not differ significantly between the 2 groups. Also, the hepatosomatic index, as an indicator of energy reserves in shrimps, was similar between healthy and affected animals. Our results indicate that the nutritional value of C. crangon is not affected by shell disease, as long as it remains superficial as in the present study.

Description: The brown shrimp, Crangon crangon, is well adapted to the variable environmental conditions in the southern North Sea. It is very abundant, has high reproduction rates, and holds a key position in coastal ecosystems. This species has very low lipid deposits in the midgut gland, suggesting that the main function of the midgut gland is metabolic turnover rather than energy storage. Based on seasonal gene expression studies and established transcriptome data, we investigated key components of lipid metabolic pathways. Gene expression of triacylglycerol lipase, phospholipase, and fatty acid desaturase were analyzed and compared with that of other digestive enzymes involved in lipid, carbohydrate, and protein catabolism. Our results suggest that gene expression of digestive enzymes involved in lipid metabolism is modulated by the lipid content in the midgut gland and is related to food availability. Brown shrimp seem to be capable of using cellular phospholipids during periods of food paucity but high energetic (lipid) requirements. Two of three isoforms of fatty acid binding proteins (FABPs) from the midgut gland involved in fatty acid transport showed specific mutations of the binding site. We hypothesize that the mutations in FABPs and deficiencies in anabolic pathways limit lipid storage capacities in the midgut gland of C. crangon. In turn, food utilization, including lipid catabolism, has to be efficient to fulfill the energetic requirements of brown shrimp.

Description: Background: Trophic interactions are key processes, which determine the ecological function and performance of organisms. Many decapod crustaceans feed on plant material as a source for essential nutrients, e.g. polyunsaturated fatty acids. Strictly herbivorous feeding appears only occasionally in marine decapods but is common in land crabs. To verify food preferences and to establish trophic markers, we studied the lipid and fatty acid composition of the midgut glands of two marine crab species (Grapsus albolineatus and Percnon affine), one semi-terrestrial species (Orisarma intermedium, formerly Sesarmops intermedius), and one terrestrial species (Geothelphusa albogilva) from Taiwan. Results: All species showed a wide span of total lipid levels ranging from 4 to 42% of the dry mass (%DM) in the marine P. affine and from 3 to 25%DM in the terrestrial G. albogilva. Triacylglycerols (TAG) were the major storage lipid compound. The fatty acids 16:0, 18:1(n-9), and 20:4(n-6) prevailed in all species. Essential fatty acids such as 20:4(n-6) originated from the diet. Terrestrial species also showed relatively high amounts of 18:2(n-6), which is a trophic marker for vascular plants. The fatty acid compositions of the four species allow to clearly distinguish between marine and terrestrial herbivorous feeding due to significantly different amounts of 16:0, 18:1(n-9), and 18:2(n-6). Conclusions: Based on the fatty acid composition, marine/terrestrial herbivory indices were defined and compared with regard to their resolution and differentiating capacity. These indices can help to reveal trophic preferences of unexplored species, particularly in habitats of border regions like mangrove intertidal flats and estuaries.

Description: A species history is characterized by fluctuations of its geographic limits, however, climate change is currently redistributing life on Earth. Particularly in the Arctic, waters are warming at faster rates and sea ice is thinning and contracting. Amphipod crustaceans of the genus Themisto are highly abundant in Arctic waters where they play a major role as food for higher trophic levels such as seabirds, fish and seals. Throughout the Arctic region, two congeneric species co-exist: T. libellula, a genuine Arctic species and T. abyssorum, considered sub-Arctic boreal. Despite their overlapping distributions, the two species seem to occupy distinct ecological niches and are preyed upon by different predators. T. libellula is bigger in size and feeds on copepods with a high lipid content. It is the main prey of some specialist feeders such as little auks that only feed on the largest size class, in which case T. abyssorum would not be a proper substitute as it is much smaller. Range expansion of T. abyssorum and retraction of T. libellula’s range is very likely to occur considering the ongoing Atlantification of the Arctic. However, many aspects of the biology, ecology and genetic connectivity of Arctic Themisto populations are still unstudied, despite their importance for a better understanding of the consequences of their potential distributional changes on the food web and biogeochemical cycles. In this context, we investigated the abundance, geographic and bathymetric distribution as well as the genetic connectivity of the two species T. abyssorum and T. libellula during two cruises with R/V Polarstern to Fram Strait and East Greenland.

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.