Predatory zooplankton on the move: Themisto amphipods in high-latitude marine pelagic food webs.

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 Based on a combination of published and unpublished occurrence data, we plot out the distributions of the seven species of Themisto.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.