Fluxes of biogenic carbon in the Southern Ocean: roles of large microphagous zooplankton

The Southern Ocean is an extreme environment, where waters are permanently cold, a seasonal ice cover extends over large areas, and the solar energy available for photosynthesis is severely restricted, either by vertical mixing to considerable depths or, especially south of the Antarctic Circle, by prolonged seasonal periods of low or no irradiance. Such conditions would normally lead to low productivity and a water column dominated by recycling processes involving microbial components of pelagic communities but this does not seem to be the case in the Southern Ocean, where there is efficient export to large apex predators and deep waters. This paper investigates the role of large microphagous zooplankton (salps, krill, and some large copepods) in the partitioning of biogenic carbon among the pools of short- and long-lived organic carbon and sequestered biogenic carbon. Large microphagous zooplankton are able to ingest microbial-sized particles and thus repackage small, non-sinking particles into both metazoan biomass and large, rapidly sinking faeces. Given the wide spatio-temporal extent of microbial trophic pathways in the Southern Ocean, large zooplankton that are omnivorous or able to ingest small food particles have a competitive advantage over herbivorous zooplankton. Krill efficiently transfer carbon to a wide array of apex predators and their faecal pellets are exported to depth during occasional brief sedimentation episodes in spring time. Salps may be a significant link towards some fish (directly) and other apex predators (indirectly) and, at some locations (especially in offshore waters) and time, they may account for most of the downward flux of biogenic carbon. Large copepods are a trophic link towards fish and at least one whale species, and their grazing activity generally impedes the export of organic particles to depth. As a result, biogenic carbon is channelled mainly towards apex predators and episodically into the deep ocean. Without these original interactions, Antarctic waters might well be dominated by microbial components and recycling processes instead of active export from the generally small primary producers towards large apex predators.     

Life cycle of Epischura baicalensis Sars (Copepoda, Calanoida) in Lake Baikal

Epischura baicalensis Sars is a dominant pelagic species of Lake Baikal zooplankton. This is endemic to Lake Baikal and inhabits the entire water column. It produces two generations per year: the winter–spring and the summer. These copepods develop under different ecological conditions and vary in the duration of life stages, reproduction time, maturation of sex products and adult males and females lifespan. The total life period of the animals from each generation is one year. One female can produce 10 egg sacks every 10–20 days during its life time. The ratio of males and females is 1:1. One of the most essential features of the ecology of E. baicalensis is the alteration of its mass inhabited areas during a year, as well as in day time. This is due to the need for various conditions for gonad maturation, reproduction, nourishing and protection from being consumed by planktivors.     

Distribution, reproduction and development of Calanus species in the Northeast water in relation to environmental conditions

The vertical and horizontal distribution of mesozooplankton biomass and its composition, together with the reproduction and development of the three dominant herbivorous copepods Calanus finmarchicus, C. glacialis and C. hyperboreus, were studied in the Northeast Water between the end of May and end of July, 1993. Biomass varied between 0.15 and 10.6 g m⁻² and was lower on the shallow banks. Highest biomass was found on the shelf slope and in the entrances to Belgica and Westwind Troughs. There was no seasonal trend during the study period. Among the zooplankton taxa, copepods were dominant, with 84% biomass of all other taxa, followed by chaetognaths with 14%. The large Calanus species made up 91% of copepod biomass. The most abundant species, C. glacialis, inhabited areas of low current speeds on Belgica and Ob Bank, C. hyperboreus dominated shelf slope and trough stations, while C. finmarchicus was most abundant in the Return Atlantic Current along the shelf slope and also eastern Belgica Trough. C. glacialis was the only Calanus species spawning during this period, but young copepodites of the other species were also abundant. Egg production of C. glacialis was at a maximum by our arrival and continued to at least mid August. According to the results from starvation experiments, its egg production was fuelled by food uptake, but was decoupled from phytoplankton chlorophyll until July, indicating ice-algae and microzooplankton as an alternative food source. Only when the polynya approached its maximum extent was a close relationship to phytoplankton established. Due to both spatial and temporal heterogeneity of the occurrence of young copepodite stages it was impossible to follow the growth of cohorts of developmental stages. Herbivorous carbon requirements estimated from egg production rates were mostly less than one third of the phytoplankton stock. From egg production and the distribution of young stages, the outer Westwing Trough seems to be the centre of biological activities. This may be related to the formation of young ice in winter in this area, which was found to carry a large mass of under-ice algae. High biomass but low production in Belgica Trough indicate this as an advective and expatriated community of C. finmarchicus and C. hyperboreus, where grazing is negligible, at least during early summer.     

Copepod diversity in the water masses of the southern East China Sea north of Taiwan

The warm oceanic current Kuroshio and the continental shelf water of the East China Sea meet in the western North Pacific, north of Taiwan and form an upwelling when they converge. The intrusion of the Kuroshio westward over the East China Sea shelf thus results in complicated exchanges of waters between these two water masses. We studied the copepods in the plankton collection taken from an east–west transect crossing these waters in April 1995 when the intrusion of the Kuroshio over the East China Sea shelf was beginning to retreat. The taxonomy of copepods was carefully treated and erroneous species records reported in the literature were guarded against. We evaluated the copepod diversity, the association of copepod species, and the association of stations in these water masses.     

Diel vertical migration and feeding in adult female Calanus pacificus, Metridia lucens and Pseudocalanus newmani during a spring bloom in Dabob Bay, a fjord in Washington USA

Diel vertical migration and feeding on phytoplankton by adult female Calanus pacificus, Metridia lucens and Pseudocalanus newmani were simultaneously measured near the end of a phytoplankton bloom. Almost the entire Calanus population migrated out of the deep layer (108–50 m) at night but only about 30% came to the surface (25–0 m). Feeding occurred only at night and was equally high in the surface and intermediate layers, in spite of much higher food concentrations in the surface. Like Calanus, the entire Metridia population was found in the deep layer during the day but unlike Calanus, 20–50% remained in the deep layer at night and most migratory Metridia were collected from the surface layer. Metridia feeding at night was highest in the surface layer but significant feeding also occurred in both the intermediate and deep layers. Migratory behavior of Pseudocalanus was weak, with the proportion of the population in the surface layer increasing from slightly <10% during the day to approx 30% at night. Feeding occurred in both surface and intermediate layers throughout the 24 h but was greater in both layers at night. The different migratory patterns are discussed in the context of our current understanding of the contributions of predator avoidance and feeding to diel vertical migration.     

Community structure of mesopelagic copepods (>500 μm) in the Ligurian Sea (Western Mediterranean)

The distribution of mesopelagic copepods between 250 and 1000 m depth was studied along a transect off the coast of Villefranche-sur-Mer in June 1991 by means of a BIONESS multiple-net sampler. Among the most abundant species at all stations were Pleuromamma gracilis, Paraeuchaeta acuta and overwintering Calanus helgolandicus CV copepodites, each species inhabiting different depth layers with maximum abundances in the 250–350 m, 450–550 m and the 700–850 m layer, respectively. Cluster analysis of the vertical distribution of all species caught revealed three distinct strata (‘TOP', ‘INTERMEDIATE', ‘BOTTOM') consisting of characteristic species that occurred in the same stratum at all stations. Differences in total abundances of the species assemblage occupying each layer are discussed with regard to predation pressure by mesopelagic macroplanktonic crustaceans and fish. We show that each stratum is characterized by distinct trophic interactions and life strategies, which play an important role in the structuring of mesopelagic zooplankton communities.