Spatio‐temporal variability of zooplankton biomass and environmental control in the Northern Benguela Upwelling System: field investigations and model simulation

Spatial and temporal distribution patterns of zooplankton are highly variable in the Northern Benguela Upwelling System. We studied the distribution of zooplankton (size class ≥ 0.33 mm) and used field data from four cruises that took place between March 2008 and February 2011, as well as simulation results of a regional ecosystem model. Remotely sensed sea surface temperatures (SST) and surface chlorophyll concentrations were analysed to investigate environmental influences on zooplankton biomass. The Intense Benguela Upwelling Index showed a distinct seasonal signal throughout the years and the highest upwelling peaks in August/September. Even though surface chlorophyll concentrations were very variable throughout the year, the highest concentrations were always detected in September, following the upwelling of nutrient‐rich water. In field catches, zooplankton biomass concentration in the upper 200 m was highest above the outer shelf and shelf‐break in December 2010 and February 2011, i.e. 6 months after the upwelling peaks. In contrast, zooplankton biomass simulated by the model in the surface water was highest in September. In March/April, biomass maxima were typically measured in the field at intermediate water depths, but the vertical distribution was also affected by extensive oxygen minimum zones. The ecosystem model reproduced this vertical pattern. Although general trends were similar, simulation data of zooplankton standing stocks overestimated the field data by a factor of 3. In upwelling systems, food webs are generally considered to be short and dominated by large cells. However, our field data indicate more small‐sized zooplankton organisms above the shelf than offshore.     

An overview of Calanus helgolandicus ecology in European waters

We review current knowledge and understanding of the biology and ecology of the calanoid copepod Calanus helgolandicus in European waters, as well as provide a collaborative synthesis of data from 18 laboratories and 26 sampling stations in areas distributed from the northern North Sea to the Aegean and Levantine Seas. This network of zooplankton time-series stations has enabled us to collect and synthesise seasonal and multi-annual data on abundance, body size, fecundity, hatching success and vertical distribution of C. helgolandicus. An aim was to enable comparison with its congener Calanus finmarchicus, which has been studied intensively as a key component of European and north east Atlantic marine ecosystems. C. finmarchicus is known to over-winter at depth, whereas the life-cycle of C. helgolandicus is less well understood. Overwintering populations of C. helgolandicus have been observed off the Atlantic coast between 400 and 800 m, while in the Mediterranean there is evidence of significant deep-water populations at depths as great as 4200 m. The biogeographical distribution of C. helgolandicus in European coastal waters covers a wide range of habitats, from open ocean to coastal environments, and its contribution to mesozooplankton biomass ranges from 6% to 93%. Highest abundances were recorded in the Adriatic and off the west coast of Spain. C. helgolandicus is generally found in 9-20 °C water, with maximum abundances from 13-17 °C. In contrast, C. finmarchicus is found in cooler water between 0 and 15 °C, with peak abundances from 0 to 9 °C. As water has warmed in the North Atlantic over recent decades, the range of C. helgolandicus and its abundance on the fringes of its expanding range have increased. This review will facilitate development of population models of C. helgolandicus. This will not only help answer remaining questions but will improve our ability to forecast future changes, in response to a warming climate, in the abundance and distribution of this important species.