Pronounced interannual variability in the abundance of medusae of the jellyfish species Aurelia aurita, Cyanea lamarckii, and Cyanea capillata (Phylum Cnidaria, Class Scyphozoa) in the North Sea was evident in data arising from the International Council for the Exploration of the Seas International 0-group Gadoid Surveys between 1971 and 1986. Possible climatic forcing of jellyfish abundance, via the North Atlantic Oscillation (NAO), was investigated with data on medusae from four areas of the North Sea (east of Scotland, north of Scotland, east of Shetland, and west of northern Denmark). There were significant inverse relationships between medusa abundance and the NAO Index (December-March) in two regions: west of northern Denmark (A. aurita r 2 ϭ 0.70, P ϭ 0.003, n ϭ 10; C. lamarckii r 2 ϭ 0.74, P ϭ 0.002, n ϭ 10) and east of Scotland (A. aurita r 2 ϭ 0.53, P ϭ 0.008, n ϭ 12). Fluctuations in the abundance of A. aurita and C. lamarckii medusae might be linked to hydroclimatic changes induced through atmospheric effects (as encapsulated in the NAO Index) on wind stress, temperature, and currents. These fundamental hydroclimatic changes alter the timing of spring phytoplankton blooms and zooplankton community composition. Predation by an abundance of medusae on zooplankton and ichthyoplankton could affect the North Sea ecosystem through top-down and bottom-up mechanisms. Because the NAO is presently in a high phase, climatic conditions could be serving to depress the abundance of medusae: a future reversal of the NAO might favor jellyfish and weaken the persistence or recovery of fisheries.

Jellyfish medusae prey on zooplankton and may impact fish recruitment both directly (top-down control) and indirectly (through competition). Abundances of Aurelia aurita, Cyanea lamarckii and Cyanea capillata medusae (Scyphozoa) in the North Sea appear to be linked to large-scale inter-annual climatic change, as quantified by the North Atlantic Oscillation Index (NAOI), the Barents Sea-Ice Index (BSII) and changes in the latitude of the Gulf Stream North Wall (GSNW). Hydroclimatic forcing may thus be an important factor influencing the abundance of gelatinous zooplankton and may modulate the scale of any ecosystem impact of jellyfish. The population responses are probably also affected by local variability in the environment manifested in intra-annual changes in temperature, salinity, current strength/direction and prey abundance. Aurelia aurita and C. lamarckii in the north-west and south-east North Sea exhibited contrasting relationships to change in the NAOI and BSII: north of Scotland, where the North Sea borders the Atlantic, positive relationships were evident between the abundance of scyphomedusae (data from 1974 to 1986, except 1975) and the indices; whereas west of northern Denmark, a region much less affected by Atlantic inflow, negative relationships were found (data from 1973 to 1983, except 1974). Weaker negative relationships with the NAOI were also found in an intermediate region, east of Scotland, for the abundance of A. aurita and C. capillata medusae (1971 to 1982). East of Shetland, the abundance of jellyfish was not correlated directly with the NAOI but, in contrast to all other regions, the abundances of A. aurita and C. lamarckii (1971 to 1986, not 1984) were found to correlate negatively with changes in the GSNW, which itself was significantly positively correlated to the NAOI with a two year lag. On this evidence, we suggest that, for jellyfish, there exist three regions of the North Sea with distinct environmental processes governing species abundance: one north of Scotland, another east of Shetland, and a more southerly group (i.e. east of Scotland and west of northern Denmark). Impacts by jellyfish are likely to vary regionally, and ecosystem management may benefit from considering this spatial variability.

This version is available at https://strathprints.strath.ac.uk/18569/ Strathprints is designed to allow users to access the research output of the University of Strathclyde. Unless otherwise explicitly stated on the manuscript, Copyright © and Moral Rights for the papers on this site are retained by the individual authors and/or other copyright owners. Please check the manuscript for details of any other licences that may have been applied. You may not engage in further distribution of the material for any profitmaking activities or any commercial gain. You may freely distribute both the url (https://strathprints.strath.ac.uk/) and the content of this paper for research or private study, educational, or not-for-profit purposes without prior permission or charge.Any correspondence concerning this service should be sent to the ABSTRACTThe population of Calanus finmarchicus in the North Sea is replenished each spring by invasion from an overwintering stock located beyond the shelf edge. A combination of field observations, statistical analysis of Continuous Plankton Recorder (CPR) data, and particle tracking model simulations, was used to investigate the processes involved in the cross-shelf invasion. The results showed that the main source of overwintering animals entering the North Sea in the spring is at depths of greater than 600 m in the Faroe Shetland Channel, where concentrations of up to 620 m 73 are found in association with the overflow of Norwegian Sea Deep Water (NSDW) across the Iceland Scotland Ridge. The input of this water mass to the Faroe Shetland Channel, and hence the supply of overwintering C. finmarchicus, has declined since the late 1960s due to changes in convective processes in the Greenland Sea. Beginning in February, animals start to emerge from the overwintering state and migrate to the surface waters, where their transport into the North Sea is mainly determined by the incidence of north-westerly winds that have declined since the 1960s. Together, these two factors explain a high proportion of the 30-year trends in spring abundance in the North Sea as measured by the CPR survey. Both the regional winds and the NSDW overflow are connected to the North Atlantic Oscillation Index (NAO), which is an atmospheric climate index, but with different time scales of response. Thus, interannual fluctuations in the NAO can cause immediate changes in the incidence of northwesterly winds without leading to corresponding changes in C. finmarchicus abundance in the North Sea, because the NSDW overflow responds over longer (decadal) time scales.

Copepodite stage V Calanus finmarchicus were collected at locations on the continental shelf north of Scotland, in the Faroe–Shetland Channel and west of Ireland on six occasions covering winter, spring and summer from October 1993 to June 1995. Oxygen consumption by the overwintering and active spring/summer population of animals was determined at temperatures close to in situ temperatures. Laboratory measurements of oxygen consumption were also made at standardized temperatures (0°C, 5°C, 7°C and 12.5°C) to determine the sensitivity of animals to temperature change in the different seasons. Rates of oxygen consumption were very low (7–30 μmol O2 gC−1 h−1) at in situ temperatures during the winter and early spring and significantly higher (105–219 μmol O2 gC−1 h−1) for the active surface population in May and June. Animals collected from the overwintering population showed no significant response to changes in temperature. Due to the low respiration rates, the calculated rate of decrease in carbon content in diapausing copepodite stage CV was very low (approximately 0.250 μgC day−1). The respiration rates were used to construct a model to estimate survival of the animals with an initial carbon content equivalent to that expected of animals in October. The results showed that in order to survive during winter and have enough energy for moulting and migration to the surface in the spring, these animals have to live at temperatures close to 0°C and be in a diapause state.

Data from plankton net and Optical Plankton Counter sampling during 12 winter cruises between 1994 and 2002 have been used to derive a multi-annual composite 3-D distribution of the abundance of over-wintering Calanus finmarchicus in a swath across the North Atlantic from Labrador to Norway. Dense concentrations occurred in the Labrador Sea, northern Irminger Basin, northern Iceland Basin, eastern Norwegian Sea, Faroe-Shetland Channel, and in the Norwegian Trench of the North Sea. A model of buoyancy regulation in C. finmarchicus was used to derive the lipid content implied by the in situ temperature and salinity at over-wintering depths, assuming neutral buoyancy. The Faroe-Shetland Channel and eastern Norwegian Sea emerged as having the highest water column-integrated abundances of copepodites, the lowest over-wintering temperature, and the highest implied lipid content. The results are discussed in the context of spatial persistence of populations, seasonal patterns of abundance, and relationships between over-wintering and lipid accumulation in the surface waters