Modelling the basin-scale demography of Calanus finmarchicus in the north-east Atlantic

In this paper, we report on a coupled physical–biological model describing the spatio-temporal distribution of Calanus finmarchicus over an area of the North Atlantic and Norwegian Sea from 56°N, 30°W to 72°N, 20°E. The model, which explicitly represents all the life-history stages, is implemented in a highly efficient discrete space–time format which permits wide-ranging dynamic exploration and parameter optimization. The underlying hydrodynamic driving functions come from the Hamburg Shelf-Ocean Model (HAMSOM). The spatio-temporal distribution of resources powering development and reproduction is inferred from SeaWiFS sea-surface colour observations. We confront the model with distributional data inferred from continuous plankton recorder observations, overwintering distribution data from a variety of EU, UK national and Canadian programmes which were collated as part of the Trans-Atlantic Study of Calanus (TASC) programme, and high-frequency stage-resolved point time-series obtained as part of the TASC programme. We test two competing hypotheses concerning the control of awakening from diapause and conclude that only a mechanism with characteristics similar to photoperiodic control can explain the test data.     

Annual weather variability and its influence on the population dynamics of Calanus finmarchicus

The reaction of the population of Calanus finmarchicus to relatively extreme annual cycles of weather in the North Sea was investigated by one-dimensional model simulations. A population dynamics model for C. finmarchicus was coupled with a physical and a biological upper-layer model for phosphate, phytoplankton and detritus to simulate the development of the successive stages of C. finmarchicus. Observed annual weather cycles were used to drive the physical water column model, the results of which were then input to the plankton model. The simulations yielded the temporal development of all stages of C. finmarchicus over an annual cycle in terms of numbers and weights. Compared with the results reported for 1984 by Carlotti and Radach (1996; Limnol . Oceanogr . 41: 522–539), it appears that the temporal range for the occurrence of the peak concentration of C. finmarchicus may be 2 weeks in the northern North Sea. The simulated variability is in accordance with observations. Years with more continuous primary production such as the year 1984, which was cloudy and windy during the summer, may then provide the optimum conditions with respect to producing an abundance of zooplankton during the following year.     

Modeling larval Calanus finmarchicus on Georges Bank: time-varying mortality rates and a cannibalism hypothesis

Throughout the North Atlantic, the copepod Calanus finmarchicus dominates the zooplankton biomass, linking primary production and higher trophic levels. On Georges Bank, the peak abundance of larval (naupliar) stages occurs in March–April and represents a potential source of prey for cod and haddock larvae. Following this maximum, naupliar abundance declines dramatically, reaching a minimum in May and increasing again in June. Explaining the naupliar seasonal cycle is critical for predicting climate effects on C. finmarchicus dynamics, including whether environmental variability may lead to a mismatch with larval fish. Here, an age-within-stage population dynamics model is used to investigate the factors controlling the temporal variation of C. finmarchicus nauplii in three Georges Bank sub-regions. The model incorporates temperature- and food-dependent development and egg production, as well as female abundance derived from the US Global Ocean Ecosystem Dynamics (GLOBEC) program. Use of field-estimated constant mortality rates overestimates May abundances by as much as an order of magnitude. These data/model discrepancies can not be explained by temperature or food-limitation effects on physiological rates. Instead, accurate simulation requires use of time-varying early stage mortalities, which differ from published estimates in both magnitude and trend. These mortality rates are correlated with C. finmarchicus female abundance, implying cannibalism as a possible regulatory factor. Thus, the biological control of predation (including cannibalism) must be considered to predict the effects of climate on C. finmarchicus and associated larval fish populations.     

Coupling of an individual-based population dynamic model of Calanus finmarchicus to a circulation model for the Georges Bank region

An individual-based life history and population dynamic model for the winter–spring dominant copepod of the subarctic North Atlantic, Calanus finmarchicus , is coupled with a regional model of advection for the Gulf of Maine and Georges Bank. Large numbers of vectors, each representing individual copepods with elements for age, stage, ovarian status and other population dynamic variables, are carried in a computation through hourly time steps. Each vector is updated at each time step according to development rate and reproductive functions derived from experimental data. Newly spawned eggs are each assigned new vectors as needed. All vectors are subject to random mortality. Thus, both life history progression and population dynamics of C. finmarchicus are represented for the temperatures in the Gulf of Maine–Georges Bank region in the active season. All vectors include elements representing depth, latitude and longitude. This allows coupling of the population dynamics to the tide- and wind-driven Dartmouth model of New England regional circulation. Summary data from the physical model are used to advance vectors from resting-stock locations in Gulf of Maine basins through two generations to sites of readiness for return to rest. Supply of Calanus stock to Georges Bank comes from all of the gulf and from the Scotian Shelf. The top of the bank is stocked from western gulf basins; the North-east Peak is stocked from Georges Basin and the Scotian Shelf. All sources contribute to stock that accumulates in the SCOPEX gyre off the north-west shoulder of Georges Bank, explaining the high abundance recurrently seen in that region. There is some return of resting stock to Wilkinson Basin in the western gulf, but other basins must mostly be restocked from upstream sources to the north-east.     

Seasonal changes in respiration rates of copepodite stage V Calanus finmarchicus (Gunnerus)

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.     

Basin-scale advection and population persistence of Calanus finmarchicus

Advection of Calanus finmarchicus in the eastern North Atlantic was analysed using a particle-tracking model based on the Hamburg Shelf Ocean Model (HAMSOM). Quasi-static seasonal mean flowfields were simulated, archived and interpolated to represent a climatological-mean annual cycle. Particles had a simple prescribed depth profile comprising deep overwintering, spring ascent, a shallow-water phase followed by descent to overwintering depth. Export routes for C. finmarchicus from the model area were identified to the south of Greenland and to the north of the Lofoten Basin. Self-sustaining overwintering areas were identified by observing how closely particles returned to their origins after one calendar year. Several such areas were found, notably in the Norway and Lofoten Basins, and in the Færoe–Shetland Channel. The particle tracking was run for up to 10 years to demonstrate persistence of these cycles. Known features of the winter and summer distributions of C. finmarchicus were reproduced by the model. The success of the HAMSOM in simulating both the shallow and deep circulation of the eastern North Atlantic and Norwegian Sea was critical to the identification of these spatio-temporal cycles of C. finmarchicus.     

Lipids,buoyancy and the seasonal vertical migration of Calanus finmarchicus

The copepod Calanus finmarchicus remains in diapause for up to 5 months in the cold (<0.5°C) deep (>700 m) waters of the Faroe–Shetland Channel of the north-western approaches to the North Sea. While in diapause, C. finmarchicus has a high lipid content, up to 76% of dry weight, mostly in the form of wax esters. The question we address here is how copepods with such a high content of buoyant lipids can remain in diapause at depth for an extended period of time? The corollary to this is how this lipid content hinders and/or assists the copepods in their seasonal vertical migration? Part of the answer is due to the physical properties of wax esters. These have a thermal expansion and compressibility higher than that of sea water. Thus, depending on their relative composition (i.e. wax esters/water/protein/chitin), a copepod that is positively buoyant in warm surface waters can become neutrally buoyant in cold deep water. We develop a simple three component physical model of a copepod to explore how and where they attain neutral buoyancy, how the lipid content can aid in their ascent, and what fraction of the lipids can be utilized in ascent in gonad/egg formation while maintaining observed ascent rates. As well as being an energy reserve, the results show that rather than being a barrier to vertical migration, wax esters serve as an important regulator of buoyancy.     

Lipids, buoyancy and the seasonal vertical migration of Calanus finmarchicus

The copepod Calanus finmarchicus remains in diapause for up to 5 months in the cold (<0.5°C) deep (>700 m) waters of the Faroe–Shetland Channel of the north-western approaches to the North Sea. While in diapause, C. finmarchicus has a high lipid content, up to 76% of dry weight, mostly in the form of wax esters. The question we address here is how copepods with such a high content of buoyant lipids can remain in diapause at depth for an extended period of time? The corollary to this is how this lipid content hinders and/or assists the copepods in their seasonal vertical migration? Part of the answer is due to the physical properties of wax esters. These have a thermal expansion and compressibility higher than that of sea water. Thus, depending on their relative composition (i.e. wax esters/water/protein/chitin), a copepod that is positively buoyant in warm surface waters can become neutrally buoyant in cold deep water. We develop a simple three component physical model of a copepod to explore how and where they attain neutral buoyancy, how the lipid content can aid in their ascent, and what fraction of the lipids can be utilized in ascent in gonad/egg formation while maintaining observed ascent rates. As well as being an energy reserve, the results show that rather than being a barrier to vertical migration, wax esters serve as an important regulator of buoyancy.     

Distribution and abundance of overwintering Calanus finmarchicus in the Faroe–Shetland Channel

The distribution and abundance of overwintering Calanus finmarchicus in the Faroe–Shetland Channel was studied during December 1994 and January 1995. Maximum abundance of animals in the Channel was approximately 50 000 m⁻², with a peak concentration of 627 m⁻³ at a depth of 930 m. Copepodite stages IV and V accounted for > 98% of the population. A clear association was found between the horizontal and vertical distribution of animals and the Arctic water masses in the bottom of the Faroe–Shetland Channel. The Wyville–Thomson Ridge formed a barrier to the southern distribution of Arctic waters and the abundance of overwintering C. finmarchicus was 25 times lower to the south of the Ridge than to the north. Spatial variability in lipid content and composition indicated that overwintering C. finmarchicus in the southern Norwegian Sea were in poorer condition with respect to wax ester content, and in a more advanced state of emergence from overwintering, than the animals within the Channel. The overwintering stock in the Channel probably originated from the Norwegian Sea or from north of the Faroe Islands. The Faroe–Shetland Channel is an important source of animals advected into the northern North Sea in the spring (March/April). The population abundance in the Faroe–Shetland Channel was estimated to be 4.5 × 10¹⁴ individuals, which is more than adequate to account for the mean concentration of adult stages observed in the northern North Sea in April.     

Lipid content of Calanus finmarchicus during overwintering in the Faroe–Shetland Channel

Lipid class contents and composition of the copepodite stage CV and adult female Calanus finmarchicus were measured from the autumn to the spring along a transect crossing the Faroe–Shetland Channel. The data give a detailed vertical resolution of lipid content of C. fin-marchicus before, during and after the time of diapause. In the deep water, where C. finmarchicus was in diapause, the wax ester (WE) content did not change between the autumn and spring with the exception of that of females, which showed a significant decrease in lipid content by March. The concentrations of lipids were lower and more variable in both CV and females in surface waters over the sampling period. In March, the lipid concentration of both CV and females decreased significantly with decreasing depth. The triacylglycerol (TAG) content of both CV and females was high in October at all depths but had almost disappeared by December. The TAG was either utilized during this time period or converted to WE. TAG was most prominent in October and again in February in copepods in the upper water column. A simple diagnostic model is presented that estimates how C. finmarchicus allocates energy reserves from lipids into various metabolic functions. The cost of diapause, molting, gonad formation and ascending from depth is estimated based on the time series data. The model indicates that the major portion of the stored lipid goes into gonad formation, in physical ascent and in basic metabolism during ascent, whereas maintenance of diapause and molting is not expensive in terms of lipid utilization.     

Climate fluctuations and the spring invasion of the North Sea by Calanus finmarchicus

The 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⁻³ 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 north-westerly winds without leading to corresponding changes in C. finmarchicus abundance in the North Sea, because the NSDW overflow responds over longer (decadal) time scales.     

Modelling the dispersal of Calanus finmarchicus on the Newfoundland Shelf: implications for the analysis of population dynamics from a high frequency monitoring site

We investigated the drift of passive particles on the Newfoundland Shelf and western Labrador Sea using numerical simulations to assess the possible sources of plankton collected at a high frequency sampling site (S27; 47.55°N, 52.59°W) located near the coast of Newfoundland, Canada. We also summarized data detailing the seasonal stage succession of Calanus finmarchicus at that site, as well as along three oceanographic sections sampled in the spring, summer and autumn across the adjacent continental shelf. Simulations indicated that the Labrador and Newfoundland Shelves represent the major sources of particles transiting through the S27 site, with relatively minor contributions from the western Labrador Sea which are significant during a few months each year. The latter point may be affected by uncertainty in the representation of cross‐shelf transport associated with seasonal or short‐term variations in atmospheric and oceanic forcing, which may also affect the strength and location of bifurcation of the inner branch of the Labrador Current around the Grand Banks. Nevertheless, our results indicated that drift along the inner shelf is likely to be the primary source of copepods collected at S27 throughout most of the year. This in turn suggested that there may be a higher degree of connectivity between conditions in coastal areas of Newfoundland and those in Baffin Bay and west Greenland than with the southern half of the Labrador Sea.     

Initial design for a fish bioenergetics model of Pacific saury coupled to a lower trophic ecosystem model

A fish bioenergetics model coupled with an ecosystem model was developed to reproduce the growth of Pacific saury. The model spatially covers three different oceanographic spatial domains corresponding to the Kuroshio, Oyashio, and interfrontal (mixed water) regions. In this coupled model, three (small, large, and predatory) zooplankton densities which were derived from the lower trophic level ecosystem model were input to the bioenergetics model of saury as the prey densities. Although certain model parameters were imposed from other species’ bioenergetics, several model parameters were estimated from observational data specific to Pacific saury. The integrated model results reproduced appropriate growth rates of Pacific saury. Model sensitivities to water temperature and prey density are examined and observational methods to evaluate the model parameters are discussed.     

A model of the spring migration into the North Sea by Calanus finmarchicus overwintering off the Scottish continental shelf

A particle tracking model was used to investigate the annual spring invasion of the North Sea by Calanus finmarchicus copepodites which overwinter in deep water off the Scottish continental shelf. Flow fields generated by a hydrodynamic model (HDM) were used to simulate the advection of zero drag particles representing the copepods. Particles were released simultaneously from a regular lattice of start positions at a given depth ( D ₁), and ascended at a fixed rate ( dD/dt ) until they reached a final depth ( D ₂) in the surface layers. The proportion of particles reaching target areas in the northern North Sea was relatively insensitive to a moderate degree of variation (±20%) around chosen default values of the vertical migration parameters ( D ₁, D ₂ and dD/dt ), derived from field data. The inclusion of horizontal diffusion velocities and diel vertical migration in surface layers did not significantly affect the results. Sensitivity to wind direction was investigated by applying flow fields from HDM runs with different wind forcing scenarios. For the default vertical migration parameters, only north-westerly winds resulted in particles entering the North Sea from release locations north of the Iceland–Scotland Ridge, where dense aggregations of overwintering copepods were encountered during field surveys. The particle tracking model predicted that the major routes for the spring Calanus invasion into the North Sea were the East of Shetland Atlantic Inflow and the Norwegian Trench Atlantic Inflow, which agreed with seasonal trends observed in Continuous Plankton Recorder data. Overall, despite its relative simplicity, particle tracking was confirmed as a robust tool to explore the causal mechanisms behind the annual invasion of the North Sea by C. finmarchicus emerging from diapause in the deep waters off the Scottish continental shelf.     

Simulations of the Black Sea pelagic ecosystem by 1-D, vertically resolved, physical–biochemical models

The present paper summarizes the results of several simulations of the central Black Sea pelagic food web using three different 1-D, physical–biochemical, water column process models. The most simplified, five-compartment version is used to explore the robust biological features of the ecosystem and the role of upper-layer physics on the evolution of the euphotic zone biological processes. The other models, introducing additional biological complexities, show how these biologically structured models become more capable of simulating intensified subsurface summer production, more dynamic plankton structure arising after the increasing role of gelatinous carnivores in the ecosystem during the 1980s, and fairly sophisticated nitrogen cycling in the water column.