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.     

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.     

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.     

A Lagrangian ensemble model of Calanus finmarchicus coupled with a 1D ecosystem model

A population dynamics model of Calanus finmarchicus based on Lagrangian particles has been coupled with a 1-D ecosystem model. Each of the particles represents a variable number of copepods which experience the same fate. Therefore all copepods of a single particle represent a cohort and are characterized by a common set of individual properties such as age, development-stage, depth, structural weight (length), lipid pool or food satiation. The physical environment is parameterized by a 1-D-water column with a vertical resolution of 1 m and a maximum depth of 800 m. Copepod food supply is provided by an interactive Eulerian NPZD model where Z represents microzooplankton. The model correctly reproduces both the dynamics of the ecosystem and the life history of the copepods in the Norwegian Sea. Simulated results of trajectories of particles in the water column, and of individual growth and stage development were analysed. Results on seasonal abundance, development time, number of generations, depth profiles, and patterns of diurnal and ontogenic migration are compared with field data from OWS India.     

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.     

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.     

Interannual differences in larval haddock survival: hypothesis testing with a 3D biophysical model of Georges Bank

The ultimate goal of early life studies of fish over the past century has been to better understand recruitment variability. As evident in the Georges Bank haddock (Melanogrammus aeglefinus) population, there is a strong relationship between recruitment success and processes occurring during the planktonic larval stage. This research sought new insights into the mechanisms controlling the recruitment process in fish populations using biological–physical modeling methods together with laboratory and field data sets. We created the first three‐dimensional model of larval haddock on Georges Bank by coupling models of hydrodynamics, lower trophic levels, a single copepod species, and larval haddock. Interactions between feeding, metabolism, growth, vertical behavior, advection, predation, and the physical environment of larval haddock were quantitatively investigated using the coupled models. Particularly, the model was used to compare survival over the larval period and the sources of mortality in 1995 and 1998, 2 years of disparate haddock recruitment. The results of model simulations suggest that the increased egg hatching rates and higher food availability, which reduced starvation and predation, in 1998 contributed to its larger year‐class. Additionally, the inclusion of temperature‐dependent predation rates produced model results that better agreed with observations of the mean hatch date of survivors. The results from this biophysical model imply that food limitation and its related losses to starvation and predation, especially from hatch to 7 mm, may be responsible for interannual variability in recruitment and larval survival outside of the years studied.     

Calanus finmarchicus egg production and food availability in the Faroe–Shetland Channel and northern North Sea: October–March

Physiological characteristics (egg production, lipid content and gonad development stage) of Calanus finmarchicus were examined on five cruises in the north-east Atlantic carried out between the months of October to March in the years 1993–95 and related to phytoplankton food availability. Appreciable egg production rates were only recorded in October and March. Published values for maximum water volume filtered daily by individual females and the standing stock of phytoplankton (estimated from chlorophyll concentration) are used to argue that ingestion of phytoplankton carbon alone was not sufficient to support the egg production rates recorded in March. Data are presented that indicate C. finmarchicus is fuelling egg production through use of internal lipids. We argue that this species exhibits a life history strategy whereby females release a limited number of eggs to the water column upon returning to the surface waters following diapause, i.e. prior to the onset of the spring bloom. This pre-spring bloom egg production will comprise only a small component of the annual Calanus egg production in shelf waters. Nevertheless, given that the recruitment of Calanus to their feeding grounds over the shelf is a function of transport of the copepods from off-shelf to shelf regions via ocean currents, a burst of egg production in the period immediately following diapause and prior to the population's transport to shelf waters, would increase the chances of large numbers of the population reaching shelf regions. Thus, even a limited pre-spring bloom burst of egg production may have important consequences for recruitment in this species.     

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.     

Juvenile cod (Gadus morhua) mortality and the importance of bottom sediment type to recruitment on Georges Bank

The demersal settlement of pelagic juvenile fish has been considered a critical period in which the final adjustment is made to the size of a year class. Distribution patterns of pelagic and recently settled juvenile cod (Gadus morhua) were examined from nine surveys on Georges Bank during the summer over 5 years, 1984–1989, to relate juvenile survival to the sedimentary environment. Pelagic juveniles were widespread across Georges Bank in June, and by mid‐July they occurred on all bottom types from sand to gravel on eastern Georges Bank. However, by late July‐early August they were mostly abundant on the northeastern edge gravel deposit, which with its complex relief, provides abundant prey and refuge from predators. A bank‐wide estimate of pelagic juvenile abundance in 1986 and 1987 was used to assess mortality of the recently settled juveniles and to evaluate the relative importance of survivors from the northeastern edge gravel area to recruitment of the Georges Bank population. Settlement mortality rates over 1–2 months on the northeastern gravel area ranged from 3 to 8% day^?1, which compared reasonably with other studies. The seasonal abundance of the pelagic juveniles was almost an order of magnitude higher in 1987 than 1986; however, recruitment at age 1 was similar, indicating that a high mortality of the demersal juveniles occurred in 1987. The limited northeastern gravel area on Georges Bank may represent a survival bottleneck depending on the variability in the distribution and abundance of juvenile cod settlement in relation to that of their predators.     

Optimum temperature and food-limited growth of larval Atlantic cod (Gadus morhua) and haddock (Melanogrammus aeglefinus) on Georges Bank

We estimated recent growth of Atlantic cod (Gadus morhua) and haddock (Melanogrammus aeglefinus) larvae collected on the southern flank of Georges Bank in May 1992–94 from the ratio of RNA to DNA (R/D) and water temperature. Growth of both species increased with water temperature to about 7°C and then decreased. The highest growth rates were observed in May 1993 at water temperatures around 7°C. These data confirm an earlier observation of comparable temperature optima for growth of Atlantic cod and haddock larvae in the north‐west Atlantic. Comparisons of field growth rates and temperature optima with data for larvae cultured at high temperatures and prey densities in the laboratory suggest that growth may have been food‐limited at higher temperatures on Georges Bank. Given that 7°C is the long‐term mean water temperature on the southern flank in May and that climate models predict a possible 2–4°C rise in water temperatures for the western North Atlantic, our findings point to a possible adverse effect of global warming on Atlantic cod and haddock.     

Sensitivity analysis of sea scallop (Placopecten magellanicus) larvae trajectories to hydrodynamic model configuration on Georges Bank and adjacent coastal regions

The previous larval-trajectory modeling studies on Georges Bank were assessed through process-oriented Lagrangian-tracking comparison experiments using the high-resolution Gulf of Maine/Georges Bank Finite-Volume Coastal Ocean Model (GOM-FVCOM). The results indicate that in a strong nonlinear system such as Georges Bank, the passive tracer movement is driven by a fully three-dimensional Lagrangian flow field that varies in space and time due to large tidal excursion and steep bottom topography. The particle-tracking methods developed based on the assumption of weak nonlinearity of the flow field are not applicable to Georges Bank. The results of previous larval transport studies driven by circulation fields constructed under the weak-nonlinearity assumption need to be interpreted with caution. In the present work, the influence of model physical setups on sea scallop larval dispersal and settlement on Georges Bank and adjacent shelf regions is examined. Distinct differences in the spatial distribution of the passive larvae predicted by the model under various physical conditions suggest that a fully nonlinear model driven by realistic spatially and temporally varying forcing should be employed for Lagrangian-based studies of fishery population dynamics on Georges Bank.     

An adjoint data assimilation approach to diagnosis of physical and biological controls on Pseudocalanus spp. in the Gulf of Maine–Georges Bank region

The underlying scientific objective here is to determine the mechanisms that control seasonal variations in the abundance of Pseudocalanus spp. in the Georges Bank–Gulf of Maine region. It is postulated that the observed distributions result from the interaction of the population dynamics with the climatological circulation. The problem is posed mathematically as a 2-D advection–diffusion–reaction equation for a scalar variable. Given an initial distribution of animals, we seek the population dynamics source term R ( x , y ) such that integration of the forward model will result in predictions3 that minimize the sum of squares of differences with observed concentrations at a later time. An adjoint data assimilation technique has been designed for these purposes.
This approach has been used to invert for the population dynamics associated with the transition between bimonthly (i.e. for 2 months) climatological Pseudocalanus spp. distributions derived from MARMAP data. Vertically averaged velocity and diffusivity fields diagnosed from hydrodynamical simulations of the climatological flow are specified. Solutions converge rapidly, and the procedure reduces the cost function by an order of magnitude within 50 iterations. The resulting population dynamics vary considerably in space and time, as does the balance between local tendency, physical transport and biological source terms. Generally speaking, the patterns in population dynamics are not inconsistent with current knowledge concerning potential controls such as predation and food limitation. Analysis of the solutions indicates that the Pseudocalanus spp. population centres located in the western Gulf of Maine and on Georges Bank may be self-sustaining, in contrast to prior studies which characterize the former as a source region for the latter.     

Growth-dependent survival mechanisms during the early life of a temperate seabass (Lateolabrax japonicus): field test of the ‘growth–mortality’ hypothesis

Three concepts based on size, time and growth rate are contained in the functional mechanisms of the ‘growth–mortality’ hypothesis, and are known as the ‘bigger is better’, ‘stage duration’ and ‘growth-selective predation’ hypotheses, respectively. Although it is sufficiently documented in fishes that faster growing and bigger larvae selectively survive, synergistic operation of the three components of the ‘growth–mortality’ hypothesis within a species have received little attention. In this study, we have tested the components of the ‘growth–mortality’ hypothesis based on the growth characteristics of original population (OP) and survivors (SV) of Japanese seabass (JSB) Lateolabrax japonicus. Larval and juvenile JSB were repeatedly sampled from the Tango Sea in 2007 and 2008. Otoliths from larvae and juveniles were analyzed to produce back-calculated daily records of size-at-age and growth rate, and were compared between OP and SV. Selective survival for fast-growing and bigger larvae was evident, and our results strongly support the ‘bigger is better’ hypothesis as well as the ‘growth-selective predation’ hypothesis. Growth rates of the individuals that metamorphosed into juveniles had significantly faster growth rates than non-metamorphosing larvae older than 48 days, the minimum age for metamorphosis, suggesting a clear relationship between growth rates and the timing of metamorphosis. On average, SV completed the larval stage ∼13 days earlier than those of OP, supporting the ‘stage duration’ hypothesis. Thus, SV of JSB exhibited traits consistent with all aspects of the ‘growth–mortality’ hypothesis: faster growth, bigger size-at-age, and shorter larval stage duration (LSD), i.e., larvae with faster growth, bigger size-at-age and a shorter LSD selectively survived the larval period. Although maternal influence on growth and survival was evident, factors that regulate growth–mortality mechanisms remain to be explored for JSB.     

Dispersal of black sea bass (Centropristis striata) larvae on the southeast U.S. continental shelf: results of a coupled vertical larval behavior – 3D circulation model

In the marine environment, pelagic dispersal is important for determining the distribution and abundance of populations, as well as providing connections among populations. Estimates of larval dispersal from spawning grounds are important to determining temporal and spatial patterns in recruitment that may have significant influences on the dynamics of the population. We present a case study of the dispersal of Centropristis striata (black sea bass) larvae on the southeast U.S. continental shelf. We use a coupled larval behavior – 3D circulation model to compare the effects of the timing and location of spawning against that of larval vertical migration on larval dispersal. Using the results of field data on larval vertical distributions, we compare the dispersal of virtual 'larvae' which have ontogenetic changes in vertical behavior with that of particles fixed near the surface and near the bottom. Larvae were released at potential spawning sites four times throughout the spawning season (February through May) for 3 yr (2002–04) and tracked for the assumed larval duration (from 27 to 37 days including the egg stage). Results indicate that adult behavior, in the form of spawning time and location, may be more important than larval vertical behavior in determining larval dispersal on the inner- and mid- continental shelves of this region.     

Effects of a peroxide-based commercial product on bacterial load of larval rearing water and on larval survival of two species of Sparidae under intensive culture: preliminary study

Larvae of two Mediterranean Sparidae species, Sparus aurata and Dentex dentex , were used to test the efficacy of a peroxide-based product (Ox-Aquaculture^©) on the reduction in bacterial load in larval rearing water and its effects on larval survival. Eleven-day-old S. aurata larvae and 15-day-old D. dentex larvae were exposed to different concentrations of Ox-Aquaculture^© (50, 100 and 200 mg L⁻¹, and 20 and 50 mg L⁻¹ respectively) for 1 h. Results indicated that 50 and 20 mg L⁻¹ were the most effective concentrations for the reduction in bacterial load (at least one order of magnitude) after 1 h treatment, without affecting larval survival and/or vitality in 11 dph S. aurata and 15 dph D. dentex larvae respectively. Ox-Aquaculture^© concentrations of 200 and 50 mg L⁻¹ during 1 h affected negatively final survival rate of the larvae of S. aurata and D. dentex respectively.     

Blue crab (Callinectes sapidus) larval settlement in North Carolina: environmental forcing,recruit–stock relationships,and numerical modeling

Ecosystem‐based fishery management requires an understanding of relationships between fisheries and environmental variability. The purposes of this study were to investigate (1) how environmental forcing drives variability in larval settlement of the blue crab Callinectes sapidus, (2) whether larval settlement data are useful for determining recruit–stock relationships, and (3) whether environmental data can be used to predict settlement. Megalopae settling on passive collectors in the Newport River estuary, NC, were collected nightly from September to November for 13 yr from 1993 to 2009. Observed settlement was compared with winds, tides, and predicted settlement (numerical model derived from environmental data) using cross‐correlation analysis. Weather events associated with settlement pulses were also identified. Recruit–stock relationships were developed by comparing observed and predicted settlement indices to lagged NC fishery data. Settlement was positively correlated with winds from the northeast, southeast and south in 12 of 13 yr and with maximum duration nighttime flood tides in all years. Settlement pulses occurred when high pressure systems formed over the southeast USA (58% of pulses) or hurricanes (25%), and nearly all occurred during nighttime flood tides (85%). Significant recruit–stock relationships occurred between observed settlement and fishery landings (r² = 0.96) and effort (r² = 0.94) in wet years and catch per unit effort (r² = 0.98) during dry years. The timing of settlement pulses, but not the magnitude of annual settlement, was successfully predicted using the settlement model. These results suggest that blue crab fisheries in NC are influenced both by larval settlement and post‐settlement processes (freshwater inflow).