FISHERIES OCEANOGRAPHY Fish. Oceanogr. 8:4, 255±263, 1999 Nonlocal wind-driven fjord±coast advection and its potential effect on plankton and ®sh recruitment LARS ASPLIN,1,* ANNE GRO VEA SALVANES2 AND JON BENT KRISTOFFERSEN2 1 Institute of Marine Research, PB 1870 Nordnes, N±5024 Bergen, Norway 2 Department of Fisheries and Marine Biology, University of Bergen, N±5020 Bergen, Norway early life stages between neighbouring fjords and thus enhance genetic exchange. Key words: advection of ®sh eggs and larvae, ®sh recruitment, fjord±coast advection, nonlocal forcing, numerical 3D simulation, wind-generated upwelling and downwelling ABSTRACT INTRODUCTION The Bergen Ocean Model (BOM), a three-dimensional physical coastal ocean model, was used for a numerical simulation experiment to investigate shortterm effects of wind-generated coastal upwelling and downwelling on the dynamics of adjacent large outer and smaller inner fjords. The effect of the real alongshore wind regime on advection for an idealized fjord topography, resembling Masfjorden, western Norway, is used as an example. This modelling exercise is a supplement to, and its predictions support, the various hypotheses investigated in ecosystem simulation studies of the Masfjorden. The model predicts that coastal winds from the north cause upwelling and transport the upper water layer out from the fjords. Winds from the south cause downwelling and transport the upper water layer into the fjords. The transport is rapid and »50% of the upper water layer may be replaced within 1±2 days. Implications of these physical processes for the dispersal and retention of planktonic organisms and the early life stages of ®sh are discussed. If strong southerly winds occur frequently, this will transport planktonic organisms into the fjord and may increase the carrying capacity for planktivorous ®sh. In contrast, frequent strong northerly winds may reduce the abundance of planktonic organisms, including the early life stages of marine ®sh, and thus possibly reduce recruitment to fjord ®sh populations. Frequent shifts between southerly and northerly winds would cause an exchange of The western coast of Norway is rugged, with numerous fjords, which are typically long and deep, with widths varying from several kilometres in the outer parts to some hundreds of metres in inner regions. Many ®sh species spawn in the fjords, with the early pelagic life stages inhabiting the upper part of the water column (Sundby, 1991; Ellertsen et al., 1992). The major proportion of mesozooplankton also occurs in the upper 30 m of the water column (Aksnes et al., 1989). Outside the fjords, the coastal waters are generally strati®ed, with the baroclinic Norwegian Coastal Current (NCC) as an important dynamic feature (Sñtre and Mork, 1981; Sñtre et al., 1988). There is normally a signi®cant exchange of water between the fjords and the NCC (Svendsen, 1980; Aksnes et al., 1989; Aure et al., 1996). Many physical processes are responsible for this water exchange (e.g. wind, tides, freshwater run-off; review: Farmer and Freeland, 1983). Of special interest for the present work is the importance of nonlocal forcing (such as coastal wind) of fjord water mass dynamics as described by Klinck et al. (1981) and Proehl and Rattray (1984). The topography of fjord areas is a complicated but important factor in the determination of water exchange, and its complexity is probably a major reason why this subject has received relatively little attention. In the Northern Hemisphere, the net wind-driven transport of water is to the right of the wind direction (Ekman transport; Gill, 1982). Along the western coast of Norway, northerly alongshore wind thus produces coastal upwelling, and southerly alongshore wind produces coastal downwelling. Associated with the upwelling and downwelling are coastal currents. The widths of the coastal currents are comparable to *Correspondence. e-mail: [email protected] Received 6 April 1998 Revised version accepted 30 September 1998 Ó 1999 Blackwell Science Ltd. 255 256 L. Asplin et al. the internal radius of deformation (i.e. the ratio of the internal wave speed and the Coriolis parameter; Gill, 1982). Along the west coast of Norway, the coastal currents associated with the upwelling and downwelling typically extend »5 km offshore. The coastal upwelling or downwelling will be reduced if the coastline is broken by a fjord mouth, and a coastal trapped, internal Kelvin wave bore will be generated. Such a Kelvin wave bore can propagate into and inside fjords having a width comparable to the internal radius of deformation. The propagation of the bore also sets up a temporary current ®eld, which generates exchange of water between the coast and the fjord, but the extent of this exchange has hitherto been largely unknown. Field studies in fjords generally report the highest biomass of mesozooplankton, including Calanus ®nmarchicus, in the upper water layers (Aksnes et al., 1989). Simulation modelling of the ecosystem dynamics of Masfjorden (Fig. 1) and other west Norwegian fjords has shown that a highly ¯uctuating transport of C. ®nmarchicus into the fjords has a pronounced effect on the carrying capacity for ®sh at higher trophic levels (Salvanes et al., 1992, 1995). Moreover, ®eld investigations in Masfjorden reveal large interannual variations in cod recruitment as Figure 1. Map of the western coast of Norway, showing the Fensfjorden and Masfjorden. Ó 1999 Blackwell Science Ltd., Fish. Oceanogr., 8:4, 255±263. Nonlocal wind-driven fjord±coast advection one-year-olds (Salvanes and Ulltang, 1992; Nordeide et al., 1994), partly caused by density-independent factors at the egg and larval stage (Ellertsen et al., 1992), but modi®ed by density-dependent predation and cannibalism after settlement (Ellertsen et al., 1992; Salvanes et al., 1994; Salvanes et al., 1995; Salvanes and BalinÄo, 1998; Giske and Salvanes, 1999). Because the early life stages of marine ®sh generally inhabit the upper water layers (Sundby, 1983; Ellertsen et al., 1992), they are affected by advection of these layers (Bailey, 1981). Hence, for local ®sh populations, wind-generated advection has the potential to transport eggs and larvae to or from suitable habitats, and similarly to transport zooplankton. The importance of advection for ®sh recruitment relative to other factors such as starvation and predation is not well known. There are indications that the in¯uence of advection can be signi®cant (Taggart and Leggett, 1987; Pepin et al., 1995; Svendsen et al., 1995), but it is not clear at what time or spatial scales advection operates. In the present paper, we demonstrate that the ¯uctuations in water transport caused by nonlocal wind conditions operate at short time scales. The modelling exercise presented is based on existing knowledge and is a natural extension of, and supplement to, the previous ecosystem simulation modelling developed for an idealized food web of Masfjorden (Salvanes et al., 1992, 1995). The ecosystem simulations of Salvanes et al. (1992, 1995) and the ®eld observations reported in Aksnes et al. (1989) provided the framework for the present modelling study by indicating connections between zooplankton advection and ®sh carrying capacity. However, the ecosystem modelling simpli®ed advection to a bordering forcing function, which changed monthly and which did not have any depth solution. Furthermore, the physics of fjord/coast water dynamics were not incorporated in the ecosystem model, because the focus was on the dynamics of the biological processes and species interactions within the ecosystem. In the present study, we use a three-dimensional numerical simulation experiment to show that the short-term effects of nonlocal wind-generated coastal upwelling and downwelling are important for water exchange between coast and fjords. The wind regimes used for the simulation experiments are frequently observed on the west coast of Norway as well as at Shetland or elsewhere in the northern North Sea. The fjord topography is idealized, keeping fjord sizes and distances from the coast resembling the real topography of Masfjorden and Fensfjorden, through which Masfjorden is connected to coastal waters of western Ó 1999 Blackwell Science Ltd., Fish. Oceanogr., 8:4, 255±263. 257 Norway. Finally, the consequences of the results for retention and dispersal of planktonic organisms and early life stages of ®sh are discussed. MATERIALS AND METHODS Model design The numerical model used for the simulations of fjord and coastal water exchange is a threedimensional, primitive equation, time-dependent, r-coordinate, ocean circulation model (the Bergen Ocean Model; BOM) developed by Berntsen et al. (1996). The prognostic variables of the model are three components of velocity, potential temperature, salinity, surface elevation and two variables representing turbulent length scale and turbulent kinetic energy. The BOM has an embedded turbulence closure submodel (Mellor and Yamada, 1982). The governing equations are the equations for conservation of mass, momentum, temperature and salinity, along with the hydrostatic equation in the vertical and an equation of state relating salinity and potential temperature to potential density. The equations are solved by ®nite difference techniques on a staggered Arakawa C-grid. The time differencing is implicit. Two numerical experiments were performed to simulate the hydrodynamic response to winds along the coast in a strati®ed, coupled coastal ocean and fjord system. The ®rst experiment simulated wind from the south, the open ocean being to the left of the southern wind component and the coast on the right. The combination of this wind direction and coastal topography generated coastal downwelling. The other experiment, with the wind in the opposite direction, generated coastal upwelling. The numerical experiments were idealized so as to focus on the physical processes and to keep the Kelvin wave problem tractable, although still having realistic characteristics of Norwegian fjord and coastal areas. The model domain consisted of a rectangular coastal ocean and a fjord system (Fig. 2). The main fjord topography (outer fjord) was idealized to 7 km wide and 15 km long, and the inner fjord arm was 1500 m wide and 12 km long, thereby resembling the topography of Masfjorden and Fensfjorden (Fig. 1). The bottom depth throughout the modelled area (i.e. both the coastal ocean and the fjords) was set constant at 100 m. The initial conditions were of no water velocities, no surface elevations and a constant temperature ®eld of 10°C. The salinity ®eld was horizontally homogeneous, but with linear strati®cation in 258 L. Asplin et al. Figure 2. Top view of the idealized model domain used for the numerical simulations. A and B refer to sections displayed in Fig. 5. Wind stress on the coastal ocean was the only forcing of the model (i.e. no local wind was speci®ed for the fjords). The wind direction was along the coast only. The total simulation period was 72 h. The simulated wind pattern was chosen, based on observations in the spring frequently registered at the Norwegian Meteorological Institute. The maximum wind speed was 16 m s±1, and the absolute wind speed values were the same for the two simulations. Wind speed was increased linearly from 0 to 16 m s±1 over the ®rst 12 h, then held constant at 16 m s±1 for the next 36 h before it was reduced linearly to 10 m s±1 through the last 24 h of the simulations. The model grid had a horizontal resolution of 500 m and was thus able to resolve adequately the internal radius of deformation (»4 km) based on a typical internal wave speed of »0.5 m s±1. Vertically, 35 grid nodes were used, with the grid size (expressed in z-coordinates, assuming no surface elevation) increasing from 0.25 m in the upper few metres to 5 m below 10 m depth. This ®ne vertical grid resolution was needed to simulate adequately the wind-driven ¯ow and internal wave activity (Asplin, 1999). The time step of the simulations was 75 s, obeying the time-step constraints of the numerical model grid. RESULTS the upper 50 m, increasing from a salinity of 33 at the surface to 35 at 50 m depth. Coastal downwelling and advection into the fjords Coastal downwelling generated by the southerly wind caused a substantial up-fjord advection of upper-layer (0±45 m depth) water masses (Table 1; Fig. 3). During the ®rst 2 d of the simulation, the internal Kelvin wave bore created by the reduced downwelling in the fjord mouth area propagated into and out of the outer fjord (with the land on its right). The Kelvin wave was Table 1. Relative distribution (%) of the tracer water masses after 72 h of the simulations of coastal downwelling (southerly wind) and upwelling (northerly wind). The initial distributions of the water masses are shown by the 100% values in parentheses, e.g. under downwelling conditions, MASS 1 was initially 100% in the outer fjord, between 0 and 45 m depth. See text for explanation. Inner fjord (Masfjorden) Outer fjord (Fensfjorden) Coastal ocean 0±45 m Simulated Conditions Water Mass 0±45 m 45±100 m 0±45 m 45±100 m Downwelling MASS 1a MASS 2b 13 28 (100) 3 42 31 (100) 6 46 24 2 0 5 0 Upwelling MASS 1a MASS 2b 1 30 (100) 0 0 42 (100) 51 0 0 55 19 2 0 a b 45±100 m Representing an idealized Fensfjorden. Representing an idealized Masfjorden. Ó 1999 Blackwell Science Ltd., Fish. Oceanogr., 8:4, 255±263. Nonlocal wind-driven fjord±coast advection 259 Figure 3. Simulated dynamic transfer and retention of tracer water masses between the large outer fjord (resembling Fensfjorden) and a small inner fjord (resembling Masfjorden) for the downwelling and upwelling simulation experiments. (a) The initial horizontal distribution of the tracer water masses; vertically, the tracer water masses extend from the surface to 45 m depth. (b and c) Changes in the distribution of the tracer water masses above 45 m depth in (b) Fensfjorden and (c) Masfjorden through the simulated period. DW refers to downwelling and coastal wind from the south. UPW refers to upwelling and coastal wind from the north. below this depth range (out of the fjords). Large horizontal variations were found in both the ¯ow ®eld and the hydrography, with in¯ow along the left fjord side (looking out from the fjord) and out¯ow along the right fjord side. About 20 h after initiation of wind forcing, the circulation connected with the Kelvin wave bore had started to push water into the upper 40±50 m of the water column of the inner fjord (1.5 km wide). Downwelling of upper-layer water took place inside the inner fjord, with out¯ow of water into the outer fjord in the lower layer (below 40±50 m depth). Seventy-two hours after the initiation of the wind, most of the water mass originally present in the upper layer of the outer fjord (MASS 1 referring to the idealized Fensfjorden, Table 1) was retained in the fjord system, with 31% still in the upper layer whereas 46% had been transported to the lower layer of the outer fjord and 13% to the upper layer of the inner fjord. Of the water mass originally in the inner fjord (MASS 2 referring to the idealized Masfjorden, Table 1), 28% was still retained in the upper layer after 72 h, 42% had been transported down to the out¯owing lower layer within the inner fjord, and 24% to the lower layer of the large outer fjord. The drift of the passive particles in the surface layer for the downwelling experiment also showed a tendency for a movement into the fjord system (Fig. 4a). approximately two-layered, with ¯ow in the direction of propagation in the upper 40±50 m of the water column (into the fjords) and in the opposite direction Ó 1999 Blackwell Science Ltd., Fish. Oceanogr., 8:4, 255±263. Coastal upwelling and advection out of the fjords Coastal upwelling, generated by a northerly wind, ¯ushed the upper water layer out of both the large outer and the small inner fjord (Table 1; Fig. 3). During the ®rst 2 d of the simulation, the internal Kelvin wave bore, created by the reduced upwelling in the fjord mouth area, propagated into and out of the outer fjord in the same direction as the bore created by the coastal downwelling (land on its right). The simulated upper-layer (0±20 m depth) ¯ow was in the opposite direction to the wave propagation, and the lower-layer ¯ow was in the same direction as the wave 260 L. Asplin et al. Figure 4. Advection of passive particles (identi®ed by number) in the surface layer (upper 1 m) for (a) the downwelling experiment (southerly coastal wind) and (b) the upwelling experiment (northerly coastal wind). The open triangles refer to start positions of the particles (at 0 h) and the open squares to end positions (after 72 h). propagation. Upwelling of lower-layer water took place inside the inner fjord, and in¯ow of water from the outer fjord was in the lower layer. During coastal upwelling, a rapid ¯ush-out of the upper layer was seen (Table 1). After 72 h, 57% of the water mass originally in the outer fjord (MASS 1) had been advected to the open ocean. Also, a signi®cant proportion of the water mass originally in the inner fjord (MASS 2) had been advected out and only 30% of MASS 2 was retained within the inner fjord; 51% had been transported to the outer fjord and as much as 19% reached the coastal ocean. The illustration of the passively drifting particles in the surface layer for the upwelling experiment also showed a tendency towards outwards drift from the fjords to the coastal ocean (Fig. 4b). Water exchange in the inner fjord (Masfjorden) Figure 5 illustrates vertical sections of current vectors and isohalines along the small inner fjord 30 h after the initiation of the coastal wind (which represents the time at which maximum advective ¯ow occurs). The maximum current speed in the inner fjord is about 0.2±0.3 m s±1 and con®ned to the upper 20 m of the water column. The results from the downwelling simulation show an in¯ow of water in the upper 40± 50 m of the water column and an out¯ow below that depth range (Fig. 5a), whereas the results from the upwelling simulation show the opposite (Fig. 5b). Note that the upper layer was thinner (20±30 m) for the upwelling simulation than for the downwelling simulation (40±50 m). Figure 6 shows that the simulated coastal downwelling generated a net positive volume ¯ux to the upper layer of the inner fjord and that the simulated coastal upwelling caused a negative volume ¯ux re¯ecting a ¯ushing-out of the upper layer of the inner fjord. DISCUSSION Previous explanations of the observations of large volume exchange between coast and fjord systems were incomplete because only selected aspects of the fjord dynamics were considered (Aksnes et al., 1989; Kaartvedt and Svendsen, 1995; Wassman et al., 1996), or the solutions were restricted to narrow fjords (nonrotating dynamics) and for time-independent dynamics (Stigebrandt, 1990; Aure et al., 1996). The present study focuses on the effects of real, but nonlocal wind-generated fjord±coast advection on the dynamics of an idealized topography of an actual fjord± coast system (Masfjord±Fensfjord±coast) and represents an alternative and more complete approach. Ó 1999 Blackwell Science Ltd., Fish. Oceanogr., 8:4, 255±263. Nonlocal wind-driven fjord±coast advection Figure 5. Vertical sections along the middle of the inner fjord (transect A±B; Fig. 2) of current vectors and isohalines after 30 h for (a) the downwelling simulation and (b) the upwelling simulation. The predictions from the simulations of the present study support the hypothesis of Aksnes et al. (1989) and Salvanes et al. (1992) that regional wind conditions are of importance in the dynamics of fjords having a greater width than the appropriate dynamic length scale (internal radius of deformation). In the Northern Hemisphere, the internal radius of deformation decreases towards the north (Gill, 1982), which means that narrow fjords at northern latitudes may have rotational dynamics whereas fjords of the same dimensions further to the south may not. Wind regimes that are common along the western Norwegian coast, as well as in areas of other North Atlantic fjord systems, are shown to generate advection of water into fjords located far from the coast and thus from the origin of the physical driving processes. Our modelling approach demonstrates that nonlocal Ó 1999 Blackwell Science Ltd., Fish. Oceanogr., 8:4, 255±263. 261 Figure 6. The calculated volume ¯ux (m3 s±1) in the upper 50 m of the water column through the mouth of the inner fjord (Masfjorden). The solid line represents the results of the downwelling experiment (southerly coastal wind) and the dashed line represents the results of the upwelling experiment (northerly coastal wind). northerly regional winds will ¯ush out the upper water layer of fjords, and that nonlocal southerly regional winds will cause a net transport into the fjords. These events take place in a remarkably short period (about 1 day) and the absolute value of the volume ¯ux is 1±2 orders of magnitude greater than typical volume ¯uxes associated with classical estuarine circulation (Stigebrandt, 1990). The validity of the simulation is supported by the good correspondence between the simulated depth range and direction of the advective layers with the vertical distribution of residual current across the sill of Masfjorden, as reported in Aksnes et al. (1989). The numerical results clearly demonstrate that studies on local fjord water mass dynamics should take into account the in¯uence of remote coastal winds. Furthermore, in numerical simulations of particular fjord systems, a better representation of the fjord topography, wind ®elds and details of the initial density distribution will improve the realism of the solutions. As to the bottom topography, details of this will have only a minor in¯uence on the simulated dynamics as long as the bottom depth is greater than »100 m. Because mesozooplankton and most ®sh eggs and larvae are generally concentrated at depths shallower than 30 m (Sundby, 1983; Aksnes et al., 1989; Ellertsen et al., 1992), it is likely that nonlocal winddriven fjord±coast advection of the surface water layers will have biological implications for these organisms. This will depend on the wind regime, for example. 262 L. Asplin et al. · Frequent strong southerly winds will transport passive particles into fjords and generate advective supply to the carrying capacity of fjords. Frequent strong northerly winds will ¯ush the surface layers out from the fjords. This implies a reduced carrying capacity for planktivorous organisms and reduced retention of the early life stages of marine ®sh. Frequent shifts between southerly and northerly winds will cause an exchange of plankton between neighbouring fjords via the coastal water and will thus counteract genetic isolation. Various strategies may have evolved to cope with such advective regimes, for example vertical migration (Eiane et al., 1998), selection of spawning grounds at locations with increased water residence time (Ellertsen et al., 1992; Brander, 1994), spawning demersal eggs, and `bet-hedging' or `spreadingthe-risk' strategies by spawning repeatedly over a prolonged period (Den Boer, 1968; Slatkin, 1974). However, the eggs and larvae of many species are concentrated in the upper advective layer, where they are vulnerable to advection. Nevertheless, for many species the depth of spawning and the buoyancy of eggs may have evolved to reduce dispersal of the youngest stages. For example, the spawning period for coastal cod (Gadus morhua) in the fjords of western Norway (February to April) coincides with frequent winds from the south, and consequently frequent downwelling. The eggs will ascend in water of salinity greater than 29 with a vertical speed (1 mm s±1) that exceeds the typical vertical speed of downwelling water (0.1 mm s±1; Sundby, 1983, 1991); hence cod eggs are expected to be retained in fjords during downwelling. The spawning of other species may be similarly adapted to upwelling situations that predominate in the summer. For example, the mesopelagic ®sh Maurolicus muelleri has a bet-hedging strategy with repeated spawnings during the summer, and, usually, a deep egg distribution (Lopes, 1979; John and Kloppmann, 1989, 1993) that may prevent advection out of a fjord during upwelling episodes. Our study suggests that the sequence of coastal wind episodes (including wind direction, strength and duration) could in¯uence the advective transport and potentially contribute to variability in ®sh recruitment, abundance of planktonic organisms and settlement areas. Pelagic early life stages of ®sh would be concentrated in inner fjords when coastal wind is from south, whereas frequent winds from the north would ¯ush out the fjords. ACKNOWLEDGEMENTS The authors received funding from The Research Council of Norway, including the Programme for Supercomputing (through a grant of computing time). Thank you, Jarle Berntsen, for providing the Bergen Ocean Model. 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