John L. Largier

Upwelling shadows as nearshore retention sites: the example of northern Monterey Bay

Abstract Periods of elevated temperatures in northern Monterey Bay suggest that this is a region of increased residence time during periods of active upwelling. Nearshore increases in near-surface temperature often coincide with offshore decreases in temperature as cold, upwelled water is advected into the center of the bay—the juxtaposition of warm and cold water enhancing the thermal signature of a feature described as an ‘upwelling shadow’. We present a variety of data collected from 1988 to 1993 invoking a shallow, stratified, cyclonic circulation in northern Monterey Bay to explain the upwelling shadow as a dynamic response to upwelling north of the bay. Residence times associated with this warm recirculation are of the order of 8 days which is comparable to semi-enclosed embayments whereas Monterey Bay is an open embayment. We estimate a continuous replacement of upwelling shadow water with offshore water at a rate of 0.083 day −1 , and we suggest that a circulation velocity of 0.1 m · s −1 , is needed to produce the thermal signature of the feature given published values of surface heat flux in the central California coastal region. The thermal signature of this upwelling shadow persists through much of the upwelling season, surviving brief relaxations in upwelling but breaking down during prolonged relaxations of the order of a week or longer. The stratification and coherent recirculation of this feature is likely to be very important for such biological processes as productivity and the dispersal and recruitment of larvae.

AVOIDING CURRENT OVERSIGHTS IN MARINE RESERVE DESIGN

The pun in the above title reflects two points. First, marine life cycles com- monly include a dispersive juvenile stage that is moved about by ocean currents. This stage often is the predominant, or only, means of dispersal that connects spatially disjunct pop- ulations. As a consequence, details of dispersal likely play a critical role in determining the effectiveness of marine reserves as a management and conservation tool. Curiously, however (and this is the second point of the title), although dozens of models for marine reserves now exist, few actually account explicitly for larval dispersal. Moreover, those that do include dispersal, do so almost exclusively by considering it to be a nondirectional spreading process (diffusion), ignoring the effects of directional transport by currents (ad- vection). Here we develop a population dynamical model for marine organisms with rel- atively sedentary adults whose larvae are transported in a simple flow field with both diffusive spreading and directional characteristics. We find that advection can play a dom- inant role in determining the effectiveness of different reserve configurations. Two of the most important consequences are: (1) with strong currents, multiple reserves can be mark- edly more effective than single reserves of equivalent total size; and (2) in the presence of strong currents, reserves can significantly outperform traditional, effort-based manage- ment strategies in terms of fisheries yield, and do so with less risk. These results suggest that successful reserve design may require considerable new efforts to examine explicitly the role of dispersal of young.

Connectivity, sustainability, and yield: bridging the gap between conventional fisheries management and marine protected areas

A substantial shift toward use of marine protected areas (MPAs) for conservation and fisheries management is currently underway. This shift to explicit spatial management presents new challenges and uncertainties for ecologists and resource managers. In particular, the potential for MPAs to change population sustainability, fishery yield, and ecosystem properties depends on the poorly understood consequences of three critical forms of connectivity over space: larval dispersal, juvenile and adult swimming, and movement of fishermen. Conventional fishery management describes the dynamics and current status of fish populations, with increasing recent emphasis on sustainability, often through reference points that reflect individual replacement. These compare lifetime egg production (LEP) to a critical replacement threshold (CRT) whose value is uncertain. Sustainability of spatially distributed populations also depends on individual replacement, but through all possible paths created by larval dispersal and LEP at each location. Model calculations of spatial replacement considering larval connectivity alone indicate sustainability and yield depend on species dispersal distance and the distribution of LEP created by species habitat distribution and fishing mortality. Adding MPAs creates areas with high LEP, increasing sustainability, but not necessarily yield. Generally, short distance dispersers will persist in almost all MPAs, while sustainability of long distance dispersers requires a specific density of MPAs along the coast. The value of that density also depends on the uncertain CRT, as well as fishing rate. MPAs can increase yield in areas with previously low LEP but for short distance dispersers, high yields will require many small MPAs. The paucity of information on larval dispersal distances, especially in cases with strong advection, renders these projections uncertain. Adding juvenile and adult movement to these calculations reduces LEP near the edges in MPAs, if movement is within a home-range, but more broadly over space if movement is diffusive. Adding movement of fishermen shifts effort on the basis of anticipated revenues and fishing costs, leading to lower LEP near ports, for example. Our evolving understanding of connectivity in spatial management could form the basis for a new, spatially oriented replacement reference point for sustainability, with associated new uncertainties.

Subtidal circulation over the northern California shelf

The circulation over the shelf and upper slope off northern California, between 38°N and 42°N, was observed using moored current and temperature recorders deployed as part of the Northern California Coastal Circulation Study (NCCCS), from March 1987 through October 1989. The results of this study provide a larger-scale view of the wind-driven circulation than that described through the 1981–1982 Coastal Ocean Dynamics Experiment (CODE), particularly with regard to alongshore and temporal variations. From an improved description of the frequency structure of wind and current, a very low frequency (VLF) signal is identified in the observed currents at frequencies below those typical of wind forcing. The majority of this VLF variance appears to be accounted for by persistent flow events associated with the presence of mesoscale circulation in the adjacent ocean. The longer duration of the NCCCS also allows an improved description of the seasonality of flow regimes off northern California. Three oceanic seasons are identified: an upwelling season (April-July), a relaxation season (August-November), and a storm season (December-March). Alongshore variations in the strength of upwelling, in the strength of the alongshore flow, both near-surface and undercurrent, and in water temperature not only are a function of latitude, as is the wind, but they also correspond to location relative to promontories, notably Cape Mendocino. Immediately south of Cape Mendocino, the near-surface flow exhibits an equatorward minimum and a temperature minimum, whereas the undercurrent exhibits a poleward maximum. Conversely, at the moorings immediately north of the cape, temperatures are a maximum and the undercurrent exhibits a minimum. The maximum in near-surface temperature relates to a minimum in upwelling; no significant correlation was found between local wind and current immediately north of Cape Mendocino. This upwelling minimum and the upwelling maximum south of the cape were also observed as persistent sea surface temperature patterns in satellite imagery.

Estuarine fronts: How important are they?

Estuarine fronts have a role in estuarine circulation, productivity, sediment dynamics, and water quality. Despite their importance, our understanding is rather modest. While some insight can be drawn from studies of oceanic fronts, the shorter time scales of estuarine fronts often exclude the role of Coriolis forcing and geostrophy from the hydrodynamics and exclude the prospect of physiological adaptation from ecosystem dynamics. Conservative processes and rapid nonconservative processes like aggregation, settling, and behavioral responses are expected to be most important at estuarine fronts. Although generally short-lived, the recurrent periodic nature of estuarine fronts does include longer time scales and some attention should be given to the importance of this recurrence to longer time scale ecological responses.

Nearshore larval retention in a region of strong upwelling and recruitment limitation

The ability of miniscule larvae to control their fate and replenish populations in dynamic marine environments has been a long-running topic of debate of central importance for managing resources and understanding the ecology and evolution of life in the sea. Larvae are considered to be highly susceptible to offshore transport in productive upwelling regions, thereby increasing dispersal, limiting onshore recruitment, and reducing the intensity of community interactions. We show that 45 species of nearshore crustaceans were not transported far offshore in a recruitment-limited region characterized by strong upwelling. To the contrary, 92% of these larvae remained within 6 km from shore in high densities throughout development along two transects sampled four times during the peak upwelling season. Larvae of most species remained nearshore by remaining below a shallow Ekman layer of seaward-flowing surface waters throughout development. Larvae of other species migrated farther offshore by occurring closer to the surface early in development. Postlarvae evidently returned to nearshore adult habitats either by descending to shoreward-flowing upwelled waters or rising to the sea surface where they can be transported shoreward by wind relaxation events or internal waves. Thus wind-driven offshore transport should not limit recruitment, even in strong upwelling regions, and larvae are more likely to recruit closer to natal populations than is widely believed. This study poses a new challenge to determine the true cause and extent of recruitment limitation for a more diverse array of species along upwelling coasts, and thus to further advance our understanding of the connectivity, dynamics, and structure of coastal populations.

The influence of spatially and temporally varying oceanographic conditions on meroplanktonic metapopulations

Abstract We synthesize the results of several modelling studies that address the influence of variability in larval transport and survival on the dynamics of marine metapopulations distributed along a coast. Two important benthic invertebrates in the California Current System (CCS), the Dungeness crab and the red sea urchin, are used as examples of the way in which physical oceanographic conditions can influence stability, synchrony and persistence of meroplanktonic metapopulations. We first explore population dynamics of subpopulations and metapopulations. Even without environmental forcing, isolated local subpopulations with density-dependence can vary on time scales roughly twice the generation time at high adult survival, shifting to annual time scales at low survivals. The high frequency behavior is not seen in models of the Dungeness crab, because of their high adult survival rates. Metapopulations with density-dependent recruitment and deterministic larval dispersal fluctuate in an asynchronous fashion. Along the coast, abundance varies on spatial scales which increase with dispersal distance. Coastwide, synchronous, random environmental variability tends to synchronize these metapopulations. Climate change could cause a long-term increase or decrease in mean larval survival, which in this model leads to greater synchrony or extinction respectively. Spatially managed metapopulations of red sea urchins go extinct when distances between harvest refugia become greater than the scale of larval dispersal. All assessments of population dynamics indicate that metapopulation behavior in general dependes critically on the temporal and spatial nature of larval dispersal, which is largely determined by physical oceanographic conditions. We therfore explore physical influences on larval dispersal patterns. Observed trends in temperature and salinity applied to laboratory-determined responses indicate that natural variability in temperature and salinity can lead to variability in larval development period on interannual (50%), intra-annual (20%) and latitudinal (200%) scales. Variability in development period significantly influences larval survival and, thus, net transport. Larval drifters that undertake diel vertical migration in a primitive equation model of coastal circulation (SPEM) demonstrate the importance of vertical migration in determining horizontal transport. Empirically derived estimates of the effects of wind forcing on larval transport of vertically migrating larvae (wind drift when near the surface and Ekman transport below the surface) match cross-shelf distributions in 4 years of existing larval data. We use a one-dimensional advection-diffusion model, which includes intra-annual timing of cross-shelf flows in the CCS, to explore the combined effects on settlement: (1) temperature- and salinity-dependent development and survival rates and (2) possible horizontal transport due to vertical migration of crab larvae. Natural variability in temperature, wind forcing, and the timing of the spring transition can cause the observed variability in recruitment. We conclude that understanding the dynamics of coastally distributed metapopulations in response to physically-induced variability in larval dispersal will be a critical step in assessing the effects of climate change on marine populations.

Observations of a pulsed buoyancy current downstream of Chesapeake Bay

A plume of low-salinity water was observed along the North Carolina coast 100 km south of the mouth of Chesapeake Bay during the Coastal Ocean Processes Pilot field program conducted from August through October 1994. The presence of the plume was episodic, occurring every 3 to 8 days. The timing was related to wind patterns, which influence both the delivery of estuarine discharge to the shelf and the plumes passage down the coast. When not affected by local winds, the low-salinity water was confined to within 7–9 km of the coast and was about 8 m deep. Downwelling winds narrow and deepen the plume, whereas upwelling winds cause it to thin and spread offshore, eventually detaching from the coast. The low-salinity plume propagated along the coast at speeds comparable to linear internal wave phase speed, except when strong downwelling wind conditions caused the plume to be in contact with the bottom. The observed propagation speeds are faster than those predicted by previous numerical modeling efforts. The plumes slowed with distance from the source, as mixing with ambient shelf water reduced the density contrast. This buoyancy source was balanced by an alongshore current with a southward velocity of 0.3 to 0.7 m s−1, bounded by a region of high horizontal velocity shear at the offshore salinity front. The currents observed in the nose of the plume are consistent with properties of an internal gravity current under rotation.

Tidal Intrusion Fronts

A tidal intrusion front forms as a dense seawater inflow plunges (subducts) beneath ambient estuarine water during flood tide. The associated foam lines and color changes have been observed on many smaller estuaries with constricted mouths. Internal hydraulic theory and laboratory experiments are reviewed and expressions are obtained for the position of plunging and the amount of associated mixing. The existence of a tidal intrusion front and its structure are discussed in terms of densimetric Froude numbers. These fronts are particularly important in smaller estuaries in which the intrusion process may dominate wind and tidal mixing and thus determine the overall stratification of the estuary. Three classes of three-dimensional plunging flow are identified and discussed. In particular, it is suggested that the peculiar, cursive V-shape plunge line is characteristic of strongly plunging flow.

Biological communities in San Francisco Bay track large-scale climate forcing over the North Pacific.

Long-term observations show that fish and plankton populations in the ocean fluctuate in synchrony with large-scale climate patterns, but similar evidence is lacking for estuaries because of shorter observational records. Marine fish and invertebrates have been sampled in San Francisco Bay since 1980 and exhibit large, unexplained population changes including record-high abundances of common species after 1999. Our analysis shows that populations of demersal fish, crabs and shrimp covary with the Pacific Decadal Oscillation (PDO) and North Pacific Gyre Oscillation (NPGO), both of which reversed signs in 1999. A time series model forced by the atmospheric driver of NPGO accounts for two-thirds of the variability in the first principal component of species abundances, and generalized linear models forced by PDO and NPGO account for most of the annual variability of individual species. We infer that synchronous shifts in climate patterns and community variability in San Francisco Bay are related to changes in oceanic wind forcing that modify coastal currents, upwelling intensity, surface temperature, and their influence on recruitment of marine species that utilize estuaries as nursery habitat. Ecological forecasts of estuarine responses to climate change must therefore consider how altered patterns of atmospheric forcing across ocean basins influence coastal oceanography as well as watershed hydrology.