Sediment distribution and transport across the continental shelf and slope under idealized wind forcing

Resuspension, transport, and deposition of sediments over the continental shelf and slope are complex processes and there is still a need to understand the underlying spatial and temporal dynamical scales. As a step towards this goal, a two-dimensional slice model (zero gradients in the alongshore direction) based on the primitive flow equations and a range of sediment classes has been developed. The circulation is forced from rest by upwelling or downwelling winds, which are spatially uniform. Results are presented for a range of wind speeds and sediment settling speeds. Upwelling flows carry fine sediments (low settling speeds) far offshore within the surface Ekman layer, and significant deposition eventually occurs beyond the shelf break. However, coarser sediments quickly settle out of the deeper onshore component of the circulation, which can lead to accumulation of bottom sediments within the coastal zone. Downwelling flows are more effective at transporting coarse sediments off the shelf. However, strong vertical mixing at the shelf break ensures that some material is also carried into the surface Ekman layer and returned onshore. The concentrations and settling fluxes of coarse sediments decrease offshore and increase with depth under both upwelling and downwelling conditions, consistent with trends observed in sediment trap data. However, finer sediments decrease with depth (upwelling) or reach a maximum around the depth of the shelf break (downwelling). It is shown that under uniform wind conditions, suspended sediment concentrations and settling fluxes decay offshore over a length scale of order τ_s/ρf|w_s|, where τ_s is the wind stress, ρ the water density, f the Coriolis parameter, and w_s is the sediment settling velocity. This scaling applies to both upwelling and downwelling conditions, provided offshore transport is dominated by wind-driven advection, rather than horizontal diffusion.     

Effect of atmospheric CO2 and solar activity on wind regime and water column stability in the major global upwelling areas

It is not clear whether global warming will favour or reduce global ocean phytoplankton productivity in coastal areas. Moreover, the relative contributions made by natural and/or anthropogenic factors to possible changes in phytoplankton productivity are not clear. As the relationship between primary production and alongshore wind forcing is well established for the Eastern Boundary Current (EBC) ecosystems, our aim is to determine whether the changes experienced over the last five decades (1958–2007) in atmospheric CO₂ and solar activity have been able to affect the wind regime and water column stability in the most biologically productive upwelling areas of California, Canary, Humboldt and Benguela. We approached the work by statistically studying the effect of solar activity and atmospheric CO₂ on surface alongshore wind stress and on water column stability. There was an increasing trend in wind stress and water column stability in all the upwelling areas over the period studied (with the single exception of stability in the California EBC system). The analysis of detrended series evidenced significant relationships between atmospheric CO₂ concentration and wind stress and water column stability in the coastal upwelling areas investigated. In addition, wind stress and stability data were found to be consistent, with negative linear relationships between wind stress and CO₂ in most of the sites in the Benguela, Canary and Humboldt regions associated, as expected, to positive relationships when water column stability is used as regressand. The results of the present study suggest that greenhouse gas forcing, independent of its well known general increasing trend, was able to decrease wind stress intensity and increase water column stability for the period 1958 to present in most of the sites of the four Eastern Boundary Ecosystems studied, with the one exception of the California region. Conversely, the impact of solar activity appeared to be quite low compared to the greenhouse gas forcing.     

The influence of mesoscale physical processes on the larval fish community in the Canaries CTZ, in summer

We have studied the relation between the hydrography, the composition and horizontal structure of the larval fish community, and the horizontal distribution patterns of larval fish abundances in an area characterised by strong mesoscale oceanographic activity, located between the Canary Islands and the African coast (the Canaries Coastal Transition Zone), during August 1999. Upwelling, upwelling filaments, cyclonic and anticyclonic eddies and island wakes are typical mesoscale features of the northwest African coast in summer. A single upwelling filament off Cabo Juby was joined in mid-August by a second that originated about 100 km to the north. The two filaments flowed together and merged 100 km offshore. The merged filament was partially entrained around a cyclonic eddy, trapped between the Canary Islands and the African coast, and interacted with cyclonic and anticyclonic eddies shed from Gran Canaria. Mesoscale oceanographic features strongly influenced the horizontal distributions of fish larvae. Eddies acted as a mechanism of concentration, while upwelling filaments were dispersive, transporting larvae from the African neritic zone into oceanic areas and towards the Canary archipelago. This transport was the major cause of the predominance of neritic larvae in the composition of the larval fish community of the area. The results also suggest: (1) that anchovy larvae are good indicators of the offshore displacement of upwelled water; (2) that the alternation between anchovy and sardine as species dominant in the larval fish community of the area during summer depends upon the water temperature in the African upwelling region, anchovy dominating at higher temperature; (3) that a coupling of anchovy and sardine spawning with the mesoscale oceanographic structure formed by the upwelling filaments and trapped eddy overcomes the negative effect that Ekman transport has on their populations.     

Poleward shift of the coastal upwelling region off the California coast

Yoshida (1967) pointed out that the coastal upwelling region may not coincide with the intense longshore wind region and shift poleward. In order to clarify this poleward shift from the existing data, the monthly mean distributions of the offshore Ekman transport and the coastal upwelling intensity are estimated along the California coast from U. S. Daily Weather Maps and from the CCOFI data in 1949, respectively. The results show that the center of the coastal upwelling region is generally shifted to the north from the position of the maximum offshore Ekman transport. The detailed discussions are given for the case of August 1949 when the shift is seen most clearly.     

Spatio‐temporal variability of zooplankton biomass and environmental control in the Northern Benguela Upwelling System: field investigations and model simulation

Spatial and temporal distribution patterns of zooplankton are highly variable in the Northern Benguela Upwelling System. We studied the distribution of zooplankton (size class ≥ 0.33 mm) and used field data from four cruises that took place between March 2008 and February 2011, as well as simulation results of a regional ecosystem model. Remotely sensed sea surface temperatures (SST) and surface chlorophyll concentrations were analysed to investigate environmental influences on zooplankton biomass. The Intense Benguela Upwelling Index showed a distinct seasonal signal throughout the years and the highest upwelling peaks in August/September. Even though surface chlorophyll concentrations were very variable throughout the year, the highest concentrations were always detected in September, following the upwelling of nutrient‐rich water. In field catches, zooplankton biomass concentration in the upper 200 m was highest above the outer shelf and shelf‐break in December 2010 and February 2011, i.e. 6 months after the upwelling peaks. In contrast, zooplankton biomass simulated by the model in the surface water was highest in September. In March/April, biomass maxima were typically measured in the field at intermediate water depths, but the vertical distribution was also affected by extensive oxygen minimum zones. The ecosystem model reproduced this vertical pattern. Although general trends were similar, simulation data of zooplankton standing stocks overestimated the field data by a factor of 3. In upwelling systems, food webs are generally considered to be short and dominated by large cells. However, our field data indicate more small‐sized zooplankton organisms above the shelf than offshore.     

Coastal upwelling in the California current system

Coastal upwelling in the California Current system has been the subject of large scale studies off California and Baja California, and of small scale studies off Oregon. Recent studies of the winds along the entire coast from 25°N to 50°N indicate that there are significant along-shore variations in the strength of coastal upwelling, which are reflected in the observed temperature distribution. Active upwelling appears to be restricted to a narrow coastal band (about 10–25 km wide) along the entire coast, but the region influenced by coastal upwelling may be much wider. Intensive observations of the upwelling zone during summer off Oregon show the presence of a southward coastal jet at the surface, a mean vertical shear, a poleward undercurrent along the bottom, and persistently sloping isopycnals over the continental shelf; most of the upwelling there occurs during relatively short periods (several days long) of upwelling-favorable winds. During the upwelling season off Oregon, the offshore Ekman transport is carried by the surface Ekman layer, and the onshore return flow occurs through a quasi-geostrophic interior. It is not known whether the structure and dynamics observed off Oregon are typical of the upwelling zone along the entire coast, though some of the same features have been observed off Baja California. Current and future research will eventually show whether the Oregon results are also applicable in the region of persistently strong upwelling-favorable winds off northern California, and in the region of complex bathymetry off central and southern California.     

Environmental influence on phytoplankton communities in the northern Benguela ecosystem

An investigation of surface phytoplankton communities was undertaken on the shelf of the northern Benguela upwelling ecosystem during austral autumn (May) and spring (September), along latitudinal transects at 20° S and 23° S, from 2 to 70 nautical miles offshore, as well as on a zigzag grid located between these transects. Microscopic identification of the phytoplankton and CHEMTAX analysis of pigment biomarkers were used to characterise the community composition. During May 2014, warmer, more-saline water with a shallower upper mixed layer corresponding to periods of less-intense offshore Ekman transport was encountered on the shelf. Satellite imagery indicated high phytoplankton biomass extending for a considerable distance from the coast, and CHEMTAX indicated diatoms as dominant at most of the stations (52–92%), although dinoflagellates were dominant at some inshore localities (57–74%). Species of Chaetoceros, Bacteriastrum and Cylindrotheca were the most abundant, with abundance of the Pseudo-nitzschia ‘seriata-group’ being particularly high at a number of stations. In September 2014, more-intense wind-forcing resulted in a deeper upper mixed layer and stronger upwelling of colder, less-saline water. Elevated phytoplankton biomass was confined close to the coast, where diatoms accounted for most of the population (54–87%), whereas small flagellates, such as prasinophytes, haptophytes and cryptophytes, as well as the cyanobacterium Synechococcus, dominated the communities (58–90%) farther from the coast. It is hypothesised that stronger upwelling and deeper vertical mixing in September of that year were not conducive for widespread diatom growth, and that small flagellates populated the water column by being entrained from offshore onto the shelf in the upwelled water that moved in towards the coast.     

Distribution and transport of organic matter along a filament-eddy system in the Canaries – NW Africa coastal transition zone region

The distribution and transport of chlorophyll a (Chla), particulate (POC) and dissolved (DOC) organic carbon, and the respiratory ETS activity of the microplankton community were studied along a filament-eddy system located in the transition zone between the NW Africa upwelling and Canary Islands waters. Two independent filaments (F1 and F2) stemming from the coastal jet, between Cape Juby and Cape Bojador, merged about 100 km offshore, turning southward and onshore forced by the circulation of a recurrent oceanic cyclonic eddy. In general, the coastal upwelling waters presented higher Chla, but lower POC, DOC and ETS activity than filament waters. However, differences in organic carbon distribution and respiratory activity were observed among stations from the two filaments. The bio-chemical fields were strongly influenced by a complex sub-mesoscale hydrography resulting from the interaction of cyclonic and anticyclonic island eddies with the filaments. The combined F1 + F2 filament system transported 97.1 kg s⁻¹ of excess (non-refractory) total organic carbon (e-TOC), a value comparable to other published estimates from upwelling filaments in the NE Atlantic. About 90% of e-TOC was exported as DOC, since eddy re-circulation precluded the offshore transport of POC. Assuming that the calculated transport of e-TOC is representative of the annual average, the yearly offshore transport (3.1 x 10⁹ kg C) would represent about 25% of the upwelling primary production of the region of study.     

Remotely sensed variability of temperature and chlorophyll in the southern Benguela: upwelling frequency and phytoplankton response

High-resolution (1km) satellite data from the NOAA AVHRR (Advanced Very High Resolution Radiometer) and OrbView-2 SeaWiFS (Sea-viewing Wide Field-of-view Sensor) are used to investigate the upper layer dynamics of the southern Benguela ecosystem in more detailed space and time scales than previously undertaken. A consistent time-series of daily sea surface temperature (SST) and chlorophyll a concentration images is generated for the period July 1998–June 2003, and a quantitative analysis undertaken. The variability in SST, upwelling and phytoplankton biomass is explored for selected biogeographic regions, with particular focus on intra-seasonal time scales. The location and emergence of upwelling cells are clearly identified along the length of the southern Benguela, being distinct on the narrow inner and the mid-continental shelves. Most notable is the rapidly pulsating nature of the upwelling, with intense warm/cold events clearly distinguished. The phytoplankton response to this physical forcing is described. Chlorophyll concentration on the inner shelf largely mirrors the pattern of SST variability, similarly dominated by event-scale processes. Over the mid-shelf, higher chlorophyll is observed throughout all seasons, although low biomass occurs during winter. The variability of the offshore extent of SST and chlorophyll is identified at locations of differing shelf width. Cooler upwelled water is confined primarily to the narrow inner-shelf, with event-scale pulses extending considerable distances offshore. Agulhas Current influences are readily observed, even on the Cape Peninsula inner-shelf. Chlorophyll concentrations vary considerably between the locations of differing shelf width. SST, upwelling and phytoplankton indices are derived for selected locations to quantify the intra-seasonal variations. The SST indices show marked temperature changes associated with rapid pulsation on the event scale. No strong seasonal signal is evident. In contrast, the upwelling indices display a strong seasonal signal, with most intense upwelling occurring in spring/summer in the south. The phytoplankton response to the seasonal upwelling index differs between the selected locations. This study concludes that, although low-resolution SST and chlorophyll data may be useful for investigating general patterns over large scales, higher resolution data are necessary to identify finer scale spatial and temporal variability, especially in the inshore coastal zones.     

Surface boundary-layer variability off Northern California, USA, during upwelling

A five-element mooring array is used to study surface boundary-layer transport over the Northern California shelf from May to August 2001. In this region, upwelling favorable winds increase in strength offshore, leading to a strong positive wind stress curl. We examine the cross-shelf variation in surface Ekman transport calculated from the wind stress and the actual surface boundary-layer transport estimated from oceanic observations. The two quantities are highly correlated with a regression slope near one. Both the Ekman transport and surface boundary layer transport imply curl-driven upwelling rates of about 3×10⁻⁴ m s⁻¹ between the 40 and 90 m isobaths (1.5 and 11.0 km from the coast, respectively) and curl-driven upwelling rates about 1.5×10⁻⁴m s⁻¹ between the 90 and 130 m isobaths (11.0 and 28.4 km from the coast, respectively). Thus curl-driven upwelling extends to at least 25 km from the coast. In contrast, upwelling driven by the adjustment to the coastal boundary condition occurs primarily inshore of the 40-m isobath. The upwelling rates implied by the differentiating the 40-m transport observations with the coastal boundary condition are up to 8×10⁻⁴ m s⁻¹. The estimated upwelling rates and the temperature–nitrate relationship imply curl-driven vertical nitrate flux divergences are about half of those driven by coastal boundary upwelling.     

Comparing trophic flows in the southern Benguela to those in other upwelling ecosystems

A balanced trophic flow model of the southern Benguela ecosystem is presented, averaging the period 1980–1989 and emphasizing upper trophic levels. The model is based largely on studies conducted within the framework of the Benguela Ecology Programme and updates the results of an expert workshop held in Cape Town in September 1989. Small pelagic fish other than anchovy Engraulis capensis and sardine Sardinops sagax, mainly round herring Etrumeus whiteheadi and mesopelagic fish, were important components of the food web in the southern Benguela. Severe balancing difficulties were encountered with respect to the semi-pelagic resources (hake Merluccius spp.) and demersal top predators (sharks), indicating the need for further research on the interaction of these groups with their ecosystem. The model is compared to other existing trophic flow models of ecosystems in major upwelling areas, i.e. the northern Humboldt Current (4–14°S), the California Current (28–42°N) and the southern Canary Current (l2–25°N), and to two independently constructed models of the northern Benguela ecosystem. These models are compared using network analysis routines of the ECOPATH software, focusing on the interactions between the five dominant fish species (anchovy, sardine, horse mackerel Trachurus trachurus capensis, chub mackerel Scomber japonicus and hake) that support important fisheries in all systems. The upwelling systems rank by size rather than species dominance. The ratio of catches and primary production differs between systems, partly because of differences in fishing regimes. Predation on the five dominant fish groups by other fish in the system was the most important cause of fish mortality in all models. Fishery catches are generally a larger cause of mortality for these groups than predation by mammals. The ecological cost of fishing appears to be comparatively low in the southern Benguela, because catches are low compared with the primary production, but also because the fishery is relatively low in the foodweb. However, in view of the very tight foodweb demonstrated in the model. it is likely that an increase in fishing pressure would cause severe trade-offs with respect to other components of the southern Benguela ecosystem.     

Ocean Color Satellite Imagery and Shipboard Measurements of Chlorophyll a and Suspended Particulate Matter Distribution in the East China Sea

Satellite-derived ocean color data of Coastal Zone Color Scanner (CZCS) on board the Nimbus-7 and Ocean Color and Temperature Scanner (OCTS) on board the Advanced Earth Observing Satellite (ADEOS) are jointly used with historical in situ data to examine seasonal and spatial distributions of chlorophyll a (Chl-a) and suspended particulate matter (SPM) concentrations in the East China Sea. Ocean color imagery showed that Chl-a concentrations on the continental shelf were higher than those of the Kuroshio area throughout the year. Satellite-derived Chl-a concentrations are generally in good accordance with historical in situ values during spring through autumn (although no shipboard in situ measurement was conducted at nearshore areas). In contrast, ocean color imagery in winter indicated high Chl-a concentrations (4–10 mg m^–3) on the continental shelf where bottom depth was less than 50 m when surface water was turbid (2–72 g m^–3 of SPM at surface), while historical in situ values were usually less than 1 mg m^–3. This suggests that resuspended bottom sediment due to wind-driven mixing and winter cooling is responsible for the noticeable overestimation of satellite-derived Chl-a concentrations. The algorithm for ocean color needs to be improved urgently for turbid water.     

Production characteristics of upwelling systems and the trophodynamic role of hake

The productivities of the Benguela, Peru and California Current upwelling systems and the trophodynamics of the associated hake stocks are compared. Within these large systems, upwelling rate, primary production and fish production are all positively correlated, as would be expected on trophodynamic grounds. Existing measurements suggest that the Benguela and the Peru Current systems are 2–4 times more productive than the California Current in terms of primary production and support 2–10 times more fish per square kilometre. The Benguela and Peru Current systems are 2–3 times more efficient in production of fish and harvestable yield to man as the California Current. The California Current appears to be less efficient primarily because a greater fraction of the annual primary production is transported offshore from the major upwelling sites by the prevailing winds and meandering structure of the currents. The strong seasonal variability in upwelling off California probably also results in poor trophodynamic coupling between the herbivores and phytoplankton. Consequently, less of the phytoplankton is consumed and, potentially, more can be exported out of the system. Hake (Merluccius spp.) are opportunistic, apex predators of euphausiids and pelagic and demersal fish in the three upwelling systems. The striking dominance of euphausiids in the diet of hake in the California Current probably reflects the lesser abundance of demersal fish in that highly advective region.     

Three-dimensional circulation in the NW Africa coastal transition zone

High-resolution data collected southeast of the Canary Islands during late winter 2006 are analyzed to describe the hydrography and three-dimensional circulation in the coastal transition zone off NW Africa. The data are optimally interpolated over a regular grid, the geostrophic velocity field is calculated and the Q-vector formulation of the omega equation is used to compute the quasi-geostrophic (QG) mesoscale vertical velocity. The coastal transition zone is divided into upwelling, frontal and offshore regions with distinct physical and dynamic characteristics. The upwelling region is characterized by cold and weakly stratified waters flowing towards the equator, with a poleward undercurrent of approximately 0.05 m s⁻¹ over the continental slope. The frontal region exhibits a southwestward baroclinic jet associated with cross-shore raising isopycnals; the jet transport is close to 1 Sv, with maximum velocities of 0.18 m s⁻¹ at surface decreasing to 0.05 m s⁻¹ at 300 db. Vertical sections across the frontal region show the presence of deep eddies probably generated by the topographic blocking of the islands to the southward current, as well as much shallower eddies that likely have arisen as instabilities of the baroclinic upwelling jet. The QG mesoscale vertical velocity field is patchy, estimated to range from −18 to 12 m day⁻¹, with the largest absolute values corresponding to an anticyclonic eddy located south of Fuerteventura Island. These values are significantly larger than estimates for other vertical velocities: diapycnal vertical velocities associated with mixing in the frontal region (a few meters per day), and wind-induced vertical velocities (non-linear Ekman pumping arising from the interaction between the wind stress and the background vorticity, maximum values of a few meters per day; linear Ekman pumping due to the divergence of Ekman transport, a fraction of a meter per day; or the coastal constraint in the upwelling region, about 0.7 m day⁻¹). However, the patchiness in both the QG mesoscale vertical velocity and the non-linear Ekman pumping velocity cause their integrated vertical transports to be one order of magnitude smaller than either coastal Ekman transport (0.08 Sv), integrated linear Ekman pumping (−0.05 Sv) or diapycnal transfer (about 0.1–0.2 Sv). The pattern of the near-surface fluorescence field is a good indicator of these different contributions, with large homogeneous values in the coastal upwelling region and a patchy structure associated with the offshore mesoscale structures.     

Effects of the Wind and Ice Conditions on the Upwelling along the Western Kamchatka Coast (Sea of Okhotsk) Based on the Satellite Data

The variability of the upwelling along the western coast of the Kamchatka Peninsula (northeastern part of the Sea of Okhotsk) has been studied based on an analysis of the multisensor satellite data. The intensity of upwelling is estimated on the basis of wind-forced offshore Ekman transport (upwelling index). The wind data for studying the seasonal variation of upwelling were collected in 1999–2009 using a Quik-SCAT/SeaWinds scatterometer. The upwelling events along the western Kamchatka coast were observed in December at the beginning of the winter monsoon period. During the period of strong winter monsoon northern winds from January to the middle of March, the drifting ice prevents the upwelling of the deep water at the western Kamchatka shelf edge under the mean conditions. The oceanographic data show that upwelling in the western coastal zone of Kamchatka was also observed during the transitional periods from winter to the summer monsoon (April). In summer, upwelling events are rarely observed in this region. The main cause of the summer upwelling is the propagation of the atmospheric cyclones over the Kamchatka Peninsula.     

Numerical investigation of the effects of upwelling on harmful algal blooms off the west Florida coast

The initiation of the toxic harmful algal bloom (HAB), Karenia brevis, along the west Florida coast has been associated with upwelling events. Upwelling processes may be responsible for the transport of nutrients or algae from deep offshore locations across the Florida shelf to the coast. The influence of coastal wind-driven upwelling on the onset and occurrences of K. brevis in this region was numerically investigated using Rutgers University's Regional Ocean Modeling System. Computations were carried out in an idealized model domain, a two-dimensional slice in the cross-shore and vertical directions. The surface forcing data used was from several offshore meteorological buoys. The motion of the algae was simulated using Lagrangian particles and a passive tracer. The numerical simulations of three K. brevis events in 2000–2002 showed that the particles respond (with upwelling/downwelling) to the along-shore wind stresses as expected and some upwelling was present during the events. Comparison of the passive tracer fields with measured fluorescence data exposed the model's sensitivity to the particular surface forcing data employed and the relatively more significant role played by surface forcing over initial conditions. The present model set-up constitutes a useful predictive tool for conditions conducive to the onset of HABs. It is planned to be used in a real-time mode to aid the NOAA HAB monitoring and forecasting system.     

Production regimes in four Eastern Boundary Current systems

High productivity (maxima 3 gCm⁻² day⁻¹) of the Eastern Boundary Currents (EBCs), i.e. the California, Peru-Humboldt, Canary and Benguela Currents, is driven by a combination of local forcing and large-scale circulation. The characteristics of the deep water brought to the surface by upwelling favorable winds depend on the large-scale circulation patterns. Here we use a new hydrographic and nutrient climatology together with satellite measurements of the wind vector, sea-surface temperature (SST), chlorophyll concentration, and primary production modeled from ocean color to quantify the meridional and seasonal patterns of upwelling dynamics and biological response. The unprecedented combination of data sets allows us to describe objectively the variability for small regions within each current and to characterize the governing factors for biological production. The temporal and spatial environmental variability was due in most regions to large-scale circulation, alone or in combination with offshore transport (local forcing). The observed meridional and seasonal patterns of biomass and primary production were most highly correlated to components representing large-scale circulation. The biomass sustained by a given nutrient concentration in the Atlantic EBCs was twice as large as that of the Pacific EBCs. This apparent greater efficiency may be due to availability of iron, physical retention, or differences in planktonic community structure.     

Comparison of assemblages and some life-history traits of seabirds in the Humboldt and Benguela systems

There are 21 and 15 species of seabirds that breed in the Humboldt and Benguela upwelling systems respectively. Only two species of gull are common to both systems, one as an endemic subspecies to the Benguela system. Eleven species and two subspecies are endemic (or nearly so) to the Humboldt system; seven species and one subspecies to the Benguela system. Each system has an endemic penguin, sulid, cormorant and tern that feed mainly on anchovy Engraulis spp., sardine Sardinops sagax or both these fish. The Peruvian pelican Pelecanus thagus also feeds primarily on these prey items. A plentiful availability of food has resulted in many of these seabirds attaining high levels of abundance. For the four pairs of species that feed on anchovy and sardine, those in the Humboldt system all have a biology that enables them to increase more rapidly than their Benguela counter-parts. This reflects the higher frequency of environmental perturbations that depress seabird populations in the Humboldt system. In addition, both systems have a small endemic cormorant that feeds near the coast and a small endemic tern that breeds in the adjacent mainland desert and feeds at the sea surface. Several seabirds endemic to a system have no obvious ecological equivalent in the other system: the pelican, a diving-petrel, four storm-petrels and a gull in the Humboldt system; a cormorant and a gull in the Benguela system. Some species with tropical or subantarctic affinities breed at the boundaries of the systems. Others breed also in freshwater systems. The grey gull Larus modestus, which feeds in the Humboldt system, breeds in montane deserts.