Influence of buoyancy and vertical distribution of sardine Sardinops sagax eggs and larvae on their transport in the northern Benguela ecosystem

In recent years, sardine Sardinops sagax spawning has been recorded inshore off central Namibia. Field observations on eggs and laboratory measurements show that spawning, demonstrated by the distribution of newly spawned eggs, takes place just below the upper mixed layer. The high positive buoyancy of the eggs causes them to ascend rapidly to the surface layer, where they are moved offshore by upwelling-induced offshore transport. However, increased wind-induced mixing also influences the vertical distribution of eggs, causing them to be partly mixed down below the layer moving offshore and into the layer moving inshore. This mechanism acts to retard the transport and offshore loss of eggs from the spawning areas. The vertical distribution of sardine larvae, with highest concentrations deeper than 20 m, indicates active movement out of the layer moving offshore, and this tendency seems to be more pronounced for older larvae. Hence, vertical migration of larvae is an additional factor mitigating their loss from nearshore. Taken together, these features seem to minimize the offshore loss of offspring, particularly in periods of low stock biomass when spawning close to the shore seems to be common.     

Currents along the Tsitsikamma coast,South Africa,and potential transport of squid paralarvae and ichthyoplankton

In July 1998, a bottom-mounted Acoustic Doppler Current Profiler was deployed at 36m depth in the centre of the Tsitsikamma National Park on the eastern Agulhas Bank, South Africa. The purpose was to investigate transport of chokka squid Loligo vulgaris reynaudii paralarvae hatched on the inshore spawning grounds (<60m) and ichthyoplankton spawned within the park. Analysis of the first 12 months of data (July 1998–June 1999) shows that surface flow was mainly eastward (alongshore), with a maximum velocity (u-component) of +115cm s^?1 and an average of +24cm s^?1. Generally, velocity decreased with depth, with a maximum bottom velocity (u-component) of +65cm s^?1 and an average of +10cm s^?1. Data from a nearby thermistor array show that the water column was usually isothermal during winter (July–September), with bottom flow in the same direction as the surface layer. In summer (December–March), vertical stratification was most intense, and surface and bottom flows differed in velocity and direction. Potential net monthly displacements calculated for three depths (5m, 23m and 31m) indicate that passive, neutrally buoyant biological material (e.g. squid paralarvae, fish eggs and larvae) would likely be transported eastwards in the surface layer for eight of the 12 months, and would generally exceed distances of 220km month–1. Displacement in the bottom layer was more evenly distributed between east and west, with net monthly (potential) transport typically 70–100km, but reaching a maximum of 200km. Wind-driven coastal upwelling, prevalent during the summer, causes the surface layer of the coastal counter-current to flow offshore for several days, resulting in potential displacement distances of 40km from the coast. These results suggest that squid paralarvae hatched on the inshore spawning grounds are not generally transported towards the 'cold ridge', a prominent semi-permanent oceanographic feature of cold, nutrient-rich upwelled water, where food is abundant, and that fish larvae, whether from the surface or bottom layer, are exported beyond the boundaries of the Tsitsikamma National Park.     

Direct hydroacoustic observations of chokka squid Loligo vulgaris reynaudii spawning activity in deep water

Unusual and distinct hydroacoustic targets were observed in November 1996, May 1998 and November 1998 during routine pelagic biomass surveys off the south coast of South Africa. During the November 1996 survey, seven such targets were observed near the bottom at depths of 115–125 m, directly south of the traditional inshore spawning grounds of chokka squid Loligo vulgaris reynaudii at Cape St Francis. The targets were close to prominent seabed ridges and extended 30–40 m off the bottom. In May 1998, three similar targets were observed at depths 55–80 m off Plettenberg Bay, another well-known squid spawning site. The shallowest target was identified, by means of a midwater trawl, as a mixture of mature male and female chokka. During the November 1998 survey, nine similar targets were again observed on the squid spawning grounds at Cape St Francis, also adjacent to seabed ridges. Drawing on fisheries hydroacoustic experience and knowledge of chokka squid spawning behaviour, the targets are believed to be aggregations of spawning squid.     

Horizontal spatial and temporal distribution patterns of nearshore larval fish assemblages at a temperate rocky shore

There have been no previous studies of the composition of nearshore larval fish assemblages along the coast of Portugal. We aimed to describe the composition and horizontal distribution patterns of larval fish assemblages and their temporal dynamics near a rocky reef at depths shallower than 13 m (inshore) and at two miles (3.70 km) from shore (offshore), as well as along transects perpendicular to the shoreline, from the reef to 10 miles offshore (18.52 km). Samples were taken using 5 min sub-surface trawls at the rocky shore of the Arrábida Marine Park (W Portugal). A total of 1021 larvae were collected, belonging to 61 taxa inshore and to 29 taxa offshore. Along transects, 626 larvae of 52 taxa were collected. Most larvae belonged to coastal species associated with rocky reefs. Total larval abundance and diversity were higher from May to July, which is consistent with the spawning activity of adults. Diversity and total larval abundance decreased significantly with increasing distance from shore, both in the inshore/offshore comparison and in the transects, where this decrease was evident at a very small spatial scale (within the first mile from the reef). Species assemblages differed in the pattern of distribution, with most species clearly associated to the extreme nearshore. The distribution patterns obtained were independent of the spawning mode of species. Results are discussed in the light of the possible physical mechanisms that can potentially act at the Arrábida Marine Park to facilitate larvae retention and the role of larval behaviour.     

Changes in potential North Sea spawning grounds of plaice (Pleuronectes platessa L.) based on early life stage connectivity to nursery habitats

We explored the hypothesis that spawning ground locations of North Sea plaice reflect the locations of nursery grounds using drift scenarios based on a baroclinic, shallow-water circulation model (HAMSOM). The transport of pelagic eggs and larvae was simulated each year from 1975 to 2006 using in situ forcing, temperature-dependent development and stage-specific behaviour of eggs and larvae. This long-term simulation period also allowed us to explore climate effects. A release position was considered a potential and suitable spawning site if larvae from that area reached coastal nurseries after the onset of metamorphosis. In general, larvae were transported in an anti-clockwise direction and settled in nurseries that were relatively close to the release positions. Spawning locations that were offshore were poorly connected to nursery grounds while those closer to the shore had higher connectivity. Simulated suitable spawning locations broadly agreed with the main centres of egg production (English Channel, Southern Bight, German Bight), except for the known spawning grounds south of Dogger Bank. Over the 31-year simulation period, positive and negative trends in transport success were found for the western and eastern parts of the North Sea, respectively. Changes in the west (Flamborough Head) were mainly due to changes in water circulation patterns whereas those in the east (northern German Bight) were induced by changes in both currents and water temperature. The implications of these findings, and the significant correlation between changes in drift and recruitment, suggest that climate-driven changes in the suitability of nursery grounds will directly affect the distribution and productivity of plaice in the North Sea.     

Spawning strategies and transport of early stages of the two Cape hake species,Merluccius paradoxus and M. capensis,in the southern Benguela upwelling system

Seasonal and short-term variability of environmental parameters influence the spawning strategies of fish species. In this study, the spawning strategies and the transport of early stages of the two Cape hake species off South Africa were investigated. Distribution of eggs and larvae of Merluccius paradoxus and M. capensis was analysed in order to derive more detailed and species-specific information on spawning season, spawning location, and transport of early stages. Samples were collected during three pilot surveys between January and October 2007 and during an extensive survey in September/October 2008 in the southern Benguela upwelling system off South Africa. Eggs and larvae of M. paradoxus were found in greater numbers than those of M. capensis during all surveys. Highest abundances were found from September to October, indicating one spawning peak for M. paradoxus during late austral winter to spring. The western Agulhas Bank was identified as the primary spawning ground, and smaller spawning events occurred on the West Coast. Larvae of both species were mainly distributed in subsurface waters between 25 and 100 m. More than 50% of all larvae caught had a total length between 3 and 4 mm and size increased significantly with decreasing latitude. Merluccius capensis were found closer inshore than M. paradoxus, indicating that early stages of the two species followed separate drift routes. We assume that this distribution pattern most likely evolved from differences in spawning location and phenology. The spawning strategies of M. paradoxus and M. capensis are well adapted to a time-frame of optimal transport conditions favourable for larval survival in the highly variable environment of the southern Benguela upwelling system, but the peak spawning of the two species is separated in time and space.     

Seasonal variation of the neustonic zooplankton community in southern waters of Korea

We investigated the seasonal variation of the neustonic zooplankton community in the surface layer (above 30 cm in depth) of 16 stations in southern waters of Korea from April 2002 to April 2003. In the neustonic community, copepods accounted for 59% of total organisms, as the most predominant group, while other dominant groups were invertebrate larvae, cladocerans, tunicates and fish eggs. Pontellid copepods, a major group of neustonic zooplankton, showed a marked seasonal variation and displayed a significant positive correlation with temperature (P < 0.05). Fish eggs appeared from spring to early summer in the inshore waters while tunicates and oceanic copepods (Eucalanus spp. and Euchaeta spp.) were dominant in the offshore waters during high temperature periods. This suggests that the distribution pattern of the neustonic zooplankton is seasonally affected by the hydrological conditions of in- and offshore waters. The abundance of zooplankton was higher in the surface layer (< 15 cm) than in the subsurface layer (15–30 cm), and was also higher at night than during the daytime. Of the dominant copepod families, the density of acartiid and pontellid copepods was higher in the surface layer than the subsurface layer. The density of acartiid and corycaeid copepods was significantly higher during daytime than nighttime. This indicates that the surface layer in temperate regions may provide a favorable habitat for neustonic copepods despite the dynamic environmental fluctuations.     

Dynamics of the pelagic component of Namibian hake stocks

A study was undertaken to investigate the spatial, diurnal and interannual dynamics of the pelagic component of Namibian hake stocks. Data were derived from trawl surveys conducted on board the R.V. Dr Fridtjof Nansen during the period 1991–1999. In addition to trawl surveys, acoustic methods were applied concurrently to quantify fish biomass off the bottom (above the headline). The pelagic component of hake was variable in both time and space – it was denser in the north and at depths of 200–500 m. Pelagic densities also correlated well with the density of adult fish (≥30 cm) at the bottom, but not with that of young hake (>18 and <30 cm) at the bottom. Pelagic densities were lowest at midday and highest at night, early morning and late afternoon densities being intermediate between the two. Incorporation of acoustic methods allows for at least partial account to be taken of diurnal variation in determining catchability. The fraction of the biomass off the bottom was not the same each year, thus affecting assessment and management advice for the stock, even if the survey results are treated only as relative indices of abundance. More field studies are needed to investigate the size range and species of hake active in vertical migration.     

Potential transport of plaice eggs and larvae between two apparently self-contained populations in the Irish Sea

A coupled physics particle-tracking model, driven by realistic meteorological forcing was used to examine the dispersal and transport of plaice eggs and larvae in the year 2000 from two spawning grounds in the Irish Sea. The model included passive transport of eggs and early stage larvae, diel vertical movements for larvae between 7 and 10.5 mm in body-length and tidally synchronised, vertical movements for larger larvae (>9 mm body-length). The year 2000 was chosen because of the availability of ichthyoplankton data with which to initialise the model. The majority of larvae originating from spawning in the eastern Irish Sea settled into nursery grounds along the Scottish, English and Welsh coasts, in agreement with previous findings. In contrast, a significant portion of larvae originating from spawning in the western Irish Sea was transported eastwards to these same nursery grounds. Transport across the Irish Sea resulted from the onset of tidally synchronised vertical behaviour encoded in the model for older larvae. Settlement of larvae into local nursery grounds along the Irish coast was limited. Because of the prevailing winds and currents in the region, plaice eggs and larvae are unlikely to be transported from east to west; in most years spawning in the western Irish Sea probably acts as an additional source of juveniles for nursery grounds along the Scottish, English and Welsh coasts.     

Early stages of Sardina pilchardus and environmental anomalies in the Northwestern Mediterranean

The influence of environmental conditions on the spatial distribution patterns of the early stages of Sardina pilchardus off the Catalan coast (Northwestern Mediterranean) during November 1998, February 1999 and November 1999 was investigated. As a general rule, eggs were found on the continental shelf, while larvae displayed a wider cross-shelf distribution. The local circulation during autumn–winter 1998–1999 was dominated by an anticyclonic eddy of new Atlantic Water that remained stationary at the centre of the Catalan Sea from September 1998 to the end of February 1999. Such a situation is very infrequent and the influence of these waters extended to the continental shelf zone, modifying the slope current by diverting the flow and even reversing the direction of the flow. On the third cruise, carried out at the beginning of the spawning season of 1999 (November), the circulation and water mass distribution in the region was typical, with the old Atlantic Water occupying the whole region. Abundance of pilchard eggs and larvae during this last cruise was higher than in the previous surveys. In November 1998, the abundance of pilchard eggs and larvae was relatively low in the zones affected by the new Atlantic Water. The most interesting result of these surveys was the very low concentrations of eggs and larvae found in February 1999, possibly due to the anomalous situation created by the eddy, which had remained almost stationary during the 1998–1999 spawning season.     

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.     

Dispersal patterns of the eggs and larvae of spring-spawning fish in the Irish Sea, UK

Many marine species produce pelagic propagules which, because of their life-history characteristics and the local hydrodynamics, can disperse considerable distances from the point of release. Distances travelled are affected by factors such as: release time and location, egg and larval stage duration, local environmental conditions and active swimming and settlement behaviours. Understanding such dispersal patterns is important for the design of effective ecosystem-conservation strategies. We used a regional scale, coupled physical-biological model for the Irish Sea to simulate the possible dispersal of eggs and larvae of five species of fish with contrasting early life histories (cod Gadus morhua, plaice Pleuronectes platessa, witch Glyptocephalus cynoglossus, sprat Sprattus sprattus and pogge Agonus cataphractus). The hydrodynamic model was forced with meteorological data for 1995, a year when extensive plankton surveys were conducted in the Irish Sea. A particle tracking method featuring particle release (spawning) and species-dependent particle development and behaviour was then run based on flow and temperature fields from the hydrodynamical model. Modelled larval distributions and settlement areas corresponded favourably with observations from field sampling. The settlement destinations (or onset of shoaling for sprat) were affected both by their initial spawning location and by the species-specific development rates and behaviours coded into the model. Eggs and larvae typically remained within 160 km of their spawning origin, although a minority travelled up to 300 km. Even in a relatively enclosed sea such as the Irish Sea, fish eggs and larvae can be dispersed over 100s of km. This provides a major challenge for the design of effective spatial management strategies if it is necessary to protect a species across its life-history stages. Further progress in the design of effective conservation measures for species or communities will need an integrated approach taking account of key aspects of early life history and behaviour.     

Patterns of variability in ichthyoplankton occurrence and abundance in Biscayne Bay,Florida

Within-day variability in ichthyoplankton and microzooplankton abundances was examined at a single station in Biscayne Bay using replicate tows of 61-cm bongo nets and Niskin bottles to determine if patchiness occurred on the 10–1000 m scale and on the minutes to hours time scale. Fish eggs and larvae often were patchy but copepod nauplii, the predominant food of larvae, usually were randomly distributed at the scales examined and over the 3.15 m-depth water column. Mean patchiness index values were of similar magnitude for fish eggs and larvae but fish eggs were patchy more often than were larvae. Individual taxa of larvae had extremely high patchiness index values on some dates. Variability in fish egg catches often reflected increasing or decreasing abundance trends during the 2.5h sampling period while fish larvae catches often appeared to be clumped within the repetitive series of tows. There was no tendency for patchiness to be correlated among taxa on collection dates nor was it correlated with abundances or wind speeds. Patchiness indices of bay anchovy Anchoa mitchilli eggs and larvae were not significantly correlated, indicating little concordance in tendency to be aggregated, suggesting that distributions were influenced by biological processes related to spawning of adults and behaviour of larvae, in addition to physical processes. Although ichthyoplankton patchiness often did exist at the 10–1000 m scale, on many days ichthyoplankton was uniformly or randomly distributed. Copepod nauplii were abundant ($\text{x}\text{=90.41}{}^{\mathrm{?1}}$), randomly distributed on most dates, and apparently readily available as fish larvae food in Biscayne Bay.     

Stochastic study of the temperature response of the upper ocean to uncertainties in the atmospheric forcing in an Atlantic OGCM

The impact of errors in atmospheric forcing on the behaviour of ocean models is a fundamental issue for ocean modellers and data assimilation and one that has yet to be fully addressed. In this study, we use a stochastic modelling approach with 50 7-months (September–March) primitive equation eddy permitting (1/4°) integrations. We investigate the response of the oceanic circulation to atmospheric uncertainties, focusing principally on their impact on the upper oceanic temperature field. The ensemble is generated by perturbing the wind, atmospheric temperature and incoming solar radiation of the ERA40 reanalysis. Each perturbation consists of a random combination of the 20 dominant EOFs of the difference between the ERA40 and NCEP/CORE reanalysis datasets. The ensemble standard deviation of various interfacial and oceanic quantities is then examined in the upper 200 m of three distinct regions of the North Atlantic: in the Gulf Stream, in the Northern Tropical band and in the North East Atlantic. These show that even a very small perturbation of the atmospheric variables can lead to significant changes in the ocean properties and that regions of oceanic mesoscale activity are the most sensitive. The ocean response is driven by vertical diffusivity and eddy activity. The role of subsurface currents is also crucial in carrying the eddy signal away from the regions of mesoscale activity. Finally, the decorrelation time scale of the mesoscale activity is critical in determining the amplitude of the oceanic response.     

Bathymetric influence on mean currents in two high-resolution near-global ocean models

The question of whether mean flow generation by eddies interacting with sloping bathymetry significantly influences World Ocean circulation is approached by examining output from two near-global circulation models, OFES and the LANL/NPS POP model, having ∼1/10° lateral resolution. In each of these vigorously eddying models, the mean currents over sloping bathymetry tend preferentially to align with the direction of topographic Rossby wave propagation, in accordance with theories of eddy-topographic interaction. This tendency, which is particularly strong near the ocean bottom and at abyssal depths, prevails both globally and within a variety of circulation regimes including the subpolar and subtropical gyres and the extra-equatorial tropics. By contrast, two coarser (∼1–2°), non-eddying models exhibit flow alignments throughout much of the abyssal ocean that are oppositely directed. This result suggests that eddies play an essential role in determining the direction of mean circulation over slopes, and that non-eddying models could benefit from a parameterization of this effect.     

Zooplankton distribution and cross-shelf transfer of carbon in an area of complex mesoscale circulation in the northern California Current

We conducted a research cruise in late summer (July–August) 2000 to study the effect of mesoscale circulation features on zooplankton distributions in the coastal upwelling ecosystem of the northern California Current. Our study area was in a region of complex coastline and bottom topography between Newport, Oregon (44.7°N), and Crescent City, California (41.9°N). Winds were generally strong and equatorward for >6 weeks prior to the cruise, resulting in the upwelling of cold, nutrient-rich water along the coast and an alongshore upwelling jet. In the northern part of the study area, the jet followed the bottom topography, creating a broad, retentive area nearshore over a submarine shelf bank (Heceta Bank, 44–44.4°N). In the south, a meander of the jet extended seaward off of Cape Blanco (42.8°N), resulting in the displacement of coastal water and the associated coastal taxa to >100 km off the continental shelf. Zooplankton biomass was high both over the submarine bank and offshore in the meander of the upwelling jet. We used velocities and standing stocks of plankton in the upper 100 m to estimate that 1×10⁶ m³ of water, containing an average zooplankton biomass of ～20 mg carbon m^?3, was transported seaward across the 2000-m isobath in the meandering jet each second. That flux equated to offshore transport of >900 metric tons of carbon each day, and 4–5×10⁴ tons over the 6–8 week lifetime of the circulation feature. Thus, mesoscale circulation can create disparate regions in which zooplankton populations are retained over the shelf and biomass can accumulate or, alternatively, in which high biomass is advected offshore to the oligotrophic deep sea.     

The Kuroshio Extension and its recirculation gyres

This paper reports on the strength and structure of the Kuroshio Extension and its recirculation gyres. In the time average, quasi-permanent recirculation gyres are found to the north and south of the Kuroshio Extension jet. The characteristics of these recirculations gyres are determined from the combined observations from the Kuroshio Extension System Study (KESS) field program (June 2004–June 2006) and include current meters, pressure and current recording inverted echo sounders, and subsurface floats. The position and strength of the recirculation gyres simulated by a high-resolution numerical model are found to be consistent with the observations. The circulation pattern that is revealed is of a complex system of multiple recirculation gyres that are embedded in the crests and troughs of the quasi-permanent meanders of the Kuroshio Extension. At the location of the KESS array, the Kuroshio Extension jet and its recirculation gyres transport of about 114 Sv. This represents a 2.7-fold increase in the transport of the current compared to the Kuroshio's transport at Cape Ashizuri before it separates from the coast and flows eastward into the open ocean. This enhancement in the current's transport comes from the development of the flanking recirculation gyres. Estimates from an array of inverted echo sounders and a high-resolution ocean general circulation model are of similar magnitude.     

The distribution of life cycle stages of two deep-water pleuronectids,Dover sole (Microstomus pacificus) and rex sole (Glyptocephalus zachirus), at the northern extent of their range in the Gulf of Alaska

Dover sole (Microstomus pacificus) and rex sole (Glyptocephalus zachirus) are both commercially valuable, long-lived pleuronectids that are distributed widely throughout the North Pacific. While their ecology and life cycle have been described for southern stocks, few investigations have focused on these species at higher latitudes. We synthesized historical research survey data among critical developmental stages to determine the distribution of life cycle stages for both species in the northern Gulf of Alaska (GOA). Bottom trawl survey data from 1953 to 2004 (25 519 trawls) were used to characterize adult distribution during the non-spawning and spawning seasons, ichthyoplankton data from 1972 to 2003 (10 776 tows) were used to determine the spatial and vertical distribution of eggs and larvae, and small-meshed shrimp trawl survey data from 1972 to 2004 (6536 trawls) were used to characterize areas utilized by immature stages. During the non-spawning season, adult Dover sole and rex sole were widely distributed from the inner shelf to outer slope. While both species concentrated on the continental slope to spawn, Dover sole spawning areas were more geographically specific than rex sole. Although spawned in deep water, eggs of both species were found in surface waters near spawning areas. Dover sole larvae did not appear to have an organized migration from offshore spawning grounds toward coastal nursery areas, and our data indicated facultative settling to their juvenile habitat in winter. Rex sole larvae progressively moved cross-shelf toward shore as they grew from April to September, and larvae presumably settled in coastal nursery areas in the autumn. In contrast with studies in the southern end of their range, we found no evidence in the GOA that Dover or rex sole have pelagic larval stages longer than nine months; however, more sampling for large larvae is needed in winter offshore of the continental shelf as well as sampling for newly settled larvae over the shelf to verify an abbreviated pelagic larval stage for both species at the northern end of their range.     

西南黄海夏季环流数值模拟研究

本文采用Regional Ocean Modeling System（ROMS）来研究西南黄海夏季环流,模拟的结果与浮标和锚定观测符合的很好。表层的海流流向东北,离岸一侧的流速大于近岸的流速,表层以下为约沿着25米等深线的气旋式流动。我们通过一系列过程导向实验研究海表风应力和底层温度锋面对环流的影响,结果表明,东南风驱动了表层东北向流动,底层温度锋面驱动了表层以下气旋式流动。底层温度锋面形成于近岸垂向混合充分水体和离岸层化水体之间。