Stratification and Tidal Current Effects on Larval Transport in the Eastern English Channel: Observations and 3D Modeling

We study how the combination of tides and freshwater buoyancy affects the marine organisms accumulation and horizontal transport in the ROFI system of the eastern English Channel. The Princeton Ocean Model coupled with a particle-tracking module is used to study the migration of fish eggs and larvae under different forcing conditions. Results of modeling are validated against observed concentrations of Flounder (Pleuronectes flesus) larvae. Numerical Lagrangian tracking experiments are performed with passive and active particles, representing sea-water organisms. Passive particles are neutrally buoyant whereas active particles are able to exercise light dependent vertical migrations equating to the swimming behavior of larvae. The experiments reveal that the strongest accumulation of particles occurs along the French coast on the margin of the ROFI. This happens because the interaction between the turbulence, the freshwater buoyancy input, and tidal dynamics, produces particle trapping and vertical spreading within the frontal convergence zone. Tides and freshwater input induce net alongshore horizontal transport toward the North. Tidal currents modulate the magnitude of horizontal transport whereas the fresh water input controls more the location of accumulation zones. Tracking experiments with active particles indicate that the vertical migration leads to a significant departure from the passive particle transport pattern. Differences lie in the shape of the particle transport pattern and the rate of the northward migration. In particular, vertically migrating particles travel slower. To find possible Flounder migration pathways, particles are released within the assumed spawning area of Flounder. The model predicts larvae drift routes and demonstrates that throughout the entire particle-tracking period the horizontal structure of the particle distribution is consistent with the larvae concentrations observed during the field experiments.     

The possible causes of diel vertical migrations of planktonic animals

Experimental measurements of the sinking velocity of anaesthetized planktonic animals carried out in the first half of this century prove that the downward migration of planktonic populations may be entirely attributed to passive sinking. This conclusion, along with data from voluminous literature devoted to circadian rhythms, supports the supposition that diel vertical migration is the result of an innate alternation of phases of high and low activity. During the phase of low locomotory activity, a passive sinking of an organism of negative buoyancy takes place; during the phase of high activity, the same organism actively climbs to the surface using the well-known mechanisms of space orientation. This hypothesis offers an explanation both of the usual course of diel migration and various deviations from this course, some of which are mentioned here. If the above considerations are correct, and if we admit that one of the trends of the evolution of pelagic organisms is directed towards the reduction of energy expenditure for the maintenance of the body in a suspended state, i.e. towards neutral buoyancy, the conclusion may be drawn that diel vertical migrations are not the result of adaptation to the planktonic mode of life.     

Vertical migration strategies with respect to advection and stratification in a semi-enclosed lough: a comparison of mero- and holozooplankton

Patterns of zooplankton vertical movement are often difficult to interpret because of multiple, complex and confounding environmental factors. Behavioural adaptations to these environmental variables are compared within and between the holo- and meroplankton constituents of a community. We used a nested design to analyse patterns at several scales in time; (semi-diel, diel, spring-neap tidal cycle and season) and two in space; (depth and site). To reduce complexity and aid interpretation we studied a semi-isolated community in a semi-enclosed, seasonally stratified sea lough (Lough Hyne Marine Nature Reserve, Ireland). In this, the main environmental gradient was water flow rate (or water residence time) caused by tidal currents. Vertical profiles of abundance showed that populations of the most abundant species of holo- and meroplankton in the lough have considerable behavioural plasticity, enabling them to switch between sedentary and migratory behaviour and patterns of migration. Some species migrate vertically in synchrony with diel cycles and others in response to semi-diel tidal currents; a few do both, but the majority did neither. It is suggested that water column structure and hydrographic discontinuities caused by flow rate and pycnocline dynamics are responsible for the variable patterns of vertical migration and distribution.Communicated by J.P. Thorpe, Port Erin     

Spring abundance and distribution of the ctenophore Pleurobrachia pileus in the Seine estuary: advective transport and diel vertical migration

Several sets of plankton and hyperbenthos samples were collected from the Seine estuary in late May-early June of 1991, 1992 and 1994, and the abundance, size-frequency, horizontal and vertical distribution, and short-term fluctuations of Pleurobrachia pileus (O.F. Müller, 1776) were determined. In the Seine estuary, P. pileus was more abundant than reported for any other European shallow waters. Its depth-averaged density was 33 individuals m^-3 in 1991 and 32 individuals m^-3 in 1992, with a maximum density of 808 individuals m^-3. The ctenophores displayed passive tidal advection and active diel vertical migration. In the outer estuary, high abundances were observed around low tide, and diel vertical migration was very pronounced. However, many individuals remained aggregated near the bottom at all times. In spring, the distribution of P. pileus coincided with the 33 and 150 S isohalines, with high abundances on the marine side of the estuary. It is suggested that aggregation of ctenophores near the bottom and diel vertical migration play an important role in the retention of P. pileus in the estuary. Two schemes of P. pileus life cycle in the Bay of Seine are proposed.     

Mechanisms to initiate a Heterosigma akashiwo red tide in Osaka bay

The ecological role of diel vertical migration of Heterosigma akashiwo Hada to initiate red tide was investigated in Tanigawa Fishing Port and Sano Harbor, Osaka Bay, Japan during red tide seasons in 1979 and 1980. This species migrated toward the surface early in the morning at a velocity of 1.0 to 1.3 m h^-1. Downward migration was found in the afternoon, and more than 2.0×10³ cells ml^-1 aggregated in the bottom layer at night. The upward migration started before sunrise and downward shifting occurred prior to sunset. The movement is presumably associated with circadian rhythm, which is known as one of the biological periodicities. H. akashiwo crossed steep temperature and salinity gradients (6.5°C and 5.7 S) during the diel vertical migration. High values of particulate organic carbon and nitrogen concentrations were obtained in dialysis bags suspended in situ at identical layers with high cell concentration, while the values for surface and bottom bags were comparatively low. The results reveal that H. akashiwo migrates toward the sea surface to carry out photosynthesis effectively, and to the bottom to utilize nutrients efficiently.     

Diurnal vertical migration of Solmissus albescens (Hydromedusae) in the Southern Adriatic

During a study on diurnal vertical migration in the hydromedusa Solmissus albescens (Gegenbaur, 1856), 112 net samples were taken from a depth of 1000 m to the surface. Sampling was performed in July, August, and September, 1969, at a station located 20 miles SSW of Dubrovnik (Yugoslavia). S. albescens migrates vertical distances of over 300 m. No significant influences on migration were noticed due to environmental factors (temperature, salinity, vertical water currents, etc.). The hydromedusa was able to approach surface layers. The recording obtained at various water-depth levels are considered in relation to illumination, mechanisms of evolution, and feeding of S. albescens.     

Langmuir circulations and plankton patchiness

A computer simulation model of oceanic Langmuir circulations illustrates the role of these vortical water movements as physical forcing mechanisms affecting the horizontal and vertical distributions of planktonic organisms. The diel migration of omnivorous and herbivorous zooplankton (approximately by literature values for the swimming speeds of Meganyctiphanes norvegica and Calanus finmarchicus, respectively) and the sinking of phytoplankton interact with the Langmuir cells, resulting in a spatially heterogeneous community. The patchy distributions, in turn, affect the prey—predator interactions in this simple food web. Situations involving threshold and non-threshold feeding strategies as well as constant and changing circulations are investigated. The simulations are restricted to twilight and nocturnal events.     

Biological and physical aspects of migration in the estuarine amphipod Gammarus zaddachi

The amphipod Gammarus zaddachi (Sexton) conducts extensive migrations along estuaries from near the limit of tidal influence in winter to more downstream reaches (where reproduction occurs) in spring. A return migration then takes place, primarily by juveniles, until the seaward areas are depopulated in winter. The present study was conducted between 1988 and 1990 in the Conwy Estuary, North Wales. This represents the first investigation on this species in a strongly tidal estuary, where the amphipods appear to migrate vertically into the water column on flood or ebb tides to control horizontal transport and to maintain preferred distributions. The timing of vertical migration seems to be largely controlled by an endogenous circatidal swimming rhythm. Phasing of peak activity relative to the time of expected high tide varies with season; upstream migrants in the autumn showed peak activity at the time of expected high tide, while in the spring at the time of downstream migration the rhythm was phase-delayed, with peak activity during the expected ebb tide. Together with the season, position along the estuary also affected the timing of peak endogenous activity; downstream migrants, originally active on the ebb tide and experimentally displaced seawards, showed a phase-advance of the rhythm relative to the time of high tide. Salinity-preference behaviour also varied between different developmental stages, with ovigerous females (downstream migrants) showing no preference between fresh and saline water, and juveniles (upstream migrants) showing a significant preference for freshwater. The interactions of endogenous rhythmicity and salinity-preference behaviour are discussed as controlling factors of migration in this species.     

Estimation of far-field horizontal and vertical turbulent diffusion coefficients from the concentration field of a wastewater plume near the Akashi Strait

We carried out a field study of the plume discharged by a near-shore wastewater outfall near the Akashi Strait, Japan. Using an Acoustic Doppler Current Profiler and a tow-body CTD, we measured the near-surface salinity and temperature fields in the region throughout an M₂ tidal cycle. We filtered the data in T–S space to remove water masses other than the wastewater, and then used the adiabatic mixing assumption to calculate the concentration of wastewater in the far field of this plume. Averaging the T–S fields of repeated surveys over a time period during which the tidal regime did not change substantially, allowed comparison of the time-averaged plume with the analytical solution for a plume diffusing in both the horizontal and vertical dimensions. The resulting vertical turbulent diffusion coefficients agreed well with those resulting from Thorpe scales determined via a vertically-profiling CTD, as well as with the canonical value for open channel flow of D _z = 0.067hu _*. The corresponding horizontal turbulent diffusion coefficients, however, were two orders of magnitude larger than those typically observed in straight channels, and an order of magnitude larger than those observed in meandering rivers. This is likely a result of enhanced horizontal mixing due to barotropic eddies generated by the interaction of strong tidal flow with headlands and levees, as well as due to the time-varying nature of tidal flow, and baroclinic spreading of the buoyant wastewater plume.     

Microbial activities as a function of water depth in the Ligurian Sea: an autoradiographic study

Fourteen species of sergestid shrimps were collected in the Sargasso Sea between the surface and 1500 m near Bermuda on 4 cruises. The vertical distribution and feeding activity of the most abundant species are discussed in relation to interspecific competition and the adaptive significance of vertical migration. Each species lives within a narrow depth range and exhibits a diel vertical migration. Sergestes splendens migrated as much as 825 m, while S. japonicus migrated less than 100 m. Neither the seasonal nor permanent thermocline influenced the migration range. The only species which occurred together both day and night were S. pectinatus with S. vigilax and S. pectinatus with S. sargassi. Morphological differences in the third maxillipeds of these species suggest differences in feeding. Although most species eat a variety of organisms, the foreguts of S. grandis, S. corniculum, and S. splendens contained euphausiids more often than those of other species, and S. grandis and S. robustus fed more frequently on fishes. In contrast, S. japonicus appears to feed on detritus. Food was found in the foreguts of most species less frequently during the day than night, but no species fed only at night. S. sargassi and S. pectinatus fed equally day and night.     

A model for the larval migration of the Japanese eel: roles of the trade winds and salinity front

A model that helps explain the mysterious long-distance migration of the Japanese eel (Anguilla japonica) is presented, based on oceanographic observations, satellite buoy drift experiments, and samplings of eel larvae taken in 1991. The trajectory of a 150 m depth buoy relased in the spawning area strongly suggests that A. japonica larvae spawned just south of the salinity front are transported westward by the North Equatorial Current (NEC). The larvae are then thought to be entrained into the Mindanao Current flowing southward along the Philippine Islands where A. japonica juveniles are scarcely distributed. These controversial results lead to the assumption that eel larvae are transferred from the NEC to the northward flowing Kuroshio, which distributes the eel larvae to the growth habitats of eastern Asia. In this eel larvae transfer model, a northward Ekman transport caused by trade winds plays an important role in explaining the wind-induced northward shift of the larvae together with the onset of diel vertical migration. Assuming that leptocephali greater than 20 mm initiate the vertical migration, a westward wind velocity greater than 5 to 10 m s^-1 should be high enough to diminish the southward current velocity. When the physical and geophysical conditions — such as the salinity front for spawning activity, the water tunnel for westward larval transport, the Ekman transport by the trade wind for transfer of the larvae from the NEC to the Kuroshio, and the strong velocity of the Kuroshio for rapid transport to growth habitats — are well matched with the timing of the onset of vertical migration, large-scale eel migration could result.     

Ontogenetic diel vertical migration of the planktonic copepod Calanus sinicus in the Inland Sea of Japan

The ontogenetic diel vertical migration of the planktonic copepod Calanus sinicus was investigated in the Inland Sea of Japan in June 1989, when the water column was thermally weakly stratified. Because of fewer eggs and less variation in their abundance, nocturnal spawning was not apparent. A pronounced upward migration occurred in NIII. NIII to CIII resided in the upper 20 m layer throughout the day, and from CIV on their median depths descended. CV and adult females underwent significant diel vertical migration, whereas adult males did not migrate. By integrating the results from the present study and those from our previous investigations (in August–September 1988, November 1988 and March 1989), we review seasonal variation in the ontogenetic diel vertical migration of C. sinicus. Spawning was largely nocturnal, reaching its maximum level around dawn, but spawning depth and fecundity changed seasonally. The distribution of pre-feeding stages, NI and NII, was similar to that of eggs. A pronounced upward migration always occurred in the first feeding stage, NIII, and late nauplii and early copepodites always resided in the food-rich upper layer, indicating that upward migration by NIII is feeding migration. As the stages progressed, they extended their vertical distribution range, and CV and adult females usually underwent diel vertical migration. However, the pattern and strength of this migration differed seasonally. Their day depths increased with the increase of relative biomass of planktivorous fish, indicating that predator avoidance induces their diurnal downward migration. High chlorophyll a concentrations in the upper layer (<15 m deep) relative to the lower layer (>20 m deep) amplified their diel vertical migrations. Diel vertical migration of C. sinicus is a phenotypic behavior.     

Vertical distribution and nocturnal migration of zooplankton in relation to the development of the seasonal thermocline in Patraikos Gulf

Vertical distribution and nocturnal migration of zooplankton species in relation to the development of the seasonal thermocline in the shallow waters (90 m) of Patraikos Gulf (Ionian Sea, Greece) were investigated using a WP-2 closing net. Juvenile and adult copepods accounted for a mean of 91% of the total collected in three sampling periods, i.e. May, July and September 1985.Ctenocalananus vanus, Paracalanus parvus andOithona plumifera were the dominant copepods. The majority of the zooplankton tend to aggregate at the thermocline layer. Among copepods the two congeneric speciesClausocalanus pergens andC. furcatus exhibited different migratory responses to the development of the thermocline.C. pergens occurred in the lower part of the thermocline andC. furcatus in the upper region or above. The diel vertical migration of all species could be divided into four types: (1) no vertical migration; (2) upward migration at night; (3) occasional migration; and (4) reverse migration (down at night). In July when the strongest thermocline developed, most zooplankters rose close to the surface at night. For most species, temperature discontinuity did not limit their diel migration.Please address all correspondence and requests for reprints to Dr J.J. Lykakis     

Endogenous swimming rhythms of blue crab, Callinectes sapidus , megalopae: effects of offshore and estuarine cues

Larvae of the blue crab Callinectes sapidus Rathbun develop on the continental shelf. The postlarval stage (megalopa) occurs near the surface and is transported shoreward by wind-driven surface currents. It then uses selective tidal stream transport for migration up an estuary. Endogenous swimming rhythms were measured under constant dark conditions in the laboratory in megalopae collected from the Newport River Estuary (North Carolina), the Delaware Bay, and offshore from the Newport River Estuary. Megalopae from all areas had a similar circadian activity rhythm, in which they swam during the time of the day phase in the field and were inactive at night. This rhythm predicts the presence of a reverse, diel, vertical-migration pattern offshore which would contribute to the location of megalopae near the surface during the day. The rhythm lacks obvious ecological significance in estuaries because it does not contribute to selective tidal stream transport and would increase vulnerability to visual predators during the day. Attempts to entrain a circatidal rhythm in swimming by cyclic and step changes in salinity were unsuccessful, as the circadian rhythm persisted. The rhythm also continued in the presence of the eelgrass Zostera marina, which is a site of settlement and metamorphosis in the field. Thus, megalopae enter estuaries with a solar day rhythm in activity. This rhythm, however, is not expressed, because light inhibits swimming during the day upon exposure to estuarine water. Since this light inhibition is removed in offshore waters, the rhythm would be expressed if, after entering an estuary, megalopae were transported back to offshore areas. Received: 19 December 1995 / Accepted: 2 August 1996     

Bioluminescence in the high Arctic during the polar night

This study examines the composition and activity of the planktonic community during the polar night in the high Arctic Kongsfjord, Svalbard. Our results are the first published evidence of bioluminescence among zooplankton during the Arctic polar night. The observations were collected by a bathyphotometer detecting bioluminescence, integrated into an autonomous underwater vehicle, to determine the concentration and intensity of bioluminescent flashes as a function of time of day and depth. To further understand community dynamics and composition, plankton nets were used to collect organisms passing through the bathyphotometer along with traditional vertical net tows. Additionally, using a moored bathyphotometer closed to the sampling site, the bioluminescence potential itself was shown not to have a diurnal or circadian rhythm. Rather, our results provide evidence for a diel vertical migration of bioluminescent zooplankton that does not correspond to any externally detectable changes in illumination.     

Vertical swimming in wave-induced currents as a control mechanism of intertidal migration by a sand-beach isopod

Both the vertical and horizontal distributions ofEurydice pulchra (Leach) within the water column of the surf zone were recorded throughout complete tidal cycles on a sandy beach in North Wales during the summer of 1989. Upon emerging from the sand, the isopods tended to swim up in the water column, where transport onshore would be facilitated by the wave-induced, onshore currents which laboratory wave-tank experiments have confirmed to occur near the water surface. This combination of active and passive transport to the water's edge results in high numbers of individuals in the narrow swash zone. At and just after the time of high tide, individuals swim to the water/sediment interface where, as again confirmed by wave-tank experiments, the predominant water movement is offshore. Continued swimming near the bottom during the ebb tide before reburrowing in the sand ensures transport downshore and avoidance of stranding above the characteristic level of zonation on the shore. Vertical migrations ofE. pulchra in the water column permit differential exploitation of up and downshore currents to achieve horizontal migration.     

Water column use and forage strategies of female southern elephant seals from Marion Island

The at-sea behaviour of marine top predators provides valuable insights into the distribution of prey species and strategies used by predators to exploit patchily distributed resources. We describe the water column usage and dive strategies of female southern elephant seals from Marion Island tracked between 2004 and 2008. Dives representing increases in forage effort were identified using a method that combines dive type analyses and the calculation of relative amounts of time that animals spend in the bottom phases of dives. Results from this analysis indicate that female elephant seals from Marion Island tend to display lower levels of forage effort closer to the island and display intensive opportunistic forage bouts that occur at a minimum distance of approximately 215 km from the island. Females from Marion Island dived deeper and for longer periods of time, compared to females from other populations. Most animals displayed positive diel vertical migration, evidently foraging pelagically on vertically migrating prey. A few animals displayed periods of reverse (negative) diel vertical migration, however, diving to deeper depths at night, compared to daytime. This behaviour is difficult to explain and prey species targeted during such periods unknown. Our results illustrate plasticity in foraging behaviour of southern elephant seals, as well as inter-population differences in forage strategies.     

Evidence of in situ diel vertical migration of a red-tide microplankton species in Ría de Vigo (NW Spain)

The migration capacity of red-tide species in the natural environment was studied at a station in the Ría de Vigo (Rías Bajas, NW Spain) over a 24 h period in September 1991. The Ría de Vigo, where red tides are frequent, normally shows a positive estuarine circulation and is subjected to seasonal upwelling and downwelling phenomena. A marked diel pattern was observed for five species that are capable of causing red tides (Ceratium furca, Scrippsiella trochoidea, Dinophysis acuminata, Mesodinium rubrum, and Eutreptiella sp.). Such diel behaviour could be clearly advantageous in a stratified environment where light and nutrients are often in two separate layers. Active movement enables species such as dinoflagellates and some ciliates to exploit high levels of irradiance at the surface during the day and to take up nutrients in deeper layers at night. Patchy distribution of phytoplankton, shear-induced horizontal dispersion and density variations were considered, but none of them accounted for the vertical changes observed. Vertical migration is thought to be one of the mechanisms that could promote blooms in nutrient-depleted surface layers.     

Effects of nitrate on the diurnal vertical migration,carbon to nitrogen ratio,and the photosynthetic capacity of the dinoflagellate Gymnodinium splendens

A non-thecate dinoflagellate, Gymnodinium splendens, was studied in a 12 d laboratory experiment in 2.0x0.25 m containers in which light, temperature, and nutrients could be manipulated. Under a 12 h light: 12 h dark cycle, the dinoflagellates exhibited diurnal vertical migrations, swimming downward before the dark period began and upward before the end of the dark period. This vertical migration probably involved geotaxis and a diel rhythm, as well as light-mediated behavior. The vertical distribution of nitrate affected the behavior and physiology of the dinoflagellate. When nitrate was present throughout the container, the organisms resembled those in exponential batch culture both in C:N ratios and photosynthetic capacity (P_max); moreover, they migrated to the surface during the day. In contrast, when nitrate was depleted, C:N ratios increased, P_max decreased, and the organisms formed a subsurface layer at a depth corresponding to the light level at which photosynthesis saturated. When nitrate was present only at the bottom of the tank, C:N ratios of the population decreased until similar to those of nutrient-saturated cells and P_max increased; however, the dinoflagellates behaved the same as nutrient-depleted cells, forming a subsurface layer during the light period. Field measurements revealed a migratory subsurface chlorophyll maximum layer dominated by G. splendens. It was just above the nitracline during the day, and in the nitracline during the night, which concurs with our laboratory observations.     

互花米草入侵盐沼湿地CH4和N2O排放日变化特征研究

甲烷（CH4）和氧化亚氮（N2O）是导致全球气候变暖的2种重要温室气体,探索其源汇及地域排放特征一直是全球变化研究领域的核心内容。CH4和N2O通量的日变化研究是正确估算大时间尺度下CH4和N2O排放量的基础。利用静态箱法原位观测了江苏沿海芦苇（Phragmites australis）、盐蒿（Suada salsa）、光滩、水面以及互花米草（Spartina alterniflora）入侵湿地CH4和N2O排放的日变化特征。结果表明,1）互花米草湿地地上部生物量为1.70 kg·m^-2,土壤有机碳质量分数为13.55 g·kg^-1;分别是芦苇和盐蒿湿地的2.50-3.43和2.15-4.15倍。2）互花米草和芦苇湿地土壤10 cm处氧化还原电位（Eh）有明显日变化,最低值出现在3：00,最高值出现在12：00;光滩和盐蒿湿地没有明显的日变化。3）互花米草湿地 CH4日平均排放通量为0.52 mg·m^-2·h^-1,是其他湿地的2.12-6.40倍;N2O日平均通量为-3.24μg·m^-2·h^-1,显著低于盐蒿湿地、光滩和水面（P＜0.05）。互花米草和芦苇湿地CH4排放通量最高值（0.73 mg·m^-2·h^-1和0.30 mg·m^-2·h^-1）出现在15：00,最低值（0.37 mg·m^-2·h^-1和0.17 mg·m^-2·h^-1）出现在3：00,均与土壤孔隙水中CH4浓度呈显著负相关（P＜0.05）。互花米草湿地CH4排放通量与10 cm土温、Eh和生态系统CO2净交换量（NEE）显著正相关（P＜0.05）。互花米草和芦苇湿地N2O通量9：00-18：00为负值,21：00—6：00为正值,均与NEE呈显著负相关（P＜0.05）。盐蒿湿地、光滩和水面CH4和N2O排放通量没有明显日变化特征。互花米草入侵提高了沿海湿地CH4排放,但降低了N2O排放,植物对CH4传输作用以及向根际传输O2和易分解有机物是导致互花米草和芦苇湿地CH4和N2O排放表现出日变化特征的原因。