A mechanism for horizontal zooplankton transport by vertical migration in tidal currents

A mechanism is proposed by which net horizontal transport can be induced in an organism which migrates vertically in a purely oscillatory, vertically sheared tidal current. The mechanism contains two elements. First, for all reasonable tidal current profiles, net horizontal transport is induced in any organism which migrates vertically with a period which is an exact multiple of the tidal period. (This is the basis for the widelyknown process of selective tidal stream transport where the migration period and the tidal period are exactly equal.) The second element in the new mechanism is the observation that diel migration, the most common form of vertical migration, has a period of 24 h and is therefore an exact multiple of the principal solar semi-diurnal tidal constituent (S2) which has a period of 12 h. This relation between the S2 and diel migration periods stems from the fact that both phenomena are independently locked to the solar cycle. Diel vertical migration can therefore interact with the S2 tidal current constituent to induce longterm horizontal transport in migrating organisms. For reasonable tidal current and vertical migration parameters it is found that horizontal transport rates of 4 km d^–1 are possible. The direction of net transport depends upon the phase of the S2 tidal current relative to local noon. The hypothesis is therefore proposed that geographical variation in S2 phase is a factor responsible for creating regions which are either retention-favorable or otherwise for diel-migrating marine organisms.     

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.     

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.     

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.     

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

The ontogenetic diel vertical migration of the planktonic copepodCalanus sinicus was investigated in the Inland Sea of Japan in summer 1988, when the water was thermally stratified with a thermocline of ca 5 °C between 35 and 45 m. Stage-specific differences in the diel vertical migration behavior ofC. sinicus were found. Eggs were spawned primarily within the surface-waters between midnight and dawn by ascending females, and sank gradually to deeper waters until they hatched into nauplii. Non-feeding nauplius stages (NI and II) were distributed throughout the water column, but the first feeding stage (NIII) performed an ontogenetic upward migration. NIV to VI and copepodite (C) stages I to III continuously aggregated in the phytoplankton-rich euphotic layer. However, the depth of the median CI to III populations descended as stage progressed. The onset of prominent diel vertical migration took place in CIV, and the amplitude of vertical migration increased with age, being maximal in adult females (CVI). Adult males (CVI), however, remained in the layer below 20 m, and did not migrate dielly. The ecological significance of ontogenetic diel vertical migration is discussed.     

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.     

Measuring the effects of thermoclines on the vertical migration of larvae of Callinectes sapidus (Brachyura: Portunidae) in the laboratory

An experimental system is described that is capable of maintaining precise and stable thermoclines of specified magnitude for laboratory study of behavior of planktonic organisms. Behavioral responses to encounters with thermoclines of various magnitudes were measured in stage I larvae of the portunid crab Callinectes sapidus Rathbun. Larvae adapted to an initial temperature of 15°C were able to swim up through a temperature gradient of 10°C, but showed increasing inhibition of upward migration when t=12.5°C. Larvae adapted to 17.5°C were unable to penetrate t=10°C. Thermoclines of 5° or 10°C slowed passive sinking rates. Results suggest that larvae avoid an upper temperature of 27.5°C by a cessation of swimming, accompanied by passive sinking. Thermal gradients of the magnitude present in the natural habitat are unlikely to alter larval vertical migration that is regulated by other exogenous stimuli. Naturally occurring thermoclines should seldom influence dispersal characteristics of C. sapidus larvae.Contribution No. 1535 of the Center for Environmental and Estuarine Studies, University of Maryland     

Contrasting diel patterns in vertical movement and locomotor activity of whale sharks at Ningaloo Reef

Activity patterns of animals often relate to environmental variables such as food availability and predation pressure. Technological advances are providing us with new tools to monitor and better understand these activity patterns. We used animal-attached data loggers recording acceleration and depth to compare activity patterns and vertical habitat use of whale sharks (Rhincodon typus) at Ningaloo Reef, Western Australia. Whale sharks showed a moderate reverse diel vertical migration but exhibited a clear crepuscular pattern in locomotory activity. Peak activity occurred at sunset, whereas vertical movement peaked prior to this. Typical ram surface filter feeding could be identified and occurred primarily during sunset and the first hours of night. At such times, direct observations indicated whale sharks were feeding on tropical krill swarms. Kinematic analysis of postural data and data from vertical movement suggests that whale sharks at Ningaloo spend ~8 min per day actively ram surface filter feeding. Considering the high biomass present in krill schools, it is estimated that whale sharks at Ningaloo have a similar energy intake as those at other aggregation sites. Diel patterns in activity and diving behaviour suggest that whale sharks have tuned their diving behaviour in anticipation of the formation of these high-density patches which appear to only be periodically, but predictably available at sunset. Our results confirm that diel patterns in vertical habitat selection and vertical movements do not necessarily reflect patterns in activity and foraging behaviour. Direct quantification of activity and behaviour is required in gaining accurate representation of diel activity patterns.     

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.     

Feeding rates and diel vertical migration of copepods near South Georgia: comparison of shelf and oceanic sites

Seventeen Longhurst Hardy Plankton Recorder profiles were taken over a diel cycle in January 1990 to study the feeding of four major copepods over the South Georgia shelf. Ontogenetic changes in vertical migration were followed and feeding cycles determined by gut fluorometry for Calanoides acutus Stage CV, Calanus sinillimus CV and CVI, C. propinquus CV and Rhincalanus gigas CV and CVI. In common with a neighbouring oceanic site visited two weeks later and reported elsewhere, all four species had a diel cycle of feeding and migration. The vertical distributions of C. simillimus (all stages), R. gigas (nauplii) and Euphausia frigida (postlarvae) were similar at both sites, the night being spent within the chlorophyll maximum at 15 to 30 m. However, the biomass dominants, C. acutus and R. gigas, dwelt below the chlorophyll maximum, about 30 m deeper than their oceanic counterparts. Unlike the oceanic site, feeding at the shelf site was not restricted to darkness, but increased 6 to 10 h before nightfall and finished at dawn; the intervening period coincided with sinking and digestion. Daylight feeding may have been induced by the shorter night, lower light levels or greater food requirements at the shelf site, despite planktonic predators being over three times more abundant. Daily ration estimates for R. gigas at both sites were only 2% body carbon per day. These low values contrast with its smaller competirors, whose rations were in the range 5.6 to 27%.     

The influence of hydrodynamic processes on zooplankton transport and distributions in the North Western Mediterranean: Estimates from a Lagrangian model

A Lagrangian module has been developed and coupled with the 3D circulation model Symphonie to study the influence of hydrodynamic processes on zooplankton transport and distributions in the North Western Mediterranean (NWM). Individuals are released every 3 days from March to August 2001 in two initial areas: around the DYFAMED sampling station in the central Ligurian Sea and in the Rhône river plume. Then the individuals are tracked for 40 days either as passive particles or with a simple diel vertical migration (DVM) pattern. The simulations suggest strong seasonal patterns in the distributions of the individuals released around the DYFAMED sampling station. Individuals spread all over the NWM basin after 40 days but different patterns occur depending on the season, the initial depths of release and the capacity of DVM. Offshore-shelf transport only occurs in April and May with particles ending up in the Gulf of Lions (GoL) in low concentrations. In other months, the Northern Current (NC) can be considered as a barrier for particles entering the GoL from the offshore sea. A quarter to a half of passive individuals released in the Rhône river plume remain in the GoL. The rest is transported by the NC towards the Catalan Sea. Applying a simple DVM scheme does not increase the retention of particles on the shelf.     

Sinking rate response to depletion of nitrate,phosphate and silicate in four marine diatoms

The effect of nutrient (N, P, and Si) depletion on sinking rates was studied for two small (Skeletonema costatum (Grev.) Cleve and Chaetoceros gracile Schütt) and two large [Ditylum brightwellii (West) Grun and Coscinodiscus wailesii (Gran et Angst)] centric diatoms obtained from stock cultures. Each diatom was examined under conditions of (1) nutrient repletion (=log growth phase), (2) nutrient depletion (48 h without a given substrate), and (3) recovery (24h after addition of limiting substrate to nutrient-deplete populations). All nutrient-replete cultures displayed low sinking rates despite large differences in cell size. In nutrient-deplete populations, sinking rate was related to the kind of nutrient depleted and varied among species. Silicate depletion elicited by far the greatest increase in sinking rates in all 4 species, indicating that biochemical aspects of silicon metabolism are more important to buoyancy regulation than density-related variations in the amount of silicon per cell. Since N- and P-depletion caused lower sinking rates in 3 of the species, this observation calls for re-evaluation of the axiom that nutrient depletion necessarily causes increased sinking rates. The exception was Coscinodiscus wailesii, which sank faster under all types of nutrient limitation. In most cases, sinking rates typical of log-phase cultures were not regained within 24 h after the addition of limiting nutrient to nutrient-depleted populations. Ultimately, the length of the recovery period may be useful in identifying the metabolic processes responsible for buoyancy regulation in actively growing cells.     

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.     

Diel vertical migration bySalpa aspera and its potential for large-scale particulate organic matter transport to the deep-sea

In mid-summer 1975 throughout the Western Slope Water of the North Atlantic Ocean, massive numbers ofSalpa aspera performed a diel vertical migration of at least 800 m. This resulted in a movement of 85 to 90% of the total zooplankton biomass out of the upper 500 m during the day. Fecal pellet production and losses from this salp population were estimated to contribute approximately 12 mg C m^-2 day^-1 to the deep planktonic and benthic populations. If all this organic matter reached the deep-sea floor, it would represent over 100% of the daily deep-sea benthic infauna energy requirements.     

Buoyancy of natural populations of marine phytoplankton

Buoyancy of natural populations of marine phytoplankton was studied in a fjord in western Norway during the diatom bloom and in autumn. The study was carried out under approximate in situ conditions by means of an apparatus described in the paper. During the spring bloom, positive buoyancy was observed only once. Sinking rates of individual fractions ranged from 0 to more than 9 m day^-1, and the mean sinking rates of the total chlorophyll content from 0 to at least 2. 2 m day^-1. The highest rates occurred in the post-bloom period, while sinking appeared negligible from the onset of the bloom up to its culmination. In autumn, the population was dominated by small, flagellated cells. Positive buoyancy, or upward migration, was then observed in two out of three experiments.     

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 November 1988 and March 1989, when the water temperature was weakly stratified in a reversed manner. In both investigations a pronounced ontogenetic difference in vertical distribution was found. Spawning always occurred during nighttime, being confined to the upper 40 m water column in November but to the layer below 35 m in March. The distribution of pre-feeding nauplius stages, NI and NII, was more or less similar to that of the eggs. The first-feeding NIII performed a marked upward migration, and late nauplius stages (NIV to NVI) and early copepodite stages (CI and CII) continuously aggregated in the upper water column where phytoplankton was abundant. CIII to CVI (adult female and male) tended to disperse in the whole water column. In November, however, they avoided the upper 10 m strate during daytime and some individuals migrated upward to the surface during nighttime. In March, CV and CVI aggregated in the layer between 5 and 15 m deep in the daytime and migrated both upward and downward at dusk, resulting in homogeneous distributions during the nighttime.     

The effects of seston food quality on planktonic food web patterns

In planktonic food webs, the conversion rate of plant material to herbivore biomass is determined by a variety of factors such as seston biochemical/elemental composition, phytoplankton cell morphology, and colony architecture. Despite the overwhelming heterogeneity characterizing the plant–animal interface, plankton population models usually misrepresent the food quality constraints imposed on zooplankton growth. In this study, we reformulate the zooplankton grazing term to include seston food quality effects on zooplankton assimilation efficiency and examine its ramifications on system stability. Using different phytoplankton parameterizations with regards to growth strategies, light requirements, sinking rates, and food quality, we examined the dynamics induced in planktonic systems under varying zooplankton mortality/fish predation, light conditions, nutrient availability, and detritus food quality levels. In general, our analysis suggests that high food quality tends to stabilize the planktonic systems, whereas unforced oscillations (limit cycles) emerge with lower seston food quality. For a given phytoplankton specification and resource availability, the amplitude of the plankton oscillations is primarily modulated from zooplankton mortality and secondarily from the nutritional quality of the alternative food source (i.e., detritus). When the phytoplankton community is parameterized as a cyanobacterium-like species, conditions of high nutrient availability combined with high zooplankton mortality led to phytoplankton biomass accumulation, whereas a diatom-like parameterization resulted in relatively low phytoplankton to zooplankton biomass ratios highlighting the notion that high phytoplankton food quality allows the zooplankton community to sustain relatively high biomass and to suppress phytoplankton biomass to low levels. During nutrient and light enrichment conditions, both phytoplankton and detritus food quality determine the extent of the limit cycle region, whereas high algal food quality increases system resilience by shifting the oscillatory region towards lower light attenuation levels. Detritus food quality seems to regulate the amplitude of the dynamic oscillations following enrichment, when algal food quality is low. These results highlight the profitability of the alternative food sources for the grazer as an important predictor for the dynamic behavior of primary producer–grazer interactions in nature.     

Behavioral responses of larvae of the crab Rhithropanopeus harrisii (Brachyura: Xanthidae) during diel vertical migration

Larval Rhithropanopeus harrisii (Gould) show nocturnal vertical migration. Larval behavioral responses to different rates of increase and decrease in light intensity were measured in an apparatus with a natural angular light distribution. A central objective was to establish whether phototaxis actually participates in vertical migration. At sunset the level of light adaptation controlled the readiness of the larvae to migrate, while an ascent was initiated by a preductable relative decrease in intensity (e.g. 4.0x10^-3s^-1). Rates of relative decrease around sunset would evoke continuous upward swimming. Gravity was the orienting cue and there was no change in swimming speed during the ascent. At sunrise, the larval descent was initiated by exposure to an absolute light intensity of about 0.23 log unit above the lower visual threshold. Light served as an orienting cue, as larvae descended by a negative phototaxis. Thus, phototaxis is not a laboratory artifact and does participate in vertical migration. A consideration of behavioral responses of other crustacean zooplankton indicates there is considerable variation in the initiating and directing cues for vertical movements. The variety of behavioral responses of R. harrisii suggests that a synthesis of hypotheses about migration may provide the proper basis for explaining the mechanisms underlying diel vertical migration.     

Direct and indirect trophic interactions of <Emphasis Type="Italic">Aurelia </Emphasis>sp. (Scyphozoa) in a stratified marine environment (Mljet Lakes,Adriatic Sea)

The pattern of diel vertical migration and the trophic interactions of moon jelly (Aurelia sp.) were investigated in the sea lakes of Mljet Island (Adriatic Sea) where this scyphomedusa is present throughout the year. Water column characteristics, plankton and in situ behaviour of Aurelia were followed over several 24-h cycles (6–8 times during each cycle) from the surface to the bottom (44 m). Aurelia exhibited a consistent pattern of diel vertical migration. Most of the time Aurelia were located at the bottom of the thermocline layer at temperatures lower than 19°C. Aurelia migrated towards the surface at dusk when the majority was found within the thermocline or just above it. During the night the medusae sank into the deepest layers below 25 m. The main medusa food items inferred from stomach contents were small adult copepods like Oithona nana and Paracalanus parvus and copepodites of small calanoids and cyclopids. In addition, in situ feeding experiments indicated high clearance rates for nauplii and naked ciliates and clear response of bacterial populations pointing to indirect cascade effects of Aurelia on microbial in addition to classical food web.     

Effects of age and food availability on diel vertical migration of Calanus pacificus

Age-specific differences in diel vertical migration behavior of Calanus pacificus were investigated in a 58 d (30 April–26 June, 1981) experiment in the Scripps Institution of Oceanography Deep Tank, La Jolla, California, USA; the experiment spanned three successive generations of copepods. The onset of vertical migration behavior occurred in the first feeding stage, Nauplius III. The amplitude of vertical migration gradually increased with age, becoming maximal in the late copepodite stages. Night depths remained constant with age while daytime depths increased. The migratory behavior of late copepodite stages was influenced by food availability. When phytoplankton was abundant and individual ingestion rates were high, copepodites performed high-amplitude migrations. As food availability declined, however, and the competition for food increased, migration amplitudes decreased and then ceased altogether so that copepodites remained in the relatively food-rich surface waters at all times. We suggest that hunger is the primary factor controlling vertical migration behavior.