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 feeding patterns of midwater fish in the central equatorial Atlantic

Fishes and zooplankton were obtained (March–April 1979 and partly in August 1974) from 45 hauls taken during the day and at night in the central equatorial Atlantic between Latitude 3°N and 2°S from the surface to 1250-m depth, using the RMT 1+8, a combined opening-closing plankton and micronekton trawl. The vertical distribution of 30 myctophid species is described. All species migrate in a diel pattern, Ceratoscopelus warmingii and Lampanyctus photonotus down to at least 1250 m. During daytime most species aggregated at 400-to 700-m depth, therefore only partly occupying the depth of the Deep Scattering Layer (400 to 500 m at 15 kHz). The feeding patterns of seven of the most abundant species were compared, with a total of 1 905 stomach contents being analysed. All seven species are regarded as opportunistic predators, which feed predominantly during the night on calanoid copepods. A total of 66 species of calanoid copepods were identified among the prey items, with smaller species definitely being in the minority. Stomachs of C. warmingii (700 to 1 250 m depth) and Lepidophanes guentheri (500 to 900 m depth) from daytime samples contained copepod species restricted to the upper 150 m of the water column, including Undinula vulgaris, Nannocalanus minor, and Euchaeta marina, thereby confirming an extended vertical migration of predators. Differences in diet and preferences between species in their total food spectrum are described.     

Vertical distribution,and seasonal and diurnal migration of Calanus helgolandicus in the Celtic Sea

The vertical distribution and migration (seasonal, diel and ontogenetic) of Calanus helgolandicus are described from the shallow (100 m) shelf-seas to the south-west of the British Isles. In 1978 and 1979, the overwintering population of C. helgolandicus consisted primarily of Stage V copepodites and adults. By late winter/early spring the copepodites had moulted to adult females (>90%), which matured and bred the first cohorts of the year, prior to onset of the spring phytoplankton bloom in April/May. C. helgolandicus reached a peak of numerical abundance in August of 20x10³ copepodites m^-2 (over the depth range sampled -0 to 70 m), which was 200 times the population in winter. The seasonal peak of abundance occurred 4 mo after the peak of the bloom of phytoplankton in spring. The yearly development of the copepod was not always out of phase with the diatom bloom, as seen when the data from 1978 was placed in the context of a longer time-series collected at 10 m over 22 yr (1960–1981, inclusive). Large vertical migrations were observed in the younger copepodites (CI and II) in May from below to above the thermocline. In the remainder of the year, the CI and CII stages behaved differently and were located above the thermocline within the euphotic zone. The largest vertical displacements of biomass were seen in the summer months due to the migrations of the CV stages and adults, which had developed from the spring cohorts. It was contended that the seasonal and vertical migrations of C. helgolandicus are part of a more complex pattern of inherent behavior than has been reported previously and that, however difficult this is to discern in the natural populations, it always expresses itself.     

Feeding patterns of Lampanyctus pusillus (Pisces: Myctophidae) throughout its ontogenetic development

The trophic ecology of the lanternfish Lampanyctus pusillus was investigated using individuals captured off the Balearic Islands (39°N, 2°E) (western Mediterranean) in December 2009. Based on gut content analyses, the trophic niche breadth, diet composition and selectivity were determined for the entire life cycle of L. pusillus. The larval stages fed actively near the surface during the day, with a feeding incidence (FI) of approximately 71 %. In contrast, the adults fed at night, both in near-surface depths and in the 400 m deep scattering layer, with a higher FI (83 %). Diet analysis revealed a shift in the prey choice throughout ontogenetic development, from preflexion individuals, which selected nauplii and small oncaeids, to postflexion larvae, which consumed a variety of calanoids, mainly Clausocalanus spp., to the adults, which preyed on large organisms, exhibiting positive selectivity for Pleuromamma spp. and euphausiids. These results show that the vertical distribution of larvae and adults is partly conditioned by their respective feeding habits, with larvae feeding on small zooplankton in the upper layer and adults preferring to consume larger taxa that perform nycthemeral migrations.     

Diel vertical migration and feeding of copepods at an oceanic site near South Georgia

Twelve Longhurst Hardy Plankton Recorder (LHPR) profiles were taken over a 16 h period in January 1990, in order to study feeding of four copepod species at an Antarctic oceanic site near South Georgia. Vertical distributions of their life stages, as well as those of dominant competitors and predators, are described in relation to the feeding cycles of Calanoides acutus CV, Calanus simillimus CV, Calanus propinquus CV and Rhincalanus gigas CIII, CV and CVI. Comparisons with vertical ring-net catches, which were used for concomitant gutevacuation experiments, demonstrated the suitability of the LHPR for these fine-scale studies. Planktonic predators, with the exception of the diel migrant Themisto gaudichaudii, resided deeper than the herbivores. During the day and around midnight, when feeding rates were low, species and stages reached their maximum vertical separation. At these times, new generation copepodites of the four species lived progressively deeper and the overwintered generation (i.e., R. gigas Stages CIV, CV, CVI) were progressively shallower. During the afternoon or evening (depending on species), all stages older than CII, as well as Euphausia frigida and T. gaudichaudii, migrated upwards, to amass in the surface mixed layer. Feeding was restricted to darkness, although R. gigas commenced several hours before dusk. In detail their migration and feeding differed widely, with combinations of unimodal and apparent bimodal cycles. As a whole, the results suggest that (1) feeding could occur during sinking as well as during upward migrations, (2) upward migrations were not always associated with feeding increases, and (3) individuals appeared to descend after filling their guts.     

New patterns of diurnal vertical migration of some deep-water copepods in the Tyrrhenian and Adriatis Seas

The diel vertical migration of copepods from intermdiate water layers of the Tyrrhenian and Adtiatic Seas has been studied on the basis of plankton samples made from depths down to 1000 m, with a Nansen net, mesh size 250 , diameter 113 cm. The species considered display two different patterns of migration: (1) nocturnal ascent, e.g. Pleuromamma abdominalis and Euchaeta acuta; (2) nocturnal descent, e.g. species of the genera Spinocalanus, Scaphocalanus, Temoropia, Mormonilla and Oncaea. These two patterns are compared on the pasis of the dimensions of the species and of their different capacities for active movement. Ililumination is considered to be the most important factor influencing vertical movements. The behaviour of Pleuromamma gracilis is also described, the adult population of which is divided in two discrete sections, each one showing a different pattern of migration.     

Diel patterns of carbon incorporation into biochemical constituents of Synechococcus spp. and larger algae in the Northwest Atlantic Ocean

Diel patterns of ¹⁴C-bicarbonate incorporation in>5 m algal communities were compared with those in cyanobacterial populations of Synechococcus spp. (0.6 to 1.0 m), collected from the surface and/or chlorophyll maximum at three stations in the Northwest Atlantic Ocean (a neritic front; in Warm-Core Eddy 84-E; and Wilkinson's Basin) from 21 July to 8 August, 1984. Cell constituents were chemically separated into four fractions: lipids, low molecular weight (LMW) metabolites, polysaccharides/nucleic acids, and proteins. The in situ diel pattern of ¹⁴C assimilation was virtually the same for >5 m algal communities adapted to different environments. Protein synthesis appeared to continue at a reduced rate at night using energy derived from the catabolism of polysaccharides and the mobilization of LMW compounds. Synechococcus spp. populations exhibited inherent physiological differences in their in situ diel pattern of carbon fixation from that in>5 m algal communities taken from the same water mass. There was no nighttime protein-synthesis in Synechococcus spp. The relative proportion of ¹⁴C-protein remained constant over night, while that of ¹⁴C-polysaccharides/nucleic acids declined and that of labelled LMW metabolites increased. Daytime light-intensity manipulations did not alter the diel pattern of carbon fixation in any of the>5 m algal assemblages, while changes in the carbon metabolism of surface and shadeadapted Synechococcus spp. populations could be rapidly induced by altering the light intensity.Bigelow Laboratory Contribution No. 86004     

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.     

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.     

Diel vertical migrations and nocturnal feeding of a dense coastal krill scattering layer (Thysanoessa raschi and Meganyctiphanes norvegica) in stratified surface waters

To study the nocturnal feeding of euphausiids during vertical migrations and its impact on the phytoplankton, a phytoplankton-rich water mass (drogue marked) drifting over a dense krill scattering layer (acoustic 104kHz) in the lower St. Lawrence estuary was monitored over 46 h in July 1982. Phytoplankton >20 m was abundant and mostly concentrated at the bottom of the photic layer above the pycnocline. Less than 42% of the particulate carbon was due to phytoplankton. The krill scattering layer was about 2 to 3 km in width, elongated along the 100-m bathymetric contour, and absent when the bottom was shallower than 50 m. Its upper day depth was 50 m. At deeper depths, its vertical distribution frequently changed from unimodal to polymodal shapes and viceversa, often with large concentrations of zooplankton just above the bottom. Typical vertical migrations were observed on both days. At night the scattering layer had a lower scattering strength. Most of it was below the thermocline but net catches showed that large concentrations of euphausiids (up to 57 individuals m^-3) crossed it. Stomach pigment content of Thysanoessa raschi was generally low, but mean stomach fullness was always high. They were more opportunistic than herbivorous. From stomach fullness and the presence of a food bolus in mouth parts, feeding in surface waters appeared to be intensive, but gut content indicated that food was not processed there. It is therefore suggested that individuals underwent vertical interchanges across the thermocline while feeding during the night. Meganyctiphanes norvegica had significant herbivorous activity during the night. The grazing pressure impact of the scattering layer on phytoplankton was negligible.Contribution to the program of GIROQ (Groupe interuniversitaire de recherches océanographiques du Québec)     

Recruitment,growth, and diel vertical migration of Euphausia pacifica in a temperate fjord

The pelagic crustacean Euphausia pacifica Hansen was sampled with a multiple-sample 1.0 m² Tucker trawl and a multiple-sample 1.0 m² vertical net in Dabob Bay, Washington on 17 dates between May 1985 and October 1987. Size (stage) structure and abundance of the population were determined for each date, while vertical distribution and diel migration were determined for 13 dates. Although internannual variability in both timing and magnitude of events occurred, consistent patterns were discernable. The population produced a large pulse of larvae (2 to 5 mm) in late spring of each year, apparently in response to the vernal phytoplankton bloom. Much lower abundances of larvae occurred during summer and autumn of each year, and larvae were completely absent during winter. Recruitment to the juvenile (6 to 9 mm) and adult (10 mm) stages was strongest during the summer, with abundances of these individuals peaking in summer and autumn. Individual growth rates, determined by modal progression analysis, were calculated for E. pacifica. Rates ranged from zero for some adult cohorts during the winter to 0.12 mm d^-1 for larvae during spring. The latter are among the highest ever reported for this species in the field. The vertical distributions and diel vertical migrations (DVM) of E. pacifica varied seasonally and between size (stage) classes. At night, all size classes were distributed in the surface layer (upper 25 m) irrespective of season or year. During the day, the larger/older stages were always distributed at middepths (50 to 125 m). In contrast, the daytime distribution of the larvae was more variable, being concentrated at the surface during spring and early summer of 1985, and at increasing depths later in the summer and autumn of 1985 and again in spring of 1986. This resulted in invariant DVM in the juveniles and adults, but variable DVM in the larvae, the latter of which is hypothesized to be a response to variable abundances of zooplanktivorous fish.     

Diel and seasonal changes in the distribution of fish on a southeast Brazil sandy beach

The effect of diel and seasonal changes in the distribution of fishes on a subtropical sandy beach on the southeastern coast of Brazil were studied. Seine netting was carried out on both seasonal and diel scales between July 1998 and June 1999. A total of 46 fish species was recorded, six being numerically dominant: Anchoa tricolor, Gerres aprion, Harengula clupeola, Atherinella brasiliensis, Mugil liza and Diapterus rhombeus. Seasonal changes in abundance of dominant species were detected. Species dominant in winter were Anchoa tricolor, H. clupeola and Atherinella brasiliensis; in spring, Anchoa tricolor and G. aprion; in summer G. aprion and D. rhombeus; and in autumn M. liza and H. clupeola. Overall, diel patterns did not reveal any significant trends; however, if we consider each season separately, an increase in A. tricolor abundance was recorded during the day in winter/spring, being replaced at night by H. clupeola in winter, and by G. aprion in spring. Increases in number of individuals and biomass at sunset, and decreases during the night were recorded. The winter/spring inshore/offshore movements at diel scale performed by the three most abundant species demonstrate that diel fluctuation acts more at a species-specific level than at a structural one; in summer there was no evidence of diel movements due to a heavy influx of G. aprion and D. rhombeus, which use the area throughout day and night, increasing overall abundance. Seasonal movements seems to be related to ontogenetic change in species, while diel movements in the fish assemblage seem to be more related to physiological requirements, such feeding activity of each particular species, than to physico-chemical conditions.Communicated by O. Kinne, Oldendorf/Luhe     

Feeding habits of the dungeness crab Cancer magister as determined by the index of relative importance

Specimens of the Dungeness crab Cancer magister, were collected subtidally and intertidally from an estuary in Washington State, USA in June and September 1980, and January, April, May and July, 1981. Gut contens of freshly collected crabs were analyzed by the Index of Relative Importance; for each prey taxon, this method measured frequency of occurrence, percentage of total biomass, and percentage of total numbers consumed. The most important higher taxon eaten was fish; however, the most important prey genus was the shrimp Crangon spp. There was greater predation on Crangon spp. at night at the intertidal site, and during winter and spring when the shrimp were most abundant there. Feeding activity, as indicated by a weight-specific gut-fullness index, showed no consistent diel pattern. There were significant ontogenetic changes in feeding patterns: first-year crabs preyed primarily on very small bivalves or small crustaceans including their conspecifics; second-year crabs preferred Crangon spp. and fish, and third-year crabs preyed less on Crangon spp. and more on fish. Such changes in feeding habits with ege could be purely due to mechanics of food handling, but might also reduce competition among age groups of crabs, possibly partitioning resources within the estuary. Findings are discussed in terms of optimal foraging and compared to other similar studies.Contribution No. 599, School of Fisheries, University of Washington, Seattle, Washington 98195, USA     

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%.     

Lunar rhythms of vertical migrations coded in otolith microstructure of North Atlantic lanternfishes,genus Hygophum (Myctophidae)

Otoliths of five Hygophum species were examined by means of light and scanning electron microscopy. In otoliths of four species (H. benoiti, H. macrochir, H. reinhardtii and H. taaningi) a strong cyclic pattern of the incremental structure was observed. In the fifth species (H. hygomii) such a pattern did not exist. An analysis of archival data on mesopelagic collections suggested three types of Hygophum spp. migratory behavior in relation to the lunar cycle which corresponded with the otolith microstructure. In H. hygomii only limited influence of moon phase on the uppermost range of night-vertical migration toward the surface was observed. The abundant nighttime occurrence of this species moved from the 0 to 50 m into the 50 to 100 m depth strata at full moon. In H. benoiti, a great part of the population, mainly juveniles, showed a tendency toward cessation of the vertical migrations during the first and fourth quarters of the lunar cycle. H. macrochir and H. taaningi had the strongest correlation of behavior with the lunar cycle. Both species showed arrested vertical migrations at the new moon phase, staying at day depths during the night, i. e., below 400 m. Thus, sequences of clear growth increments in otoliths represented a fast-growth period associated with the night migration to the warm surface layers, while bands without easily distinguishable incremental structure were interpreted as a period of slow growth in deep, cold waters due to limitation of the upward migration range occurring approximately at new moon.     

Diel feeding behavior of neritic copepods during spring and fall blooms in Akkeshi Bay,eastern coast of Hokkaido,Japan

In situ diel feeding behavior of neritic copepods was investigated using the gut fluorescence method, during spring and fall bloom periods in Akkeshi Bay, on the eastern coast of Hokkaido, Japan. Acartia omorii and Paracalanus sp. were the dominant species during the fall, and Pseudocalanus spp. and A. longiremis during the spring. During both bloom periods, diel rhythms were always observed for the gut pigment contents of these dominant copepods, although there were interspecific differences in the pattern. The maximum gut pigment content was always observed during the night and the minimum during the day. For all species, except Paracalanus sp., the average gut pigment content during the night was significantly higher (p<0.05) than during daytime by factors of between 1.5 and 2.7. There were no significant differences between the gut evacuation rate constants determined during the day and the night, and initial gut pigment content had no effect on the value of gut evacuation rate constants. The instantaneous ingestion rates of individual copepods calculated from gut pigment and the mean value of gut evacuation rate constants followed the same diel rhythms as gut pigment contents. Copepod daily ingestion rates were higher than the daily requirements for respiration during both bloom periods. Estimated daily ration was 40 to 91% of body carbon during the fall bloom, and 17 to 28% during the spring bloom. The higher daily rations during fall were probably due to the difference in in situ temperature (ca. 14°C).     

Vertical distribution and feeding of the copepod<Emphasis Type="Italic"> Chiridius armatus</Emphasis>

Adult females of the copepod Chiridius armatus inhabited the lower half of a 200-m water column during winter and spring. Their distribution became gradually shallower during summer and autumn. Female C. armatus carried out diel vertical migrations, with shallower population distribution at night compared to day, although normally with few individuals captured in the upper 50 m. Enumeration of fecal pellets produced by freshly collected individuals showed that C. armatus was foraging throughout the year. Production of fecal pellets was at a minimum in February and at a maximum in April. Individuals from all depths produced fecal pellets, but the numbers seemed to decrease by depth. Numbers of fecal pellets were greatest for individuals captured at night. The omnivorous C. armatus consistently produced many more fecal pellets than the concurrently studied, strictly carnivorous copepod Paraeuchaeta norvegica. We ascribe the relatively high number of fecal pellets to the higher content of indigestible components in plant material than in animal food. The assumption of phytoplankton as a prevailing food source got qualitative support from scanning electron microscope pictures, showing that fecal pellets mainly contained phytoplankton.Communicated by L. Hagerman, Helsingør     

Vertical distributions of eggs,nauplii and copepodites of Calanus helgolandicus (Copepoda: Crustacea) in the Celtic Sea

The vertical distributions of eggs, nauplii, copepodites and adults of Calanus helgolandicus (Claus) from five oblique plankton-net hauls taken in May (1980), March and September (1981) and January (1982) at a site in the shelf sea to the south-west of the United Kingdom are described. The water depth is approximately 95 m and becomes thermally stratified during the summer months when a thermocline of ∼6 C° develops. In early spring when the water column was isothermal (∼8 °C), the development of the eggs and nauplii took place below 60 m and a single ontogenetic migration was observed between Nauplius VI (NVI) and Copepodite I (CI). As the temperature of the water increased, this migration occurred in progressively earlier naupliar stages. The eggs were distributed throughout the water column in the profile taken in early May when a 1 C° thermocline occurred between 30 to 40 m. The majority of the NI to NIV stages occurred below 40 m, with the ontogenetic migration taking place in the NIV stage; the NV and NVI stages were found above the thermocline. In September, the eggs were again distributed throughout the water column (101 490 m^-2), with a maximum number of >4 500 m^-3 occurring in the surface to 5 m depth interval. Nauplius I and II were found at all depths, demonstrating that hatching occurred throughout the water column. The ontogenetic migration in these late-summer profiles took place between the NII and NIII stages, the remainder of the nauplii being found above the thermocline in the top 20 m. This is the first time that an ontogenetic migration, similar to the developmental ascent observed in the naupliar stages of the euphausiid Euphausia superba in the deep ocean, has been shown for a copepod nauplius.     

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.     

Diel changes in gut-cell morphology and digestive activity of the marine copepod Acartia tonsa

Morphometric measurements of the gut cells of the marine calanoid copepod Acartia tonsa were made over a 24 h diel cycle to test the hypothesis that feeding rhythms in this copepod are limited by the cycles of blishter-like cells (B-cells) in the gut. Copepods were collected from Long Island Sound, New York, USA, in June 1991. Experimental treatments included low (1 g chlorophyll al^-1) and high (10 g chlorophyll al^-1) food concentrations. Copepods were fixed and embedded in Epon at six time intervals over the 24 h period, and semi-thin sections through the length of Midgut Zone II were analyzed for gut-cell area and vacuole area, with area measurements integrated to yield estimates of gut-cell volume and vacuole volume. Concurrent measurements of gut fluorescence, a measure of gut fullness, and digestive enzyme activities also were made. A diel cycle in gut fluorescence was observed, most notably in the low-food treatment, which exhibited a 3-fold increase in gut pigments. There was little consistent change in digestive enzyme activities over this time span. Gaps in the gut wall indicative of spent B-cells were observed in 17 of 39 copepods, with no trend over time or food treatment. Generally, only a single gap was seen in any one copepod. B-cells were vacuolated throughout the 24 h period. Morphometric analyses revealed a correlation between gut fluorescence and both maximum vacuole area and vacuole volume, as well as percent vacuolated cell volume, in the low-food treatment. The high-food treatment, which had a relatively smaller increase in gut fluorescence (1.5-fold) during the night cycle, showed no significant increase in vacuole size. B-cells appear to have a life-cycle greater than the diel feeding period, and while B-cell vacuoles respond to the ingested food during the diel cycle, production of B-cells does not appear to be a limiting factor in A. tonsa's feeding cycle.