Natural diets of vertically migrating zooplankton in the Sargasso Sea

The feeding preferences of three common diel vertically migrating zooplankton were investigated from December 1999 to October 2000 at the U.S. JGOFS Bermuda Atlantic Time-Series Study (BATS) station in the Sargasso Sea. Gut content analysis of the copepods Pleuromamma xiphias (Giesbrecht) and Euchirella messinensis (Claus) and of the euphausiid Thysanopoda aequalis (Hansen) indicated that all three species fed on a wide variety of phytoplankton, zooplankton, and detrital material. Diet changes generally reflected seasonal trends in phytoplankton community structure. However, species-specific feeding preferences and differences in feeding selectivity among the three species were evident, and in general agreement with feeding habits predicted from the analysis of mouthpart morphology. The euphausiid T. aequalis fed equally on more different food types compared to both copepod species. The copepod P. xiphias consumed a diverse assemblage of phytoplankton from late winter through the summer (78-93% of gut items, by number, were phytoplankton) and based its diet more strongly on carnivorous feeding in autumn and early winter (31% and 61% of gut items were phytoplankton, respectively). E. messinensis showed the greatest feeding specialization, with a strong preference for pennate diatoms in winter and spring and for coccolithophorids during late summer and fall (constituting 67-93% of gut items by number). All three species consumed diatoms more than other phytoplankton taxa, even though diatoms form only a small fraction of the phytoplankton biomass in the Sargasso Sea. Although the majority of gut items identified were phytoplankton cells, the relative biomass contribution of these small cells may be lower than that of zooplankton and detritus. Zooplankton on which the three species primarily preyed were protozoans and crustaceans, but also included other metazoans such as chaetognaths and cnidarians. Marine snow was also an important component of the diet in all three species, with typically >50% and rarely <20% of the gut content being olive-green debris. Marine snow from larvacean houses was found in the guts of all three species, while E. messinenis appeared to selectively consume marine snow aggregates enriched with bicapitate Nitzschia spp. Large cyanobacteria (>4 µm in diameter) found in guts were also likely consumed with marine snow. The species-specific differences in the diets of these three migrating species suggest that an individual species approach is important in determining how feeding habits affect the structure of pelagic food webs and carbon cycling in the sea. Electronic supplementary material to this paper can be obtained by using the Springer LINK server located at http://dx.doi.org/10.1007/s00227-002-0815-8.     

Fungi from the Sargasso Sea

Fungi inhabiting Sargassum spp. from the Sargasso Sea were examined. Three ascomycetes were found, two of which are new records for this environment. Haloguignardia oceanica (Ferdinandsen et Winge) Kohlm. comb. nov. was collected only once before, over 50 years ago. Haloguignardia tumefaciens (Cribb et Herbert) Cribb et Cribb was known only from Australia and New Zealand. Imperfect states of these ascomycetes and fruiting bodies (acervuli) of a hyperparasitic deuteromycete (cf. Sphaceloma sp.) were found for the first time in the galls of the two species. Lindra thalassiae Orpurt et al., previously known only from the sea grass Thalassia testudinum Köngig, develops in air bladders of Sargassum sp., causing a shedding of infested vesicles. Possible reasons for the rarity of fungal species and diseased plants in the Sargasso Sea are diseussed.Dedicated respectfully to my teacher, Professor Dr. H Drawert (Universität Famburg), on the occasion of his 60th birthday.     

Diel patterns of planktonic bioluminescence in the northern Sargasso Sea

Day-night changes in the vertical distribution, intensity, and size of bioluminescence flashes were investigated during a series of cruises to the northern Sargasso Sea in 1987 and 1988. Overall, depth integrated bioluminescence potential and flash density estimated from in situ measurements with a pumping bathyphotometer were 2 to 5 x higher at midnight than at midday. Depths from 50 to 100 m exhibited the most substantial day to night increases in bioluminescence potential and flash density. When classified by flash size (photon output per flash event), the increase from day to night was significant for all flash sizes, but was most dramatic for small flashes producing <7 x 10⁸ photons flash^-1. Bioluminescence potential and flash density increased 2 to 3 x during bathyphotometer measurements made at dusk. Bioluminescent light budgets estimated from day and night net collections in May and August 1987 also predicted 2.5 x higher nighttime than daytime mesoplankton bioluminescence. However, large bioluminescent taxa (mesoplankton) capable of significant vertical migrations only contributed on the order of 15% of the total bioluminescence in surface waters. Our results do not support the idea that most of the nightly increase in bioluminescence potential and flash density are due to vertical migration of bioluminescent organisms; rather they are consistent with an alternate view that photoinhibition of bioluminescent flashing by dinoflagellates may be primarily responsible for the diel patterns.     

Mesopelagic fishes and thermal fronts in the western Sargasso Sea

Analysis of 25 midwater trawl collections, disposed along the meridian 70°20′W from off Hispaniola to the Gulf Stream, showed a change in the mesopelagic fish fauna at about 27°N. The point of faunal change corresponded to a change in the temperature structure of the upper part of the water column, i.e., at a (the?) so-called “thermal front”, perhaps identical to the so-called “North Atlantic subtropical convergence”. Of 44 species occurring in four or more collections, 13 species were collected only to the north of the front and 1 species only to the south. By most criteria, north-of-the-front collections were larger than southern ones. This is in accord with the north-south difference in primary production noted by other workers which, in turn, seems atributable to the north-south difference in temperature structure. To the north the upper part of the water column is well stratified in summer only, while to the south it is well stratified at all seasons. Taken altogether, what is now known suggests that the thermal front logically divides the upper Sargasso Sea into northern and southern portions that differ in many ways.     

Spectral composition of bioluminescence of epipelagic organisms from the Sargasso Sea

The spectral characteristics of single identified epipelagic sources of bioluminescence from the western Sargasso Sea were measured with an optical multichannel analyzer (OMA) system during the April, 1985, Biowatt cruise. The emission spectra of specimens representing 45 species from 8 phyla were measured. Peak bioluminescence emissions typically occurred between 440 and 500 nm, in the blue region of the visible spectrum. Three exceptions involved emission in the green, yellow, and red spectral regions. Intraspectific variability in spectra, was noted in several species. One shrimp species exhibited two modes of light emission, each with different emission spectra. Other cases involved dynamic color shifts of 10 to 14 nm; the source of the spectral variability is unknown, but may involve optical filtering or differences in the color of luminescence from multiple sites of light emission. Measurements from independent samples of unsorted plankton revealed different spectral distributions. This suggests that the spectral emissions of bioluminescence in the upper water column will vary, based on species assemblage.     

Temporal variability of deep-sea zooplankton in the Arabian Sea

Mesozooplankton samples from two stations in the Arabian Sea (WAST, 4,050 m, 16°15′N, 60°20′E; CAST, 3,950 m, 14°30′N, 64°30′E) were collected from the surface down to 20 m above bottom during three monsoon periods: the autumn intermonsoon in October 1995, the spring intermonsoon in April 1997, and the NE monsoon in February 1998. The main goal of this study is to enhance our knowledge on the effect of spatial and temporal differences in primary production and particle flux rates on the abundance and distribution of mesozooplankton, with special attention to the deep sea. Literature data indicate episodically high rates of primary production and particle flux in the region during the SW monsoon and the autumn intermonsoon. Set in this context, the zooplankton showed an in-phase coupling in biomass and abundance with the primary production in the surface 150 m. In the mesopelagic realm (150–1,050 m), the seasonal coupling was less clear. In the bathypelagic zone, below 1,050 m, there was no evidence of in-phase coupling, though temporal differences in the distribution of zooplankton abundance and biomass with depth between seasons could be shown by an analysis of covariance and an a posteriori test. The results suggest that the bathypelagic community responds to increased particle flux rates, but with longer time gaps than in the epipelagic zone. This is probably due to longer development and response times of zooplankton in the cold, deep-water environment, independent of possible lateral advection processes.     

Depth-dependent changes in chlorophyll fluorescence number at a Sargasso Sea station

Chlorophyll a-specific in vivo fluorescence exhibited depth-dependent changes in a Sargasso Sea phytoplankton community, decreasing from a maximum value at the surface to a minimum at 90m, and then increasing again below 90 m. This distribution pattern was not explained by irradiance conditions, diurnal variability, senescence in the deep population, or changes in light-absorption efficiency of chlorophyll a. However, a significant positive correlation was found between mean phytoplankton cell size and fluorescence number in the upper euphotic zone, where nutrient concentrations were low. We hypothesize that the direct cause for this observed correlation was nutrient limitation. In this picoplankton-dominated community, packaging effect was minimal. Under nutrient-limiting conditions, as mean cell size increases photosynthetic efficiency decreases and therefore fluorescence number increases. In the lower euphotic zone where nutrients were not limiting, changes in fluorescence number exhibited weak size-dependence and appeared to be related to species compositional changes.     

Pattern of planktonic bioluminescence in the northern Sargasso Sea: Seasonal and vertical distribution

The vertical structure of bioluminescence potential (BPOT) and flash density (FD) were measured on five cruises to the northern Sargasso Sea in 1987 and 1988. Depth-integrated (0 to 150 m) BPOT did not vary seasonally, remaining within the range 9 to 15 × 10¹⁵ photons m^–2 in all months sampled. Conversely, depth-integrated FD was significantly higher (> 2 × 10⁵ flashes m^–2) during winter (November and March) than during summer (< 9 × 10⁴ flashes m^–2 in May and August). The vertical patterns of BPOT and FD were well correlated within a single profile, more highly so in summer than in winter. Despite intracruise variability in the vertical pattern of BPOT and FD, there were clear summer-winter differences in the vertical distribution of BPOT and FD. During winter, BPOT and FD were maximal and relatively uniform throughout the surface mixed layer; for example in November they declined sharply within the thermocline at 130 to 150 m. During summer, BPOT and FD were greatest (12 to 25 × 10¹³ photons m^–3 and 600 to 1 200 flashes m^–3, respectively) at subsurface depths. Commonly in summer, the upper depth limit of high BPOT and FD occurred at the base of the surface mixed layer (10 to 40 m) and the lower depth limit was located at the base of the subsurface fluorescence maximum (usually at 100 to 120 m).     

Deep distributions of oceanic cirripede larvae in the Sargasso Sea and surrounding North Atlantic Ocean

From deep ( 1000 m), multi-depth zooplankton samples taken in the Azores frontal region from November 1980 to June 1981, high numbers of the two larval forms ofLepas pectinata, the commonest lepadid cirripede in this area, were sampled. The nauplii and cyprid larvae are large and long-lived. Nauplii were sampled where they feed, in the upper 150 m, closely associated with the fluorescence maximum. The cyprids were mainly sampled between 300 and 400 m, with a sharp cut-off in numbers just below the distribution peak. This deep distribution is intriguing, as the cyprids, which were not migrating diurnally, are non-feeding. To complete their life cycle they must settle on floating debris at the surface, where very few were sampled. They are negatively buoyant and there is no apparent physical reason for this deep distribution. There was no obvious pattern of changes in individual length or length/dry weight distribution with depth, the population appeared homogeneous. There was also no accumulation of other zooplankton at the same depth, or evidence from gut contents of large carnivores that the distribution pattern was caused by selective predation. A close association exists betweenL. pectinata andSargassum spp. weed, the cyprids preferentially settling on small fragments of weathered weed, the supply of which may be enhanced after winter. During winter, the harsh environment at the surface of the northern Sargasso Sea may discourage cyprid settlement. The deep distribution may be an ontogenetic migration conferring some survival advantage and saving energy due to the lower temperature and oxygen levels at depth. If it is a behavioural mechanism, duration of the cyprid stage will be extended until a more clement time of year, allowing them to settle over a longer period. The cyprids of other species were also sampled at depth, so the phenomenon appears to be widespread. If subsequent research confirms these observations as an ontogenetic migration, our current understanding of the fouling behaviour of oceanic lepadid cirripedes would be considerably modified.     

Morphology,nutrition and taxonomy of an Aplanochytrium sp. from the Sargasso Sea

Aplanochytrium sp. 2400/57 was isolated from samples of sargasso weed collected in the Sargasso Sea on April 4, 1979. The monocentric eucarpic thallus of this organism produced aplanospores by successive bipartitioning. The cell wall contained only 20% polysaccharide, composed of fucose, galactose, xylose, glucose, and galacturonic acid. The guanine+cytosine content of the DNA (mol% G+C) was 37.6%. L-isoleucine, L-leucine, and L-proline were utilized as nitrogen sources in synthetic media of defined composition.     

Dependence of dissolved carbohydrate concentrations upon small scale nanoplankton and bacterioplankton distributions in the western Sargasso Sea

Significant correlations between trophic components of the <20 m microbial plankton and the concentrations of total dissolved carbohydrate (TCHO) and polysaccharide (PCHO) have been found at two drogued buoy stations which were sampled at 3 or 4-h intervals over diel cycles. An attempt was made to sample on two pre-selected isotherms (approximating isopycnals) at each station. Significant inverse correlations of TCHO and of PCHO with numbers of phototrophic nanoplankton (PNAN) were found along both isotherms at a station off the Carolina coast. Off central Florida, TCHO and/or PCHO were significant multiple linear functions of PNAN, heterotrophic bacterioplankton (BAC) and sometimes heterotrophic nanoplankton (HNAN) counts. Partial regression coefficients for PNAN were always negative and those for HNAN were positive. Inverse trends dominated the relations of BAC with PCHO and TCHO. The combined data from the two stations (300 nautical miles apart) produced very similar multiple linear relations. This suggests that the observed relationships are real, resulting from general physiological processes and interactions of the microbial plankton groups rather than from chance occurrences at a particular station, and that the combined activities of the <20 m plankton actively regulate dissolved carbohydrate concentrations in the Sargasso Sea.     

Spring and autumn spatial distribution of zooplankton carbon requirement across the Mediterranean Sea

Zooplankton represents a key contributor to the ocean biological pump through its consumption of sinking and suspended carbon. A specific and highly sensitive method to evaluate zooplankton carbon requirement from the sinking flux is through the estimation of the activity of the electron transport system. The present study was carried out from samplings in 2006, and it was focused on the spatial 200–0?m zooplankton carbon demand across 24 sampling stations, along the Mediterranean Sea, from the island of Crete to the Strait of Gibraltar. Its potential day/night variability was evaluated. The zooplankton composition, abundance and biomass were investigated. The carbon demand per unit zooplankton biomass indicates geographical and diel differences among the sampling stations. A higher mean carbon demand was seen for the western Mediterranean with respect to the eastern Mediterranean, which can be justified through the observed ratio of gelatinous:crustacean taxa and the water temperatures recorded. Higher carbon demand was measured in samples collected during the dark hours. The relation to the presence and abundance of actively migrating euphausiids and copepods was discussed. A comparison with data from another of our study carried out in the same study area but in another seasonal period was done.     

Nodularin concentrations in Baltic Sea zooplankton and fish during a cyanobacterial bloom

Toxic cyanobacterial blooms, dominated by Nodularia spumigena, are a recurrent phenomenon in the Baltic Sea during late summer. Nodularin, a potent hepatotoxin, has been previously observed to accumulate on different trophic levels, in zooplankton, mysid shrimps, fish as well as benthic organisms, even in waterfowl. While the largest concentrations of nodularin have been measured from the benthic organisms and the food web originating from them, the concentrations in the pelagic organisms are not negligible. The observations on concentrations in zooplankton and planktivorous fish are sporadic, however. A field study in the Gulf of Finland, northern Baltic Sea, was conducted during cyanobacterial bloom season where zooplankton (copepod Eurytemora affinis, cladoceran Pleopsis polyphemoides) and fish (herring, sprat, three-spined stickleback) samples for toxin analyses were collected from the same sampling areas, concurrently with phytoplankton community samples. N. spumigena was most abundant in the eastern Gulf of Finland. In this same sampling area, cladoceran P. polyphemoides contained more nodularin than in the other areas, suggesting that this species has a low capacity to avoid cyanobacterial exposure when the abundance of cyanobacterial filaments is high. In copepod E. affinis nodularin concentrations were high in all of the sampling areas, irrespective of the N. spumigena cell numbers. Furthermore, nodularin concentrations in herring samples were highest in the eastern Gulf of Finland. Three-spined stickleback contained the highest concentrations of nodularin of all the three fish species included in this study, probably because it prefers upper water layers where also the risk of nodularin accumulation in zooplankton is the highest. No linear relationship was found between N. spumigena abundance and nodularin concentration in zooplankton and fish, but in the eastern area where the most dense surface-floating bloom was observed, the nodularin concentrations in zooplankton were high. The maximum concentrations in zooplankton and fish samples in this study were higher than measured before, suggesting that the temporal variation of nodularin concentrations in pelagic communities can be large, and vary from negligible to potentially harmful.     

Metabolism and elemental composition of zooplankton from the Barents Sea during early Arctic summer

Rates of oxygen consumption, ammonia excretion and phosphate excretion were measured on a hydromedusae (Aglantha digitale), pteropods (Limacia helicina, Clione limacina), copepods (Calanus finmarchicus, C. glacialis, C. hyperboreus, Metridia longa), an amphipod (Parathemisto libellula), a euphausiid (Thysanoessa inermis) and a chaetognath (Sagitta elegans), all of which were dominant species in the Barents Sea during early summer 1987. Water and ash contents and elemental composition (C and N) were also analysed on the specimens used in these metabolic experiments. Between species variations were 67.8% to 94.7% of wet weight in water content, 6.4% to 56.5% of dry weight in ash content, 16.7% to 61.0% of dry weight in carbon content, and 4.3% to 11.2% of dry weight in nitrogen content. Oxygen consumption rates ranged from 0.33 to 13.8 l O₂ individual^-1 h^-1, ammonia excretion rates, from 0.0072 to 0.885 gN individual^-1 h^-1 and phosphate excretion rates, from 0.0036 to 0.33 g P individual^-1 h^-1. In general, higher rates were associated with larger species, but considerable differences were also seen between species. The ratios between the rates (O : N, N : P, O : P) exhibited a wide species-specific variation, indicating differences in dominant metabolic substrates. Typical protein oriented metabolism was identified only in S. elegans. From the results of metabolic rate measurements and elemental analyses, daily losses of body carbon and nitrogen were estimated to be 0.50 to 4.15% and 0.084 to 1.87%, respectively, showing faster turnover rates of carbon than that of nitrogen. Comparison of daily loss of body carbon of the Barents Sea zooplankton with that of the Antarctic zooplankton indicated reduced rates of the former (63% on average).     

Surface zooplankton dynamics and community structure in the Gulf of Aqaba (Eilat), Red Sea

Winter and summer zooplankton maxima were observed on both near-reef and offshore sampling sites in the northern part of the Gulf of Aqaba, with summer maxima smaller than those of winter and more characterized by larval forms. Near-reef zooplankton biomass was generally several times greater than that observed 2 km offshore. During 1987, a near-reef maximum of 155 ind. or 12.2 g wet biomass m^–3 was observed in March, while 103 ind. or 8.5 g wet biomass m^–3 was observed in July. In the same year, 2 km offshore a maximum of 53 ind. or 2.5 g wet biomass m^–3 was observed in February, while a maximum of 33 ind. or 0.5 g wet biomass m^–3 was noted in July. The following year, 1988, the near-reef zooplankton abundances were little changed, but offshore zooplankton abundances were much higher (317 m^–3). During 1987, the dominant winter (March) forms near the reef were gammarid amphipods, at maximum concentrations of 100 ind. m^–3, where the summer (July) maximum was composed primarily of mysids (34 m^–3), gammarid amphipods (30 m^–3), and fish eggs (24 m^–3). The offshore winter zooplankton fauna was characterized by copepods and appendicularians, each at a maximum concentrations of ca 13 ind. m^–3, while the summer maximum was dominated by brachyuran zoea (31 m^–3). Though the 1988, winter near-reef zooplankton community compositions were similar to those of 1987, the offshore zooplankton fauna was dominated by ostracods, which were relatively rare in previous years. Preliminary data suggests that holoplanktonic forms like chaetognaths, copepods and appendicularians, at an offshore site exhibit different patterns of vertical migration than those near the reef. This different behavior may result from different species compositions of these taxa or from high concentrations of pseudoplanktonic bentho-neritic peracarid crustaceans.Please address correspondence and reprint requests to T. Echelman, Marine Science Research Center, State University of New York, Stony Brook, New York 11794-5000, USA     

Distribution and trophic links of gelatinous zooplankton on Dogger Bank,North Sea

The ecology of small, gelatinous zooplankton is not integrated into management of Dogger Bank (54° 00′ N, 3° 25′ E to 55° 35′ N, 2° 20′ E). In pursuit of this goal, gelatinous zooplankton and their potential prey were sampled along a transect across the bank on June 10–16, 2007. Eleven species of small medusae and ctenophores were collected, with six abundant taxa occurring in greater numbers below the thermocline and in the shallower, southeastern portion of the bank. There were no statistically significant diel changes in distribution. In contrast, potential prey were distributed more evenly across the bank and throughout the water column. Isotopic analyses revealed that gelatinous zooplankton fed on both smaller (100–300 μm) and larger (>300 μm) mesozooplankton, but also potentially on each other. These ecological insights suggest that small medusae and ctenophores should be integrated into sustainable management of Dogger Bank.     

Relationship between the zooplankton,phytoplankton, particulate matter and dissolved free amino acids in the Celtic Sea

Estimates of the biomass of zooplankton, phytoplankton and particulate matter collected in the Celtic Sea during mixed-water conditions (on 8 and 9 April 1983) were compared to the concentration and diversity of sixteen dissolved free amino acids (DFAA) measured in seawater and in particles. During a day profile, variations of dissolved amino acids with depth reflected the feeding activity of copepods. The relationship was not apparent in a night profile and other processes, such as heterotrophic utilization of dissolved nitrogen by microorganisms, were thought to be involved. The ratios of total DFAA concentration (nM litre^-1) in the particulate phase over the concentration in seawater ranged from 1 to 200 within the water column. Of the sixteen amino acids measured, ornithine, a decomposition product of arginine, was responsible for more than 70% of the total concentration of DFAA in seawater. In the particles, phenylalanine ranged from 30 to 88% of total DFAA. In seawater this amino acid occurred in the 20 to 40 m depths (1.3 to 9.9% of total DFAA) in the day profile and at 5 m (12.4%) and 80 m (6.4%) in the night profile. Previously it has only been found in very low concentrations (<5%) in seawater, and its presence is considered to be the result of zooplankton feeding.     

Distribution and abundance of ctenophores and their zooplankton food in the Black Sea. II. Mnemiopsis leidyi

The distribution of Mnemiopsis leidyi Agassiz, 1865 in the Black Sea was determined using plankton samples collected above the anoxic zone (maximum depth 200 m) in the summer, winter, and spring from 1991 to 1995. Distribution was patchy. Average biomasses of 15 to 500 g m⁻² were measured, and abundances varied from 10 to 180 ind m⁻². Biomass and abundance peaked in winter, and there was a secondary peak in the summer. The distribution of M. leidyi was correlated with hydrographic features in the Black Sea with higher concentrations in anticyclonic gyres. The centers of the two main cyclonic gyres generally had a low biomass of M. leidyi. From July 1992 to March 1995, the populations were largely offshore. M. leidyi were confined to the upper part of the mixed layer both day and night. Some individuals displayed a negative taxis to daylight and were concentrated below the thermocline at night. Smaller M. leidyi (1.5 to 2 cm) were present in the winter, and individuals reached maximum size in the summer. Although reproduction was continuous throughout the year, there were two distinct peaks: the larger peak in the summer and the smaller peak in the winter. Microscopic analysis of stomach contents showed that copepods and molluscs form their main diet. Received: 1 November 1997 / Accepted: 30 August 1999     

Distribution and abundance of moon jellyfish (Aurelia aurita) and its zooplankton food in the Black Sea

The distribution of moon jellyfish (Aurelia aurita Linnaeus, 1758) in the Black Sea was determined from plankton samples collected above the anoxic zone (maximum depth 200 m) in the summer, winter and spring during 1991–1995. Distribution was patchy. Average biomass ranged from 98 to 380 g m⁻², and abundance varied from 2 to 14 individuals m⁻². Biomass and abundance peaked in late spring and summer. The distribution was correlated with hydrographic features in the Black Sea, with higher concentrations occurring at the peripheries of anticyclonic eddies. Centers of the two main cyclonic gyres generally had a low biomass of A. aurita. From July 1992 to March 1995, the populations were largely concentrated in offshore regions. A. aurita were confined to the upper part of the mixed layer. Smaller A. aurita (≤1 cm) were present in early spring (March), and individuals reached maximum size in the summer. Release of the epyhrae occurred in spring on the northwestern shelf of the sea when the seawater temperature was 11–12 °C. Microscopic analysis of stomach contents showed that copepods and mollusks form their main diet. Received: 3 September 2000 / Accepted: 29 September 2000