The feeding habits of three species of lanternfishes (family myctophidae) off oregon,USA

The common myctophids Stenobrachius leucopsarus, Diaphus theta and Tarletonbeania crenularis were found to feed primarily on the euphausiid Euphausia pacifica, the copepods Metridia lucens and Calanus spp., and the amphipod Parathemisto pacifica. The diets of these species of fishes were diverse and overlapped broadly, suggesting that they are feeding generalists. Most stomachs contained either all copepods or all euphausiids. Euphausiids were the most important prey on the basis of biomass. They comprised over one-half the weight of the stomach contents in over 40% of individuals of each of the three fishes. Stomach fullness and state of digestion of stomach contents differed over the diel period, but not enough to indicate feeding only during the nighttime. Average stomach fullness was greatest during the morning and night and well-digested material, which predominated in most stomachs, was most prevalent in the morning and afternoon. Either some feeding occurs throughout the day or digestion rates are slow.     

Feeding ecology of five fishes from the mid-slope micronekton community off southern Tasmania, Australia

We examine the feeding ecology of characteristic micronekton species inhabiting the mid-slope region (~700 to 1,500 m depths) off southern Tasmania. Five fishes, Diaphus danae, Hygophum hanseni, Lampanyctus australis (Myctophidae), Phosichthys argenteus (Phosichthyidae) and Chauliodus sloani (Chauliodontidae), were sampled by depth-stratified midwater trawling on a diel and seasonal basis. Overall, 74 prey taxa were identified from 2,132 stomachs. Euphausiids (mostly Euphausia spinifera and E. similis) and calanoid copepods (mostly Pleuromamma species) were the main prey of the three myctophids; P. argenteus ate fishes and decapods in addition to the euphausiids, while C. sloani ate only fishes. Copepods were less important in the diets of larger D. danae, L. australis and P. argenteus and were replaced by euphausiids in the myctophids and by fishes in P. argenteus. In autumn, when euphausiid biomass increased six-fold, all three myctophids and P. argenteus fed most intensively and consumed a high proportion of euphausiids. The three myctophids appeared to feed nocturnally. Differences in the timing and duration of feeding corresponded to differences in their spatio-temporal overlap with key prey. Daily rations of H. hanseni (1.93% dry-weight biomass) and L. australis (1.43%), estimated from data on stomach fullness, were typical for temperate myctophids and higher than that of the non-migratory P. argenteus (0.43%). The vertical flux of near-surface plankton production to the mesopelagic food web is based primarily on diel feeding in the upper water column (<500 m) rather than consumption of species that migrate seasonally into the deeper mesopelagic zone. Because species such as P. argenteus and C. sloani feed above the third trophic level, their predators, including the commercially important orange roughy, are feeding between levels four and five.     

Feeding ecology of the lanternfish Benthosema pterotum from the Indian Ocean

The feeding ecology of lanternfish Benthosema pterotum (Alcock) from the north Arabian Sea, Mozambique and the Bay of Bengal was studied. Samples were collected on cruises carried out by R.V. Dr. Fridtjof Nansen during the period 1978 to 1983. A wide variety of zooplankton organisms were identified in the diet of B. pterotum with crustanceans dominating the diet. Copepods constituted ca. 40 to 90% of the diet. Dry weight analyses of the stomach contents from the Gulf of Oman in February 1983 showed copepods to be 35 to 55% in weight (average in samples). Ontogenetic differences were observed in the diet. Prey size increased as the fish length increased, but the largest fish did not exclude the smaller prey organisms from their diet. Regional variation in diet was also shown in B. pterotum. The degree of filling and the state of digestion of stomach contents revealed that this species feeds intensively at night in the epipelagic layer. All copepods indentified were epipelagic species, providing additional evidence of diurnal pattern in the feeding chronology of B. pterotum. Identification of copepods from the Gulf of Oman in February 1983, revealed that herbivorous species dominated in biomass. Quantitative analyses show that B. pterotum probably have a daily food intake of ca. 4.5% of the body weight.     

Feeding ecology of three species of midwater fishes associated with the continental slope of eastern Tasmania,Australia

The feeding ecology of Maurolicus muelleri, Lampanyctodes hectoris and Diaphus danae was examined from samples collected from continental slope waters of eastern Tasmania between April 1984 and April 1985. A total of 2 232 stomachs was analysed. M. muelleri, L. hectoris and D. danae fed primarily on euphausiids and secondarily on copepods, although larger D. danae (> 60 mm standard length) fed on other lanternfish (chiefly L. hectoris). The diets of M. muelleri and L. hectoris overlapped substantially. Diet overlaps between D. danae and the former species was low, however, due to the large biomass of fish present in D. danae. The prey taxa consumed changed with time of year and predator size. Stomach fullness (feeding intensity) varied seasonally in all three species, but only M. muelleri showed significant diel differences in fullness. The synchronization of the size structure of the predator populations and their feeding intensity, with seasonal variations in preferred prey, is proposed as a mechanism whereby each species maximizes its share of the available food resources.     

Spatial overlap and trophic interactions between pelagic fish and large jellyfish in the northern California Current

Recent studies have indicated that populations of gelatinous zooplankton may be increasing and expanding in geographic coverage, and these increases may in turn affect coastal fish populations. We conducted trawl surveys in the northern California Current and documented a substantial biomass of scyphomedusae consisting primarily of two species (Chrysaora fuscescens and Aurelia labiata). Spatial overlap of these jellyfish with most pelagic fishes, including salmon, was generally low, but there were regions of relatively high overlap where trophic interactions may have been occurring. We compared feeding ecology of jellyfish and pelagic fishes based on diet composition and found that trophic overlap was high with planktivorous species that consume copepods and euphausiid eggs such as Pacific sardines (Sardinops sagax), northern anchovy (Engraulis mordax), Pacific saury (Cololabis saira), and Pacific herring (Clupea pallasi). Moreover, isotope and diet analyses suggest that jellyfish occupy a trophic level similar to that of small pelagic fishes such as herring, sardines and northern anchovy. Thus jellyfish have the potential, given their substantial biomass, of competing with these species, especially in years with low ecosystem productivity where prey resources will be limited.     

Impact of copepod grazing on the red-tide flagellate Chattonella antiqua

Although planktonic copepods are major suspension feeders in the sea, the impact of their grazing pressure upon red-tide flagellates has not been fully investigated. In the present study, the grazing of adult females of several copepod species is examined using three food types: viz. natural suspended particles, natural suspended particles mixed with cultured Chattonella antiqua, and cultured C. antiqua. The functional response on C. antiqua was investigated for five species of copepods (Acartia erythraea, Calanus sinicus, Centropages yamadai, Paracalanus parvus and Pseudodiaptomus marinus). Ingestion rates increased linearly with increasing cell concentrations until a maximum level was reached, beyond which the rates were constant. This cell concentration was higher for larger copepods. The weight-specific maximum ingestion rates were higher in the small species. In general, copepods tended to feed selectively on larger particles when feeding on natural particles. This tendency was strongest in a simulated red-tide environment. Thus, it can be surmised that copepods may selectively graze on C. antiqua during the outbreak of a red tide. Grazing pressure by the natural copepod community in Harima Nada, the Inland Sea of Japan, was calculated by integration of the laboratory determined feeding rates and field measurements of zooplankton biomass. The daily removal rate was 3.4 to 30.8% (mean: 12.3%) of C. antiqua biomass at 20 cells ml^-1 and decreased to 0.6–4.3% (mean: 1.8%) at 500 cells ml^-1. Therefore, the grazing pressure by the copepod community is important at the initial stage of the red tide.     

Morphology and sinking velocities of fecal pellets of copepod, molluscan, euphausiid, and salp taxa in the northeastern tropical Atlantic

Morphological characteristics and sinking velocities of naturally occurring fecal pellets of copepods, euphausiids, salps, and pelagic mollusks collected in the northeastern tropical Atlantic were investigated during the period of May-June 1992. The fecal pellets of copepods and euphausiids were cylindrical and distinguished only by their size. Those of salps were, in general, rectangular, and slight differences were noted according to the species. The fecal pellets of the molluscan pteropod Clio sp. were conical, while those of the molluscan heteropod Carinaria sp. were spiral. The sinking velocities ranged from 26.5 to 159.5 m day^-1 for copepod fecal pellets, from 16.1 to 341.1 m day^-1 for euphausiid pellets, from 43.5 to 1167.6 m day^-1 for salps' pellets (Cyclosalpa affinis, Salpa fusiformis, Iasis zonaria, and two unidentified species), from 65 to 205.7 m day^-1 for Clio sp. pellets, and from 120.3 to 646.4 m day^-1 for Carinaria sp. fecal pellets. The measured sinking velocities were compared with estimates predicted using the equations of Komar et al. (1981; Limnol Oceanogr 26:172-180), Stokes' law, and Newton's second law, using either a constant density of fecal pellets (1.22 g cm^-3) or densities estimated with the three different equations. When a constant density was used, the three equations overestimated the sinking velocities; Stokes' law resulted in the largest overestimation, and Newton's second law, the smallest. At the taxa level, the overestimation was greatest for euphausiid 1 fecal pellets and smallest for copepod fecal pellets. When the three equations were used to estimate fecal pellet density, the density estimated using the equation of Komar et al. was the greatest, and that using Stokes' law, the smallest, resulting in over- and underestimation of sinking velocities, respectively. Newton's second law resulted in an intermediate density and gave the closest estimate of sinking velocities. We propose that measurement of sinking velocities of a portion of the fecal pellets might guide in choosing an appropriate equation to be used for a reasonable interpretation of vertical mass flux.     

Vulnerability of the copepod Acartia tonsa to predation by the scyphomedusa Chrysaora quinquecirrha : effect of prey size and behavior

Although scyphomedusae have received increased attention in recent years as important predators in coastal and estuarine environments, the factors affecting zooplankton prey vulnerability to these jellyfish remain poorly understood. Current models predicting feeding patterns of cruising entangling predators, such as Chrysaora quinquecirrha (Desor, 1948), fail to account for the selection of fast-escaping prey such as copepods. Nevertheless, our analysis of gastric contents of field-collected medusae showed that this scyphomedusa fed selectively on the calanoid copepod Acartia tonsa (Dana, 1846) and preferentially ingested adult over copepodite stages. We measured feeding rates in a planktonkreisel while simultaneously videotaping predator–prey interactions. C. quinquecirrha consumed adult A. tonsa ten times faster than copepodites. Differences in prey behavior, in the form of predator–prey encounter rates or post-encounter escape responses, could not account for the elevated feeding rates on adults. Prey size, however, had a dramatic impact on the vulnerability of copepods. In experiments using heat-killed prey, feeding rates on adults (1.5 times longer than copepodites) were 11 times higher than on copepodites. In comparison, medusae ingested heat-killed prey at only two to three times the rate of live prey. These results suggest that during scyphomedusan–copepod interactions, prey escape ability is important, but ultimately small size is a more effective refuge from predation. Received: 26 September 1997 / Accepted: 20 May 1998     

Zooplankton feeding ecology: contents of fecal pellets of the copepods Temora turbinata and T. stylifera from continental shelf and slope waters near the mouth of the Mississippi River

In-situ feeding habits of the copepods Temora turbinata and T. stylifera were investigated by scanning electron microscope examination of fecal pellets, the contents of which reflected copepod gut contents upon capture. Pellet contents were compared with assemblages of available phytoplankton in the water column at the times of zooplankton sampling. Samples were collected in continental shelf and slope waters of the Gulf of Mexico near the mouth of the Mississippi River. Both species ingested a wide size range and taxonomic array of phytoplankters, and to a lesser extent, other crustaceans. Fecal pellets contained primarily the remains of the phytoplankters that were most abundant in the water at times of collection. There was considerable overlap in the food items ingested by adult females of both copepod species, or two stages of T. turbinata copepodites. Thus, T. turbinata and T. stylifera are omnivores, but primarily opportunistic herbivores.     

Omnivorous feeding behavior of the Antarctic krill Euphausia superba

Feeding experiments were conducted at Palmer Station from December 1985 to February 1986 to examine the potential role of copepod prey as an alternative food source for Euphausia superba. Copepod concentration, copepod size, phytoplankton concentration, the duration of krill starvation and the volume of experimental vessels were altered to determine effects on ingestion and clearance rates. Krill allowed to feed on phytoplankton and copepods in 50-litre tubs showed greatly increased feeding rates relative to animals feeding in the much smaller volumes of water traditionally used for krill-feeding studies. Clearance rates on copepods remained constant over the range of concentrations offered, but clearance rates on phytoplankton increased linearly with phytoplankton concentration. Feeding rates increased when larger copepods were offered and when krill were starved for two weeks prior to experiments. Clearance rates of krill feeding on copepods were higher than, but not correlated with, their clearance rates on phytoplankton in the same vessel. E. superba may have a distinct mechanism for capturing copepods, perhaps through mechanoreception. Although our observed clearance rate of 1055 ml krill^-1 h^-1 indicates that krill can feed very efficiently on copepod prey, such feeding would meet less than 10% of their minimum metabolic requirements at the typical copepod concentrations reported for Antarctic waters. However, substantial energy could be gained if krill fed on the patches of high copepod concentrations occasionally reported during the austral summer, or if krill and copepods were concentrated beneath the sea ice during the winter or spring months. Our results, indicating efficient feeding on zooplankton and higher clearance rates on phytoplankton than previously believed, represent a step towards balancing the energy budget of E. superba in Antarctic waters.     

Coexistence: Symbiotic sharing of feeding territories and algal food by some coral reef fishes from the Western Indian Ocean

A symbiotic relationship between a damselfish (Stegastes fasciolatus) and 2 surgeonfishes (Acanthurus lineatus and A. leucosternon) is described. The damselfish, which is about 1/10 the size of the surgeonfishes, is concentrated in, and appears to prefer to be in, the feeding areas of the surgeonfishes. There is an average of one adult damselfish per adult surgeonfish feeding area. All 3 species defend their feeding areas against conspecifics and various other fishes that have similar diets, but there are few aggressive interactions between the cohabitants. Most of the interspecific defense of cohabited areas is provided by the surgeonfishes. The cohabitant species eat the same types of benthic microalgae, but only the damselfish eats microcrustaceans. It appears that the net cost to a surgeonfish of having a damselfish in its feeding area is very low because (1) biomass density of the damselfish is low, (2) it uses some food that the surgeonfishes do not use, and (3) it makes a small contribution to the defense of shared feeding areas. Even if there is a cost to the surgeonfish, the small size of the damselfish would allow it to take shelter from the surgeonfish, thus making it too costly for the surgeonfish to exclude it. Such cohabitation relationships, which may be quite common among reef fishes, represent a means by which the coexistence of species that use the same limiting resources is achieved.     

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.     

Food selection by laboratory-reared larvae of the scaled sardine Harengula pensacolae (Pisces,Clupeidae) and the bay anchovy Anchoa mitchilli (Pisces,Engraulidae)

Food selection by laboratory-reared larvae of scaled sardines Harengula pensacolae, and bay anchovies Anchoa mitchilli, was compared. Natural plankton was fed to the larvae during the 22 days following hatching. Food levels in the rearing tanks were maintained at an average of 1,600 to 1,800 potential food organisms per liter. Larvae of both species selected as food copepod nauplii, copepodites, and copepods; initial feeding was on organisms of 50 to 75 body width. Larvae of H. pensacolae averaged 4.2 mm in total length at hatching and those of A. mitchilli about 2.0 mm. H. pensacolae larvae grew about 1.0 mm per day and A. mitchilli 0.70 mm per day. The mean number of food organisms in each digestive tract was greater in H. pensacolae than in A. mitchilli, and the difference in number increased as the larvae grew. Average size of food organisms eaten increased for both species with growth, because of selection by the larvae; the average size of copepodites and copepods in digestive tracts increased at a faste rate in H. pensacolae than A. mitchilli. A. mitchilli longer than 8 mm did not eat copepod nauplii.Contribution No. 170, Bureau of Commercial Fisheries Tropical Atlantic Biological laboratory, Miami, Florida 33149, USA.     

Fate of domoic acid ingested by the copepod <Emphasis Type="Italic">Acartia clausi</Emphasis>

Two important issues in the studies of harmful algae include ecological role of the toxic compounds and their fate through the food web. The aims of this study were to determine whether the production of domoic acid is a strategy evolved to avoid predation and the role of copepods in the fate of this toxic compound through the food web. The copepod Acartia clausi was fed with single and mixed cultures of the toxic diatom Pseudo-nitzschia multiseries and the non-toxic diatom Pseudo-nitzschia delicatissima. Ingestion rate as a function of diatom abundance was the same for the toxic and non-toxic Pseudo-nitzschia species, indicating no selective feeding behaviour against P. multiseries. The toxins ingested by the copepods did not affect mortality, feeding behaviour, egg production and egg hatching of the copepods. Copepods assimilated the 4.8% of the total domoic acid ingested. Although the amount of toxins daily detoxificated by the copepods was 63.6%, the copepods accumulated domoic acid in their tissues. We conclude that domoic acid is not toxic for copepods and, probably for this reason, this toxin does not act as feeding deterrent for copepods. However, even though the production of domoic acid has apparently not evolved to deter predation, copepods may play an important role on the fate of this toxic compound through the marine food web.     

Phytoplankton pigments in the gut concents of planktonic copepods from coastal waters off southern California

Phytoplankton xanthophylls in the gut contents of the copepods Calanus pacificus, Corycaeus anglicus, and Paracalanus parvus, collected from 5 stations off San Onofre, California, in June 1982, were measured by reverse phase, high-performance liquid chromatography (HPLC). The dinoflagellate pigment, peridinin, was usually the most abundant xanthophyll in the guts of all three species of copepods. Evidently, feeding was principally on dinoflagellates (which dominated the phytoplankton biomass). The level of feeding activity, rather than the class of phytoplankton ingested, seemed to differentiate the behaviors of the copepods. Xanthophyll content per unit copepod wet weight was higher in Corycaeus anglicus and Paracalanus parvus than in Calanus pacificus. Chlorophyll a fluorescence of the copepod gut contents was measured in conjunction with the analysis of gut xanthophylls. The xanthophyll content of the gut varied directly with the concentration of chlorophyll a in the gut. Xanthophyll content was not related to the concentration of pheopigments in the gut. Apparently, the xanthophylls that were detected were due to the presence of recently ingested phytoplankton biomass.     

Fatty acid signature analysis documents the diet of five myctophid fish from the Southern Ocean

Fatty acid (FA) and fatty alcohol (FAlc) compositions of both total lipid and neutral lipid fractions were studied for five myctophid species sampled in Kerguelen waters. Both qualitative and quantitative FA signature analyses were then performed to investigate their diet over longer time scales than the conventional stomach content analysis. Regarding their lipid class, FA and FAlc compositions, the five species could be discriminated into two groups: wax-ester-rich species (Electrona antarctica, Krefftichthys anderssoni) characterised by large amounts of monounsaturated FAs (>73% of total FAs) and triacylglycerol-rich species (Electrona carlsbergi, Gymnoscopelus nicholsi, Protomyctophum bolini) with major amounts of saturated and monounsaturated FAs (>29 and >46% of total FAs, respectively). Qualitative and quantitative FA analyses showed that K. anderssoni mainly preyed upon copepods, E. antarctica upon copepods and more euphausiids and P. bolini and E. carlsbergi mainly upon euphausiids with some copepods, while G. nicholsi had a more diverse diet. This study shows the usefulness of quantitative statistical analysis to determine the diet of Antarctic and sub-Antarctic predators and stresses the need of increasing the lipid and FA analyses of more zooplanktonic and micro-nektonic marine species.     

Spatial and temporal feeding patterns of copepods from the North Pacific centra gyre

Spatial and temporal feeding patterns (determined from an index of gut fullness) are described for 10 typical species of calanoid copepods collected from the North Pacific central gyre (September 1968 to June 1977), an area where the zooplankton is food limited and there were a-priori reasons to suspect that feeding and competition for food were important in regulating zooplankton community structure. Over 100 samples from 11 cruises to the eastern part of the gyre were examined, and patterns of gut fullness were related to environmental variables and the copepod species structure. The copepods studied all tended to be omnivores and food generalists. Males had lower indices of gut fullness than females but both males and females of a species had similar spatial and temporal feeding patterns. Guts were usually fuller at night than during the day, even in nonmigrating species; however, within nighttime depth distributions, no depths were preferred for feeding. There were also differences between species in mean gut fullness, but different species tended to have similar spatial and temporal feeding patterns. There was considerable spatial variability, and locales could be identified in which most species had higher indices of gut fullness. The copepods were not necessarily more abundant in these locales, nor did these tend to be areas of above average chlorophyll concentration. These patterns were consistent with relatively nonselective feeding, and there was no evidence that these species separate their niches by feeding at differing places or times.     

Flux of cadmium through euphausiids

Flux of the heavy metal cadmium through the euphausiid Meganyctiphanes norvegica was examined. Radiotracer experiments showed that cadmium can be accumulated either directly from water or through the food chain. When comparing equilibrium cadmium concentration factors based on stable element measurements with those obtained from radiotracer experiments, it is evident that exchange between cadmium in the water and that in euphausiid tissue is a relatively slow process, indicating that, in the long term, ingestion of cadmium will probably be the more important route for the accumulation of this metal. Approximately 10% of cadmium ingested by euphausiids was incorporated into internal tissues when the food source was radioactive Artemia. After 1 month cadmium, accumulated directly from water, was found to be most concentrated in the viscera with lesser amounts in eyes, exoskeleton and muscle, respectively. Use of a simple model, based on the assumption that cadmium taken in by the organism must equal cadmium released plus that accumulated in tissue, allowed assessment of the relative importance of various metabolic parameters in controlling the cadmium flux through euphausiids. Fecal pellets, due to their relatively high rate of production and high cadmium content, accounted for 84% of the total cadmium flux through M. norvegica. Comparisons of stable cadmium concentrations in natural euphausiid food and the organism's resultant fecal pellets indicate that the cadmium concentration in ingested material was increased nearly 5-fold during its passage through the euphausiid. From comparisons of all routes by which cadmium can be released from M. norvegica to the water column, it is concluded that fecal pellet deposition represents the principal mechanism effecting the downward vertical transport of cadmium by this species.     

Accumulation of paralytic shellfish poisoning toxins in planktonic copepods during a bloom of the toxic dinoflagellate<Emphasis Type="Italic"> Alexandrium tamarense</Emphasis> in Hiroshima Bay,western Japan

Concentrations of paralytic shellfish poisoning (PSP) toxins in toxic dinoflagellate cells and in marine planktonic copepods were monitored during the bloom of Alexandrium tamarense in Hiroshima Bay, western Japan. Concentration of the toxins retained by copepods was a function of the ambient toxin concentration, i.e. the product of A. tamarense cell density and cellular toxicity. The toxin concentration in copepods increased with the increase of toxicants in the seawater then leveled off, but decreased significantly at higher concentrations. In the field, the maximum toxin concentration was 1.2 pmol ind^-1, whereas in the laboratory, the copepod Acartia omorii accumulated a much higher concentration of PSP toxins (24 pmol ind^-1). Feeding avoidance against Alexandrium tamarense and a shift to alternative food sources such as diatoms in the field might keep their toxin levels lower than their potentially maximum level. The copepod toxin levels in the field were not so high as to cause an instantaneous lethal effect on their predator fishes but may reach possibly lethal levels after a few days' continuous feeding. Overall toxin retention by copepods after 12 h feeding and 2 h starvation was only 2.5% of total ingested toxins, which suggested that a significant amount of toxins was released into the seawater. Measurements of toxin reduction and gut evacuation suggested that the toxins were removed through both fecal evacuation and metabolism (e.g. excretion, decomposition and transformation). The results, as a whole, imply that copepods can be a link for PSP toxin flux in both pelagic and benthic food webs and can also be a sink for toxins by metabolizing and removing them from the environment.Communicated by T. Ikeda, Hakodate     

Recherches sur la situation trophique d'un groupe d'organismes pélagiques (Euphausiacea). VI. Conclusions sur le rôle des euphausiacés dans les circuits trophiques de l'océan Pacifique intertropical

The role of euphausiids in the food webs of the Intertropical Pacific Ocean is defined through analysis of their nutrition, vertical distributions and migrations, and their utilization by pelagic predators. It is suggested that the abundance of the group, the extensive vertical migrations of many species and the fact that feeding takes place mainly in subsurface layers, result in a leading role of euphausiids in energy transfer between different bathymetric levels. For night-time feeding predators, they represent a noticeable food source only in the 0 to 300 m water layer, as 97% of the euphausiid biomass concentrates in this layer at night. In the daytime, only the smaller specimens (chiefly genus Stylocheiron), accounting for 10 to 15% of the whole biomass of the group, remain available for epipelagic (0 to 400 m) predators, larger individuals dwelling deeper. Euphausiids account for 8 to 10% of the food ingested by micronektonic fishes, but the species are not the same for different categories of fishes. Migrating fishes caught by pelagic trawls, more or less connected with the deep scattering layer, feed on migrating species in subsurface layers at night as well as in deeper layers during the daytime, and on non-migrating species inhabiting shallower and intermediate layers. On the other hand, fishes which comprise the prey of large long-line tunas, which are not caught by trawls because they are fast swimmers, feed almost solely on species which remain above 400 m in the daytime. These results suggest a certain degree of independance between the trophic webs which concern, on the one hand, epipelagic ichthyofauna (including tuna), and, on the other hand, migrating and deep-living faunas. Migrating populations are able to feed at night upon subsurface organisms, a part of this resource being then transmitted during the day to the deep-living fauna; but the epipelagic ichthyofauna, with a feeding activity restricted to light hours, has few possibilities to benefit from the migrating or deepliving biomass. Therefore, energy transfers seem to be intense only from subsurface (0 to 400 m) to deeper layers. From a more general point of view, these investigations suggest that, in the pelagic system, vertical distributions and migrations, and feeding rhythms, are the main factors determining the structure of the food webs.