Feeding habits of the spotted flounder Citharus linguatula off the eastern coast of Spain

The composition of the diet of Citharus linguatula (L.) off the coast of the Gulf of Valencia, Spain, was determined between October 1989 and October 1990. The percentage of empty stomachs remained constant throughout the year, except for the period August to September, when a maximum was recorded coinciding with the reproductive period. Crustaceans (Mysidacea and Decapoda) and teleosts constituted the main prey. The composition of the prey ingested varied with predator size; small specimens contained a greater number of mysids in their stomachs, while decapods and fishes were more abundant in the stomachs of larger specimens. Diets varied seasonally: mysids were more important during autumn, whereas decapods were more important during winter and spring. Fishes were numerically most important in the diet in summer, but the frequency of occurrence was constant throughout the year.     

Food and feeding dynamics of the larval Antarctic fishNototheniops larseni

The feeding dynamics of larvae of the Antarctic fishNototheniops larseni were analyzed from data collected over three years in Bransfield Strait and adjacent waters (Antarctica). Seasonal feeding was examined from 1977/1978 (November–March). The diel feeding cycle was investigated during a 96 h station established in February 1976, while food selection was analyzed using larvae and zooplankton samples collected in February 1982. Hatching occurs in early spring, and larvae fed on eggs of calanoid copepods and on cyclopoid copepods. Copepod eggs were the principal food near the pack ice, and cyclopoids in open waters. Cyclopoids were the staple food in summer. Eggs of the Antarctic krillEuphausia superba were ingested selectively and formed major portions of the larval summer diet in neritic (Joinville Island) and oceanic (Elephant Island) spawning areas ofE. superba. In the fall, copepods predominated in the diets. Most abundant and most frequently ingested prey in summer and fall wereOncaea spp. Feeding commenced at dawn and continued at least until dusk. Krill eggs were taken chiefly during morning hours and egg incidence declined during the day, suggesting that eggs were ingested soon after spawning. Prey size at the onset of feeding was estimated as 0.130 to 0.330 mm. Size-selective feeding was evident in small larvae, while in larger larvae median prey length remained constant. High feeding incidence among yolk-sac larvae in spring, high overall feeding incidence in summer, and size-selective foraging of small larvae suggested favorable feeding conditions in the 1977/1978 season. Yolk-absorption times in Antarctic fish larvae vary on a scale of weeks and may be further retarded due to early feeding. Hence, year-to-year variability of yolk incidence inN. larseni indicated variable biotic environments of early feeding larvae rather than temporal shifts of hatching periods. As hatching periods are constant between years in contrast to the variable retreat of the pack ice and subsequent onset of the production cycle in space and time, maternal yolk reserves are probably utilized to compensate for such variations.     

Feeding habits of the Madeira rockfish <Emphasis Type="Italic">Scorpaena maderensis</Emphasis> from central Mediterranean Sea

In order to assess diet composition and niche breadth of this species, we analysed the stomach content of 182 specimens collected monthly along the eastern coast of Sicily (Central Mediterranean Sea). Overall, 50 prey taxa belonging to five major groups (algae, gastropods, crustaceans, polychaetes, fishes) were identified in 102 full stomachs. Benthic or epibenthic crustaceans, such as decapods, amphipods and isopods were the most important prey, whereas algae, gastropods, polychaetes and fishes were only occasionally ingested. In terms of composition by species, the diet of Scorpaena maderensis was characterized by a variety of rare or unimportant prey, which was consumed by few individuals only, although sometimes in large amount. As a result, S. maderensis can be considered a generalized and opportunistic feeder. The feeding intensity followed roughly a seasonal trend, with a minimum food intake in summer. The individual fish size was the most important factor affecting diet. According to the observed ontogenetic shift, small-sized individuals fed primarily on small crustaceans (i.e. amphipods and isopods), whereas large-sized specimens consumed preferably bigger and more vagile prey, such as walking and swimming decapods. No significant differences in diet were observed in relation to sex of predator and sampling season.     

Feeding selectivity in larvae of the European hake (<Emphasis Type="Italic">Merluccius merluccius</Emphasis>) in relation to ontogeny and visual capabilities

Feeding ecology was analysed for the first time in the larvae of the European hake (Merluccius merluccius) to determine whether their diet and selectivity were constrained by environmental conditions and how these feeding characteristics were related to ontogeny, prey availability and visual capabilities. Larvae collected during both day and night were analysed, and it was found that feeding incidence was high, regardless of the time of day. Examination of the visual system corroborated the hypothesis that hake larvae should be able to cope with a wide range of photic conditions and to forage even at low light intensity. A clear preference for adult calanoid copepods and, especially, for Clausocalanus spp. was observed in all sizes analysed. Prey number increased with larval size, but prey size did not. This finding indicates that hake larvae behave as selective and specialist predators that consume an increasing number of prey rather than larger prey during larval growth.     

Temporal changes in feeding habits and daily rations of<Emphasis Type="Italic"> Hoplostethus mediterraneus</Emphasis> in the bathyal Ionian Sea (eastern Mediterranean)

Data on the diet, feeding habits and daily rations of Hoplostethus mediterraneus Cuvier, 1829 in the bathyal eastern Ionian Sea (Mediterranean Sea) are presented. A total of 430 specimens collected by bottom trawls at depths ranging from 473 to 603 m during four 24-h day–night sampling cycles covering the four annual seasons was examined. H. mediterraneus diet consisted of pelagic and vagile epibenthic prey, mainly crustaceans, and was dominated by benthopelagic natantian decapods (83.35% IRI, index of relative importance). Seasonal changes in diet were apparent and related to seasonal fluctuations in suprabenthic and zooplanktonic prey in the environment. Diel patterns in stomach fullness and trends in diel feeding cycles are discussed in relation to the vertical migratory movements of available prey (i.e. suprabenthos and zooplankton). Daily-ration estimates were determined by evacuation-rate models and ranged from 0.143% to 0.397% WW/WW. Overall, daily-ration estimates were within the range of the daily consumption of other deep-sea fish. Deduced from diet contents, we found a constant gross energy intake (305–316 kcal g^–1) during all seasons. As a possible response to the reproductive peak of mature females observed in summer, H. mediterraneus increases its food consumption, which, in turn, is coupled with an increase in food availability.Communicated by S.A. Poulet, Roscoff     

Observations on the nocturnal feeding of some mesopelagic decapod crustacea

Feeding in relation to temporal changes in the depth distribution of predator and prey is described for 9 species of mesopelagic decapods from an examination of 268 foreguts. Intensive nighttime feeding appears to be the rule in all species. The smaller decapods Sergestes (Sergestes) atlanticus, Sergestes (Sergestes) sargassi and Sergestes (Sergestes) pectinatus exploit the smaller prey, principally copepods and to a lesser extent ostracods. Larger decapod species Sergestes (Sergestes) henseni, Sergestes (Sergestes) curvatus, Sergestes (Sergia) grandis, Systellaspis debilis, and Acanthephyra purpurea mainly prey on macrozooplankton and micronekton, i.e., chaetognaths, euphausiids, decapods and fish, but copepods also occur in the foreguts. Gennadas valens is exceptional for the high incidence of foraminiferal remains, and a predator-prey relationship seems probable. All 9 decapod species have mixed diets, and pronounced feeding preferences are not evident. However, a high incidence of “secondary” feeding or “dietary contamination” has been deduced from the frequent occurrence of remains of the copepods Pleuromamma spp. and Oncaea spp. in the foreguts of the larger decapod species. Direct feeding cannot have occurred, since the depth distributions of these copepods and decapods are disjunct by day and night. It is concluded that the remains of Pleuromamma probably represent the food of the larger prey such as chaetognaths etc. which are eaten by the decapods. The presence of Oncaea is speculatively attributed to a possible ectoparasitic relationship with the larger prey items, but confirmatory evidence is required. These anomalies suggest that caution must be exercised in deducing predator-prey relationships simply from gut contents without consideration of distributional factors.     

Feeding habits of pasiphaeid shrimps close to the bottom on the Western Mediterranean slope

The composition of the diet and daily cicle of predatory activity of pasiphaeid shrimps in the Northwestern Mediterranean were established; special attention was focussed on nocturnal feeding habits close to the bottom. Daily activity in both species was studied in two continuous 24-h sampling periods. Samples were obtained using bottom trawls between 1988 and 1990. Both species fed on benthic prey items at night. The nocturnal diet of Pasiphaea multidentata consisted of gammarid amphipods, isopods (Cirolana borealis) and macruran decapods (Calocaris macandreae). Nocturnal feeding activity was carried out only by large specimens (cephalothorax length >28 mm), which stayed close to the bottom during the nighttime. In contrast, the specimens collected during the daytime exhibited highly digested remains of pelagic prey (hyperiids, fishes, euphausiids, chaetognaths) ingested as a result of predatory higher activity in the water column the night before. The feeding strategy of P. sivado was parallel to that of P. multidentata. At night large specimens were located near the bottom and fed on suprabenthic gammarid amphipods. This nocturnal feeding activity by both these mesopelagic pasiphaeid species furnishes evidence of energy transfer from the benthos to the planktonic system in bathyal communities. Changes were observed in the diet of P. multidentata with depth. Crustaceans made up a larger share of the diet of P. multidentata on the lower slope than on the upper middle slope, probably because of changes undergone by bathyal communities with increasing depth. The feeding rate was higher in the submarine canyons, where the diet is also more specialised. Dietary overlap between the two pasiphaeids was very low, due to the different size range of prey exploited.     

Cercopagis pengoi and Mysis spp. alter their feeding rate and prey selection under predation risk of herring (Clupea harengus membras)

The anti-predator behaviour of Baltic crustacean planktivores was studied in feeding experiments under predation pressure of herring. The experiments were conducted with pelagic mysids: Mysis mixta and Mysis relicta, and with Cercopagis pengoi, a non-indigenous cladoceran, which invaded the Baltic Sea in 1992. Zooplankton was offered as prey. Two kinds of experiments were performed in the absence and presence of chemical predator cues: (1) two-prey experiments with prey, which have poor or good escape responses and all three planktivores and (2) natural prey experiments with mysids in natural zooplankton assemblages. The results showed that all three species reacted to the chemical cue of herring by decreasing their feeding rate and altering prey selection. C. pengoi selected easily captured prey (rotifers) in two-prey experiments under predation risk while selection for any prey was evident in mysids in natural prey experiments only in the absence of predator cues. This indicates that planktivores have different anti-predator strategies, which are modified by their own prey capture abilities. C. pengoi was a very efficient predator on small prey with size-specific prey consumption rate 5 to 18 times the rate of mysids. Results show that the studied planktivores are capable of adjusting their feeding behaviour to decrease their conspicuousness in order to increase survival under predation risk. Further, results support the view that C. pengoi has adapted well to the Baltic ecosystem, sharing food niche with pelagic mysids and most probably having a strong influence on the whole pelagic food web.     

Degradation of mangrove leaf litter by the tropical sesarmid crab Chiromanthes onychophorum

The food and feeding habits of 3 species of gadoid larvae — the cod Gadus morhua Linnaeus, 1758, the whiting Merlangius merlangus (Linnaeus, 1758), and the bib Trisopterus luscus (Linnaeus, 1758), collected in the eastern English Channel and Southern Bight during the spring of 1971 are described. All 3 species began to feed in the yolk-sac stage on diatoms, dinoflagellates and tintinnids, but the principal food was the nauplii and copepodites of calanoid copepods, particularly of Pseudocalanus minutes, but also of Paracalanus parvus, Temora longicornis and Acartia clausii. Pseudocalanus minutus and Paracalanus parvus were eaten mainly early in the season and T. longicornis later when it became more abundant. The larvae discriminated for prey size as growth proceeded. They sometimes took the largest prey available to them, but in general the size of the prey was considerably less than the maximum size which could have been swallowed. Feeding larvae were found at all times of the day, but the incidence of feeding was lowest before dawn. Feeding increased at sunrise, declined until late in the morning, and then increased again to a maximum around sunset. There was evidence of feeding by moonlight, particularly by whiting and bib larvae. There was little difference between the English Channel and Southern Bight in regard to the food eaten.     

Locomotor activity patterns and feeding habits in the prawn Palaemon xiphias (Crustacea: Decapoda: Palaemonidae) in Alfacs Bay,Ebro Delta (northwest Mediterranean)

Diel activity rhythms of the prawn Palaemon xiphias Risso, 1818 from Alfacs Bay, Ebro Delta, were studied by time-lapse videorecordings. Activity displayed an endogenous circadian rhythm, with maximum activity at night. Feeding habits were studied by frequency of occurrence and by the points method. The food of P. xiphias consisted mainly of crustaceans; the remains of amphipods, isopods, mysids, copepods, decapods and ostracods were identified. The remaining items consisted of molluscs, polychaetes, ophiuroids, plant material, sand, and unidentified organic debris. The results indicate that P. xiphias is a predator of benthic invertebrates rather than a scavenger or detritus feeder. Diet composition changes with increasing size of the prawn.     

Diets of deep-sea brachyuran crabs in the Western Mediterranean Sea

This study examines the feeding habits of Paromola cuvieri (Risso, 1816) and Geryon longipes A. Milne Edwards, 1881, the only two common deep-sea brachyuran crabs inhabiting the bathyal mud assemblages in the Catalan Sea (Western Mediterranean). Samples were obtained by bottom trawls at depths between 360 and 1871 m during 1983 to 1992. Both species had highly diverse diets, but very low feeding activity, as reflected by the high proportion of empty stomachs. Both characteristics may be important factors enabling deepsea crabs to adapt to bathyal zones, where trophic resources are scarce. The most important food items found in P. cuvieri were fish remains (teleost, sharks) and benthic decapods (Monodaeus couchii, Munida tenuimana). Scavenging activity plays an important role in this species. The diet of G. longipes included a broad range of benthic invertebrates. In the upper middle slope, the bivalve Abra longicallus, decapods (Calocaris macandreae and Monodaeus couchii), echinoderms and polychaetes were the dominant prey, with epibenthic peracarids as a secondary resource. On the lower middle slope, the incidence of decapod crustaceans (C. macandreae, Pontiphilus norvegicus) and peracarids in the diet declined. Small macrobenthic prey (glycerids, cumaceans or amphipods) were rare in the diet of both species, in accordance with the large size of the crab specimens studied. The absence of preferred prey items and the lack of food items of an optimum size on the lower slope may contribute to the progressive decline in abundance of P. cuvieri and G. longipes with increasing depth.     

Predatory feeding of two marine mysids

Predatory feeding of the marine mysids Mysidopsis bigelowi and Neomysis americana on several species of co-occurring copepods was examined in laboratory experiments. M. bigelowi exhibited a curvilinear functional response; there was a negative logarithmic relationship between prey density and clearance rates. N. americana also exhibited higher clearance rates at lower prey densities. Increased clearance rates at lower prey densities were probably due to increased swimming speed or reaction distance as hunger increased. This response occurred only when mysids could visually locate prey; in complete darkness clearance rates were significantly lower and independent of prey density. Feeding rates on different prey species were only partially dependent on prey size; prey movement patterns and escape behavior also strongly affected feeding rates. M. bigelowi showed active prey selection when offered a choice of different prey species. Estimates of predation rates of estuarine mysid populations indicate that they could have a significant effect on co-occurring copepod populations.     

Predation by the epipelagic heteropod mollusk Carinaria cristata forma japonica

In surface waters off Southern California (USA), Carinaria cristata forma japonica van der Spoel, 1972 feeds on a variety of zooplankton, although thaliaceans, chaetognaths, and copepods predominate numerically in the diet. Feeding intensity is greatest on the most abundant of two species of thaliaceans, depending on which one dominates in the plankton at the time. Some cannibalism occurs, with the prey being about one half the size of the predator. Feeding intensity is greatest during the day, possibly because heteropods depend on vision to locate prey and because prey species are more available by day. Comparisons of the proportion of each prey species in the diet and in the plankton indicate preferential feeding on thaliaceans, chaetognaths, and mollusks; in contrast, crustaceans and especially the copepods are non-preferred prey. These preference patterns may reflect differences among prey species in the ability to escape capture. Predator and prey size are positively correlated for Doliolum denticulatum gonozoids and oozoids, Thalia democratica aggregates, and Sagitta spp. Smaller individuals of D. denticulatum gonozoids and Sagitta spp. are selectively preyed on, resulting in size refuges for larger individuals.     

Tidal and diurnal influence on food consumption of a salt marsh killifish Fundulus heteroclitus

Feeding patterns during four 24-h periods, sampled at 3-h intervals, were investigated for the mummichog Fundulus heteroclitus, in a Delaware, USA tidal marsh. Two factors potentially influencing feeding patterns, time of day and tide height, were examined. On 2 of the sampling periods a low tide occurred in the morning, while on the other 2 sampling periods a high tide occurred in the morning. Results are reported as g-dry wt. of food per g-dry wt. of fish. F. heteroclitus is primarily a daytime feeder that most actively feeds at high tide, regardless of whether or not the high tide inundates marsh surface areas. When tide height was sufficient to inundate the marsh surface, fish invaded these areas and consumed prey characteristic of the marsh surface. F. heteroclitus is an important link in energy transfers between the marsh surface and subtidal systems, enhancing its own energy supplies by consuming marsh surface prey whenever available.     

Spatio-temporal variability in fatty acid trophic biomarkers in stomach contents and muscle of Iberian sardine (<Emphasis Type="Italic">Sardina pilchardus</Emphasis>) and its relationship with spawning

Temporal variation in the fatty acid (FA) composition of stomach contents of Iberian sardines was compared to the relative contribution to dietary carbon made by different prey types for fish from two areas off Portugal. The effect of the FA content of the diet on sardine muscle FA composition was also studied, aiming at (1) analysing if FA biomarkers can be used as a complementary technique for the study of sardine diet and (2) to relate spatial and temporal variations of prey FA content with sardine condition and reproduction. Significant spatial differences in the FA composition of sardine diet occurred with concentrations of n-3 polyunsaturated FA, namely eicosapentaenoic acid [EPA, 20:5n-3] and linolenic acid 18:3n-3, being significantly higher in the diet of sardines from the west coast, whilst the diet of sardines from the south coast was richer in monounsaturated fatty acids (MUFA), namely the carnivory biomarker oleic acid 18:1n-9. These results are in agreement with the higher contribution made by diatoms and dinoflagellates to the diet of sardines off the west coast. Spatial variation in sardine dietary FA was also detected in their muscle composition, specifically for EPA, and the eicosapentaenoic/docosahexaenoic acid and (n-3)/(n-6) ratios, which were higher in sardines from the west coast. No difference in FA composition was detected between sexes, and the seasonal variability in sardine total FA concentration was primarily related to the seasonality of spawning. Sardines accumulate high concentrations of FAs during the resting stage of reproduction when the feeding intensity is similar or lower to that observed during the spawning season. Additionally, sardines show a high selective retention of MUFA and polyunsaturated FA (PUFA) throughout the year except at the beginning of the spawning season, when these FAs are largely invested in the formation of the gonads. Therefore, temporal and regional differences of prey environments are strong enough to be reflected in fish body composition, namely on the accumulation of essential FAs, which can have a strong impact on reproduction success for this species.     

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     

Diet of orange roughy,Hoplostethus atlanticus (Pisces: Trachichthyidae) on the Challenger Plateau,New Zealand

Stomach contents of 687 orange roughy [Hoplostethus atlanticus (Collett, 1889)] from the Challenger Plateau, sampled in March, July and November 1984, were examined. The relative importance of different components of the diet was assessed using three feeding indices which combine, in different ways, percentage frequency of occurrence, percentage number, and percentage weight of prey categories. For both sexes, for all size classes and at any time of the year, the most frequent and abundant components of the diet were prawns, followed by fish, squid, amphipods and mysids. The main natant decapod families were Oplophoridae, Pasiphaeidae, and Sergestidae. Fish, mainly Chauliodontidae and Myctophidae, were the most important prey by weight. Most prey species were benthopelagic and mesopelagic organisms which move towards the surface at night. However, H. atlanticus can be caught by bottom trawl (between 750 and 1 200 m) during any 24 h period, and there was no evidence of vertical migration in search of their prey. When the fish grow in length, there is a transition in the diet from prawns, mysids, and fish, to prawns, fish and squid. Squid were not found in the stomach contents of fish smaller than 20 cm. These dietary changes may be linked to modifications in morphology with growth. The relative proportions of the main dietary items, and in particular the natant decapod families, varied with time of the year. Also, the stomach data seem to indicate an increasing importance of fish and squid in the diet in deeper water. H. atlanticus appears to be an opportunistic predator, consuming a wide variety of invertebrates and fishes. Our results provide evidence to support the notion that benthopelagic predators which consume vertically migrating mesopelagic fish, have an important role in the transfer of energy to the benthos.     

Diet and food preferences of the adult horseshoe crab Limulus polyphemus in Delaware Bay,New Jersey,USA

Adult horseshoe crabs Limulus polyphemus (L.) feed on a wide variety of infaunal and epifaunal invertebrates during their spring spawning migration in Delaware Bay, New Jersey, USA. Comparison of the gut contents with estimates of available prey showed that the most abundant potential prey item, the bivalve Gemma gemma, was avoided. The thinner shelled but comparatively scarce clam Mulinia lateralis was a preferred prey item. In the laboratory, crabs fed on G. gemma when it was the only available item but not when M. lateralis or soft-shell clams, Mya arenaria, were offered in conjunction. Large M. lateralis (>10mm) were preferred to small M. lateralis; there was no discrimination between M. lateralis and M. arenaria of the same size. Male and female horseshoe crabs had similar gut contents and laboratory feeding preferences, despite the fact that females are larger than males. Crabs spawning later in the summer contained more food than did crabs collected at the peak of spawning activity.     

Variability of albacore (<Emphasis Type="Italic">Thunnus alalunga</Emphasis>) diet in the Northeast Atlantic and Mediterranean Sea

This study aims to describe the variability of albacore (Thunnus alalunga) diet in the Northeast Atlantic and Mediterranean Sea and to identify possible relationships between this variability and the features of different feeding areas, the behavior, and the energetic needs of albacore. Stomach contents from albacore caught in five zones of the Bay of Biscay and surrounding waters (n = 654) and three zones of the Mediterranean Sea (n = 152) were analyzed in terms of diet composition and stomach fullness. Carbon and nitrogen stable isotope and C/N ratios were measured for white muscle and liver from albacore in the Bay of Biscay (n = 41) and Mediterranean Sea (n = 60). Our results showed a spatial, seasonal, inter-annual, and size-related variability in the diet of albacore. Albacore diet varied by location in the Mediterranean Sea, with a particularly high proportion of cephalopods, and low δ¹⁵N values in the Tyrrhenian Sea. In the Northeast Atlantic, albacore consumed a higher proportion of crustaceans and a lower proportion of fishes in the most offshore sampling zone than inshore. The digestion states of the major prey reflected a diurnal feeding activity, indicative of feeding in deeper waters offshore, whereas on the continental slope, feeding probably occurred in surface waters at night. Important seasonal and inter-annual diet variability was observed in the southeast of the Bay of Biscay, where preferred albacore prey appeared to be anchovy (Engraulis encrasicolus). Stomach fullness was inversely related to body size, probably reflecting higher energetic needs for smaller individuals. Albacore from the Bay of Biscay had significantly lower δ¹³C and higher δ¹⁵N values compared with albacore from the Mediterranean Sea, indicative of regional baseline shifts, and trophic position and muscle lipid stores in albacore increased with body size.