Diet of the northern fulmar Fulmarus glacialis: reliance on commercial fisheries?

The massive expansion in breeding numbers and range of the northern fulmar Fulmarus glacialis over the last two centuries is generally attributed to an increased availability of fish offal and discarded fishes from commercial fisheries. This implies that discards should be a major component in fulmar diets in the more recently colonised areas in the south of their range. This paper examines the contemporary diet of the northern fulmar at three major breeding sites, Fair Isle (Shetland, UK), Iceland and Disko Fjord (western Greenland). At Fair Isle, 89% of regurgitates contained fishes, with sandeels (Ammodytidae) recorded in 37%; 32% contained crustaceans (mainly decapods); 8% contained squid. There was also a temporal trend; sandeels declining and crustaceans increasing in the diet from July to August. In Iceland, fishes were also the main prey (47 to 93% of wet mass), with sandeels common in the south and west, and capelin (Mallotus villosus) in the north and east. Other fish species were mainly discards, and together with discarded decapods and fish offal formed 5 to 72% of the diet, depending on the sector. Euphausiids, amphipods and copepods constituted 3 to 13% of the diet. At Disko, 39% of samples contained fishes, 64% contained crustaceans (mainly amphipods), 22% contained squid, and 16% contained pteropods. There was a clear temporal trend, with the bulk of the samples made up of crustaceans in mid-June, capelin from late June to late July, and crustaceans and pteropods from late July to late August. An extensive review of published studies was also carried out. The general pattern was for birds in more southerly populations to consume more discarded fishes, fish offal and benthic invertebrates. However, a considerable proportion of their diet also consisted of juvenile gadids, sandeels, capelin and pelagic zooplankton, which fulmars catch for themselves, and we suggest that breeding adults in the south are less dependent on fishing waste than is generally assumed. Received: 25 September 1998 / Accepted: 7 May 1999     

Inter- and intra-year variation in foraging areas of breeding kittiwakes (Rissa tridactyla)

While seabird conservation efforts have largely focused on protection from threats at the colony (e.g. reducing disturbance and predation), attention is increasingly being given to implementing protection measures for foraging areas at sea. For this to be effective, important foraging areas must be identified. Although numerous studies have examined seabird foraging behaviour, information is still lacking on the variability in area utilisation within and among breeding seasons. GPS devices were attached to adult black-legged kittiwakes breeding at an expanding North Sea colony (55°20′N, 1°32′W) during both incubation and chick-rearing in 2012 and during chick-rearing in 2011, to determine whether foraging areas remained consistent and to identify the oceanographic characteristics of areas used for foraging. The type and size of prey items consumed at different stages of the breeding cycle was also examined. During incubation (April–May 2012), kittiwakes foraged substantially further from the colony and fed on larger sandeels than when feeding chicks, and there was significant inter-annual variation in foraging areas used during the chick-rearing period (June–July 2011 and 2012). Foraging areas were characterised by cooler sea surface temperatures and areas of high chlorophyll a concentration, although association with specific oceanographic features changed within the breeding season and between years. These results emphasise the importance of considering how foraging areas and reliance on specific oceanographic conditions change over time when seeking to identify important marine areas for seabirds.     

Importance of the Japanese anchovy (Engraulis japonicus) to breeding rhinoceros auklets (Cerorhinca monocerata) on Teuri Island, Sea of Japan

Temporal variation in the diet and chick growth of rhinoceros auklets (Cerorhinca monocerata), on Teuri Island, Sea of Japan, was studied to understand how local marine environmental changes affect the reproduction of this piscivorous seabird. The food delivered by parents to chicks was sampled every 1-2 weeks from late May to July, 1994-1998. Overall, the diet of nestling rhinoceros auklets consisted of (by mass) 61% Japanese anchovy (Engraulis japonicus), 18% Japanese sand lance (Ammodytes personatus), 18% Japan Sea greenling (Pleurogrammus azonus), 2% other fish and 1% squid. Among years, the contribution of anchovy ranged from 16% to 93%. Once anchovy occurred in the diet, it dominated (80% on average) thereafter. Accordingly, when anchovy appeared in the diet early in the chick-rearing season (1994, 1998), the contribution of anchovy overall was large. The first appearance of anchovy in the diet of auklets late in the summer of 1997 was possibly related to negatively anomalous sea-surface temperature. Food loads composed of anchovy (34.0 g) were heavier than those of sand lance (22.5 g) and greenling (28.5 g). The energy density of anchovies also was higher: 6.3 kJ g^-1 wet mass compared to 0+ greenling (4.78 kJ g^-1) and 0+ sand lance (3.78 kJ g^-1). Thus, a high proportion of anchovy in the diet resulted in high food load mass, high daily growth rates of chicks and high fledging success. This study highlighted the importance of the time of arrival of migratory high-lipid prey, which is influenced by local oceanographic conditions, to the reproductive performance of a piscivorous seabird.     

Size matters: variation in the diet of chick and adult crested terns

We investigated ontogenetic, temporal and spatial patterns in the composition and size of prey in the diet of crested terns, Sterna bergii. Diet analyses indicated that crested terns are a generalist predator on surface-schooling clupeids (Australian anchovy Engraulis australis, sardine Sardinops sagax and blue sprat Spratelloides robustus), Degens leatherjacket Thamnaconus degeni, southern sea garfish Hyporhamphus melanochir, Australian herring Arripis georgianus, slender bullseye Parapriacanthus elongatus and barracouta Thyrsites atun. Ontogenetic differences in prey size indicated that adults are constrained in their foraging behaviour during the early chick-provisioning period by the need to self feed and select smaller prey that can be ingested by their chicks. Chicks consumed significantly higher proportions of clupeids than adults, which consumed mainly Degens leatherjackets and barracouta, suggesting that adults may select higher quality prey for their chicks compared to what they consume themselves. Spatial differences in prey composition were driven by differing proportions of sardine, Australian anchovy and Degens leatherjacket and could reflect local differences in the abundances of these prey. The size of prey taxa consumed by adults also reflected a North–South gradient in prey size. The large component of juvenile sardine in the diet of crested terns suggests future dietary measures may inform fisheries managers about changes in local juvenile sardine abundance. These data could assist in highlighting any fishery-related decreases in sardine recruitment and help ensure commercial fishing practices address principals of Ecologically Sustainable Development developed for Australian fisheries.     

Flexible foraging strategies of gentoo penguins <Emphasis Type="Italic">Pygoscelis papua</Emphasis> over 5 years in the South Shetland Islands,Antarctica

Gentoo penguins Pygoscelis papua show considerable plasticity in their diet, diving, and foraging behaviors among colonies; we expected that they might exhibit similar variability over time, at a single site, since flexible foraging habits would provide a buffer against changes in prey availability. We examined interannual changes in the foraging strategies and diet of gentoo penguins in the South Shetland Islands, Antarctica, over 5 years with variable prey abundance. Antarctic krill Euphausia superba was the primary diet item, and fish the secondary, though the importance of these items varied among years. Diving behavior also varied over time: different dive depth distributions were observed among years. Nonetheless, chick-rearing success remained relatively constant, indicating that gentoo penguins were able to maintain chick provisioning by altering their foraging strategy among years. Variable abundance of krill in the region did not have observable impacts on the diet, foraging behaviors or chick-rearing success of gentoo penguins. We suggest that foraging plasticity may be one reason that gentoo penguin populations have remained stable in the region, while their congeners (P. antarctica and P. adeliae) with less flexible foraging strategies have declined.     

Seasonal changes in the diet of a critically endangered seabird and the importance of trawling discards

Pelagic seabirds obtain food from oceans where the availability of their prey changes rapidly both seasonally and spatially. Here, we investigated changes in the trophic habits of the critically endangered Balearic shearwater (Puffinus mauretanicus) through the breeding season and tested for dietary differences between sexes and age classes. We analysed δ¹⁵N and δ¹³C values in blood of adults during the pre-incubation, incubation and chick-rearing periods and of their chicks. Using a two-isotope mixing model, we estimated dietary contributions based on isotope values from potential prey species which included small pelagic species available naturally and demersal fish species available only from trawling discards. Balearic shearwaters showed clear isotopic and dietary variation through the breeding season. During pre-incubation, breeding adults appeared to exploit demersal fish, whereas during the incubation and chick-rearing period, they fed mainly on pelagic anchovies (Engraulis encrasicolus) and pilchards (Sardina pilchardus). Similarly, chicks were fed mainly with anchovies, a resource with a high energetic value. This variation in the dietary habits of adult shearwaters during the breeding season was probably related to both natural and fishery-induced seasonal changes in the availability of potential prey species within their main feeding grounds. However, changes in the nutritional requirements of the shearwaters could also play an important role. Indeed, diet differed between sexes during pre-incubation: females fed less on trawling discards and more on small pelagic fish than males. This sexual segregation in diet could be the consequence of higher nutritional requirements of females during this period. Our study reveals the differential importance of both trawling discards and small pelagic fish species for a pelagic seabird depending on the breeding period and illustrates the importance of considering the entire breeding season when making inferences about the importance of specific prey in seabird dietary studies.     

Trophic ecology of Grey-headed albatrosses from Marion Island, Southern Ocean: insights from stomach contents and diet tracers

During chick-rearing, albatrosses can alternate between long foraging trips that provide the main source of food for the adults and short foraging trips that they use to feed their young. This flexibility in foraging behaviour can lead to differences in diet composition between adults and chicks and implies that they may be vulnerable in different ways to food shortages. The trophic ecology of the Grey-headed albatross Thalassarche chrysostoma was investigated at the sub-Antarctic Prince Edward Islands during the chick-rearing period in April 2006 using a combination of approaches. Diets of adults and chicks were assessed using stable isotope ratios and fatty acid (FA) profiles of blood and/or stomach oils, in addition to stomach contents analysis. Fish from the family Macrouridae and cephalopods (particularly the onychoteuthid Kondakovia longimana) were the primary prey, whereas crustaceans (krill Euphausia superba) represented a smaller proportion of the stomach contents. Stomach oil FA profiles contained more monounsaturated FA than the profiles of plasma, which were richer in saturated FA and arachidonic acid (20:4n-6). There was also a distinct separation of adults from chicks, with higher levels of monounsaturates in chick plasma, and higher saturated FA levels (particularly 16:0) in the adult plasma. Stable carbon isotope ratios of whole blood were similar in adults and chicks, whereas stable nitrogen isotope ratios showed significant enrichment by >1‰ in chicks. The combined FA, stable isotopes and stomach contents analyses suggest clear differences in diet quality between adults and chicks, with chicks feeding at a higher trophic position through feeding more on highly nutritious fish and adults keeping much of the less nutritious zooplankton for themselves.     

Do activity costs determine foraging tactics for an arctic seabird?

How energy costs affect foraging decisions is poorly understood for marine animals. To provide data relevant to this topic, we examined the relationship between activity levels and foraging behavior by attaching activity recorders to 29 chick-rearing wing-propelled diving birds (thick-billed murres, Uria lomvia) in 1999–2000. We connected the activity during the final dive bout with the prey item we observed being fed to the chicks. After accounting for changes in activity level with depth, activity was highest during the final dive of a dive bout, reflecting maneuvring during prey capture. Pelagic prey items, especially invertebrates (amphipods), were associated with higher depth-corrected activity, leading to shorter dives for a given depth (presumably due to higher oxygen consumption rates) and, thus, shorter search times (lower bottom time for a given depth). Pelagic prey items were likely captured during active pursuit, with the birds actively seeking and pursuing schooling mid-water prey. In contrast, benthic prey involved low activity and extended search times, suggesting that the birds slowly glided along the bottom in search for prey hidden in the sediments or rocks. We concluded that activity levels are important in determining the foraging tactics of marine predators. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Balancing between mutualism and exploitation: the symbiotic interaction between Lasius ants and aphids

Nests of Lasius niger (L.) ants were given varied food regimens to test whether their behaviour towards an aphid partner, Aphis fabae (Scop.), changed with alternative food supplies. Honeydew collection and predation on aphids were measured by video monitoring the movement of ants between their nest and an aphid aggregation. Data collected from the aphid aggregations enabled comparisons between remaining aphid biomass and between the tending intensities of the ants. I tested how ant behaviour was influenced by their access to alternative prey and sugar. The results showed that ants accepted a honey solution as a substitute for the honeydew produced by aphids. Ants not only attended their aphid partners, but also preyed on them. The average predation rate increased eightfold when ants were offered the alternative of sugar, whereas alternative prey had no significant effect. In contrast, ant-tending intensity decreased with alternative sugar whereas alternative prey elicited no effect.     

Inter-annual changes in diet and foraging trip lengths in a small pelagic seabird, the thin-billed prion Pachyptila belcheri

Central place foragers are constrained in their foraging distribution by the necessity to return to their nest site at regular intervals. In many petrels that feed on patchily distributed prey from the sea surface over large foraging areas, alternating long and short foraging trips are used to balance the demands of the chick with the requirements of maintaining adult body condition. When the local conditions are favourable for prey density and quality, adults should be able to reduce the number of long foraging trips. We studied the flexibility in foraging trip lengths of a small pelagic petrel, the thin-billed prion Pachyptila belcheri, over three breeding seasons with increasingly favourable, cold-water conditions. During a warm-water influx in February 2006, chicks were fed less frequently and adults carried out foraging trips of up to 8 days. When conditions became more favourable with colder water temperatures in 2007 and 2008, thin-billed prions decreased trip lengths, more often attended their chick every day, and long foraging trips of six to eight days were not registered during 2008. Chick growth rates mirrored this, as chicks grew poorly during 2006, intermediate during 2007 and best during 2008. Thin-billed prions preyed mainly on squid during incubation and mainly on amphipods and euphausiids during chick-rearing. In the poorest season only, the diet was substantially supplemented with very small copepods. Together, the present results indicate that during warm-water conditions, thin-billed prions had difficulties in finding sufficient squid, amphipods or euphausiids and were forced to switch to lower trophic level prey, which they had to search for over large ocean areas.     

Foraging, prey capture, and gut contents of the mesopelagic narcomedusa Solmissus spp. (Cnidaria: Hydrozoa)

Narcomedusae are the most common group of medusae in the mesopelagic depths of Monterey Bay, California. Numerous capture events of various prey taxa were recorded in situ and analyzed using the Monterey Bay Aquarium Research Institute's remotely operated vehicle "Ventana". In situ video analysis of the stomach contents of 82 Solmissus incisa and S. marshalli revealed 88 identifiable prey items. Most (88%) were gelatinous animals. Of these gelatinous prey, 60.3% were ctenophores, 20.5% were cnidarians, 12.8% were salps, 3.8% were chaetognaths, and 2.6% were polychaetes. Euphausiids accounted for 11.4% of the diet, but they were probably captured adventitiously, as the gut contents of ctenophore prey. The tentacle-first foraging behavior of the narcomedusae is an effective way to capture large, comparatively fast-moving prey, because the fluid disturbance caused by swimming is decoupled from the area of prey capture. This behavior contrasts with the prevailing models of feeding behavior in medusae. Stealth predation may be the dominant mode of capturing prey by medusae in the mesopelagic depths of the oceans.     

Depth utilisation by breeding Adélie penguins,Pygoscelis adeliae,at Esperanza Bay,Antarctica

Miniature depth gauges were attached in December 1987 and January 1988 to Adélie penguins,Pygoscelis adeliae, breeding at Esperanza on the Antarctic Peninsula. Results from 34 birds showed that foraging penguins with eggs and with brooded and crèching chicks spent mean periods away from the nest of 96, 36 and 21 h, respectively, during which time means of 29.0 h (30%), 11.2 h (31%) and 2.7 h (13%), respectively, were spent under water at depths > 5 m. Time under water was positively correlated with time absent from nest. Maximum depth reached was 170 m but overall birds spent most time at shallower depths. Birds foraging for brooded chicks dived deeper than birds foraging for crèching chicks. Stomach-pumping indicated that the principal prey caught at this time was krill,Euphausia superba. Mean mass changes of adults during single foraging trips indicated that krill were caught at a mean rate of 7.2 g min^–1 spent under water.     

Tuna food habits related to the micronekton distribution in French Polynesia

Stomach content analyses are commonly used to study both fish feeding behaviour and trophic conditions. However, the interpretation of such data depends on fish foraging behaviour for a given environment and how representative the stomach contents are to the prey distribution. Tuna feeding behaviour was studied within the context of a research programme conducted in French Polynesia. Tuna prey distribution was characterised using acoustic measurements and pelagic trawls; thereafter, this distribution was compared with the stomach contents of tuna caught using an instrumented longline. Acoustic, pelagic trawling and stomach content analyses give complementary elements to describe the pelagic trophic habitat and to better understand tuna-prey relationships. The classic concept of a reduced food availability for tunas in the tropical pelagic environment seems relative. Tunas able to dive enough during daytime to exploit the migrant micronektonic species secure a source of regular food. This is particularly true of bigeye tuna (Thunnus obesus), which have ecophysiological capacities for this purpose. The behaviour of albacore tuna (T. alalunga), which dive >400 m in depth, remains less clear, as little is known about their vertical behaviour. Lastly, yellowfin tuna (T. albacares), which are distributed in more superficial waters, can better exploit the biomass of juvenile fish and crustaceans exported from the reefs. Analysis of the stomach fullness of tuna caught by longline, a passive gear, generally showed an empty state. This result suggests that most tuna foraging on large prey aggregations present in the study area are quickly satiated and escape longline capture and sampling. A consequence is that studies of tuna feeding behaviour based on longlining may be biased, particularly when large aggregations of prey are present such as in convergence zones. Another potential consequence is that longline tuna catch rates could differ according to prey richness. Longline tuna catch rates may sometimes reflect the relative abundance of prey rather than relative tuna abundance. Electronic supplementary material to this paper can be obtained by using the Springer LINK server located at http://dx.doi.org/10.1007/s00227-001-0776-3.     

Brood neglect and contingent foraging behavior in a pelagic seabird

Among species where there is a risk to leaving offspring unattended, parents usually take alternating shifts guarding their young. However, they may occasionally exhibit brood neglect by leaving their offspring unattended at the nest. To investigate this phenomenon further, we examined the foraging behavior of the northern gannet (Morus bassanus) during chick-rearing. This species has a prolonged nestling period (13 weeks) and the single chick is usually guarded by one or other of its parents, because unattended chicks are frequently attacked by conspecifics. We tested the prediction that the foraging behavior of adults when they left their offspring alone at the nest (unattended trips) would differ in character to when adults left offspring with their partner (attended trips). Brood neglect typically occurred after a longer-than-average attendance period at the nest. Unattended trips were, on average, about half the duration of attended trips, and therefore much closer to the colony. There was also a difference in departure direction between attended and unattended trips, with unattended trips tending to be northeast of the colony. Chicks were fed by parents after both attended and unattended trips, but the frequency and the duration of unattended trips increased as chicks grew older whereas the duration of attended trips decreased as chicks grew. These results indicate that parents may be making a trade-off between risk of attack to their offspring and food-intake rate, and that the solution to this trade-off is dependent on chick age.Communicated by C. Brown     

Compensating for the costs of polygyny in hen harriers Circus cyaneus

In polygynous species, the adults are faced with a dilemma during chick rearing. Males must decide how to distribute food between their females and food allocation patterns are often highly unequal. In turn, the females that receive less food from males have to decide how much time to invest in additional hunting. If they spend more time hunting, then they leave their young exposed to weather and predators. However, if they stay at the nest, they increase the risk of their chicks starving. One way that birds may compensate for reduced provisioning is by increasing the size of prey caught. We tested this hypothesis by comparing prey deliveries to nests of hen harriers, Circus cyaneus, with females of different breeding status. As expected, male harriers delivered less food items to the nests of polygynous females, and especially their secondary, or β females. However, both sexes were able to compensate by delivering larger items and there was no difference in the overall mass of food delivered to nests. Moreover, females spent a similar amount of time at the nest, irrespective of status, and there were no overall differences in breeding success. Our results show that polygynous female harriers can compensate for the costs of polygyny, but we suggest that their ability to do so will vary according to the abundance of both large prey and predators.     

Food supply differentially affects sibling negotiation and competition in the barn owl (Tyto alba)

In contrast to most birds, nestling barn owls (Tyto alba) vocalise not only when parents are at the nest but also in their absence. Calls produced in their absence have been shown to facilitate sibling negotiation over the impending food resource. Since nestlings vocalise more vigorously in the presence of parents, they may be calling not to negotiate resources but to compete amongst each other over parental food distribution. A critical issue is to determine whether offspring need differentially affects sibling negotiation and sibling competition, that is vocalisation in the absence and presence of parents. To answer this question, I manipulated the food supply of 26 broods by adding or removing prey items. In the absence of parents, food-added broods vocalised at a significantly lower level than food-removed ones. In contrast, once a parent arrived at the nest, the vocalisation level was not significantly related to the manipulation of brood food supply. This suggests that in the absence of parents, it is more important for food-removed nestlings to vocalise intensely, and that in their presence, the relationship between begging and offspring need tends to vanish. In other words, brood food supply may affect sibling negotiation to a larger extent than sibling competition.     

Elysia timida (Risso, 1818) (Gastropoda, Opisthobranchia): relationship and feeding deterrence to a potential predator on the south-western Mediterranean coast

The relationship between the sacoglossan Elysia timida and the ornate wrasse Thalassoma pavo was studied in two laboratory experiments using artificial models. A feeding-preference experiment was conducted to determine whether mollusc extract deterred feeding by T. pavo, by using a "realistic" mollusc model (imitating the colour pattern of E. timida) coated with mollusc extract, and a reference model (without imitation or extract), and fishes collected from two locations. It was observed that fish approached, attacked and ate more reference models than mollusc models. A second feeding-preference experiment was designed with four different coloured models: "realistic" (W), green (G), red (R) and chequered (S) pattern. Both fish populations clearly rejected the S model, and differed in their colour preferences. Although both populations seemed to prefer the R and G models equally, the W model was clearly rejected by the fish that coexist with the mollusc at one site (Mazarrón), but was not rejected by the other population of fish which does not coexist with it (Cabo de Palos). Mazarrón fishes would identify the W model with the presence of a toxic compound during their coexistence, and therefore avoid attacking conspicuously coloured E. timida models as a response to their visual signals. Therefore, it was concluded that extract of E. timida is a deterrent for T. pavo, and its effect is sufficiently noxious that the fish tend to avoid it, so that the ability of the fishes to learn to recognise colours and identify certain colour patterns associated with obnoxious prey provides the molluscs the opportunity to survive by exhibiting a conspicuous coloration.     

Foraging behavior of juvenile Carcinus maenas (L.) and Cancer pagurus L.

Information concerning the way juvenile crabs choose their diet from a variety of prey types can be useful for a better understanding of community dynamics, as well as for the adequate management of natural resources. Prey size and species selection by juvenile Carcinus maenas (15-35 mm carapace width, CW) and Cancer pagurus (20-40 mm CW) feeding on four bivalves of contrasting shell morphology were investigated. When offered a wide size range of Mytilus edulis, Ostrea edulis, Crassostrea gigas, and Cerastoderma edule presented individually, crabs generally showed evidence of size-selective predation. Cancer pagurus selected larger mussels relative to the size of their chelae (relative prey size, RPS) than did Carcinus maenas of similar and even larger carapace width. However, the RPS of selected O. edulis and Cerastoderma edule were similar for all crabs, suggesting that certain prey features constitute effective barriers even to the powerful chelae of Cancer pagurus. When offered a wide size range of mussels and oysters simultaneously, all crabs consistently selected mussels. When offered O. edulis and Crassostrea gigas, crabs consumed both these oyster species in similar numbers. Carcinus maenas consumed similar numbers of mussels and cockles; Cancer pagurus, however, showed no preference for either prey in the smaller size classes but selected more mussels than cockles as prey increased in size. Although previous studies report that adult Carcinus maenas select prey species according to their profitability (amount of food ingested per unit of handling time, milligrams per second), consumption rates of the size classes of prey selected by juvenile shore crabs did not always parallel prey value. Although variations in crab strength can account for many of the differences between the foraging strategy of juvenile and adult C. maenas, our results suggest that juvenile crabs are less species selective than adults as a result of the restrictions imposed on small individuals that have limited access to larger prey.     

Body condition of predatory fishes linked to the availability of sandeels

Lesser sandeels Ammodytes marinus are eaten by a range of predatory fishes including commercially fished species, but are also exploited at large scale by industrial fisheries. Is availability of sandeels, as key prey source, linked to the body condition of predatory fishes? In the North Sea, the largest sandeel biomass is concentrated in the Dogger Bank region. Here we studied predator–sandeel interactions at two sites differing widely in sandeel abundance and local sandeel fishing effort. Surveys took place in 2004, 2005, and 2006, years when local sandeel densities observed at these sites were low, intermediate, and high, respectively. Five predator species––whiting, lesser weever, grey gurnard, plaice, and haddock––showed better body condition indices in either the years or study area (or both) characterised by higher local sandeel densities, when compared to sandeel-poorer conditions. Moreover, whiting, weever, and gurnard condition was better for those individuals actually observed to have eaten sandeels (based on stomach contents) than for those that had not. As body condition relates to growth, reproduction, and survival, predators in sandeel-rich conditions may be inferred to have a higher fitness. These links between sandeel availability, sandeel consumption, and predator condition hint that, if large-scale localised depletions of sandeels were to occur, negative indirect effects on predatory fish might become apparent, underlining the importance of considering the sandeel fishery in an ecosystem context.     

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.