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.     

Combining DNA and morphological analyses of faecal samples improves insight into trophic interactions: a case study using a generalist predator

The diet of pinnipeds is most commonly inferred from morphologically diagnostic remains of prey in their scats. Although this method can generate quantitative estimates of diet simply, important prey types may not always be detected. DNA-based methods improve detection of prey in scats, but they are not quantitative. While some studies have combined morphological and DNA-based methods, these have only assessed prey that are represented by their hard remains in scats. To overcome this bias, we apply molecular and morphological analyses to the soft and hard portions of faecal samples respectively, to estimate the diet of lactating Antarctic fur seals (Arctocephalus gazella) on Heard Island. The diet of this population is of particular interest because it is expanding rapidly and may rely to some extent on mackerel icefish (Champsocephalus gunnari), which are subject to commercial fisheries. Based on results from morphological analysis and likely important prey types, we tested for DNA remains of C. gunnari, myctophids and squid in faecal samples. The proportion of samples (n = 54) yielding no dietary information was reduced from around 25.9% using either method alone, to 9.3% when combined. Detection of all prey types tested for was notably improved by integrating molecular and morphological data. Data from either method alone would have underestimated the number of animals consuming C. gunnari by around 25.7%. Detection of multiple prey types in samples increased from 9.3% when using morphological analysis only, to 33.3% when using DNA only, to 46.3% when using both methods. Taken in isolation, morphological data inferred that individual seals consume either C. gunnari or myctophids, probably foraging in separate locations characteristic of those prey. Including molecular data demonstrated that while this may be true of some individuals, many other seals consume a mixed diet of at least C. gunnari, myctophids and squid. This new approach of combining DNA-based and morphological analyses of diet samples markedly increased the number of samples yielding dietary information, as well as increasing the amount of information attained from those samples. Our findings illustrate the broad potential of this technique to improve insight into trophic interactions in marine ecosystems.

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Sympatry and allopatry in the deeply divergent mitochondrial DNA clades of the estuarine pulmonate gastropod genus <Emphasis Type="Italic">Phallomedusa</Emphasis> (Mollusca,Gastropoda)

DNA sequences of the mitochondrial cytochrome c oxidase subunit I (COI) gene were collected from estuarine snails in the genus Phallomedusa to examine the effects of estuarine isolation on population structure and gene flow. Three clades were recovered, one corresponding to Phallomedusa austrina and two others with the morphology of Phallomedusa solida. The haplotype diversity in all three clades indicated recent population expansion. Phallomedusa austrina was restricted to the west of a previous land bridge in the Bass Strait between mainland Australia and Tasmania, and P. solida to its east and to northern Tasmania. Phylogeographic analysis of P. austrina and P. solida shows strong geographic separation of species, but no local genetic structure indicative of regional or estuarine isolation. The clades of P. solida exhibit substantial genetic divergence and were sympatric across their entire distribution in eastern Tasmania and mainland Australia. Such a situation, which has not previously been observed in phylogeographic studies of southeast Australia, suggests that P. solida has had a complex refugial history during periods of environmental challenge.     

Localization of infaunal prey by the sea star<Emphasis Type="Italic"> Leptasterias polaris</Emphasis>

We performed field and laboratory studies to investigate how large adult Leptasterias polaris detect and locate their major prey, large infaunal bivalves, in the sediment bottom community. A field survey using SCUBA diving showed that 95% of the locations where L. polaris dug into the sediment bottom were over bivalves and this success rate was much greater than if digging was done at random (22%). Furthermore, when sea stars were provided with a low density of randomly distributed prey in a laboratory arena, they dug exclusively in locations where a clam had been buried. These observations indicated that L. polaris locates infaunal prey prior to investing energy into digging. Studies in a laboratory flow tank showed that L. polaris readily detected and moved towards its preferred prey Ensis directus whereas its responses to less preferred prey Mya truncata and Spisula polynyma were much weaker. The degree to which it oriented towards these three common prey seemed to reflect potential energy intake relative to foraging costs (which likely increase with the depth of the different prey) and risks from interactions with other carnivores (which are greatest when feeding on large prey). This is the first study to clearly demonstrate that sea stars use prey odours to locate infaunal prey.Electronic Supplementary Material Supplementary material is available in the online version of this article at http://dx.doi.org/10.1007/s00227-004-1497-1Communicated by R.J. Thompson, St. Johns     

Behavioral ecology of the neotropical termite-hunting ant Pachycondyla (= Termitopone) marginata: colony founding,group-raiding and migratory patterns

This study provides the first detailed field account of colony founding, group-raiding and migratory habits in the neotropical termite-hunting ant rPachycondyla marginata, in a semi-deciduous forest in south-east Brazil. New colonies can originate by haplometrosis, pleometrosis, or colony fission. Incipient colonies with multiple foundresses persisted longer in the field, and most excavated nests contained more than one dealated female. A total of 202 group raids by P. marginata were registered, and in all cases the raided termite species was Neocapritermes opacus. Nearly 20% of the workers within a colony engage with raiding activity. Colonies of P. marginata hunt for termites approximately every 2–3 weeks, and group-raids may last for more than 24 h. Target termite nests are up to 38 m from the ant colony, and occasionally two nests are simultaneously raided by one ant colony. Raiding ants carry 1 or 2 paralysed prey, and nearly 1600 termites can be captured during a 9-h raid. Migration by P. marginata colonies lasted over 2 days and covered distances of 2-97 m (n = 48). Average residence time at a given location was 150 days. Three basic migratory patterns were noted: colony fission (only part of the colony moves), long-distance migrations, and short-distance migrations. Both raiding and migratory activities appeared to be strongly affected by seasonal factors. The group raiding and migratory patterns of P. marginata are compared with other ant taxa with similar habits. It is concluded that P. marginata presents a rudimentary form of the so-called army ant behavior, which is highly developed in the subfamilies Dorylinae and Ecitoninae. The extremely specialized diet of P. marginata and the associated high costs of migration are features likely to prevent it from evolving a full army ant life pattern.     

Diet of the social groups of long-finned pilot whales (<Emphasis Type="Italic">Globicephala melas</Emphasis>) in the Strait of Gibraltar

The Strait of Gibraltar is inhabited throughout the year by a group of pilot whales (Globicephala melas), but their spatial distribution varies between Summer and Autumn. In this paper, we have used carbon (¹³C/¹²C) and nitrogen (¹⁵N/¹⁴N) stable isotope signatures to investigate the differences in diet amongst seasons, sex and stable social units. Skin samples were collected from 56 individually photo-identified pilot whales during Autumn 2005 and Summer 2006. These individuals were genetically sexed and their isotopic signature determined. The level of inter-individual association both within and between stable social units were compared to Euclidean distances between individual isotopes signatures. No differences in either δ¹⁵N or δ¹³C were found according to the sex of individuals, but significant seasonal differences were found in δ¹⁵N, although not in the δ¹³C values. This suggests that pilot whales are resident year round in the Strait, a finding supported by independent photo-identification. The variation in δ¹⁵N could reflect a shift in pilot whale diet through the year, with pilot whales feeding at a higher trophic level in Autumn compared to Summer. This could also represent a change in the diet of pilot whale prey species. The δ¹³C values were significantly different amongst the four stable social units sampled and individual δ¹³C values were significantly related to the level of inter-individual association, while no relationship was found for δ¹⁵N. These results suggest that within the same general area (i.e. the Strait of Gibraltar), there is some level of specialisation in habitat or prey choice between pilot whales social units.     

Influence of pycnocline topography and water-column structure on marine distributions of alcids (Aves: Alcidae) in Anadyr Strait,Northern Bering Sea,Alaska

Systematic ship-board surveys were used to simultaneously record seabird abundances and resolve coarse-scale (3 to 10 km) horizontal and fine-scale (1 to 10 m) vertical variability in water-column structure and bathymetry for portions of the coastal zone in Anadyr Strait near western St. Lawrence Island, northern Bering Sea, Alaska, during August and September 1987. Three plankton-feeding alcids, parakeet (Cyclorrhynchus psittacula), crested (Aethia cristatella) and least (A. pusilla) auklets, each exhibited distinct associations for different pycnocline characteristics. Least auklets were more abundant in mixed water, but they also occurred within stratified water where the pycnocline and upper-mixed layer were shallow (8 m) and thin (10 m), respectively. Low body mass (85 g), high buoyancy, and relatively poor diving ability may have restricted this auklet to areas where water-column strata nearly intersected the surface, or to areas from which strata were absent altogether due to strong vertical mixing. Parakeet and crested auklets, which are larger-bodied (ca. 260 g) planktivores with presumably greater diving ability, were more abundant in stratified water, and both species exhibited less specific affinities for water-column characteristic at intermediate and shallow levels. All three auklets avoided locations with strong pycnocline gradients (0.22 _t m^–1), a crude index of the strong, subsurface shear in water velocities characteristic of this region. Auklet distributions in Anadyr Strait were consistent with: (1) strata accessibility, as estimated from relationships between body mass and relative diving ability, (2) possible avoidance of strong subsurface water motions, and (3) habits and distributions of plankton prey. In contrast, largebodied (>450 g) alcids [i.e., common (Uria aalge) and thick-billed (U. lomvia) murres, pigeon guillemots (Cephus columba), tufted (Fratercula cirrhata), and horned (F. corniculata) puffins feeding on fish or benthic invertebrates] showed no consistent relationships with either the pycnocline or upper-mixed layers. All large alcids were more common in stratified than in vertically-mixed waters, but differences in abundance between mixing regimes were small or equivocal. The only measured variable with which all large alcids were associated was total water-column depth: murres, guillemots, and puffins each used areas with shallow sea floors and avoided areas with deeper sea floors. Failure of large alcids to discriminate among foraging areas in Anadyr Strait as a function of pycnocline topography and strength may be attributable to: (1) greater reliance on large pelagic and benthic prey not associated with the pycnocline; (2) higher body mass, lower buoyancy, and greater diving ability; (3) foraging over a uniquely shallow continental shelf where all vertical strata, including the sea floor, are potentially accessible from the ocean surface.     

Diet and trophic position of Atlantic bluefin tuna (<Emphasis Type="Italic">Thunnus thynnus</Emphasis>) inferred from stable carbon and nitrogen isotope analysis

Stable ¹³C and ¹⁵N isotope analyses of scale, bone, and muscle tissues were used to investigate diet and trophic position of North Atlantic bluefin tuna (Thunnus thynnus Linnaeus) during residency in the northwest Atlantic Ocean off the northeast coast of the United States. Adult bluefin tuna scales collected from fish between June and October 2001 were significantly enriched in ¹³C compared to both muscle and bone across all months, while muscle was significantly enriched in ¹⁵N compared to either bone or scale throughout the same period. In muscle tissue, there was evidence of a shift over the summer from prey with ¹³C values (–17 to –18) that were characteristic of silver hake (Merluccius bilinearis) to species with ¹³C values of –20 to –21 that were similar to Atlantic herring (Clupea harengus) and sandlance (Ammodytes americanus). Depletion of ¹⁵N values in adult scales and bone compared to muscle tissue may be explained by bone and scale samples representing juvenile or life-long feeding habits, isotopic routing, or isotopic differences in amino acid composition of the three tissue types. Adult bluefin tuna were estimated to be feeding at a trophic position similar to pelagic sharks in the northwest Atlantic Ocean, while the trophic positions of yellowfin tuna (Thunnus albacares), albacore tuna (Thunnus alalunga), and juvenile bluefin tuna were indicative of a diet of up to a full trophic position below adult bluefin tuna. The close relationship between the juvenile bluefin ¹⁵N values and those of suspension feeders suggests that nektonic crustaceans or zooplankton may contribute significantly to the diet of bluefin tuna, a food source previously overlooked for this species in the northwest Atlantic Ocean.Communicated by J.P. Grassle, New Brunswick     

Responses of Asterias rubens to bivalve prey in a Y-maze

Experiments carried out in a Y-maze apparatus using a constant number of mussels, Mytilus edulis L., as prey showed the presence of a food-seeking period in the predatory starfish Asterias rubens L. collected in the Menai Strait, Anglesey, North Wales. The starfish orientated towards the mussels in the experiment conducted from Novemeber-December to May. The food-seeking activity was not significantly different from random expectation between June and October. It is suggested that this seasonal change in the behaviour of A. rubens can be due either to a lowering of chemo-sensitivity and failure to respond by A. rubens, or to a seasonal change in production of attractant(s) by the prey, or to a combination of the above. A. rubens does not move towards a bait of oyster spat (Ostrea edulis L.) in the Y-maze unless it has been previously conditioned to eat the oysters for about a month. In the Menai Strait, oysters would rarely be encountered by A. rubens and, therefore, the experiments confirmed the concept of ingestive conditioning (Wood, 1968).     

Anatomical and experimental evidence for particulate feeding in<Emphasis Type="Italic"> Lucinoma aequizonata</Emphasis> and<Emphasis Type="Italic"> Parvilucina tenuisculpta</Emphasis> (Bivalvia: Lucinidae) from the Santa Barbara Basin

Previous nutritional models for adults of the lucinid bivalve Lucinoma aequizonata contend that symbiotic chemoautotrophic bacteria provide most of the organic carbon for the host. The existence of this symbiosis, coupled with the hosts distinctive anatomical features, shaped the impression that particulate feeding was not a significant part of L. aequizonata nutrition. Here, we use several techniques to show that particulate feeding is a consistent and important part of the L. aequizonata nutritional strategy. Histological and scanning electron microscopy observations reveal that the gills of L. aequizonata, like those of the lucinid Parvilucina tenuisculpta, have functional mucociliary epithelia, able to transport captured particles to the mouth. Observations of gut content and radiolabeled feeding experiments indicate that L. aequizonata does ingest and assimilate carbon from particulate organic matter. Furthermore, molecular identification of a broad spectrum of organisms in the guts of native adult specimens demonstrates that L. aequizonata is non-selective when ingesting organic material, and has a mixotrophic diet.Electronic Supplementary Material Supplementary material is available in the online version of this article at Communicated by P.W. Sammarco, Chauvin     

Vertical distribution and feeding patterns of midwater fish in the central equatorial Atlantic II. Sternoptychidae

The vertical distribution of seven sternoptychid species was examined from RMT 1+8 samples collected aboard R. V. Meteor in March-April 1979 and from Royal Research Ship R.R.S. Discovery in July 1974 in the central equatorial Atlantic. During daytime sternoptychids occupied depths between 200 and 1250 m, with Sternoptyx pseudobscura living deepest, centering between 800 and 900 m, and Argyropelecus sladeni most shallow, aggregating predominantly at 300 and 400 m. They are all considered limited or partial migrants, ascending only some 100 and 200 m towards the surface at night. Only A. sladeni was observed to enter the epipelagic zone (0 and 200 m).-Feeding patterns were investigated from stomach content analyses of Sternoptyx diaphana, S. pseudobscura, Argyropelecus sladeni and A. affinis. Additional stomach contents were analysed from samples of S. diaphana, A. hemigymnus and A. olfersi collected in June 1985 from F.R.V. Walther Herwig in the temperate NE Atlantic at 46°N, 17°W by means of the Engel Trawl. The food spectrum of the six species is generally described, and additional dietary evidence regarding calanoid copepod prey is provided for four of these taxa. All sternoptychid species investigated were planktivorous, feeding predominantly on copepods and ostracods, except for the largest size class, which preyed heavily on euphausiids and amphipods. The relationship of predator size towards prey type and prey size is analysed for both Sternoptyx species. Of these, S. pseudobscura in particular exhibits taxonomic selectivity towards polychaete prey. The diet of both species of Sternoptyx included a number of epipelagic or even neustonic calanoid copepod species which contributed more than 50% of the total copepod population by numbers. So far it is not known how the predators find access to prey organisms of the upper 200 m, as netfeeding is considered unlikely. Cyclopoid copepods of the genus Sapphirina were observed as dietary component particular of S. diaphana.     

Feeding ecology of a size-structured predator population,the South American sun-star Heliaster helianthus

Patterns of feeding in a population of Heliaster helianthus (Lamarck), a common and dominant species of starfish indigenous to the Pacific South American coast, were investigated in an intertidal habitat in central Peru from October 1986 to April 1987. The H. helianthus population comprised individuals of 3.5 to 30.2 cm body size (diameter) with two modal size classes. The number of rays ranged between 18 and 40, and individuals with 31 to 33 rays accounted for ca. 42% of the total population. There was a higher rate of increase in ray number with body size amongst small individuals(<13.0 cm diam). H. helianthus is capable of feeding on more than one prey item at a time (average of 5.6 to 13.2 prey items handled, with several predators observed to hold >100), and both the number of prey individuals captured and the total prey biomass were significantly correlated with predator size. Amongst a total of 1132 feeding observations, the largest number of predators (an average of 85.4% of those feeding) were preying on the mussel Semimytilus algosus whilst another mussel, Perumytilus purpuratus, ranked second with 21,9% of predators feeding. The proportion of S. algosus in the diet increased from 65.4% in the smallest predator size-group (10.9 cm diam) to 91.2% in the largest (19.0 cm). In contrast, P. purpuratus and barnacles were more highly represented in the diet of small H. helianthus. The smallest size-group (10.9 cm) had low dietary overlap with larger sizes and less specialized prey utilization. Two geographically separated populations of H. helianthus in Peru and Chile showed contrasting patterns of prey utilization. S. algosus and P. purpuratus comprised 85.5 and 6.5% by number in the diet of the Peruvian population, respectively, whilst corresponding figures for the Chilean population were 8.3 and 60.5%, with barnacles attaining a higher share (22.6%). However, the total number of prey individuals per feeding predator was almost the same in Peru and Chile, with 10.0 and 10.7 individuals, respectively. H. helianthus individuals of different sizes occupy slightly different microhabitats within the intertidal area, which, coupled with differential spatial distribution of prey species, results in the predator population being able to utilize a wide range of resources.     

Diet of the hatchetfish Sternoptyx diaphana

Sternoptyx diaphana Hermann is a non-migrating hatchetfish inhabiting the mesopelagic zone between 300 and 1500 m in temperate to tropical oceanic regions. An analysis is presented of the diet of this species, collected from 5 oceanic faunal provinces—the Pacific Subantarctic, and 4 subregions of the North Atlantic Ocean. Stomach analyses of 20 to 40 mm specimens revealed considerable intra-trawl (fish to fish), intraregional (Station to station), and inter-regional variation in size and composition of the diet. Generic composition of stomach contents differed noticeably from region to region, with the highest degree of faunal affinity being only 38%. The dominant food items both in terms of biomass and abundance also varied inter-regionally, with fish, euphausiids, or decapods predominant by weight; and euphaussids, copepods, or amphipods most prevalent by number. Differences in size distribution of food items from region to region were also noted. S. diaphana from regions of cooler overlying water masses fed on fewer but larger prey items, and diet was less diverse than that of fish from warmer waters. The diet consists primarily of omnivorous and carnivorous prey, and there is evidence that larger specimens feed on larger food items. The broad size-distribution and taxonomic spectrum of the prey, features of functional morphology, and low density as inferred from trawl catches, indicate that S. diaphana is a predator of limited pursuit capability. Feeding strategy appears to involve capture of the nearest available prey within its immediate vicinity.     

Temporal variability in zooplankton prey capture rate of the passive suspension feeder<Emphasis Type="Italic"> Leptogorgia sarmentosa</Emphasis> (Cnidaria: Octocorallia), a case study

There is increasing evidence that suspension feeders play a significant role in plankton–benthos coupling. However, to date, active suspension feeders have been the main focus of research, while passive suspension feeders have received less attention. To increase our understanding of energy fluxes in temperate marine ecosystems, we have examined the temporal variability in zooplankton prey capture of the ubiquitous Mediterranean gorgonian Leptogorgia sarmentosa. Prey capture was assessed on the basis of gut content from colonies collected every 2 weeks over a year. The digestion time of zooplankton prey was examined over the temperature range of the species at the study site. The main prey items captured were small (80–200 µm), low-motile zooplankton (i.e. eggs and invertebrate larvae). The digestion time of zooplankton prey increased when temperature decreased (about 150% from 21°C to 13°C; 15 h at 13°C, 9 h at 17°C, and 6 h at 21°C), a pattern which has not previously been documented in anthozoans. Zooplankton capture rate (prey polyp^–1 h^–1) varied among seasons, with the greatest rates observed in spring (0.16±0.02 prey polyp^–1 h^–1). Ingestion rate in terms of biomass (g C polyp^–1 h^–1) showed a similar trend, but the differences among the seasons were attenuated by seasonal differences in prey size. Therefore, ingestion rate did not significantly vary over the annual cycle and averaged 0.019±0.002 g C polyp^–1 h^–1. At the estimated ingestion rates, the population of L. sarmentosa removed between 2.3 and 16.8 mg C m^–2 day^–1 from the adjacent water column. This observation indicates that predation by macroinvertebrates on seston should be considered in energy transfer processes in littoral areas, since even species with a low abundance may have a detectable impact.Communicated by S.A. Poulet, Roscoff     

Prey selection by larval weakfish (Cynoscion regalis): the effects of prey size,speed, and abundance

We examined feeding by larval weakfish, Cynoscion regalis (Bloch and Schneider), in laboratory experiments conducted during the 1991 spawning season. under natural conditions weakfish larval development is ca. 3 wk, and we ran separate experiments with larvae of five different ages (5, 8, 11, 14, and 17 d post-hatching). We used two different size classes of rotifers (Brachionus plicatilis) and brine shrimp nauplii (Artemia sp.) as prey organisms. Contrary to results of previous research, weakfish larvae did not select prey based on size alone. When prey abundance was above 100 itemsl^-1 weakfish, larvae always chose large rotifers (length = 216 m) over small rotifers (length = 160 m). At 11 d post-hatching, larvae switched their diet from large rotifers to small brine shrimp nauplii (length = 449 m); however, when fed small rotifers and small brine shrimp nauplii the change in diet occurred at 14 d post-hatching. This pattern of selectivity was maintained in each larval age class. Early-stage larvae (5 and 8 d post-hatching) did not feed selectively when prey abundance was less than 100 itemsl^-1. Late-stage larvae (17 d post-hatching) fed selectively at abundances ranging from 10 to 10000 items^-1. Lwimming speeds of prey items, which ranged from 1 to 6 mms^-1, had no consistent effect on prey selection. These results suggest that weakfish larvae are able to feed selectively, that selectivity changes as larvae age, and that selectivity is also influenced by prey abundance.     

Diet of the Humboldt penguin (<Emphasis Type="Italic">Spheniscus humboldti</Emphasis>) in northern and southern Chile

The diet of the Humboldt penguin (Spheniscus humboldti) was examined and compared in two colonies in Chile. Field work was conducted on Pan de Azúcar Island in northern Chile in the breeding season 1998/1999 and on the Puñihuil Islands in southern Chile over two successive breeding seasons during 1997/1998 and 1998/1999. Penguin diet was studied by stomach-pumping birds and analysed by species composition, size and mass of prey. Fish were the dominant prey item at both sites, the contribution of cephalopods and crustaceans varying between sites. The fish prey consisted predominantly of school fish, but there were clear latitudinal differences in fish prey taken. Penguins in the northern colony consumed primarily garfish (Scomberesox saurus), while birds at the southern colony of Puñihuil fed primarily on anchovy (Engraulis ringens), Araucanian herring (Strangomera bentincki) and silverside (Odontesthes regia). The results showed significant differences in terms of numbers of fish taken between the two breeding seasons at Puñihuil. In 1997/1998 penguins consumed almost exclusively anchovy, while they fed primarily on silversides in the successive year. Almost all prey, except stomatopods, were characterised as being pelagic species that occur in relatively inshore water, consistent with the foraging behaviour of Humboldt penguins. The dependence of Humboldt penguins on commercially exploited, schooling prey species makes the species particularly susceptible to changes in prey stocks, due to non-sustainable fisheries management.Communicated by O. Kinne, Oldendorf/Luhe     

Daily ration and specific daily ration of the chaetognath Sagitta enflata

Digestion times, obtained directly from laboratory feeding, were used in conjunction with a day/night series of field samples to estimate the daily ration and specific daily ration of Sagitta enflata Grassi, the most abundant chaetognath in the Gulf Stream near Miami, Florida (USA), during winter and early spring. Feeding was independent of time of day and increased with increasing chaetognath size from 1.7 prey day^-1 for the 11.5 mm size class to 2.9 prey day^-1 for the 21.5 mm class. The diet of s. enflata consisted of 94.8% copepods having a mean weight of 21.8 g, and 5.2% prey chaetognaths estimated to be two-thirds the length of the predators. The daily ration of S. enflata from 12.5 to 20.5 mm in length increased from 48.3 to 143.3 g of prey dry weight and from 18.5 to 48.1 g of prey carbon. The specific daily ration declined from 0.12 to 0.08 dry weight basis, and from 0.26 to 0.14, carbon basis, over the same size range. Prey chaetognaths contributed as much as 51% of the ration on a carbon basis.     

Assessing feeding of a carnivorous copepod using species-specific PCR

The polymerase chain reaction (PCR) offers a sensitive and selective way to detect trace amounts of biological remnants. Here, we show that this simple molecular technique can be applied to identify prey copepods in the fecal pellets of carnivorous zooplankton. Using variation in the mitochondrial cytochrome C oxidase subunit I (mtCOI) sequence, we developed a species-specific oligonucleotide PCR primer (COI-2026) for Calanus helgolandicus. In a touch-down PCR, Calanus DNA was amplified from pellets collected from freshly incubated individuals of the carnivorous copepod Pareuchaeta norvegica. Positive results could easily be detected by agarose gel electrophoresis.Electronic Supplementary Material Supplementary material is available in the online version of this article at Communicated by M. Kühl, Helsingør     

Dietary constraints on reproductive periodicity in two sympatric deep-sea astropectinid seastars

Examination of the diet of two sympatric species of seastar, Bathybiaster vexillifer and Plutonaster bifrons from 2200 m depth in the Rockall Trough, NE Atlantic Ocean (57°18'N; 10°28'W), suggested that diet may determine the different reproductive patterns found between these two species. In the non-seasonally breeding B. vexillifier, the diet showed a high Shannon-Wiener prey diversity index, the dominant prey being the irregular echinoid Hemiaster expergitus together with a variety of prosobranch gastropods and protobranch bivalves. By contrast, the prey diversity in the seasonally breeding P. bifrons was significantly lower than that of B. vexillifer. In addition, organic carbon content in the sediment residue in the stomachs of P. bifrons displayed a seasonal cycle, while no such seasonality was detected in B. vexillifer. The stomachs of P. bifrons also contained a higher proportion of scavenged material, including the seasonally available remains of mesopelagic blue whiting. These data, together with Bathysnap (time-lapse camera) observations of feeding behaviour in both species, suggest that B. vexillifer is a predator feeding deep in the sediment, whereas P. bifrons feeds close to the sediment surface where it is affected by the seasonal availability of phytodetritus and fish carcasses.     

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.