Local changes in the composition and community structure of suprabenthic peracarid crustaceans on the bathyal Mediterranean: influence of environmental factors

Deep-water assemblages of suprabenthic peracarids were analyzed in the SW Balearic Islands (Algerian Basin, southwestern Mediterranean) between 249 and 1,622 m depth; the patterns of species composition, possible zonation, and trophic structure found in this area were compared with those exhibited by peracarids in the mainland side of the Catalan Sea slope (northwestern Mediterranean). One hundred and four peracarid species (plus one leptostracan) were identified on the Balearic Islands slope, amphipods being the most diversified taxon (45 species). On the Balearic slope, two distinct depth assemblages were distinguished: one at the upper slope (US), between 249 and 402 m depth and the second at the deep slope, between 543 and 1,620 m depth. A remarkable species substitution occurred at depths between 402 and 638 m. In the Catalan Sea, in addition to the US assemblage occupying depths between 208 and 408 m, a second boundary of faunal change was found around 1,250 m. Suprabenthos biomass increased from 242 to approximately 500 m. Suprabenthos attained the highest biomass values (100 g wet weight/10,000 m²) at intermediate depths between 504 and 1,211 m, as also occurred with the associated zooplankton collected with suprabenthos (peak biomass between 502 m and 898 m). Suprabenthos biomass did not show any significant correlation with any environmental water-column variable. In contrast, zooplankton (especially small fish and decapod crustaceans) showed a significant positive correlation with fluorometry and turbidity at different levels of the water column. The feeding guilds of species showed important differences between the two areas only on the US, with a higher abundance of deposit feeders in the Catalan Sea (20.4%) than in the Balearic Islands (4.2%). The low contribution of deposit feeders in the SW Balearic Islands may ultimately be a consequence of the lack of river discharges in this area.Communicated by S.A. Poulet, Roscoff     

Predator-prey relationships between the rock lobster Jasus lalandii and the mussel Aulacomya ater at Robben Island on the Cape West Coast of Africa

The natural diet and mode of feeding of the rock lobster Jasus lalandii (H. Milne Edwards) was determined in a rock-lobster sanctuary near Cape Town, South Africa. Field observations were tested and confirmed by means of aquarium studies. Rock lobsters feed mainly upon ribbed mussels Aulacomya ater (Molina), which comprise the largest component of the sessile benthic fauna. Mussel remains were found as the major constituent in 97% of the rock-lobster stomachs examined. The density of rock lobsters averaged 8,100 per hectare (0.81 m^-2), while mussel biomass averaged more than 5 kg (wet whole weight) m^-2 within the same depth range (12 to 30 m). More than 80% of the mussel biomass comprised large individuals between 60 and 90 mm in length. Large rock lobsters (mainly males) were capable of feeding on all sizes of mussels, although many of these were inaccessible to predation. Smaller rock lobsters became progressively more limited in the size range of mussels which they could crack open and consume. Competition between rock lobsters for small mussels appeared to be intense, as mussels of suitable size for feeding were generally in short supply to most of the rock-lobster population. Hence, feeding and growth rates of rock lobsters are likely to be affected by the relative population densities of predator and prey. Growth rates of rock lobsters could be limited by food supplies even in areas where mussel biomass is comparatively large.     

A model of nematode dynamics in the Westerschelde Estuary

We developed a time dynamic model to investigate the temporal dynamics of nematode community in the brackish zone of the Westerschelde Estuary. The biomass of four nematode feeding groups observed from March 1991 to February 1992 is used to calibrate the model. Using environmental data as the input, the model predicts the temporal modification and interrelation of four nematode feeding groups. Nematodes achieve a dominant position in the community because of their lower loss rate (in respiration, excretion and natural death). Predators which are deposit-feeding macrobenthos control the variations of dominant nematodes, such as omnivores and non-selective deposit feeders. Food availability causes modification only for rare nematodes such as epigrowth feeders and selective deposit feeders. Temperature is a factor affecting both predation death and a loss including respiration, excretion and natural death. Overall, the modification of nematode community by food availability is much lower than by predator. The macrobenthos in the Westerschelde Estuary decrease from upstream to the estuarine mouth. The stability and standing stock of nematode population follow the opposite gradient of their predators. They increase from upstream to the estuarine mouth.     

Quantitative relationships among phytoplankton body size classes and production processes in the North Adriatic frontal region

The variation patterns of phytoplankton standing crop and productivity in the North Adriatic frontal region and the relative importance of pico-, nano-, micro-phytoplankton are shown. Data of standing crop (chlorophyll a - Chl a ) and productivity ( 14 C assimilation) and PAR radiation ( w Em m 2 r s m 1 ) were collected during the four oceanographic cruises of the PRISMA II project. Average standing crop and productivity in the study area were 1.41 - 0.42 r mg r m m 3 Chl a and 1.23 - 0.37 r mg r C r m m 3 r h m 1 ; average assimilation number (P/B) was 0.872 - 0.589 r mg r C (mg-Chl a ) m 1 h m 1 and average photosynthetic efficiency (PE) was 0.020 - 0.054 (mg r C(mg-Chl a ) m 1 r h m 1 ) [ w Em m 2 r s m 1 ] m 1 . Phytoplankton biomass and productivity showed significant patterns of variation with the distance from the coast, with increasing depth and decreasing light intensity. The same patterns were shown by the three phytoplankton size classes. The spatio-temporal variations were significantly larger within the microplankton than within the pico- and nano-plankton size classes. Planktonic guilds were dominated by picoplankton, both as standing crop and productivity, in the northern stations (0.539 - 0.21 r mg r m m 3 Chl a and 0.572 - 0.25 r mg r C r m m 3 r h m 1 ) and in those more offshore, while microplankton was more important in the coastal and southern stations (0.727 - 0.58 r mg r m m 3 Chl a and 0.63 - 0.28 r mg r C r m m 3 r h m 1 ). In relative terms, picoplankton accounts for the 53% and 46% of biomass and primary production, while the microplankton account for the 43.6% and 48%. Assimilation number and photosynthetic efficiency did not show spatio-temporal variations but PE was inversely related with PAR radiation for all the size classes. Data suggest that the spatio-temporal patterns observed in this study are affected by the competitive relationships among body size classes in the phytoplankton guilds.     

Combining gut fluorescence technique and spatial analysis to determine <Emphasis Type="Italic">Littorina littorea</Emphasis> grazing dynamics in nutrient-enriched and nutrient-unenriched littoral mesocosms

Spatiotemporal distribution patterns in relation to feeding behavior of herbivorous gastropods have been studied extensively, but still knowledge about small-scale patterns is limited in relation to eutrophication. This experimental study aimed to describe the small-scale distribution of Littorina littorea in nutrient-enriched and nutrient-unenriched mesocosms in a merely atidal region and relate the distribution to food abundance and possible competing organisms, while checking simultaneously for feeding activities. The latter part was accomplished through the “gut fluorescence technique” GFT (which, to our knowledge, has not previously been used for benthic grazers) to estimate per capita grazing rates and the former part through monitoring of spatial heterogeneity of L. littorea and co-variation with sessile organisms (using semivariograms and cross-semivariograms, respectively). After 5 months of nutrient addition, the abundance and biomass of L. littorea had increased in enriched systems, which also had significantly higher total biomass of green algae. Gut pigment content was higher in L. littorea from enriched mesocosms, and gut depletion rate was higher in L. littorea from unenriched mesocosms. Spatial analysis showed that L. littorea exhibited generally random patterns (suggesting feeding activities) but sometimes (often in the morning) spatial patchiness (clumped distribution) in both enriched and unenriched conditions. There was mainly positive co-variation between L. littorea and biofilm, while different nutrient conditions exhibited contrasting co-variation between L. littorea and barnacles (positive co-variation in enriched and negative co-variation in unenriched mesocosms). The study offered insights into how feeding behavior and spatial distribution of a species may interact with community components differently under different nutrient regimes. The applied methodology can be useful for purposes of faster examination of grazing effects among different regions and also to compare grazing intensities and interactions between grazers and the benthic communities in disturbed (including pollution and nutrient enrichment) and non-disturbed systems, as well as in up-welling versus non-upwelling areas.     

Distribution of deep-water gorgonian corals in relation to benthic habitat features in the Northeast Channel (Atlantic Canada)

The distribution and abundance of deep-water gorgonian corals were investigated along 52 transects at 183–498 m depth in the Northeast Channel, between Georges Bank and Browns Bank in the northwest Atlantic, using a remotely operated vehicle and a towed video-camera system. Three species (Paragorgia arborea, Primnoa resedaeformis, and Acanthogorgia armata) were observed. Primnoa occurred on 35 transects below 196 m depth, with highest local abundance in stands of 104 colonies per 100 m². Paragorgia was present on 21 transects deeper than 235 m, with highest local abundance of 49 colonies per 100 m². Acanthogorgia was observed at only four transects between 231 m and 364 m, with a local maximum abundance of 199 colonies per 100 m². The maximum abundance averaged for whole transects was 19.2 and 6.2 colonies per 100 m² for Primnoa and Paragorgia, respectively. The corals were more common in the outer part of the channel along the shelf break and slope than on the shelf in the inner part. All three species showed a patchy distribution with no signs of competitive exclusion at any spatial scale. Transects with high abundance of corals were characterised by depths greater than 400 m, maximum temperatures less than 9.2°C, and a relatively high percentage coverage of cobble and boulder (more than 19% and 6%, respectively). High temperatures probably control the upper depth limit of the corals, and Primnoa seems to tolerate slightly higher temperatures than Paragorgia. Abundance of both species was negatively correlated with average temperature and positively with cobbles. Together, temperature, percentage cobble and salinity accounted for 38% of the variance of Primnoa. The comparable figure for Paragorgia was 15%. The observed distribution indicated that the abundance of coral is controlled by additional factors such as larger-scaled topographic features governing the current regimes and thus also the supply of food and larvae.Communicated by R. J. Thompson, St. Johns     

Deep-sea harpacticoid copepod diversity maintenance: The role of polychaetes

Several models to account for enhanced diversity in the deep sea have been proposed, but the available natural history information has been inadequate to distinguish among them. In particular, few data exist on patterns of co-occurrence among species. At 1220 m depth in the San Diego Trough (32°35.75N; 117°29.00W), harpacticoid copepod species covary significantly with polychaetes when the polychaetes are combined into functional groups on the basis of feeding type and mobility. In particular, harpacticoid species tend to avoid polychaetes which are sessile surface-deposit feeders. The results provide support for models in which disturbance/predation plays an important role in maintaining deep-sea diversity.Contribution No. 15 from Expedition Quagmire.     

Dietary contribution of the microphytobenthos to infaunal deposit feeders in an estuarine mudflat in Japan

The food sources of benthic deposit feeders were investigated at three stations in an estuarine mudflat (Idoura Lagoon, Sendai Bay, Japan) during July and August 2005, using δ¹³C and δ¹⁵N ratios. Sediment at the stations was characterized by low chlorophyll (chl) a content (0–1 cm depth, <4 μg cm⁻²) and the dominance of riverine–terrestrial materials (RTM) in the sediment organic matter (SOM) pool. Surface-deposit feeders (Macoma contabulata, Macrophthalmus japonicus, and Cyathura muromiensis) exhibited much higher δ¹³C values (−18.4 to −12.4‰) than did the SOM pool (<−25‰). A δ¹³C-based isotopic mixing model estimated that benthic diatoms comprised 45–100% (on average) of their assimilated diet, whereas RTM comprised a lesser fraction (29% maximum). The major diet of the deep-deposit feeding polychaetes Notomastus sp. and Heteromastus sp. was benthic diatoms and/or marine particulate organic matter (POM), with little RTM assimilated (39% maximum). The consumers appeared to lack specific digestive enzymes and to use detritus-derived carbon only after its transfer to the microbial biomass. The isotopic mixing model also showed that the dietary contribution of RTM increased slightly (15% maximum) in the vicinity of freshwater input, suggesting that spatial changes in RTM supply affect the dietary composition of deposit feeders. These results clearly demonstrate that deposit feeders selectively ingest and/or assimilate the more nutritious microalgal fractions in the SOM pool. Such adaptations may allow enhanced energy gain in estuarine mudflats that are rich in vascular plant detritus with low nutritive value.     

Particle size-selection of two deposit feeders: the amphipod Corophium volutator and the prosobranch Hydrobia ulvae

The feeding biology of the deposit-feeding amphipod Corophium volutator is compared to that of the coexisting, deposit-feeding prosobranch Hydrobia ulvae. Regarding ingestion of particles, both forms show size selection which alone can explain their coexistence. Particle size-selection also explains some qualitative differences in the composition of the food of the two forms; thus, diatoms play a relatively larger role in the diet of H. ulvae than in the diet of c. volutator, whereas bacteria are probably relatively more important for the latter. Results of experiments with feeding of C-14 labelled microorganisms are in accordance with the findings on particle size-distribution of the gut contents, and show that (1) C. volutator can only utilize bacteria adsorbed to particles within the size range 4 to 63 (this is why the presence of clay and silt particles in the sediment are necessary for efficient feeding of this amphipod); (2) C. volutator can utilize bacteria suspended in the water pumped through its burrow for respiration if silt and clay particles are present in the sediment. (3) H. ulvae can utilize large particles, and also browses on surfaces, and some evidence is brought forward that it also utilizes mucus for trapping microorganisms. The coexistence of deposit-feeding animals is discussed. It is concluded that the number of coexisting, closely related species is usually small, and that their resource partitioning is probably mainly based on particle-size selectivity. In the case of unrelated forms (e.g. H. ulvae and C. volutator, a number of behavioural, physiological and morphological differences, and also the widespread ability of deposit feeders to utilize alternative feeding mechanisms may also lead to resource partitioning. Thus, there are often several niche dimensions related to feeding allowing a certain diversity of coexisting deposit feeders.     

Direct versus indirect methods of quantifying herbivore grazing impact on a coral reef

Two methods were used to assess the grazing impact of roving herbivorous fishes across a coral reef depth gradient within Pioneer Bay, Orpheus Island, Great Barrier Reef. The first technique employed was a method traditionally used to quantify herbivory on coral reefs via the (indirect) inference of herbivore impact from biomass estimates and reported feeding rates. The second method (one of a range of direct approaches) used remote underwater video cameras to film the daily feeding activity of roving herbivores in the absence of divers. Both techniques recorded similar patterns and relative levels of herbivore biomass across five reef zones at the study site. Indirect estimates of the grazing impact across the reef depth gradient of the three predominant species of herbivore broadly coincided with levels quantified directly by remote underwater video, indicating that, to a large extent, presence does correspond to function. However, the video data suggested that, for individual species in particular reef zones, the absolute level of impact may be less than that inferred from presence. In the case of the parrotfish Scarus rivulatus, the video recordings suggested that, at the reef crest, an average of 52% (±18 SE) of each m² area of reef would be grazed each month, compared with an area of 109% (±41 SE) suggested by inferring grazing activity from presence alone. Potential biases associated with remote video recorders may explain some of the discrepancy between values. Overall, the results suggest that, for some fish groups, the indirect method of inferring function from presence can provide a good indication of relative levels of herbivore impact across a coral reef. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Biodiversity and trophic structure of nematode assemblages in seagrass systems: evidence for a coupling with changes in food availability

Nematode assemblage composition, trophic structure and biodiversity were followed over an annual cycle in a Posidonia oceanica bed of the NW Mediterranean to test the response to temporally changing food availability (sediment organic matter, bacterial and microphytobenthic biomass). The sediment-water interface of the seagrass meadow was characterised by high particulate organic matter concentrations. Also, seagrass sediments had high organic matter content, chloropigment concentrations and bacterial biomass. All organic matter components (i.e. carbohydrates, proteins and lipids) changed temporally, with higher concentrations in winter and spring and lower concentrations in summer; however, overall, large amounts of organic compounds were potentially available for (seagrass) benthic consumers throughout the year. Nematode assemblage in the P. oceanica bed maintained a high genus number (88 genera), and a trophic structure tightly coupled with the composition of the potential food sources. In agreement with the relevance of microphytobenthos, epigrowth-feeder nematodes (2A) represented the dominant trophic guild. The biomass of predator nematodes (2B) was significantly correlated with the biomass of other nematodes. Non-selective deposit feeders (1B) displayed a close relationship with the concentration of proteins, carbohydrates and biopolymeric carbon (i.e. labile organic detritus). Conversely to what was expected, epigrowth feeders were not correlated with chloropigment concentrations, but showed a significant relationship with the number of dividing bacteria, possibly suggesting a trophic plasticity of the 2A feeding guild. The coupling between temporal changes in food sources and temporal variability of the nematode trophic structure was highlighted by a CANOCO analysis, which allowed us to identify and associate, at each sampling time, nematode genera and their feeding habits, with specific environmental variables and food indicators. Furthermore, species diversity (H') and evenness (J) calculated on nematodes identified to the genus level displayed evident temporal changes, also reflected by the index of trophic diversity (ITD). Both structural and functional diversity were coupled with high concentration and highly heterogeneous composition of the food sources (including organic detritus, microphytobenthic algae and bacteria) potentially available to nematodes. These data suggest that temporal changes in quantity and quality of food sources do not only influence nematode dynamics and trophic composition, but also influence nematode structural and functional diversity.     

Gonad morphology and oocyte development in<Emphasis Type="Italic"> Pseudocalanus</Emphasis> spp. in relation to spawning activity

Gonad maturation processes were studied in Pseudocalanus spp. females from Georges Bank and Cape Cod Bay (northwest Atlantic) using a combination of morphological analysis and experiments. For light microscopy of the oocytes, females of different maturation stages were preserved immediately after capture. The maturation processes during the spawning cycle were described from observations of live females which were exposed to feeding and starvation at two temperatures, 8 and 15°C, for 12 days. The gonad morphology of these females was examined in 24 h intervals, and spawning events were recorded. Both light microscopy and whole animal observation revealed that oocytes during maturation change in shape and size, in the morphology of the nucleus, and in the appearance of the ooplasm. Due to these modifications of oocyte morphology and due to oocyte migration, the morphology of the gonads changed distinctly during a spawning cycle. Five oocyte development stages were identified by light microscopy and related to a macroscopic system of four gonad development stages, that can be applied to whole animals and allows the identification of females ready to spawn. The experiment showed that food and temperature had strong effects on gonad maturation processes. High proportions of mature females were found when food was available, whereas the proportion of immature females increased shortly after exposure to starvation. Compared to 15°C, gonad maturation at 8°C was prolonged and thus spawning frequency was lower. The final maturation processes at food saturated conditions were slower than the embryonic development, and no indication was found that mature oocytes are stored in the diverticula waiting to be released. The duration of the interspawning interval would thus be determined by the duration of final oocyte maturation which is dependent on both temperature and food supply.Communicated by O. Kinne, Oldendorf/Luhe     

Population dynamics of Eudendrium glomeratum (Cnidaria: Anthomedusae) on the Portofino Promontory (Ligurian Sea)

Eudendrium glomeratum Picard, in the Ligurian Sea, is one of the major components of hard-bottom sessile zoobenthos in the cold season. It settles mainly between 10 and 40 m depth, forming a seasonal facies. The presence of E. glomeratum has been evaluated by measuring in situ the height of the colonies present within a standard surface of 1 m². Observations were carried out every other week for a whole year. The height of the colonies was correlated with their biomass (wet weight) and trophic functionality (number of polyps). The quantitative evaluation of the population shows that the situation is stable, with almost identical values at the extremes of the observed yearly time series. The species is present from October to April–May, and it is absent in the summer. The formation and disappearance of the E. glomeratum population are sudden. The colonies are sexually mature from their appearance in October to December–January. Different categories of colonies are recognizable; and are essentially referable to two groups: (1) colonies deriving from regeneration of colonies present in the previous year (45%); (2) colonies deriving from planula settlement (43%). A third group (12%) is of uncertain origin and may belong to either of the abovementioned categories. Mortality is 60% in the first group, and 82% in the second one. By dead colonies is meant those which do not reappear after a yearly cycle.     

The capacity of the filter-feeding bivalve Mya arenaria L. to affect water transport in sandy beds

Benthic filter feeding macrofauna organisms may be an important factor linking sediment and water column. Transport of water and concomitantly of suspended matter is directly related to the size of the benthic filter feeding population. This paper aims to determine the potential for water transport by the bivalve Mya arenaria along a coastal stretch of roughly 100 km length in the southern Baltic Sea. Quantification of population filtration rates specific to the area is based on distribution, abundance and biomass of M. arenaria and calculated according to previously published filtration rate–biomass relations. Calculated rates range up to >8 m³ m^–2 day^–1 (at 5–20 m water depth in sandy sediment) with the potential to locally process a volume of water equivalent to the water column within <1 day. Data from 1991–2002 at one site suggest that the area-specific potential population filtration rate remains remarkably constant in time despite changes in population structure of M. arenaria. The related impact on pore water exchange within the permeable sediment associated with the leakage of water from the gape of M. arenaria valves is discussed.Communicated by L. Hagerman, Helsingør     

Surface sequestration of chemical feeding deterrents in the Antarctic sponge<Emphasis Type="Italic"> Latrunculia apicalis</Emphasis> as an optimal defense against sea star spongivory

Latrunculia apicalis is a spherically shaped demosponge that previous investigations have shown is rarely preyed upon by sea stars which are the dominant spongivores in antarctic benthic communities. Prior studies have also demonstrated that L. apicalis produces organic compounds that elicit a tube foot retraction response in the keystone spongivorous sea star Perknaster fuscus that can be used as a reliable assay for feeding deterrence. L. apicalis is known to contain discorhabdin alkaloids which serve, among other roles, as the source of its green coloration. To assess the defensive nature of the discorhabdin alkaloids toward P. fuscus, we have determined discorhabdin G concentrations in discrete sponge layers and evaluated those concentrations in the P. fuscus bioassay. In discorhabdin G-bearing sponges, we found a gradient of discorhabdin G that falls off rapidly toward the center of the sponge. On average, 52% of total discorhabdin G in a given sponge was found within 2 mm of the sponge surface. Tube foot retraction responses to extracts from the surface tissues (0–2 mm depth) of L. apicalis were compared to those of an inner layer (8–10 mm depth) and to a sample comprised of the same inner layer spiked with discorhabdin G at a concentration equivalent to that of the surface tissues. Tube foot retraction response times to extracts of the surface layers and the spiked inner layers were not statistically different, but were significantly greater than responses to the unaltered inner layer and controls. These results support the predictions of the optimal defense theory as L. apicalis sequesters its defensive chemistry (discorhabdin G) in its most vulnerable surface tissues, where the likelihood of predation from sea stars is highest. As antarctic sponges are generally preyed upon by extraoral feeding sea stars rather than deeper biting predators such as fish, surface sequestration may be uniquely adaptive in sessile macroinvertebrates occurring in antarctic marine benthic environments.Communicated by P.W. Sammarco, Chauvin     

Potential impact of a toxic dinoflagellate (Alexandrium excavatum) bloom on survival of fish and crustacean larvae

During the last 20 yr the western half of the Dutch Wadden Sea has undergone significant eutrophication: concentrations of P and N compounds and planktonic algae have roughly doubled, as has primary production. Though oxygen levels are often low in summer, anoxic areas are small and rare due to strong tidal mixing. During the 1970 to 1990 period, macrozoobenthos was sampled annually at 15 stations at Balgzand, a 50-km² tidal-flat area in the westernmost part of the Wadden Sea. Not only did the estimates of total numbers, biomass, and production double during these two decades, but significant changes in the composition of the benthic community were observed, too: (1) the numerical proportion of polychaetes increased at the expense of molluscs and crustaceans, (2) the overall mean weight per individual of the macrozoobenthos decreased (numbers of individuals of small-sized species increased more rapidly than those of large-sized species), and (3) though absolute numbers and biomass of all feeding types increased, the share of carnivores declined and that of deposit feeders increased; the proportion of suspension feeders showed little change. This study refers to true macrobenthos only (1-mm sieve) and further excludes two taxa (Corophium spp. andHydrobia ulvae) which occasionally exercised an undue influence on numbers. Mass mortalities caused by low oxygen concentrations were of a small-scale nature only. Total number of species fluctuated without a clear trend. As a consequence of the increasing numerical densities, trends in species numbers were slightly increasing when expressed per unit of area and slightly decreasing when estimated per 100 individuals (by rarefaction).     

Contrasting diel patterns in vertical movement and locomotor activity of whale sharks at Ningaloo Reef

Activity patterns of animals often relate to environmental variables such as food availability and predation pressure. Technological advances are providing us with new tools to monitor and better understand these activity patterns. We used animal-attached data loggers recording acceleration and depth to compare activity patterns and vertical habitat use of whale sharks (Rhincodon typus) at Ningaloo Reef, Western Australia. Whale sharks showed a moderate reverse diel vertical migration but exhibited a clear crepuscular pattern in locomotory activity. Peak activity occurred at sunset, whereas vertical movement peaked prior to this. Typical ram surface filter feeding could be identified and occurred primarily during sunset and the first hours of night. At such times, direct observations indicated whale sharks were feeding on tropical krill swarms. Kinematic analysis of postural data and data from vertical movement suggests that whale sharks at Ningaloo spend ~8 min per day actively ram surface filter feeding. Considering the high biomass present in krill schools, it is estimated that whale sharks at Ningaloo have a similar energy intake as those at other aggregation sites. Diel patterns in activity and diving behaviour suggest that whale sharks have tuned their diving behaviour in anticipation of the formation of these high-density patches which appear to only be periodically, but predictably available at sunset. Our results confirm that diel patterns in vertical habitat selection and vertical movements do not necessarily reflect patterns in activity and foraging behaviour. Direct quantification of activity and behaviour is required in gaining accurate representation of diel activity patterns.     

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

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

Feeding ecology and niche segregation in oceanic top predators off eastern Australia

We examined the feeding ecology and niche segregation of the ten most abundant fish species caught by longline operations off eastern Australia between 1992 and 2006. Diets of 3,562 individuals were examined. Hook timer data were collected from a further 328 fish to examine feeding behaviour in relation to depth and time of day. Prey biomass was significantly related to predator species, predator length and year and latitude of capture. Although the fish examined fed on a mix of fish, squid and crustacea, fish dominated the diet of all species except small albacore (Thunnus alalunga) which fed mainly on crustacea and large swordfish (Xiphias gladius) and albacore which fed mainly on squid. Cannibalism was observed in lancetfish (Alepisaurus spp.). Multidimensional scaling identified three species groups based on their diet composition. One group consisted of yellowfin tuna (T. albacares), striped marlin (Tetrapturus audax) and dolphinfish (Coryphaena hippurus); a second group consisted of bigeye tuna (T. obesus), swordfish and albacore; and a third consisted of southern bluefin tuna (T. maccoyii) and blue shark (Prionace glauca). Of note was the separation of mako shark (Isurus oxyrhynchus) and lancetfish from all other predators. Prey length generally increased with increasing predator length although even large predators fed on a wide range of prey lengths including very small prey. Overall, differences in prey type and size, feeding times and depths were noted across the range of species examined to the extent that predators with overlapping prey, either in type or size, fed at different times of the diel period or at different depths. Taken together these data provide evidence for feeding niche segregation across the range of oceanic top predators examined.     

Long-term changes in the Georges Bank food web: trends in stable isotopic compositions of fish scales

Fish scales from seven species of demersal fish in an archival collection were analyzed for stable isotopic compositions of carbon and nitrogen to study long-term changes in trophic structure of the Georges Bank food web. Nitrogen isotopic compositions are often used to infer trophic level. In the case of haddock, Melanogrammus aeglefinus (Linnaeus), there was a trend towards feeding at 2/3 of one trophic level (2.45% in ¹⁵N) lower in 1987 than in 1929. Values of °¹³C, frequently employed to identify sources of organic carbon to consumers, declined by 1.5 from 1929 to 1960, and then increased again toward the present, suggesting changes in the food web at the level of the primary producers. Superimposed on long-term isotopic trends were short-term variations (1 to 10 yr).To identify potential causes for these isotopic trends, canonical correlation analysis was performed between isotopic data and a suite of environmental and population factors including sea surface temperature, the Greenland Regional Pressure Anomaly (GRPA), the North Atlantic Oscillation (NAO), and the following haddock stock parameters; stock size, fishing mortality, recruitment, and weight-at-age-2 (a measure of growth rate). Isotopic variation was significantly correlated with a combination of environmental and population variables: GRPA, NAO, weight-at-age-2, stock size, and fishing mortality. On the basis of published gut content analyses, the seven fish species were predicted to vary in trophic level (TL) from American plaice, Hippoglossoides platessoides (Fabricius), (TL 2.9) to summer flounder, Paralichthys dentatus (Linnaeus), (TL 4.5), whereas measured °¹⁵N values suggested smaller differences in the trophic levels of these species (less than one TL). Four species showed good agreement between gut-predicted and measured ¹⁵N values, while three species did not. Inclusion of information on ontogenetic dietary shifts in our predictions improved the agreement in some cases but not in other. Differences between stable isotope analysis and gut content analysis in terms of what they measure, i.e., integrated assimilated diet vs short-term ingested diet, respectively, may account for some of the differences in results. Based on our analyses and previous studies, feeding habits of these fish may undergo considerable year-to-year and geographic variation, some of which may have been missed in gut content analyses. To the extent that these fish are representative members of the food web, trophic variation in these fish may indicate more general changes in the food web.