Trophic niche overlap between round sardinella (Sardinella aurita) and sympatric pelagic fish species in the Western Mediterranean

The northward expansion of round sardinella (Sardinella aurita) in the Mediterranean Sea, together with declines and fluctuations in biomass and landings of European sardine (Sardina pilchardus) and anchovy (Engraulis encrasicolus) observed in recent decades, may suggest potential inter‐specific competition in the pelagic domain. The coexistence of sympatric zooplanktivorous fish species might therefore be exposed in part to trophic niche overlap and competition for food. Combining visual diet characterization under the microscope with DNA metabarcoding from stomach contents of fish collected in spring results show that predation on relatively large krill is equally important for sardinella than for the other two niche overlapping species. Furthermore, an important overlap is found in their isotopic niche, especially with anchovy, using nitrogen (δ¹⁵N) and carbon (δ¹³C) stable isotopes in muscle tissue. In fact, the three fish species are able to feed effectively in the whole prey size spectrum available during the sampled season, from the smallest diatoms and copepods to the larger prey (i.e., decapods and euphausiids), including fish larvae. Moreover, effective predation upon other large prey like siphonophores, which is observed only when multi‐proxy analyses in stomach contents are applied, might also be relevant in the diet of sardinella. The overlapping diet composition in spring, together with the effective use of food resource by sardinella, can be of special interest in potential future scenarios with warmer water temperature leading to lower zooplankton and/or higher jellyfish availability, where sardinella may take advantage over other species due to its feeding plasticity.     

Diet composition of bluefin tuna (Thunnus thynnus L. 1758) in the Eastern Mediterranean Sea,Turkey

This study gives relevant information on the diet composition of the bluefin tuna (Thunnus thynnus) during the spawning period in the eastern Mediterranean Sea. The stomach contents of 218 bluefin tuna were sampled from 2003 to 2006 during the fishing season (May–June) aboard purse seiners operating in the northern Levantine Sea off the coast of Turkey. Stomachs were removed from the fish soon after landing and kept frozen at ?18°C until analysis. Prey items were classified into large taxonomic categories and preserved in 70% ethanol. A total of 745 different prey specimens belonging to 47 taxa were identified, including 34 species of fish, 11 of squid, and two of crustaceans. The most important fish and cephalopod prey belonged to the families Myctophidae, Carangidae, Chauliodontidae, Paralepididae, and Octopoda. This study marks the observation of myctophid fish in the stomach contents of bluefin tuna from the Mediterranean Sea. The paper offers some new information of regional importance and compares the feeding habits of the species to other regions, bringing confirmation on the opportunistic feeding ecology of the species in the enclosed Mediterranean Sea, where bluefin tuna seasonally occur as a strong cohort. New information on the diet composition of T. thynnus in the eastern Mediterranean Sea is revealed; the findings indicate that, depending on the abundance of the different prey species in the habitat, the dominant prey species can be distinctive.     

Scale morphometry allows discrimination of European sardine Sardina pilchardus and round sardinella Sardinella aurita and among their local populations

Landmark‐based geometric morphometric analysis revealed differences in scale shape between European sardine Sardina pilchardus and round sardinella Sardinella aurita as well as among the local populations of each species. Fish scale measurements from four different areas in the central and eastern Mediterranean Sea showed that the mean scale shape of the two species using landmark data could be differentiated with high certainty. Populations of S. aurita from the central and eastern Mediterranean Sea could be separated reliably (P < 0·001) with an average discrimination rate of 91%, whereas the average discrimination of the S. pilchardus populations was lower (80%), albeit still high.     

Sea warming and fish distribution: the case of the small pelagic fish, Sardinella aurita, in the western Mediterranean

This study analyses the temporal and spatial changes in abundance and distribution of the warm water species round sardinella (Sardinella aurita) in the western Mediterranean over the last decades in relation to sea water temperature. In the western Mediterranean basin (1950–2003), a significant positive relationship was found between round sardinella landings and temperature anomalies. Along a latitudinal gradient off the Mediterranean Iberian coast (1989–2004), a gradual increase in species abundance was observed from south to north, with a certain time lag going northwards, associated with the increase in sea water temperature. The abundance of round sardinella in the two warmest and southernmost areas was positively and significantly correlated with sea surface temperature registered during the start of gonad maturation the previous year. In addition, the positive relationship established between water temperature and abundance of round sardinella in the coldest and northernmost study area demonstrates that there is a temperature limit for the distribution of this species in the western Mediterranean. In addition, this study analyses round sardinella larvae distribution and abundance in the summers of 2003 and 2004, and conducts a comparison with the situation 20 years ago (summer 1983). Results show a marked increase in larval abundance during the last decades and the present appearance of larvae in the northernmost study areas, where they did not occur 20 years ago. This indicates the successful reproduction of round sardinella in the northern part of the Mediterranean, where the species has expanded, confirming its establishment in the area.     

A phenotypic trade-off between previous growth and present fecundity in round sardinella <Emphasis Type="Italic">Sardinella aurita</Emphasis>

The decision of how to allocate surplus energy to reproduction and growth can have important effects on fish population dynamics as well as on other life history traits. The natural examples on the interrelationship between maternal growth and number of offspring produced in fishes are scarce. We tested the hypothesis that these traits are competing for resources by estimating maternal previous growth decisions, using back-calculation, and present reproduction, expressed as absolute fecundity, of female round sardinella (Sardinella aurita) in the northeastern Mediterranean Sea. Despite the overall increasing trend of fecundity with age, individual fecundity was negatively related to individual specific growth rate between the most recent annulus formation and spawning within ages. A decreasing trend between previous growth rate and present fecundity emerged, showing that round sardinella allocate increasingly less energy to growth with age and more into reproduction and that the previous growth decisions determine present fecundity.     

Ocean acidification responses in paralarval squid swimming behavior using a novel 3D tracking system

The study presents the biomass and spatial distribution of Sardinella aurita in the Libyan, Maltese and southern Sicily waters, estimated during four acoustic surveys in summer 2008 and 2010. Strong differences in terms of both spatial distribution and total biomass between years and areas were found, with higher total biomass in 2010 than in 2008 in both southern and northern areas of the central Mediterranean. Habitat suitability and changes in the spatial dynamics of round sardinella among areas and years are then explored in relation to total biomass variation and environmental factors. The area of presence of S. aurita increased in the Maltese and Sicilian waters in 2010 with respect to 2008, while most of the total biomass observed in the two years occupied the same proportion of the continental shelf in Libyan waters. The link between environmental conditions and S. aurita area of presence and aggregation was investigated by means of generalized additive models (GAM). The application of GAM singled out the key role of depth and temperature in driving higher round sardinella aggregations, as they were able to explain in both years about the 48% of total deviance in the case of strictly positive values. Favourable habitat for round sardinella was found in waters shallower than 60 m depths, with a clear peak around 40 m depth. Favourable temperature values were above 22°C for presence/absence case and above 24°C when GAM was applied on strictly positive values. Overall S. aurita biomass and distribution were discussed in the light of the classical models proposed to describe the fish spatial dynamics in relation to an increase in total biomass.     

Feeding habits of Schizothorax oconnori Lloyd, 1908 in the Yarlung Zangbo River,Tibet

Seasonal, ontogenetic and sexual feeding habits of Schizothorax oconnori in the Yarlung Zangbo River were studied using percentages by number, weight and index of relative importance (IRI). The Schoener overlap index was applied to compare diets related to season, fish size and sex. The feeding intensity exhibited seasonal trends, with a minimum food intake in summer. Overall, 103 prey taxa belonging to seven prey categories (diatoms, green algae, cyanophytes, other algae, small invertebrates, macroinvertebrates and remains) were identified in 136 guts. Highest feeding diversity and evenness were recorded in summer, while lowest values of both indices occurred in autumn. S. oconnori fed almost exclusively on periphyton. Diatoms were the most important prey in terms of number, weight and IRI, followed by macroinvertebrates; other prey were also occasionally ingested. S. oconnori is a generalized and opportunistic feeder relying upon a wide trophic spectrum, but mainly feeds on diatoms as they are the most prevalent periphyton. There were high diet overlaps among seasons, fish sizes and between sexes. S. oconnori showed no dietary preference for any of the other prey available in the periphyton community.     

Trophic diversity of the otter (Lutra lutra L.) in temperate and Mediterranean freshwater habitats

Aim To analyse the geographical patterns in the composition and diversity of otter's (Lutra lutra L.) diet and their relationship with climatic characteristics. Location European freshwater habitats under Mediterranean and temperate climatic regimes. Methods Thirty‐seven otter diet studies were reviewed, twenty‐one from temperate and sixteen from Mediterranean areas. All studies were based on spraint analysis and their results expressed as relative frequency of occurrence of seven main prey categories. Principal Component Analysis was performed to extract the main gradients of diet composition. Pearson's correlation and t‐tests were used to assess the relationship between diet characteristics (composition, diversity and taxonomic richness) and geographical and climatic variables. Results A clear latitudinal gradient in diet composition was observed. Otter diet was more diverse and featured more prey classes in southern localities, while the species was more piscivorous towards the north, where it predated upon a higher number of fish families. This pattern was similar when temperate and Mediterranean localities of Europe were compared. Mediterranean otters behaved as more generalist predators than temperate ones, relying less on fish, and more on aquatic invertebrates and reptiles. Main conclusions Geographical differences in otter feeding ecology in Europe seem to be related with the two contrasted climatic conditions affecting prey populations. The otter can act as a highly specialized piscivorous predator in temperate freshwater ecosystems, which do not suffer a dry season and have a comparatively stable water regime compared to Mediterranean ones. However, the unpredictable prey availability in Mediterranean areas, affected by strong spatial and temporal water shortages, favours a diversification of the otter's diet.     

Comparison of the feeding apparatus and diet of European sardines Sardina pilchardus of Atlantic and Mediterranean waters: ecological implications

In this study, the feeding apparatus (gill rakers, GR) and the diet composition of European sardine Sardina pilchardus populations living in two contrasting environments were compared: the upwelling area off western Iberia and the comparatively less productive region of the north‐western Mediterranean Sea. The importance of local adaptations in the trophic ecology of this species was estimated. Sardina pilchardus from the Atlantic Iberian coast and from the north‐western Mediterranean Sea have clear differences in the feeding apparatus and diet compositions. Those from the Atlantic Iberian coast have significantly more GRs than S. pilchardus of the same size range in the Mediterranean Sea. While S. pilchardus from the Mediterranean Sea mostly depend on prey ranging between 750–1500 and 3000–4000 µm, corresponding mostly to cladocerans, decapods and copepods, those from the Atlantic depend on smaller prey (50–500 and 1000–1500 µm) that include phytoplankton and copepods, particularly during summer months, and S. pilchardus eggs during the winter. The marked difference between the trophic ecology of S. pilchardus in the two areas studied appears to have originated from different dietary strategies that the two populations have adopted in contrasting feeding environments. These differences are shown to profoundly affect the size and quality of prey consumed, and the effect of cannibalism on the populations.     

Feeding ecology of the shortnose greeneye Chlorophthalmus agassizi Bonaparte, 1840 (Pisces: Chlorophthalmidae) in the eastern Ionian Sea (eastern Mediterranean)

The feeding ecology of Chlorophthalmus agassizi from the North Ionian Sea was examined in specimens caught during daylight hours on a seasonal basis from December 1996 to November 1997, at depths between 250 and 800 m (92 stations, total). Chlorophthalmus agassizi showed a mixed feeding strategy, exploiting a wide range of prey including mesopelagic, benthic and endobenthic organisms. Crustaceans and fishes were dominant food items in the diet year round. Diet composition showed slight seasonal fluctuations; these variations were correlated with food availability and reproductive activity. However, ontogenetic changes in the diet were relatively clear, despite the high overlap observed between small, medium and large individuals. Larger individuals are more efficient hunters that exhibit selection of prey with good swimming capabilities; smaller individuals consume prey with low mobility. Chlorophthalmus agassizi seem to be adapted to a food‐scarce environment, as typified in the deep‐water habitats of the Ionian Sea, and exploit all available niches.     

Diet of the Atlantic bonito Sarda sarda (Bloch, 1793) in the Northeast Aegean Sea

Seasonal and size‐related patterns in feeding habits of the Atlantic bonito (Sarda sarda) in the Northeast Aegean were investigated over a 2‐year period May 2007 – May 2009 based on analysis of stomach contents of 134 fish ranging from 266–655 mm fork length. The fish were caught by purse‐seine boats operating around the islands of Lesvos and Chios (Greece). Of the stomachs examined, 46% contained at least one prey item and significant seasonal and size‐related patterns were found in the occurrence of non‐empty stomachs. The overall diet composition in terms of numerical importance was 71% fish, 28% crustaceans and 1% cephalopods. Of the fish, Engraulidae (Engraulis encrasicolus) were the dominant prey taxon, followed by Centracanthidae (Spicara spp.) and then Clupeidae (Sardina pilchardus). Statistically significant monthly variation in diet was apparent, suggesting that S. sarda take advantage of seasonally abundant prey fish in the study area, but no size‐related variation in the importance of the main prey species was detected.     

Feeding habits and ontogenetic diet shifts of hairtail, Trichiurus margarites, in the Beibu Gulf of the South China Sea

The monthly and ontogenetic variations in diet of hairtail Trichiurus margarites Li (1992) [20] (Perciformes, Trichiuridae) were examined in the Beibu Gulf of the South China Sea from October 2008 to September 2009. Stomachs of 666 hairtail ranging in size from 121 mm to 561 mm pre-anal length (PL) were sampled monthly from commercial fish catches. Fish were the primary prey, with Decapterus maruadsi (Perciformes, Carangidae) averaging 23% by number, 34% by weight, and 49% by Index of Relative Importance (IRI). Other important prey in the diet included the small benthic fish, Bregmaceros rarisquamosus and Bregmaceros nectabanus (Gadiformes, Bregmaceroridae) and the decapod shrimp, Acetes chinensis, with IRI percentages of 8%, 4%, and 3%, respectively. Low rates of cannibalism and predation on 30 additional species of prey were observed. Trophic diversity (Brillouin Index, H_z) was moderate throughout the year. Mean feeding intensity (Repletion Index, RI) varied considerably by month and increased with the gonad maturity stages of hairtail; conversely, the proportion of empty stomachs decreased with maturity stages, from 60.9% empty at Stage I to 41.6% empty at Stage VI. Prey weight and numbers differed significantly among the 14 PL size classes and increased with body size. Cluster analysis of prey IRI values also indicated differences in the feeding habits of hairtail by size class. Understanding the monthly and ontogenetic changes in hairtail diet may assist in the development and improvement of marine ecosystem modeling for the Beibu Gulf, South China Sea.     

The diet of whiting Merlangius merlangus in the western Baltic Sea

The diet of whiting Merlangius merlangus in the western Baltic Sea was investigated and compared to the diet in the southern North Sea. Clupeids were important prey in both areas, but especially in the western Baltic Sea where they constituted up to 90% of the diet of larger individuals. Gobies, brown shrimps and polychaetes were the main prey of juveniles in the western Baltic Sea, while a wider range of species were consumed in the North Sea. The shift to piscivory occurred at smaller sizes in the western Baltic Sea and the fish prey consumed was proportionately larger than in the southern North Sea. Estimates of prey abundance and food intake of M. merlangus are required to evaluate its predatory significance in the western Baltic Sea, but its diet suggests that it could be just as significant a fish predator here as in the southern North Sea.     

Modeling and predicting potential spawning habitat of anchovy (Engraulis encrasicolus) and round sardinella (Sardinella aurita) based on satellite environmental information

Anchovy and round sardinella are two important small pelagic species in the Mediterranean that spawn during the summer period. This is a first attempt to model and predict the two species’ potential spawning habitats in this area. Generalized additive models (GAMs) were constructed based on satellite environmental variables and presence/absence egg data, available from ichthyoplankton surveys conducted in the North Aegean Sea during early summer (June 2003–2006). These models were subsequently used to predict the probability of anchovy and round sardinella spawning in the Greek Seas as well as the entire Mediterranean and Black Sea during the same month of the year. The interaction of bottom depth and chlorophyll explained most of the deviance in the presence/absence GAMs of both species, indicating spawning over continental shelf areas with increased surface chlorophyll values. Round sardinella spawned closer to coast than anchovy. Predicted potential spawning areas for anchovy and round sardinella in unsampled areas of the Greek Seas and the entire Mediterranean and Black Sea were in good agreement with existing information on the distribution and extent of the spawning grounds, especially for anchovy. Modeling the species’ reproductive activity in relation to easily accessible environmental information and applying the models in a predictive way could be an initial, low-cost step to designate potential spawning fish habitats. Guest editor: V. D. Valavanis Essential Fish Habitat Mapping in the Mediterranean     

Diet of the invasive piscivorous fish <Emphasis Type="Italic">Fistularia commersonii</Emphasis> in a recently colonized area of the eastern Mediterranean

The composition of the diet of the Indo-Pacific cornetfish Fistularia commersonii from the SE Aegean Sea is described. The stomach contents of 245 specimens collected between September 2004 and March 2005 were examined. Dietary comparisons were made reflecting the relationship between diet composition, time of year, and fish size. Correlation between predator length and prey length was significant. The diet of the blue cornetfish consists of 96% by number and 99.95% by weight of fish. Size classification and habitat of prey groups (benthic, supra-benthic, and pelagic) showed that with increased body length the blue cornetfish extended its diet to larger prey and more generalist feeding. Spicara smaris, Boops boops, and Mullidae spp. were the most abundant prey by weight whereas a variety of small benthic fish (especially gobiids) and newly hatched fish constituted the largest number of prey items. Length–weight relationships for the cornetfish were investigated.     

Long-Term Changes in the Diet of Gymnogobius isaza from Lake Biwa,Japan: Effects of Body Size and Environmental Prey Availability

Body size and environmental prey availability are both key factors determining feeding habits of gape-limited fish predators. However, our understanding of their interactive or relative effects is still limited. In this study, we performed quantitative dietary analysis of different body sizes of goby (Gymnogobius isaza) specimens collected from Lake Biwa between 1962 and 2004. First, we report that the diet was composed mainly of zooplankton (cladocerans and copepods) before the 1980s, and thereafter, shifted to zoobenthos (gammarids). This foraging shift coincided with, and thus can be linked to, known historical events in the lake at that time: decrease in zooplankton abundance with the alleviation of eutrophication, increase in fish body size resulting from fish population collapse, and increase in gammarid abundance due to reduced fish predation pressure. Supporting this view, our data analyses revealed how the long-term changes in the diet composition would be co-mediated by changes in fish body size and environmental prey availability. Specifically, while zoobenthos abundance strongly affected the fish diet composition, larger (smaller) fish preferred zoobenthos (zooplankton). Furthermore, the body size effects were stronger than those of prey availability. These results provide the best long-term evidence that fish feeding habits vary over decades with its body size and prey community due to anthropogenic disturbances.     

Diet of alewives,Alosa pseudoharengus and blueback herring,A. aestivalis (Pisces: Clupeidae) in Minas Basin,Nova Scotia,a turbid,macrotidal estuary

Synopsis Stomach contents of anadromous alewives, Alosa pseudoharengus and blueback herring, A. aestivalis, obtained from brush weir and drift net collections in Minas Basin, N.S., were examined. Diets showed much overlap in terms of resource use, but the dietary importance of major prey categories differed substantially between species. Alewives favoured larger, more benthic prey (e.g. amphipods, mysids and crangonids), while blueback herring appeared to concentrate their feeding on microzooplankters (e.g. calanoid copepods, cypris larvae and molluscan veligers). Interspecific differences in diet composition are largely attributed to the planktivorous feeding habits of small (81–155 mm fork length) blueback herring. Differences in prey suggest that alewives utilize a particulate feeding strategy while blueback herring are predominantly filter-feeders. Although competition for food in the Basin seems unlikely since high secondary production yields a superabundance of prey, differences in feeding behaviour between younger, smaller individuals of both species could be a means of avoiding competitive interactions in an environment where there are space/access limitations imposed by the tidal cycle.     

Feeding ecology of Mediterranean common dolphins: The importance of mesopelagic fish in the diet of an endangered subpopulation

The Mediterranean subpopulation of common dolphin (Delphinus delphis) is classified as endangered by the IUCN. Still, information about their diet in the Mediterranean is scarce. Stomach contents of 37 common dolphins stranded in the Alboran Sea and Strait of Gibraltar were analyzed. A total of 13,634 individual prey of 28 different taxa were identified. For fish, Myctophidae was the most important family as indicated by the highest index of relative importance (IRI = 8,470), followed by the family Sparidae (IRI = 609). The most important Myctophidae species was Madeira lantern fish (Ceratoscopelus maderensis) and for Sparids, the bogue (Boops boops). Cephalopods, instead, were found in low quantities only with 31 prey from the Loliginidae, Ommastrephidae, and Sepiolidae families. Overall, our results indicate that common dolphins are mainly piscivorous (99.77%N, 94.59%O, 99.73%W), feeding mostly on mesopelagic prey. Although common dolphins inhabit mainly coastal waters in the study area, the narrow continental shelf seems to facilitate the availability of Myctophids and other members of the mesopelagic assemblage to dolphins when the assemblage migrates to the surface at night. Our results represent the first attempt at quantifying the diet of this predator in the Alboran Sea and Strait of Gibraltar.     

Diet and foraging opportunism of the Guiana dolphin (Sotalia guianensis) in the Abrolhos Bank,Brazil

Diet analysis allows exploring how coastal dolphins interact with the environment and their role in the marine food webs. We studied the diet and feeding ecology of the Guiana dolphin, Sotalia guianensis, through analysis of stomach content from 42 animals stranded on the eastern coast of Brazil. A total of 1,336 semidigested prey items (fish, otoliths, cephalopod beaks, and crustaceans) were identified. Teleost fish comprised the most frequent food item (92% of the total), followed by cephalopods, and crustaceans. Prey belonged to 34 taxa and richness in individual stomachs varied from 1 to 15 prey taxa. Prey were generally small, but showed a significant trend to increase in size with dolphin length. The main prey of Guiana dolphins were demersal, estuarine, and sound-making fish such as catfish and sciaenids. No sex-related differences in diet were found. Diet composition varied seasonally and occurrence of prey items was coherent with breeding or high abundance periods of some fish species and squids. Our study shows the importance of demersal prey from estuarine and soft-sediment habitats to Guiana dolphin in the Abrolhos Bank and reveals that feeding habits are generalist and opportunistic, with diet reflecting the seasonal abundance and availability of prey.