Diet overlap among flatfish species in the southern North Sea

The diets of two non-commercial flatfish species (solenette Buglossidium luteum and scaldfish Arnoglossus laterna) and two commercial flatfish species (dab Limanda limanda and plaice Pleuronectes platessa) were compared in a study area in the German Bight (southern North Sea) to investigate prey-resource partitioning between these species. The diets of A. laterna and B. luteum mainly comprised crustaceans (harpacticoids, amphipods, cumaceans and decapods), whereas the diet of L. limanda and P. platessa consisted mainly of polychaetes. The Schoener index, calculated for different fish size classes between these flatfish species, showed a biologically significant diet overlap between small-sized L. limanda and P. platessa and B. luteum and A. laterna, using similar prey resources of smaller prey (e.g. amphipods, harpacticoids and juvenile bivalves). In contrast, with increasing body size, a change in the diet of L. limanda and P. platessa towards larger prey occurred (e.g. polychaetes and decapods), resulting in low diet overlap values with B. luteum and A. laterna. Due to these size-related differences in resource use, it is assumed that there is reduced interspecific competition for prey between larger L. limanda and P. platessa and both non-commercial flatfishes, probably facilitating resource partitioning within the same area. In contrast, smaller L. limanda and P. platessa may compete directly for the same prey resources with B. luteum and A. laterna. Furthermore, prey availability of most important prey items of the studied flatfishes was relatively low in the study area. Therefore, increasing abundances of B. luteum and A. laterna in the southern North Sea since the late 1980s, owing to fishing effects and climate change, might affect the population dynamics of L. limanda and P. platessa.     

Feeding ecology and growth of O-group flatfish (sole, dab and plaice) on a nursery ground (Southern Bight of the North Sea)

The food composition of O-group sole Solea solea , dab Limanda limanda and plaice Pleuronectes platessa on a nursery ground at Gravelines, France, included 17–25 taxa. Sole (new settlers) fed mainly on harpacticoid copepods and when 50 mm in size, on polychaetes (Terebellidae). Dab (<40 mm) consumed mainly polychaetes (Magelonidae and Spionidae), and later amphipods, polychaetes (Spionidae) and Hydrozoa. O-group plaice diet was dominated by polychaetes (Terebellidae), crustaceans and molluscs at all sizes. O-group sole, dab and plaice did not compete for food resources, each species being specialized in different prey items. Growth rates during May-July 1998 varied between 0·5 and 0·67 mm day⁻¹ for sole, 0·12 and 0·24 mm day⁻¹ for dab and 0·55 and 0·81 mm day⁻¹ for plaice. For sole and plaice, these estimates were similar to those recorded in other nurseries and also close to the maximal growth predicted by experimental models. This suggests that their growth was not limited by food during the first summer of life.     

On the morphology of the alimentary tract of flatfishes (Pleuronectiformes)

The changes during the life cycle in the relative length of certain parts of the alimentary tract, buccal-pharyngeal cavity, oesophagus-stomach, intestine, rectum—are investigated in the five North Sea flatfish species, viz. turbot, brill, plaice, dab, sole. According to the average ratio between the length of the alimentary tract and the food preference three main groups are distinguished.     

0-group flatfish growth conditions on a nursery ground (Bay of Canche, Eastern English Channel)

The settlement and growth conditions of three 0-group flatfish species, sole (Solea solea L.), plaice (Pleuronectes platessa L.) and dab (Limanda limanda L.) in the Bay of Canche, French coast of the Eastern English Channel, were investigated during summer 1997. 0-group dab were the more abundant of the three species (75.4%) followed by plaice (21%) and sole (3.6%). Analyses of fish size distribution indicated that settlement of plaice occurred before sole and dab. Growth of sole was faster than that of plaice, which in turn was faster than that of dab (0.68, 0.43 and 0.33 mm d^?1 respectively). Sole differed from plaice and dab by their larger mean length attained at the end of their first summer stay in the nursery ground. Growth performances of 0-group sole and plaice were analysed by comparing estimated growth in the field with predicted maximum growth according to temperature-growth rate models from experimental studies of growth with unlimited food supply. The data of this study suggest that plaice growth may have been limited during late summer, whereas sole growth does appear to have been determined only by water temperature.     

The feeding ecology of 0 year-group turbot Scophthalmus maximus and brill Scophthalmus rhombus on Irish west coast nursery grounds

On sandy beach nursery grounds along the west coast of Ireland, 0 year-group turbot Scophthalmus maximus were found to consume six types of crustaceans, in addition to polychaetes. The 0 year-group brill Scophthalmus rhombus fed almost exclusively on mysids, even though nine taxonomic prey groups were identified in the sediment across the investigated beaches. Both species avoided non-motile organisms such as gastropods and bivalves, which were present in high abundances in the sediment and their growth and condition was not significantly related to the quantity or type of prey consumed, temperature or salinity. A high incidence of feeding was detected for both species over the duration of the study, suggesting that food was not limiting on west of Ireland nursery grounds. Temporal partitioning of settlement was detected between S. maximus and S. rhombus, indicating that inter-specific competition for food does not occur between these two flatfish species on west of Ireland nursery grounds.     

Effects of bottom trawling on fish foraging and feeding

The effects of bottom trawling on benthic invertebrates include reductions of biomass, diversity and body size. These changes may negatively affect prey availability for demersal fishes, potentially leading to reduced food intake, body condition and yield of fishes in chronically trawled areas. Here, the effect of trawling on the prey availability and diet of two commercially important flatfish species, plaice (Pleuronectes platessa) and dab (Limanda limanda), was investigated over a trawling intensity gradient in the Irish Sea. Previous work in this area has shown that trawling negatively affects the condition of plaice but not of dab. This study showed that reductions in local prey availability did not result in reduced feeding of fish. As trawling frequency increased, both fish and prey biomass declined, such that the ratio of fish to prey remained unchanged. Consequently, even at frequently trawled sites with low prey biomass, both plaice and dab maintained constant levels of stomach fullness and gut energy contents. However, dietary shifts in plaice towards energy-poor prey items were evident when prey species were analysed individually. This, together with a potential decrease in foraging efficiency due to low prey densities, was seen as the most plausible cause for the reduced body condition observed. Understanding the relationship between trawling, benthic impacts, fish foraging and resultant body condition is an important step in designing successful mitigation measures for future management strategies in bottom trawl fisheries.     

Macrobenthic prey availability and the potential for food competition between 0 year group Pleuronectes platessa and Limanda limanda

This study describes the feeding habits of plaice Pleuronectes platessa and dab Limanda limanda during early juvenile development and relates differences between nursery grounds and sampling years to spatial and temporal variabilities in macrobenthic prey availability. The main prey taxa of both species were copepods, bivalves, amphipods, polychaetes and oligochaetes and size-related variation in diet was found. Despite considerable similarity in the prey items, differences in food composition between the two species were observed and spatial variability in diet confirmed their opportunistic feeding behaviour. A high degree of dietary overlap was found in June and decreased steadily throughout the season. The prey composition in the guts of both species largely reflected the composition of the main macrobenthic taxa in the sediment. The overall data suggest that resources were not limiting in the littoral sandy nursery areas in the west of Ireland and no indications were found that exploitative competitive forces upon the benthic prey assemblages occurred between P. platessa and L. limanda. Feeding success, Fulton's K condition and dietary overlap, however, showed spatial and temporal variations, and were probably affected by the availability and density of macrobenthic prey.     

Allozyme variation and genetic inter-relationships between seven flatfish species (Pleuronectiformes)

A total of 11 enzymes coded by 14 loci were assayed for each of 17 populations (from the north east Atlantic and Mediterranean) of seven flatfish species, including representatives of three of the four European families of the order Pleuronectiformes. Diagnostic alleles were observed for each species and there were fixed differences between the species at many loci. Thus all species were genetically distinct, although there were some common alleles. H_obs was higher than the average of a range of fish species and was also higher than that of vertebrate species as a whole. It seems that flatfish as a group may show higher levels of genetic variation than other fish. Values of genetic identity for all pairwise comparisons fell comfortably within the ranges expected. The data support the grouping of plaice and flounder into a single genus, Pleuronectes , but it is concluded that the retention of the dab in a separate genus, Limanda , is justified. A high level of genetic divergence was found between Dover and thickback soles. Genetic divergence data support the hypothesis that Pleuronectes flesus luscus in the Aegean Sea is a distinct subspecies of Pleuronectes flesus (flounder). The data show a clear separation of the Mediterranean Dover sole population from those in the Atlantic. Low genetic divergence was observed between the Aegean Sea and Atlantic brill populations. It is speculated that about 5 Mya the families Pleuronectidae and later Soleidae evolved from the ancestral Scophthalmidae.     

The feeding ecology of Pleuronectes platessa L., Limanda limanda (L.) and Scophthalmus rhombus (L.) in Carmarthen Bay, South Wales, U.K.

To assess the feeding habits of plaice, Pleuronectes platessa L., and dab, Limanda limanda (L.), within Carmarthen Bay, three sampling techniques were used: benthic trawling and surf-zone push-netting were undertaken during the feeding season for the gut contents of plaice and dab (age groups 0-IV) and of 0-group brill, Scophthalmus rhombus , to be analysed in detail, and a benthic macro-fauna grab survey covered the trawled ground to assess the distribution of food species in relation to gut contents. Plaice consumed commonly-occurring food species whereas dab took any available food. The main items were the same for both species, but secondary food preferences differed. Variations in gut contents within species, for the same sized fish, chiefly resulted from the spatial distribution of fish and not seasonal variations in the benthic macro-fauna. The length and not the age of fish primarily determined the selection of food organisms.     

Comparative study of growth of flatfishes on the west coast of Brittany

Lusitanian and boreal flatfish species live in the Bay of Douarnenez (Brittany, France); ten are abundant enough to be the subject of this growth investigation. In all species, females attain greater ultimate size than males, and oldest specimens are frequently female. Species are segre-gated to their longevity: those rapidly reaching their asymptotic length have the shortest life spans and also the highest values of the Von Bertalanffy parameter K .
In the bay, species of the boreal fish fauna (plaice, dab) find suitable environmental conditions for rapid growth; fish of the Lusitanian fauna (sand soles) grow slower than those living in southern areas.
In the Atlantic Ocean, the growth of the sole is better than in the North Sea and the Mediterranean.     

Drinking rates and Na+ effluxes in response to temperature change in two species of marine flatfish: dab,Limanda limanda and plaice,Pleuronectes platessa

The relationship between fish mass and drinking rate in two species of flatfish, dab and plaice, weighing between 1 and 150 g was investigated. Both plaice and dab showed increased drinking rates with increasing fish mass, although — when calculated on a weight-specific basis — the increase was negligible. Fish were acclimated to winter and summer temperatures of 9 and 14°C, respectively. In winter both species were acutely transferred to 5, 14 and 21°C and in summer to 5, 9, 21 and 25°C. Drinking rates, Na⁺ efflux and body ion content were measured. Dab showed lower drinking rates than plaice (e.g. the weight-specific drinking rates of summer-and winter-acclimated dab were 0.12±0.01 and 0.06±0.006 ml·h^-1·100 g^-1, respectively, compared to that of plaice which were 0.25±0.02 ml·h^-1·100 g^-1 in summer and 0.17±0.02 ml·h^-1·100g^-1 in winter). Summer dab exhibited decreased weight-specific drinking rates at 5, 9 and 25°C, while winter dab increased drinking at 21°C. Winter plaice also showed increased drinking at 21°C and a decrease at 5°C, but in contrast summer plaice did not increase drinking at either 21 or 25°C but showed a decrease at 5 and 9°C. Winter dab and plaice showed similar Na⁺ efflux rates but summer dab showed higher efflux at all temperatures except 5°C. The data indicates that (a) the osmoregulatory function of plaice is much weaker than that of dab at higher temperatures (>20°C) and (b) mass has a greater effect on drinking and Na⁺ efflux rates than temperature; although when calculated on a weight-specific basis neither drinking nor efflux showed any variation with fish mass suggesting that these functions occur at similar intensities across the entire weight range.Abbreviations ANOVA analysis of variance - ⁵¹Cr-EDTA ⁵¹chromium ethylenediaminetetra-acetic acid - SW sea water     

Habitat and infauna prey availability for flatfishes in the northern Bering Sea

Yellowfin sole (Limanda aspera), northern rock sole (Lepidopsetta polyxystra), and Alaska plaice (Pleuronectes quadrituberculatus) are valuable flatfishes in the southeastern Bering Sea (EBS) bottom trawl fishery. The northern Bering Sea (NBS) is near their northern distribution limit. We conducted the first assessment of NBS habitat suitability for these benthivorous flatfishes from the perspective of prey availability in 2010. Benthic samples were collected at 12 trawl stations along a meridional transect extending from 60.5°N to 64.5°N east of St. Lawrence Island to characterize the prey environment. Stomach contents from the flatfishes were concomitantly collected to relate diets to prey fields. The diet compositions did not correspond spatially with the infauna communities. The flatfishes elected a prey group regardless of its relative availability. The spatial mismatch between diet and infauna compositions suggests that prey availability was high in the NBS. The flatfishes generally have versatile diets, but they were more selective of their prey here than in the EBS. The biomass and the abundance of the infauna along the transect were comparable with the EBS. Although niche overlap was high in the NBS, competition for food was likely lower than in the EBS because of the lower density of flatfish. The bottom temperatures in the NBS were in the same range as the EBS during the summer of 2010. The NBS appears to be suitable flatfish habitat at least during the summer ice-free period. The effects of climate warming on the prey environment and on the production and distribution of flatfish are complex and difficult to predict, but if the NBS were to shift over time toward milder winter conditions that more resemble the EBS, its suitability as flatfish habitat would presumably increase based on the present prey availability.     

The diets of two co-occurring marine teleosts, Parequula melbournensis and Pseudocaranx wrighti, and their relationships to body size and mouth morphology, and the season and location of capture

The dietary compositions of the co-occurring gerreid Parequula melbournensis and the carangid Pseudocaranx wrighti have been determined, using samples collected seasonally from a 200 km stretch of coastal water in temperate Australia, in which these species are very abundant. Although the small representatives of P. melbournensis and P. wrighti both fed to a large extent on copepods, the latter ingested a wider variety of prey and thus had a greater dietary breadth. The diets of both species changed markedly as body size increased. The larger representatives of P. melbournensis fed mainly on onuphid and other polychaetes, whereas those of P. wrighti consumed a considerable volume of crustaceans, molluscs, polychaetes other than onuphids, and echinoderms. The above differences account for the dietary breadth being far greater in large P. wrighti than large P. melbournensis. Schoener's index and classification showed that dietary overlap between P. melbournensis and P. wrighti was low, suggesting that food resources were partitioned between these two demersal species. Intraspecific overlap was less amongst the length classes of small fish than among those of larger fish, indicating that any competition for food would be less among small fish, when growth would have been most rapid. The larger P. melbournensis fed mainly on prey types which were relatively sessile and can protrude from the substrate, such as tube-dwelling onuphid polychaetes, whereas the larger P. wrighti fed on a variety of epibenthic and/or more mobile species, such as mysids, amphipods, bivalves, gastropods, nereid polychaetes and echinoderms. Although P. melbournensis foraged to a far greater extent on a sessile fauna that occurs on or in the substrate, it ingested a far smaller amount of sand than P. wrighti, even though this latter species fed on a more epibenthic fauna. The presence of smaller amounts of sand in the stomach contents of P. melbournensis than of P. wrighti, presumably reflects the possession of a smaller and far more protrusible mouth, which enabled its sessile prey to be targetted more precisely. The dietary composition of P. melbournensis and P. wrighti underwent some seasonal changes, presumably reflecting seasonal fluctuations in the densities of prey species, and that of P. melbournensis differed slightly between some sites.     

Effect of light on the feeding rates of pelagic and littoral mysid shrimps: a trade-off between feeding success and predation avoidance

It has been shown that freshwater pelagic mysid shrimps capture zooplankton at a higher rate in light than in darkness. This has been suggested to be due to facilitation of visual predation on evasive zooplankton prey. To test this hypothesis with Baltic mysid shrimps, and to see whether pelagic (migrating) and littoral (non-migrating) mysids differ in this respect, we compared the feeding rates of Mysis mixta and Praunus flexuosus on the copepod Acartia spp. As light conditions change radically from the beginning of summer to late autumn at the Baltic latitudes, we conducted the experiments during three different times of the year to determine if there was a seasonal response to light in mysids. The feeding rates of pelagic mysids were significantly higher in total darkness than in light. In contrast, the feeding rates of littoral mysids did not differ in the dark and the light in the three different seasons. The decreased feeding of pelagic mysids under well-lit conditions may be an adaptation to avoid visual predation by pelagic fish. In contrast, littoral mysids, which live in the well-lit layer throughout the day, do not show a similar response. The fact that light did not increase feeding in either of the mysid species indicates that these mysid species do not use vision in capturing prey, but rely mainly on mechano-reception.     

Food habits of the sandpaper skate, Bathyraja kincaidii (Garman, 1908) off central California: seasonal variation in diet linked to oceanographic conditions

The stomachs of 130 sandpaper skates, Bathyraja kincaidii (Garman, 1908), were sampled from off central California to determine their diet composition. The overall diet was dominated by euphausiids, but shrimps, polychaetes and squids were also important secondary prey. A three-factor MANOVA demonstrated significant differences in the diet by sex, maturity status and oceanographic season using numeric and gravimetric measures of importance for the major prey categories. These three main factors explained more variation in diet than interactions between the factors, and season explained the most variance overall. A detailed analysis of the seasonal variation among the prey categories indicated that abundance changes in the most important prey, euphausiids, were coupled with seasonal changes in the importance of other prey. When upwelling occurred and productivity was great (Upwelling and Oceanic seasons), euphausiids were likely highly abundant in the study area and were the most important prey for B. kincaidii. As productivity declined (Davidson Current season), euphausiids appeared to decrease in abundance and B. kincaidii switched to secondary prey. At that time, gammarid amphipods and shrimps became the most important prey items and polychaetes, mysids and euphausiids were secondary.     

The biology of the scaldfish, Arnoglossus laterna (Walbaum) on the west coast of Scotland.

The biology of a Scottish population of the small bothid flatfish, Arnoglossus laterna , was studied from January 1975 until September 1976. The data were taken from monthly samples totalling over 500 fish trawled in 18–36 m on a soft mud bottom. Otoliths were used for age determination and a growth curve was constructed which showed that most growth occurs in the first 2–3 years of life. The maximum age recorded was 8+ years. The fish first mature sexually in their second year at a standard length of 6–7 cm and the short spawning season lasts from the late June to August. Fecundity is length-dependent and the relationship could be described by the regression equation: log fecundity = 3·3472 log standard length (mm) -2·1064. The diet consists mainly of decapod crustaceans (particularly crangonid shrimps), polychaetes, mysids and small fish.     

Analysis of diet composition and overlap in four species of the genus Diplectrum (Perciformes: Serranidae) in the Mexican Central Pacific

Analysis of diet composition and overlap in four species of the genus Diplectrum (Perciformes: Serranidae) in the Mexican Central Pacific. The information of trophic interactions among species is essential to understand ecosystem function. To assess this in four Diplectrum species, we analyzed the stomach contents of 397 individuals caught using shrimp trawling nets off the coasts of Jalisco and Colima, Mexico. Main food component of D. eumelum were fish of the Order Pleuronectiformes, followed by shrimps (Metapenaeopsis spp). D. euryplectrum feeds mainly on crustaceans, with stomatopods (Squilla mantoidea) as main food items followed by shrimps and brachyurans. Fish and polychaetes had a low representation in the diet of this species. D. labarum feeds mainly on crustaceans, mollusks and fish. The stomatopod Eurysquilla veleronis contributed to with a high diet percentage by weight, followed by shrimps (mostly Solenoceraflorae) and brachyurans, as well as the squid Lolliguncula diomedae. Fishes (Ophidion spp.) were also an important component of the diet of this species. The most common preys of D. rostrum were crustaceans, mollusks, polychaetes and fish. The squid Lolliguncula diomedae and fishes of genus Ophidion were also well represented in the diet. Diet breadth index yielded significant differences between species. The number of diet items varied from 16 in D. euryplectrum to 6 in D. eumelum. The average value of overlap (0.247) was not significantly different (p = 0.118) from the expected one for a null model (0.174). The observed variance of the overlap index (0.071) was significantly higher (V = 0.025, p = 0.0004) than the value expected for a null model.     

Food habits and trophic status of Limanda sakhalinensis (Pleuronectidae) on the western Kamchatka shelf in the summer period

Based on materials of expeditions of TINRO-Center on the western Kamchatka shelf in July 2005, 2008, and 2010, we analyze data on feeding of Sakhalin flounder. Sakhalin flounders are consumers of euphausiids, copepods, amphipods, polychaetes, and juvenile fish. The calculated values of the trophic levels are 3.58–3.95. Consumption of maturing and mature Sakhalin flounder primarily plankton prey leads to divergence of ecological niches with morphologically close yellowfin sole, extends prey species, and apparently provides the basis for the dominance of the western Kamchatka shelf community. Mature and maturing Sakhalin flounders feed predominantly on plankton prey, which leads to a divergence of their ecological niche with morphologically close yellowfin sole and expanding food supply and apparently is the basis of the dominance of flatfish community of the western Kamchatka shelf.     

Comparative Feeding Ecology of Two Sympatric Greenling Species, <Emphasis Type="Italic">Hexagrammos otakii</Emphasis> and <Emphasis Type="Italic">Hexagrammos agrammus</Emphasis> in Eelgrass <Emphasis Type="Italic">Zostera marina</Emphasis> Beds

Synopsis Feeding ecology was compared between sympatric greenling species, Hexagrammos otakii and H. agrammus in the eelgrass beds in Jindong Bay, Korea, from January to December. These two species had similar diets composed of crustaceans, polychaetes, gastropods and fishes; both species consumed primarily crustaceans throughout study periods. H. otakii, however, fed a greater proportion of polychaetes and fishes than H. agrammus. H. agrammus had a greater proportion of gastropods in their diets. The diet of both species underwent size-related changes; smaller individuals of H. otakii and H. agrammus consumed amphipods (gammarid amphipods and caprellid amphipods), while larger individuals of H. otakii ate polychaetes and fishes and those of H. agrammus fed mainly on gastropods and crabs. The diet of H. otakii underwent seasonal changes; H. otakii consumed mainly polychaetes and fishes during January and February 2002 but amphipods during March and May 2002. H. agrammus, however, ate mainly gastropods and crabs all seasons. H. otakii underwent also a significant diel changes that could be related to difference of prey availability. Thus the nocturnal emergence of gammarid amphipods, polychaetes and fishes explained their greater consumption by H. otakii. Dietary breadth of both species was lower in the smallest individuals (<5 cm SL) and in March and April 2002. This was due to the disproportionate dry mass attributable to the consumption of amphipods by both H. otakii and H. agrammus. Dietary overlap of both species was relatively moderate to high, in particular in <9.9 cm SL (0.62 – 0.71) from May to July 2002 (0.63 – 0.71). This is coincident with higher abundances of crabs, caridean shrimps and polychaetes in the study area, and it was assumed that these prey species were not limited resources. Higher dietary overlap was correlated with an abundance of a shared resource and did not indicate the interspecific competition between H. otakii and H. agrammus.     

Sexual, ontogenetic, temporal and spatial effects on the diet of Urotrygon rogersi (Elasmobranchii:Myliobatiformes)

The food habits and trophic ecology of Urotrygon rogersi were analysed to ascertain sexual, ontogenetic, temporal and spatial effects on the diet. These effects were examined with contingency tables, simple correspondence analysis and MANOVA. The trophic relationships of the species were determined with Levin's niche amplitude index and Pianka's diet overlap index. Urotrygon rogersi is a predator of benthic organisms, especially shrimp and polychaetes. Analysis of sex and time showed no significant differences, but differences among class sizes were found. A strong trend towards diet specialization and a strong overlap between sexes and between class sizes were found. Finally, fishing activity on shrimps in the study area could force U. rogersi to change its trophic spectrum over time as an adaptation to the considerable reduction in its principal prey.