Diet of two commercially important shark species in the United Arab Emirates: milk shark,Rhizoprionodon acutus (Rüppell, 1837), and slit‐eye shark,Loxodon macrorhinus (Müller & Henle, 1839)

The diets of the milk shark, Rhizoprionodon acutus, and the slit‐eye shark, Loxodon macrorhinus, landed from the artisanal fishery in the Arabian Gulf waters of the United Arab Emirates were investigated to determine their dietary preferences. Stomach contents from 57 milk sharks and 53 slit eye sharks were collected from Abu Dhabi (R. acutus, n = 23), Dubai (R. acutus, n = 5; L. macrorhinus, n = 15) and Ras Al Khaimah (R. acutus, n = 29; L. macrorhinus, n = 38) during fishery surveys from January to May 2012. Prey items were identified to the lowest possible taxonomic level, grouped into five categories including ‘teleost fish’, ‘cephalopods’, ‘crustaceans’, ‘invertebrates’, and ‘other’. The diets of both species were described using the numeric, frequency and weight methods, and the index of relative importance (IRI). The majority of stomachs for both species had food, with 66.6% of milk shark stomachs and 90.5% of slit‐eye shark stomachs containing prey items, both dominated by small teleosts. Rhizoprionodon acutus fed on a wide variety of teleost species, primarily Engraulidae (anchovies) (28%), Gerreidae (mojarras) (5.6%) and Carangidae (jacks) (1.6%) with occasional crustacean and cephalopod prey (8%). On the other hand, L. macrorhinus seemed to have a preference for one species in terms of teleosts (anchovies) (35.1%) and fed on a wider variety of crustaceans and cephalopods (22.6%). There was little overlap in the diets of these two species, suggesting that they may either be using different habitats or that in these waters, the milk shark is a generalist species while the slit‐eye is a specialist feeder.     

Stomach content analysis of juvenile, scalloped hammerhead shark Sphyrna lewini captured off the coast of Mazatlán, Mexico

We quantified the diet of juvenile, scalloped hammerhead shark Sphyrna lewini in the area off Mazatlan, Sinaloa, Mexico, to understand their feeding ecology this shark. The prey species of Sphyrna lewini were identified and quantified from stomach content analysis. In addition, we determined the variations between genders. During two fishing seasons (2000–2001 and 2001–2002), we analyzed 232 stomachs, of which 85% contained food. The trophic spectrum was composed of three species of cephalopods, six of crustaceans and 19 species of fish from mainly pelagic and benthic habitats. According to the Index of Relative Importance (%IRI), the cephalopod Loliolopsis diomedeae with IRI = 18%, fish of the family Carangidae IRI = 25% and family Synodontidae IRI = 19% constituted the main prey in general. The trophic niche width was <0.4, which indicated that S. lewini juveniles in this area feed on a wide range of prey items, though they showed a preference for a few prey items.     

Temporal variation in chum salmon, Oncorhynchus keta, diets in the central Bering Sea in summer and early autumn

Seasonal, ontogenetic, and diel variations in the diets of chum salmon, Oncorhynchus keta, were examined by analyzing the stomach contents of 1398 fish (300–755 mm fork length) collected in the Bering Sea during summer and early autumn of 2002. Whereas mesozooplankton, including euphausiids, hyperiids, and gastropods, constituted the greatest portion of the stomach contents during the summer, forage fishes (Stenobrachius leucopsarus and Atka mackerel, Pleurogrammus monopterygius) were the most important items during early autumn. Although no apparent diel trend was found in feeding intensity, distinct diel differences in prey composition were observed. Chum salmon caught in the morning contained Stenobrachius leucopsarus, whereas those caught in the afternoon had mainly fed on euphausiids. Thus, chum salmon diets change temporally because of changes in prey availability that result from differences in the annual life cycles and diurnal vertical migrations of prey species.     

Feeding habits of the king soldier bream,Argyrops spinifer (Forsskål, 1775) (Perciformes: Sparidae), in the northern Persian Gulf

The feeding habits of the king soldier bream Argyrops spinifer (Forsskål, 1775), were examined in the northern part of the Persian Gulf (coastal waters of Bushehr, Iran). The stomach contents of 622 specimens, 13.5–64.2 cm in total length, collected by a trawl boat from June 2010 to May 2011, were analysed to assess the influence of size, season and sex on their diet composition. Amongst the examined stomachs, 57 were empty (vacuity index = 9.2%), which varied significantly (P < 0.05) throughout the year. According to the results, this species is a generalist carnivorous feeder with a diet consisting of 52 different prey items belonging to nine main categories. Bivalvia, Gastropoda, Malacostraca, Crustacea and Teleostei were the most important prey items. Fish size and gender showed no significant effects on diet composition. Moreover, in the main food catagories no seasonal variations were detected.     

Feeding habits of anglerfish,Lophius budegassa (Spinola, 1807) in the Adriatic Sea,north‐eastern Mediterranean

Stomach contents of 283 anglerfish (Lophius budegassa), collected in the Adriatic Sea (north‐eastern Mediterranean) from experimental trawl surveys in 2005 and 2006, were examined to assess their diet as influenced by fish size, sex and water depth. Changes in food quantities in the stomach were observed, the percentage of empty stomachs decreasing with increasing body size and depth. No differences were found between male and female diets. The anglerfish is essentially an ‘opportunistic feeder’ that consumes mainly fishes, crustaceans and molluscs. Fishes were the primary food consumed by all size classes; the favourite prey was Merluccius merluccius (TL >150 mm for medium and large anglerfish) and Gaidropsarus biscayensis (TL <150 mm for smaller anglerfish). However, the proportion of fish species and molluscs changed with the body size of the anglerfish. The trophic level of L. budegassa estimated in the study area reached a mean value of 4.38, confirming that the species was a carnivore with a preference for large decapodes, cephalopods and fish.     

Diet assessment of the Atlantic Sea Nettle Chrysaora quinquecirrha in Barnegat Bay,New Jersey,using next‐generation sequencing

Next‐generation sequencing (NGS) methodologies have proven useful in deciphering the food items of generalist predators, but have yet to be applied to gelatinous animal gut and tentacle content. NGS can potentially supplement traditional methods of visual identification. Chrysaora quinquecirrha (Atlantic sea nettle) has progressively become more abundant in Mid‐Atlantic United States’ estuaries including Barnegat Bay (New Jersey), potentially having detrimental effects on both marine organisms and human enterprises. Full characterization of this predator's diet is essential for a comprehensive understanding of its impact on the food web and its management. Here, we tested the efficacy of NGS for prey item determination in the Atlantic sea nettle. We implemented a NGS ‘shotgun’ approach to randomly sequence DNA fragments isolated from gut lavages and gastric pouch/tentacle picks of eight and 84 sea nettles, respectively. These results were verified by visual identification and co‐occurring plankton tows. Over 550 000 contigs were assembled from ~110 million paired‐end reads. Of these, 100 contigs were confidently assigned to 23 different taxa, including soft‐bodied organisms previously undocumented as prey species, including copepods, fish, ctenophores, anemones, amphipods, barnacles, shrimp, polychaete worms, flukes, flatworms, echinoderms, gastropods, bivalves and hemichordates. Our results not only indicate that a ‘shotgun’ NGS approach can supplement visual identification methods, but targeted enrichment of a specific amplicon/gene is not a prerequisite for identifying Atlantic sea nettle prey items.     

Feeding strategy of silverside species in eutrophic Lake Pátzcuaro,Mexico

The trophic guild, niche breadth, and trophic position of two different ontogenetic phases were measured according to the size of Chirostoma spp. captured in different zones of Lake Pátzcuaro, Mexico. Analyses were carried out at the genus level due the difficulties to find diagnostic characters to separate the three small silverside species accurately, since even the molecular analyses of mitochondrial DNA for a subsample of the most morphologically differentiated specimens were conducted without success. Fish were captured with a seine net during the wet (September and November 2009) and dry (February and June 2010) seasons at four environmentally different sites. Gut contents of 480 Chirostoma specimens were divided into size‐class intervals of 38–51 mm and 66–71 mm for analyses. The contribution of each food item was quantified using the frequency of occurrence and area percentage. Fish were categorized by size, and diets were compared between fish sizes and sites. Dorsal muscle tissue and zooplankton were obtained for nitrogen isotope signature (δ¹⁵N) analyses. The trophic guild was determined using a modified quadrant method for gut content analysis; the importance of each prey item was estimated according to a modified index of relative importance. Niche breadth was calculated by the Levins’ Index. The trophic position was assessed using the TrophLab Program and stable isotope analysis. The index of relative importance among study sites and fish size classes was determined by a multivariate approach. Silversides represent a secondary and tertiary consumer in Lake Pátzcuaro that feed mainly on evasive prey, such as copepods (55%) and on non‐evasive prey, such as cladocerans (25%) at sites with the lowest transparency in the water column (from 350 mm to 150 mm). Fish scales (11%) and insects (6%) were secondary food items. Ingestion of juvenile silversides was rare (1%). Chirostoma spp. prey on insects on the lake bottom and on water hyacinth roots. Based on δ¹⁵N analysis, the trophic position of silversides changes between wet and dry seasons. Their opportunistic and specialized predatory feeding strategy indicates the capability of silversides to resist anthropic changes in the lake.     

Veligers of an introduced bivalve,Limnoperna fortunei,are a new food resource that enhances growth of larval fish in the Paraná River (South America)

1. Larvae of ‘sábalo’, Prochilodus lineatus, whose adults represent over 60% of overall fish biomass in the Río de la Plata Catchment, have been observed to feed intensively on veligers of the exotic bivalve Limnoperna fortunei. 2. To assess the effects of this dietary shift on the growth of P. lineatus, 28‐day laboratory experiments were carried out feeding newly hatched P. lineatus larvae with three diets: zooplankton artificially enriched with L. fortunei veligers; natural zooplankton; and zooplankton artificially enriched with cladocerans and copepods. The average length, weight and gut contents of the fish larvae were assessed weekly and metabolic rates of fish larvae were measured. 3. Proportions of veligers in gut contents were always higher than those in the experimental diet: 100, 76 and 21% for veliger‐enriched, natural and low‐veliger diets, respectively. Larvae fed a veliger‐enriched diet grew to a significantly larger size than larvae fed the other two diets. In energetic balance comparisons using metabolic rates and prey energy content, all three diets were sufficient to support metabolism and growth. The greatest values of excess energy at the end of each week were in the veliger‐enriched experiments. 4. Feeding on veligers of L. fortunei significantly enhances the growth of P. lineatus larvae and supports the idea that this new and abundant resource is selectively preyed upon by P. lineatus during its larval stage. Higher growth rates may stem from the higher energy contents of veligers compared to crustaceans and/or from the lower energy costs of capturing slower prey.     

Diet niche relationships among predator and prey fish species in their early life stages in Lake Võrtsjärv (Estonia)

In Lake Võrtsjärv pikeperch was observed not to shift to piscivory in their first autumn of life, although juvenile stages of a variety of fish species were abundant in the lake. It was hypothesized that the diets of predator and prey fish fry overlap and that coarse fish species are important food competitors for juvenile piscivores and thus, pikeperch and perch fry do not shift to piscivory during their first growing season. To discover the possible linkages in this pattern, in 2009 the feeding relationships of pikeperch, perch, ruffe and roach fry were analysed. The stomach content analyses showed that in the summer period, Mesocyclops leuckarti was the most frequent prey for perch and ruffe, pikeperch consumed Leptodora kindti in large quantities, and roach ate mostly plant material. Towards autumn, M. leuckarti was the most abundant prey for all percids. However, average stomach content weight and the number of prey items eaten by ruffe were considerably higher than for other fish fry. Since the feeding opportunities of fish fry are considered poor in the examined lake, the prey has the potential to restrict the recruitment to piscivory of their predators, as prey fish seem to have better abilities to persist in this ecosystem. Furthermore, supposed competition in the juvenile stage may result in a reduced top‐down effect on coarse fish.     

Ontogenetic dietary shifts and morphological correlates in striped mullet,Mugil cephalus

Synopsis Ontogenetic dietary shifts are well studied in fishes. However, these shifts are rarely correlated with habitat and morphology. This study investigates: (1) ontogenetic dietary changes in mullet; and if dietary shifts do occur (2) whether they are correlated with feeding location, feeding behavior and/or feeding morphology. Stomach contents from eight size classes, each with 25 individuals, were analyzed using a gravimetric sieve fractionation procedure, Czechanowski's Quantitative Index and cluster analysis. Core samples (5 mm deep) were also analyzed in the same manner. Morphological characters were measured to detect differences among the size classes. Cluster analysis showed three distinct trophic units for Mugil cephalus in Cross Bayou: (1) 20.0 mm standard length — 30.0 mm SL; (2) 30.0 mm SL — 40.0 mm SL; and (3) 40.0 mm SL — > 100 mm SL. Comparison of the gut contents to the sediment contents for each size class showed an increase in similarity as the fish grew. Mullet gut contents from fish 20–25 mm SL were only 45% similar based on the weights of the prey items to the sediment subsample prey items, while stomach contents from fish > 100 mm SL were 84% similar to the sediment subsample contents. Gill raker length and interraker spacing of four gill arches were allometric with respect to standard length. Prey particle size within size classes appeared to be more correlated with feeding behavior than gill interraker spacing because smaller particles than predicted by the interraker spacing were ingested. It is suggested that smaller fish may be browsing more selectively than larger fish which graze.     

A kinematic investigation into the feeding behavior of the Goliath grouper <Emphasis Type="Italic">Epinephelus itajara</Emphasis>

Goliath grouper Epinephelus itajara suffered significant overfishing in the United States until they were protected from harvest in 1990. As the population recovers, interactions between Goliath grouper and anglers have increased, and are often reported to management agencies as complaints after grouper predation upon hooked fish. Goliath grouper are generally characterized as opportunistic predators capable of consuming a wide variety of prey types; however, minimal data are available regarding the prey capture behavior of this species. Kinematic analyses of adult Goliath grouper feeding events demonstrated the capacity of individuals to modulate feeding behavior based upon the mobility and position of ‘prey’ items. Mobile epibenthic food (tethered swimming fish) elicited larger maximum gapes, faster times to food capture, shorter times to mouth closing, and more rapid total bite durations than food items that were not moving (cut dead bait). Feeding sequences involving mobile food items were characterized by a significantly higher degree of ram feeding behavior, while immobile food elicited primarily suction feeding and were preceded by a slower and closer approach to the food item prior to the onset of mouth opening. The findings are discussed in light of predation upon angled species and demonstrate the ability of Goliath grouper to adjust their feeding strategy based upon prey type and condition. This behavior likely allows for the exploitation of a wide variety of prey and provides an expansive dietary breadth for these opportunistic predators.     

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.     

Diet and feeding habits of Economidichthys pygmaeus (Perciformes: Gobiidae) in Lake Pamvotis,NW Greece

The purpose of the study was to define the diet composition, feeding preferences, seasonal, size and sex related changes in diet, feeding strategy and diel cycle of Economidichthys pygmaeus. Important features of the overall feeding patterns of the goby include (i) feeding activity, (ii) bottom habits and (iii) consumption of food items to identify the dietary breadth. Stomach contents of 533 specimens, 13.42–48.65 mm total length (TL), collected by hand net (2 mm mesh size) from October 2006 to September 2007 were analyzed. The percentage of 76 empty stomachs (14.26%) did not vary significantly with season. The food composition suggested that the goby is a carnivorous fish. Prey items identified in the stomachs belonged to four major groups: Crustaceans, Insects, Mollusca and Plants. Copepods were the most important (%IRI = 57.51), especially in fish larger than 23 mm TL. Chironomids (%IRI = 43.23) constituted the main prey for fish <23 mm TL. Diet composition showed little seasonal variation. Copepoda dominated the diet in autumn and winter (43.4%; 82.4%) and were replaced by Chironomidae larvae in spring (39.7%) and summer (47.7%). During the reproductive season, large males show a narrow food spectrum as opposed to large females, reflecting the different activity pattern exhibited by males and females during the breeding season, since the former supports parental care. According to the modified Costello graphical method, specialist individuals form the goby population that feeds on two preferential prey types (e.g. Copepods, Chironomids). Nevertheless, they consume some occasional prey (e.g. Cladocera, Insects, Gastropods). The species foraged chiefly at night and early morning, during lower light intensity. The highest feeding activity recorded at night (02.00 hours) and during early morning (08.00 hours) could be related to the cryptic behaviour displayed by the species.     

Effects of ram speed on prey capture kinematics of juvenile Indo-Pacific tarpon,Megalops cyprinoides

We examined the effects of variation in swimming speed, or ‘ram speed’, on the feeding kinematics of juvenile Indo-Pacific tarpon, Megalops cyprinoides. Tarpon were filmed feeding on non-elusive prey at 500 images s^?1. Prey items were offered at one end of the filming tank, the opposite end where tarpon grouped, to encourage them to use a ram strategy to capture their prey. We describe tarpon as ram-suction feeders. Ram speed varied among strikes from 0.19 to 1.38 m/s and each individual produced speeds that spanned at least 0.9 m/s across trials. Although suction distances were much less variable, prey movement towards the predator was present in all feeding trials. There was a strong positive relationship between initial predator – prey distance and ram speed (r²=0.72, P<0.001). When tarpon initiated their strike from further away, they achieved higher ram speeds, but also took longer to capture their prey. All other timing variables were unaffected by ram speed whereas at higher ram speeds tarpon exhibited greater expansion of the mouth and buccal cavity. Greater buccal expansion accomplished in the same period of time implies that both the total volume of water captured and the water flow rate entering the mouth was greater in strikes at higher ram speeds. Our results demonstrate how feeding kinematics may vary as a function of ram speed, and how fish predators that lack jaw protrusion and have a large gape capacity can maximize their feeding success by altering their swimming speed.     

Utilizing stomach content and faecal DNA analysis techniques to assess the feeding behaviour of largemouth bass Micropterus salmoides and bluegill Lepomis macrochirus

In this study, the feeding behaviour of the non‐native invasive predatory fishes largemouth bass Micropterus salmoides and bluegill Lepomis macrochirus was studied in the Ezura River, a northern tributary of Lake Biwa, Japan. Prey composition was estimated based on visual examination of stomach contents and faecal DNA analysis to determine feeding habits of these predatory fishes. Stomach content analysis showed that native fishes (e.g. ayu Plecoglossus altivelis and gobies Rhinogobius spp.) and shrimps (e.g. Palaemon paucidens) were the major prey items for M. salmoides, while snails, larval Chironomidae and submerged macrophytes were the dominant prey items of L. macrochirus. Micropterus salmoides tended to select larger fish in the case of crucian carp Carassius spp., but smaller fishes in the case of P. altivelis and Rhinogobius spp. Faecal DNA analyses revealed prey compositions similar to those identified in predator stomach contents, and identified additional prey species not detected in stomach content inspection. This study demonstrated that both stomach content inspection and DNA‐based analysis bear several inherent shortcomings and advantages. The former method is straightforward, although identification of species can be inaccurate or impossible, whereas the latter method allows for accurate species identification, but cannot distinguish prey size or stage. Hence, integration of morphology‐based and DNA‐based methods can provide more reliable estimates of foraging habits of predatory fishes.     

Food habits of small fishes in a common reed <Emphasis Type="Italic">Phragmites australis</Emphasis> belt in Lake Shinji,Shimane, Japan

To clarify the feeding habits of reed fishes, the gut contents of 13 fish species collected in a Phragmites australis belt in Lake Shinji were examined. Six species showed ontogenetic and/or seasonal changes in food use patterns. Smaller individuals generally preyed on small planktonic items (e.g., calanoid and cyclopoid copepods) or small crustaceans (gammaridean amphipods), subsequently changing to other prey items (e.g., mysids and filamentous algae) with growth. The most important dietary items for the reed fish assemblage comprised planktonic copepods, gammaridean amphipods and mysids. However, the relative importance of these changed seasonally, gammaridean amphipods being the most important in autumn and winter, and planktonic copepods and mysids the most important in spring and in summer. Cluster analysis based on dietary overlaps showed that the reed fish assemblage comprised five feeding guilds (planktonic-copepod, mysid, gammaridean-amphipod, filamentous-algae, and detritus feeders). Of these, the three former guilds were the most abundantly represented, whereas detritivores were represented by a single species.     

Constraints affecting partial prey consumption by a crab spider,Diaea sp. indet. (Araneae: Thomisidae)

Summary The influence of feeding constraints on the feeding behaviour of Diaea sp. indet., was investigated. Diaea is a crab spider which ambushes its prey and practises extraintestinal digestion. A laboratory study was carried out using fruit flies, Drosophila immigrans, as prey. Diaea feeds from two sites on the prey — initially the head, followed by the posterior abdomen, with most of the prey's contents being extracted from the head. If additional prey are available, Diaea will, instead of switching to the posterior abdomen, catch a new prey item. The efficiency with which Diaea can extract food is influenced by changes which occur in the prey as a consequence of it being killed and fed on. Evaporative fluid loss from prey is an important constraint on food uptake because in influences the viscosity of the prey's contents. Regardless of whether a new prey item arrives, Diaea discards the prey item on which it is feeding before all of the available food has been extracted from it. The fluid content of the prey is not only part of the food the spider extracts, it is also a resource enabling efficient transfer of food from prey to predator. The value of the prey's fluid content as a resource decreases as a function of feeding time and as a consequence of the spider feeding on the prey.     

Diets and food consumption rates of pelagic fish in Lake Malawi, Africa

• 1 Studies of diel feeding periodicity, rates of food consumption and diet composition of pelagic fish were undertaken to resolve the dispute regarding the existence of a vacant niche for a pelagic zooplanktivore in Lake Malawi.
• 2 Six species of zooplanktivore were abundant in the offshore pelagic zone, these were: Diplotaxodon limnothrissa, D.‘bigeye’, Rhamphochromis longiceps and Copadichromis quadrimaculatus (all haplochromine cichlids), Synodontis njassae (Mochokidae) and Engraulicypris sardella (Cyprinidae). The main piscivores were cichlids of the genus Rhamphochromis.
• 3 All cichlids were daytime feeders; some showed peaks in feeding activity at dawn and dusk that were related to vertical migration patterns of their prey. Synodontis njassae was able to feed throughout the day, but fed most actively at night.
• 4 Food consumption: biomass ratios (Q/B yr^-1) calculated from diel stomach contents data were 5.87–7.13 for D. limnothrissa, 12.79 for D.‘bigeye’, 4.20–24.7 for R. longiceps and 6.45 for S. njassae. Annual Q/B ratios for other species, which ranged from 4.74 to 9.28, were obtained from an empirical model relating food consumption to fish morphology. Published estimates were used for E. sardella and Opsaridium microcephalum (Cyprinidae). An estimate of total prey consumption by the pelagic fish population was obtained from Q/B ratios and fish biomass estimates from acoustic surveys covering the entire offshore pelagic zone of the lake (24 000 km²).
• 5 Diplotaxodon limnothrissa, R. longiceps and C. quadrimaculatus had a broad range of diets, with the ability to switch from crustacean zooplankton to larvae and pupae of Chaoborus edulis (Diptera) or larval and juvenile E. sardella. Diplotaxodon‘bigeye’ and S. njassae fed almost exclusively on fourth-instar larvae and pupae of C. edulis. Engraulicypris sardella fed mainly on crustacean zooplankton. The main prey of the large piscivorous Rhamphochromis species were Diplotaxodon spp. and E. sardella.
• 6 The fish community consumed only 3% of estimated crustacean zooplankton production directly. Predation pressure was intense (> 80% of estimated production consumed by predators) on late instars of the zooplanktivorous C. edulis larvae and on E. sardella larvae. Rhamphochomis spp. consumed 22–43% of estimated production of the zooplankton-eating fishes.
• 7 Although many members of the fish community do not feed directly on crustacean zooplankton, so that the food-chain supporting fish production involves an extra trophic level, it is considered that the endemic fish community is well adapted to feeding on the low-density prey in the pelagic zone, and that there is no vacant feeding niche.

     

Bioenergetic model of planktivorous fish feeding, growth and metabolism: theoretical optimum swimming speed of fish larvae

The feeding activity of an individual fish larva is described by an equation which includes parameters for the area successfully searched, probability of food capture multiplied by the cross-sectional perceptive visual field, larval swimming speed and the time required to consume a unit of food energy. The proportion of ingested food energy used for metabolism increases exponentially with increasing swimming speed. The model predicts that food consumption rate increases asymptotically whereas metabolic rate increases exponentially. This results in a predicted growth rate curve that reaches a maximum at a certain swimming speed and decreases at both higher and lower speeds. The model can be used to predict the influence of type of prey, prey density, water temperature etc. on larval growth. An expression describing how many hours per day fish larvae must forage in order to grow at a certain daily body weight gain allows the limits of environmental conditions for positive, zero and negative growth rate to be set. Results of simulations demonstrated that the optimum swimming speed for maximum growth of coregonid larvae increased with an increase in food density, decrease in water temperature or decrease of prey vulnerability. At optimum ‘theoretical’ swimming speed an increase in water temperature from 5 to 17° C required the food density to be increased from 20 to 80 copepods l^?1 in order to maintain a daily growth increment of 2%. The minimum Artemia density required for maintenance metabolism increased from 10 to 30 items 1¹ over the same temperature increase from 5 to 17° C, and food densities required for 8% growth rates were 26 and 56 Artemia nauplii l^?1 at 5 and 17° C, respectively. Contrary to previous findings, results of the present study suggest that metabolic rates of actively feeding fish larvae may be from 5 to 50 times the standard metabolic rate: earlier studies suggested that a factor of 2–3 may be generally applicable.