Fish eggs and larvae in the diet of herring Clupea harengus membras Linnaeus, 1758 and the sprat Sprattus sprattus balticus (Schneider, 1904) (Clupeidae) in the Southeastern Baltic Sea

The qualitative and quantitative composition of fish eggs and larvae in the stomach contents of Baltic herring and sprat was examined. The occurrence of fish eggs and larvae in the diet of herring was markedly higher, compared to sprat. Eggs of four and larvae of nine fish species were found in the herring stomachs. The prey species composition of the herring and sprat diets was related to the ichthyoplankton composition and to the degree of overlapping of the prey and predator distributions. Sprat eggs were the predominant prey in the herring diet in May and sprat larvae prevailed in August. The daily ration estimates indicate that the sprat larvae consumption by herring in August exceeded the sprat egg consumption in May by many times. Cod eggs in the herring stomachs were only found at the optimal oxygen content and salinity for the cod eggs in the near-bottom water layer.     

Antipredator responses to alarm pheromone in groups of young and/or old thrips larvae

Many prey species suffer from different predators in the course of their ontogeny. Hence, the alarm signal a small prey individual sends can have a different meaning than the signal a large prey individual sends, both for small and for large receivers. Larvae of Western Flower Thrips face predators that attack only small larvae, or predators that attack small larvae and large larvae. Furthermore, thrips larvae release a two‐component alarm pheromone, which varies in composition with larval age. Here, we study whether their response to alarm pheromone varies with composition of the pheromone. First, we confirmed that large and small larvae respond when nearby larvae of both sizes were prodded with a brush to induce alarm pheromone excretion. Subsequently, we tested whether thrips larvae of a given size respond differentially to alarm pheromone excreted by a small or large companion larva. We analyzed two types of behavior used in direct defense against a predator and one type of escape response. Only small (not large) larvae attempted to escape more frequently in response to excretions from a large larva. This difference in response could have been due to the alarm pheromone or to the companion larva in the vicinity. We subsequently tested for, but did not find, an effect of size of the companion larva on the behavior of the test larva when exposed to synthetic pheromone mimicking that of a large larva. Finally, we tested how pheromone composition affects antipredator behavior by exposing thrips larvae to synthetic pheromones differing in amount and ratio of the two components. Only for small larvae, we found significant changes in escape behavior with pheromone amount, and a trend with the ratio. Overall, we conclude that small thrips larvae respond differentially to alarm pheromones excreted by small and large larvae and that this differential response is due to differences in pheromone quantity and possibly also quality. Our results suggest that responses to alarm signals can vary with the chemical composition of those alarm signals.     

Susceptibility of Atlantic herring and plaice larvae to predation by juvenile cod and herring at two constant temperatures

Predation encounters were staged in the laboratory to compare prey responsiveness, predator error rate, and predator capture success for juvenile cod Gadus morhua (a suction feeder) and herring Clupea harengus (a biting predator) preying on herring and plaice Pleuronectes platessa larvae. Trials were conducted at near natural temperature extremes for the larvae (8 and 13°C) to assess the importance of water temperature to the interaction. Herring larvae were significantly more responsive to attacks by herring than were plaice larvae (5·7 vs 0'0%). The two prey species were equally responsive to attacks by cod (2–6 vs 10%). Cod caught 91% of herring larvae attacked and juvenile herring caught 87%. Cod were successful in 96% of attacks on plaice, but juvenile herring caught significantly fewer (83%) plaice larvae. For each predator species, capture success did not vary significantly with prey species. Overall capture success for herring was significantly lower than that for cod. Responsiveness of herring larvae to attacks by juvenile herring increased with temperature, but predator error rate and capture success were not altered by water temperature.     

Diet changes in prey size selectivity in larvae of perch <Emphasis Type="Italic">Perca fluviatilis</Emphasis>

A hypothesis on change in prey size selectivity in relation to illumination level was tested on the basis of data on weight and size composition of the content of the digestive tract of larvae of perch Perca fluviatilis and on zooplankton in the layer 0–6 m (Wallersee Lake, Austria). Larvae foraging in the twilight-night period had almost two times more food in the intestine than those foraging in the daytime. The size composition of perch larvae and concentration and size composition of zooplankton hardly differed in the daytime and twilight-night samples. For the first time, it is shown on field material that more intensive feeding of larvae at twilight is related to selection of significantly larger prey than in daytime feeding. In the day-time, the larvae consumed more prey but their maximum size did not exceed 0.6 of the diameter of mouth opening of the fish; at twilight it was over 0.8. In case of feeding on so large prey, not only the weight of the consumed feed increases but the time used for capture and swallowing also considerably increases. The larvae which in the period of investigation did not yet form schools, which perform the principal defensive function, were especially vulnerable for predators feeding on relatively large prey. The decrease of the part of small-sized prey at twilight is not related to their lesser availability due to a low illumination. It is assumed that feeding on energetically more valuable but less available prey is shifted to the period of low illumination when the larvae are less exposed to predation risk. The obtained results are discussed from positions of the triotroph concept (Manteifel’, 1961).     

Prey selection by the dobsonfly larva, Protohermes grandis (Megaloptera: Corydalidae)

SUMMARY 1. Prey selection by the dobsonfly larva, Protohermes grandis (Thunberg), was studied in stony riffles of the Yataro River, central Japan. The density, size distribution and taxonomic composition of available prey were assessed for 2 years. In order to know the encounter rate between prey and this ambush predator, prey mobility was also estimated from patterns of colonization of experimentally detiuded stones.
2. Foregut analyses revealed that maximum size of prey eaten increased with larval size, and large larvae did not take the smallest prey in spite of high availability in all seasons.
3. Charnov's (1976) optimal diet model quantitatively predicted such size-selective feeding under seasonally fluctuating conditions of water temperature and prey availability. Larvae maximized the feeding rate by selecting prey.
4. Maximum width of prey eaten coincided approximately with larval mandible length. Mandible size seemed to play an important role in the selection of prey in the optimal size range.     

Potential predation pressure of littoral mysids on herring (Clupea harengus membras L.) eggs and yolk-sac larvae

The predation potential of littoral mysid shrimps (Mysidacea) on Baltic herring (Clupea harengus membras L.) eggs and yolk-sac larvae was studied experimentally. The results showed that littoral mysids feed actively on both eggs and yolk-sac larvae. It was shown that Neomysis integer preys on eggs, which are not attached to the substrate. Alternative food (yolk-sac larvae or zooplankton) did not decrease feeding rate on eggs. Only gravel as a bottom material lowered the ingestion rate to nearly zero. The largest of the mysid species Praunus flexuosus ate yolk-sac larvae more than other mysids and most efficiently. Mysids switched to feed on eggs when larvae and eggs were offered simultaneously, thus predation focused on eggs. It is possible that hydrodynamic signals of moving larvae induced mysids to prey and eggs were easier prey to catch as well as more numerous. In addition egg size is optimal and the nutritive value (measured as C:N ratio) is better compared with larvae. The results indicate that mysids may have local effects on populations of Baltic herring by eating the early life stages, mostly eggs. Especially when large swarms of N. integer shoal in the spawning areas. However, the effect on recruitment of herring is still hard to evaluate.     

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.     

DIET OF HARBOR PORPOISES IN THE KATTEGAT AND SKAGERRAK SEAS: ACCOUNTING FOR INDIVIDUAL VARIATION AND SAMPLE SIZE

Stomach contents of 112 bycaught harbor porpoises ( Phocoena phocoena ) collected between 1989 and 1996 in the Kattegat and Skagerrak seas were analyzed to describe diet composition and estimate prey size, to examine sample size requirements, and to compare juvenile and adult diets. Although porpoises preyed on a variety of species, only a few contributed substantially to the diet. Atlantic herring ( Clupea harengus ) was the dominating prey species for both juveniles and adults. Our results, in combination with those from previous studies, suggest that where herring is a dominant food source, porpoises prey primarily on size classes containing mature or maturing individuals. Further, we also show that Atlantic hagfish ( Myxine glutinosa ) may be an important food resource, at least for adult porpoises. Examination of sample size requirement showed that, depending on the taxonomic level used to describe the diet, a minimum of 35–71 stomachs are needed to be confident that all common prey species will be found.     

The effects of thin layers on the vertical distribution of larval Pacific herring, Clupea pallasi

Temporal and spatial heterogeneity of resources is likely to have dramatic effects on the behaviors of zooplankton and ichthyoplankton, which in turn are likely to have strong effects on ecological dynamics such as predation, growth, and mating. The objective of this study was to determine whether vertically thin layers of extreme prey concentration affect the vertical distribution of larval Pacific herring (Clupea pallasi). We employed 2-m tall experimental tanks equipped with video cameras that scanned the vertical extent of the tanks to investigate the effects of thin layers on the vertical distribution of 5- and 10-day-old herring larvae. Three treatments were established: (1) a thin layer of prey (rotifers, Brachionus plicatilis) through density (salinity) stratification, (2) homogeneous vertical distribution of both prey and density, and (3) density (salinity) stratification, but with a homogeneous distribution of prey. We found that in all treatments the majority of larval herring were at the surface, near the light, despite the absence of a peak in rotifer abundance at this depth in some instances. However, there were also clear effects of the thin layers—secondary subsurface peaks in herring abundance occurred at the mid-depths in the stratified tanks, in and around the thin layers. In addition, our results provide some evidence that thin layers specifically, rather than prey patches generally, influence the vertical distribution of larval herring, i.e., larvae may use the physical properties of thin layers to locate and distribute themselves, instead of reacting solely to the prey patches. Thus thin layers can affect directly the vertical distribution of larval herring, and perhaps indirectly their horizontal distribution, as herring larvae live in environments (e.g., estuaries) where advective transport is also often vertically heterogeneous.     

Temperature-driven changes in early life-history stages influence the Gulf of Riga spring spawning herring (<Emphasis Type="Italic">Clupea harengus m</Emphasis>.) recruitment abundance

Processes occurring during early life-history stages influence the year-class abundance of marine fish. We found that the abundance of 1-year-old spring spawning herring is statistically significantly determined by the number of post-flexion herring larvae in the Gulf of Riga (Baltic Sea). The abundance of consecutive developmental stages of larvae: yolk-sac, pre-flexion, flexion and post-flexion strongly correlated with each other, indicating that factors which already influence the yolk-sac stage are important in determining the abundance of post-flexion herring larvae. Winter air temperature before spawning determined the timing of maximum abundance of pre-flexion herring larvae, but not their main prey: copepod nauplii, implying that different mechanisms governing major preconditions for the formation of year-class strength. The abundance of post-flexion larvae displayed a potential dome-shaped relationship with sea surface temperature experienced after hatching. We suggest that increased summer temperatures, which exceed the physiological optimum negatively, affect the survival of post-flexion herring larvae. Overall, future climate warming poses an additional risk to larval herring survival and this may lead to a reduction in those herring stock which rely on recruitment from shallow coastal areas.     

Modelling the attack success of planktonic predators: patterns and mechanisms of prey size selectivity

A mathematical model of the attack success of planktonic predators(fish larvae and carnivorous copepods) is proposed. Based ona geometric representation of attack events, the model considershow the escape reaction characteristics (speed and direction)of copepod prey affect their probability of being captured.By combining the attack success model with previously publishedhydrodynamic models of predator and prey perception, we examinehow predator foraging behaviour and prey perceptive abilityaffect the size spectra of encountered and captured copepodprey. We examine food size spectra of (i) a rheotactic cruisingpredator, (ii) a suspension-feeding hovering copepod and (iii)a larval fish. For rheotactic predators such as carnivorouscopepods, a central assumption of the model is that attack istriggered by prey escape reaction, which in turn depends onthe deformation rate of the fluid created by the predator. Themodel demonstrates that within a species of copepod prey, theability of larger stages to react at a greater distance fromthe predator results in increased strike distance and, hence,lower capture probability. For hovering copepods, the vorticityfield associated with the feeding current also acts in modifyingthe prey escape direction. The model demonstrates that the reorientationof the prey escape path towards the centre of the feeding current'sflow field results in increased attack success of the predator.Finally, the model examines how variability in the kineticsof approach affects the strike distance of larval fish. In caseswhere observational data are available, model predictions closelyfit observations.     

Diet breadth variability in larval blue whiting as a response to plankton size structure

Diet breadth (measured as the S.D. of the log of prey size per larvae; SLH) of blue whiting micromesistius poutassou larvae followed a quadratic equation with larval size. In small larvae, diet breadth in terms of size (SLH), the mean and the maximum of the log of prey size per larvae (MLH and XLH, respectively) increased with larval size as prey size selection shifted to larger prey. In contrast, large larvae tended to reduce diet breadth of prey sizes ingested, focusing on the larger prey that were abundant, instead of raising the upper limit of prey sizes because of the low abundance of larger prey. Except for larvae at the onset of first feeding, number of prey stayed constant or decreased in relation to larval size. Both patterns (in small and large larvae) maintained a constant rate of increase of gut carbon content with increase in larval size. Large larvae appear to maintain the increase in gut carbon content during ontogenetic development by reducing diet breadth (SLH) and increasing selection towards the larger prey that are abundant.     

Effects of alternative prey on predation intensity from herring Clupea harengus and sandeel Ammodytes marinus on capelin Mallotus villosus larvae in the Barents Sea

Stomach contents from lesser sandeel Ammodytes marinus and herring Clupea harengus caught at one and three stations, respectively, were analysed to investigate predation intensity on capelin Mallotus villosus larvae. Most capelin larvae were found in the anterior sections of the stomachs close to the oesophagus, indicating that rapid digestion rates of larvae reduced the abundance in the posterior sections. The anterior sections of the stomachs had alternating layers of either copepods or capelin larvae and krill. This shows that the individual predators switched between feeding on either copepods or on krill and capelin larvae. A total of 549 capelin larvae was found in 440 fish stomachs. Capelin larvae were found in 20% of the sandeel stomachs, and 24, 34 and 62% of the stomachs from the three herring stations, respectively. Many of the predator stomachs contained more than five larvae, and up to 28 larvae were found in one herring stomach. The herring appeared to prey selectively on the largest capelin larvae, and the number of larvae per stomach was much higher in stomachs where krill had recently been eaten than where copepods were dominant. The predation intensity measured in this study is much higher than reported in earlier studies, and it is suggested that the predators were feeding using a searching image, and that the concentrations of alternative prey (copepods and krill) varied around a switching threshold.     

Effect of larval ontogeny, turbulence and light on prey attack rate and swimming activity in herring larvae

The effects of ontogeny (larval size), light and turbulence on the attack rate and swimming activity (proportion of time swimming and duration of swimming bout) of herring larvae (15-28 mm TL) have been investigated. Emphasis was put on the experimental design in order to create a set-up where the turbulence intensity distribution could be accurately measured as well as controlled in the entire experimental tank.Both larval size (ontogeny) and light had a significant positive effect on prey attack rate. Likewise, an intermediate increase in turbulence had a positive effect on prey attack rate, but this effect was dependent of light intensity and larval size.At low light (1.5 μE m² s⁻¹) intermediate turbulence increased the prey attack rate significantly for larger larvae (26 and 28 mm), while at high light (18 μE m² s⁻¹) intermediate turbulence had only a significant positive effect on the attack rate of smaller larvae 20 and 23 mm.In general, our data show a dome-shaped response of turbulence on attack rate and a U-shaped response of turbulence on swimming activity.For herring larvae >20 mm, the maximum (attack rate) and minimum (swimming activity) response of turbulence were found at intermediate turbulence intensities (energy dissipation rates between 7∗10⁻⁸ and 1∗10⁻⁶ W/kg). The highest turbulence level tested (8∗10⁻⁶ W/kg) showed only negative effects, as attack rates where at the lowest and swimming activity at the highest.Swimming activity increased with larval size or light, and decreased at intermediate turbulence. Compared to turbulent intensities under natural conditions this implies that larger herring larvae at 10 m depth have to be exposed to wind speeds of more than 17 m/s before negative effects on attack rate and swimming activity occurs.     

Predation by sprat and herring on pelagic fish eggs in a plaice spawning area in the Irish Sea

Predation on planktonic fish eggs was examined by stomach content analysis of sprat and herring sampled in a plaice spawning area to the east of the Isle of Man in March 1993. Plankton samples were taken to examine prey selection. The clupeids showed a selection for the later developmental stages of plaice eggs. Plaice eggs had a refuge in size from predation by sprat <80 mm total length which selected smaller non-plaice eggs. However, herring and sprat >80 mm total length showed a strong selection for plaice eggs, presumably due to their large size.     

Effects of prey size structure and turbulence on feeding and growth of anchovy larvae

Foraging processes in plankton and planktivorous fish are constrained by relative prey and predator size and therefore, these are important variables to include in a foraging model. The distribution of prey biomass across different size classes can be characterized by a size spectrum slope. We present a foraging model for anchovy larvae including the most relevant processes such as prey encounter, capture- and pursuit success, all influenced by light, turbulence and prey characteristics. We modelled ingestion rates and specific growth rate by coupling the foraging model with an existing bioenergetic model, and performed a sensitivity analysis of prey ingestion in turbulent environments assuming either hemispherical or conical perceptive volume. Our results suggest that turbulence has no positive effect because of the low capture ability, small prey size and small visual volume for anchovy larvae. The predicted ingestion is too low to sustain the growth potential of larvae when assuming conical perceptive volume even under prey densities substantially higher than normally found in the field. Ingestion rate is sensitive to the total biomass and the slope of the prey size spectra, specifically because it determines the abundance of prey around the optimal size for the larvae. The model also suggests that small larvae benefit from a prey size structure with steep prey size-spectra slope while a large larva benefit from less steep slopes. The model can act as a link between size-spectra measurements from the field and the foraging conditions of larval anchovies.     

The importance of small-scale turbulence in the feeding of herring larvae

Herring larvae form a large, persistent and well-defined aggregateoff southwest Nova Scotia coincident with a region of strongtidally generated turbulence. Our objective is to test the hypothesisthat turbulent, tidally well-mixed regions constitute a preferentialfeeding environment for herring larvae, and are thus the proximatecause for the apparent retention of larvae in these areas. Dimensionalanalysis, coupled with relevant biological parameters and fielddata, are used to derive two non-dimensional numbers from existingpredator-prey encounter models, and to propose a third numberwhich incorporates the minimum encounter rate required for larvalmaintenance. The results of this analysis show that when root-mean-squareturbulent velocities are of the same order of magnitude as theprey velocities, they start to influence encounter rates significantly;and that when turbulent velocities are an order of magnitudegreater than the predator velocities, they dominate the encounterprocess. At reported natural food concentrations, food itemsare on average one order of magnitude away from the larvae'sreach. The importance of turbulence as a mechanism to bringfish larvae closer to their prey is revealed by the findingthat in tidally well-mixed regions herring larvae would requireone order of magnitude less food than their counterparts inthe stratified regions. The model shows that the feeding environmenton the adjacent stratified waters off southwest Nova Scotiais detrimental for herring larvae, and that high mortality wouldbe expected if larvae had to overwinter in this region. Ouranalysis also shows that the growth pattern observed for autumn-spawnedlarvae is explained by the combination of the relevant physicaland biological scales. The overall results of this analysislend support to the hypothesis that turbulent, tidally well-mixedareas provide ideal feeding conditions for herring and may bethe proximate cause for an apparent retention.     

Facilitation of fisheries by natural predators depends on life history of shared prey

Predators commonly share prey with human exploiters, intuitively suggesting that there is an inherent human–predator conflict through competition for prey. Here we studied the effects of fishing and predation mortality on biomass distributions and yields of shared prey using a size‐structured model of competing populations, describing the life histories of Baltic Sea sprat and herring. Whereas both species responded in a similar fashion to increased fishing mortality, with decreasing juvenile and adult biomasses, we found that responses to predation mortality differed between species. Sprat only display weak compensatory responses with increasing predation mortality, while over a substantial range of mortalities there was a strong increase in adult (and total) herring biomass, i.e. overcompensation. The observed biomass overcompensation results from relaxed intraspecific competition as predation mortality increased, allowing for faster individual growth rates that in turn lead to a change in population composition (juvenile:adult biomass ratio). Our results suggest that the potential for biomass overcompensation is higher for species exhibiting substantial growth after maturation. Differences in size‐selectivity of predators and fishing mortality resulted in a positive effect of predation mortality on fisheries yields, which can be explained by an overcompensatory response in adult herring biomass. Thus, somewhat counter intuitive, our results suggest that fishermen, depending on prey life history, may actually benefit from allowing for a higher abundance of predators, despite competing for shared prey.     

Lake Whitefish and Lake Herring Population Structure and Niche in Ten South-Central Ontario Lakes

Synopsis This study compares simple fish communities of ten oligotrophic lakes in south-central Ontario. Species densities and population size structure vary significantly among these lake communities depending on fish species present beyond the littoral zone. Lake whitefish are fewer and larger in the presence of lake herring than in their absence. Diet analysis indicates that lake whitefish shift from feeding on both plankton and benthic prey when lake herring are absent to a primarily benthic feeding niche in the presence of lake herring. When benthic round whitefish are present, lake whitefish size and density decline and they move lower in the lake compared to round whitefish. Burbot are also fewer and larger in lakes with lake herring than in lakes without herring. Burbot, in turn, appear to influence the population structure of benthic coregonine species. Lower densities of benthic lake whitefish and round whitefish are found in lakes containing large benthic burbot than in lakes with either small burbot or where burbot are absent. Predation on the pelagic larvae of burbot and lake whitefish by planktivorous lake herring alters the size and age structure of these populations. As life history theory predicts, those species with poor larval survival appear to adopt a bet-hedging life history strategy of long-lived individuals as a reproductive reserve.     

Fatty acid content of eggs determines antipredator performance of fish larvae

Recent work has suggested that provisioning of eggs with certain critical nutrients could be a more meaningful measure of maternal investment and correlate of offspring fitness than traditional measures of egg size. The aim of our study was to assess variability in egg quality and larva quality and to identify connections between them and the implications for larval survival. Egg size, proximate composition, and fatty acid composition were measured for 40 batches of eggs from 8 captive pairs of red drum (Osteichthyes: Sciaenops ocellatus). We reared larvae from these batches of eggs to a common size (10 mm total length, 2-3 weeks posthatching) and assessed routine activity and escape response performance of 671 individuals. Egg fatty acid composition varied more than egg size or proximate composition. Concentrations of certain long chain, highly unsaturated essential fatty acids (e.g., arachidonic acid and docosahexaenoic acid) were the only egg traits that were significantly related to larva quality (measured as escape performance). Reduced escape performance of larvae from eggs with low fatty acid concentrations was not compensated by 3 weeks of feeding on a diet enriched with fatty acids, suggesting irreversible developmental effects. Since fatty acids in eggs originate from the maternal diet, offspring survival may be determined in part by availability of nutrient-rich prey to pre-spawning adults. Migrations, regime shifts, and exploitation of marine communities could operate through this mechanism to influence recruitment in fish populations. Our findings underscore the importance of non-genetic maternal contributions to egg quality and the linkage between environmental conditions experienced by adult females and offspring fitness.