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.     

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.     

Vulnerability of late larval and early juvenile Atlantic herring, Clupea harengus, to predation by whiting,Merlangius merlangus

Laboratory experiments investigated changes in the vulnerability of herring to predation by whiting during growth from <34 to >80mm in length, Premetamorphic herring (<50mm) failed to react to c . 50% of attacks by the predator, but this was reduced to c . 20% in postmetamorphic fish. Premetamorphic herring failing to react had c. 30% probability of survival due to unforced errors by the predator (unsuccessful attacks that did not elicit a reaction by the prey), but these errors did not occur after herring metamorphosis. The loss of the advantage of lower conspicuousness following metamorphosis was counterbalanced by increased reactivity. Antipredator benefits should increase with schooling behaviour (first evident in experiments at c . 50 mm), so predation mortality should decrease in postmetamorphic herring in the sea. Metamorphosis itself would be associated with high predation risk since conspicuousness is enhanced but reactivity and schooling behaviour are still not fully developed.     

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.     

Spatially structured interactions between a migratory pelagic predator, the Norwegian spring-spawning herring Clupea harengus L., and its zooplankton prey

Spring-spawning herring Clupea harengus was patchily distributed over large parts of the Norwegian Sea in May 1995–2005, during the early phase of the annual feeding migration. Overall, herring tended to be found in areas with intermediate biomasses of zooplankton prey, intermediate water temperatures and relatively high salinities. Herring had more food in their stomachs in areas of relatively low water temperature and high herring abundance. Hydrographical conditions revealed that herring was feeding mainly within Atlantic water masses, and more intensely in western and northern regions of the Norwegian Sea. Zooplankton biomass was patchily distributed, and was generally higher towards the western parts of the Norwegian Sea. Here, zooplankton biomass in year _{i +1} was also negatively associated with herring spawning stock biomass in year _i , while there was no evidence for such an association in the eastern region; indicating that herring may have a geographically structured 'top–down' effect on the recruitment of its zooplankton prey. The fact that herring was not typically associated with the areas containing the greatest zooplankton biomasses may reflect that the fish had not yet reached the most profitable feeding grounds or alternatively that herring was depleting zooplankton biomass.     

Growth, ingestion rates and metabolic activity of walleye in lakes with and without lake herring

Growth efficiencies, ingestions rates and activity levels of walleye Sander vitreus were compared in lakes with and without lake herring Coregonus artedi . Yellow perch Perca flavescens were the main prey in lakes without lake herring. Walleye were sampled in September and October from 38 lakes in Ontario in 1998 and 1999, using multimesh monofilament gillnets. Ingestion rates were estimated from annual increments in somatic mercury and body mass, and the mercury content of yellow perch and lake herring. Walleye had higher growth efficiencies, and lower ingestion and activity rates in lakes with lake herring. Lake herring grow larger than yellow perch and therefore could provide more profitable prey for larger walleye. The results are consistent with optimal foraging theory that predicts that walleye feeding on optimal prey sizes should grow more efficiently, if the ratio of feeding benefit (energy) to cost (search and seizure) is a function of the ratio of predator and prey size.     

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.     

The relative profitability of particulate- and filter-feeding in the herring, Clupea harengus L.

Analysis of laboratory video-recordings of herring feeding by biting and filtering on Calanus finmarchicus and three sizes of Artemia enabled the capture rates of the two methods to be estimated at different prey concentrations. At low concentrations the fish feed by selective capture of individual particles, but the capture rate achievable by this method is constrained by the maximum rate at which they can bite. Filter-feeding is not subject to this constraint because capture rate is directly proportional to prey concentration and above a critical prey concentration its capture rate exceeds that of biting. The possession of two feeding methods allows the fish to maximize prey intake over a wide range of concentrations and the phenomenon of switching between feeding methods can be explained by their relative profitabilities at different concentrations. The observation that less than 50% of fish are filtering when capture rates by the two methods are equal suggests that filtering is energetically more costly than biting. Estimates of the energy cost of filtering indicated that it may be from 1.4 to 4.6 times higher than that of biting.     

Metapopulations and the Atlantic herring

Two opposing concepts of Atlantic herring, Clupea harengus L., population structure are critically reviewed with the objective of unifying these divergent views under the metapopulation concept. It is concluded that neither the discrete population concept nor the dynamic balance concept adequately explains all the data associated with herring population structure and dynamics, including meristic and morphometric measurements, life- history traits, homing, year-class twinning, and biochemical analyses. However, the available information does suggest that Atlantic herring population structure and dynamics are well described within the metapopulation concept. The example of sympatric seasonal-spawning populations is used to illustrate the strategy, opportunity and mechanism by which local population integrity and persistence are maintained within the adopted- migrant hypothesis. Local population integrity is maintained through behavioural isolation, i.e. repeat rather than natal homing to spawning areas, while local population persistence is ensured through the social transmission of migration patterns and spawning areas from adults to recruiting individuals     

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.     

Vertebrate exploitation of pulsed marine prey: a review and the example of spawning herring

Short-term bursts of prey availability occur in many ecosystems and have potential important consequences for both predator biology and ecosystem function. Examples of prey ‘pulses’ in marine ecosystems include spawning runs of several anadromous and marine fishes, horseshoe crab spawning, and salmonid juvenile outmigrations, which are exploited by numerous species of vertebrate predators. In a few cases, the fitness or demographic consequences of such predator–prey interactions are known or inferred, but too often that information remains unknown. We explored the extent of temporal and spatial variation in one example of a pulsed marine resource: the spawning of Pacific herring (Clupea pallasii). Spawning herring provide a rich, aggregated resource to which dozens of species of vertebrate predators often exhibit strong numerical responses. However, the spawning events are often variable in both time (annual differences of several to many weeks) and space (both regional and more local differences in size and timing of events). Such variability must affect more mobile predators less than area-restricted predators, and thus its effect would vary not only among species but also within species, depending on constraints of the predator life history. Unpredictability of the prey concentrations, whatever their proximate causes, may contribute to maintenance of metapopulations of prey such as herring, if unpredictability lessens the impact of predation.     

Development, morphometry and particle retention capability of the gill rakers in the herring, Clupea harengus L.

The development of gill rakers in the herring, Clupea harengus L. was followed from the larva to the adult. The first rakers appear on the gill arches at a total length ( t.l .) of about 16 mm. Their number then increases rapidly until the fish are about 50 mm t.l . when the rate of addition becomes much slower. The length of individual rakers and the space between them continues to increase throughout life. The rakers on the first gill arch account for almost 60% of the whole filtering area. The observed particle retention capabilities of the fish when filter-feeding were lower than those expected on the basis of the estimated spaces between the rakers. Several reasons are suggested for the discrepancy.     

Resource sharing among cod, haddock, saithe and pollack on a herring spawning ground

Cod Gadus morhua , haddock Melanogrammus aeglefinus , pollack Pollachius pollachius and saithe Pollachius virens were abundant fishes on herring Clupea harengus spawning grounds o. southwestern Norway. Their diets were species specific and ontogenetic changes were similar to patterns observed in other areas, indicating both inter- and intraspecific resource sharing. However, strong interspecific diet overlap occurred when cod and pollack preyed upon sandeel Ammodytes marinus , and when all the gadoids fed on herring eggs during the herring spawning season in the spring. These prey items were very abundant and the extensive diet overlap may not reflect strong competitive interaction.     

The role of escape reactions in determining the size distribution of prey captured by herring larvae

Synopsis Fish larvae are selective planktivores, and size is an important factor in prey selection. However, for herring larvae, a selection model based solely on the principle of optimising calorific gain per unit energy expenditure consistently overestimates the mean size of ingested prey. Most such models ignore the escape capabilities and shape of the prey, and the potential role of escape on capture success. In this paper, a static foraging model incorporating both selection and escape is described and tested against published data on the composition of herring larvae stomach contents. The results indicate that prey escape is a major factor structuring the diet composition of the larvae.     

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.     

The spatial patchiness of Pacific herring larvae

Synopsis The spatial patchiness at age was measured for Pacific herring,Clupea harengus pallasi, larvae of a small coastal inlet. Lloyd's index of patchiness decreased from 3.8 at hatching to a minimum of 1.5 at 25–30 d and then increased to 2.5 at 50 d. This two-cycle pattern resembles that measured by others for the larvae of other species of schooling pelagic fishes. It differs by being several times lower in magnitude. This is attributed to two factors: (1) a rapid reduction in patchiness from a relatively high level in the demersal egg interval to a lower level in the planktonic larval interval as larvae are released into the water column over a period of 2–4 days, and (2) foraging by larval herring for prey that are larger, more dilute, and less patchy than the prey of other, smaller pelagic fish larvae.     

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.     

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.     

Effects of temperature, prey type and prey size on gastric evacuation in small cod and whiting

Gastric lavage was used to investigate the effects of temperature, prey type and prey size on gastric evacuation in small cod Gadus morhua and whiting Merlangius merlangus . The fish were fed to satiation and subsequently the stomach contents were sampled to determine the rate at which food was evacuated. Satiation meal size was positively related to temperature and differed between prey types. The gastric evacuation rate (GER) also tended to increase with temperature and varied with prey type. GER at temperatures of 11.3–12.7° C averaged 1.5–1.7 times higher than at 6–9.5° C. There was no significant difference (P>0.05) in the evacuation of lugworm Arenicola marina , sandeel Ammodytes spp., and herring Clupea harengus , but the GER of brown shrimp Crangon vulgaris was much slower (P<0.05). No significant changes in GER were observed when fish were fed on three different size groups of either herring or brown shrimp. In whiting, there was no significant difference in the GER of individual herring or brown shrimp when they were fed as single species meals or incorporated in meals containing a mixture of prey species.     

Feeding ecology of mackerel and dietary overlap with herring in Icelandic waters

The Northeast Atlantic mackerel (Scomber scombrus) has been extending its summer feeding distribution north and west, including around Iceland, since around 2006. The objective of this study is to quantify the weight gain and total food consumption of mackerel and to evaluate the food competition between mackerel and herring (Clupea harengus) feeding in the marine ecosystem around Iceland during the summers 2009–2011. Mackerel feeding in Icelandic waters gained around 43% on average in weight during these summers. Based on swept-area abundance estimates of mackerel from an international survey in 2011 and available estimates of food conversion efficiency in mackerel, the weight gain in Icelandic waters in 2011 corresponded to a total consumption of around 3.4 (2.4?4.5) million tonnes. Overall, 98% of 2314 mackerel, 91% of 398 Icelandic summer-spawning herring and 96% of 424 Norwegian spring-spawning herring stomachs, collected over the summers 2009?2011, contained food. The mean weight of the stomach content of mackerel was higher than for herring in all the years. While the stomach content weight of mackerel was generally highest in southeastern and southwestern areas, it was highest for herring in western, eastern and northern areas. Analysis of stomach contents showed that Copepoda were the most important food of mackerel in most areas, while Copepoda and Euphausiacea were the most important food items for herring. Fish prey contributed a higher proportion in stomachs of mackerel than herring, although relatively low for both species.