The predation impact of juvenile herring Clupea harengusand sprat Sprattus sprattus on estuarine zooplankton

The consumption of estuarine copepods by juvenile herring and sprat during estuarine residency was estimated using fish biomass data and daily rations calculated from two models of feeding in fish: a bioenergetic model and a gastric evacuation model. The bioenergetic model predicted daily rations that were, on average, three times higher than those estimated by a model based on field records of stomach contents. The biomass of herring and sprat in the estuary was negatively correlated with the daily ration suggesting that the clupeid fish populations were resource-limited. Copepod production decreased towards the winter and peaked in spring and summer. The relative importance of predation changed seasonally in function of the migration pattern of herring and sprat. In the spring and the summer, in situ production ofcopepod biomass was higher than the in situ consumption by fish. During the fall and the winter, consumption exceeded production. This suggests that top–down control exerted by marine pelagic fish may be an important force structuring estuarine copepod populations.     

Growth variability of juvenile soles Solea solea and Solea senegalensis, and comparison with RNA : DNA ratios in the Tagus estuary, Portugal

Growth variability and condition of juvenile soles Solea solea and Solea senegalensis , were assessed through RNA : DNA estimates and compared to absolute growth rates. Higher mean cohort RNA : DNA ratios were observed for cohort I at the beginning of estuarine occurrence for both species (4·42 and 4·87, for S. solea and S. senegalensis respectively). Despite different estuarine colonization habits, no significant differences were observed between RNA : DNA monthly variation for both sole species within the same year ( P  > 0·05 for 2003 and 2004). Juvenile S. senegalensis showed significant differences between RNA : DNA ratios obtained for the two nursery areas ( P  < 0·001). The decrease of seasonal growth rates with fish age was similar to seasonal variation of mean RNA : DNA values. Thus the RNA : DNA pattern of juvenile S. solea and S. senegalensis reflected growth and estuarine colonization patterns.     

Seasonal variations in the energy density of fishes in the North Sea

The energy density ( E _D, kJ g^-1 wet mass) of saithe Pollachius virens , haddock Melanogrammus aeglefinus , whiting Merlangius merlangus , Norway pout Trisopterus esmarki , herring Clupea harengus , sprat Sprattus sprattus , sandeel Ammodytes marinus and pearlsides Maurolicus Muelleri , from the North Sea, increased with total length, L _T. However, there was not always a significant ( P> 0·05) linear relationship between L _T and E _D. Seasonal differences in E _D were obvious in mature fish, while geographical differences were insignificant. For all species there was a highly significant correlation ( P< 0·0001) between the percent dry mass of the fish ( D S) and E _D. A general relationship was established for gadoids and sandeel E _D=–3·1492+0·3459 D _S and herring E _D=–4·6395+0·4170 D _S. Thus seasonal and size-specific data on E _D needed for bioenergetics and gastric evacuation models can be determined simply from D _S, which is considerably less costly and time consuming than calorimetry or proximate analysis.     

The role of salinity in structuring the fish assemblages in a tropical estuary

The present study describes the seasonal changes of the fish species composition in three areas of the main channel of the Caeté River estuary, Brazil. The fish faunas of each habitat differed in density, biomass and species composition. Mean fish density and biomass for the Caeté River estuary channel was 0·25 individuals m⁻² and 0·9 g m⁻² respectively. Analysis of catch data showed that the number of species, total density and total biomass differed significantly between areas and seasons. For the most important species, the mean density of Cathorops spixii, Aspredinichthys filamentosus, Aspredo sp. 2, Pimelodus blochii, Pseudauchnipterus nodosus and Macrodon ancylodon , differed significantly between seasons while the mean density of Stellifer rastrifer , Stellifer microps, Aspredo aspredo , Aspredo sp. 1 and Cynoscion acoupa did not. The mean biomass of these species, with exception of S. microps and Aspredo sp. 1, also differed significantly between seasons. In the Caeté estuary seasonal salinity fluctuations appeared to be the main factor that structured the fish assemblage in the entire estuarine system. At least 85% of the species captured by the artisanal and subsistence fisheries in the Bragantine region required estuarine conditions to complete their life cycle.     

Feeding ecology of central Baltic Sea herring and sprat

A unique dataset of stomach contents sampled between 1977 and 1999 in the central Baltic Sea was used to perform a comprehensive study of the feeding ecology of Central Baltic herring Clupea harengus and sprat Sprattus sprattus . Both fish species were mainly preying upon calanoid copepods with Pseudocalanus sp. dominating the diet of herring, whereas sprat generally preferred Temora longicornis . Sprat preyed upon older copepodite stages, indicating size‐selective particulate feeding, whereas herring additionally fed on smaller copepodite stages, indicating occasional low food supply inducing filter‐feeding. Additional food sources other than copepods were mysids in winter and autumn for medium to large herring, as well as cladocerans for sprat in spring and summer, determined by the seasonal occurrence of these plankton species. Seasonally the highest feeding activity of both fishes species occurred in spring and summer, the main reproductive periods of calanoid copepods. The most important food item for both predators in spring was Pseudocalanus sp. In summer sprat switched to T. longicornis and Acartia spp. Since the late 1970s, the total stomach fullness decreased and the fraction of empty stomachs increased. In parallel the amount of Pseudocalanus sp. in the diets of both fish species decreased. Further, a considerable dietary overlap between both species in spring indicated considerable competition for food resources, especially due to an enlarged sprat stock. The results of this study support the hypothesis that growth reductions observed in Baltic herring and sprat are due to combination of a change in food availability and an increase in density‐dependent competition.     

Herring and Sprat Abundance Indices Predict Chick Growth and Reproductive Performance of Common Terns Breeding in the Wadden Sea

Herring (Clupea harengus) and sprat (Sprattus sprattus) are the key prey resources of common terns (Sterna hirundo) breeding in the Wadden Sea. Breeding success of the terns has been below average since 2002, coinciding with exceptionally low herring recruitment and sprat abundance. Time series of herring and sprat abundance in the North Sea and in the Wadden Sea were analyzed to explain long-term breeding success and chick development at two common tern breeding colonies. North Sea herring recruitment and sprat abundance in the Wadden Sea explained the largest part of common tern breeding success, both as single variables and in a multiple regression approach. Breeding success showed stronger correlations with herring recruitment indices derived from the North Sea region compared to the Wadden Sea. Also, herring and sprat abundance data explained more variability in breeding success than of more directly responding measures such as growth rate and maximum weight of chicks. Despite spatial and temporal incoherences between fish surveys and the common tern breeding season, breeding success of common terns reflected the abundance of their key prey fish beyond their foraging range and breeding season. We argue that the ecological connectivity between large- and small-scale herring abundance and the responsiveness of common tern breeding success is strong enough to establish a fish–seabird indicator system to be potentially valuable in monitoring and conservation.     

Fatty acid composition of sprat (<Emphasis Type="Italic">Sprattus sprattus</Emphasis>) and herring (<Emphasis Type="Italic">Clupea harengus</Emphasis>) in the Baltic Sea as potential prey for salmon (<Emphasis Type="Italic">Salmo salar</Emphasis>)

Sprat (Sprattus sprattus) and small herring (Clupea harengus) are the dominant prey fish of Atlantic salmon (Salmo salar) in the Baltic Sea. If the fatty acid (FA) proportions of sprat and herring differ, the dietary history of ascending salmon could be determined from their FA profiles. Therefore, we investigated the FA composition of several age groups of whole sprat and small herring, caught from the three main feeding areas of salmon in autumn and spring. Oleic acid (18:1n-9) was the most prevalent FA in sprat and characteristic of this species. In herring, palmitic acid (16:0) was the most common FA, but herring lipid was characterized by n-6 polyunsaturated FAs, and moreover, by palmitoleic acid (16:1n-7) and vaccenic acid (18:1n-7). Due to the higher lipid content of sprat, the concentrations of all other FAs, excluding these, were higher in sprat than in herring. The concentration of docosahexaenoic acid (DHA, 22:6n-3) increased with an increase in the lipid content and was consequently highest in the youngest specimens, being in young sprat almost double that of young herring, and 2.6 times higher in the sprat biomass than in that of herring. As a result of a decrease in the DHA concentration with age, the ratio thiamine/DHA increased with respect to age in both species, and was lower in sprat than in herring. It is concluded that an abundance of DHA in the diet of salmon most likely increases oxidative stress because of the susceptibility of DHA to peroxidation, and thus decreases thiamine resources of fasting, prespawning salmon. Because the FA composition of sprat and herring differs, and the relative abundancies of prey fish differ between the feeding areas of salmon, the feeding area of ascending salmon can most probably be derived by comparing their FA profiles.     

River flow,zooplankton and dominant zooplanktivorous fish dynamics in a warm‐temperate South African estuary

The possible links between river flow, zooplankton abundance and the responses of zooplanktivorous fishes to physico‐chemical and food resource changes are assessed. To this end, the seasonal abundance, distribution and diet of the estuarine round‐herring Gilchristella aestuaria and Cape silverside Atherina breviceps were studied in the Kariega Estuary. Spatio‐temporal differences were determined for selected physico‐chemical variables, zooplankton abundance and zooplanktivorous fish abundance and distribution. Results indicated that, following a river flood event in winter (>30 m³ s^?1), altered physico‐chemical conditions occurred throughout the estuary and depressed zooplankton stocks. Abundance of G. aestuaria was highest in spring, with this species dominant in the upper and middle zones of the estuary, while A. breviceps was dominant in summer and preferred the middle and lower zones. The catch per unit of effort of both zooplanktivores also declined significantly following the flooding, thus suggesting that these fishes are reliant on zooplankton as a primary food source for healthy populations. Copepods dominated the stomach contents of both fish species, indicating a potential for strong interspecific competition for food, particularly in the middle reaches. Temporal differences were evident in dietary overlap between the two zooplanktivorous fish species and were correlated with river flow, zooplankton availability and fish distribution. The findings of this study emphasize the close trophic linkages between zooplankton and zooplanktivorous fishes under changing estuarine environmental conditions, particularly river flow and provide important baseline information for similar studies elsewhere in South Africa and the rest of the world.     

Differential Use of Nursery Areas Within the Tagus Estuary by Sympatric Soles, Solea solea and Solea senegalensis

The spatio-temporal pattern of estuarine use by both Solea solea and Solea senegalensis was investigated between April 1995 and November 1996 by beam trawl surveys. Spatial and seasonal distribution and abundance were different for each species. The highest densities of S. solea were recorded in deeper, warmer, low salinity areas and where the sediment consisted of a high proportion of fine sand and presented a high abundance of amphipods. The 0-group individuals of this species first occurred on the nursery ground in April. Numbers peaked in June and July, and from October to April abundance was low. S. senegalensis had a wider distribution and its abundance was related to food availability. The 0-group individuals of this species occurred in the nursery areas later than S. solea, from June to August.     

Interannual and regional variations in body length, weight and energy content of age-0 Pacific herring from Prince William Sound, Alaska

Age-0 Pacific herring were surveyed in October of 4 years in a large northern Gulf of Alaska estuary, to determine the range of variations in length, weight and whole body energy content (WBEC). These parameters reflect their preparedness for surviving their first winter's fast. During the surveys there were distinct regional and interannual variations in all three parameters for individual groups of herring in Prince William Sound. Likewise, with each collection there was typically a large range of size and WBEC values. The average standard length was (±S.D.) 80±13 mm (range=40–118), the mean whole body wet weight was 5·7±3·0 g (range=0·7–29·2) and the average WBEC of all age-0 herring captured, regardless of year or site (n=1471), was 5·4± 1·0 kJ g⁻¹ wet weight (range=2·4–9·4). The large range of WBEC and size indicates that age-0 herring at different capture sites were not all equally prepared for surviving their first winter.     

Diet comparison between pelagic and demersal whiting in the North Sea

Pelagic North Sea whiting Merlangius merlangus preyed upon (pelagic) sprat and sandeel, while demersal whiting preyed upon (demersal) Norway pout and herring. Values of diet breadth were low for both feeding groups using Levin's index. Diet overlaps within layers were low ( D <0·25), while the between layer food overlap was moderate ( D =0·25–0·74) to high ( D >0·74) using Schoener's index. Selection of prey was density dependent. However, prey size also played an important role. The diet of whiting shifted from amphipods and mysids to fish with increasing predator length, and the length of prey consumed increased significantly with length of whiting. The fact that the stomach contents differed between the feeding groups demonstrates the need for a sampling design that includes both pelagic and demersal habitats when trying to quantify the diet of whiting.     

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.     

Variation in the calorific value and total energy content of the lesser sandeel (Ammodytes marinus) and other fish preyed on by seabirds

Wet and dry calorific values (energy densities) and total energy content of lesser sandeel Ammodytes marinus , sprat Sprattus sprattus , Atlantic herring Clupea harengus , whiting Merlangius merlangus , saithe Pollachius virens and cod Gadus morhua , were measured. Calorific values varied both within and between species. Larger fish tended to have higher calorific values than small ones, particularly when considered in terms of wet weight, but there was considerable temporal variation. The calorific values and body weights of sandeels larger than 10 cm showed marked seasonal trends and in consequence the total energy content of a sandeel of given length in summer is approximately double the spring value. The calorific values of herring and sprat also varied from month to month but seasonal cycles were less obvious. Whiting varied least. Both calorific values and total energy content of individual sprat were very variable in summer (the spawning season of this species). Because there is so much intraspecific variation, care must be taken when assessing the relative merits (in terms of energy) of different species as prey. The practical difficulties of obtaining reliable data on wet calorific values are discussed and the use of dry calorific values and dry weight/length relationships is recommended.     

The forth estuary: a nursery and overwintering area for North Sea fishes

The fish community of the Forth estuary, Scotland, has several components — estuarine resident species, diadromus migratory species, marine and freshwater adventitious species, marine juvenile migrants using the area as a nursery, and adults of marine species with seasonal migrations. The population changes during the period 1981–1988 in six species representative of the last two categories are described here. The species are the juvenile gadoids, whiting, Merlangius merlangus (L.) and cod, Gadus morhua L., juvenile flatfish, plaice, Pleuronectes platessa L. and common dab, Limanda limanda (L.), and the adult clupeids, sprat, Sprattus sprattus (L.) and herring, Clupea harengus L.The species' population changes are described in relation to aspects of wider significance such as commercial fishing for these species in North Sea areas. In particular, the underlying decline of the juvenile cod populations since 1981 in the Forth Estuary is discussed in relation to the recent reduction in commercial fishing quotas.     

The composition and community structure of the ichthyofauna of the upper Scheldt estuary: synthesis of a 10-year data collection (1991–2001)

Fish collections on the cooling water filter screens of the Doel power plant were used to analyse temporal trends in estuarine fish populations of the upper Scheldt estuary, Belgium. Between 1991 and 2001, 62 species shared the estuary, with 15 species present at any one time. Identified were 23 marine stragglers, 16 freshwater species and four diadromous species, but their contribution to the total catch was insignificant. In terms of numbers, marine estuarine opportunists (17 species) mainly occurring as juveniles and estuarine residents (two species) were the dominant life cycle categories. Six fish species contributed to >97% of the total catch: three Pomatoschistus species [P. minutus (32%), P. microps (30%) and P. lozanoi (6%)], two Clupeidae [Clupea harengus (16%) and Sprattus sprattus (8%)] and Syngnathus rostellatus (5%). Abundance of almost all species in the estuary was highly seasonal and characterized by pronounced abundance peaks. Ordination of the data showed that variability in the temporal structure of estuarine fish community was first determined by differences in annual recruitment and then by predictable, cyclical patterns of species abundance.     

Spatial and temporal variations in fish populations in the upper Thames estuary

Between February 1989 and August 1990, the upper Thames estuary contained 23 species of fish. Fish numbers were higher and relatively constant in the uppermost part of the estuary. Number of species was augmented in summer from fresh water and from downstream, coinciding with high temperature, low flow and high salinity. The eight most abundant species contributed to 98·5% of the total number. Flounder Pleuronectes flesus , dace Leuciscus leuciscus and perch Perca fluviatilis , recruited from May to August, and common goby Pomatoschistus microps , roach Rutilus rutilus and chub Leuciscus cephalus , from August to November. The upper estuary (salinity 0·34–2·96 p.s.u.) formed a species transition area between the freshwater but salinity-resistant roach, chub, and gudgeon Gobio gobio upstream, and the estuarine eurhyhaline common goby and flounder downstream. The three-spined stickleback Gasterosteus aculeatus and cyprinids were more abundant at upstream while perch was more abundant at downstream sites. High abundances of gudgeon, chub and roach were associated with high transparency and dissolved oxygen and low salinity, while high abundances of perch were associated with high salinity and low transparency. Dace and three-spined stickleback were associated with high dissolved oxygen and low pH, and common goby with high pH. Flounder showed no clear preferences.     

Disused docks as habitats for estuarine fish: a case study of Preston dock

A netting and hydrographic study of Preston dock was undertaken between May 1987 and April 1988. Fish populations were assessed using Lundgren monofilament multimesh gillnets. The dock was a low-salinity environment regularly replenished with estuarine water on spring tides. Influxes of higher-salinity water caused intermittent saline wedges in the deeper regions. The dock water was poorly mixed in the summer, resulting in bottom water anoxia. The water was eutrophic, and blue-green algal blooms were common for much of the year.
Fourteen freshwater, marine-estuarine, anadromous and catadromous species were present in the dock. No seasonal patterns in diversity were apparent, due to the dominance of the herring population. Changes in species composition and abundance seem to relate to seasonal migratory patterns and, in the case of the freshwater fish, estuarine salinity, rather than to water quality.
The suitablity of docks for sampling estuarine fish assemblages is discussed.     

Comparison of fish impingement by a thermal power station with fish populations in the Ems Estuary

On an annual basis >10 million individuals of c . 35 fish species are impinged at the EC20 unit of the Eems power station located on the Ems Estuary. The most abundant are: herring Clupea harengus , gobies ( Gobiidae ), Nilsson's pipefish Sygnathus rostellatus , three-spined stickleback Gasterosteus aculeatus and sprat Sprattus sprattus . The impingement shows a seasonal pattern which reflects the presence of the fishes in the estuary. The results are compared with other impingement studies at power stations on the Belgian and Dutch estuaries.
In 1995, the cooling water entrance was displaced from the shoreline to 300 m off the coast at the edge of the tidal channel. The effect of this relocation on the level of fish impingement, determined by comparing the ratio of fish density in the cooling water and in the estuary before (1992–1993) and after (1996–1997) the relocation of the intake canal is discussed.     

Inter‐annual variation in herring,Clupea harengus,and sprat,Sprattus sprattus,condition in the central Baltic Sea: what gives the tune?

The Baltic Sea ecosystem has undergone large changes during the last two decades, including a severe reduction in cod and herring biomass but, at the same time, a large increase in sprat abundance. The lower trophic levels of the Baltic Sea also changed due to environmental fluctuations, including variations in salinity and in volume of oxygenated water. In this apparently shifting environment, the conditions of herring and sprat have undergone large inter-annual variations during the past 15–20 years. In this study, we explore how abiotic factors (i.e. salinity and temperature) and biotic factors (biomass of the copepods Pseudocalanus elongatus , Temora longicornis , Acartia spp. and of cladocerans as well as clupeid abundance) in different seasons (May and August) affect clupeid body condition. Our analyses suggest that data of zooplankton biomass and abiotic factors in August have higher predictive power than May data. Although our analysis suggests that salinity (a bottom-up process) has an effect on sprat condition, total abundance of clupeids (a top-down process) is by far the most significant predictor of both herring and sprat condition. The strong correlation between clupeid abundance and total zooplankton biomass points to food competition and to top-down control by herring and sprat on common food resources. Furthermore, clupeid condition co-varied with the changes in the weight of zooplankton in the stomachs, which further suggest food competition being the main mechanism behind the changes in clupeid condition during the last two decades. Hence, our results are not in agreement with most of the current literature that has suggested that clupeid growth is regulated by environmentally mediated bottom-up processes acting on the abundance of copepods. This is, to our knowledge, the first evidence of food resources mediated density-dependent fish growth in a large marine ecosystem.     

Associations between wintering avian predators and schooling fish in the Skagerrak-Kattegat suggest reliance on predictable aggregations of herring Clupeaharengus

We investigated the distributional dynamics of piscivorous seabirds in relation to the distribution of young fish outside the breeding season in the Skagerrak and Kattegat, using synoptic bottom trawl and seabird surveys. Kittiwake Rissa tridactyla , Common Guillemot Uria aalge and Razorbill Alca torda were significantly correlated with herring Clupea harengus abundance at coarse spatial scales (>20 km). The association between the three seabird species and other abundant fish species like sprat Sprattus sprattus , whiting Merlangius merlangus and Norway pout Trisopterus esmarki was considerably weaker. In the Skagerrak and Kattegat, the location of aggregations of immature herring during winter is relatively stable and predictable. The correlation with herring and the lack of correlation with the abundant, but less stable occurrence of sprat of similar caloric value suggest that the tendency among piscivorous seabirds to depend on predictable concentrations of prey during the breeding season is a common feature during the non-breeding season as well. The results are discussed in relation to ecological and winter habitat characteristics of wintering piscivorous seabirds in the eastern North Atlantic, and in relation to the management and conservation of nursering herring in the Skagerrak and Kattegat.