Factors controlling long-term changes of the eutrophicated ecosystem of Pärnu Bay,Gulf of Riga

Phytoplankton, mesozooplankton, mysids and fish larvae were studied during 15–29 annual cycles measured weekly to monthly in Pärnu Bay, the Gulf of Riga. The monthly variability of the biological data was related to temperature, ice conditions, salinity, influx of nutrients, the North Atlantic Oscillation (NAO) index, cloudiness and solar activity. Phytoplankton development was mainly a function of the NAO index. For the whole study period the abundance of zooplankton increased with increasing water temperature and solar activity. Significant correlations between phytoplankton and zooplankton densities were found until 1990. After the invasion of the predatory cladoceran Cercopagis pengoi in 1991, the zooplankton community was likely to be regulated by the introduced species rather than phytoplankton dynamics. The increased abundances of rotifers and copepods triggered the increase in mysid densities. The development of herring larvae was positively affected by the high density of copepods and rotifers but also by increased eutrophication. Until 1990 there was no significant relationship between the density of zooplankton and herring larvae. A negative relationship between the density of zooplankton and herring larvae in the 1990s suggests that the major shift in zooplankton community resulted in food limitation for herring larvae. The results indicated that (1) atmospheric processes in the northern Atlantic explain a large part of the interannual variation of the local phytoplankton stock, (2) trophic interactions control the development of pelagic communities at higher trophic levels, and (3) the introduction of an effective intermediate predator has repercussions for the whole pelagic food web in Pärnu Bay.     

Direct and indirect effects of predatory young‐of‐year fishes in a dryland river food web

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Trophic shift in young‐of‐the‐year Mugilidae during salt‐marsh colonization

This study investigated the trophic shift of young‐of‐the‐year (YOY) thinlip grey mullet Liza ramada and golden grey mullet Liza aurata during their recruitment in a salt marsh located on the European Atlantic Ocean coast. Stable‐isotope signatures (δ¹³C and δ¹⁵N) of the fishes followed a pattern, having enrichments in ¹³C and ¹⁵N with increasing fork length (L_F): δ¹³C in fishes < 30 mm ranged from ?19.5 to ?15.0‰, whereas in fishes > 30 mm δ¹³C ranged from ?15.8 to ?12.7‰, closer to the level in salt‐marsh food resources. Large differences between the δ¹⁵N values of mugilids and those of food sources (6·0‰ on average) showed that YOY are secondary consumers, similar to older individuals, when feeding in the salt marsh. YOY mugilids shift from browsing on pelagic prey to grazing on benthic resources from the salt marsh before reaching 30 mm L_F. The results highlight the role of European salt marshes as nurseries for juvenile mugilids.     

Wasp-waist interactions in the North Sea ecosystem

Background

In a “wasp-waist” ecosystem, an intermediate trophic level is expected to control the abundance of predators through a bottom-up interaction and the abundance of prey through a top-down interaction. Previous studies suggest that the North Sea is mainly governed by bottom-up interactions driven by climate perturbations. However, few studies have investigated the importance of the intermediate trophic level occupied by small pelagic fishes.

Methodology/Principal Findings

We investigated the numeric interactions among 10 species of seabirds, two species of pelagic fish and four groups of zooplankton in the North Sea using decadal-scale databases. Linear models were used to relate the time series of zooplankton and seabirds to the time series of pelagic fish. Seabirds were positively related to herring (Clupea harengus), suggesting a bottom-up interaction. Two groups of zooplankton; Calanus helgolandicus and krill were negatively related to sprat (Sprattus sprattus) and herring respectively, suggesting top-down interactions. In addition, we found positive relationships among the zooplankton groups. Para/pseudocalanus was positively related to C. helgolandicus and C. finmarchicus was positively related to krill.

Conclusion/Significance

Our results indicate that herring was important in regulating the abundance of seabirds through a bottom-up interaction and that herring and sprat were important in regulating zooplankton through top-down interactions. We suggest that the positive relationships among zooplankton groups were due to selective foraging and switching in the two clupeid fishes. Our results suggest that “wasp-waist” interactions might be more important in the North Sea than previously anticipated. Fluctuations in the populations of pelagic fish due to harvesting and depletion of their predators might accordingly have profound consequences for ecosystem dynamics through trophic cascades.     

A thermal effluent as a sporadic cornucopia: effects on fish and zooplankton

Synopsis We report effects of a thermal effluent on fish and zooplankton in an impoundment in South Carolina. The effects are not directly due to heated effluent but are indirect effects on the trophic dynamics of the system and depend on the physical structure of the system, season, and geography. As effluent from a nuclear reactor cooling pond (Pond C) enters the larger Par Pond it sporadically carries with it zooplankton and/or dead fish. Pond C produces high zooplankton densities in two situations: (1) high and hot effluent flow during cold winter weather; and (2) low (cool) effluent flow during warmer seasons. High zooplankton densities attract Blueback herring, Alosa aestivalis, which attract largemouth bass, Micropterus salmoides. Dying and dead bluegills, Lepomis macrochirus from Pond C (heat killed) are eaten by Par Pond bass which swim into effluent temperatures as high as 46°C to take these easy prey. Blueback herring and largemouth bass distribution are affected by the presence of food rather than by temperature. Par Pond blueback herring appear to be available to bass mainly when herring are near the effluent from Pond C and when they spawn in spring in the littoral zone. Because Blueback herring in Par Pond live only one year and because their presence in bass habitat is seasonal there is a strong seasonal component to bass food abundance. A previously reported annual oscillation in bass condition (K), with a peak in winter, occurs throughout Par Pond but is extreme in the vicinity of the effluent. The strong seasonality of food abundance for bass at the effluent correlates the winter peak in bass condition.     

Mesozooplankton Grazing on Picocyanobacteria in the Baltic Sea as Inferred from Molecular Diet Analysis

Our current knowledge on the microbial component of zooplankton diet is limited, and it is generally assumed that bacteria-sized prey is not directly consumed by most mesozooplankton grazers in the marine food webs. We questioned this assumption and conducted field and laboratory studies to examine picocyanobacteria contribution to the diets of Baltic Sea zooplankton, including copepods. First, qPCR targeting ITS-1 rDNA sequence of the picocyanobacteria Synechococcus spp. was used to examine picocyanobacterial DNA occurrence in the guts of Baltic zooplankton (copepods, cladocerans and rotifers). All field-collected zooplankton were found to consume picocyanobacteria in substantial quantities. In terms of Synechococcus quantity, the individual gut content was highest in cladocerans, whereas biomass-specific gut content was highest in rotifers and copepod nauplii. Moreover, the gut content in copepods was positively related to the picocyanobacteria abundance and negatively to the total phytoplankton abundance in the water column at the time of sampling. This indicates that increased availability of picocyanobacteria resulted in the increased intake of this prey and that copepods may rely more on picoplankton when food in the preferred size range declines. Second, a feeding experiments with a laboratory reared copepod Acartia tonsa fed a mixture of the picocyanobacterium Synechococcus bacillaris and microalga Rhodomonas salina confirmed that copepods ingested Synechococcus, even when the alternative food was plentiful. Finally, palatability of the picocyanobacteria for A. tonsa was demonstrated using uptake of ¹³C by the copepods as a proxy for carbon uptake in feeding experiment with ¹³C-labeled S. bacillaris. These findings suggest that, if abundant, picoplankton may become an important component of mesozooplankton diet, which needs to be accounted for in food web models and productivity assessments.     

Trophic polymorphism in bluegill sunfish (<Emphasis Type="Italic">Lepomis macrochirus</Emphasis>) introduced into Lake Biwa: evidence from stable isotope analysis

Trophic polymorphism was recently reported in introduced bluegill (Lepomis macrochirus) in Lake Biwa, Japan, where three morphs are specialized in benthic invertebrates (benthivorous type), submerged aquatic plants (herbivorous type), and zooplankton (planktivorous type). We evaluated the long-term effects of food resource utilization by these trophic morphs using stable isotope ratios, δ¹⁵N and δ¹³C. A significant difference in δ¹⁵N was found between the benthivorous and planktivorous types. The planktivorous type had the higher δ¹⁵N value, which corresponded with the value expected from its prey, zooplankton. The lower δ¹⁵N value of the benthivorous type would be derived from the lower δ¹⁵N values of benthic prey organisms compared to zooplankton. These results support previous findings that the benthivorous and planktivorous types have different food resource utilization. In contrast, the δ¹⁵N and δ¹³C values of the herbivorous type were distinctly different from the expected values, indicating that this type was unlikely to utilize aquatic plants substantially, contradicting the results of the dietary analysis.     

Metabolism and foraging strategies of mid‐latitude mesozooplankton during cyanobacterial blooms as revealed by fatty acids,amino acids,and their stable carbon isotopes

Increasing sea surface temperatures (SST) and blooms of lipid‐poor, filamentous cyanobacteria can change mesozooplankton metabolism and foraging strategies in marine systems. Lipid shortage and imbalanced diet may challenge the build‐up of energy pools of lipids and proteins, and access to essential fatty acids (FAs) and amino acids (AAs) by copepods. The impact of cyanobacterial blooms on individual energy pools was assessed for key species temperate Temora longicornis and boreal Pseudo‐/Paracalanus spp. that dominated field mesozooplankton communities isolated by seasonal stratification in the central Baltic Sea during the hot and the cold summer. We looked at (a) total lipid and protein levels, (b) FA trophic markers and AA composition, and (c) compound‐specific stable carbon isotopes (δ¹³C) in bulk mesozooplankton and in a subset of parameters in particulate organic matter. Despite lipid‐poor cyanobacterial blooms, the key species were largely able to cover both energy pools, yet a tendency of lipid reduction was observed in surface animals. Omni‐ and carnivory feeding modes, FA trophic makers, and δ¹³C patterns in essential compounds emphasized that cyanobacterial FAs and AAs have been incorporated into mesozooplankton mainly via feeding on mixo‐ and heterotrophic (dino‐) flagellates and detrital complexes during summer. Foraging for essential highly unsaturated FAs from (dino‐) flagellates may have caused night migration of Pseudo‐/Paracalanus spp. from the deep subhalocline waters into the upper waters. Only in the hot summer (SST>19.0°C) was T. longicornis submerged in the colder subthermocline water (~4°C). Thus, the continuous warming trend and simultaneous feeding can eventually lead to competition on the preferred diet by key copepod species below the thermocline in stratified systems. A comparison of δ¹³C patterns of essential AAs in surface mesozooplankton across sub‐basins of low and high cyanobacterial biomasses revealed the potential of δ¹³C‐AA isoscapes for studies of commercial fish feeding trails across the Baltic Sea food webs.     

Size‐based interactions and trophic transfer efficiency are modified by fish predation and cyanobacteria blooms in Lake Mývatn,Iceland

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Biochemical tracers reveal intra-specific differences in the food webs utilized by individual seabirds

Food web structure regulates the pathways and flow rates of energy, nutrients, and contaminants to top predators. Ecologically and physiologically meaningful biochemical tracers provide a means to characterize and quantify these transfers within food webs. In this study, changes in the ratios of stable N isotopes (e.g., δ¹⁵N), fatty acids (FA), and persistent contaminants were used to trace food web pathways utilized by herring gulls (Larus argentatus) breeding along the shores of the St Lawrence River, Canada. Egg δ¹⁵N values varied significantly among years and were used as an indicator of gull trophic position. Temporal trends in egg δ¹⁵N values were related to egg FA profiles. In years when egg δ¹⁵N values were greater, egg FA patterns reflected the consumption of more aquatic prey. Egg δ¹⁵N values were also correlated with annual estimates of prey fish abundance. These results indicated that temporal changes in aquatic prey availability were reflected in the gull diet (as inferred from ecological tracer profiles in gull eggs). Analysis of individual eggs within years confirmed that birds consuming more aquatic prey occupied higher trophic positions. Furthermore, increases in trophic position were associated with increased concentrations of most persistent organic contaminants in eggs. However, levels of highly brominated polybrominated diphenyl ether congeners, e.g, 2,2′,3,3′,4,4′,5,5′,6,6′-decabromoDE (BDE-209), showed a negative relationship with trophic position. These contrasting findings reflected differences among contaminant groups/homologs in terms of their predominant routes of transfer, i.e., aquatic versus terrestrial food webs. High trophic level omnivores, e.g., herring gulls, are common in food webs. By characterizing ecological tracer profiles in such species we can better understand spatial, temporal, and individual differences in pathways of contaminant, energy, and nutrient flow.     

River habitat homogenisation enhances trophic competition and promotes individual specialisation among young of the year fish

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Relationships between microzooplankton and mesozooplankton: competition versus predation on natural assemblages of the Gulf of Trieste (northern Adriatic Sea)

We performed, on a seasonal basis, 16 dilution experiments and,simultaneously, copepod or cladoceran grazing experiments onnatural assemblages from Gulf of Trieste (northern AdriaticSea). The autotrophic fraction was almost entirely composedof diatoms in late winter. As the seasons progressed, relativeabundance of nanoplankton and cyanobacteria increased. Microzooplanktonwas always present in the diet of mesozooplankton, even if inpercentages usually not exceeding 6% of diet intake on carbonbasis. Microzooplankton took advantage of ephemeral increasesof autotrophic biomass when prey were in the optimal size rangebut did not consume diatoms when these were large. When autotrophicresources were scarce, micrograzers used heterotrophic biomasswhich, in turn, fuelled the upper trophic levels through predationby mesozooplankton on microzooplankton. Microzooplankton grazingwas the most important loss term of primary production in theGulf of Trieste (on average, microzooplankton consumed 100%of primary production, while mesozooplankton only 76%), whichcan be considered a mesotrophic coastal system. This paper is one of six on the subject of the role of zooplanktonpredator–prey interactions in structuring plankton communities.     

A review of the adaptive significance and ecosystem consequences of zooplankton diel vertical migrations

Diel vertical migration (DVM) by zooplankton is a universal feature in all the World's oceans, as well as being common in freshwater environments. The normal pattern involves movement from shallow depths at night to greater depths during the day. For many herbivorous and omnivorous mesozooplankton that feed predominantly near the surface on phytoplankton and microzooplankton, minimising the risk of predation from fish seems to be the ultimate factor behind DVM. These migrants appear to use deep water as a dark daytime refuge where their probability of being detected and eaten is lower than if they remained near the surface. Associated with these vertical movements of mesozooplankton, predators at higher trophic levels, including invertebrates, fish, marine mammals, birds and reptiles, may modify their behaviour to optimise the exploitation of their vertically migrating prey. Recent advances in biotelemetry promise to allow the interaction between migrating zooplankton and diving air-breathing vertebrates to be explored in far more detail than hitherto.     

Carbon sources and trophic structure in a macrophyte‐dominated polyculture pond assessed by stable‐isotope analysis

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Species‐specific separation of lake plankton reveals divergent food assimilation patterns in rotifers

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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.     

Feeding differences among common littoral mysids, <Emphasis Type="Italic">Neomysis integer,Praunus flexuosus</Emphasis> and <Emphasis Type="Italic">P. inermis</Emphasis>

The energy flow in aquatic food webs and their structures are largely determined by food utilisation of predators. Mysid shrimps are important predators in various aquatic ecosystems. We studied the stomach contents of three common littoral mysids from the Baltic Sea. The aim was to study whether the diets differ between species and size classes inhabiting shallow coastal areas. The effects of season (spring, summer, autumn) and habitat were also explored. Results showed that all species were highly omnivorous, utilising various phyto- and zooplankton prey, algal filaments and dead organic material through the growing season. No ontogenetic diet shifts were observed although different size classes preferred slightly different prey. The amount of detritus increased in the diets during growth. In addition, large mysids ate more macro- and less microzooplankton compared with the small ones. There were also species-specific differences in the food utilisation. Neomysis integer ate more benthic material, Praunus flexuosus more macrozooplankton and P. inermis more phytoplankton compared with the others. These differences reflect microhabitat differences and probably also size differences of the studied species. Seasonal variation was also observed in the diets. Food utilisation followed the changes in the food availability, e.g. phytoplankton spring bloom and zooplankton peak abundances in late summer. Results confirm the omnivorous nature of mysids showing the importance of a diversity of prey as energy sources during growth.     

Observations on the biology and seasonal variation in feeding of the east coast round herring Etrumeus wongratanai (Clupeiformes), off Scottburgh,KwaZulu-Natal,South Africa

The basic biology and ecology of the South African east coast round herring Etrumeus wongratanai was investigated from samples of fish collected between 2013 and 2016. This species is short-lived and reaches a maximum of 3 years of age, with rapid growth in its first year of life. It reproduces from June to December (austral summer) and condition factor was lowest in May through to August and increased from September, probably reflecting the physiological strain before and during spawning. Fish larvae were the most important food items consumed during summer, whereas eucalanid copepods were the most important prey at other times of the year. Stable-isotope data suggest that there are gradual changes in the trophic level with increasing fish size, δ¹⁵N and δ¹³C values also differed between seasons. The results obtained here are compared with those of other Etrumeus species, regionally and globally.     

Variable littoral‐pelagic coupling as a food‐web response to seasonal changes in pelagic primary production

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Evaluating zooplankton indicators using signal detection theory

Indicators are used to help managers to conserve biodiversity and guarantee the sustainable use of marine resources. Good indicators are scientifically valid, ecologically relevant, respond to pressures, and it is possible to set target levels for them. Zooplankton is an important link in the food web as it transfers energy from primary producers (phytoplankton) to planktivorous fish, including the commercially important herring in the Baltic Sea. Eutrophication is known to increase zooplankton abundance and decrease the mean size of zooplankton individuals, while particularly herring prefers larger zooplankton as prey. Therefore, both the abundance/biomass and the size structure of the zooplankton community are highly relevant for the functioning of the pelagic food web, and their combination has been proposed as an indicator of the status of the pelagic food web. In this study, we evaluated the indicator performance of zooplankton abundance and mean size in the northern Baltic Sea using signal detection theory and annual zooplankton monitoring data from years 1979–2014. Herring weight-at-age and chlorophyll a levels were used to estimate the reference periods or “gold standard” of the food web status. The sensitivity and specificity of the indicator was evaluated using ROC curves. Thresholds were set and evaluated with positive and negative predictive values for the zooplankton mean size in three sub-basins of the Baltic Sea. The results suggest that zooplankton mean size is able to reflect the state of the food web in the Baltic Sea. Our study also confirms that signal detection theory is useful in evaluating ecological indicators with clear pressures. However, with parameters that are affected by multiple contrary pressures, such as zooplankton abundance, ROC curves cannot offer enough information about the performance of an indicator parameter.