Seasonal depth distribution and thermal experience of the non-indigenous round goby Neogobius melanostomus in the Baltic Sea: implications to key trophic relations

Biological Invasions - Native to the Ponto-Caspian region, the benthic round goby (Neogobius melanostomus) has invaded several European inland waterbodies as well as the North American Great Lakes...     

Invasion of round goby to the temperate salmonid streams in the Baltic Sea

Ichthyological Research -     

Predator and prey: the role of the round goby Neogobius melanostomus in the western Baltic

Different studies on the position of the non-indigenous species Neogobius melanostomus within the coastal food web of the Pomeranian Bay (western Baltic) were performed, resulting in a quantitative and qualitative species list of prey organisms found in the stomachs of the invader and an estimation concerning the importance of round goby as prey for different resident predators. It seems that the colonization process is not fully completed yet, but the results reveal that the species is already established in the food web 16 years after the first observation within the study area. The results show that N. melanostomus feed upon a wide range of different resident organisms. While a direct predation effect on native fish species appears rather unlikely, indirect effects such as competition cannot yet be excluded. In addition, our results reveal an ontogenetic diet shift and that the round goby itself already serves as an important prey for piscivorous fish and seabirds. Finally, we formulate different hypotheses based on our results which will require further research.     

Ecological niche differentiation between native and non-native shrimps in the northern Baltic Sea

Invasions of non-native species are modifying global biodiversity but the ecological mechanisms underlying invasion processes are still not well understood. A degree of niche separation of non-native and sympatric native species can possibly explain the success of novel species in their new environment. In this study, we quantified experimentally and in situ the environmental niche space of caridean shrimps (native Crangon crangon and Palaemon adspersus, non-native Palaemon elegans) inhabiting the northern Baltic Sea. Field studies showed that the non-native P. elegans had wider geographical range compared to native species although the level of habitat specialization was similar in both Palaemon species. There were clear differences in shrimp habitat occupancy with P. elegans inhabiting lower salinity areas and more eutrophicated habitats compared to the native species. Consequently, the non-native shrimp has occupied large areas of the northern Baltic Sea that were previously devoid of the native shrimps. Experiments demonstrated that the non-native shrimp had higher affinity to vegetated substrates compared to native species. The study suggests that the abilities of the non-native shrimp to thrive in more stressful habitats (lower salinity, higher eutrophication), that are sub-optimal for native shrimps, plausibly explain the invasion success of P. elegans.     

Fifty years later: trophic ecology and niche overlap of a native and non-indigenous fish species in the western basin of Lake Erie

Since the introduction of white perch (Morone americana) into Lake Erie over 50 years ago, the population size of native yellow perch (Perca flavescens) has decreased up to 79 % and significant changes to the ecosystem have occurred. We examined long-term population estimates and used stable isotopes of carbon (δ ¹³C) and nitrogen (δ ¹⁵N) paired with stomach content analysis to quantify the trophic ecology and niche overlap of adult yellow perch and white perch in the western basin of Lake Erie. We found that changes in yellow perch abundance since 1979 appeared to be better correlated with changes in fishery exploitation rates than with food competition effects from white perch. At the time of this study, yellow perch were found to have higher δ ¹³C values, indicating greater utilization of benthic food resources than white perch, and white perch occupied higher trophic positions based on δ ¹⁵N. The diets of both species varied spatially and seasonally based on stable isotopes and stomach contents, likely driven by changes in prey abundance. Comparison of niche widths using stable isotope population metrics and Schoener diet similarity index suggested a low to moderate degree of niche overlap between species. Isotopic niches of white perch were generally larger than those of yellow perch demonstrating broader resource utilization by this non-indigenous species. We submit that isotopic niche overlap comparisons are more appropriate for studies seeking to understand interactions among populations over course temporal scales, while diet overlap indices, such as the Schoener index provide a means to study fine-scale interactions such as ontogenetic and seasonal diet shifts.     

Establishment of trophic continuum in the food web of the Yellow Sea and East China Sea ecosystem: Insight from carbon and nitrogen stable isotopes

Stable carbon and nitrogen isotope ratios (δ¹³C and δ¹⁵N) are used to study the trophic structure of food web in the Yellow Sea and East China Sea ecosystem. The trophic continuum of pelagic food web from phytoplankton to top preyer was elementarily established, and a trophic structure diagram in the Yellow Sea and East China Sea was outlined in combination with carbon isotopic data of benthic organisms, which is basically consistent with and makes some improvements on the simplified Yellow Sea food web and the trophic structure diagram drawn based on the biomass of main resource population during 1985–1986. This result indicates that the stable isotope method is a potential useful means for further studying the complete marine food web trophic continuum from viruses to top predators and food web stability.     

Establishment of trophic continuum in the food web of the Yellow Sea and East China Sea ecosystem: Insight from carbon and nitrogen stable isotopes

The determination of trophic level for the biology in a marine ecosystem is very important as alteration of its structure and function may be reflected in the tro-phic level of component species. A change in trophic level indicates variation in an organism’s feeding bi-ology or in the pathway of energy flow from primary producers to the consumer. The gut content analysis is a traditional method for studying trophodynamics of food web in marine ecosystems. Species composition and amounts in al…     

Trophic niche differences between coexisting omnivores silver carp and bighead carp in a pelagic food web

Understanding how omnivorous consumers are affected by their resources and how this is expressed through the food chain is a fundamental issue in ecology. We used stable isotope analysis of archived scales of two pelagic single-chain omnivorous fish species, bighead carp (Hypophthalmichthys nobilis) and silver carp (H. molitrix), to reconstruct historical trophic interactions patterns along a gradient of resources. We found that, although bighead carp and silver carp utilize the similar resources from the pelagic food chain, they can coexist and persist not only by regulating their trophic position and trophic dissimilarity, but also by regulating trophic niche width. Omnivorous fish often exhibit flexible foraging strategies, which is closely related to the availability of ecologic context. We found a positive relationship between trophic dissimilarity and zooplankton density, which may indicate that the competitive interactions induce strong top-down effects on zooplankton, and/or that high zooplankton availability release the between-population trophic interaction through bottom-up effect. The trophic niche width of bighead carp was positively related with zooplankton availability, probably reflecting that the niche of an omnivore at a higher trophic position is more sensitive to high quality resources. Our results indicate how different aspects of the trophic partitioning of coexisting omnivores may be regulated by different ecological contexts. These alternatives are not mutually exclusive and further theoretical work should include both these mechanisms to re-evaluate the effects of omnivory on food web properties.     

Differential modes of selection on the rhodopsin gene in coastal Baltic and North Sea populations of the sand goby,Pomatoschistus minutus

An excellent model to elucidate the mechanisms and importance of evolution in the marine environment is the spectral tuning mechanism of the visual pigment in vertebrates. In the sand goby Pomatoschistus minutus (Teleostei; Gobiidae), a distribution‐wide study showed that spatial variation at the rhodopsin gene (RH1) matches the characteristics of specific light environments. This match suggests that populations are locally adapted to selective light regimes targeting the RH1 gene. If so, then the direction of selection should depend on the regional spatial and temporal stability of the light conditions. We tested this prediction by comparing goby populations from two regions: the Baltic Sea, characterized by divergent, but temporally stable light conditions, and the North Sea, characterized by locally heterogeneous and temporally variable light conditions. RH1 sequences of 491 Pomatoschistus minutus individuals from 15 locations were analysed. We found that variation at the RH1 gene in the Baltic populations showed signatures of diversifying selection, whereas the RH1 gene in the North Sea showed signatures of stabilizing selection. These different modes of selection are consistent with the regional light conditions and hence support our predictions, but may also be influenced by migration between the open sea and more turbid estuarine environments. An interesting observation is that within one gene, synonymous and non‐synonymous SNPs show a totally different pattern between populations. Population differentiation based on non‐synonymous SNPs of the RH1 gene correlated with spectral variation of the local environment of the sand goby populations. In contrast, the differentiation based on synonymous SNPs of RH1 reflects more the neutral historical pattern of the species.     

Genetic divergence and a hybrid zone between Baltic and North Sea Mytilus populations (Mytilidae: Mollusca)

Populations of the common mussel ( Mytilus edulis ) from the North Sea area (Skakerrak-Kattegat) and those from the Baltic Sea are almost diagnostically differentiated at five out of 22 studied allozyme loci; at a further seven loci, alleles predominant or common in one area are nearly absent in the other. Genetic distance was estimated at 0.28; this is similar to the distances of these populations to the Mediterranean mussel M. galloprovincialis. The three mussel types obviously represent equal evolutionary divergence from one another, and should also be taxonomically equally separated; a semispecies rank within a more comprehensive M. edulis complex or superspecies is suggested. The age of the Baltic mussel type ( 'M. trossulus' ), as an independent evolutionary lineage, is probably far greater than that of the post-glacial Baltic Sea.
Allele frequencies change gradually and in parallel when entering from the Kattegat through the Sound into the Baltic. Only a slight Wahlund effect at the strongly diverged Gpi and Pgm loci was found in intermediate populations, indicating that extensive hybridization of the two taxa takes place in the area. However, strong interlocus genotypic associations suggest that selection against hybrids is intense in later generations; the c. 100 km wide hybrid zone is narrow relative to the dispersal distance. The genotypic structure of the Lap locus does not conform with those of the other loci studied in the hybrid zone; it cannot be viewed merely as a neutral marker of the process of hybridization.     

13C-isotope analyses reveal that chemolithoautotrophic Gamma- and Epsilonproteobacteria feed a microbial food web in a pelagic redoxcline of the central Baltic Sea

Marine pelagic redoxclines are zones of high dark CO₂ fixation rates, which can correspond up to 30% of the surface primary production. However, despite this significant contribution to the pelagic carbon cycle, the identity of most chemolithoautotrophic organisms is still unknown. Therefore, the aim of this study was to directly link the dark CO₂ fixation capacity of a pelagic redoxcline in the central Baltic Sea (Landsort Deep) with the identity of the main chemolithoautotrophs involved. Our approach was based on the analysis of natural carbon isotope signatures in fatty acid methyl esters (FAMEs) and on measurements of CO₂ incorporation in ¹³C-bicarbonate pulse experiments. The incorporation of ¹³C into chemolithoautotrophic cells was investigated by rRNA-based stable isotope probing (RNA-SIP) and FAME analysis after incubation for 24 and 72 h under in situ conditions. Our results demonstrated that fatty acids indicative of Proteobacteria were significantly enriched in ¹³C slightly below the chemocline. RNA-SIP analyses revealed that two different Gammaproteobacteria and three closely related Epsilonproteobacteria of the Sulfurimonas cluster were active dark CO₂-fixing microorganisms, with a time-dependent community shift between these groups. Labelling of Archaea was not detectable, but after 72 h of incubation the ¹³C-label had been transferred to a potentially bacterivorous ciliate related to Euplotes sp. Thus, RNA-SIP provided direct evidence for the contribution of chemolithoautotrophic production to the microbial food web in this marine pelagic redoxcline, emphasizing the importance of dark CO₂-fixing Proteobacteria within this habitat.     

Complex life cycles in a pond food web: effects of life stage structure and parasites on network properties,trophic positions and the fit of a probabilistic niche model

Most food webs use taxonomic or trophic species as building blocks, thereby collapsing variability in feeding linkages that occurs during the growth and development of individuals. This issue is particularly relevant to integrating parasites into food webs because parasites often undergo extreme ontogenetic niche shifts. Here, we used three versions of a freshwater pond food web with varying levels of node resolution (from taxonomic species to life stages) to examine how complex life cycles and parasites alter web properties, the perceived trophic position of organisms, and the fit of a probabilistic niche model. Consistent with prior studies, parasites increased most measures of web complexity in the taxonomic species web; however, when nodes were disaggregated into life stages, the effects of parasites on several network properties (e.g., connectance and nestedness) were reversed, due in part to the lower trophic generality of parasite life stages relative to free-living life stages. Disaggregation also reduced the trophic level of organisms with either complex or direct life cycles and was particularly useful when including predation on parasites, which can inflate trophic positions when life stages are collapsed. Contrary to predictions, disaggregation decreased network intervality and did not enhance the fit of a probabilistic niche model to the food webs with parasites. Although the most useful level of biological organization in food webs will vary with the questions of interest, our results suggest that disaggregating species-level nodes may refine our perception of how parasites and other complex life cycle organisms influence ecological networks.     

Assessing the trophic impacts of marine mammals: From metabolism to food web indices

Marine mammals are an important part of ecosystems, and their trophic role and potential impact have been increasingly studied. One key question is how these large animals interact with fisheries or compete for similar resources. Consequently, some models once used only for fisheries management are now including pinnipeds and cetaceans. However, fish and marine mammals do not share the same ecology and bioenergetics, and complex ecosystem models may not be the best way to assess the impact of pinnipeds or cetaceans in food webs. Indeed, simpler methods based on thermodynamics might give us reasonable answers with limited amounts of data. Here, we present an assessment of two different approaches to assess the trophic role of marine mammals in the northern Gulf of St. Lawrence (Canada): mixed trophic impacts (MTI) based on ecosystem modeling and surface index (SI) impact based on bioenergetics. Our results show that while modeling represents a good way of getting a holistic view of the role of marine mammals in ecosystems, trophic impact estimates based on fundamental thermodynamics principles can also give us answers requiring less data. The body surface area approach presented here might provide a practical tool for ecologists, who are not necessarily ecosystem modelers, to study this issue.     

Sedimentation of Nodularia spumigena and distribution of nodularin in the food web during transport of a cyanobacterial bloom from the Baltic Sea to the Kattegat

Nodularia spumigena is a toxic cyanobacteria that blooms in the Baltic Sea every year. In the brackish water of the Baltic Sea, its toxin, nodularin, mainly affects the biota in the surface water due to the natural buoyancy of this species. However, the fate of the toxin is unknown, once the cyanobacteria bloom enters the more saline waters of the Kattegat. In order to investigate this knowledge gap, a bloom of N. spumigena was followed during its passage, carried by surface currents, from the Baltic Sea into the Kattegat area, through the Öresund strait. N. spumigena cells showed an increased cell concentration through the water column during the passage of the bloom (up to 130 10³ cells ml⁻¹), and cells (4.2 10³ cells ml⁻¹) could be found down to 20 m depth, below a pycnocline. Sedimentation trap samples from below the pycnocline (10–12 m depth) also showed an increased sedimentation of N. spumigena filaments during the passage of the bloom. The toxin nodularin was detected both in water samples (0.3–6.0 μg l⁻¹), samples of sedimenting material (a toxin accumulation rate of 20 μg m^-2 day⁻¹), zooplankton (up to 0.1 ng ind.⁻¹ in copepods), blue mussels (70–230 μg kg⁻¹ DW), pelagic and benthic fish (herring (1.0–3.4 μg kg⁻¹ DW in herring muscle or liver) and flounder (1.3-6.2 μg kg⁻¹ DW in muscle, and 11.7-26.3 μg kg⁻¹ DW in liver). A laboratory experiment showed that N. spumigena filaments developed a decreased buoyancy at increased salinities and that they were even sinking with a rate of up to 1,7 m day⁻¹ at the highest salinity (32 PSU). This has implications for the fate of brackish water cyanobacterial blooms, when these reach more saline waters. It can be speculated that a significant part of the blooms content of nodularin will reach benthic organisms in this situation, compared to blooms decaying in brackish water, where most of the bloom is considered to be decomposed in the surface waters.     

The influence of a parasitoid's response to temperature on the performance of a tri‐trophic food web

1. Food web interactions are complex and can respond to environmental changes in unpredictable ways that do not necessarily equate to the individual responses of each of the components of the food web. 2. Biomass can be used to evaluate the productivity of the three individual trophic levels, in the form of the Net Generational Productivity (NGP) and the performance of the entire food web with the newly developed tri‐trophic food web performance ratio (?_3t). 3. These parameters were used to evaluate the performance of nine plant‐based tri‐trophic food webs composed of: potato, Solanum tuberosum L. and two cultivars of bell pepper Capsicum annuum L; three biotypes of the aphid Macrosiphum euphorbiae Thomas (Hemiptera: Aphididae) and the parasitoid wasp Aphidius ervi (Haliday) (Hymenoptera: Braconidae). 4. The NGP showed that the thermal window for biomass productivity for each trophic level is different and is reduced by approximately 4 °C with respect to the inferior level. Aphidius ervi had the smallest thermal window for biomass productivity and development. 5. The present results showed that the performance (?_3t) of the tested food webs is influenced in a top‐down fashion, where the intra‐specific variation of the food web, namely the host plant, played a major role in the productivity of each of the subsequent trophic levels. 6. The ?_3t suggested that exposure to high and low temperatures might severely affect the effectiveness of A. ervi as a biocontrol agent of the aphid M. euphorbiae in bell pepper and potato crops.     

The role of notothenioid fish in the food web of the Ross Sea shelf waters: a review

The Ross Sea, a large, high-latitude (72–78°S) embayment of the Antarctic continental shelf, averages 500 m deep, with troughs to 1,200 m and the shelf break at 700 m. It is covered by pack ice for 9 months of the year. The fish fauna of about 80 species includes primarily 4 families and 53 species of the endemic perciform suborder Notothenioidei. This review focuses on the diet and role in the food web of notothenioids and top-level bird and mammal predators, and also includes new information on the diets of artedidraconids and bathydraconids. Although principally a benthic group, notothenioids have diversified to form an adaptive radiation that includes pelagic and semipelagic species. In the southern Ross Sea, notothenioids dominate the fish fauna at levels of abundance and biomass >90% and are, therefore, inordinately important in the food web. Antarctic krill (Euphausia superba) and mesopelagic fishes are virtually absent from the shelf waters of the Ross Sea. Of the four notothenioid families, nototheniids show the most ecological and dietary diversification, with pelagic, cryopelagic, epibenthic and benthic species. Neutrally buoyant Pleuragramma antarcticum constitutes >90% of both the abundance and biomass of the midwater fish fauna. Most benthic nototheniids are opportunistic and feed on seasonally or locally abundant zooplanktonic prey. Artedidraconids are benthic sit-and-wait predators. Larger bathydraconids are benthic predators on fish while smaller species feed mainly on benthic crustaceans. Channichthyids are less dependent on the bottom for food than other notothenioids. Some species combine benthic and pelagic life styles; others are predominantly pelagic and all consume euphausiids and/or fish. South polar skuas, Antarctic petrels, Adélie and emperor penguins, Weddell seals and minke and killer whales are the higher vertebrate components of the food web, and all prey on notothenioids to some extent. Based on the frequency of occurrence of prey items in the stomachs of fish, bird and mammal predators, P. antarcticum and ice krill E. crystallorophias are the key species in the food web of the Ross Sea. P. antarcticum is a component of the diet of at least 11 species of nototheniid, bathydraconid and channichthyid fish and, at frequencies of occurrence from 71 to 100%, is especially important for Dissostichus mawsoni, Gvozdarus svetovidovi and some channichthyids. At least 16 species of notothenioids serve as prey for bird and mammal predators, but P. antarcticum is the most important and is a major component of the diet of south polar skua, Adélie and emperor penguins and Weddell seals, at frequencies of occurrence from 26 to 100%. E. crystallorophias is consumed by some nototheniid and channichthyid fish and can be of importance in the diet of emperor and Adélie penguins, although in the latter case, this is dependent on location and time of year.Unlike the linear phytoplanktonE. superbaconsumers of the E. superba food chain hypothesized for much of the Southern Ocean, the food web of the Ross Sea shelf is non-linear, with complex prey-predator interactions. Notothenioid fish play a key role: as predators, they occupy most of the trophic niches available in the ecosystem, relying on benthic, zooplanktonic and nektonic organisms; as prey, they are important food resources for each other and for most top predators living and foraging on the shelf. They also constitute the major link between lower (invertebrates) and higher (birds and mammals) levels of the food web. This is especially true for P. antarcticum. Along with E. crystallorophias, its ecological role in the Ross Sea is equivalent to that of myctophids and E. superba elsewhere in the Southern Ocean.