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Marine communities undergo rapid changes related to human-induced ecosystem pressures. The Baltic Sea pelagic food web has experienced several regime shifts during the past century, resulting in a system where competition between the dominant planktivorous mesopredatory clupeid fish species herring ( Clupea harengus ) and sprat ( Sprattus sprattus ) and the rapidly increasing stickleback ( Gasterosteus aculeatus ) population is assumed to be high. Here, we investigate diet overlap between these three planktivorous fishes in the Baltic Sea, utilizing DNA metabarcoding on the 18S rRNA gene and the COI gene, targeted qPCR, and microscopy. Our results show niche differentiation between clupeids and stickleback, and highlight that rotifers play an important role in this pattern, as a resource that is not being used by the clupeids nor by other zooplankton in spring. We further show that all the diet assessment methods used in this study are consistent, but also that DNA metabarcoding describes the plankton-fish link at the highest taxonomic resolution. This study suggests that rotifers and other understudied soft-bodied prey may have an important function in the pelagic food web and that the growing population of pelagic stickleback may be supported by the open feeding niche offered by the rotifers.

Coexistence of fish populations (= stocks) of the same species is a common phenomenon. In the Baltic Sea, two genetically divergent stocks of Atlantic cod ( Gadus morhua ), Western Baltic cod (WBC) and Eastern Baltic cod (EBC), coexist in the Arkona Sea. Although the relative proportions of WBC and EBC in this area are considered in the current stock assessments, the mixing dynamics and ecological mechanisms underlying coexistence are not well understood. In this study, a genetically validated otolith shape analysis was used to develop the most comprehensive time series of annual stock mixing data (1977–2019) for WBC and EBC. Spatio-temporal mixing analysis confirmed that the two stocks coexist in the Arkona Sea, albeit with fluctuating mixing proportions over the 43-year observation period. Depth-stratified analysis revealed a strong correlation between capture depth and stock mixing patterns, with high proportions of WBC in shallower waters (48–61% in <20m) and increasing proportions of EBC in deeper waters (50–86% in 40-70m). Consistent depth-specific mixing patterns indicate stable differences in depth distribution and habitat use of WBC and EBC that may thus underlie the long-term coexistence of the two stocks in the Arkona Sea. These differences were also reflected in significantly different proportions of WBC and EBC in fisheries applying passive gears in shallower waters (more WBC) and active gears in deeper waters (more EBC). This highlights the potential for fishing gear-specific exploitation of different stocks, and calls for stronger consideration of capture depth and gear type in stock assessments. This novel evidence provides the basis for improved approaches to research, monitoring and management of Baltic cod stocks.

Marine food webs are highly compartmentalized, and characterizing the trophic niches among consumers is important for predicting how impact from human activities affects the structuring and functioning of marine food webs. Biomarkers such as bulk stable isotopes have proven to be powerful tools to elucidate trophic niches, but they may lack in resolution, particularly when spatiotemporal variability in a system is high. To close this gap, we investigated whether carbon isotope (δ13C) patterns of essential amino acids (EAAs), also termed δ13CAA fingerprints, can characterize niche differentiation in a highly dynamic marine system. Specifically, we tested the ability of δ13CAA fingerprints to differentiate trophic niches among six functional groups and ten individual species in the Baltic Sea. We also tested whether fingerprints of the common zooplanktivorous fishes, herring and sprat, differ among four Baltic Sea regions with different biochemical conditions and phytoplankton assemblages. Additionally, we investigated how these results compared to bulk C and N isotope data for the same sample set. We found significantly different δ13CAA fingerprints among all six functional groups. Species differentiation was in comparison less distinct, due to partial convergence of the species' fingerprints within functional groups. Herring and sprat displayed region‐specific δ13CAA fingerprints indicating that this approach could be used as a migratory marker. Niche metrics analyses showed that bulk isotope data had a lower power to differentiate between trophic niches than δ13CAA fingerprinting. We conclude that δ13CAA fingerprinting has a strong potential to advance our understanding of ecological niches, and trophic linkages from producers to higher trophic levels in dynamic marine systems. Given how management practices of marine resources and habitats are reshaping the structure and function of marine food webs, implementing new and powerful tracer methods are urgently needed to improve the knowledge base for policy makers. Our study uses stable isotope fingerprinting of amino acid to characterize resource utilization and niche segregation among consumers in the Baltic Sea. We demonstrate that the isotope fingerprinting method (a) can resolve dietary niches at a much higher resolution than bulk stable isotope analysis (b). This methodological advance can improve our understanding of how management practices affect marine ecosystem functioning.

The trophic mode is one of the most important functional traits of organisms, determining their position in the food web and their role in the ecosystem. Under the classical concept, phytoplankton was considered to consist exclusively of phototrophic and microzooplankton exclusively of heterotrophic organisms. However, it is now increasingly recognized that mixotrophy (e.g. combining photo- and phagotrophy) occurs among both photo- and microzooplankton species, and that in extension, trophic diversity and relationships among plankton are expected to be more extensive and complex than previously thought. To enhance understanding of plankton trophic modes, diversity and relationships during spring bloom period in Bornholm Basin, central Baltic Sea, we categorized here the temporal succession in (1) the aquatic protist community by trophic modes, and (2) the community composition in terms of taxonomic groups and feeding mode, and its correlation with environmental factors and available prey-size. Our results show that the trophic mode composition of the community changed drastically over the course of the spring season, representing a high trophic complexity and more complex dynamics than previously suggested. The heterotrophic community was characterized by a high diversity of species and groups, with heterotrophic ciliates showing a clear seasonal succession in body size-classes, switching from the smallest sized-fraction (10-20 µm) in winter to an increasingly amount of larger-sized species of 30-55 µm and >55 µm with progression of the spring period. Changes in ciliate community composition were correlated with sea surface temperature, shifting from a cold-associated to a warm-associated community over the course of the spring season. Results further suggest that in communities including a larger mixotrophic component, size trait-based trophic relations between heterotrophic groups and their prey are complex, potentially due to similar prey-size preferences among heterotrophic and mixotrophic species. Overall, our findings emphasize the importance of accounting for the trophic modes of species to enhance the understanding of trophic relations and dynamics within bloom events.

Assessments of ecosystem functioning are a fundamental ecological challenge and an essential foundation for ecosystem‐based management. Species trophic position (TP) is essential to characterize food web architecture. However, despite the intuitive nature of the concept, empirically estimating TP is a challenging task due to the complexity of trophic interaction networks. Various methods are proposed to assess TPs, including using different sources of organic matter at the base of the food web (the ‘baseline’). However, it is often not clear which methodological approach and which baseline choices are the most reliable. Using an ecosystem‐wide assessment of a tropical reef (Marquesas Islands, with available data for 70 coral reef invertebrate and fish species), we tested whether different commonly used TP estimation methods yield similar results and, if not, whether it is possible to identify the most reliable method. We found significant differences in TP estimates of up to 1.7 TPs for the same species, depending on the method and the baseline used. When using bulk stable isotope data, the choice of the baseline significantly impacted TP values. Indeed, while nitrogen stable isotope (δ15N) values of macroalgae led to consistent TP estimates, those using phytoplankton generated unrealistically low TP estimates. The use of a conventional enrichment factor (i.e. 3.4‰) or a ‘variable’ enrichment factor (i.e. according to feeding guilds) also produced clear discrepancies between TP estimates. TPs obtained with δ15N values of source amino acids (compound‐specific isotope analysis) were close to those assessed with macroalgae. An opposite seasonal pattern was found, with significantly lower TPs in winter than in summer for most species, with particularly pronounced differences for lower TP species. We use the observed differences to discuss possible drivers of the diverging TP estimates and the potential ecological implications. Based on a case study from a remote coral reef ecosystem in the Pacific Ocean, this work analyses how several potential sources of organic matter (i.e. ‘baselines’) and calculation methods strongly affect assessments of trophic positions. We also demonstrated that seasonal temporality and feeding guild are important parameters to be considered.

Atlantic cod (Gadus morhua) is a species of great ecological and economical importance in the Baltic Sea. Here, two genetically differentiated stocks, the western and the eastern Baltic cod, display substantial mechanical mixing, hampering our understanding of cod ecology and impeding stock assessments and management. Based on whole-genome re-sequencing data from reference samples obtained from the study area, we designed two different panels of Single Nucleotide Polymorphisms markers (SNPs), which take into account the exceptional genome architecture of cod. A minimum panel of 20 diagnostic SNPs and an extended panel (20 diagnostic and 18 biologically informative SNPs, 38 in total) were developed and validated to distinguish unambiguously between the western and the eastern Baltic cod stocks and to enable studies of local adaptation to the specific environment in the Baltic Sea, respectively. We tested both panels on cod sampled from the southern Baltic Sea (n = 603) caught in 2015 and 2016. Genotyping results showed that catches from the mixing zone in the Arkona Sea, were composed of similar proportions of individuals of the western and the eastern stock. Catches from adjacent areas to the east, the Bornholm Basin and Gdańsk Deep, were exclusively composed of eastern Baltic cod, whereas catches from adjacent western areas (Belt Sea and Öresund) were composed of western Baltic cod. Interestingly, the two Baltic cod stocks showed strong genetic differences at loci associated with life-history trait candidate genes, highlighting the species' potential for ecological adaptation even at small geographical scales. The minimum and the extended panel of SNP markers presented in this study provide powerful tools for future applications in research and fisheries management to further illuminate the mixing dynamics of cod in the Baltic Sea and to better understand Baltic cod ecology.

We studied the food web structure and functioning of a coral reef ecosystem in the Marquesas Islands, French Polynesia, characterized by low coral cover, high sea surface temperature and meso- to eutrophic waters. The Marquesas constitute a relevant ecosystem to understand the functioning of low diversity reefs that are also subject to global change. A multi-tracer assessment of organic matter pathways was run to delineate ecosystem functioning, using analysis of fatty acids, bulk and compound specific stable isotope analysis and stable isotopes mixing models. Macroalgae and phytoplankton were the two major food sources fueling this food web with, however, some marked seasonal variations. Specifically, zooplankton relied on phytoplankton-derived organic matter and herbivorous fishes on macroalgae-derived organic matter to a much higher extent in summer than in winter (~ 75% vs.  ~ 15%, and ~ 70 to 75% vs.  ~ 5 to 15%, respectively) . Despite remarkably high δ 15 N values for all trophic compartments, likely due to local dynamics in the nitrogen stock, trophic levels of consumers were similar to those of other coral reef ecosystems. These findings shed light on the functioning of low coral cover systems, which are expected to expand worldwide under global change.

Genetic data have great potential for improving fisheries management by identifying the fundamental management units—that is, the biological populations—and their mixing. However, so far, the number of practical cases of marine fisheries management using genetics has been limited. Here, we used Atlantic cod in the Baltic Sea to demonstrate the applicability of genetics to a complex management scenario involving mixing of two genetically divergent populations. Specifically, we addressed several assumptions used in the current assessment of the two populations. Through analysis of 483 single nucleotide polymorphisms (SNPs) distributed across the Atlantic cod genome, we confirmed that a model of mechanical mixing, rather than hybridization and introgression, best explained the pattern of genetic differentiation. Thus, the fishery is best monitored as a mixed‐stock fishery. Next, we developed a targeted panel of 39 SNPs with high statistical power for identifying population of origin and analyzed more than 2,000 tissue samples collected between 2011 and 2015 as well as 260 otoliths collected in 2003/2004. These data provided high spatial resolution and allowed us to investigate geographical trends in mixing, to compare patterns for different life stages and to investigate temporal trends in mixing. We found similar geographical trends for the two time points represented by tissue and otolith samples and that a recently implemented geographical management separation of the two populations provided a relatively close match to their distributions. In contrast to the current assumption, we found that patterns of mixing differed between juveniles and adults, a signal likely linked to the different reproductive dynamics of the two populations. Collectively, our data confirm that genetics is an operational tool for complex fisheries management applications. We recommend focussing on developing population assessment models and fisheries management frameworks to capitalize fully on the additional information offered by genetically assisted fisheries monitoring.

Range expansions can lead to increased contact of divergent populations, thus increasing the potential of hybridization events. Whether viable hybrids are produced will most likely depend on the level of genomic divergence and associated genomic incompatibilities between the different entities as well as environmental conditions. By taking advantage of historical Baltic cod (Gadus morhua) otolith samples combined with genotyping and whole genome sequencing, we here investigate the genetic impact of the increased spawning stock biomass of the eastern Baltic cod stock in the mid 1980s. The eastern Baltic cod is genetically highly differentiated from the adjacent western Baltic cod and locally adapted to the brackish environmental conditions in the deeper Eastern basins of the Baltic Sea unsuitable for its marine counterparts. Our genotyping results show an increased proportion of eastern Baltic cod in western Baltic areas (Mecklenburg Bay and Arkona Basin)—indicative of a range expansion westwards—during the peak population abundance in the 1980s. Additionally, we detect high frequencies of potential hybrids (including F1, F2 and backcrosses), verified by whole genome sequencing data for a subset of individuals. Analysis of mitochondrial genomes further indicates directional gene flow from eastern Baltic cod males to western Baltic cod females. Our findings unravel that increased overlap in distribution can promote hybridization between highly divergent populations and that the hybrids can be viable and survive under specific and favourable environmental conditions. However, the observed hybridization had seemingly no long‐lasting impact on the continuous separation and genetic differentiation between the unique Baltic cod stocks.