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Abstract Anthropogenic deoxygenation of the Baltic Sea caused major declines in demersal and benthic habitat quality with consequent impacts on biodiversity and ecosystem services. Using Baltic cod otolith chemical proxies of hypoxia, salinity, and fish metabolic status and growth, we tracked changes from baseline conditions in the late Neolithic (4500 BP) and early twentieth century to the present, in order to understand how recent, accelerating climate change has affected this key species. Otolith hypoxia proxies (Mn:Mg) increased with expanding anoxic water volumes, but decreased with increasing salinity indexed by otolith Sr:Ca. Metabolic status proxied by otolith Mg:Ca and reconstructed growth were positively related to dissolved oxygen percent saturation, with particularly severe declines since 2010. This long-term record of otolith indicators provides further evidence of a profound state change in oxygen for the worse, in one of the world’s largest inland seas. Spreading hypoxia due to climate warming will likely impair fish populations globally and evidence can be tracked with otolith chemical biomarkers. 

Chronological records of elemental concentrations in fish otoliths are a widely used tool to infer the environmental conditions experienced by individual fish. To interpret elemental signals within the otolith, it is important to understand how both external and internal factors impact ion uptake, transport and incorporation. In this study, we have combined chronological records from otoliths and archival data storage tags to quantify the influence of internal (sex, size, age, growth) and external (temperature, depth, salinity) conditions on otolith elemental chemistry of cod (Gadus morhua) in natural settings of the Baltic Sea. This study focused on elements primarily under physiological control: Phosphorus (P), magnesium (Mg) and zinc (Zn); and elements under environmental control: Strontium (Sr), barium (Ba) and manganese (Mn). Based on known spatial and temporal patterns in environmental conditions and fish size, growth, and maturity, we posed a series of hypotheses of expected otolith element patterns. Partial effects of internal and external drivers on element concentration were analyzed using a Linear Mixed Model approach with random variables (fish and year). Predicted effects of otolith concentrations of all elements under physiological control (P, Mg, Zn) showed similar trends, with distinct seasonal patterns (lowest concentration in late spring, highest concentrations in winter), and a positive correlation with water temperature, in addition to higher Zn and lower P in spawning individuals. Predicted effects of otolith concentrations of elements expected to be predominantly under environmental control showed the predicted geographic and depth-related trends based on ambient salinity (Ba) and coastal hypoxia (Mn). However, contrary to expectation, Sr was unrelated to salinity. Predicted otolith Ba, Sr and Mn concentrations also exhibited pronounced seasonal patterns that were out of phase with each other but appeared to be partly explained by spawning/feeding migrations. While performing laboratory validation studies for adult fish is typically not possible, these results highlight the importance of assessing local water chemistry and freshwater endmembers in one’s study system before otolith elemental chemistry can be reliably used to reconstruct fish habitat use and environmental histories. 

Abstract. During the past 20 years, hypoxic areas have expanded rapidly in theBaltic Sea, which has become one of the largest marine “dead zones” in theworld. At the same time, the most important commercial fish population ofthe region, the eastern Baltic cod, has experienced a drastic reduction inmean body condition, but the processes behind the relation betweendeoxygenation and condition remain elusive. Here we use extensive long-termmonitoring data on cod biology and distribution as well as on hydrologicalvariations to investigate the processes that relate deoxygenation and codcondition during the autumn season. Our results show that the depthdistribution of cod has increased during the past 4 decades at the sametime of the expansion, and shallowing, of waters with oxygen concentrationsdetrimental to cod performance. This has resulted in a progressivelyincreasing spatial overlap between the cod population and low-oxygenatedwaters after the mid-1990s. This spatial overlap and the actual oxygenconcentration experienced by cod therein statistically explained a largeproportion of the changes in cod condition over the years. These resultscomplement previous analyses on fish otolith microchemistry that alsorevealed that since the mid-1990s, cod individuals with low condition wereexposed to low-oxygen waters during their life. This study helps to shedlight on the processes that have led to a decline of the eastern Baltic codbody condition, which can aid the management of this population currently indistress. Further studies should focus on understanding why the codpopulation has moved to deeper waters in autumn and on analyzing the overlapwith low-oxygen waters in other seasons to quantify the potential effects ofthe variations in physical properties on cod biology throughout the year. 

Deoxygenation worldwide is increasing in aquatic systems with implications for organisms' biology, communities and ecosystems. Eastern Baltic cod has experienced a strong decline in mean body condition (i.e. weight at a specific length) over the past 20 years with effects on the fishery relying on this resource. The decrease in cod condition has been tentatively linked in the literature to increased hypoxic areas potentially affecting habitat range, but also to benthic prey and/or cod physiology directly. To date, no studies have been performed to test these mechanisms. Using otolith trace element microchemistry and hypoxia-responding metrics based on manganese (Mn) and magnesium (Mg), we investigated the relationship between fish body condition at capture and exposure to hypoxia. Cod individuals collected after 2000 with low body condition had a higher level of Mn/Mg in the last year of life, indicating higher exposure to hypoxic waters than cod with high body condition. Moreover, lifetime exposure to hypoxia was even more strongly correlated to body condition, suggesting that condition may reflect long-term hypoxia status. These results were irrespective of fish age or sex. This implies that as Baltic cod visit poor-oxygen waters, perhaps searching for benthic food, they compromise their own performance. This study specifically sheds light on the mechanisms leading to the low condition of cod and generally points to the impact of deoxygenation on ecosystems and fisheries. 

Accurate age data are essential for reliable fish stock assessment. Yet many stocks suffer from inconsistencies in age interpretation. A new approach to obtain age makes use of the chemical composition of otoliths. This study validates the periodicity of recurrent patterns in 25 Mg, 31 P, 34 K, 55 Mn, 63 Cu, 64 Zn, 66 Zn, 85 Rb, 88 Sr, 138 Ba, and 208 Pb in Baltic cod (Gadus morhua) otoliths from tag–recapture and known-age samples. Otolith P concentrations showed the highest consistency in seasonality over the years, with minima co-occurring with otolith winter zones in the known-age otoliths and in late winter – early spring when water temperatures are coldest in tagged cod . The timing of minima differs between stocks, occurring around February in western Baltic cod and 1 month later in eastern Baltic cod; seasonal maxima are also stock-specific, occurring in August and October, respectively. The amplitude in P is larger in faster-growing western compared with eastern Baltic cod. Seasonal patterns with minima in winter – late spring were also evident in Mg and Mn, but less consistent over time and fish size than P. Chronological patterns in P, and to a lesser extent Mg and Mn, may have the potential to supplement traditional age estimation or to guide the visual identification of translucent and opaque otolith patterns used in traditional age estimation. 

Abstract Fish otoliths' chronometric properties make them useful for age and growth rate estimation in fisheries management. For the Eastern Baltic Sea cod stock (Gadus morhua), unclear seasonal growth zones in otoliths have resulted in unreliable age and growth information. Here, a new age estimation method based on seasonal patterns in trace elemental otolith incorporation was tested for the first time and compared with the traditional method of visually counting growth zones, using otoliths from the Baltic and North seas. Various trace elemental ratios, linked to fish metabolic activity (higher in summer) or external environment (migration to colder, deeper habitats with higher salinity in winter), were tested for age estimation based on assessing their seasonal variations in concentration. Mg:Ca and P:Ca, both proxies for growth and metabolic activity, showed greatest seasonality and therefore have the best potential to be used as chemical clocks. Otolith image readability was significantly lower in the Baltic than in the North Sea. The chemical (novel) method had an overall greater precision and percentage agreement among readers (11.2%, 74.0%) than the visual (traditional) method (23.1%, 51.0%). Visual readers generally selected more highly contrasting zones as annuli whereas the chemical readers identified brighter regions within the first two annuli and darker zones thereafter. Visual estimates produced significantly higher, more variable ages than did the chemical ones. Based on the analyses in our study, we suggest that otolith microchemistry is a promising alternative ageing method for fish populations difficult to age, such as the Eastern Baltic cod.