Stratification-induced hypoxia as a structuring factor of macrozoobenthos in the open Gulf of Finland (Baltic Sea)

Long-term (1965–2000) changes of macrozoobenthos and hydrography have been studied in the Gulf of Finland (GoF). For the first time, statistical multivariate time series analysis is applied to Baltic Sea data to verify the relationship between biota and interacting environmental factors causing large-scale hypoxia in the open sea. For macrozoobenthos, a consistent long-term development of the assemblages was found over the study area. In the period before the 1990s, very sparse macrozoobenthos prevailed, followed by a notable expansion of macrofauna between the late 1980s and early 1990s and leading to a maximum of total abundance and species number between 1991 and 1996. After that, a sudden collapse of the communities took place in 1996–1997. The hydrographical changes included a continuous decrease in salinity and density stratification until the early 1990s, after which an increase took place again. In contrast, low mean and minimum dissolved oxygen concentrations were observed at the beginning of the study period, followed by increasing values in the late 1980s and early 1990s, and a simultaneous decline of oxygen conditions in 1996. Based on non-linear trends estimated by dynamic factor analysis (DFA), high and significant correlations were found between total macrofauna abundance, number of species, salinity, oxygen conditions, strength of stratification and freshwater run-off. The results confirm that oxygen is obviously a fundamental factor that determines the state of the macrozoobenthos in the deep GoF, overruling other abiotic factors. However, the improvement of the oxygen conditions is apparently caused by the long-term decrease of salinity and loss of stratification in the relatively shallow GoF, reflecting large-scale changes in hydrography of the Baltic Sea during the long 1977–1993 stagnation period. Thus the development in GoF is opposite to the deeper basins in the central Baltic. We conclude that salinity and stratification are probably linked with climatic variability via freshwater run-off, which may be important in regulating the oxygen conditions and state of macrozoobenthos in GoF.