Fourth-corner analysis with 5,000 permutations demonstrating pairwise correlations between traits, environmental variables and fishing pressure. The p-values have been corrected for multiple comparison using false discovery rates. Cells in red indicate positive correlations and cells in blue negative correlations. The significance level is given by asterisks (‘•’: p-value < 0.1, ‘*’: < 0.05, ‘**’: < 0.01). Non-significant correlations are in light grey. SBT: sea bottom temperature, SBS: sea bottom salinity, Chl: chlorophyll a concentration, SBT.var: seasonality in sea bottom temperature, Chl.var: seasonality in chlorophyll a concentration.