Distribution and co-occurrence patterns of charophytes and angiosperms in the northern Baltic Sea

The distribution data of 11 soft substrate charophyte and angiosperm species were analyzed. Our study aimed to elucidate the co-occurrence patterns among these sympatric macrophyte species and quantify their distribution areas. The central hypothesis of this study proposed that the observed co-occurrence patterns among the studied species deviate from what would be expected by random chance. Macrophyte occurrence data was derived from an extensive field sampling database. Environmental variables available as georeferenced raster layers including topographical, hydrodynamic, geological, physical, chemical, and biological variables were used as predictor variables in the random forest models to predict the spatial distribution of the species. Permutation tests revealed statistically significant deviations from random co-occurrence patterns. The analysis demonstrated that species tended to co-occur more frequently within their taxonomic groups (i.e., within charophytes and within angiosperms) than between these groups. The most extensive distribution overlap was observed between Chara aspera Willd. and Chara canescens Loisel., while Zostera marina L. exhibited the least overlap with the other species. The mean number of co-occurring species was the highest in Chara baltica (Hartman) Bruzelius while Z. marina had the largest share of single-species occurrences. Based on the distribution models, Stuckenia pectinata (L.) Börner had the largest distribution area.