Species richness correlations among primary producers in boreal forests

Close correlations in species numbers may make it possible to indirectly assess the species richness of difficult taxonomic groups by investigating indicator groups, for which data are more easily collected. We asked if species numbers correlate among the four dominating groups of primary producers in boreal forests (liverworts, macrolichens, mosses, and vascular plants) and if substrate affiliation of species (ground vs. other substrates), sample plot size (0.01-1000 m(2)), and stand age (young vs. old) influence correlation strength. We used three sets of study plots from northern Sweden each including wide ranges of species richness. Although there are large differences in the ecophysiology and substrate use of vascular plants and the two bryophyte groups (mosses and liverworts), we found strong positive correlations among them not previously reported from boreal forests. In contrast, no correlation in total species richness was found between macrolichens and the two bryophyte groups, despite large overlaps in their ecology. We suggest that the positive correlations among land plants (liverworts, mosses, and vascular plants) are linked to positive relationships between site moisture and species number for all three groups. In contrast, total species number of macrolichens has not been shown to be strongly associated with moisture. However, ground-living lichens and mosses correlated negatively in old forests. This may relate to the inability of macrolichens to exploit shaded and wet old forest ground, a habitat that is used by many moss species. Furthermore, lichens and mosses of 'other substrates' correlated positively in old forests, probably because the amount of boulders was positively related to species richness in both groups. Generally, correlations became stronger with increasing plot size, whereas stand age had relatively little influence. We conclude that vascular plants could be used as an indicator group for species richness of mosses and liverworts in boreal landscapes.