Bacterial communities in an optional lichen symbiosis are determined by substrate, not algal photobionts

Borderline lichens are simple mutualistic symbioses between fungi and algae, where the fungi form loose mycelia interweaving algal cells, instead of forming a lichen thallus. Schizoxylon albescens shows two nutritional modes: it can either live as a borderline lichen on Populus tremula bark or as a saprotroph on Populus wood. This enables us to investigate the microbiota diversity in simple fungal–algal associations and to study the impact of lichenization on the structure of bacterial communities. We sampled three areas in Sweden covering the distribution of Schizoxylon, and using high-throughput sequencing of the 16S rRNA gene and fluorescence in situ hybridization we characterized the associated microbiota. Bacterial communities in lichenized and saprotrophic Schizoxylon were clearly distinct, but when comparing the microbiota with the respective substrates, only the fruiting bodies show clear differences in composition and abundance from the communities in the substrates. The colonization by either lichenized or saprotrophic mycelia of Schizoxylon did not significantly influence the microbiota in the substrate. This suggests that in a morphologically simple form of lichenization, as represented by the Schizoxylon–Coccomyxa system, algal–fungal interactions do not significantly influence bacterial communities, but a more complex structure of the lichen thallus is likely required for hosting specific microbiota.