Fungal interactions with vascular and non-vascular plants: an investigation of mutualisms and their roles in heathland regeneration.

Mycorrhizal mutualisms between aboveground vascular plant communities, which reward their belowground fungal associates with photosynthates in return for growth-limiting nutrients such as phosphate, are widely recognized as stable long-term interactions which helped plants colonize land. Pezoloma ericae (D.J. Read) Baral, an ascomycete mycorrhiza-forming fungus present amongst plants in the Ericales, such as heathers, also forms associations in several families of non-vascular leafy liverworts. Whether there is a mutually beneficial functional relationship between these leafy liverworts and the fungus growing in their rhizoids was previously unconfirmed. Furthermore, an ecological role of this ‘shared’ mycobiont and its link between vascular (Ericaceae) and non-vascular (liverworts) plants was also unknown. Thus the main questions asked in this dissertation are: 1) Is there a measurable mutually beneficial relationship between a liverwort and its fungal partner?; and, 2) Can liverworts harbouring the ericoid mycorrhiza P. ericae act as inoculum that facilitates the re-establishment of Ericaceae - and henceforth be proposed as a practical tool in a restoration ecology context. This is the first time British species of leafy liverworts are conclusively identified to harbour the ericoid mycorrhizal fungus Pezoloma ericae using molecular identification. I have demonstrated a mutualism occurring between the leafy liverworts and their fungal symbiont in two independent microcosm growth experiments and confirmatory reciprocal trophic exchanges between phosphorus and carbon and the two organisms. Glasshouse experiments demonstrated P. ericae originating from leafy liverwort rhizoids, can repeatedly colonize Ericaceae plant roots. Under realistic ecological circumstances (further tested at two field sites), liverworts delivered mycorrhizal inoculum and improved the resilience and growth of vascular plants. By providing this novel source of mycorrhizal inoculum, symbiotic non-vascular plants can contribute to the restoration of plant communities dominated by Ericaceous plants. This research leads to broader knowledge about the function of ericoid mycorrhizas in ecosystems with multi-trophic non-vascular – fungi - vascular community interactions, both above- and belowground.