Pine-fungal co-invasion alters whole-ecosystem properties of a native eucalypt forest

Pine-fungal co-invasions into native ecosystems are increasingly prevalent across the southern hemisphere. In Australia, invasive pines slowly spread into native eucalypt forests, creating novel mixed forests. We sought to understand how pine-fungal co-invasions impact interconnected above- and belowground ecosystem characteristics. We sampled beneath mature Pinus radiata and Eucalyptus racemosa in a pine-invaded eucalypt forest in New South Wales, Australia. We measured microbial community composition via amplicon sequencing of 16S, ITS2, and 18S rDNA regions, microbial metabolic activity via Biolog plate substrate utilization, and soil, leaf litter, and understory plant characteristics. Pines were associated with decreased topsoil moisture, increased pine litter, and decreased eucalypt litter total phosphorus content. Soils and roots beneath pines had distinct microbial community composition and activity relative to eucalypts, including decreased bacterial diversity, decreased microbial utilization of several C- and N-rich substrates, and enrichment of pine-associated ectomycorrhizae. Introduced suilloid fungi were abundant across both pine and eucalypt soils and roots. Many ecosystem impacts increased with pine size. Invasive pines and their ectomycorrhizae have significant impacts on eucalypt forest properties as they grow. Interconnected impacts at the scale of individual trees should be considered when managing invaded forests and predicting effects of pine invasions.