Earthworms increase the ratio of bacteria to fungi in northern hardwood forest soils, primarily by eliminating the organic horizon

The exotic earthworm invasion in hardwood forests of the northern United States is associated with many ecosystem-level changes. However, less is known about the effects of the invasion on the composition of the soil microbial community through which ecosystem-level changes are mediated. Further, earthworm effects on soil microbial community composition have not been well studied in the field. To evaluate changes in bacterial and fungal abundance associated with the earthworm invasion we quantified bacterial and fungal biomass by microscopic counts in paired earthworm-invaded (earthworm) and earthworm-free (reference) plots in five forest stands in central New York (USA). Earthworms significantly increased the ratio of bacteria to fungi on an area basis (per m2), by more than two times in mid-summer and early autumn. While this effect was associated primarily with the lack of the fungal-dominated organic horizon in earthworm plots, a higher ratio of bacteria to fungi in the surface 5 cm mineral soil also contributed as it developed between spring and mid-summer. Earthworm reduction of fungal biomass was confirmed by substantially lower growth of fungal hyphae into mesh sand bags in earthworm compared to reference plots. Burrowing activity by the earthworm Lumbricus terrestris increased the ratio of bacteria to fungi over the short-term within earthworm plots, introducing small-scale spatial heterogeneity associated with burrows. Our study suggests that the exotic earthworm invasion in these northern hardwood forests markedly increased the ratio of bacteria to fungi by eliminating the fungal-rich organic horizon, and was associated localized increases in bacterial vs. fungal abundance in mineral soil, setting the stage for future research into linkages between the earthworm invasion, bacterial and fungal abundance, and ecosystem processes.

► Earthworms increased bacteria: fungi in northern hardwood forest soil. ► Growth of hyphae into mesh bags was reduced by earthworms. ► Effects at a landscape scale were related mostly to elimination of the organic horizon. ► Bacteria:fungi in mineral soil showed within-season and burrow-associated effects. ► Earthworms likely affect nutrient processing by changing fungal translocation.