The second wave of earthworm invasion: soil organic matter dynamics from the stable isotope perspective

Through transformation of plant litter into soil organic matter (SOM) and translocation of ingested organic material among different soil depths, soil organisms, especially earthworms, are one of the major factors affecting SOM dynamics. In North America temperate soil, historical human activity has lead to invasion of European earthworms into habitats that were previously earthworm-free or inhabited only by native species. By consuming leaf litter and SOM, burrowing, and casting, invasive earthworms have been known for reducing the understory vegetation and leaf litter layer while increasing the thickness of organic soil, causing changes in the soil habitat and the distribution of SOM. Recently, another group of invasive earthworm, namely Amynthas from Asia, has been reported invading habitats already dominated by European species, causing a 'second wave of invasion' where the soil ecosystem, already modified by European species, is going through another transition. The mechanisms through which these functionally (ecologically) different species affect C and N transformation could be better understood by tracing the carbon and nitrogen derived from 13C- and 15N-labeled leaf litter into earthworm tissues and SOM. The objective of this study is to understand how earthworm species that differ ecologically, including the Asian Amynthas, interact with each other and how these interactions affect SOM dynamics. We hypothesized that 1) species feeding on different food resources will have different isotopic signature and their tissue 13C and 15N values will change due to facilitation or interspecific competition on food resources, and 2) the short-term fate of litter-derived carbon differs depending on the presence or absence of different earthworm species. These hypotheses were tested by field sampling and lab mesocosm experiments using 13C and 15N double-enriched Tulip Poplar leaf litter (mean 13C = 124‰, mean 15N = 1667‰) produced from tree saplings growing in an airtight chamber. Stable isotope mass balance calculation is used to estimate the recovery of litter-derived carbon from three pools (earthworm tissue, SOM, remaining litter), the loss of litter-derived carbon through soil respiration, and the contribution of different carbon sources to soil CO2 efflux in different earthworm treatments. Our results show that earthworm species recognized as 'soil feeders' have 13C and 15N values that are 1.2‰ and 3.8‰ higher than those of 'litter feeders', and 15N also differ significantly amount different soil feeders, suggesting different food resource usage even within the same functional group. There are strong species effects on both leaf litter disappearance rate and CO2 efflux rate, both being high when Amynthas earthworms are present. Our results suggest that changing earthworm species composition leads to changing resource use, which alters the fate of organic carbon in the forest floor and soil and could potentially affect long-term SOM dynamics in temperate forests.