Earthworm invasion alters enchytraeid community composition and individual biomass in northern hardwood forests of North America

Abstract

European earthworms are invading many ecosystems worldwide and fundamentally transform habitats by acting as dominant ecosystem engineers. However, there is little knowledge of the consequences of earthworm invasion on the composition and diversity of native soil organisms. Particularly functionally similar groups, such as enchytraeids (Annelida: Enchytraeidae), may be affected through changes in the chemical and physical properties of the soil, but also due to competition for resources. In 2010–2011, we studied the impact of earthworm invasion on enchytraeids at two sites in the northern hardwood forests of North America: one site within the Chippewa National Forest in northern Minnesota and one site in the Chequamegon-Nicolet National Forest in northern Wisconsin, USA. At each site, three plots were sampled along a transect, representing (1) a non-invaded or very slightly invaded area, (2) the leading edge of earthworm invasion and (3) a heavily invaded area with an established population of the anecic earthworm Lumbricus terrestris (among other species). In total, 29 enchytraeid (morpho)species were identified (some yet to be formally described, several first or second records for the continent); of those 24 occurred at the Minnesota site and 17 at the Wisconsin site. The structure of enchytraeid assemblages differed significantly among the three invasion stages, although this was not equally pronounced at the two sites. Each stage was characterized by one or several indicator species. Mean enchytraeid densities (10,700–30,400individuals/m2) did not differ significantly among the invasion stages, but were lowest at the leading edge of earthworm invasion at both sites. In the heavily invaded plot at the Minnesota site, the mean enchytraeid density and biomass in L. terrestris middens were significantly higher than in soil in-between the middens. This was due to a pronounced effect of L. terrestris middens in the uppermost 3cm of soil. Differences in biomass among earthworm invasion stages were most apparent for mean individual biomass. This was significantly higher in the heavily invaded area than at the leading edge or in the non-invaded area at the Minnesota site. Compositional changes of the enchytraeid assemblage are likely to result in changes in the functioning of soil foods webs. Our results suggest that earthworm invasions can cause a loss of native species in soil, including heretofore unknown ones, that might go unnoticed.