Earthworms are not just "earth" worms: Multiple drivers to large diversity in deadwood

Earthworms are ecosystem engineers associated with important soil functions. Despite the large amount of literature on earthworm ecology, relatively few studies have examined earthworms in deadwood or quantified their importance in this habitat. We investigated earthworm communities in decaying deadwood and disen-tangled how their community dynamics are influenced by variation in tree species, wood decomposition stage, and forest environment. Decaying logs (of standardised size) of ten common, temperate tree species were laid out to decay on the soil surface for four years. The experiment was carried out in the "tree cemetery" experiment LOGLIFE, in two contrasting temperate forests in the central Netherlands. The decaying logs yielded surprisingly rich earthworm populations, with on average 19 individuals per meter of log and in total belonging to 12 different species. Our findings highlighted that earthworm communities in deadwood in terms of composition and abundance were influenced by tree species, wood decomposition stage and forest type with different soil properties, as well as their interactions. After one and two years of decay, earthworm abundance was higher in the logs of Picea abies and relatively fast decomposing Populus spp. than in other trees, while this pattern changed after four years with higher earthworm abundance observed in the other tree species. Overall, Populus spp. supported the highest earthworm abundance, followed by Picea abies and Quercus robur. The earthworm com-munity composition in the logs in the two forest sites had broadly similar dynamic trends of first becoming very dissimilar between one and two years, then relatively more similar from two to four years of decay, although the community composition differed between sites. The interacting influences of tree species, wood decay stage and forest environment on earthworm communities strongly suggest that diversity in deadwood resources contributes to earthworm diversity in forests. Thus, by mixing different tree species and logging gradually through the years, forest managers could enhance the diversity of this abundant and understudied component of deadwood invertebrate diversity. Further research should study the feedback loop between earthworm communities and decomposition of deadwood.