Earthworms as colonizers of natural and cultivated soil environments

For cultivated soils, the important function of earthworms as ecosystem engineers and their major contribution to the composition and functioning of soil ecosystems with a varying species diversity has been extensively addressed. However, the role of earthworms as colonizers of virgin, uncultivated soil in the process of soil formation has been little researched and long underrated. To better understand this role, the following questions need to be considered: (1) what makes an early colonizer successful, what are its characteristics, and which species are the most successful and under what circumstances are they successful?; (2) what are the limiting factors in these colonization processes with respect to environmental conditions and also to interspecific interactions?; (3) what do earthworms contribute to the further colonization by other soil animals?; and (4) how do they impact the soil itself and what could therefore be the consequences for soil management and restoration? These questions have recently been addressed from the perspective of new (or 'alien') earthworm species invading ecosystems, suggesting a massive influx of species, competitive to the originally present fauna. This idea is, however, contrary to colonization, which suggests a gradual exploration of a previously uninhabited area. Unlike recent research, this review approaches colonization primarily as a spatial dispersal process and part of natural succession processes, and is mainly illustrated with examples of Palearctic species, either in Europe or introduced elsewhere. To begin, the various stages of colonization: dispersal, establishment, population growth and interspecies relations are analysed. Next, the colonization processes, the possible limiting environmental factors and the sequence of the appearance and establishment of species are described. Dispersal rates and sequences of colonization by different earthworm species are given for different soil ecosystems. For colonization, limiting environmental factors such as pH, soil type and heavy metal contents as well as the presence of organic matter seem to play a more important role than inherent ecological characteristics like r/K selection. Finally, the role of earthworms in the early colonization of soils that are earthworm-free because of non-cosmopolitan distribution, drained former sea bottom, permanently water-logged soils or anaerobic, acid peaty soils are reviewed. If we understand the role of earthworms in succession, we will be able to improve their role in soil restoration and soil management.