Earthworms and collembola relationships: effects of predatory centipedes and humus forms

Relationships between anecic earthworms (Lumbricus terrestris and Aporrectodea giardi) and the collembolan species Heteromurus nitidus (Templeton, 1835), which is known to be attracted to earthworms, were investigated in an 8-week laboratory experiment. Our aims were (1) to assess whether earthworms influence the population dynamics of H. nitidus, and (2) to study pathways of influence and how earthworm effects are modified by humus forms and predators. Using microcosms with three defaunated humus forms, then provided with earthworms and predators, we intended to demonstrate that, amongst possible favourable effects of earthworms on springtail populations, earthworm activity may provide greater access and more pathways for springtails to explore soil and avoid predation. We expected that the effects of predators (centipedes) on the abundance of H. nitidus would increase from less (calcic mull) to more (moder) compact soil, and we hypothesized that earthworms would reduce predation pressure on H. nitidus by providing escape routes through increased macroporosity. Humus forms and earthworms only affected the population size of H. nitidus under high predation pressure, when collembolan numbers were higher in calcic mull than in moder, and were increased by the presence of earthworms. These results corroborate the hypothesis that earthworms, by increasing soil macroporosity, improve the escape routes for Collembola and thus evade predation. In moder humus earthworms increased the density of H. nitidus whether predators were present or not, so we cannot exclude that earthworms were also directly beneficial to H. nitidus. However, the hypothesis of a functional relationship mediated by soil macroporosity seems relevant since it was supported by differences observed when considering body size. When two size classes were distinguished within populations of H. nitidus (1) the positive effect of earthworms in moder was observed only on larger Collembola (>1 mm), (2) the density of the larger Collembola was decreased by predation only in moder and not in mull, (3) the effects of predators on the smaller individuals were not influenced by the presence of earthworms whatever the humus form, and was not decreased by the presence of earthworms. Nevertheless, factors other than macroporosity may operate as the presence of earthworms in acidic mull led to an unexplained decrease in the abundance of small-sized H. nitidus.