Downed deadwood habitat heterogeneity drives trophic niche diversity of soil-dwelling animals

Habitat heterogeneity is one of the important drivers of biodiversity, but the underlying mechanisms are still vague. Increasing habitat heterogeneity often coincides with greater habitat space and/or greater food availability for organisms, but it is unclear whether and how these components are related to habitat selection. Downed dead wood (i.e., dead wood laying on the ground) is an important but understudied component of dead plant matter that creates spatial habitat heterogeneity for soil fauna on the ground. To unravel the role of dead wood for providing niche dimensionality for soil fauna via variance in resource use, we sampled Collembola communities from moss growing on bark and from bark on the logs of two tree species (pedunculate oak and Norway spruce) and from soil next to the logs. We assessed habitat selection of Collembola species combining abundance data with trophic position data using stable isotope ratios (i.e., delta C-13 and delta N-15). Based on the Bray-Curtis dissimilarity, Collembola community composition differed significantly among the three habitats (i.e., moss, bark and soil). Collembola community-weighted mean delta C-13 and delta N-15 values also differed among the three habitats (average delta C-13 values of moss: -25.94 parts per thousand, bark: -25.30 parts per thousand, soil: -25.60 parts per thousand; average delta N-15 values of moss: -4.80 parts per thousand, bark: -3.83 parts per thousand, soil: -5.11 parts per thousand), partly reflecting isotopic differences of the habitats themselves (delta C-13 of moss: -29.83 parts per thousand, bark: -28.26 parts per thousand, soil: -29.04 parts per thousand; delta N-15 of moss: -7.59 parts per thousand, bark: -4.18 parts per thousand, soil: -7.84 parts per thousand). These findings indicate that Collembola depend on food sources derived from the habitats they inhabit at least at the community level. Collembola community-weighted variance of both delta C-13 and delta N-15 values ranked as bark > moss > soil. Species-level data suggested that the bark community included both bark-specific food specialists as well as species which likely use bark only as a temporary habitat based on their morphological traits supporting high mobility. These findings together indicate that the trophic niche variation of the Collembola community is larger in bark than in soil. Our results stress the importance of downed dead wood as a source of habitat heterogeneity, which in turn promotes soil animal diversity, likely via heterogeneity in food sources. Furthermore, the presence of dead wood on the ground could have consequences for organic matter processing by the soil animal community through sustaining a larger trophic niche variation even at a small spatial scale.