Vertical stratification of invertebrate assemblages in water-filled treeholes of a temperate deciduous forest

Water-filled treeholes provide temporal habitats and resources to detritus-based aquatic organisms in the aboveground forest strata. Treeholes are found at different vertical positions, and are often surrounded by dense understory vegetation, which may affect water volume and litter weight. Differences in water volume and litter weight are indicative of the habitat quantity and the quality of treehole communities. Therefore, in the present study, we used containers as artificial treeholes to examine the effects of these characteristics on treehole invertebrate assemblages in a broad-leaf deciduous forest in central-eastern Japan. We first generated two models: a direct-effect model that reflected the physical and chemical properties of treeholes (water volume, litter weight, and other measurable properties that might directly influence invertebrate survival and fitness); and an indirect-effect model that reflected the differences in vertical position and surrounding understory vegetation per se. We compared these models and found that species richness is better explained by the direct-effect model, whereas the indirect-effect model plausibly explains the differences in invertebrate abundances. Further analyses revealed that some species such as Tripteroides bambusa utilized lower treeholes, while higher treeholes had a greater abundance of Ceratopogonidae sp. A within dense understory vegetation. Our study demonstrates that treehole invertebrates are not only influenced by simple physicochemical properties, but also by ambient conditions. However, the response patterns were highly variable across species. Our approach provides insight for elucidating the key drivers of treehole detritivore diversity in vertically stratified environments.