Trickle-down ecology: Vertical stratification of temperate forests drives asymmetric cross-layer effects on consumer communities

Abstract

O_LIForests are vertically structured ecosystems where light attenuation, microclimatic buffering and resource availability occur along continuous gradients from the canopy to the understorey layers. Despite this complexity, studies rarely integrate trophic interactions across vertical layers, overlooking how vertical forest structure shapes consumer communities through abiotic and biotic pathways. C_LIO_LIIn this study, we combined layer-specific measurements of plant diversity, structure and microclimate with arthropod sampling in the canopy and understorey, as well as bird survey data, in a temperate forest. Applying Bayesian structural equation models with explicitly defined directional pathways, we modelled both consumer biomass and abundance across trophic levels and vertical layers. C_LIO_LIAbundance measures were predominantly filtered by local layer conditions, while biomass responded to conditions across layers, reflecting stand-level energy flow. This suggests that these two metrics capture fundamentally different ecological processes. Canopy conditions consistently predicted understorey arthropod abundance across trophic levels, while the reverse was not observed, suggesting a strong asymmetric downward propagation of canopy-driven effects. Furthermore, trophic interactions between arthropod primary and secondary consumers remained largely stratified within vertical layers, suggesting a vertical food web compartmentalisation rarely shown in structurally complex aboveground systems. C_LIO_LIPlant diversity, structure and microclimate shaped consumer communities mainly through the modulation of resource availability and plant apparency, with effects varying across vertical layers, trophic levels and taxonomic groups. Through complementary mechanisms, plant diversity likely determined the variability of resources available to consumers at different trophic levels. Structural properties, in contrast, potentially drove the spatial redistribution of these resources through light attenuation and microclimatic buffering, which in turn influenced the physiological capacity of consumers to access and exploit available resources. C_LIO_LIThese findings demonstrate that vertical stratification mediates trophic pathways in a highly directional manner, with canopy characteristics playing a disproportionate role in structuring the forest community across layers. Integrating layer-specific structural and trophic indicators into forest biodiversity assessments and management strategies is therefore essential to fully evaluate biodiversity dynamics and multifunctionality in structurally complex forest ecosystems. C_LI