Tree roots in canopy soils of old European beech trees—An ecological reassessment of a forgotten phenomenon

Abstract

The formation of adventitious roots in humus accumulations in tree canopies is widely acknowledged from tropical and temperate rainforests, while the occurrence of those canopy roots in temperate tree species under mesic climates has been largely disregarded for ca. 100 years. Moreover, almost nothing is yet known of the ecological growth conditions or the structure or morphology of such canopy root systems. This study reports on the occurrence of tree fine roots in crown humus pockets of old European beech ( Fagus sylvatica L.) trees. The aim was to compare these canopy roots with the fine roots in the terrestrial organic layer soil in terms of fine root biomass density, root morphological traits, ectomycorrhizal colonisation and chemical composition of the root tissue, and to relate these root traits to the chemical properties of the respective soils. Fine root biomass density in crown humus pockets was ca. 7 times higher than in the terrestrial organic layer, even though soil chemical properties of both rooting media were similar. Fine roots in the canopy differed from terrestrial fine roots by lower specific root tip abundance, specific root length, and specific root surface area, all of which points to a longer lifespan of the fine roots in the canopy. Moreover, canopy roots revealed a lower percentage of root tips colonised by ectomycorrhizal fungi than terrestrial roots (87% vs. 93%). Chemical composition of the root tissue in canopy and terrestrial soils was similar for most elements, but canopy roots showed lower P, Fe, and Al concentrations and a higher N/P ratio than terrestrial roots. Root P concentrations of both canopy and terrestrial fine roots were closely related to soil P concentration, but not to soil C/P or N/P ratios. On the other hand, tissue N of canopy roots, but not of terrestrial roots, revealed a clear dependence on soil N and C/N values, suggesting a more limited N availability in the canopy soil compared to the terrestrial organic layer. However, the overall small differences in soil chemical properties between canopy and terrestrial organic layer soil cannot explain the markedly higher volumetric root density in the crown humus and the differences in ecomorphological traits between canopy and terrestrial soil. Instead, it is speculated that these differences are more likely a result of temporarily high water availability in crown humus pockets due to high water flow along the surface of branches to the central crown parts of the beech trees.