Functional Crown Architecture of Five Temperate Broadleaf Tree Species: Vertical Gradients in Leaf Morphology, Leaf Angle, and Leaf Area Density

The morphology, inclination, and spatial distribution of leaves in different parts of tree crowns are important determinants of the radiation, momentum, and gas exchange between the canopy and the atmosphere. However, it is not well known how these foliage-related traits vary among species differing in successional status. We measured leaf size, leaf mass area (LMA), leaf inclination (angle to the horizontal), leaf area density (LAD), total leaf area (leaf area index, LAI), and leaf area distribution across the crown in adult trees of five common, early to late-successional tree species (Betula pendula Roth, Quercus petraea (Matt.) Liebl., Carpinus betulus L., Tilia cordata Mill., and Fagus sylvatica L.) using different canopy access techniques and the harvest of foliated trees (29 trees in total). Leaf size increased continuously with crown depth in B. pendula and T. cordata but peaked at mid-crown in Q. petraea, C. betulus, and F. sylvatica to decrease toward the shade crown. By contrast, LMA and leaf angle decreased continuously with crown depth in all species, but the pattern of vertical change varied. The mid/late- and late-successional species had higher LAI, lower shade-leaf LMA, lower leaf angles (shade and sun crown), and higher LAD in the uppermost sun crown in comparison to early successional B. pendula. We assume that the most peripheral sun leaf layer is partly acting as a shield against excess radiation, with foliage properties depending on the structure of the shade crown. We conclude that the vertical change in leaf morphology, inclination, and spatial distribution in tree crowns is highly species specific, with partial dependence on the species' position in succession.