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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. peerReviewed

The optical properties of leaves and canopies determine the availability of radiation for photosynthesis and the penetration of light through tree canopies. How leaf absorptance, reflectance and transmittance and radiation transmission through tree canopies change with forest succession is not well understood. We measured the leaf optical properties in the photosynthetically active radiation (PAR) range of five Central European early-, mid- and late-successional temperate broadleaf tree species and studied the minimum light demand of the lowermost shade leaves and of the species&rsquo, offspring. Leaf absorptance in the 350&ndash, 720 nm range varied between c. 70% and 77% in the crown of all five species with only a minor variation from the sun to the shade crown and between species. However, specific absorptance (absorptance normalized by mass per leaf area) increased about threefold from sun to shade leaves with decreasing leaf mass area (LMA) in the late-successional species (Carpinus betulus L., Tilia cordata Mill., Fagus sylvatica L.), while it was generally lower in the early- to mid-successional species (Betula pendula Roth, Quercus petraea (Matt.)Liebl.), where it changed only a little from sun to shade crown. Due to a significant increase in leaf area index, canopy PAR transmittance to the forest floor decreased from early- to late-successional species from ~15% to 1%&ndash, 3% of incident PAR, linked to a decrease in the minimum light demand of the lowermost shade leaves (from ~20 to 1%&ndash, 2%) and of the species&rsquo, saplings (from ~20 to 3%&ndash, 4%). The median light intensity on the forest floor under a closed canopy was in all species lower than the saplings&rsquo, minimum light demand. We conclude that the optical properties of the sun leaves are very similar among early-, mid- and late-successional tree species, while the shade leaves of these groups differ not only morphologically, but also in terms of the resource investment needed to achieve high PAR absorptance.