Influence of light availability on growth, leaf morphology and plant architecture of beech ( Fagus sylvatica L.), maple ( Acer pseudoplatanus L.) and ash ( Fraxinus excelsior L.) saplings

In a field study, we measured saplings of beech, ash and maple growing in a fairly even-aged mixed-species thicket established by natural regeneration beneath a patchy shelterwood canopy with 3–60% of above canopy radiation reaching the saplings. Under low light conditions, maple and ash showed a slight lead in recent annual length increment compared with beech. With increasing light, ash and maple constantly gained superiority in length increment, whereas beech approached an asymptotic value above 35% light. A suite of architectural and leaf morphological attributes indicated a more pronounced ability of beech to adapt to shade than ash and maple. Beech displayed its leaves along the entire tree height (with a concentration in the middle crown), yielding a higher live crown ratio than ash and maple. It allocated biomass preferentially to radial growth which resulted in low height to diameter ratios, and expressed marked plagiotropic growth in shade indicating a horizontal light-foraging strategy. In addition, beech exhibited the highest specific leaf area, a greater total leaf area per unit tree height, a slightly greater leaf area index, and a greater plasticity to light in total leaf area. Ash and maple presented a “gap species” growth strategy, characterized by a marked and constant response in growth rates to increasing light and an inability to strongly reduce their growth rates in deep shade. In shade, they showed some plasticity in displaying most of their leaf area at the top of the crown to minimize self-shading and to enhance light interception. Through this, particularly, maple developed an “umbrella” like crown. These species-specific responses may be used for controlling the development of mixed-species regeneration in shelterwood systems.     

Growth of European beech (Fagus sylvatica L.) saplings is limited by elevated atmospheric vapour pressure deficits

There is an ongoing debate about how European beech might be affected by a future drier climate. While numerous studies have examined the effects of soil drought on beech growth and development, studies investigating the effects of elevated atmospheric water vapour pressure deficit (VPD) are lacking. By increasing VPD in climate and open-top chamber experiments, with moisture in the rooting medium near optimum, we tested the hypothesis that increased VPD negatively affects the growth and development of European beech saplings. In the climate chambers, a reduction in relative air humidity by 40% resulted in a 68% reduction in productivity. Similarly, in the open-top chamber experiment conducted on the forest floor, biomass declined by 30% when relative air humidity was 15% lower. The reduction in biomass was mainly a consequence of a dramatically reduced leaf growth of beech in the elevated VPD treatments. Our results show that growth and development of beech saplings strongly depend not only on soil moisture but also on the prevailing VPD level. We conclude that the vapour pressure deficit is a widely ignored factor which influences the growth and vitality, and possibly also the distribution of European beech. Future forest management schemes under an altered climate should take this factor into account.     

Crown plasticity and neighborhood interactions of European beech (Fagus sylvatica L.) in an old-growth forest

Competition for canopy space is a process of major importance in forest dynamics. Although virgin and old-growth European beech (Fagus sylvatica L.) forests in Europe have been studied for many years, there are to date no studies of individual-tree crown plasticity and the way this is influenced by local neighborhood interactions in these forests. In this study, we analyzed crown plasticity and local neighborhood interactions of individual trees in the upper canopy of the old-growth beech forests of Serrahn, northeast Germany. In a 2.8-ha sample plot, we measured crown radii of all upper canopy trees and analyzed the direction and extent of crown asymmetry. Size, relative position, and distance of neighboring trees were used to construct vectors of neighborhood asymmetry within different distances from target trees. The crowns of beech trees showed strong morphological plasticity. Mean absolute and relative displacement of crown centers from the stem base were 1.95 m and 0.37, respectively. Circular–circular rank correlation coefficients between the direction of crown displacement and the direction of neighborhood pressure showed that trees strongly positioned their crowns away from local neighbors. Highest correlation coefficients were obtained when basal area and relative position of neighboring trees within a radial distance of 12 m were considered. Clark and Evans index and Ripley’s K-function showed that crowns were more regularly distributed than stems. Projected canopy cover was about 10% higher than canopy cover with simulated circular crowns. We conclude that the crowns of older beech trees have a high ability to plastically respond to changes in the local canopy conditions, enabling very effective exploitation of canopy space.     

The impact of roe deer (Capreolus capreolus), seedbed, light and seed fall on natural beech (Fagus sylvatica) regeneration

The aim of the present study was to quantify the effects of roe deer browsing relative to the effects of soil preparation, shelterwood density (light) and seed fall. Ultimately, the goal was better silvicultural guidelines to support the use of natural beech regeneration in the close-to-nature forestry context. In spring 1995, an experiment on natural regeneration with three experimental stands (0.28–0.6 ha) was installed in a beech (Fagus sylvatica) dominated broadleaved forest inhabited by a dense roe deer (Capreolus capreolus) population (24 deer km⁻²). The autumn of 1995 offered a large beech mast (stand average 307–1168 beechnuts m⁻²).Treatments installed included fencing to exclude deer, soil preparation, shelterwood thinning, and the relevant control treatments. Roe deer and soil preparation had dramatic effects on regeneration from the moment the seedlings sprouted in the spring 1996. The regeneration sprouted only sparsely in the unprepared seedbed producing only 5 seedlings m⁻² (average across fence treatments) after 2 months, which was reduced to 1 sapling m⁻² 8 years later. Deer had no significant effect on the regeneration density of the unprepared seedbed. In contrast, the cultivated seedbeds produced initially (after 2 months) up to 191 seedlings m⁻² protected by fence, whereas the densities in the unfenced treatments peaked by 22 seedlings m⁻². By the end of the study, these densities were reduced to 22 saplings m⁻² in the permanently fenced mineral soil seedbed and to 2 saplings m⁻² in the unfenced mineral soil seedbed. Regeneration height outside the permanent fence was generally only half the height of the regeneration inside. Additionally, we found significantly positive effects of increasing light and seed fall on regeneration density and of light on regeneration growth.We conclude that the roe deer only reduced the regeneration density of the dense regenerations established in the cultivated seedbeds. However, the regeneration density of the unprepared seedbed was not sufficient to support a future high quality stand. We found no treatment that within the timeframe of this study could support successful regeneration establishment outside the permanent fences. Whether this will change in the future with more time given for the regeneration to establish is yet to be revealed.     

The relationship between site factors and white ash (Fraxinus americana L.) decline in Massachusetts

Forest inventory records incorporating individual tree data are an important source of information about the extent and severity of past rates of forest decline, thus providing a temporal perspective for contemporary observations. We demonstrate the potential of this approach by using continuous forest inventory (CFI) data to reconstruct the extent, severity, pattern of development and site-factor associations for white ash decline in Massachusetts. White ash (Fraxinus americana L.) increased in basal area on Massachusetts CFI plots at about 1.5% per year from 1962 to 1979. Ash decline is, however, locally severe. Plot decline status was estimated, based on vigor rating, and growth and mortality rates of white ash. Plots with a mean change in basal area per year of −0.5% and/or with a mean vigor rating of 2.0 (on a scale of 0–4) were classified as ‘decline’. Using these criteria, 20% of the 82 CFI plots with 10% total basal area of white ash in 1962 were classified as ‘decline’ in both 1979 and 1991. These results suggest that statewide there has been no net loss of ash basal area since 1962. The total forest area affected by ash decline has not increased in the past decade.

Analyses of the CFI data indicate that decline was most prevalent on mesic sites, high on the landscape and/or on steep slopes. Such sites are potentially subject to large fluctuations in soil moisture availability during drought periods. Relationships identified between the prevalence of ash decline and site factors were further evaluated within a 2 ha intensived study site in the center of a 13 ha white ash stand affected by ash decline in 1990. Decline was heterogeneously distributed within the stand, with declining trees most frequent on the ‘decline-prone’ site-type identified through CFI plot analyses.     

Influence of slope position, stand type and rhododendron (Rhododendron ponticum) on litter decomposition rates of Oriental beech (Fagus orientalis Lipsky.) and spruce [Picea orientalis (L.) Link]

Oriental beech (Fagus orientalis Lipsky.) and Oriental spruce [Picea orientalis (L.) Link] are the two most common tree species in northeast Turkey. Their distribution, stand type and understorey species are known to be influenced by topographical landforms. However, little information is available as to how these changes affect litter decomposition rates of these two species. Here, we investigated the effects of slope positions (top 1,800 m, middle 1,500 m and bottom 1,200 m), stand type (pure and mixed stands) and purple-flowered rhododendron (Rhododendron ponticum) on litter decomposition rates of Oriental beech and spruce for 4 years using the litterbag technique in the field. Among these three factors, stand type had the strongest influence on litter decomposition (P < 0.001, F = 58.8), followed by rhododendron (P < 0.001, F = 46.8) and slope position (P < 0.05, F = 11.6). Litter decomposition was highest under mixed beech/spruce forest, followed by pure beech and spruce forest. Beech and spruce litter decomposed much faster in mixed bags (beech–spruce) than they did separately under each stand type. Purple-flowered rhododendron significantly reduced litter decomposition of Oriental beech and spruce. Beech and spruce litter decomposed much slower at top slope position than at either bottom or middle position. Differential litter decomposition of Oriental beech and spruce was mainly due to adverse conditions in spruce forest and the presence of rhododendron on the ground which was associated with lower soil pH. Higher elevations (top slope position) slowed down litter decomposition by changing environmental conditions, most probably by decreasing temperature as also other factors are different (pH, precipitation) and no detailed investigations were made to differentiate these factors. The adverse conditions for litter decomposition in spruce forest can be effectively counteracted by admixture of beech to spruce monoculture and by using the clear-cutting method for controlling rhododendron.