Forest structure and C stocks in natural Fagus sylvatica forest in southern Europe: The effects of past management

Managed forests often differ substantially from undisturbed forests in terms of tree structure and diversity. By altering the forest structure, management may affect the C stored in biomass and soil. A survey of 58 natural stands located in the south-westernmost limit of European beech forests was carried out to assess how the C pools are affected by the changes in tree structural diversity resulting from past management. The mean tree density, basal area and the number of large trees found in unmanaged forests were similar to those corresponding to virgin beech forests in Central Europe, whereas large live trees were totally absent from partially cut stands. Analysis of the Evenness index and the Gini coefficient indicated high structural diversity in the three stand types. The results of the Kolmogorov–Smirnov test used to compare the diameter distributions of each group revealed significant differences between stand types in terms of distributions of total tree species and of Fagus sylvatica.

The mean C stocks in the whole ecosystem – trees, litter layer and mineral soil – ranged from 220 to 770 Mg ha⁻¹ (average 380 Mg ha⁻¹). Tree biomass (above and belowground), which averaged 293 Mg C ha⁻¹, constituted the main C pool of the system (50–97%). The statistical test (Kolmogorov–Smirnov) revealed differences in the distribution of C pools in tree biomass between unmanaged and partially cut stands. As a consequence of the presence of large trees, in some unmanaged stands the C stock in tree biomass was as high as 500–600 Mg C ha⁻¹. In the partially cut stands, most of the C was mainly accumulated in trees smaller than 20 cm dbh, whereas in unmanaged stands the 30% of tree C pool was found in trees larger than 50 cm dbh. Furthermore, many unmanaged stands showed a larger C pool in the litter layer. The C content of mineral soils ranged from 40 to 260 Mg C ha⁻¹ and it was especially high in umbrisols. In conclusion, the implementation of protective measures in these fragile ecosystems may help to maintain the highly heterogeneous tree structure and enhance the role of both soils and trees as long-term C sinks.     

Estimation of litter fall and seed production of Acacia mangium in a forest plantation in South Sumatra, Indonesia

Annual litter fall of Acacia mangium in the period of September 1995 to August 1996 was estimated at 5939 kg ha⁻¹ year⁻¹ and from September 1995 to August 1996 at 6048 kg ha⁻¹ year⁻¹, with the highest seasonal production in the dry season. The litter fall was dominated mainly by leaves, 4446 kg (75%) and 4137 kg (68%), respectively. Seed production in the litter fall was estimated at 42.4 kg ha⁻¹ year⁻¹ (4.1 million seeds ha⁻¹) and 39 kg ha⁻¹ year⁻¹ (3.8 million seeds ha⁻¹), with the highest in the dry season from June to October. The accumulated litter fall in the forest floor together with shrubs and grass provide a high fuel load, increasing fire risk.     

Effects of vegetation control on nutrient removal and Fagus orientalis, Lipsky regeneration in the western Black Sea Region of Turkey

Dense Rhododendron ponticum (L.) understories of eastern beech (Fagus orientalis Lipsky) stands in the Black Sea Region (BSR) of Turkey create challenging forest vegetation management problems relative to beech regeneration. Rhododendron traditionally is controlled in Turkish forests with grubbing and bulldozing. The effects of these practices on nutrient removal and natural beech regeneration have not been quantitatively studied. Two woody vegetation control treatments (bulldozing and hand-grubbing) were installed during late summer, 2002 in three mature beech stands with dense rhododendron understories in the Düzce Forest Management Directorate, in the Turkish western BSR. Aboveground biomass of each vegetation component, total aboveground vegetation biomass, nutrient concentrations, organic matter (OM) removal, and total amount of OM nutrients were determined for each woody vegetation control treatment. Soil bulk density and nutrient content, and beech seedling biomass, nutrient content, and natural regeneration also were studied. One year after treatment, the machine site preparation by bulldozing (MSP) that removed understory vegetation and attached roots, reduced mean forest floor OM content by about 84%, when compared to hand-grubbing. Mean soil C, N, K and Mg concentrations on the bulldozed sites were 36, 27, 50 and 55% less, respectively, than those on the grubbed sites. Total C and Mg amounts at the 5–10 cm soil depth were 24 and 47% lower, respectively, for mechanical site preparation (MSP) sites, when compared to grubbed sites. Overall, soil bulk density did not differ significantly between the grubbing and MSP treatments. Frequent passes on designated transects on MSP sites resulted in a significant (P < 0.01) one-third increase in bulk density of the first 20 cm soil depth, when compared to grubbed sites. No natural beech regeneration occurred on untreated control sites. Mineral soil exposure on the MSP sites increased beech seedling germination substantially. One year after treatment, the mean number of naturally regenerated seedlings for MSP sites was 2.5 times greater than for grubbed sites. Woody control treatments had significant effects upon beech seedling chemistry, with N, P, K, Ca and Mg concentrations averaging 35, 47, 12, 33 and 25% lower, respectively, for MSP sites, when compared to grubbed sites. However, mean seedling biomass and nutrient content were significantly greater (P < 0.05) on MSP sites. Long-term effects from windrowing on MSP site residues, associated topsoil and nutrients need to be evaluated.     

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.     

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.     

Roles of seed and pollen dispersal in natural regeneration of Castanopsis fargesii (Fagaceae): Implications for forest management

Natural regeneration is an important process to reverse the loss of forests. Understanding the process of natural regeneration is crucial for achieving sustainable forest management. In this study, we examined the effects of seed and pollen dispersal in naturally regenerated populations of Castanopsis fargesii. Genetic variation in six populations along two successional series (three successional stages in each series: early, pre-climax, and climax) was assayed using RAPD (random amplified polymorphic DNA) markers. High genetic variability was observed as measured with Shannon's information index. A majority of genetic variation was distributed within populations (Φ_st = 0.1271) and significant isolation by distance existed among populations. A contrasting pattern of genetic variation along these two series was observed, representing different scenarios of natural regeneration processes. The ratio of the number of migrants between the mature populations to the number of migrants from the mature to immature populations was estimated as 1.146 ± 0.099 to 1.981 ± 0.164, implying that comparable seed and pollen dispersal might exist at a fine spatial scale (∼2 km²). The results suggest the critical role of seed dispersal in shaping genetic composition and diversity in the second-growth forests. Barriers to seed dispersal from a variety of genetic sources could result in low genetic diversity in naturally regenerated populations. Management that facilitates the connectivity of newly regenerated stands with mature forests could be effective for natural regeneration given the predominant role of short-distance dispersal of seeds in this species.     

Differences in abundance,species richness,and body size of ground beetles (Coleoptera: Carabidae) between beech (Fagus sylvatica L.) forests on Podzol and Cambisol

Ten forest sites with a tree layer of almost pure beech in southern-central Scania, south Sweden, were studied. They were all located in the interior of large forest stands. Five of the stands were developed on mor Podzols with scanty or almost no ground layer vegetation. The other five sites were located on mull Cambisols with a rich ground layer of many vascular plant species and a shrub layer of varying density. In total, 42 carabid beetle species and 9260 individuals were captured with pitfall traps (15 per site, diameter 87 mm) during 12 weeks in April, July and September 2007. Composition of the carabid fauna differed considerably between the two types of sites. In the Cambisol sites 39 carabid species were found, compared to only 21 species in the Cambisol sites. Mean carabid species richness per site differed significantly between Cambisol and Podzol sites (n = 20.4 and 13.2, respectively). The difference was wholly due to a lower number of small carabid species (mean body length <12 mm) in the Podzol sites. Of four variables (pH-KCl, Clay, Organic matter, and Sum of plant cover percentages below tree canopy) the sum of plant cover variable usually accounted for more (>50%) of the carabid beetle variability compared to the three other variables. Carabus glabratus and Pterostichus oblongopunctatus were almost exclusively found on Podzol. Carabus coriaceus, Pterostichus melanarius and in particular Carabus nemoralis were more abundant on Cambisol, whereas Carabus hortensis and Pterostichus niger were almost ubiquitous and attained the highest total numbers. There was no significant difference between Podzol and Cambisol sites in the total number of carabid individuals captured. C. coriaceus, C. hortensis, and Carabus violaceus, species that were abundant enough in all sites for a comparison, had significantly larger mean body sizes in the Podzol than in the Cambisol sites, a finding valid for both sexes. There were more males than females of C. coriaceus, C. violaceus, and C. nemoralis in the catches, whereas sex distribution of C. hortensis was equal. Possible explanations for the findings are differences in microclimate and food web characteristics related to differences between Podzols and Cambisols in understorey vegetation and soil or litter properties.     

The influence of silvicultural treatments and site conditions on American chestnut (Castanea dentata) seedling establishment in eastern Kentucky, USA

After more than 50 years of research and selective breeding, blight-resistant American chestnut (Castanea dentata) trees will soon be available for planting into the species’ pre-blight range. Increased understanding of the regeneration requirements of pure American chestnut (C. dentata [Marsh.] Borkh.) will increase the success of future efforts to establish blight-resistant chestnut. We quantified survival and initial growth of bare-root American chestnut seedlings at five locations in eastern Kentucky, USA. We used a split-plot design to compare seedlings planted within adjacent mesic and xeric sites treated with either a two-age shelterwood overstory treatment or a midstory removal treatment. The silvicultural treatments and topographic settings allowed us to evaluate chestnut seedling performance under four light and site productivity combinations. Seedling survival was 57% and seedling height averaged 94 cm following two growing seasons. Seedling survival was negatively related to sand and coarse fragment content, but was unrelated to silviculture treatment or topographic position. Chestnut seedlings grew best in shelterwood overstory treatments areas on mesic sites. Seedlings growing in shelterwood overstory treatment areas added 3- and 3.5-times more height and stem increment compared to seedlings planted after midstory removal. Seedling leaf mass and foliar nitrogen (N) content were also greatest in shelterwood plantings on mesic sites. The high-light environment created by shelterwood overstory removal resulted in better initial seedling growth, but the moderate-light of the midstory removal treatment may ultimately provide chestnut seedlings a greater advantage over competing vegetation.