Diversity and succession of adventive and indigenous vascular understorey plants in Pinus radiata plantation forests in New Zealand

The vegetation of Pinus radiata plantation forests in New Zealand was studied to examine how the indigenous flora has responded to this novel habitat. A chronosequence of stands about 5, 16 and 27 years was assessed in each of four different biogeographic regions to test the effects of several stand and site factors on the succession of vascular understorey plant communities. A total of 202 indigenous and 70 adventive vascular plant species were found across all study areas, with considerable geographic variation among forests in species composition, species richness (range 48–135 species), and the percentage of indigenous species (50–86%). Both richness and cover of adventive species decreased significantly over time, whereas richness and cover of indigenous species was highest in the oldest stands, and overall species richness was lowest at mid-rotation. The guild composition changed from dominance of grasses and forbs in young stands to dominance of ferns and understorey trees in mature stands. These temporal changes were accompanied by a decrease in light-demanding pioneer species and an increase in shade tolerant, later seral species adapted to a forest environment. Measurements of the degree of canopy closure in stands with low or high stocking and modelling of temporal changes of canopy closure indicated that these understorey plant dynamics are influenced by changes in light availability as stands age. Despite the successional changes within forests, geographic variation more strongly influenced understorey communities because stands within a forest area were grouped together in DCA and TWINSPAN analyses, along rainfall and temperature gradients. Although the canopy species of such intensively managed plantation forests is an alien element in the New Zealand flora, the sheltered forest environment of older stands allows the establishment of a mostly indigenous forest understorey community with considerable similarities to indigenous forests located nearby.     

Effects of artificial afforestation and successional stage on a lowland forest bird community in southern China

To examine the relationship between forest succession after severe logging forestry practices and the composition of avian communities, we investigated how forest bird composition and guild structure change as a function of structural properties along a successional gradient, including a climax mature forest (>400 years), a rehabilitated mixed forest (50-70 years), and a disturbed Masson pine forest (70 years) of the Dinghushan Nature Reserve, Guangdong Province, China. Of a total of 51 resident species recorded, mixed forests hosted the highest numbers of individuals and species, reflecting the high species richness of both forest and non-forest species. For forest-dependent species, however, mature stands had the highest observed and estimated species richness. Of 6 habitat-use guilds identified, vertical-profile generalists and understory-birds formed the two dominant guilds, accounting for 54.0% and 38.7% of all individuals respectively. The results of canonical correspondence analysis (CCA) clearly showed that most forest-dependent species were associated with high proportions of native canopy cover and the mean density of dead trees and large trees, which are characteristic of old-growth mature forests (horizontal heterogeneity) at stand level. Accordingly, conservation efforts should focus on the specialized requirements of the most habitat-restricted species in the future, especially for understory insectivores (Babblers) and large-tree users in mature subtropical monsoon forests of southern China. Moreover, since regenerating mixed forests are very similar to mature forests in both vegetation structure and bird community composition, we recommend that logging cycles (>50 years) be increased to a minimum of 50 years in southern China, so that a balance between economic and ecological interest can be reestablished.     

Composition,structure, and management potential of secondary dry tropical vegetation in two abandoned henequen plantations of Yucatan,Mexico

Data on the composition and stand dynamics of abandoned henequen (Agave fourcroydes) fields of different ages are presented and their management potential discussed. The study was carried out in the Hacienda of Kancabchen, Yucatan, Mexico in 12- and 26-year-old secondary forests. Species richness and diversity were greater as the stand grew older. Even though 73% of species have the capacity to regenerate from coppiced shoots, only between 30 and 50% of the individuals actually regenerate from coppiced shoots in both communities, respectively. According to the Sørensen similarity index both stands are similar in composition (73%), while the Motyka index indicates that they differ in biomass distribution (17%). This could be due to soil conditions and the management of these areas, or a combination of both. The 26-year-old abandoned henequen stand still has pioneer species dominating and has very few late successional species present. Slow-growing tree species on the 26-year-old stand were expected, but their absence could be due to the lack of seed sources, a limited dispersal capability, or to an arrested succession due to human perturbation and management. Eight species dominated the stands, accounting for more than 60% of the total importance values. Most species present in both stands are widely used locally and are commercialized inside the rural communities. Potential management strategies could involve forest thinning to remove competition and favor those species used by the communities, those with potential to increase soil fertility, and those that could have semi-commercial purposes. Management could also include enrichment with other species such as Cedrela odorata and Apoplanesia paniculata that are widely used and were previously present as part of the native vegetation of the area.     

Silviculture as a disturbance regime: the effects of clear-cutting,planting and thinning on polypore communities in mixed forests

The structure of modern forest landscapes is profoundly affected by human-caused disturbances, particularly forest management; however, the effects and prospects of individual silvicultural techniques are insufficiently understood. This study distinguishes the effects of clear-cutting, planting and thinning on species richness and community composition of polypore fungi. In 2008–2009, 181 forest compartments (ranging from naturally regenerated deciduous stands to planted Picea abies stands and 0–137 years post clear-cutting) were explored in a hemiboreal landscape subjected to even-aged management. Altogether 104 polypore species were recorded. For species richness, time since clear-cutting was the most influential factor at both stand and landscape scales, followed by thinning. Clear-cuts had distinct polypore communities (including several red-listed species) whose species richness declined in time. Following 20 years post clear-cutting, species richness started to increase along different community–composition pathways determined by regeneration type. The communities developed after planting were moderately species rich at stand scale but homogeneous over larger areas. Thus, at landscape scale, mature unmanaged naturally regenerated stands hosted most species; thinning reduced species richness by approximately 15%, and among thinned stands, planted areas had a further 9–22% fewer species than naturally regenerated areas. In such variably managed landscape, silviculture appeared to create particularly distinct communities in young stands on nutrient-rich soils, which naturally provide polypores with a rich supply of small deciduous snags absent from stands artificially planted with P. abies and intensively thinned.     

Centuries-old logging legacy on spatial and temporal patterns in understory herb communities

Understory herb communities in the Southern Appalachians are among the highest biodiversity plant communities in North America. In the mid-1990s, a debate began over whether understory herb communities recover to their pre-disturbance states following logging. Studies showing reduced herb-layer diversity in previously logged forests were criticized for not accounting for intersite environmental heterogeneity. More recent studies have addressed environmental heterogeneity, but have neglected long-term recovery by using “mature forests” as young as 80 years old as the benchmark for diversity comparison, even though old growth stands have disturbance return intervals exceeding 500 years. Here we address concerns clouding previous studies of high-diversity Appalachian herb communities and investigate their long-term recovery by comparing paired sites of old growth forest and forest logged 100–150 years ago. We found that species richness and individual abundance is greater in old growth forests than mature forests and that species composition differed significantly between the two. Turnover in species among old growth and mature forests accounted for 11% of the total species richness and was significantly greater than expected. Species turnover at intermediate (5–50 m) and landscape-scales (>10 km) contributed the most towards total species richness. Herb communities in rich cove forests have successional trajectories that exceed 150 years, with important community changes still occurring long after the forest returns to what has been previously termed a “mature” state. To conserve the diverse herb layer, we conclude that mature forest stands are too young to serve as baselines for recovery, landscape-scale preservation of multiple forest stands is needed to maximize species richness, and maintaining 100–150-year logging rotations will likely lead to loss of biodiversity.     

Deer density effects on vegetation in aspen forest understories over site productivity and stand age gradients

Studies examining the interacting effects of ungulate herbivore pressure and site productivity on vegetation are mostly on grassland-grazing systems and have shown conflicting patterns. Here we examine the effects of deer density (>30 years differences in density between two landowners), site productivity (site index, SI) and stand age on subcanopy vegetation characteristics in 60 closed canopy, clear-cut origin Populus tremuloides dominated stands, Michigan, USA. Stand age effects were included because age varied among stands and can affect subcanopy vegetation patterns. Compared with fewer deer, stands with more deer had greater total forest floor vegetation mass, and its major components bracken fern (Pteridium aquilinum), sedge (mostly Carex pensylvanica) and trees/shrubs <0.25 m tall, but lower forb mass and lower forest floor vegetation species richness and diversity. Deer density and SI had strong interacting effects on total forest floor mass, forb mass, and species richness. Forb mass increased with SI, but only in stands with fewer deer, whilst total vegetation mass was greater in stands with more deer at lower SI and declined with SI more sharply than for stands with fewer deer. Species richness increased with SI but more so at lower than higher deer density. Deer density and age had interacting effects on mass of trees/shrubs <0.25 m tall and sedge. Compared with fewer deer, stands with more deer had greater sedge and tree/shrub mass, and sedge mass decreased and tree/shrub mass increased more sharply with age. In lower deer stands there was a dense subcanopy tree and shrub strata within and beyond the reach of deer 0.9-10 m tall whereas in higher deer stands this vegetation layer was nearly absent. We conclude that higher deer browse pressure in early successional Populus stands (1) strongly limits the recruitment of woody stems to larger (>0.9 m tall) size classes, which could affect long-term successional trajectories, and (2) diminishes forb density and species richness, especially at higher site productivity, but increases total forest floor vegetation mass (mostly bracken fern and sedge), especially at lower site productivity. Given associations of bracken fern and sedge with poorer and/or more open sites and assuming high palatability of forbs, this pattern may result from the combination of selective herbivory and higher light availability caused by limited recruitment of trees and shrubs to taller strata.     

Successional process of Picea crassifolia forest after logging disturbance in semiarid mountains: A case study in the Qilian Mountains,northwestern China

Selective logging is the most widely employed method of commercial timber production in Asia, and its impact on forest structure, composition, and regeneration dynamics is considerable. However, the successional processes in forest communities after logging in semiarid mountains are poorly understood. To provide more information on these processes, we used data from tree rings, direct and indirect age determinations, and field measurements of stand structure to reconstruct the historical disturbance regime, stand development patterns, and successional processes in a natural Picea crassifolia forest community in the Qilian Mountains of northwestern China. The results showed that the density of P. crassifolia forest increased significantly after logging. The densities of second growth forests 30 and 70 years after logging disturbance had increased to 2874% and 294% of primary forest's density, respectively. Logging disturbance did not alter tree species composition of logged stands. However, the diversity of understory species changed significantly among the successional phases. Logging disturbance decreased the spatial heterogeneity of second growth forest. The spatial distributions of recruitment were affected by the location of the remaining trees. There was less recruitment near the remaining trees than near forest that had been cut. In addition, logging disturbance also induced a growth release for the trees on the sites sampled. Our results imply that the succession and regeneration of P. crassifolia forest may be improved if the remaining trees could be retained relative uniform distribution pattern, thinning or selective logging could be performed to height density, exotic shrubs could be removed or the shrubs cover could be reduced during the earlier successional stages.     

Patterns of field layer invertebrates in successional stages of managed boreal forest: Implications for the declining Capercaillie Tetrao urogallus L. population

Capercaillie (Tetrao urogallus L.) populations in Finland have decreased markedly during past decades. One of the assumed reasons is the decreased quality of brood feeding grounds since current forest management transfers mature forests to younger successional stages. We studied how different types of managed Finnish forests offer resources for capercaillie broods by comparing the vegetation and invertebrate fauna of four successional stages in the two most common forest types in Finland, spruce dominated Vaccinium myrtillus (MT) and pine dominated Vaccinium vitis-idaea (VT) type. Forest age class had a significant effect on the cover of bilberry. There was a positive correlation between bilberry cover and the biomass of larvae and of all invertebrates (including all developmental stages) both at the stand and the sample level. Both forest type and age class significantly affected the biomass of larvae. Mature stands and young stands had the highest biomass of larvae in both MT and VT. In both forest site types the sapling stands hosted the smallest biomass of larvae. The results suggest that successional stages that follow clear cutting seriously lowers the food availability for capercaillie chicks and also reduces the shelter that field layer might provide. However, already the young stages that follow sapling phases seem to provide both bilberries and invertebrates so it is quite unlikely that the bilberry abundance alone could explain the dramatic decline of the capercaillie.     

Composition and structure of natural mixed-oak stands in northern and central Portugal

Stand composition and structure of natural mixed-oak stands of common-oak (Quercus robur L.) and pyrenean-oak (Quercus pyrenaica Willd.) were studied. Diverse compositional and structural elements in early and late successional stand stages were analysed. The study was conducted in north and central Portugal where different natural mixed oak forests types are located. The following mixed-oak forest types involving common-oak and pyrenean-oak were studied: common-oak & other hardwoods; common-oak & cork-oak (Quercus suber L.); ash (Fraxinus angustifolia Vahl) & pyrenean-oak; and pyrenean-oak & madrone (Arbutus unedo L.). Measurements were made in early and late successional stand stages on the different mixed oak forest types. Different stand characteristics and indices were used to describe and compare stand structure and composition. The study showed changes in species diversity and stand structure. Most tree species in mature stands are present in early stages but with higher abundance. Shannon diversity index may change between 0.798 and 1.915. Significant differences on species diversity and abundance were found depending on the forest type and successional stage. Mature mixed-oak forests have high species diversity with an abundance of small to medium tree size species. Species distribution and diameter differentiation indices range from 0.30 to 0.70 and 0.52 to 0.82, respectively, revealing significant structural complexity. The average number of standing and downed dead trees was 265 and 83 trees ha⁻¹ for early and late stage, respectively, with 6.9 and 65.4 m³ ha⁻¹. Higher values of stand diversity index were 41 and 53 in more complex and developed forests. Later stand stages have complex structure, with a wider range of tree diameter distribution and higher degree of irregularity.     

Do position and species identity of neighbours matter in 8–15-year-old post harvest mesic stands in the boreal mixedwood?

Neighbourhood competition indices (NCI), where position and species identity of neighbours are known, have been used to investigate growth and competitive interactions among adult trees. In this study, we used NCI in 8–15-year-old stands following clear-cutting in a boreal mixedwood forest of eastern Canada to improve our understanding of early successional forest dynamics. Trees of increasing diameter from the center (≥1 cm) to the edge (≥5 cm) were mapped in twenty-five circular 450 m² plots. Target trees (DBH ≥ 1 cm) were sampled in plot center to determine their annual radial stem growth. For each species, we compared a set of growth models using either a spatially explicit NCI or a non-spatial competition index. Both types of indices estimated a species-specific competition coefficient for each pair of competitor–target species. NCI were selected as the best competition model for all target species although differences in variance explained relative to the non-spatial index were small. This likely indicates that competition occurs at the local level but that the high density and the relative uniformity of these young stands creates similar neighbourhoods for most trees in a given stand. The effective neighbourhood radius for competitors varied among species and was smaller for shade tolerant species. Intraspecific neighbours were the strongest competitors for most species. Aspen (Populus tremuloides) was a weak competitor for all species as opposed to balsam fir (Abies balsamea) which was a strong competitor in all cases. These results are in contradiction with some widely used forest policies in North America (e.g. free-to-grow standards) that consider broadleaf species, such as aspen, as the strongest competitors. For these early successional forests, the decision regarding the use of spatial or non-spatial competition indices should rest on the intended use. For even-age management, spatial indices might not justify their use in high-density stands but they are needed for the simulation of novel harvest techniques creating complex stand structure.     

晋西黄土区不同森林树种及其林地土壤养分含量的变化

[目的]探究晋西黄土区不同树种营养器官(叶、枝、干、根)及其土壤的养分含量变化特征。[方法]通过野外样地调查和取样分析方法,对3种典型森林树种(刺槐、侧柏、辽东栎)不同器官及其林地土壤有机碳、全氮、全磷含量进行了研究,并探讨了不同树种叶片与其土壤C、N、P化学计量的关系。[结果]辽东栎、刺槐和侧柏的叶片有机碳含量分别为468.43、454.96、438.53 g·kg-1,刺槐、辽东栎和侧柏叶片的全氮含量分别为27.52、20.74、12.73 g·kg-1,辽东栎、侧柏和刺槐叶片的全磷含量分别为2.73、2.15、1.35 g·kg-1,叶片C∶N值为侧柏辽东栎刺槐,C∶P、N∶P值分别为刺槐侧柏辽东栎、刺槐辽东栎侧柏;3个树种树枝的有机碳含量均最大,树叶的全氮含量均最大,而树干的全氮含量均最小。侧柏、辽东栎、刺槐树枝的全磷含量分别为3.07、3.07、1.87 g·kg-1,显著比其它器官的大;3种森林土壤的C、N、P含量均随土层的加深而降低,且0 10 cm土层的含量最大,C∶N、C∶P、N∶P值随土层深度的变化不一致;刺槐叶片的有机碳、全磷含量与土壤C∶N值显著或极显著负相关;侧柏叶片的有机碳含量均与土壤C∶N、C∶P、N∶P值极显著负相关,叶片N∶P值与土壤C∶N、C∶P、N∶P值均极显著正相关;辽东栎叶片的有机碳含量、C∶P值与土壤有机碳含量极显著负相关。[结论]辽东栎、刺槐和侧柏的有机碳含量均较高,分别为468.43、454.96、438.53 g·kg-1;辽东栎和刺槐对晋西黄土区干旱环境的防御能力比侧柏强,当地环境较适宜辽东栎的生长发育,而刺槐和侧柏的生长分别受土壤P和N的限制。     

宁夏六盘山区辽东栎林的空间分布及林分特征

基于2005年森林资源一类清查数据,分析了六盘山区辽东栎林分结构和空间分布特征。结果表明:辽东栎林在六盘山主要分布于海拔1 900-2 300 m范围内,坡度为20-35°的半阴坡、半阳坡和阴坡上,与山杨林的分布生境相近。现有的辽东栎林多为幼龄林,平均林龄30 a,林分密度较大,平均1 046株·hm^-2,树木平均高度7.8 m,平均胸径12.4 cm。胸径5-10 cm的辽东栎株数占50％以上,平均木材蓄积量仅为 39.9 m³·hm^-2。随着海拔升高、坡位从坡上部到坡下部,坡向从半阴坡到半阳坡,林分密度增大,平均树高和胸径增加,这主要是因为沿此梯度林龄在增加,而且随着年龄的增加,辽东栎林的平均胸径以0.8 cm·a^-1的速度增大,作为主要先锋伴生树种的山杨在林内的比重则下降,说明六盘山区的现有辽东栎林正处于正向演替阶段,更趋稳定。建议对辽东栎林采取近自然林业的管理方式,减少不利的人工干预,加速培养近自然森林。     

The ecology of tree regeneration in mature and old forests: combined knowledge for sustainable forest management

We focused our attention on quantifying the factor complex of forest regeneration in 423 mature and old stands with contrasting environmental conditions. We recorded the microhabitat selection of tree recruits, the frequency of tree seedlings, and evaluated the drivers of sapling abundance and diversity. The majority of forest regeneration was established on undisturbed forest floor. Dead wood was a frequent substrate in spruce-(co)dominated forests. Seedling frequency within a stand was related to the site-type specific productivity gradient of stands—pine seedlings were common in low-productivity and spruce in high-productivity boreal forests. Seedlings of temperate broad-leaved trees dominated in productive boreonemoral forests, except for oak, which showed a uniform distribution of abundance in all forest site-types. Sapling abundance was dictated by forest site-type, and facilitated by stand diversity, variability in stand closure, lying dead wood, abundant moss, and a thick organic layer. Only in boreal forests was sapling abundance suppressed by the abundant spruce and younger trees. Upon considering the relationship between sapling abundance and species richness, sapling diversity was dependent on forest site-type, suppressed by stand density and dead wood (old gap) abundance, and facilitated by stand diversity. In addition, boreonemoral stands, competition from herbs, and facilitation by mosses occurred. The observed pattern of tree recruitment points to the importance of top-down effects of the overstory, competing or facilitating interactions with forest floor vegetation, and availability of regeneration microhabitats, which in complex make their ecology comparable with forest herbs. Natural forest regeneration can be enhanced if silvicultural methods support mixed stands and enhance field layer diversity. Oak can provide the universal tree species to improve stand structure over a wide range of habitats.     

Retention of aspen (Populus tremulae) at final cuttings - The effect of dead wood characteristics on saproxylic beetles

To preserve biodiversity in managed forest landscapes dead and living trees are retained at final cuttings. In the present study we evaluated the effect of these practices for saproxylic (wood-dependent) beetles inhabiting dead aspen trees (Populus tremulae). For saproxylic beetles, tree retention at final cuttings can be expected to be especially valuable for species adapted to sun-exposed dead wood, a substrate that only rarely occurs in well managed forest stands. Therefore, the current evaluation was conducted as a comparison of species richness, species density (number of species per sample), assemblage composition and occurrence of individual species between clear-cuts, where aspen trees were retained, and closed forest stands with aspen trees. The study was conducted in central Sweden and the beetles were sampled by sieving of bark from CWD (coarse woody debris) of aspen. There was no significant difference in rarefied species richness between forest and clear-cut sites. Species composition differed significantly between the two stand types. Generalized linear mixed-effects models predicted the species density to be 34% lower in CWD objects in forest sites than on clear-cuts. This pattern could partly be explained by differences in CWD diameter, decay class and bark types between the two stand types (clear-cut/forest). Stand type was a significant predictor of occurrence in individual CWD objects for 30% of analysed individual beetle species. For all species except one, the variable stand type predicted higher occurrence on clear-cuts than in forest stands. To conclude, our results demonstrate that retention of aspen on clear-cuts contributes to population recruitment of a different assemblage of species than CWD within stands.     

Boreal bird community response to jack pine forest succession

The objective of this study was to examine bird communities in regenerating (5–25 years) and mature (40–100 years) jack pine (Pinus banksiana) forest in boreal Ontario. The study area was located near White River in north central Ontario with an area of 187,800 ha. We explored the response of bird community structure to stand age, and the influence of stand age on the distribution of individual species. We were interested in two principal questions. The first was how unique are the bird communities to specific age classes. If bird communities are highly specific to age classes then alterations to the age class distribution of the forest can have important impacts on the overall bird community composition and structure. The second question was how specific are individual species to age classes. Species that are highly specific to a single age class are expected to be highly sensitive to the amount and potentially the configuration of that age class on the landscape. We sampled birds for three breeding seasons. The number of bird species increased with stand age. Tree species composition did not change as stands aged, but there were distinctive changes in vegetation structure through succession. For example, the total amount of vertical vegetation structure increased significantly with age. More than half of the bird species examined were significant indicators of individual age classes. Blue-headed vireo, brown creeper, black-throated green warbler, golden-crowned kinglet, ovenbird and red-breasted nuthatch were all significant indicators of the mature age class. The bird assemblage of mature stands was significantly different from that of regenerating forest and within regenerating forest, 3–5-year-old stands contained a significantly different bird assemblage to that of 8–25-year-old regenerating forest. These results suggest that the distribution of forest age classes on the landscape is a critical element in determining habitat availability and therefore the viability of boreal bird populations in managed forests.     

Influence of fire intensity on structure and composition of jack pine stands in the boreal forest of Quebec: Live trees,understory vegetation and dead wood dynamics

North American jack pine (Pinus banksiana Lamb.) stands are generally characterized by an even-aged structure resulting from high intensity fires (HIF). However, non-lethal fires of moderate intensity (MIF), which leave behind surviving trees, have also been reported. The objectives of this study were two-fold: (1) assess the concurrent dynamics of live trees, understory vegetation and different types of coarse woody debris (CWD) during succession after HIF; and (2) document how MIF affects stand structure component dynamics compared to HIF. Stands affected by both HIF and MIF were selected. Tree characteristics and age structure, understory biomass, and CWD volume were assessed. Our results suggest that the structural succession of jack pine stands following HIF comprises three stages: young stands (<48 years), premature and mature stands (58–100 years) and old stands (>118 years). Canopy openness and jack pine density significantly decreased with time since HIF, while black spruce density and CWD volume significantly increased. The highest structural diversity was measured in the premature and mature stands. Compared to HIF, MIF increased mean jack pine basal area, decreased average stand density, delayed the replacement of jack pine by black spruce replacement in the canopy, decreased CWD volume, and significantly increased bryophytes mass. MIF increased the diversity of live trees and generally decreased CWD structural diversity. The study confirms the diversity of natural disturbance magnitude and successional processes thereby initiated. Thereafter, it appeared to be relevant for adjustment of disturbance emulating forest-management systems.     

麻池背油松天然林林分生长结构的研究

依据历年调查资料,对麻池背油松天然林的林分生长结构进行了研究.该林分的植物种类丰富;直径、树高、林龄等林分结构呈正态分布、近似正态分布和多峰状分布;近熟林的林分蓄积生长率为0.82%,材积生长量69年生时连年生长量仍大于平均生长量.由于林分密度过大,油松形状比值偏高,林木自然稀疏强烈,枯损率达32.7%,其中小径木占86.3%,病虫害感染枯死木占29.4%.油松天然更新不良,辽东栎不能成为更新树种,林下有青木杄、落叶松幼树分布.     

Vegetation structure, floristic composition and growth characteristics of Aucoumea klaineana Pierre stands as influenced by stand age and thinning

The coastal forest of Gabon abounds in monospecific secondary Aucoumea klaineana stands derived from natural regeneration after shifting cultivation. This paper aims to describe the changes in the structure and dynamics of these stands with age. It then assesses the impact of selective thinning in the upper storey on both structural and dynamic parameters.

The experiment consisted of 34 Permanent Plots in stands from establishment to more mature stages (ca. 50 years old). Thirteen plots (17–45 years old) were thinned. More than 80% of the removal came from supernumerary dominant A. klaineana.

A. klaineana represented 60% of the total density in stands ca. 15 years old but >90% of the dominant trees in older stands. The changes with age in the floristic composition of the unthinned stands showed three successional stages during which pioneer species associated with A. klaineana (from establishment to ca. 15 years) were progressively replaced by mature forest species.

Basal area increased and density decreased with age before reaching stable values at ca. 40–45 years. Mortality was very high in young stands but decreased in the older ones. Mortality generally affected small diameter individuals in the dominated storey. Diameter and basal area increments showed that the stand growth resulted from the growth of dominant A. klaineana. Diameter increments of A. klaineana were elevated during the first years of colonisation (1.9 cm/year) and were still ca. 0.7 cm/year for 50-year-old dominant trees.

Thinning did not increase the mortality of the dominant population. It favoured the individual growth of A. klaineana. The gain was substantial for dominated trees and small dominant trees (from 60 to 100%) but was lower for large dominant trees (ca. 25–30%). Therefore, stimulation of individual growth did not compensate for the loss of basal area at the stand level.     

Epiphytic lichen diversity in old-growth and managed Picea abies stands in Alpine spruce forests

In the last decades, a large body of literature has grown to evaluate the impact of forest management on epiphytic lichens in boreal coniferous forests. However, information is still lacking on coniferous forests of the Alps. This study compares lichen diversity between spruce forest stands of four successional stages: (1) young, (2) intermediate, (3) mature forests managed for timber production with a rotation cycle of 120–180 years, and (4) old-growth protected forests. The emphasis was placed on the occurrence of nationally rare and calicioid species (lichens and fungi traditionally referred to as Caliciales, known to be indicative of forest age and continuity). For each forest successional stage, four plots were selected. In each plot, 7 spruce individuals were surveyed for epiphytic lichens according to a standardised sampling method. Species richness increased from young to mature stands, while no difference was detected between mature and old-growth stands. This pattern was also confirmed for rare and calicioid species which are, however, more frequent in old-growth stands. Differences in species composition were also found between the different forest successional stages. Mature and old-growth plots slightly overlap, indicating that to some extent comparable lichen assemblages could be found in these stands. A nested pattern of species assemblages was found, old-growth stands hosting most of the species which were also found in stands belonging to the previous forest successional stages. Our results support the hypothesis that the management regime applied to spruce forests of the Italian Alps renders mature stands managed for timber production somewhat similar to old-growth stands as lichen habitat. However, we found a higher complexity in old-growth forests, and many species of conservation concern clearly preferred old-growth stands. In this perspective, a further prolongation of the normal cycle it is likely to be a most favourable conservation-oriented management to be recommended at least within protected areas and Natura 2000 sites, where conservation purposes should receive a high priority.     

Stand structure and successional trends in virgin boreal forest reserves in Sweden

Fire history and stand structure was examined in twelve virgin forest stands situated within forest reserves in northern Sweden. The selected stands represented fire refuges as well as different successional stages after fire. Six of the stands were dominated by Norway spruce (Picea abies L. Karst.), three were dominated by Scots pine (Pinus sylvestris L.), and three were dominated by hairy birch (Betula pubescens Ehrh.) or aspen (Populus tremula L.). In 3 of the southernmost stands, the average fire interval was 34 to 65 years during the late 1600s to late 1800s, but since 1888 no fires had occurred in any of the stands. The absence of fire disturbance since 1888 is probably caused by the fire suppression in the overall landscape. The standing volume of living trees ranged between 87 and 511 m³ ha⁻¹ while the volume of dead trees, including both snags and logs, ranged between 27 and 201 m³ ha⁻¹. The volume of dead trees constituted ca. 30% of the total stem volume. In the conifer dominated stands, there was a statistically significant relationship between total stem volume, including both living and dead trees, and site productivity. A comparison between the amount of dead and living trees indicated substantial changes in tree species composition in several stands. It is suggested that data on the amount of dead trees, especially logs, and its distribution over decay classes could be used to examine the continuity of certain tree species. All stands had a multi-sized tree diameter distribution, which in most cases was similar to a reversed J-shaped distribution. In general spruce was numerous in the seedling cohort and in small diameter classes, indicating that its proportion in the stands was stable, or was increasing at the expense of pioneer tree species such as pine, aspen and silver birch (Betula pendula Roth.). The most numerous species in the seedling cohort, rowan (Sorbus aucuparia L.), was almost totally missing in the tree layer, indicating a high browsing pressure preventing rowan seedlings from growing into trees. The general increase of spruce and the sparse regeneration of pioneer species, in the stands previously affected by fire, are discussed in relation to natural disturbance regimes, biological diversity and nature conservation policies. It is proposed that reintroduction of fire disturbance is a necessity for future management plans of forest reserves. Other management practices to increase species diversity within forest reserves are also discussed.