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.     

Modelling growth behaviour in monoculture in subtropical eastern Australia of two eucalypt species that differ in shade tolerance

Growth behaviour of the relatively shade-intolerant species Eucalyptus dunnii and the rather more shade-tolerant Corymbia citriodora was considered during several growth periods over 8–14 years of age in experimental plantation monoculture in subtropical New South Wales. Larger trees dominated growth in any stand; this was consistent with the principle that inter-tree competition in such forests is for light and is asymmetric in that larger trees shade smaller trees but the reverse does not occur. Once competition was established, competition-induced mortality of C. citriodora was less than that of E. dunnii. A model relating tree diameter growth rate to tree diameter, competitive position in the stand and stand basal area was developed. The model showed that smaller, hence partially shaded, trees in stands of C. citriodora were better able to maintain their growth rates than equivalent trees in stands of E. dunnii. It is hypothesised that this reflects the greater shade tolerance of C. citriodora. This pattern of growth led to a more even distribution of tree size classes across the stand in the more shade-tolerant species.     

基于竞争指数的杉木林分生长可视化模拟研究

以我国南方速生丰产林——杉木纯林为研究对象,采用改进过的Hegyi的简单竞争指数模型对杉木纯林间的竞争强度进行定量分析,并建立单木生长模型,再通过径阶模型将其应用于全林分,将单木生长模型研究扩展到全林分生长模型研究。同时,通过可视化模拟技术,为更加逼真的恢复和重建林分空间结构提供理论依据。     

Analysis of tree interactions in a mixed Mediterranean pine stand using competition indices

Studying species interactions in mixed forests allows us to assess their potential benefits and adapt current silvicultural tools developed in monospecific stands to multi-specific stands. We analyzed tree interactions in a Pinus halepensis Mill. and Pinus pinea L. mixed plantation using individual tree neighborhood models and competition indices that accounted for symmetric and asymmetric competition, to analyze whether the growth of each species was better explained by symmetric or asymmetric competition. We also split the competition indices into their intra- and interspecific forms, to test for competition effects on growth change based on competitor identity. Finally, we analyzed whether P. halepensis and P. pinea trees had different growth responses to competition. When calculating competition indices, we explored how spatial information and size of competitor trees contributed to the quantification of the process. Competition measurements were optimized to more precisely describe interactions. Results showed that the inclusion of competition indices generated important improvements in growth models. The main mode of competition was symmetric, which could be related to water restrictions typical of the Mediterranean climate. Considering competitor identity did not improve the growth models, while measurement without discriminating competitors by species generated more parsimonious models. P. halepensis and P. pinea trees had similar growth responses to competition, indicating that the two species cope with competition in similar ways. However, P. pinea showed lower average growth than P. halepensis in the period analyzed. Results suggest that preventing the onset of intense interspecific competition processes could help slow down the long-term replacement of P. pinea by P. halepensis and could have benefits for silvicultural management in systems with two species that share ecological niches but are capable of generating different goods and services.     

The effects of species mixture on the growth and yield of mid-rotation mixed stands of Scots pine and silver birch

The effect of tree species mixture on stand volume yield and on tree-species-specific diameter and height growth rates were analysed in managed mixed stands of Scots pine (Pinus sylvestris L.) and silver birch (Betula pendula Ehrn.).Data were obtained from 14 repeatedly measured stands located in Southern Finland on mineral soil sites with varying admixture of Scots pine and silver birch. Statistical analysis was carried out for studying the effect of species mixture on the development of stand characteristics. For the analysis, the plots were categorised into three groups (plot types) according to the species dominance. In order to analyse species-specific growth rates, individual-tree mixed linear growth models for tree diameter and height growth were developed for both tree species.The results clearly show that the yield of the managed mid-rotation, mixed stands was greater for stands dominated by Scots pine than for stands dominated by birch, and the stand volume increment decreased with an increasing proportion of silver birch. Analysis of diameter and height growth by tree species revealed that the main reason for this pattern is the negative impact of birch competition on the growth of pine trees. The increase in diameter of pine was clearly hampered if the proportion of birch was high. An abundance of birch also slightly decreased the growth in height of Scots pine, although the effect was less than on diameter growth. Species mixture did not affect the diameter growth of birch but did have a significant effect on height development. Height growth of birch was considerably greater in pine-dominated stands than in birch-dominated stands. In pine-dominated mixed stands, the height growth of birch was quite close to that of dominant pine trees, and birches can endure in competition with pines for light.The results apply for even-aged and single-storey managed stands, where stocking density and structure are controlled with pre-commercial and commercial thinnings. The results are not applicable to unmanaged mixed stands undergoing self-thinning. This study provides new information on mixed stands from a silvicultural perspective, which can be applied in decisions involving the management of mixed stands.     

Crown plasticity reduces inter-tree competition in a mixed broadleaved forest

The objective of the study was to analyse to which extent horizontal crown plasticity reduces inter-tree competition at stand scale, and how it relates to species growth strategy. Two components of crown plasticity defined at the individual tree level (crown shape distortion, CSD and crown displacement relative to stem, CRD) were analysed and their relative importance in the reduction of competition was quantified. Inter-tree competition at stand scale was estimated using spatial pattern analysis and crown overlap estimation. Measurements were performed in a mixed broadleaved stand of Western Europe. Crown plasticity was shown to regularise the spatial distribution of crowns in comparison with the corresponding stems and to reduce inter-tree competition by optimising space occupation. A significant reduction in crown overlap was observed, mainly due to CRD and secondarily to CSD. At the species level, CSD and CRD were positively correlated. In addition, both were negatively correlated with species shade-tolerance scores. In particular, three European temperate tree species showed contrasting responses, which were related to their known specific ecological strategies. Fagus sylvatica, known to be a highly shade-tolerant species, showed large crowns, low CSD and CRD, indicating a low plasticity and suggesting a strong competitive ability. At the opposite, Quercus sp., known to be clearly less shade-tolerant, exhibited a reduced growth, associated with high CSD and CRD, indicating high crown plasticity and probably a lower competitive ability. For this species, plasticity could be described as passive. Last, Carpinus betulus, known to have a relatively good shade-tolerance, showed a contrasting behaviour with high CSD and CRD and a strong presence in the stand, suggesting high crown plasticity and a good competitive ability. In that case, plasticity was described as rather adaptive.     

Spatially explicit competition in a mixed planting of Araucaria cunninghamii and Flindersia brayleyana

? Context

A 20-year-old Nelder wheel planted with hoop pine (Araucaria cunninghamii Aiton ex D.Don) and Queensland maple (Flindersia brayleyana F.Muell.) in 18 spokes and 8 rings represents nominal point densities of 3,580, 2,150, 1,140, 595, 305, 158, 82, and 42 stems/ha and offers an opportunity to examine competition and spatial interaction between these two species.

? Aims

This study aimed to evaluate the intraspecific and interspecific competition between two contrasting tree species and to determine the distance over which competition can be observed.

? Methods

Competition was estimated using Hegyi’s index, implemented using the Simile visual modeling environment, and calibrated using nonlinear least squares with PEST.

? Results

Interactions were detected between pairs of stems closer than D _ij ?<?40(d _i ?+?d _j ) where D is distance (in centimeters) and d is stem diameter (in centimeters diameter at breast height). F. brayleyana trees surrounded by A. cunninghamii trees experience negligible competition, whereas A. cunninghamii surrounded by F. brayleyana trees suffer strong competition.

? Conclusion

Forty times diameter offers a useful guide to the extent of competition in even-aged stands planted with these species. Competition can be observed empirically when pairs of trees are closer than 40 times the sum of their diameters, but the intensity of the competition may vary considerably with species.     

Stand structure, competition and growth of Scots pine (Pinus sylvestris L.) in a Mediterranean mountainous environment

The relationship between competition and tree growth was studied in four stands of Pinus sylvestris L. occurring in a continental Mediterranean mountain area (in the Guadarrama range, Spain), i.e., an uneven-aged stand, a stand with oak (Quercus pyrenaica Willd.) understorey, a plantation, and a mature even-aged stand. Competition was measured by a simple size-ratio distance-independent index and was negatively associated with tree diameter. This negative association was stronger in the uneven-aged, plantation and mature even-aged stands than in the stand with oak understorey. Competition was also negatively associated with current diameter increment. This relationship was moderately strong in the mature even-aged stand and weak in the uneven-aged stand and the plantation. In the uneven-aged and the mature even-aged stands, a weakly significant relationship was found between diameter growth and tree size, whereas these parameters were not associated in the stand with oak understorey. The competition index provided a better prediction of growth rate than the alternative use of diameter. Both diameter and basal area growth were greater in the uneven-aged than in the even-aged stands.     

Models for predicting height and diameter of individual trees in young Picea abies (L) karst. stands

Models for predicting height and diameter of individual trees in young Picea abies (L.) Karst. stands were developed. Data collected in a large survey of young forest stands in Sweden (the HUGIN young stand survey) were used in the construction of the models. Models were developed both with and without competition indices included. When constructing the competition indices trees within three metres from the subject tree were regarded as competitors. Functions with competition indices included (distance dependent) will be useful in analyses of the development of stands with different stand structure, whereas functions without competition indices (distance independent) will be useful in systems for long‐term forecasts of yield.     

不同类型毛竹林植物物种多样性研究

通过对不同类型毛竹林植物物种多样性的研究,结果表明:毛竹林内物种是丰富的,但较大部分物种是脆弱的;乔木层植物物种多样性指数与林下木本植物物种多样性指数相关显著,与草本植物物种多样性指数相关不显著,乔木层物种多样性指数小于林下植物物种多样性指数;木本植物物种多样性指数,竹阔混交林＞竹针混交林＞毛竹纯林,粗放经营竹林＞中等集约经营竹林＞集约经营竹林;林下草本植物物种多样性指数,竹针混交林＞毛竹纯林＞竹阔混交林,中等集约经营竹林＞粗放经营竹林＞集约经营竹林;竹阔混交林向毛竹纯林转型时,林下木本植物部分丧失,而草本植物将可能增加,也可能减少.     

Spatial point-pattern analysis for detecting density-dependent competition in a boreal chronosequence of Alberta

In the boreal forest of Alberta, fire and wind often open gaps in the canopy where late-successional species can establish and over time cause a shift in the species distribution from deciduous (e.g., trembling aspen) dominated to mixedwood, to shade-tolerant conifer (e.g., white spruce) dominated stands. This study attempted to understand the change of density-dependent competition in a boreal chronosequence and the role of tree competition in affecting stand structure and mortality. Four 1-ha stem-mapped plots were established to represent a chronosequence comprised of aspen dominated, mixedwood, and spruce dominated stands in Alberta. Second order spatial point-pattern analysis using Ripley's K(t) function showed that intraspecific competition is a prevailing force causing conspecific tree mortality and thus shaping the stand structure. The results of bivariate K(t) function analysis did not reveal sufficient evidence of interspecific competition. This suggested that competitive interaction among heterospecific trees was not strong enough to cause significant tree mortality, but the analysis of marked correlation function revealed that interspecific competition could have a negative impact on tree growth. This study highlights the importance of density-dependent competition in understanding stand dynamics of boreal forests over succession.     

Stand characteristics of mixed oriental beech (<Emphasis Type="Italic">Fagus orientalis</Emphasis> Lipsky) stands in the stem exclusion phase,northern Iran

The structure of forest stands changes through developmental phases. This study is carried out in the unmanaged, oriental beech (Fagus orientalis Lipsky) stands in the north of Iran. The aim of this research was to quantify structural characteristics of stands in the stem exclusion phase using common structural indices, which include mingling, tree–tree distance, stem diameter, and tree height differentiation. According to our measurements from three stands, naturally regenerated stands tend to be mixed in species composition have slightly heterogeneous diameter distributions and uniform tree height. The average distance between trees was 3.3 m. Stocking volume of the stands had an average of 540 m³ ha^?1 and 412 stem ha^?1. Dead wood volume was 24 m³ ha^?1, and as a standing volume, the most frequent species in dead wood pool was oriental beech (F. orientalis) (48 %). The common form of dead trees was snag (41 %). The mean value of mingling and tree-to-tree interval indices revealed that beech was mixed intensively with hornbeam and appears to be a more successful competitor for space and light compared with hornbeam; moreover, we found relatively high evidence of inter-species competition in this phase. A better understanding of stand characteristics in the stem exclusion phase as a critical part of the natural dynamics of forest ecosystems could facilitate predictions about the future changes within the stand.     

Five-year radial growth of red oaks in mixed bottomland hardwood stands

We studied the relationships among 5-year radial (diameter and basal area) growth of red oak (genus Quercus, subgenus Erythrobalanus) crop trees and predictor variables representing individual tree vigor, distance-dependant competition measures, and distance-independent competition measures. The red oaks we examined are representative of the commercially and ecologically important oak species of the bottomland hardwood forests of the southeastern US. The crown class score, a quantitative measure of crown class and tree vigor, performed best in accounting for the variability in tree diameter growth. Plot-level variables failed to account for a significant proportion of the variability in tree radial growth. The basal area of the first-order neighbors that were taller than the crop trees and located within 2.4 times the mean overstory crown radius had the highest negative correlation with crop tree 5-year radial growth. Red oaks were a major part of these competitors and likely exerted the greatest competitive pressure. However, crop tree radial growth was positively associated with the basal area of the red oaks which were indirect (second order) neighbors and which were taller than the crop trees. It is possible that indirect neighbors do not compete with the crop trees, but they likely compete with the direct competitors of the crop trees, thus having an indirect positive influence on crop tree growth. Such reasoning is consistent with previously observed spatial dependence up to four times the mean overstory crown radius. The findings may have implications for thinning hardwoods stands and crop tree management in that foresters need to take into account (1) oak intra-genus competition, (2) the negative competitive effect of direct neighbors, and (3) the potentially positive effect of the indirect neighbors, the competitors’ competitors.     

Prediction of individual tree growth in managed stands of mixed Picea abies (L) Karst,and Betula pendula Roth & Betula pubescens Ehrh.

The individual tree growth in stands of mixed Norway spruce (Picea abies (L.) Karst.) and birch (Betula pendula Roth & Betula pubescens Ehrh.) is estimated using basal area and height growth functions for each species separately. The individual tree growth models are distant dependent and the number, size and proximity of neighbours are expressed as size‐ratio competition indices. The competition indices were calculated using a basal area factor gauge to define competitors. The tree growth functions are based on data from nine mixed stands of Norway spruce and birch. The recursive multivariate regression approach is used. The growth functions have standard deviation about the function/standard deviation about the mean (sf/sm) values between 31 and 61% and the evaluations made with root mean square error (rmse) give estimates which vary between 8 and 45 % of the observed mean value. These values are comparable with the precision reported in other studies. In the present investigation the distant dependent indices are important independent variables. It seems suitable to describe the change in growth conditions from retarded to released growth by means of size ratio competition indices. For birch, a positive effect on growth is obtained the more the total competition consists of Norway spruce. For Norway spruce a negative effect on growth is obtained the more the total competition consists of Norway spruce. The lower competitors have a positive effect on the growth of the spruce trees. The interpretation should be that it is better to have a small competitor than a larger one, not that small competitors as such have a positive effect on growth of Norway spruce.     

The interaction between competition in interior Douglas-fir plantations and disease caused by Armillaria ostoyae in British Columbia

Interior Douglas-fir trees in plantations were assessed for size differences related to the level of diseased neighbours infected with Armillaria ostoyae. The four Douglas-fir stands studied ranged from 25- to 34-year-old, and represented the oldest accessible planted stands in the Interior Cedar Hemlock (ICH) ecosystem in British Columbia. Twenty-three to 25, 10-m radius plots were established in each stand. The spatial coordinates, total height, and diameter at breast height of all live and dead trees in the plots were recorded. Subject trees whose competitors were contained in the 10-m radius plots were also identified. Trees were pulled out of the soil using a mechanical excavator and the root systems were surveyed for evidence of infection by A. ostoyae. Stem disks were taken from each tree at 1.3 m above the ground for a determination of basal area. Increasing proportion of diseased trees in the plots resulted in less total plot basal area, but did not affect the mean basal area or height. Individual subject tree basal area was negatively related to the level of disease in surrounding competitors, opposite to expectations; however, diseased subject trees had reduced height and basal area compared to disease-free subject trees. Increasing competition reduced both the height and basal area of the trees, while regular distribution of all trees increased both total and mean plot basal area but not height. Disease incidence at the plot level and in individual subject trees was mainly affected by the neighbourhood conditions in which it grew, and was also related to disease intensity in the tree root systems. Although disease may alter resource partitioning among trees, the utilization of these resources is mostly limited by the increasing disease incidence as the stands age, the higher probability of larger trees being diseased with time, the occurrence of dead trees in clumps, and the high probability that dead trees will eventually infect live neighbours. The widespread belowground incidence of A. ostoyae in the ICH, its rapid colonization of stumps, and its wide host range can reduce site potential in managed stands.     

Stand age and habitat influences on salamanders in Appalachian cove hardwood forests

We surveyed cove hardwood stands aged 15, 25, 50, and ≥85 years following clearcutting in the southern Appalachian Mountains of northern Georgia to assess the effects of stand age and stand habitat characteristics on salamander communities using drift-fence array and pitfall methodologies from May 1994 to April 1995. Over a 60,060 pitfall trapnight effort, we collected 3937 salamanders represented by Desmognathus aeneus, Desmognathus monticola, Desmognathus ocoee, Desmognathus quadramaculatus, Eurycea bislineata, Gyrinophilus porphyriticus, Pseudotriton ruber, Plethodon glutinosus, Plethodon serratus, and Notophthalmus viridescens. Analysis of covariance with pitfall array to stream distance as the covariate showed that salamander species richness and diversity measures and numbers of Desmognathus aeneus and Desmognathus ocoee were highest in stands ≥85 years. Eurycea bislineata and Plethodon glutinosus were more abundant in stands ≤50 years old than in stands ≥85 years. Within cove hardwood stands, species richness and diversity measures and relative abundances of Desmognathus spp. and Gyrinophilus porphyriticus were negatively correlated with distance to stream. Species richness and diversity were positively correlated to amounts of emergent rock. Species richness, diversity and relative abundances of Desmognathus spp. were correlated with basal area within stands and extent of connected mesic, cove hardwood habitat and amount of cove habitat within 1 km radius among stands. Eurycea bislineata was negatively correlated with landform index, a measure of surrounding landform sheltering, and Plethodon glutinosus was positively correlated with elevation in cove hardwood stands. Our research indicates stand age is an important factor in explaining the abundance and community composition of salamanders in southern Appalachian cove hardwood communities. Because southern Appalachian woodland salamander communities are slow to recover and are substantially changed following disturbances such as clearcutting, populations in small, isolated cove hardwood stands might be more vulnerable to extirpation or may require longer recovery times than those in larger coves. Managers may need to assess habitat features such as cove extent and habitat connectivity to minimize impacts on these taxa by forest management activities in southern Appalachian cove hardwood communities.     

Using a retrospective dynamic competition index to reconstruct forest succession

Understanding forest dynamics and stand structures is crucial for predicting forest succession. However, many forests have been altered due to century-long land-use practices, which complicates the reconstruction of past and current successional trajectories. For a better understanding of successional processes, we suggest studying the intra- and interspecific competition among single trees across time. We introduce a tree-ring based competition index to reconstruct the competitive dynamics of individual trees over time. This new retrospective dynamic competition index combines a temporal and a spatial component by calculating the yearly ratio between the basal area increments (bai) of the neighbouring trees and the subject tree. The new index is applied to mixed Scots pine (Pinus sylvestris L.) and pubescent oak (Quercus pubescens Willd.) stands in the inner-Alpine dry-valley Valais, for which a change in species composition is hypothesised. The aim is to analyse current stand structures in terms of recent changes in the competitive interactions at the single tree level and to relate these competitive dynamics to land-use change and increasing drought due to climate change. On five plots, the positions of 456 trees were recorded and increment cores were taken to derive bai data. The individual dynamic competition index curves were aggregated in clusters, which define typical patterns of competitive dynamics in both tree species. A large percentage of the trees (87% in oak, 70% in pine) were clustered into a group of trees with constant competition at a relatively low level. However, a smaller group of pines (20%) had recently faced increasing competition. In addition, stand structure analyses indicated a change towards a higher proportion of oak. This change in the competitive ability between oak and pine was found to be related to drought, in that oak had a competitive advantage in dry years. Furthermore, the high proportion of dead branches in pines with decreasing competitive abilities indicated increasing competition for light as a consequence of natural development towards a later successional stage that favours the more shade-tolerant oak. The new retrospective dynamic competition index proved to be promising in studying forest succession. The tree-ring based method allows us to identify changes in the competitive ability of single trees with a high temporal resolution and without repeated assessments.     

基于分层的典型中亚热带天然阔叶林林木竞争关系研究

[目的]通过分析不同林层林木之间的竞争情况,以期揭示林木间相互竞争的类型、林木受到的竞争压力与林木垂直分布之间的关系,为森林资源的保护与利用提供科学依据。[方法]在福建万木林自然保护区内设5块典型中亚热带天然阔叶林样地,采用Hegyi的竞争指数,按分层和不分层两种情形研究林木间的竞争强度。[结果]在1—5号样地,当取样半径超过4 m后,林木间的竞争强度随取样半径的增大而减小的趋势变缓,4 m是分析林木竞争的适宜半径;以4 m为半径分析林木间的竞争,发现有一半以上的组成树种只发生种间竞争,剩余同时发生种间和种内竞争的树种,其竞争木构成中数量占优势的普遍是与其相异的树种,且种间竞争强度普遍强于种内竞争;各样地的乔木层都可以划分出由高到低的3个林层,在平均水平,越高层的林木距离竞争木越远、在竞争木的组成中较矮小的第3林层林木数量也越多;越高层的林木,平均每株对象木与其全部竞争木之间的竞争压力越小。[结论]在调查的典型群落中,普遍发生的是种间竞争,而林木受到的竞争压力及其竞争木的组成都与它们的垂直分布关系密切,在森林资源保护和经营中应对垂直分布予以更多的关注;乔木层中的优势树种未发生激烈种间或种内竞争;可以发生激烈种间竞争的乔木树种有米槠、庆元冬青、少叶黄杞和杜英,不存在特定的竞争木与它们竞争,它们与竞争木相近、主要分布在第2或第3林层;种内竞争过于激烈的有山杜英、华杜英、山黄皮、野含笑和羊舌树,也主要分布在第2或第3林层。     

Canopy gap dynamics and development patterns in secondary Quercus stands on the Cumberland Plateau, Alabama, USA

Forest disturbances of various spatial extents and magnitudes shape species composition, structure, and stand development patterns. The disturbance regimes of most complex stage hardwood stands of the deciduous forests of eastern North America are typified by asynchronous and localized disturbance events. The overwhelming majority of gap-scale disturbance studies in hardwood forests of the region have analyzed late-successional stands. As such, there is a paucity of data on gap dynamics in hardwood stands prior to a complex developmental stage. We quantified biophysical characteristics of 60 canopy gaps in secondary Quercus stands on the Cumberland Plateau in Alabama to analyze gap-scale disturbance processes in developing systems. We found most gaps (90%) were caused by the removal of a single tree. Of the three gap formation mechanisms, snag-formed gaps were most common (40%). However, based on the number of uprooted and snapped stems we speculate that wind was also an important disturbance agent in these stands. Gap size and shape patterns were similar to what has been reported in other hardwood forests of the southern Appalachian Highlands. We did not find differences in gap size or shape based on formation mechanisms; a finding that may be related to the number of single-tree gap events. Gaps projected to close via subcanopy recruitment were significantly larger than those projected to close through lateral crown expansion. Most gaps (65%) were projected to close by lateral crown expansion of gap perimeter trees. However, the number of gaps projected to fill by subcanopy recruitment indicated the stands were approaching a transition in their developmental stage. Gap-scale processes modify residual tree architecture and stand structure. Through time these alterations result in progressively larger gaps, eventually reaching a size when most will fill by subcanopy recruitment, thus marking the complex stage of development. Gap capture by Quercus was restricted to relatively xeric sites that did not contain abundant shade-tolerant mesophytes in the understory. However, the majority of gaps contained abundant subcanopy Fagusgrandifolia, Acer saccharum, and Acer rubrum leading us to project that the forest will undergo a drastic composition shift under the current disturbance regime. Liriodendron tulipifera was projected to capture several relatively small gaps illustrating the role of topography on gap closure mechanisms.     

How to quantify forest management intensity in Central European forests

Existing approaches for the assessment of forest management intensity lack a widely accepted, purely quantitative measure for ranking a set of forest stands along a gradient of management intensity. We have developed a silvicultural management intensity indicator (SMI) which combines three main characteristics of a given stand: tree species, stand age and aboveground, living and dead wooden biomass. Data on these three factors are used as input to represent the risk of stand loss, which is a function of tree species and stand age, and stand density, which is a function of the silvicultural regime, stand age and tree species. Consequently, the indicator consists of a risk component (SMI_r) and a density component (SMI_d). We used SMI to rank traditional management of the main Central European tree species: Norway spruce (Picea abies [Karst.] L.), European beech (Fagus sylvatica L.), Scots pine (Pinus sylvestris L.), and oak (Quercus robur L. and Quercus petraea L.). By analysing SMI over their whole rotation period, we found the following ranking of management intensity: oak<beech<pine?spruce. Additionally, we quantified the SMI of actual research plots of the German Biodiversity exploratories, which represent unmanaged and managed forest stands including conifer forests cultivated outside their natural range. SMI not only successfully separate managed from unmanaged forests, but also reflected the variability of forest management and stand properties across the entire sample and within the different management groups. We suggest using SMI to quantify silvicultual management intensity of stands differing in species composition, age, silvicultural system (even-aged vs. uneven-aged), thinning grade and stages of stand conversion from one stand type into another. Using SMI may facilitate the assessment of the impact of forest management intensity on biodiversity in temperate forests.