Bottomland hardwood forest recovery following tornado disturbance and salvage logging

Catastrophic wind events, including tornado, hurricane, and linear winds, are significant disturbances in temperate forested wetlands. Information is lacking on how post-disturbance salvage logging may impact short and long-term objectives in conservation areas where natural stands are typically managed passively. Woody regeneration and herbaceous cover were assessed for three years in a bottomland hardwood forest across a gradient of damage from an F4 tornado, with and without subsequent salvage logging. Soil disturbance intensity and recovery associated with salvage logging within wind-disturbed sites were also assessed. Woody stem density and proportion of potential overstory species (species with the potential to occupy a position in the canopy) increased as a function of wind disturbance intensity. Stem density, proportion of overstory trees, or species diversity did not differ between wind + salvage and wind-disturbed-only plots. Significant dissimilarity occurred among soil disturbance classes within salvaged sites. By the third growing season, vegetation in soil disturbance classes in wind + salvage areas was converging toward undisturbed conditions and bottomland hardwood forest recovery was underway in all vegetation disturbance types and soil disturbance classes. Post-tornado salvage logging, applied judiciously, may contribute to microsite and vegetation diversity.     

Canopy light transmittance in Douglas-fir--western hemlock stands

We measured vertical and horizontal variation in canopy transmittance of photosynthetically active radiation in five Pseudotsuga menziesii (Mirb.) Franco-Tsuga heterophylla (Raf.) Sarg. (Douglas-fir-western hemlock) stands in the central Cascades of southern Washington to determine how stand structure and age affect the forest light environment. The shape of the mean transmittance profile was related to stand height, but height of mean maximum transmittance was progressively lower than maximum tree height in older stands. The vertical rate of attenuation declined with stand age in both the overstory and understory. A classification of vertical light zones based on the mean and variance of transmittance showed a progressive widening of the bright (low variance and high mean) and transition (high variance and rapid vertical change) zones in older stands, whereas the dim zone (low variance and mean) narrowed. The zone of maximum canopy surface area in height profiles, estimated by inversion of transmittance profiles, changed from relatively high in the canopy in most young stands ("top-heavy") to lower in the canopy in older stands ("bottom-heavy"). In the understory, all stands had similar mean transmittances, but the spatial scale of variation increased with stand age and increasing crown size. The angular distribution of openness was similar in all stands, though the older stands were less open at all angles than the younger stands. Understory openness was generally unrelated to transmittance in the canopy above. Whole-canopy leaf area indices, estimated using three methods of inverting light measurements, showed little correspondence across methods. The observed patterns in light environment are consistent with structural changes occurring during stand development, particularly the diversification of crowns, the creation of openings of various sizes and the elaboration of the outer canopy surface. The ensemble of measurements has potential use in distinguishing forests of differing ages that have similar stature.     

Predicting understory maximum shrubs cover using altitude and overstory basal area in different Mediterranean forests

In some areas of the Mediterranean basin where the understory stratum represents a critical fire hazard, managing the canopy cover to control the understory shrubby vegetation is an ecological alternative to the current mechanical management techniques. In this study, we determine the relationship between the overstory basal area and the cover of the understory shrubby vegetation for different dominant canopy species (Pinaceae and Fagaceae species) along a wide altitudinal gradient in the province of Catalonia (Spain). Analyses were conducted using data from the Spanish National Forest Inventory. At the regional scale, when all stands are analysed together, a strong negative relationship between mean shrub cover and site elevation was found. Among the Pinaceae species, we found fairly good relationships between stand basal area and the maximum development of the shrub stratum for species located at intermediate elevations (Pinus nigra, Pinus sylvestris). However, at the extremes of the elevation-climatic gradient (Pinus halepensis and Pinus uncinata stands), stand basal area explained very little of the shrub cover variation probably because microsite and topographic factors override its effect. Among the Fagaceae species, a negative relationship between basal area and the maximum development of the shrub stratum was found in Quercus humilis and Fagus sylvatica dominated stands but not in Quercus ilex. This can be due to the particular canopy structure and management history of Q. ilex stands. In conclusion, our study revealed a marked effect of the tree layer composition and the environment on the relationship between the development of the understory and overstory tree structure. More fine-grained studies are needed to provide forest managers with more detailed information about the relationship between these two forest strata.     

Height-to-diameter ratios with temporal and dendro/morphometric variables for Brazilian pine in south Brazil

Height-to-diameter ratios(HD) are an important measure of the stability,density and competition of forest stands.It reflects the vertical growth of the trees,the vulnerability of the forest canopy structure and influences volumetric production.HD ratios vary according to tree size,availability of resources for growth,stand density and species composition.Data were taken from 210 trees and a regression technique of generalized linear models for the HD ratio applicable for forest structure conservation was developed.The objective of this study was to model the HD ratios of dominant and co-dominant trees of Araucaria angustifolia according to morphometric,dendrometric,annual diameter increment,stand density,and age variables in three sites in southern Brazil.The results show that the HD ratio decreases with increasing age,crown area and basal area,and increases with stand density and annual diameter increment.Accuracy of the developed equations was demonstrated by the values of deviation,Bayesian and Akaike criteria.The results are of interest to forest managers since they make decisions about silvicultural operations.Growth continuity and forest production indicate that any intervention should be directed at younger trees of smaller sizes,and that one of the main management factors for stand stability and growth is the formation of the stand and its capture of light.     

林分密度对蒙古栎林下草本植物多样性的影响研究

以内蒙古赤峰市阿鲁科尔沁旗沙日温都栎林自然保护区蒙古栎林为主要研究对象,研究不同林分密度对林下草本植物多样性的影响。结果表明,研究区内共有13科、14属、16种草本植物;Simpson指数、Shannon-Wiener指数、Pielou均匀度指数和物种丰富度指数均随着林分密度的增加呈先增大后减小的趋势,当林分密度为750株/hm²时,林下草本植物多样性指数达到最大值;林分密度与Shannon-Wiener多样性指数、Pielou均匀度指数、Simpson多样性指数、物种丰富度指数、树高、胸径均呈极显著负相关,与郁闭度呈极显著正相关,与枝下高不相关。综上所述,最合理的蒙古栎林密度为750株/hm²,该密度下最有利于蒙古栎林及林下草本植物生长发育。     

SMALL-DIAMETER KOREAN PINES AND THEIR ROLES INNATURAL KOREAN PINE FOREST

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Trade-offs between seedling growth,thinning and stand stability in Sitka spruce stands: a modelling analysis

The forest industry is increasingly adopting alternative silvicultural systems, involving regeneration beneath an existing forest canopy, rather than clear-felling and replanting. To apply these silvicultural systems in windy regions such as Britain and Ireland, it is essential that the interactions between thinning intensity, stand stability and seedling growth are properly understood. Here, we present a modelling analysis of the three key relationships between: (i) stand density and the proportion of incident radiation transmitted through a forest canopy as a stand is thinned; (ii) transmitted radiation and seedling growth, and (iii) stand density and stand stability. These relationships were examined using separate models of radiative transfer (MAESTRO), seedling growth, and stand stability/wind risk (ForestGALES). Output from the three models was synthesised to calculate whether a given stand thinned to a pre-defined stability limit would allow sufficient light to penetrate the canopy for seedling growth. A minimum transmittance of 20% was identified as a requirement for seedling growth, which corresponds to removing 45% of stand basal area. A thinning of this intensity left some stands susceptible to unacceptable wind damage, especially in old or previously thinned stands on soils where rooting is impeded. The results emphasised the fact that rooting conditions, thinning history and age of intervention are major constraints on the silvicultural options. In general, older stands are not suitable for conversion to continuous cover forestry (CCF) systems, and the transformation process should begin at pole stage, when heavy thinning does not leave the stand unstable. The analysis approach used here illustrates the potential for combining models to address complex forest management issues.     

不同林型对林下光照和辐射消减的影响

在中山市长江库区水源林市级保护区内选择4种不同的林分类型,进行每木检尺和冠层影像拍摄,处理得到林下光照因子和辐射消减因子,分析不同林型对林下光照和辐射消减的影响。结果表明:不同林型对林冠结构和林下光照影响显著,针叶林与针阔混交林的林下光照接近,林冠较差,沟谷季雨林与常绿阔叶林接近,林冠较好;辐射消减在不同林型间差异显著,针叶林和针阔混交林辐射消减较差,常绿阔叶林和沟谷季雨林辐射消减较好;林下光照与辐射消减相关性强,特别是两个林冠结构因子与辐射消减关系密切。在进行林分改造时,建议选择叶片较大较厚,冠幅大而优美的常绿阔叶树种进行更新,天然起源的林分与常绿阔叶林的林冠结构较好,有效维持了森林生态系统稳定。     

Changes in the spatial structure of oak carbon-based secondary compounds after pine harvesting

In natural plant populations, leaf polyphenols show high intraspecific variation that occurs both temporally and spatially. Leaf phenolics may be induced by diverse ecological factors such as light, nitrogen availability or herbivory attack. Both light and nitrogen availability can show spatial structure in forested stands, meaning that they each have a high degree of autocorrelation, which can determine the appearance of spatial structure in leaf polyphenols. However, the availability of these resources may be drastically changed by forest disturbance, and little is known about the effect of forest disturbance on the spatial pattern and scale of leaf secondary compounds. We hypothesise that the spatial structure of leaf polyphenols in understory vegetation will disappear due to forest harvesting, because these compounds depend on light availability, yet it will remain unaltered for those compounds that either depend on the availability of other resources or are under major genetic control. The study was performed in young pedunculate oak (Quercus robur) populations growing either under a pine canopy (Pinus pinaster) stand or in a pine harvested stand in NW Spain. The spatial structures of green and senescent leaf polyphenols, tannins, non-tannin polyphenols and nitrogen were analysed in both stands using geostatistical analysis. The spatial structures observed for green and senescent leaf polyphenols and tannins in the forested stand disappeared in the harvested stand. However, non-tannin polyphenols, as well as nitrogen, showed spatial structure in both stands. Understanding these changes may be important for the successful recovery of native oak populations growing under pine forests in NW Spain, one of the priorities of the local government. Our results showed that changes in the concentration of leaf secondary compounds after disturbance may be accompanied by differences in their spatial properties, which may have important consequences for ecosystem function.     

Predicting natural forest regeneration: a statistical model based on inventory data

Regenerating forest stands through natural seedlings is a commonly accepted silvicultural strategy in Germany. To plan for and increase natural regeneration within a given stand, foresters must be able to estimate whether the stand in its current state can produce sufficient saplings to replace the overstory. In this paper, we present two approaches to building a model that can estimate the probability of natural regeneration occurring, based on variables that are typically readily available from forest inventories. To estimate model parameters we used the large database of the third National Forest Inventory, which covers forest stands and sites across the whole of Germany, as well as weather and soil data. We examined how these variables impact the emergence of natural regeneration, ultimately fitting a model that can predict the occurrence of natural regeneration in 72% of cases. The influence of the variables on the predicted occurrence of natural regeneration was mixed, with most stand variables contributing only minor impact and most likely influencing natural regeneration via complex interactions. The exception was vertical structure (number of stand layers), which accounted for a large proportion of the goodness-of-fit of the model. An important finding was that forest ownership structure is a key variable for the prediction of the presence of regeneration. Data from this study support the assumption that some forest owners manage their stands in a way that fosters natural regeneration.     

森林冠层结构的生态学研究现状与展望

林冠是树木光合作用的主要层次,林冠结构造成的林下光环境差异对于植物的生长和群落的更新演替有着重要的生态学意义。冠层结构的重要作用已引起全球生态学家的高度重视,国际科学界已举行了三届国际冠层大会,认为合作并发展全球冠层研究计划是必须的。文章较系统地介绍了国内外冠层结构、林内环境、林下植被组成及土壤环境之间的一系列相互关系的研究进展,并将近年来有关森林冠层结构最新研究方法作简单介绍,为今后的同类研究提供参考。     

Small-scale variation of vegetation in a mixed forest understorey is partly controlled by the effect of overstory composition on litter accumulation

We investigated how richness and composition of vascular plant species in the understory of a mixed hardwood forest stand varied with respect to the abundance and composition of the overstory. The stand is in central Spain and represents the southernmost range of distribution of several tree and herbaceous species in Europe. Understory species were identified in 46 quadrats (0.25 m²) where variables litter depth and light availability were measured. In addition, we estimated tree density, basal area, and percent basal area by tree species within 6-m-radius areas around each plot. Species richness and composition were studied using path analysis and scale-dependent geostatistical methods, respectively. We found that the relative abundance of certain trees species in the overstory was more important than total overstory abundance in explaining understory species richness. Richness decreased as soil litter depth increased, and soil litter increased as the relative proportion of Fagus sylvatica in the overstory increased, which accounted for a negative, indirect effect of Fagus sylvatica on richness. Regarding understory species composition, we found that some species distributed preferentially below certain tree species. For example, Melica uniflora was most frequent below Fagus sylvatica and Quercus petraea while the increasing proportion of Q. pyrenaica in the overstory favored the presence of Cruciata glabra, Arenaria montana, Prunus avium, Conopodium bourgaei, Holcus mollis, Stellaria media and Galium aparine in the understory. Overall, these results emphasize the importance of individual tree species in controlling the assemblage and richness of understory species in mixed stands. We conclude that soil litter accumulation is one way through which overstory composition shapes the understory community.     

Differential overstory leaf flushing contributes to the formation of a patchy understory

Understory individuals were found to form patches in a 100-year-old deciduous broad-leaved forest. The closed forest canopy was uniform, and so the light conditions at various locations across the forest floor differed little after the leaf flush of the overstory. To explain the distribution pattern in the understory, a hypothesis was proposed: in spring, the forest floor is divided into patches according to the timing of leaf flush of the overstory individuals, and the light conditions are more favorable for understory plants under the crowns of trees with later-flushing leaves. In the plot, three groups of early, intermediate, and late, were recognized in the overstory concerning the timing of leaf flush. As for the start of leaf flush, a difference of 31.6 days was recognized among tree species, and for the end of leaf flush, a difference of 40.3 days. In the spring of 1998, the relative photosynthetic-photon-flux density under an intensively studiedCastanea crenata tree (late-flushing species) usually showed higher values than that under a similarly studiedAcer mono tree (early-flushing species). Analysis of the spatial-distribution pattern using Morisita’s1δ index revealed that the understory community had an aggregated distribution. In the overstory, the late- and the intermediate-flushing-species groups showed aggregated distributions, while the early-flushing-species group showed random distribution. Spatial correlation between the understory and the overstory was analyzed by using Morisita’sRδ index. The distribution of whole understory community spatially co-occurred with that of the late-flushing-species group of the overstory. In contrast, the understory community was less developed below the members of the early-flushing-species group of the overstory. We consider that the data presented here support our hypothesis, and we suggest that the growth and survival of understory individuals were promoted in the places receiving light for long periods in spring.     

Effects of single-tree selection harvesting on hymenopteran and saproxylic insect assemblages in the canopy and understory of northern temperate forests

Insects respond to changes in microhabitat caused by canopy disturbance, and thus can be used to examine the ecological impacts of harvesting. Single-tree selection harvesting is the most common silvicultural system used to emulate local small-scale natural disturbance and maintain uneven-aged forest structure in temperate forests. Here, we test for differences in richness, abundance, and composition of hymenopteran and saproxylic insect assemblages at four different taxon levels (selected insect orders; and all hymenopteran families, and braconid subfamilies and morphospecies) between the canopy and understory of unharvested and single-tree selection harvested sites in a northern temperate forest from central Canada. Harvesting had no effect on insect assemblage richness, composition or abundance at the three highest taxon levels (order, family and subfamily). Similarly, richness and abundance at the lowest-taxon level (braconid morphospecies) were similar, although composition differed slightly between unharvested and harvested stands. Insect assemblages were vertically stratified, with generally higher abundance (for Diptera, Hymenoptera, some hymenopteran families and braconid subfamilies) and richness (for braconid morphospecies) in the understory than the canopy. In particular, composition of the braconid morphospecies assemblage showed relatively low similarity between the understory and canopy. Single-tree selection harvesting appears to influ-ence wood-associated insect taxa only subtly through small changes in community composition at the lowest taxon level, and thus is recommended as a conservative approach for managing these northern temperate forests.     

Comparison of stand dynamics after dieback caused by pine wilt disease among pine forests with different management regimes in western Japan

In pine forests damaged by pine wilt disease, in western Japan, the effect of protection regimes of pine trees on the stand dynamics were examined in the following four stands: (1) lightly damaged stand (age 30–40 years) with no procedure in operation for protecting pine trees; (2) severely damaged stand (age 30–40 years) with no procedure in place for protecting pine trees; (3) severely damaged stand (age 50 years) with a selective cutting of infected trees; (4) severely damaged stand (age 30–40 years) with a selective cutting of infected trees. All the stands had been abandoned before the pine wilt disease damage. The understory structure of the severely damaged stand with no protection procedure was similar to that of the lightly damaged stand. Frequent invasion by tree species and acceleration in the growth of understory trees occurred after the dieback in the selective cutting stand. These results suggest that a deficiency in the canopy layer caused by the dieback resulted in low disturbance intensity in the early stages after the dieback, but the selective cutting increased the intensity by the reduction in the understory as well as the canopy layer. The intensity of the disturbance in the selective cutting stands was larger in the younger stand because it had a higher density of selectively cut pine trees. The different stand structure of pine forests occurred after the dieback because the intensity of the disturbance varied as a result of the selective cutting operation and the stand age.     

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.     

Influence of tree species on understory vegetation diversity and mechanisms involved—A critical review for temperate and boreal forests

Tree species composition is a primary attribute of forest ecosystems, and is often manipulated by silvicultural practices. Forest management to diversify tree species is now being promoted to favor biodiversity. To assess the soundness of this policy we reviewed and analyzed the literature on the relationship between tree species composition and floristic diversity, including the mechanisms involved therein. Coniferous forests generally provide less diversified vascular understories than broadleaved forests. At the tree species scale, there are not enough reports to draw firm conclusions on the effect of any particular species. Mixing of deciduous and coniferous tree species generally affects understory diversity, but in almost all cases maximum diversity is observed in one of the pure stands, not in mixed stands. Understory vegetation is influenced by overstory composition and structure through modifications of resource availability (light, water and soil nutrients) and other effects, such as physical characteristics of the litter layer. Overstory light transmittance and diverse properties of forest litter are factors that have been most fully studied to date, but other factors such as throughfall water quantity and chemistry may also play a role. While the relative importance of mechanisms that account for the effect of overstory on understory biodiversity has often been discussed, these mechanisms have rarely been the subject of formal experiments. Overall, varying management practices and site attributes make it difficult to generalize results. They combine with the effects of tree species in influencing understory vegetation diversity, but they have been rarely considered. Future research is needed to gain a better understanding of the relationship between overstory and understory diversity and establish general laws.     

Resilience of stand structure and tree species diversity in subtropical forest degraded by clear logging

Subtropical forests in the Ryukyu Islands have been degraded by silvicultural practices, and thus their structural attributes are being shifted to other states dominated by a few tree species. This study clarified the mechanisms of the change, and examined the effect of clear logging on the resilience of a subtropical forest. Sprouting regeneration and typhoon disturbance were introduced into an individual-based model, SEIB-DGVM, for describing stand development and succession. The regeneration dynamics from young secondary to old-growth stands were reproduced fairly well with the model. Sprouting recruitment produced high stem density at the beginning of stand development, which caused a self-thinning trajectory following the −3/2 power law. In the late development stage after 70 years, tree species diversity fluctuated because of the regenerative response of sprouting species and the facilitatory effect of typhoon disturbance on the coexistence of subordinate species. The death of canopy trees because of typhoon disturbances reduced the dominance of Castanopsis sieboldii, and depressed its dominance in the understory. Consequently, the understory species could establish by virtue of fallen canopy trees, and tree species diversity increased at the stand level. Clear logging experiments in the model revealed that species diversity deteriorated, especially in the stand dominated by sprouting species. Resilience of subtropical forests was determined by initial species composition before clear logging. Our simulation results suggest that repeated logging drives subtropical forests with high species diversity to a stand monopolized by C. sieboldii.