Canopy gaps in an old-growth fir-beech forest remnant of Western Carpathians

The old-growth forest remnants of Western Carpathians provide a unique possibility to study the disturbance regimes of forest ecosystems without human influence. This study investigated the gap dynamics in beech-dominated old-growth forest Badínsky prales in Central Slovakia. Considering the decline of silver fir in last decades, the study analyzed the main characteristics of disturbance regime with the emphasis on the role of fir. On a 5-ha research plot, the dominant tree species was beech, the proportion of fir reached about 10%. However, a significantly higher proportion of fir (>30%) was observed in the coarse woody debris. In total, 45 canopy openings were recorded. Canopy gaps and expanded gaps covered 11.3 and 37.9% of the forest area, respectively. Despite the highest frequency of small gaps <100 m², their proportion of the overall gap area reached only 20%, what suggests the important role of intermediate and large gaps in the gap dynamics as well. The analysis of gapmakers’ crown projections confirmed a rather low contribution of fir (14.6%) to the gap formation despite its relatively intensive mortality. A high variability of the next generation age between the gaps (6–44 years) was recorded what suggests a large temporal variation of the disturbance events. The lateral expansion of adjacent trees was found to be the determining process for the closure of small canopy openings. The intermediate and large gaps are more likely closed by the height growth of natural regeneration and understory trees that are present on the majority of the area.     

Correlations between canopy gaps and species diversity in broad-leaved and Korean pine mixed forests

Regeneration of tree species associated with canopy gaps in broad-leaved Korean pine forests was investigated. Species diversity in gaps and under closed canopy was compared, the relationship between biodiversity and gap structure was analyzed. Results indicate that there were significant differences between tree species diversity in gaps and that under canopy (p<0.01). In terms of Shannon-Wiener index, evenness index, and abundance index, the biodiversity in gap community were higher than those under forest canopy in regeneration layer. In terms of Simpson’s dominance index, the dominance of certain species in the regeneration layer increased from gaps to closed canopy (p<0.01). In contrast, trends of biodiversity changes of succession layer in gaps and under closed canopy were opposite. Tree species diversity of different layers reacted directly to the change of gap size class. For example, Shannon-Wiener index and abundance index is higher and Simpson’s dominance index is the lowest in succession layer of medium-size gap (100–250 m²) in the broad-leaved Korean pine forest of Changbai Mountains. Shannon-Wiener index reached the highest in a size of ≥250 m² and <100 m², reached the lowest in a size of 200–250 m² in the regeneration layer. Simpson’s dominance index reached its maximum when the gap size was between 200 and 250 m². Generally, species of different layers reacted differently to the changes of gap size classes. The gap size class with more seedlings did not correspond to size class containing more medium-size trees. Tree species diversity indices in the two layers behaved reciprocally during the development process of forest gaps. __________ Translated from Chinese Journal of Applied Ecology, 2005, 16(12): 2,236–2,240 [译自: 应用生态学报, 2005, 16(12): 2,236–2,240]     

Forest gap dynamics and tree regeneration

When one or a few canopy trees die (or are injured) in a forest, small openings, which are called ‘gaps,’ are formed in the forest canopy and are then filled with other trees. This sort of forest dynamics is termed gap dynamics; a large number of papers and data on gap dynamics have been accumulated since the 1970’s, and gap dynamics has been described in many forest types. In this review, I introduce the basic concepts of gap dynamics and summarize major issues on gap dynamics relating to tree regeneration, with many references. Although enormous studies on gap dynamics of natural forests have been conducted, applications of gap dynamics to forest practice are limited. However, accumulated knowledge on gap dynamics should be useful for sustainable forest ecosystem management, as much of the literature suggests. Recipient of the Japanese Forestry Society Award 2000.     

Long-term changes of structure and tree species composition in Dinaric uneven-aged forests: are red deer an important factor?

Uneven-aged silver fir-European beech forest stands were studied to (1) analyse the dynamics of diameter structure and tree species composition in the past two centuries and (2) determine the impact of red deer on the regeneration and recruitment of silver fir. The study used current data on forest stands, archival data from old forest management plans for the period 1789–2004, and red deer harvesting records for the period 1907–2006. During the observation period, the silver fir population aged and silver fir and European beech alternated in dominance. The study revealed a strong impact of red deer on the composition and recruitment of tree regeneration, especially on silver fir regeneration. The drastic changes in red deer density (from extermination up to 5.8 animals km⁻²) and past forest management practices were apparently the main factors driving the population dynamics of silver fir (regeneration, recruitment, and diameter structure) in the study area during the past two centuries.     

Tree recruitment in gaps of various size,clearcuts and undisturbed mixed forest of interior British Columbia,Canada

Tree seedling recruitment was monitored after various types of logging in mixed conifer and deciduous forests of northern British Columbia, Canada. Predicting tree seedling recruitment after disturbance is fundamental to understanding forest dynamics and succession and is vital for forest management purposes. Seedling recruitment success in multi-species northern latitude forests varied as a function of mature tree canopy cover, gap size and position in a gap. Recruitment was abundant within canopy gaps across a wide range of gap sizes (20–5000 m²), but recruit numbers dropped off rapidly under the closed forest canopy and in the open conditions of clearcuts. Inside canopy gaps, recruitment was similar by gap position in small gaps (<300 m²) but, in these northern latitude forests, exhibited a trend of increasing density from the sunny north to shady south end of larger gaps. This was true for all tree species regardless of their shade tolerance ranking. There was no evidence of gap partitioning by any of the tree species during the regeneration phase suggesting that adaptation to the subtleties of gap size during early recruitment are not well developed in these tree species. Favorable locations for emergence and early establishment of germinants were less favorable for growth and survival of established seedlings, i.e. the regeneration niches in these forests were discordant. Tree abundance and species diversity appears to be controlled more by differentiation among growth and survival niches than by the regeneration niches. From the perspective of forest management, abundant natural regeneration of all the dominant tree species of these mixed-species forests can be obtained after partial cutting.     

Gap phase regeneration recruitment of mixed conifer-broadleaf forests in Wolong Nature Reserve

Wolong Nature Reserve is the largest reserve for protecting the endangered wild giant panda. Due to historical factors, even in many sections of the core protection area, the forests have been seriously destroyed and natural forests are poorly restored. However, the relative importance of the determinants for recruitment of communities under disturbance is rarely explored. In our study the endogenous and exterior factors in a forest gap that affect the conifer-broad-leaved mixed forest regeneration were investigated near Wuyipeng, one of the observation stations at Wolong, to explore which determinant had the greatest effect on gap regeneration and to discover the recruitment of seedling establishment in forest gaps. With a linear sampling method, environmental factors, gap characteristics and recruitment of new individuals were measured and examined in every forest gap along three sampling lines. Data of environmental factors in the gaps were collected for a Pearson correlation analysis in order to explore the disturbance and preprocessed characteristics of the gaps, using principal component analysis in SPSS. Correlation analysis was applied to further explore the relationship between changes in the gaps and the response of the regenerating seedlings. The results show that a range of natural and human disturbances affected the pattern and characteristics of the forest gaps in this area. The richness in the composition of the seedlings was higher than that of gap makers, but the order of dominance of the composition was different between seedlings and gap makers. The success of dominant species in establishing themselves was affected by different environmental factors. For instance, the establishment of Betula spp. was correlated significantly with topographic factors, while that of Abies faxoniana was affected by soil characteristics and that of Rhododendron spp. correlated significantly with topographic factors and characteristics of gap makers. Moreover, all the biodiversity indices of regenerating seedlings were significantly correlated with environmental principle components mostly reflecting soil properties. From this we can infer that soil characteristics are the most important factors affecting the regeneration of dominant species and seedling diversity under gap disturbances. __________ Translated from Journal of Plant Ecology (Chinese Version), 2007, 31 (3): 425–430 [译自: 植物生态学报]     

A review of the roles of forest canopy gaps

Treefall gap, canopy opening caused by the death of one or more trees, is the dominant form of disturbance in many forest systems worldwide. Gaps play an important role in forest ecology helping to pre- serve bio- and pedo-diversity, influencing nutrient cycles, and maintain- ing the complex structure of the late-successional forests. Over the last 30 years, numerous reviews have been written describing gap dynamics. Here we synthesize current understanding on gap dynamics relating to tree regeneration with particular emphasis on gap characteristics consid- ered critical to develop ecologically sustainable forest management sys- tems and to conserve native biodiversity. Specifically, we addressed the question： how do gaps influence forest structure？ From the literature re- viewed, the size of gaps induces important changes in factors such as light intensity, soil humidity and soil biological properties that influence tree species regeneration and differ in gaps of different sizes. Shade- tolerant species can colonize small gaps; shade-intolerant species need large gaps for successful regeneration. Additionally, gap dynamics differ between temperate, boreal, and tropical forests, showing the importance of climate differences in driving forest regeneration. This review summa- rizes information of use to forest managers who design cutting regimes that mimic natural disturbances and who must consider forest structure, forest climate, and the role of natural disturbance in their designs.     

Canopy gaps and regeneration in old-growth Oriental beech (Fagus orientalis Lipsky) stands, northern Iran

Virgin beech Fagus orientalis forests in northern Iran provide a unique opportunity to study the disturbance regimes of forest ecosystems without human influence. The aim of this research was to describe characteristics of natural canopy gaps and gap area fraction as an environmental influence on the success of beech seedling establishment in mature beech stands. All canopy gaps and related forest parameters were measured within three 25 ha areas within the Gorazbon compartment of the University of Tehran’s Kheyrud Experimental Forest. An average of 3 gaps/ha occurred in the forest and gap sizes ranged from 19 to 1250 m² in size. The most frequent (58%) canopy gaps were <200 m². In total, canopy gaps covered 9.3% of the forest area. Gaps <400 m² in size were irregular in shape, but larger gaps did not differ significantly in shape from a circle. Most gaps (41%) were formed by a single tree-fall event and beech made up 63% of gap makers and 93% of gap fillers. Frequency and diversity of tree seedlings were not significantly correlated with gap size. The minimum gap size that contained at least one beech gap-filling sapling (<1.3 m tall) was 23.7 m². The median gap size containing at least one beech gap-filling sapling was 206 m² and the maximum size was 1808 m². The management implications from our study suggest that the creation of small and medium sized gaps in mixed beech forest should mimic natural disturbance regimes and provide suitable conditions for successful beech regeneration.     

雪灾干扰下林窗对木荷幼苗更新的影响

林窗干扰是维持森林生态系统的重要驱动力之一,对种子萌发、幼苗等自然更新过程、森林物种组成和动态、森林生物多样性的维持具有重要作用。本研究以2008年雪灾干扰后的浙江江郎山木荷林为研究对象,对木荷林窗大小结构、幼苗更新、生长等进行调查研究,结果表明:扩展林窗以50 100 m2的林窗个数最多(占总数的45.45%),各等级林窗中以50 100 m2的林窗占总面积比例最大(占总面积的30.31%)。林窗中木荷幼苗的平均高度和地径较对照林分分别高1.44 cm和0.61 mm,幼树在林窗中的平均高度和地径则比对照林分中分别高45.37 cm和5.00 mm且差异显著;林窗大小对木荷幼苗、幼树的高度和地径生长影响显著,中林窗中幼苗的高度和地径均高于小林窗和大林窗中的幼苗且差异均显著(F=4.893,P=0.007;F=5.203,P=0.004;n=357);幼树的地径在不同大小林窗中差异显著(F=3.569,P=0.037;n=43)。林窗幼苗的更新密度随着林窗面积的增大而增大,在林窗面积达到76 m2时,更新密度达到最大值,而后随着林窗面积的增大下降;中林窗和小林窗中更新苗木以低矮植株(1级、2级)为主,面积100 m2大林窗中,木荷幼苗生长较快。与他人研究的森林天然林窗相比,雪灾干扰后改变了林窗的大小分布结构和面积,50 100 m2的林窗比例较大,一定程度上更利于幼苗更新,具有相对较大的林窗幼苗更新密度;不论林窗大小,林窗内的更新幼苗都比林内多,郁闭度较大的林内或大面积的空地上都不利于更新幼苗的生长。因此,从受灾木荷林窗大小结构、幼苗更新、生长等来看,中林窗是幼苗适宜更新的面积,为木荷灾后恢复与重建提供了科学依据。     

Gap evapotranspiration and drainage fluxes in a managed and a virgin dinaric silver fir–beech forest in Slovenia: a modelling study

The monthly water balance in gaps in a managed Dinaric silver fir–beech forest and a virgin forest remnant located in SE Slovenia was modelled using a capacity water balance model for two growing seasons. Two gaps of different size (ca. 0.07 and 0.15 ha) were selected in each forest and plots for soil moisture monitoring were established in each gap (2–4) and in the surrounding forest (2–3). We report on the modelled actual evapotranspiration (AET) and potential evapotranspiration (PET) and drainage fluxes (DF) from the rooting zone at the plots. Precipitation over the 2001 growing season (May–October) was considerably drier than average and that for 2002 growing season was wetter than average. Modelled AET for the 2001 growing season varied between 88% and 96% of PET for the managed forest plots and between 90% and 100% for the virgin forest plots. The values for the gap plots varied between 87% and 100% at the managed forest site and between 92% and 96% for the gaps in the virgin forest site. Monthly AET values declined to 57–59% of PET at plots in the centre of the gaps in the managed forest site and to 63–74% in the gaps in the virgin forest site (July), indicating that the highest drought stress occurs in gap centres. For 2002 growing season, AET for all plots was 100% of PET. Modelled DF values in 2001 were 13–35% of rainfall for plots in the gap centres and 12–16% for plots in the forest at the managed forest site. On an average, gap DF values were 20% (154 mm) higher than the forest plot values. DF values in 2002 were similar for all plots at the managed forest site, 31–33% of rainfall. At the virgin forest site, 2001 growing season, DF values varied between 12% and 30% of rainfall at the forest plots and between 12% and 32% at the gap plots. In 2002, DF values for all plots in the virgin forest site varied between 24% and 38% of growing season rainfall. The varying development of vegetation and forest in the gaps of the virgin forest remnant resulted in more variable evapotranspiration and DF during the drought year 2001, with values not as clearly related to distance from the gap centre as in the managed forest site.     

Spatial variation of seed rain and seed banks in gaps of karst forest in the Maolan Nature Reserve, Guizhou Province

Based on an investigation on gaps and non-gap stands of the Maolan National Karst Forest Nature Reserve, Guizhou Province, quantitative characteristics and dynamic changes of seed rain and seed banks in gaps were analyzed. The results show that the total amount of seed rain was 117.4 ± 32.6 seeds/m² during the period of observation. The number of immature seeds was 56.3 ± 10.3 seeds/m², that of mature damaged seeds was 15.7 ± 4.7 seeds/m², and the number of mature germinated seeds was 45.4 ± 8.2 seeds/m². It is suggested that the seed number is rich for gap regeneration. Seed rain in gaps has spatial and temporal heterogeneities which deeply affect regeneration patterns of gap plants. Along a gradient from the gap center to a non-gap stand, seed density in the litter layer, the number of species, and the Shannon-Wiener diversity index were gradually reduced, but these indices increased in the soil. The seed density in the gap center was 2415 ± 639 seeds/m², near the gap center was 2218 ± 421 seeds/m² and at the gap border area 1815 ± 311 seeds/m². This shows that plants in gaps have good latent regeneration potential. In both gaps and non-gap stands, the Jaccard similarity index of seed in litter layer was the largest, second largest at 5–10 cm soil depth, and the least at the 0–5 cm soil layer the index. The Jaccard index between the soil seed bank and the present plant community was large in the litter layer, but decreased with soil depth both in gaps and non-gap stands. The results show that soil seed banks are the main source of gap regeneration in the karst forests of Maolan and contribute significantly to gap regeneration. __________ Translated from Acta Botanica Yunnanica, 2007, 29(3): 327–332 [译自: 云南植物研究]     

Spatial pattern and regeneration dynamics in a temperate Abies–Tsuga forest in southwestern Japan

This article reports the regeneration dynamics of a temperate Abies–Tsuga forest in Kirishima Yaku National Park, southwestern Japan, and examines the influence of species coexistence mediated by gap disturbances on biomass production. All trees taller than 2 m in a 1-ha plot were monitored over four growing seasons. Three growth-form groups occupied different vertical layers. Evergreen conifers and deciduous broad-leaved trees tended to be spatially segregated from evergreen broad-leaved trees, which formed thickets in the understorey. The regeneration of understorey evergreen broad-leaved trees was affected by canopy gaps. The recruitment of conifers and deciduous broad-leaved species was not observed during the four growing seasons. This suggests that regeneration is sporadic and the present environmental conditions are not favorable for these canopy species. The mortality and unsuccessful recruitment of conifers and deciduous trees appeared to cause fluctuations in the productivity of the stand. However, an abundance of canopy gaps accelerates the regrowth of shorter species, and the fluctuation of productivity resulting from the population dynamics of canopy species would be partly mitigated by the regeneration of evergreen understorey species. The horizontal and vertical heterogeneity of the temperate mixed forest was a result of the patch structures of the three growth-form groups. The different regeneration patterns among the three groups, which were driven by interactions of species-specific regeneration niches and disturbance regimes, might be an important factor in maintaining the aboveground productivity in a transitional mixed forest between warm-temperate and cool-temperate zones.     

A new calculation method to estimate forest gap size

The distance and direction coordinates from the center of each canopy gap to its edge in sixteen compass directions were measured in the secondary forests. Total of 21 gaps of different sizes were measured. The gap size was determined using different calculation methods: octagon method (OM), which was applied with two compass directions (OM_1 and OM_2); sixteen-gon method (SM); and elliptical-sector method (ESM), a new calculation method proposed in this work, including 8-elliptical-sector (8ESM) with two different compass directions (8ESM_1 and 8ESM_2) and 16 elliptical sector (16ESM). The results indicated that within the same gap delimitation, the average estimated gap size varied from 122.0 to 164.4 m² with different calculation methods. The estimated area of the same gap among all calculation methods and between pairs exhibited significant differences (p < 0.05). Particularly, the estimated area for the same gap was significantly different between OM_1 and OM_2 (p < 0.005), indicating that the starting direction of the OM significantly affected the gap size. However, there were no significant differences among applications of the new method. When SM is used as a standard, the gap size calculated by OM was 10.0% less but, by ESM, was 11.6% larger. Therefore, the new method is proposed for future research involving the gap size calculation. Finally, we concluded that a comparison of forest gap sizes, particularly using OM, should be conducted when discussing the effects of gaps on forest regeneration and succession. Based on Chinese Journal of Ecology, 2007, 26(3): 455–460     

Patterns of regeneration and ground vegetation associated with canopy gaps in a managed beechwood in southern England

In autumn 2001, 15 canopy gaps were selected for study in RumerhedgeWood, a semi-natural, mesotrophic beechwood in southern England.The gaps were located in mature, beech-dominated stands, andhad originated from openings created during a thinning in theearly 1980s and wind damage in 1987/1990 and/or the consequentsalvage operations. The extent of each gap and surrounding treeswere mapped. Tree/shrub regeneration, ground vegetation, bareearth, leaf litter and canopy openness (using a canopy-scope)were measured within and around the gaps using a 5 x 5-m gridand placing a 1 x 1-m quadrat at each grid intersection (totalnumber of quadrats = 400). Most of the gaps were <75 m² inarea. The largest was 241 m². They were generally irregularin shape and there was little or no understorey present. Mostsurrounding trees were beech Fagus sylvatica L. Bramble Rubusfruticosus L. formed a moderate to dense ground vegetation belowmost gaps and declined around the edges only once the gap openingwas substantially obscured. Apart from a few larger saplings,most regeneration was small and of beech. Most of the latterappeared to be in their fifth or sixth growing season, were10–35 cm tall, had an erect base and flat top, had increasedby <5 cm in height during 2001 and were not browsed by deer.Their height and growth form was related to (1) their positionwithin gaps, (2) the degree of canopy openness and (3) the coverof ground vegetation. This was translated into the followingzonation—(1) around the centre of larger gaps: canopyopenness increased to >15 per cent; bramble cover was nearcomplete; litter depth was low; many places had no beech seedlings,but some of the few present were among the tallest, most uprightand fastest growing; (2) towards the edges of the large gapsor directly below smaller gaps: canopy openness was about 4–10per cent; bramble cover was slightly less; beech seedlings weremoderately abundant but patchy, generally shorter, more flattopped and slow growing than in the gap centre, albeit somewere still among the tallest, most upright and fastest growing;(3) beyond the edge of the large/medium gaps (with the gap onlypartially visible) or directly below very small gaps: canopyopenness was only about 2–3 per cent; bramble was muchreduced; beech seedlings were at their most abundant but stillpatchy in distribution and even shorter, more flat topped andslower growing than in the above zone; (4) in an outer zonebeyond or almost beyond the sight of the gaps: canopy opennesswas <2 per cent; bramble was weak and sparse; beech seedlingswere mostly at low densities and predominately short, slow growingand flat topped. This ring pattern of beech regeneration appearedto relate mainly to (1) differences in light availability affectingthe survival, growth rate and form of seedlings; (2) competitionfrom bramble and possibly (3) limited dispersal of beech seedinto gaps. Recommendations are given for managers who wish touse natural regeneration to restock beech woodland.     

Canopy gap characteristics and their implications for management in the temperate rainforests of southeast Alaska

In the coastal temperate rainforests of southeast Alaska, much progress has been made in describing landscape-level natural disturbances and formulating management systems that emulate those disturbances. Little is known, however, concerning canopy gaps, the dominant form of natural disturbance in the region. During June–August, 1991–1993, we characterized canopy gap patterns and dynamics at three sites in the western hemlock/blueberry/shield fern plant association in the northern portion of the Tongass National Forest.Forest area in canopy gaps ranged from 5.8 to 12.6% and averaged 8.7%. The proportion of forest area in expanded gaps ranged from 18.1 to 43.9% and averaged 27.4%. Gap and gapmaker (tree whose death or crown displacement results in the creation or expansion of a canopy gap) characteristics were generally similar among sites. The majority of canopy gaps were <50 m² in area, had a D/H ratio <0.50, were created from the death of one or two gapmakers, and had experienced gap expansion. The majority of expanded gap areas were <200 m². Gapmakers were usually snapped, had recently died (<20 years ago), and tended to be larger in diameter than surrounding overstory trees. Species composition of gapmakers was similar to surrounding overstory trees. A substantial amount of gap infilling takes place between 20 and 80 years following tree death, but gaps, or portions of gaps, can persist for >80 years. Forest turnover time was estimated to range from 230 to 920 years, and average 575 years. Canopy residence time was estimated to range between 210 and 840 years, and averaged 525 years.To emulate canopy gap dynamics in the plant association studied, forest managers should: (1) maintain a small proportion of a stand in openings within an otherwise undisturbed canopy; (2) use a combination of single tree selection and small group selection systems; (3) re-enter stands every 20–80 years; (4) select larger than average diameter crop trees in proportion to the species composition of the stand; (5) minimize soil disturbance and (6) select crop trees during re-entry so that the creation of new gaps and the expansion of old gaps is accomplished in approximately equal proportions.     

林窗效应研究综述

森林群落常发生一些小规模的内源干扰,从而形成林窗。林窗的形成对推动森林群落的演替更新和生态系统发展至关重要。林窗面积的大小与树木的倒伏方式和林冠冠幅及大小有关。林窗面积及林窗内位置的不同,导致其小气候和土壤理化性质等环境因子发生改变,进而影响到林窗内树种更新和物种组成、林窗植被的物种多样性及其微生物和土壤动物的种类、数量等方面。未来林窗研究重点应该放在次生林和人工林的林窗效应,林窗对森林生态系统碳储量影响机制,林窗凋落物分解因子间的相互关系、作用机理和养分循环,不同树种的林窗与最适更新面积的关系,林窗的边缘效应,林窗的土壤动物和微生物动态及过程。     

The effect of single-tree selction system on soil properties in an oriental beech stand of Hyrcanian forest,north of Iran

A case study was conducted in beech forests of northern Iran to determine the effect of the created gaps on some soil properties in beech stand. Changes of soil properties in small (60 150 m 2 ), medium (151 241 m 2 ), large (242 332 m 2 ) and very large (333 550 m 2 ) gaps, as well as under closed stands were studied eight years after the gap creation. Soil samples were taken from three depths, 0 10, 10 20 and 20 30 cm. The gaps were different from their around undisturbed stands in terms of the following soil parameters: Mg +2 concentration of 0 10 cm at medium gap size, bulk density of 10 20 cm at very large gap size as well as K + and Ca +2 concentrations at 20 30 cm at small and large gap sizes, respectively. Furthermore, the size of the gaps had no effect on soil characteristics through the whole profile. Water saturation percent (Sp %) at 0 10cm as well as P and Mg +2 at 20 30 cm was different amongst undisturbed stands around different gap sizes. The center and the edges of the gap were different only in terms of organic carbon at the depth of 10 20 cm. Significant differences were observed between gaps andclosed canopy regarding P and Ca +2 at depth 0 10 cm and 10 20 cm, respectively. It can be concluded that applied silvicultural system for harvesting trees which created these gaps might be suitable for conservation and forest management in the region.     

Effects of gaps on regeneration of woody plants: a meta-analysis

Forest gaps, openings in the canopy caused by death of one or more trees, have a profound effect on forest regeneration and drive the forest growth cycle. It is therefore necessary to understand the effects of forest gaps on regeneration for modern forest management. In order to provide a quantitative assessment of the effects of forest gaps on regen-eration of woody plants, we conducted this review of gap effects on woody plant regeneration on the basis of 527 observations from 42 indi-vidual papers, and reported the results of these data in a meta-analysis. Overall, densities of regenerated woody plants were significantly greater （359%） in forest gaps than on the closed-canopy forest floor. The regen-eration density in gaps of plantation forests was significantly greater （P＆lt;0.05） than that of natural forest because the regeneration in gaps of plan-tation forests was improved by both gap effects and experimental meas-ures. Similarly, in comparison to natural gaps, regeneration was better enhanced in artificial gaps. Regeneration density exhibited a significantly positive correlation with gap size, but a negative correlation with gap age because the gap size decreased with increasing gap age. Shade tolerance of woody plants affected regeneration density in gaps and understory. Average regeneration density of shade-tolerant species exhibited a sig-nificantly positive response to gaps but densities remained lower in total than those of intermediate and shade-intolerant species. Gap effects on regeneration decreased in response to increasing temperature and pre-cipitation because of the limiting effects of lower temperature and moisture on woody plant regeneration. In summary, forest gaps enhance woody plant regeneration, and the effects of gaps varied by forest type, gap characteristics, environmental factors and plant traits. The results of this meta-analysis are useful for better understanding the effects and roles of gaps on forest regeneration and forest management.     

The dynamics of biomass production in relation to foliar and root traits in a grey alder (Alnus incana (L.) Moench) plantation on abandoned agricultural land

The dynamics of the above-ground biomass production of a greyalder plantation on abandoned farmland was investigated during11 years after establishment. In the 12-year-old stand, thetotal biomass of the above-ground part of the stand was 68.8t dry matter (DM) ha^–1 and the current annual production(CAP) was 14.0 t DM ha^–1 year^–1. The predicted meanannual increment (MAI) reached is maximum at the age of 16 years,which indicates bulk maturity (the stand age when CAI = MAI)and appropriate rotation time for obtaining maximum biomassproduction. In the case of short-rotation forestry, initialstand density should not be higher than 6500–6000 treesper hectare. Below-ground biomass accounted for 18 and 16 percent of total stand biomass at a stand age of 5 and 10 years,respectively. The biomass of the nodules was estimated at 155± 63 kg DM ha^–1 and the biomass of the fine rootswas estimated at 870 ± 130 kg DM ha^–1 in the 10-year-oldgrey alder stand. Of the fine roots, 80 per cent and almostall nodules were located in the upper 0–20 cm soil layerin both the 5-year-old and the 10-year-old stand. The valueof leaf area index increased with stand age, ranging between1.38 and 5.43 m² m^–2 during the development of the stand.Specific leaf area varied in different years from 11.1 to 13.5m² kg^–1.     

Gap-Disturbance Regimes in Different Forest Types of Japan

Abstract

In this study, I defined a gap as a small opening formed in a forest canopy (area < 0.1 ha) and tried to synthesize gap-disturbance regimes of primary mature stands in different forest types of Japan, such as warm temperate evergreen broad-leaved (4 stands in 3 localities), cool temperate deciduous broad-leaved (10 in 5) and subalpine evergreen coniferous (3 in 1) forests. Mean percentage of the total gap area within the total forested area (percentage gap area) in each forest type was 17.0% in warm temperate (number of surveyed gaps was 161), 13.8% in cool temperate (278 gaps), and 8.0% in subalpine (100 gaps) forests. Mean gap density (ha^-1) and mean gap size (m²) were 19.5 and 77.1 in warm temperate, 16.4 and 92.0 in cool temperate, and 19.1 and 41.9 in subalpine forests, respectively. These figures indicate that gap density is not substantially different among the forest types, but the mean gap size of subalpine forests is smaller than the other two, resulting in lower percentage gap area of this forest type. The gap size distributions were similar among the forest types; smaller gaps were much more frequent than larger ones, and gaps > 400 rrr were rare in warm temperate and cool temperate forests. In subalpine forests gaps > 200 m² were rare. Gaps formed by multiple gapmakers comprised 19.9% of all gaps in warm temperate, 9.9% in cool temperate, and 44.9% in subalpine forests, which implies that gap formation by simultaneous tree fall or gap enlargement is more frequent in subalpine forests. Canopy trees died less often by uprooting in every forest type; dominant mode for the death of canopy trees was by leaving standing-dead or with broken trunks in every forest type. Since typhoons are obviously a chief agent of forest disturbance in Japan, frequency or magnitude of typhoon disturbance may influence these differences in the gap-disturbance regimes. In addition to the disturbance, tree architectures seem to affect some of these differences; narrower crown size of conifers compared with broad-leaved trees is considered one major cause for smaller gap size of the conifer forest.