Methods for studying treefall gaps: A review

As silvicultural objectives have changed over the last several decades, managers are increasingly designing cutting regimes that mimic natural disturbance with the hopes that such systems will restore forests to a more natural condition while optimizing harvest yield. Treefall gaps, canopy openings caused by the death of one or more trees, are the dominant form of disturbance in many forest systems worldwide. These gaps play an important role in forest ecology by helping to maintain bio- and pedo-diversity, influencing nutrient cycling, and preserving the uneven-age nature of late-successional forests. In gap literature, there are inconsistencies with regard to gap terminology, methods for identifying and studying gaps, and modeling gap disturbances. From the papers reviewed, the size of treefall gaps ranges widely from 10 to >5000 m²; we suggest that the maximum gap size should be set at 1000 m². Larger openings tend to have microclimates and return intervals significantly different than smaller treefall gaps. Two main definitions of treefall gaps exist: canopy gap: a ‘hole’ in the forest through all levels down to an average height of 2 m above ground and extended gap: canopy gap plus the area that extends to the bases of surrounding canopy trees. Although researchers have assumed a variety of gap shapes to simplify measuring gap size, gaps are often irregularly shaped and so we recommend that gap areas and shapes be determined from detailed field measurements. Gap age may be determined from tree ring analysis of released trees in or near the gap edge, the spacing of whorls on released saplings, or from decomposition of gap-making trees. Windthrow is the main cause of canopy gaps in a variety of ecosystems; other causes include insects, diseases, acidic deposition, drought, and climate change. Treefall-gap models have been developed to predict the following processes during gap making or infilling: (i) gap abundance, (ii) forest structure, (iii) spatial and temporal variations in light levels, (iv) canopy dynamics, and (v) soil nutrient and water regimes. We recommend a protocol for gap studies and identify future research topics.     

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.     

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.     

Patterns in spatial extent of gap influence on understory plant communities

Gap formation in forests can have impacts on forest ecosystems beyond the physical boundary of the canopy opening. The extent of gap influence may affect responses of many components of forest ecosystems to gap formation on stand and landscape scales. In this study, spatial extent of gap influence on understory plant communities was investigated in and around 0.1 and 0.4 ha harvested canopy gaps in four young Douglas-fir (Psuedotsuga menziesii) dominated stands in western Oregon. In larger gaps, the influence of gap creation on understory plant communities in surrounding forests was minimal. The area showing evidence of gap influence extended a maximum of 2 m beyond the edge of the canopy opening, suggesting that the area affected by gap creation did not differ greatly from the area of physical canopy removal. In smaller gaps, influence of the gap did not extend to the edge of the canopy opening. In fact, the area in which understory vegetation was influenced by gap creation was smaller than the physical canopy opening. Gap influence appears to be limited to areas where ruderal or competitor species are able to replace stress-tolerator species, likely due to elimination or reduction of these species by physical disturbance or competition. The limited gap influence extent exhibited here indicates that gap creation may not have a significant effect on understory plant communities beyond the physical canopy opening. This suggests a limited effectiveness of gaps, especially smaller gaps, as a tool for management of understory plant diversity, and perhaps biodiversity in general, on a larger scale.     

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.     

Effect of natural disturbances on the abundance of Norway spruce (Picea abies (L.) Karst.) regeneration in nemoral forests of the southern boreal zone

The impact of natural disturbances on the canopy (trees ≥14 m high) and sapling stratum (>0.3 and ≤14 m high) composition was studied in nemoral old-growth forests located within the southern boreal zone in Central Russia (Central Forest Reserve, 32°29′–33°01′E, 56°26′–56°31′N). I hypothesized that the current disturbance regime does not allow the maintenance of current spruce abundance in the canopy, and, as a result, there is a continuous shift in the canopy composition towards a greater abundance of deciduous species. Three 300×20 m² transects were established to estimate the proportions of stand under non-closed unexpanded canopy gaps. Data on sapling composition of 49 canopy gaps were used to analyze pattern of gap refuting in these forests. Additionally, data from three forest inventories showed changes in canopy composition over a period from 1972 to 1990.

The current status of nemoral forests is characterized by the high proportion of stand area under treefall gaps (71%). The loss of spruce from the canopy caused by treefalls (53% of the total basal area of gap-makers) was slightly greater than its canopy abundance (45%). Canopy gaps of all sizes encouraged spruce regeneration which might be due to a decrease in sapling mortality and/or more active recruitment of spruce seedlings. After a gap was formed, the presence of spruce in sapling strata increased. However, within both small (<200 m² in size) and large (>200 m²) gaps, tall (>6 m) spruce saplings did not reach the level of its abundance in the tree canopy. In gaps, tall (>6 m) saplings of lime (Tilia cordata) and elm (Ulmus glabra) grew more quickly than those of spruce and maple. These data suggested a decrease in canopy spruce and an increase in deciduous species in the near future which supported the original hypothesis. Analysis of forest inventory records revealed similar changes in the canopy structure over the past two decades. However, the observed high proportion of stand area under gaps implies that for the next few decades large areas of nemoral communities will be occupied by relatively young stands. This may, in turn, decrease the frequency of large-scale treefalls revegetated mainly by deciduous saplings.     

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.     

Thirty years of gap dynamics in a central european beech forest reserve

Investigation of gap characteristics and tree regeneration patternsis central for our understanding of forest dynamics. By integratingaerial photograph analyses and ground surveys, we provide astudy of long-term canopy gap dynamics and tree regenerationpatterns in a Hungarian beech forest reserve. We found (1) thatin spite of the overall increase of gap area during the investigated30-years (from 2.5 to 7.7 per cent), total gap area and averagegap size (40–93 m²) were remarkably similar to those foundin other temperate and tropical forests, (2) if the fate ofindividual gaps was followed, two to three times more intensivecanopy dynamics (gap creation, closure and expansion) couldbe recognized than simple change of gap area indicated, and(3) average seedling density was considered to be sufficientfor natural regeneration. However, it was apparent that recentincreased deer browsing had prevented establishment of youngertrees of 1–2 m in height, as taller saplings were recordedonly in old gaps. Our results not only provide useful informationon forest dynamics but can also contribute to understandingthe potential roles that small forest reserves can play in providingessential reference data for nature-based forest managementof this forest type.     

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     

Comparison of fixed area and distance sampling methods in open forests: case study of Zagros Forest,Iran

The main aim of this study was to evaluate methods for fixed area and distance sampling in the Zagros open forest area in western Iran.Basic forest management and planning required appropriate quantitative and qualitative information.Two sampling methods were compared on the basis of the actual means of characteristics derived from the 100 % survey.In total,37 sampling plots were systematically installed with a grid of 100 m 9 100 m in the study area.Density,crown canopy,and basal area of the stands were measured.The 100 % survey showed that tree density above 12.5 cm diameter at breast height was 68.04 stem ha~(-1),basal area was 15.16 m~2ha~(-1) and crown canopy percentage was 35.71 % ha~(-1).The values for the traits determined by the two sampling methods differed significantly(P=0.05).When the time required for the methods was compared,transect sampling required less than systematic-random sampling.Therefore,the transect sampling method was the more economical method for the Zagros open forests.The transect sampling method was statistically defensible and practical for quantitating characteristics of the Zagros open forests.     

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.     

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。     

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.     

Canopy tree development and undergrowth bamboo dynamics in old-growth Abies–Betula forests in southwestern China: a 12-year study

Interactions between forest canopy characteristics and plants in the forest understory are important determinants of forest community structure and dynamics. In the highlands of southwestern, China the dwarf bamboo Bashania fangiana Yi is an understory dominant beneath a mixed canopy of the evergreen Abies faxoniana (Rheder & Wilson) and the deciduous Betula utilis (D. Don). The goal of this study was to better understand the role of bamboo dominance, canopy characteristics, and periodic bamboo dieback on forest development. To achieve this goal, we measured tree seedling, tree saplings, and trees, forest canopy characteristics, and bamboo cover in permanent forest (n = 4) and gap plots (n = 31) in a mixed A. faxoniana and B. utilis forest in Sichuan, China. Dwarf bamboos died off in 1983 in the gap plots, and in three of the four forest plots. Forest development was assessed for the period 1984–1996. The seedling bank in forest and gap plots increased after bamboo die-off. A. faxoniana seedlings increased more than B. utilis in forest plots; the opposite pattern characterized gap plots. The proportion of seedlings on raised micro-sites on the forest floor also changed and new seedling were more abundant on the forest floor. By 1996, bamboo seedling cover and biomass had recovered to ca. 45% or their pre-flowering values. Rates of bamboo seedling recovery were faster beneath canopy gaps and deciduous trees than beneath forest or evergreen trees. Tree mortality exceeded recruitment in plots with dense bamboo; the opposite pattern was found in the plot with little bamboo. The mortality rate for B. utilis trees (2.4% year⁻¹) was higher than that for A. faxoniana (0.8% year⁻¹) and forests with dense bamboos became more open over the census period. Tree mortality was size-dependent and intermediate sized trees had the lowest rates of mortality. Stand basal area increased mainly due to greater basal area gain than loss for A. faxoniana. Interactions between tree species life history, canopy type, and bamboo life-cycles create heterogeneous conditions that influence tree and bamboo regeneration and contribute to the coexistence of A. faxoniana and B. utilis in old-growth forests in southwestern China.     

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.     

Effects of treefall gaps created by windthrow on bat assemblages in a temperate forest

Determining the way in which spatial distribution and diversity of forest-dwelling mammals varies with natural disturbance is essential to understanding the spatial dynamics of mammal assemblages in forests. Bats are the only forest-dwelling mammals capable of true flight. At a local scale, bat flight ability, which may be related to ecomorphological traits, is an important factor influencing spatial distribution. We tested two postulates: (1) the spatial distribution of bats is affected by sizes of forest gaps created by natural disturbances and (2) species-specific responses can be predicted from bat ecomorphological traits (aspect ratio (AR) and wingtip shape index (WT)) that influence bat flight ability. We found that sizes of forest gaps affected the occurrence of each bat species and species richness of bats at local scales; species-specific responses were related to the ecomorphological traits of individual species. Bat species with high AR and low WT were not affected by variation in canopy gap size. In contrast, bat species with low AR and high WT responded negatively to gap size, and those with intermediate AR and WT responded positively to canopy gap size at sites with small-sized gaps but responded negatively to large-sized gaps. Overall bat species richness responded negatively to gap size. Thus, ecomorphological traits may be important determinants of bat spatial distributions and species diversity at local scales in disturbed habitats. In this study, forest edges might have been undersampled due to the location of bat detectors. However, this potential undersampling should not have affected the interpretation of occurrence patterns of bat species responding to gap size. Our results imply that bat conservation efforts in forest lands should take into consideration specific responses related to ecomorphological traits of species inhabiting an area. The results also suggest that quantifying the effects of natural disturbances on bat assemblages may provide a knowledge base for forest management to minimize the impacts of unavoidable anthropogenic disturbances on bat species diversity. Rare or infrequent natural disturbances can provide models for forest management aimed at maintaining bat species diversity.     

Canopy gap characteristics and its influence on the regeneration of broad-leaved Korean pine forests in Changbai Mountain

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Regeneration of Norway spruce in canopy gaps in Sphagnum-Myrtillus old-growth forests

Gap-associated spruce (Picea abies (L.) Karst.) regeneration in Sphagnum-Myrtillus stands of south taiga forests (Central Forest Biosphere reserve, Tver region, Russia) was studied to evaluate the role of different disturbances in spruce dynamics. Sampled gaps (n=70) ranged from 40 m² to 1.7 ha in size, and from 1 to 70 years since disturbance moment. Formation of gaps lead to increase in the number of stems per ha in all gap size classes (small: 40–200 m², medium: 200–3000 m², and large: >3000 m² gaps). Spruce was the most important species in gap refilling, although its role was not the same in different gap classes. The highest values of relative abundance (compared to other species) were recorded in small gaps, and much lower values – in middle and large gaps. However, as refilling of gaps proceeded, spruce showed rather active regeneration in middle and large gaps and partly regained its abundance in middle-age disturbances. In general, all types of gaps studied supported spruce regeneration into the forest canopy. Almost perfect correlation between predicted outcome of spruce dynamics in gaps and its current role in the canopy of Sphagnum-Myrtillus stands suggests a good adaptation of this species to the current disturbance regime and a steady state of the these forests.     

Characteristics of typhoon disturbed gaps in an old-growth tropical montane rainforest in Hainan Island,China

Disturbances that create gaps can shape the structure and function of forests. However, such disturbance regimes in Asian tropical montane rainforests remain largely unquantified. Least studied are typhoon disturbances that are attributable to climate change. We investigated gap characteristics in terms of size, age, and gapmaker to quantify the gap disturbance regimes in an intact old-growth tropical montane rainforest on Hainan Island,China. The intensity of typhoons has increased since 1949,and typhoon winds blow mostly(45.5%) from the northeast corner of Hainan Island, resulting in a higher frequency of gaps in the northeast. A total of 221 gap-makers(trees that fell to create canopy gaps) and 53 gaps were observed in a 3.16 ha old-growth rainforest. Most canopy gaps(85%)were 200 m~2. The average size of canopy gaps was smaller in the rainforest than in other tropical forests, while the average size of expanded gaps was similar to those in other tropical forests. The maximum age of gaps was 23.5 years indicating that gaps had more rapid turnover than other parts of tropical forests. The frequency distribution of gap-makers followed a lognormal distribution with a distinctive peak at three gap-makers, which was different from the inverse J-shaped curve typical of other tropical forests. Gaps were recorded mainly on slopes between 20° and 35° and wood density of gap-makers was between 0.6 and 0.7 g cm~(-3). Our results suggest that small-scale disturbance was the dominant agent of gap formation in this old-growth rainforest that is subject to increasing typhoon disturbances.     

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]