Gap characteristics and gap regeneration in subalpine old-growth coniferous forests,central Japan

Gap characteristics and gap regeneration were studied in three old-growth stands of subalpine coniferous forests in the northern Yatsugatake and the northern Akaishi mountains, central Japan. With the results of the present study and those of a previous study conducted in another locality, general features of gap characteristics and gap regeneration behavior of major tree species in subalpine coniferous forests of central Japan were summarized and discussed. Of the total 237 gaps investigated in the 14.48 ha of forested area, the percentage gap area to surveyed area, gap density and mean gap size were 7.3%, 17.2 ha⁻¹, and 43.3 m², respectively. The gap size distributions were similar among stands and showed a strong positive skewness with a few large and many small gaps; gaps <40m² were most frequent and those >200 m² were rare. Gaps due to the death of multiple canopy trees comprised 44.7% of the total ones. Canopy trees died in various states; standing dead (42.6%) or trunk broken (43.7%) were common and uprooted (12.2%) was an uncommon type of death of canopy trees. These figures indicate that general features of gap characteristics in this forest type are the low proportion of gap area and the high proportions of small gap size and multiple-tree gap formation. In general, shade-tolerantAbies frequently, andTsuga, infrequently, regenerate in gaps from advance regenerations recruited before gap formation, whilePicea and shade-intolerantBetula possibly regenerate in gaps from new individuals recruited after gap formation. Gap successors of conifers occurred in a wide range of gap size and did not show the clear preference to species specific gap size. In old-growth stands without large-scale disturbance (≥0.1 ha in area) of subalpine coniferous forests of central Japan, major tree species may coexist with their different gap-regeneration behaviors and, probably, different life history traits.     

Gap dynamics in climaxFagus crenata forests

Gap characteristics and gap regeneration were studied in several climaxFagus crenata forests in Japan. 278 gaps were observed. Gaps covered 12% of the total land area of 20.05 ha. Gap density was 13.9 gaps per ha and, mean gap size was 92.0 m². Smaller gaps were much more frequent than larger ones. Gaps larger than 400 m² were rare. Most gaps were created by the death of single trees. Canopy trees died more often standing or with broken trunks than by uprooting, although uprooted trees were relatively abundant in the site with poor soil drainage and in the site on upper slope. Differences of gap regeneration behaviour were recognized among tree species.F. crenata regenerates in gaps from saplings recruited before gap creation and can replace not only its own gaps but also gaps of other species. Most species other thanF. crenata andMagnolia obovata could not regenerate in their own gaps. More successful regeneration ofF. crenata may occur in gaps smaller than 200 m², althought it regenerated in a wide range of gap size. However, increased relative density ofF. crenata in the canopy layer seems to prevent its successful regeneration. Gap regeneration of other species did not clearly depend on a species-specific gap size.     

Gap characteristics and gap regeneration in primary evergreen broad-leaved forests of western Japan

Gap characteristics and regeneration in gaps were studied in some primary evergreen broad-leaved forests of the warm temperate zone in western Japan. Total observed 161 gaps covered 15.7% of the total land area of 8.23 ha. Gap density was 19. 6 gaps ha⁻¹ and mean gap size was 80.3 m². Smaller gaps (<80 m²) were much more frequent than larger ones, and gaps larger than 400 m² were rare. Gaps created by the death or the injury of single trees were 79.5%. Canopy trees died most often with broken trunks and not so often by uprooting or leaving standing-dead. Different types of gap regeneration behaviour were recognized among the major canopy tree species, though gap regeneration of the common evergreen broad-leaved tree species did not clearly depend on a species-specific gap size.Distylium racemosum, which occurred in equal importance (about 25%) in the canopy layer of each study stand, regenerates in gaps from saplings recruited before gap creation and can replace not only its own gaps but also gaps of other species. Therefore, it can be considered a typical climax species in this forest type of western Japan.Persea thunbergii, which can reproduce vegetatively, showed a similar type of gap regeneration behaviour.Castanopsis cuspidata can replace itself with relatively higher frequency by means of vegetative reproduction (stump sprouting) after gap creation.Quercus acuta andQuercus salicina did not regenerate under the current gap-disturbance regime. Though the frequency of uprooting is low, soil disturbance by uprooting seems to be important for the perpetuation of the pioneer tree species,Fagara ailanthoides, which recruits from buried seeds in the soil     

Regeneration in subalpine coniferous forests

The regeneration process of a subalpine coniferous forest, a mixed forest ofTsuga diversifolia (dominant species),Abies veitchii, Abies mariessi, andPicea jezoensis var.hondoensis, was studied on the basis of annual ring data. The age class distribution was discontinuous and four age groups occurred in the study plot (30m×30m). The canopy layer was a mosaic of patches (83.8–133.7 m² patch area), which had different mean ages. The recruitment of canopy trees was carried out only by advance regeneration in the plot. The diameter growth ofAbies andPicea exceeded diameter growth ofTsuga in the gap.Abies lived for 200–300 years and their trunks were susceptible to heart rot.Picea lived for 300–400 years andTsuga for more than 400 years. The regeneration process derived from the analysis of the plot consisted of three phases leading to the development of a even-aged patch; (1) the establishment of saplings before a gap opening, (2) the opening of a gap in the canopy and repair of the canopy by advance regenerated saplings dominated by rapid growth species,Abies andPicea, and (3) the dying off of canopy trees as each species reached the end of its life-span, resulting in pure patches of long-livedTsuga.     

Gap formation, microsite variation and the conifer seedling occurrence in a subalpine old-growth forest, central Japan

To evaluate the effects of canopy gaps and forest floor microsites (soil, fallen logs, root-mounds, buttresses and stumps) on regeneration of subalpine forests, the gap regeneration and seedling occurrence of conifers (Abies mariesii, Abies veitchii, Picea jezoensis var. hondoensis and Tsuga diversifolia) were studied in two stands of a subalpine old-growth forest, central Japan. The percentage of gap area to total surveyed area was 11.2–11.3% in the stands. Gap regeneration was not common for P. jezoensis var. hondoensis and T. diversifolia. In contrast, gap regeneration by advanced regeneration was relatively common for Abies. Seedling occurrence of P. jezoensis var. hondoensis and T. diversifolia was restricted on elevated surfaces such as stumps and root-mounds, while Abies seedlings could occur on soil as well as on elevated surfaces. Rotten stumps were the most favorable microsites for conifer seedling occurrence, which covered small area in the forest floor. Although canopy gaps were not always favorable for seedling occurrence, all conifer seedlings were larger under canopy gaps than under closed canopy. Canopy gaps and forest floor microsites clearly affected seedling occurrence and growth of conifers. This suggests that regeneration of conifers is related to the difference of growth advantage under canopy gaps and favorable microsites for seedling occurrence.     

Succession after stand replacing disturbances by fire,wind throw,and insects in the dark Taiga of Central Siberia

The dark taiga of Siberia is a boreal vegetation dominated by Picea obovata, Abies sibirica, and Pinus sibirica during the late succession. This paper investigates the population and age structure of 18 stands representing different stages after fire, wind throw, and insect damage. To our knowledge, this is the first time that the forest dynamics of the Siberian dark taiga is described quantitatively in terms of succession, and age after disturbance, stand density, and basal area. The basis for the curve–linear age/diameter relation of trees is being analyzed. (1) After a stand-replacing fire Betula dominates (4,000 trees) for about 70 years. Although tree density of Betula decreases rapidly, basal area (BA) reached >30 m²/ha after 40 years. (2) After fire, Abies, Picea, and Pinus establish at the same time as Betula, but grow slower, continue to gain height and eventually replace Betula. Abies has the highest seedling number (about 1,000 trees/ha) and the highest mortality. Picea establishes with 100–400 trees/ha, it has less mortality, but reached the highest age (>350 years, DBH 51 cm). Picea is the most important indicator for successional age after disturbance. Pinus sibirica is an accompanying species. The widely distributed “mixed boreal forest” is a stage about 120 years after fire reaching a BA of >40 m²/ha. (3) Wind throw and insect damage occur in old conifer stands. Betula does not establish. Abies initially dominates (2,000–6,000 trees/ha), but Picea becomes dominant after 150–200 years since Abies is shorter lived. (4) Without disturbance the forest develops into a pure coniferous canopy (BA 40–50 m²/ha) with a self-regenerating density of 1,000 coniferous canopy trees/ha. There is no collapse of old-growth stands. The dark taiga may serve as an example in which a limited set to tree species may gain dominance under certain disturbance conditions without ever getting monotypic.     

Community dynamics of evergreen broadleaf forests in southwestern Japan

The process and rate of revegetation in gaps in an evergreen oak forest were studied by comparing the species composition, tree density, frequency distribution of tree height, and relation between diameter at breast height and tree height among different aged stands. For estimating stand ages, the ages of gap indicators, such as,Symplocos prunifolia andAcer rufinerve, were very useful. It took about 70 years for gaps to be filled by large fully-grown trees. Since the mean residence time of the forest canopy was 180 years, the trees that attain the forest canopy were expected to be canopy trees for 110 years on the average. Tree densities of all broadleaved evergreens exceptS. prunifolia, were independent of stand age. On the other hand, densities of gap indicators,S. prunifolia andA. rufinerve, decreased as stand age increased. Other deciduous broadleaf and coniferous species were scarce as a whole. According to the frequency distributions of height of live and dead trees in different aged stands, it was suggested that shorter trees were more susceptible to death than taller trees. The self-thinning in revegetation process in gaps approximately followed the 3/2 power law, though the power was larger (−1.32) than expected from the law.     

Spatial dynamics of regeneration in a conifer/broad‐leaved forest in northern Japan

Abstract. This study deals with stand dynamics over a 6‐yr period in a conifer/broad‐leaved mixed forest in Hokkaido, northern Japan. The annual rates of gap formation and recovery were 81.3 m²/ha and 66.7 m²/ha, respectively and turnover time of the canopy was 125 yr. The recruitment processes of the component species in this cool‐temperate forest were governed by different canopy types: gap, canopy edge and closed canopy. Magnolia obovata regenerated in canopy edges, and Acer mono and Prunus ssiori regenerated in canopy edges and gaps. The results suggested that the mosaic structure made up of closed canopy, canopy edge and gap was related to various regeneration niches. Abies sachalinensis had high mortality rates, initiating gap expansion. The transition probabilities from closed canopy or canopy edge to gap for deciduous broad‐leaved trees were lower than for A. sachalinensis, which implies that the difference in degeneration patterns of conifer and broad‐leaved canopies contributes to the heterogeneity of spatial structure in the mixed forests. Spatial dynamics were determined by a combination of gap expansion by A. sachalinensis (neighbour‐dependent disturbance) and gap formation by deciduous broad‐leaved trees (random disturbance).     

Loss of oak dominance in dry-mesic deciduous forests predicted by gap capture methods

Gap capture methods predict future forest canopy species composition from the tallest trees growing in canopy gaps rather than from random samples of shaded understory trees. We used gap capture methods and a simulation approach to forecast canopy composition in three old oak forests (Quercus spp.) on dry-mesic sites in southern Wisconsin, USA. In the simulation, a gap sapling is considered successful if it exceeds a threshold height of 13–17 m (height of maximum crown width of canopy trees) before its crown center can be overtopped by lateral crown growth of mature trees. The composition of both the tallest gap trees and simulated gap captures suggests that 68–90% of the next generation of canopy trees in the stands will consist of non-Quercus species, particularly Ulmus rubra, Carya ovata and Prunus serotina. Quercus species will probably remain as a lesser stand component, with Quercus alba and Quercus rubra predicted to comprise about 19% of successful gap trees across the three stands. Several methods of predicting future canopy composition gave similar results, probably because no gap opportunist species were present in these stands and there was an even distribution of species among height strata in gaps. Gap trees of competing species already average 11–13 m tall, and mean expected time for these trees to reach full canopy height is only 19 years. For these reasons, we suggest that dominance will shift from oaks to other species, even though late successional species (e.g., Acer and Tilia) are not presently common in the understories of these stands.     

The dynamics of old-growth Pseudotsuga forests in the western Cascade Range,Oregon, USA

Size and age structure, spatial analysis, and disturbance history were used to analyse the population structures and regeneration patterns of 8 conifer stands in the central western Cascade Range, Oregon, USA. Variation in forest structure reflected the effects of frequent (20–50 yr) low-intensity fires and treefalls, infrequent (100–200 yr) localised, intense fires, and extensive fires that resulted in stand replacement (every ca 400 yr?).The amount of canopy removed and the size of openings formed by fires and treefalls were important determinants of subsequent forest establishment. Single or several species stands of Pseudotsuga and/or Abies procera, or mixed species stands of Pseudotsuga, Abies procera, Tsuga heterophylla and Abies amabilis established in openings where intense fires had removed most of the canopy trees over several ha. Multi-tiered and multi-aged stands, often containing 400–500 yr-old Pseudotsuga and variously-sized more or less even-aged patches of younger shade tolerant Tsuga heterophylla and/or Abies amabilis, occurred where lower-intensity fires did not kill all overstorey trees or where treefalls occurred after the initial fire.Current regeneration processes are influenced by overstorey composition, the availability and size of canopy openings, and the availability of substrates suitable for regeneration. Tsuga heterophylla and Abies amabilis established under Pseudotsuga menziesii and Abies procera canopies and in small canopy openings (<400 m²) created by windfalls, but rarely under Tsuga. Down logs and stumps were favoured establishment sites for Tsuga.The disturbance regime of fires of low-to moderate-intensity, windfalls, and occasional fires that result in extensive stand replacement contrasts with the pattern of infrequent, catastrophic disturbances proposed for other areas of the Pacific Northwest. Although fires at stand establishment commonly determine much of the composition, structure, and subsequent stand development, canopy replacement by shade tolerant species occurs as the different life histories of the species are expressed in response to various disturbances differing in intensity and frequency. Such a non-equilibrium view of vegetation change is consistent with many other fire-dominated forests of the western United States.     

Reactions of the herb and moss layer,tree saplings and the shrub layer to tree deaths in forests of the Wielkopolska National Park (Western Poland)

This study examines the impact of canopy and canopy gaps on the development of lower forest layers in five protected phytocoenoses of oak-hornbeam habitats (natural and regeneration stands) and oak-pine habitats in the Wielkopolska National Park (WPN). In the studied forests the most common form of dead trees are those which are uprooted (45–59%), while the most frequently dying tree is pine (40–88%). The total area of gaps in relation to the studied forest area ranges from 329 to 2356 m²/ha.     

Effects of disturbance and size structure on the regeneration process in a sub-boreal coniferous forest,northern Japan

The stand structure and disturbance history in a sub-boreal coniferous forest dominated byPicea jezoensis, Picea glehnii andAbies sachalinensis were investigated in four study plots set up in Taisetsuzan National Park, Japan. The effect of stand characteristics on the growth and mortality rates of understory trees was examined. Although all the stands showed inverse J-shape d.b.h. (diameter at breast height) distributions, the age structure and disturbance history differed amongst the stands. The stands with wide d.b.h. distribution (i.e. large CV and skewness) were more uneven-aged than those with narrow d.b.h. distribution (i.e. small CV and skewness). The disturbance-return interval based on the model of Hett and Loucks was 31 to 65 years. The gap ratio in the canopy was also different among the stands. These suggest that the variations in stand structure represent different occurrences of natural disturbances. Furthermore, the structural features such as size structure, canopy gap ratio and density of canopy trees also affected the growth dynamics of understory trees (≥2 m in height and <10 cm in diameter at breast height). The growth and mortality rates of understory trees changed with the canopy gap ratio and canopy tree density. The understory trees of stands with wide canopy d.b.h. distribution had higher growth and canopy recruitment rates than those of stands with narrow canopy d.b.h. distribution, contributing to the maintenance of continuous stand stratification. The understory trees of stands with narrow canopy d.b.h. distribution showed lower growth and higher mortality rates than those of stands with narrow canopy d.b.h. distribution, leading to the formation of a single-canopy structure. It is suggested that natural disturbance governs the regeneration process in the future by affecting the growth and mortality patterns of understory trees through the stand structure (size and age structure, canopy tree density, canopy gap ratio).     

Nitrogen dynamics of a boreal black spruce wildfire chronosequence

This study examined the nitrogen (N) dynamics of a black spruce (Picea mariana (Mill.) BSP)-dominated chronosequence in Manitoba, Canada. The seven sites studied each contained separate well- and poorly drained stands, originated from stand-killing wildfires, and were between 3 and 151 years old. Our goals were to (i) measure total N concentration ([N]) of all biomass components and major soil horizons; (ii) compare N content and select vegetation N cycle processes among the stands; and (iii) examine relationships between ecosystem C and N cycling for these stands. Vegetation [N] varied significantly by tissue type, species, soil drainage, and stand age; woody debris [N] increased with decay state and decreased with debris size. Soil [N] declined with horizon depth but did not vary with stand age. Total (live + dead) biomass N content ranged from 18.4 to 99.7 g N m⁻² in the well-drained stands and 37.8–154.6 g N m⁻² in the poorly drained stands. Mean soil N content (380.6 g N m⁻²) was unaffected by stand age. Annual vegetation N requirement (5.9 and 8.4 g N m⁻² yr⁻¹ in the middle-aged well- and poorly drained stands, respectively) was dominated by trees and fine roots in the well-drained stands, and bryophytes in the poorly drained stands. Fraction N retranslocated was significantly higher in deciduous than evergreen tree species, and in older than younger stands. Nitrogen use efficiency (NUE) was significantly lower in bryophytes than in trees, and in deciduous than in evergreen trees. Tree NUE increased with stand age, but overall stand NUE was roughly constant (∼ ∼150 g g⁻¹ N) across the entire chronosequence.     

Tree species regeneration in a mid-elevation,temperate rain forest in Isla de Chiloé, Chile

The regeneration of canopy and subeanopy species in a mid-elevation, primary rain forest in the Coastal Range of Isla de Chiloé (42°30S), in the cold-temperate region of Chile, was studied by comparing seedling and sapling abundances under the forest canopy, and within 36 tree-fall gaps. The forest was dominated byAmomyrtus luma andLaurelia philippiana (33 and 32% of the main canopy individuals), and two subcanopy species (Myrceugenia ovata, andMyrceugenia planipes) were also important. Uncommon species in the canopy wereDrimys winteri, Amomyrtus meli, andRaphithamnus spinosus. Tree-fall gaps were created generally by the fall of several trees, and the main canopy species were the principal gap-makers. Gap sizes varied between 28 and 972 m², with a mean of 197 m². Seedling and sapling abundances indicate that the dominant species are capable of regenerating below the canopy, but they also germinate and show enhanced growth within small light gaps. For one of the common subcanopy species (M. planipes) and the two infrequent canopy species (D. winteri, andA. meli) regeneration seems to depend entirely on tree-fall gaps. Thus, in this forest, light gaps allow the persistence of infrequent canopy species, but seem less important for the regeneration and maintenance of dominant canopy species.     

Autotrophic and heterotrophic respiration in needle fir and Quercus-dominated stands in a cool-temperate forest, central Korea

To investigate annual variation in soil respiration (R _S) and its components [autotrophic (R _A) and heterotrophic (R _H)] in relation to seasonal changes in soil temperature (ST) and soil water content (SWC) in an Abies holophylla stand (stand A) and a Quercus-dominated stand (stand Q), we set up trenched plots and measured R _S, ST and SWC for 2 years. The mean annual rate of R _S was 436 mg CO₂ m⁻² h⁻¹, ranging from 76 to 1,170 mg CO₂ m⁻² h⁻¹, in stand A and 376 mg CO₂ m⁻² h⁻¹, ranging from 82 to 1,133 mg CO₂ m⁻² h⁻¹, in stand Q. A significant relationship between R _S and its components and ST was observed over the 2 years in both stands, whereas a significant correlation between R _A and SWC was detected only in stand Q. On average over the 2 years, R _A accounted for approximately 34% (range 17–67%) and 31% (15–82%) of the variation in R _S in stands A and Q, respectively. Our results suggested that vegetation type did not significantly affect the annual mean contributions of R _A or R _H, but did affect the pattern of seasonal change in the contribution of R _A to R _S.     

Gap regeneration in primary evergreen broad-leaved forests with or without a major canopy tree,Distylium racemosum,southwestern Japan: A comparative analysis

Gap regeneration was studied in a typical primary evergreen broad-leaved forest withoutDistylium racemosum, at the Kasugayama Forest Reserve, southwestern Japan and the results were compared with those from other primary evergreen broad-leaved forests in southwestern Japan, whereD. racemosum was the dominant species. Several common types of gap regeneration behavior were recognized among the major tree species and forests with or withoutD. racemosum consisted of three typical regeneration guilds which could be detected in the principal component analysis.Castanopsis cuspidata frequently regenerated in gaps from saplings recruited before gap formation in the forest withoutD. racemosum, although elsewhere, in forests withD. racemosum, it lacked advanced regeneration and regenerated in gaps from saplings recruited after gap formation. Some evergreenQuercus had their regenerations in gaps of the forest withoutD. racemosum, although elsewhere, in forests withD. racemosum, evergreenQuercus might not regenerate. The results indicate that tree species may change their regeneration behavior depending on the presence or absence of another key dominant species. This suggests that the presence and the dominance of a potential competitor induces shifts in the regeneration niche of other coexisting tree species.     

Growth and survival of <Emphasis Type="Italic">Abies sachalinensis</Emphasis> seedlings for three years after selection harvesting in northern Hokkaido,Japan

Previous studies have indicated that recruitment of Abies sachalinensis, a representative conifer species of northern Japan, decreased following single-tree selection harvesting in stands with dense dwarf bamboo understory. We tested the hypothesis that growth and survival of A. sachalinensis seedlings are reduced by canopy opening in that type of stand. A 0.75 ha study plot was examined, and all the seedlings (defined as trees with height 0.5–2 m) were identified and their shoot extensions measured for three years after single-tree selection harvesting (26% intensity in terms of basal area). The leader extensions of A. sachalinensis seedlings that experienced canopy opening were greatly improved. However, a negative effect on survival was also apparent; nearly 40% of seedlings died at the sites that experienced canopy opening. These results were supported by generalized linear models that examined variations of local harvesting intensity for individual seedlings. Despite the ability of A. sachalinensis to respond rapidly to exposure, some physiological stresses may have appeared, and presumably were amplified by co-occurring dwarf bamboos. With regard to the low seedling density (156 stems ha⁻¹), common in this type of stands, local harvesting intensity in the selection system should be reduced to maintain survival of advanced regeneration, thus sustaining stand structure and composition.     

Regeneration process of coniferous forests in northern Hokkaido I.Abies sachalinensis forest andPicea glehnii forest

Regeneration of natural forests was studied in the Nakagawa Experiment Forest of Hokkaido University using age distribution surveys made by the clear felling method. In Plot 1 (30 m × 65 m),Abies sachalinensis dominated the canopy layer but there were also a fewBetula ermanii trees.Sasa senanensis densely covered the forest floor. Most of the canopy trees were from 122 to 195 years old. Seedlings younger than 50 years old ofA. sachalinensis were found on fallen logs and root bases. There were, however, few trees from 50 to 120 years old. The present canopy trees seemed to have regenerated after competitive pressure from old canopy andSasa disappeared 180 years ago. Plot 2 (50 m × 100 m) on serpentinite soil was dominated byPicea glehnii. Sasa kulirensis covered the floor but not as densely asS. senanensis in Plot 1. The ages ofP. glehnii ranged from 1 to 586 years old, and the age distribution ofA. sachalinensis was L-shaped. A small gap in the canopy formed about 290 years ago, and it gradually extended. Conifers regenerated continuously in the extending gap butB. ermanii did not. One hundred thirty years ago, part of Plot 2 was again destroyed andA. sachalinensis andB. ermanii regenerated. Thus, two types of regeneration were found. One regenerated both conifers andBetula after a sudden disturbance of canopy layer or death ofSasa, and the other, under an extending gap, regenerated only conifers.     

小兴安岭阔叶红松混交林林隙特征

对小兴安岭阔叶红松混交林林隙基本特征进行了研究。结果表明:林隙的线状密度为31.78个/km,冠空隙和扩展林隙所占的面积比例分别为15.71%和30.78%;冠空隙的年干扰频率为0.46%,干扰轮回期约为434.8a。冠空隙的大小变化在42.12—372.52m2之间,平均为153.37m2;扩展林隙的大小变化在98.65m2—633.10m2之间,平均为300.44m2。冠空隙和扩展林隙面积分布格局均符合Weibull分布。林隙形成方式主要为干基折断,占总形成木总数的35.29%,其次为掘根风倒,占28.43%。平均每个林隙的形成木为4.98株,由红松、白桦、枫桦、冷杉形成,径级在20—30 cm之间,高度在15—30 m之间。冠空隙的直径与高度比值的相对频率的分布呈单峰型曲线,当比值为0.30—0.45时,出现峰值;而扩展林隙的直径与高度比值的相对频率的分布呈双峰型曲线,当比值分别为0.75—0.90和1.05—1.15时,出现峰值。林隙边缘木胸径级的多度分布和高度级多度分布符合Weibull分布,但不符合正态分布。约13.41%的边缘木未出现偏冠现象,偏冠率在0.5—0.7之间的边缘木占70.49%。     

Structure and tree-fall gap dynamics of old-growth Nothofagus forests in Tierra del Fuego,Argentina

Abstract. Tree size and age structure, tree-fall and gap characteristics, and regeneration in gaps were studied in Nothofagus-dominated old-growth forests in Tierra del Fuego, Argentina. Gap-phase regeneration has resulted in all-aged populations for N. pumilio, N. betuloides, and Drimys winteri, and regeneration in gaps appears to be maintaining coexistence between species in mixed stands. N. betuloides fills many gaps via advance regeneration and some individuals persist for > 150 yr in the understory. Multiple periods of release and suppression indicate that N. betuloides may take advantage of several gap events to reach the main canopy. Likewise, Drimys grows well under closed canopy and can rapidly respond to gap formation, sometimes impeding the regeneration of N. betuloides. In contrast, N. pumilio regenerates in gaps mainly from seed or from advance regeneration of small, ephemeral seedlings. Gap turnover times in Fuegian forests were estimated at 300 - 500 yr, although gap formation was highly episodic and possibly associated with regionally extensive windstorms, earthquakes, and stand-level dieback. 92 % of gaps involved multiple tree-falls, and at least 53 % involved secondary expansion. Gap and tree-fall characteristics in Tierra del Fuego were similar to results from northern Patagonia, Chile, and New Zealand; however, we emphasize that regeneration of Nothofagus spp. and Drimys winteri in gaps depends on associated vegetation and varies along both local and regional environmental gradients.