Natural disturbance and tree species coexistence in an old-growth beech - dwarf bamboo forest,southwestern Japan

Abstract. The structure and composition of a cool-temperate old-growth beech (Fagus crenata) - dwarf bamboo (Sasa spp.) forest, partially affected by landslide disturbance, in the Daisen Forest Reserve of southwestern Japan, were investigated in relation to forest floor and canopy conditions. All stems ≥ 4 cm DBH were mapped on a 4-ha plot and analyses were made of population structure, spatial distribution and spatial association of major tree species. The dominant species, F. crenata, which had the maximum DBH among the species present, had the highest stem density. However, for other species, larger-sized species had lower stem density with few smaller stems or saplings, while smaller-sized species had higher stem density with many smaller stems or saplings. Canopy trees of F. crenata were distributed randomly in the plot, while its stems in the other layers and all other species were distributed patchily. Small patches represent gap-phase regeneration. Larger patches correlate with landslide disturbance, difference in soil age, or the presence/absence of Sasa. Cluster analysis for spatial associations among species and stems in the different layers revealed that the forest community consists of several groups. One main group was formed on sites not covered with Sasa. This group contained a successional subgroup (from Betula grossa to Acer mono and/or F. crenata) initiated by landslide disturbance and a subgroup of tree species that avoid Sasa. Another group was formed on sites with mature soils covered largely with Sasa. This contained associations of canopy trees of F. crenata and smaller-sized tree species such as Acanthopanax sciadophylloides and Acer japonicum. It is found that the community of this old-growth beech forest is largely organized by natural disturbance and heterogeneous conditions of the forest floor (difference in soil age and presence/absence of Sasa). The existence of these different factors and the different responses of species to them largely contribute to the maintenance of tree species diversity in this forest.; Keywords: Cluster analysis; Fagus crenata; Forest dynamics; Gap; Landslide; Spatial pattern.     

Heterogeneous genetic structure in a Fagus crenata population in an old-growth beech forest revealed by microsatellite markers

The within-population genetic structure of Fagus crenata in a 4-ha plot (200 x 200 m) of an old-growth beech forest was analysed using microsatellite markers. To assess the genetic structure, Moran's I spatial autocorrelation coefficient was calculated. Correlograms of Moran's I showed significant positive values less than 0.100 for short-distance classes, indicating weak genetic structure. The genetic structure within the population is created by limited seed dispersal, and is probably weakened by overlapping seed shadow, secondary seed dispersal, extensive pollen flow and the thinning process. Genetic structure was detected in a western subplot of 50 x 200 m with immature soils and almost no dwarf bamboos (Sasa spp.), where small and intermediate-sized individuals were distributed in aggregations with high density because of successful regeneration. By contrast, genetic structure was not found in an eastern subplot of the same size with mature soils and Sasa cover, where successful regeneration was prevented, and the density of the small and intermediate-sized individuals was low. Moreover, genetic structure of individuals in a small-size class (diameter at breast height < 12 cm) was more obvious than in a large-size class (diameter at breast height >/= 12 cm). The apparent genetic structure detected in the 4-ha plot was therefore probably the result of the structure in the western portion of the plot and in small and intermediate-sized individuals that successfully regenerated under the favourable environment. The heterogeneity in genetic structure presumably reflects variation in the density that should be affected by differences in regeneration dynamics associated with heterogeneity in environmental conditions.     

Fire disturbance and forest structure in an old‐growth Pinus ponderosa forest,southern Cascades,USA

Questions: Did fire regimes in old‐growth Pinus ponderosa forest change with Euro‐American settlement compared to the pre‐settlement period? Do tree age structures exhibit a pattern of continuous regeneration or is regeneration episodic and related to fire disturbance or fire‐free periods? Are the forests compositionally stable? Do trees have a clumped spatial pattern and are clumps even‐ or mixed‐age? How might information from this old‐growth forest inform current restoration and management practices? Location: A 235‐ha old‐growth forest in the Ishi Wilderness, southern Cascade Mountains, California. Methods: Age, size, and spatial pattern of trees were quantified in seven stands. Fire history was reconstructed using fire scar dendrochronology. The influence of fire on stand structure was assessed by comparing fire history with age, size, and spatial structure of trees and identifying and measuring trees killed by two recent fires. Results: Species composition in plots was similar but density and basal area of tree populations varied. Age structure for P. ponderosa and Quercus kelloggii showed periods of episodic recruitment that varied among plots. Fire disturbance was frequent before 1905, with a median period between fires of 12 years. Fire frequency declined after 1905 but two recent fires (1990, 1994) killed 36% and 41% of mostly smaller diameter P. ponderosa and Q. kelloggii. Clusters of similar age trees occurred at scales of 28‐1018 m² but patches were not even‐aged. Interactions between tree regeneration and fire promoted development of uneven age groups of trees. Conclusions: Fire disturbance strongly influenced density, basal area, and spatial structure of tree populations. Fire exclusion over the last 100 years has caused compositional and structural changes. Two recent fires, however, thinned stands and created gaps favorable for Q. kelloggii and P. ponderosa regeneration. The effects of infrequent 20th century fire indicate that a low fire frequency can restore and sustain structural characteristics resembling those of the pre‐fire suppression period forest.     

Influence of site conditions on the survival of Fagus sylvatica seedlings in an old‐growth beech forest

Abstract. Following a full mast production in autumn 1995 in the old‐growth beech forest of la Tillaie (France), cupules, live and dead 1‐yr old seedlings were counted in September 1996 in 40 plots, representing a wide range of ecological conditions. The influence of geomorphology and forest stage on mast production and on seedling establishment and survival was studied. Maximum seedling abundance did not occur in areas with the largest cupule abundance, but this result was not statistically significant. The presence of a shallow sandstone table under Fontainebleau sand, where beech is far from its ecological optimum, positively influenced the production of mast but negatively influenced seedling survival. Contrary to expectation, the early biostatic rather than the late biostatic phase seemed to be the best stage for beech regeneration. Gaps also provide good environmental conditions for seedling establishment. Humus and light conditions, water availability and competition between beech individuals may explain the results. Light was the main factor influencing mast production in mature stages, and drought was an important factor causing seedling mortality. This study points out the importance of seedling survival for regeneration patterns in beech forests.     

Tree competition and species coexistence in a cool-temperate old-growth forest in southwestern Japan

Abstract. The growth dynamics and mode of competition between adult trees ≥ 4 cm in DBH (stem diameter at breast height 1.3 m) of eight abundant species occupying ca. 90 % of the total basal area were investigated in a 4-ha study plot (200 m × 200 m) of a cool-temperate, old-growth forest on Mount Daisen, southwestern Japan. In the study plot, 30 tree species with individuals ≥ 4.0 cm DBH co-occurred. A bimodal DBH distribution showing upper and lower-canopy layers was found for the most dominant and largest species, Fagus crenata (ca. 78 % of the total basal area), whilst other tree species showed unimodal DBH distributions corresponding mostly to the lower-canopy layer. We developed a model for individual growth incorporating both intra and interspecific competition and the degree of competitive asymmetry. Onesided interspecific competition was detected only from Fagus crenata (upper-canopy species) to Acer japonicum and Acanthopanax sciadophylloides (lower-canopy species) on the scale of the 4-ha study plot. Only Acanthopanax sciadophylloides showed symmetric intraspecific competition. However, a positive (non-competitive) interspecific relationship between adult trees prevailed over a competitive relationship; for example, individual DBH growth rate of Fagus crenata (especially lower-canopy trees) was correlated with the abundance of Acer mono. The positive relationship represented a group of species with similar habitat preference [soil type (mature or immature) caused by landslide disturbance and the presence/absence of Sasa dwarf bamboos in the understorey], where tree densities were not so high as to bring about competition. Competitive interactions between adult trees ≥ 4 cm in DBH occurred only locally between a few specific species and were suggested to be almost irrelevant to the variation in species coexistence on the 4-ha scale of cool-temperate forest. Rather, the coexistence of 30 tree species (species diversity) on this large scale was suggested to be governed by the regeneration pattern of each component species (habitat preference, seedling establishment, sapling competition) with respect to landslide disturbance.     

Distance decay in an old‐growth neotropical forest

Abstract. Questions: Does distance decay exist in an old‐growth neotropical forest? Is this distance decay stronger than expected due to environmental heterogeneity alone? At what spatial scales are distance decay manifested? Location: La Selva Biological Station, Costa Rica, Central America. Methods: An index of distance decay is applied appropriate for small quadrats (the probability of encountering a conspecific tree) to a grid of 1170 0.01‐ha plots. A null model is provided that accounts for environmental heterogeneity. Results: Significant, but weak, distance decay is found. After correcting for known patterns of environmental heterogeneity, the distance decay almost disappears, except for fine spatial scales. Conclusions: These results are inconsistent with models that predict distance decay at all spatial scales. However, biological processes leading to distance decay may be more relevant and detectable at scales broader than this study. Research utilizing objectively‐located samples over much broader scales is necessary to evaluate the generality and magnitude of distance decay.     

Fire disturbance and forest structure in old‐growth mixed conifer forests in the northern Sierra Nevada,California

Question: This study evaluates how fire regimes influence stand structure and dynamics in old‐growth mixed conifer forests across a range of environmental settings. Location: A 2000‐ha area of mixed conifer forest on the west shore of Lake Tahoe in the northern Sierra Nevada, California. Methods: We quantified the age, size, and spatial structure of trees in 12 mixed conifer stands distributed across major topographic gradients. Fire history was reconstructed in each stand using fire scar dendrochronology. The influence of fire on stand structure was assessed by comparing the fire history with the age, size, and spatial structure of trees in a stand. Results: There was significant variation in species composition among stands, but not in the size, age and spatial patterning of trees. Stands had multiple size and age classes with clusters of similar aged trees occurring at scales of 113 ‐ 254 m². The frequency and severity of fires was also similar, and stands burned with low to moderate severity in the dormant season on average every 9–17 years. Most fires were not synchronized among stands except in very dry years. No fires have burned since ca. 1880. Conclusions: Fire and forest structure interact to perpetuate similar stand characteristics across a range of environmental settings. Fire occurrence is controlled primarily by spatial variation in fuel mosaics (e.g. patterns of abundance, fuel moisture, forest structure), but regional drought synchronizes fire in some years. Fire exclusion over the last 120 years has caused compositional and structural shifts in these mixed conifer forests.     

A tree‐ring reconstruction of wind disturbances in a forest of the Slovakian Tatra Mountains,Western Carpathians

Question: Have past windstorm events influenced the structure and composition of mountain forests in the Tatra Mountains? Can severe and infrequent wind disturbances lead to dynamic coexistence of two tree species with different ecological requirements? Location: Subalpine mixed spruce‐larch forest at 1200‐1300 m a.s.l. in the Slovakian Tatra Mountains. A forested site affected by catastrophic large‐scale windthrow on 19 November 2004. Methods: Sixty‐seven spruce and 30 larch cross‐sections from the oldest cohorts were collected in a regular pattern in a 100‐ha plot. Tree‐ring series were analysed to reconstruct growth releases associated with past windthrows. A boundary‐line release criterion was applied to detect disturbance year. Spatial patterns of release signals were statistically detected with Mantel's test. We compared reconstructed years of disturbance events with historical records. Results: Releases in both species showed three main pulses. More than 85% showed major or moderate releases in 1865‐1879, 48% in 1915‐1924, and 25% in 1940‐1949. All of these disturbance events affected the whole 100‐ha area. Releases were spatially patterned in the first disturbances, but distributed randomly in the last. Releases co‐occurred in time with enhanced production of compression wood, suggesting disturbances were of wind origin. Reconstructed dates of windthrows were confirmed using historical data on storms. Conclusions: At least three windthrows of major and moderate severity took place in the last 150 years on southern slopes of the Tatra Mountains. This disturbance regime may contribute to coexistence of spruce and larch through differences in vulnerability and response to heavy windstorms.     

Earthquake impacts in old‐growth Nothofagus forests in New Zealand

Abstract. Six stands located on different land forms in mixed old‐growth Nothofagus forests in the Matiri Valley (northwest of South Island, New Zealand) were sampled to examine the effects of two recent large earthquakes on tree establishment and tree‐ring growth, and how these varied across land forms. 50 trees were cored in each stand to determine age structure and the cores were cross‐dated to precisely date unusual periods of radial growth. The 1968 earthquake (M = 7.1, epicentre 35 km from the study area) had no discernible impact on the sampled stands. The impact of the 1929 earthquake (M = 7.7, epicentre 20 km from the study area) varied between stands, depending on whether or not they had been damaged by soil or rock movement. In all stands, the age structures showed a pulse of N. fusca establishment following the 1929 earthquake, with this species dominating establishment in large gaps created by landslides. Smaller gaps, created by branch or tree death, were closed by both N. fusca and N. menziesii. The long period of releases (1929–1945) indicates that direct earthquake damage was not the only cause of tree death, and that many trees died subsequently most likely of pathogen attack or a drought in the early 1930s. The impacts of the 1929 earthquake are compared to a storm in 1905 and a drought in 1974–1978 which also affected forests in the region. Our results confirm that earthquakes are an important factor driving forest dynamics in this tectonically active region, and that the diversity of earthquake impacts is a major source of heterogeneity in forest structure and regeneration.     

Structure,dynamics and dendroecology of an old‐growth Fagus forest in the Apennines

Abstract. Question: Which are the structural attributes and the history of old‐growth Fagus forest in Mediterranean montane environments? What are the processes underlying their structural organization? Are these forests stable in time and how does spatial scale affect our assessment of stability? How do these forests compare to other temperate deciduous old‐growth forests? Location: 1600–1850 m a.s.l., Fagus forest near the tree line, central Apennines, Italy. Methods: An old‐growth Fagus forest was studied following historical, structural and dendroecological approaches. History of forest cover changes was analysed using aerial photographs taken in 1945, 1954, 1985 and 1994. The structural analysis was carried out in the primary old‐growth portion of the forest using 18 circular and two rectangular plots. Dendroecological analyses were conducted on 32 dominant or co‐dominant trees. Results: These primary old‐growth Fagus remnants consist of four patches that escaped logging after World War II. Both living and dead tree components are within the range of structural attributes recognized for old‐growth in temperate biomes. Dendroecological analyses revealed the roles of disturbance, competition and climate in structuring the forest. We also identified a persistent Fagus community in which gap‐phase regeneration has led to a mono‐specific multi‐aged stand at spatial scales of a few hectares, characterized by a rotated‐sigmoid diameter distribution. Conclusion: Even at the relatively small spatial scale of this study, high‐elevation Apennine Fagus forests can maintain structural characteristics consistent with those of old‐growth temperate forests. These results are important for managing old‐growth forests in the Mediterranean montane biome.     

Tree mortality and effects of release from competition in an old‐growth Fagus‐Abies‐Picea stand

Abstract. In a montane mixed Fagus‐Abies‐Picea forest in Babia Gora National Park (southern Poland), the dynamics of an old‐growth stand were studied by combining an 8‐yr annual census of trees in a 1‐ha permanent sample plot with radial increments of Abies and Picea growing in the central part of the plot. The mortality among the canopy trees was relatively high (10% in 8 yr), but the basal area increment of surviving trees slightly exceeded the losses caused by tree death. DBH increment was positively correlated with initial diameter in Abies and Picea, but not in Fagus. For individual trees smaller than the median height, basal area increment was positively related to the basal area of old snags and the basal area of recently deceased trees in their neighbourhood, but negatively related to the basal area of live trees. Dendrochronological analysis of the past growth patterns revealed numerous periods of release and suppression, which were usually not synchronized among the trees within a 0.3 ha plot. The almost normal distribution of canopy tree DBH and the small number of young individuals in the plot indicated that stand dynamics were synchronized over a relatively large area and, hence, were consistent with the developmental phase concept. On the other hand, the lack of synchronization among periods of growth acceleration in individual mature Abies and Picea trees conforms more closely to the gap‐dynamics paradigm.     

Availability of coarse woody debris in a boreal old‐growth Picea abies forest

Abstract. This study reports temporal (based on cross‐dated dead trees) and spatial patterns of availability of coarse woody debris (CWD) from Picea abies in a Swedish boreal landscape with discrete old‐growth forest patches in a wetland matrix. Data were collected from 29 patches ranging in size from 0.3 to 28 ha. A total of 897 dead trees with a minimum diameter of > 15 cm occurred on the 7.2 ha area analysed. The year of death was established for 50% of these trees. CWD volume ranged from 17 to 65 m³/ha for downed logs and from 0.5 to 13 m³/ha for standing snags. CWD of all decay stages and diameter classes occurred abundantly and the probability of finding logs of all decay stages and sizes was very high at the scale of single hectares. Tree mortality differed among 5 yr periods. However, during the last 50 yr no 5 yr period produced less than 3 logs/ha. Decay rates were highly variable among different logs. Logs with soft wood and some wood pieces lost (decay stage 5) died ca. 34 years ago. This suggests a fairly rapid decay in this northern forest. The data indicate a high and continuous availability of CWD of all types. It is likely, therefore, that selection pressures for efficient dispersal among CWD dependent species may not be very high. Consequently, species with narrow habitat demands and/or low dispersal ability may have evolved and this may contribute to the decrease of certain species in the managed landscape.     

Long-term growth dynamics of natural forests in Hokkaido,northern Japan

Abstract. The long-term growth dynamics of natural forest stands on the island of Hokkaido were described on the basis of an analysis of data from 38 permanent plots spanning 15–22 yr. Stand structure was characterized by basal area, stem density and tree size variability. To detect trends in stand structure, regression models for recruitment rate (per ha per yr), mortality rate and the rate of change in stem density and tree size variability were developed by a stepwise method using initial basal area, stem density, tree size variability, species composition summarized by LNMDS ordination, altitude, annual mean temperature, annual precipitation, type of understorey vegetation, topography and slope aspect as candidates for predictor variables. The same analyses were conducted for basal area increment (net growth) and its components: survivor growth = basal area gain by growth of surviving individuals and mortality = basal area loss by death of individuals. Stem density remained generally unchanged; recruitment was relatively low even in very sparse stands. Stand basal area generally increased as survivor growth was approximately double the mortality. Recruitment rate was strongly affected by the presence of dwarf bamboo (Sasa spp.) vegetation on the forest floor which inhibited tree regeneration. Mortality rate was density-dependent; dense stands had higher mortality than sparse stands. Density change rate (recruitment rate - mortality rate) was, therefore, determined by both the type of understorey vegetation and stem density. Survivor growth was high in stands with high stem density and basal area. Mortality was dependent on basal area and altitude. Net basal area increment (net growth) was dependent only on stem density with other factors that influenced survivor growth and mortality omitted. Tree size variability decreased in stands with high tree size variability whereas it increased in stands with low size variability. Based on the obtained models for density change rate and net basal area increment, trajectories of stands were illustrated on a log-log diagram of stem density and basal area. The predicted differences in trajectories as affected by the understorey vegetation type indicated the importance of dwarf bamboo vegetation for forest dynamics on Hokkaido.