The effects of Sasa and canopy gap formation on tree regeneration in an old beech forest

Abstract. We examined the response of tree seedling emergence and survival to the dieback of Sasa and canopy gap formation in an old‐growth forest near Lake Towada, northern Japan. Synchronous death of Sasa occurred in 1995. We established four types of sampling sites differing in forest canopy conditions (Closed or Gap) and Sasa status (Dead or Live). Gap‐Dead sites had the highest light levels and the greatest fluctuation in soil temperatures. The death of Sasa alone facilitated the emergence (Acer japonicum, Fagus crenata, Fraxinus lanuginosa, and Tilia japonica) and survival (Acanthopanax sciadophylloides, F. crenata, F. lanuginosa, Kalopanax pictus, and Sorbus commixta) of species with a seedling bank strategy. Cercidiphyllum japonicum grew at all sites at a higher density than other species, but survived well only in Gap‐Dead sites. This behaviour was associated with a seed rain strategy. The additive effects of Sasa death and canopy gap formation promoted seedling emergence of pioneer tree species (Betula maximowicziana, Lindera umbellata, and Magnolia obovata), probably through break of dormancy by the large temperature fluctuation. In addition, the scarcity of advance regeneration in canopy gaps due to Sasa cover facilitates the regeneration of pioneer species. The dominance and dieback cycle of Sasa contributes to species diversity in this forest.     

Seed and seedling demography of invasive and native trees of subtropical Pacific islands

Abstract. Bischofia javanica is an invasive tree of the Bonin Islands in the western Pacific, Japan. This species has aggressive growth, competitively replacing native trees in the natural forest of the islands. The aim of this study was to examine seed and seedling factors which might confer an advantage to the establishment of Bischofia over native trees. During a 5‐yr period we compared the demographic parameters of early life history of Bischofia and Elaeocarpus photiniae‐folius, a native canopy dominant, in actively invaded forests. Predation of Elaeocarpus seeds by in troduced rodents was much higher before (27.9–32.9%) and after (41.3–100%) dispersal of seeds than that of B. javanica. Most Elaeocarpus seeds lost viability ca. 6 mo after burial in forest soil while some seeds of Bischofia remained viable for more than 2 yr. Seedling survival in the first 2 yr was much higher in Bischofia (16%) than in Elaeocarpus (1.3%). The high persistence of Bischofia in the shade, coupled to its rapid acclimation to high light levels, is an unusual combination because in forest tree species there is generally a trade‐off between seedling survival in the shade and response to canopy opening. Compared with a native canopy dominant, greater seed longevity, lower seed predation by introduced rodents, longer fruiting periods and the ability to form seedling banks under closed canopy appear to have contributed to the invasive success of Bischofia on the Bonin Islands.     

Structure,dynamics and disturbance regime of temperate broad-leaved forests in Japan

Abstract. Structure, diversity and dynamics of five Japanese temperate old-growth forests were compared, differing from each other in, i.a., climate, dominant tree type, topography, disturbance regime. The forests were Aya, Ogawa, Kanumazawa Riparian, Kanumazawa upland and Senju. A permanent plot (1–6 ha) was established in each forest and trees were censused several times at intervals of two years. Mean annual recruitment rates and mortality rates in these forests were both in the range of 0.5 to 4.6 %/yr at the community level. Analyses of the structure and dynamics of populations showed that the underlying process was different among the forests. Some forests experienced compositional shifts in their canopies, others had a constant canopy composition but appeared to lack effective regeneration in recent years. The recruitment rate appeared to be strongly affected by competitive undergrowth vegetation such as dwarf bamboo which has been controlled by natural disturbance or human impact. It is likely that the forests with mostly low recruitment rates had a low species diversity. The current variation in structure, diversity and dynamics of the studied forests might have been determined not only by physical conditions (e.g. climate) but also by chance factors (e.g. disturbance, outbreak of deer population).     

Plant traits as predictors of woody species dominance in climax forest communities

Abstract. The dominance of a given tree or shrub species in a particular forest community may be determined by many ecological traits of the target species, as well as those of the surrounding species as its potential competitors. The present study was conducted to evaluate the possibility of predicting community status (species composition and dominance) on the basis of traits of local flora using statistical methods, and to visualize the mathematical function which determines species dominance. A general linear model and logistic regression were used for the statistical analysis. Dependent variables were designated as dominance and presence/absence of species in climax forest, with independent variables as vegetative and reproductive traits. Subalpine, cool‐temperate, warm‐temperate and subtropical climax rain forests in East Asia were studied. Quantitative prediction of climax community status could readily be made based on easily measured traits of local flora. Species composition and 74.6% of the total variance of species dominance were predicted based on two traits; maximum height and shade tolerance. Through application of this method, the capacity of an alien species to invade a climax forest community could possibly be predicted prior to introduction of the alien species.     

Tropical forest structure and diversity in relation to altitude and disturbance in a Biosphere Reserve in central India

Questions: Is species diversity affected in protected areas where human activities are permitted or tolerated? On plots of a fixed size, does stem density alone predict number of species? Are differences in density related to disturbance and altitude? Location: Achanakmar‐Amarkantak Biosphere Reserve, central India. Methods: 42 sites, each with three replicate 10‐m radius plots, were examined. All trees (≥ 30 cm GBH) in each plot were measured for girth at breast height. α‐diversity, species richness and evenness were calculated for each site. The sites were ordinated by Nonmetric Multidimensional Scaling (NMS) using relative importance values of component species. Correspondence Analysis was used to broadly delineate communities. Anthropogenic disturbances were recorded in terms of percentage of trees lopped, scale of lopping, number of domestic livestock dung piles and foot trails (both livestock and people) for each plot. Results: The NMS analysis exhibited a near linear arrangement of sites with no evidence of discrete vegetation zones. NMS axes were significantly related to altitude and disturbance scores. With increasing elevation, basal area increased but number of species, α‐diversity and its components declined monotonically. The number of species and indices of species diversity were positively associated with tree lopping and also with total disturbance. Number of species was controlled by stem density only in plots not dominated by Shorea robusta. Conclusions: Recent levels of human disturbance are associated with higher species diversity in this biosphere reserve. There is some evidence that stands at all altitudes follow the same successional pattern to dominance by Shorea, a successional pattern that also results in decreased diversity without disturbance.     

Effects of disturbance intensity and frequency on early old‐field succession

Abstract. Early old‐field succession provides a model system for examining vegetation response to disturbance frequency and intensity within a manageable time scale. Disturbance frequency and intensity can interact with colonization and competition to influence relative abundance of earlier and later successional species and determine, respectively, how often and how far succession can be reset. We tested the joint effects of disturbance frequency and intensity on vegetation response (species richness, abundance, canopy structure) during the first six years of succession by clipping the dominant species (D) or all species (T) in spring and fall of each year (S), once per year in summer (Y1), each two years in summer (Y2), or each four years in summer (Y4). Vegetation response reflected disturbance effects on expansion of a later monospecific dominant perennial herb, Solidago altissima, and persistence of the early, richer flora of annuals. A more abundant and taller top Solidago canopy developed on plots clipped each 2 yr or less frequently. Plots clipped yearly or seasonally were richer, but had less abundant, shorter, and differently stratified canopy. Disturbance mediated the relative abundance of early and later successional species; however, frequency and intensity effects were not completely congruent. Persistence of a richer early successional flora increased through the most frequent disturbance (S), and was magnified by disturbance intensity. Disturbance as extreme as clipping all vegetation twice yearly did not cause a drop in species richness, but maintained the early successional community over the first six years of succession. We conclude that clipping disturbance influenced the rate of succession, but the early community could rebound through the range of disturbance frequency and intensity tested.     

Density-dependence and co-existence of conifer and broad-leaved trees in a Japanese northern mixed forest

Abstract. The growth and survival of coniferous trees (particularly Abies sachalinensis) and broad-leaved trees (particularly Quercus crispula) were followed over a 15-yr period in a 15.5-ha area in a northern mixed forest in Japan, and the coexistence of the two groups was simulated by a density-dependent projection matrix model. The density-dependent model assumes that the density effect of mother trees due to one-sided competition for light on smaller-sized tree regulates the demographic functions. The mother tree densities of conifers and broad-leaved trees have stronger negative effects on the recruitment and survival of seedlings of their own group than of the other group. These results support the idea of reciprocal replacement for conifer and broad-leaved trees. Simulations using the density-dependent model showed that the two groups will co-exist within a particular range of recruitment rates. However, the density of both groups did not affect the growth rate of any tree, and equilibrium DBH-distributions from density-dependent matrices were quite different from present distributions both for conifers and broad-leaved trees. On the other hand, equilibrium DBH-distributions of conifer and broad-leaved trees from density-independent matrices were quite distinct from each other, reflecting different survivorship curves of the two dominants. These results suggest that density-dependent processes are not so important for shaping population structures in this northern mixed forest.     

Seedling establishment of deciduous trees in various topographic positions

Abstract. We investigated the effect of topography‐related environmental factors (i.e., ground‐surface stability and soil moisture) on seedling establishment of 8 deciduous tree species. A field experiment was carried out using canopy species, which were classified into 3 groups based on the spatial distribution of adult trees (ridge, slope and valley). Demographic parameters were compared among species during the early stage of seedling establishment among 3 topographic positions in combination with gap and canopy conditions. The percentage of emerging seedlings tended to be lower on the ridge irrespective of the adult topographical distribution patterns. There was no clear trend in seedling emergence among the species groups classified by their spatial distribution. Seedling survival during 2 growing seasons was significantly different among species, topographic positions and light conditions. On the ridge, seedlings of the species dominating ridge tops had greater survival than those of other species, probably due to differences in demand for soil moisture. On the slope, frequent physical damage caused by surface material movement was observed and some species showed greater adaptability to the disturbed slope habitat. Survival of all seedlings was highest in the valley plots. Light conditions were the critical factor for seedling survival in some species. The results of this study suggest that topography creates diverse habitats for the establishment of tree seedlings. In addition to soil moisture, surface material movement may be a significant factor affecting seedling establishment.     

Effects of selective logging on tree diversity,composition and plant functional type patterns in a Bornean rain forest

Abstract. The effects of selective logging on tree diversity, changes in tree species composition and plant functional types were studied with the use of seven permanent plots in virgin and in logged forest. All plots were located in a lowland dipterocarp rain forest in East Kalimantan on the island of Borneo. Just after logging and during the following 20 yr tree diversity measured as Fishers’α was not significantly affected in logged forest plots. Temporal shifts in tree species composition were analysed with Principal Component Analysis (PCA). Logged forest plots had much larger changes over time than virgin forest plots. In the smallest diameter class, some logged forest plots showed a distinct trajectory in PCA space compared to virgin forest plots, while in larger diameter classes movement of logged plots in PCA space was random. This suggests that there is no predetermined community to which logged forest plots tend to shift when recovering from logging. We found a significant negative correlation between diameter increments and the species‐specific wood densities of tree species. Species‐specific wood density and potential tree height were used to assign species to five PFTs. As expected, logging increased the fraction of softwood stems in small diameter classes. In the largest diameter classes (≥ 50 cm DBH) a strong decrease of softwood emergent stems was found in logged forest plots. After more than 20 yr no recruitment was found of softwood emergent stems in selectively logged forest.     

Establishment of Fagus sylvatica and Fraxinus excelsior in an old-growth beech forest

Abstract. Distribution of tree seedlings, forest architecture, light conditions, ground vegetation and humus conditions were studied in a 45 m × 100 m area including multiple gaps in an old-growth beech forest. Gaps were created after some beech trees had been felled in severe storms in February 1990. A group of adult ash trees is found near the study site. The data were analyzed by Correspondence Analysis. Young seedlings (< 4 yr), of both Fraxinus (a sun species) and Fagus (a shade species), were most abundant under the crown of beech trees in semi-shade conditions, and where beech litter did not accumulate. Differences in the dissemination of Fraxinus and Fagus explained differences in the establishment of the two species. In contrast, older seedlings of beech established before the storms were more numerous in the gaps, suggesting a change in the ecological requirements of beech seedlings in the course of time.     

Seed bank spatial pattern in a temperate secondary forest

Abstract. Seed bank spatial pattern was studied in a secondary forest dominated by Fagus sylvatica and Betula celtiberica in the Urkiola Natural Park (N Spain). Soil samples were taken every 2 m in a regular grid (196 points) and divided into two fractions (0–3 cm and 3–10 cm deep). The viable seed bank was studied by monitoring seedling emergence for ten months. The effect of different factors on seed bank composition and patterning was analysed using constrained ordination as a hypothesis testing tool. Furthermore, the existence of spatial autocorrelation was evaluated by geostatistical analysis. Seed density was high, 7057 seed.m^?2, with a few species dominating. Species composition in the various layers were significantly correlated. The seed bank showed significant spatial structure, which was partially explainable by the spatial structure of the canopy and understorey vegetation. Spatial clumping from 0–8 m was observed in seed bank density and composition, mainly due to the pattern of two abundant taxa Juncus effusus and Ericaceae. The Ericaceae seed bank was related to the spatial distribution of dead stumps of Erica arborea. J. effusus was not present in the above‐ground vegetation, which indicates that its seed bank was formed in the past. As expected, the seed bank of this forest reflects its history, which is characterized by complex man‐induced perturbations. The seed bank appears to be structured as a consequence of contrasting driving forces such as canopy structure, understorey composition and structural and microhabitat features.     

Spatio‐temporal colonization patterns of forest plant species in a mixed deciduous forest

Abstract. An integrated analysis of the colonization patterns of forest plant species was carried out in a 34‐ha, mixed deciduous forest in northern Belgium. First, we sought to describe the relationships between land use history and environmental conditions. Land use history and soil type were related and negative correlations between pH and secondary forest age were found. The density of the shrub layer increases with secondary forest age. Litter quantity and cover of Urtica dioica were mainly indirectly influenced by land use history. Litter starts accumulating at low pH values and high shrub density and Urtica dioica grows vigorously on nutrient enriched soils where much light can reach the ground. Next, the importance of these human‐altered environmental conditions for the colonization of forest plant species was assessed relative to the importance of dispersal limitation. Therefore, the distribution of 16 forest species was mapped and species‐specific spatio‐temporal isolation measures were calculated. The analysis revealed that the colonization patterns of the slowly colonizing species (i.e. ‘ancient forest plant species’) are best explained by a combination of spatio‐temporal isolation, soil type, pH and the (non‐)cover of Urtica dioica. By contrast, spatio‐temporal isolation was never a limiting factor for good colonizing forest species. Our results suggest that colonization of ‘ancient forest plant species’ is hampered by a combination of dispersal‐ and recruitment limitation and that the relative importance of both factors is species‐specific.     

Relationships among the seed rain,seed bank and vegetation of a Hawaiian forest

Abstract. Hawaiian ecosystems are prone to invasion by alien plant species. I compared the seed rain, seed bank, and vegetation of a native Hawaiian forest to examine the potential role that seed ecology plays in allowing alien species to invade native forest. Absolute cover of seed plants in the forest was 126 %, annual seed rain was 5 713 seeds m^-2 yr^-1, and the mean density of seedlings emerging from the seed bank averaged across four seasons was 1 020/m². The endemic tree Metrosideros polymorpha was the most abundant species in the vegetation, seed rain and winter seed bank. Overall, native seed plants comprised 95 % of the relative cover in the vegetation and 99 % of the seeds in the seed rain, but alien species comprised 67 % of the seeds in the seed bank. Alien species tended to form persistent seed banks while native species formed transient or pseudo-persistent seed banks. Dominance of the seed bank by alien species with persistent seed banks suggests that aliens are favorably placed to increase in abundance in the vegetation if the forest is disturbed.     

Effects of trampling and vegetation removal on species diversity and micro-environment under different shade conditions

Abstract. The effects of disturbance by trampling and vegetation removal on herbaceous communities and their micro-environments were examined at two sites with different levels of shade. The dominant species of the original vegetation were the perennial herb Artemisia princeps at the sunny site and the dwarf-bamboo Pleioblastus chino at the shady site. With no disturbance and marked dominance of these species, diversity was low. After vegetation removal there was a rapid recovery, leading to a more diverse vegetation with many more species, each with a lower dominance. Continuous trampling induced a short vegetation while the light intensity under the foliage was increased. At the sunny site, species richness was depressed by trampling because the soil water potential decreased markedly during summer and only the stress-tolerant annual Digitaria adscendens grew vigorously. Thus, the lowest species diversity was observed in the heavily trampled vegetation in late summer. At the shady site, soil water availability was not affected by trampling. This allowed the survival of many species and prevented a strong decline in diversity. The results suggest that the pattern of change in diversity in communities subjected to various disturbances, was always determined by the original environments.     

Altitudinal zonation of evergreen broad-leaved forest on Mount Lopei,Taiwan

Abstract. Evergreen broad-leaved forest was studied in a transect on the northwestern slope of Mount Lopei in order to reveal altitudinal zonation in structure and floristic composition and the decisive environmental factors. 20 plots of 20 m × 20 m at altitudes from 540 m to 1320 m were analysed. 144 woody species were found. The results of a DCA ordination clearly pointed to a single dominant altitudinal gradient. Nevertheless, wind-exposure associated topography was found to account for additional variation for a given altitudinal range. Along the altitudinal gradient, four dominance-based forest types were recognized. Tree density, species diversity and evenness of the four types differed significantly but total basal area and tree volume were not significantly different. The 95 % turnover range for woody species as measured by the Community Coefficient was calculated as 1030 m, and the 50 % turnover range as 238 m. For the understorey, the change in species composition with altitude was less obvious. Species population structures of 57 sufficiently abundant species revealed four characteristic patterns, but most species showed a good fit to the negatively exponential or power function distribution and thus appeared to have good reproduction and regular recruitment. Both ANOVA and Redundancy Analysis (RDA) showed that significant differences among forest types were found for most soil variables. Organic C, exchangeable Na and K tended to increase with altitude, while pH and available N showed a reverse trend. There was little evidence that the differences in soil pH and available N were responsible for the variation in forest growth.     

Effects of age and distance on the composition of mixed deciduous forest fragments in an agricultural landscape

Abstract. Forest patches in central Belgium were inventoried twice for the presence or absence of forest plant species to study the effects of age and distance on species composition. All forests in the study area were subdivided based on their land use history. To avoid effects of autocorrelated environmental characteristics on species composition, habitat homogeneity was indirectly investigated using a TWINSPAN classification of the vegetation data. Two major habitats (alluvial and non‐alluvial forests) were distinguished and analysed separately. Patterns of species composition were investigated at the landscape level using Mantel tests. Species composition similarity measures were calculated between all pairs of fragments based on the floristic data. A highly significant correlation was found between species composition similarity and inter‐patch distance. Difference in age, which we used as a measure for habitat quality, was less important in explaining species composition patterns. The effects of spatial configuration became significant when difference in age was accounted for, and a partial correlation was calculated between inter‐patch distance and species composition similarity. Different results were found for alluvial and non‐alluvial forest types. Alluvial forests were more influenced by the spatial configuration than the non‐alluvial. For the non‐alluvial forest type effects measured with the difference in age between forest fragments obscured the effects of inter‐patch distance. Based on our findings we suggest that species composition is not only internally generated, but external processes such as differential colonization caused by varying degrees of isolation may be of overriding importance.     

西藏冷杉原始林林隙对草本植物和灌木树种多样性的影响

研究了西藏东南部亚高山原始冷杉林林隙对草本植物和灌木树种多样性的影响．结果表明,西藏东南部亚高山原始冷杉林林隙与非林隙林分内草本植物和灌木树种的种类组成、数量、密度和物种多样性均存在较大的差异．由于林隙内光照条件的改善,草本植物和灌木树种更加丰富;林隙内有喜光的草本植物和灌木,也有耐荫性较强的草本植物和灌木,而非林隙林分内仅有耐荫和较耐荫的草本植物和灌木;大部分林隙内的草本植物和灌木树种的物种多样性和均匀度大于非林隙林分;林隙内与非林隙林分的群落相似系数较小,表明林隙与非林隙林分草本植物和灌木群落组成存在差异．林隙大小在174．2～562．2m^2之间,群落相似系数与林隙大小无明显关联．林隙促进了草本植物和灌木树种的繁衍,在维持藏东南亚高山原始冷杉林的物种多样性方面起着重要作用．     

Early post-fire regeneration of a Pinus halepensis forest on Mount Párnis,Greece

Abstract. The post-fire regeneration of a 45-yr-old Pinus halepensis (Aleppo pine) forest, burned in July 1989, has been studied on Mount Párnis, Attiki, Greece. Four experimental plots at various slopes and exposures were established at altitudes of 400 - 450 m, and monitored for 3 yr at 3-month intervals. Early regeneration took place abundantly, through both resprouting and seed germination of mostly hard-seeded herbs and shrubs; the floristic richness was high with 80 taxa. Pine seedling emergence took place during the winter of the first post-fire year. The mean pine seedling density by the end of the recruitment period (March 1990) was 5–6 seedlings/m². This density decreased slightly during late spring and considerably during summer. During the second post-fire year only a relatively slight decline was observed; thereafter the density was stabilized to 1 - 2 seedlings/m². Mortality follows a negative exponential curve that levels off at ca. 20 %. Height distributions throughout the three post-fire years were all positively skewed as a result of the presence of few very tall saplings. A considerable fraction (20 %) of very short (5–15 cm) saplings were still alive 39 months after the fire; these may constitute the sapling bank. Based on the analysis of height distribution curves, it is concluded that the taller seedlings survived significantly better than the shorter ones.     

Soil seed bank and floristic diversity in a forest‐grassland mosaic in southern Spain

Abstract. Soil seed bank and floristic diversity were studied in a forest of Quercus suber, a forest of Quercus canariensis and a grassland, forming a vegetation mosaic in Los Alcornocales Natural Park, southern Spain. The soil seed bank was estimated by the germination technique. In each community patch, diversity, woody species cover and herbaceous species frequency was measured. Three biodiversity components – species richness, endemism and taxonomic singularity – were considered in the vegetation and the seed bank. Forest patches had a soil seed bank of ca. 11 200–14 100 seed.m^?2 and their composition had low resemblance to (epigeal) vegetation. The grassland patch had a more dense seed bank (ca. 31 800 seed.m^?2) and a higher index of similarity with vegetation, compared with the forests nearby. The complete forest diversity was 71–78 species on 0.1 ha, including 12–15 species found only in the seed bank; the grassland species richness was higher (113 species on 0.1 ha). We discuss the role of soil seed banks in the vegetation dynamics and in the complete plant biodiversity of the mosaic landscape studied.     

Growth of Fagus sylvatica saplings in an old-growth forest as affected by soil and light conditions

Abstract. Studies were conducted on 41 five yr-old common beech (Fagus sylvatica) saplings collected in an old-growth beech wood (Fontainebleau forest, biological reserve of La Tillaie, France), under varying humus and light conditions, following gypsy moth (Lymantria dispar) caterpillar injuries. Aerial and subterranean parts of each sapling were described by means of 34 parameters and environmental conditions at the microsite, where each sapling was excavated, were characterized by 23 parameters. The development of beech saplings is strongly affected by microsite conditions. An increase in sapling size was associated with darkness of the A-horizon, typical of zones with poor mineralization of organic matter. Light conditions were more important in influencing the development of the root system than that of the aerial parts. Rooting depth was shallower and rate of mycorrhiza development by the black ascomycete Cenococcum geophilum was lower in microsites receiving incident light during the morning than in those never receiving incident light during this period. Results are discussed in the frame of survival of young beech individuals in varying environmental conditions, when submitted to competition by other vegetation and adverse climate conditions.