Distribution patterns of tree species in an evergreen broadleaved forest in eastern China

Ecological assembly rules in evergreen broad-leaved forest are far from clear understanding. Spatial dispersion of individuals in a species is central in ecological theory. We analyzed the spatial patterns as well as associations between adult and juvenile of each tree species in a 5-ha subtropical evergreen broad-leaved forest plot in eastern China. Out of the 74 species occurring with more than 10 individuals, 88.4% of these species are aggregated. Most of them are aggregated from small to large scales. Spatial distributions of some species corre-spond with topography. Many bad dispersed species in the Baishanzu exhibit a highly aggregated distribution at small scales. These suggest that environmental heterogeneity and/or dispersal limitation may be the most important mechanisms that control the distribution patterns of these species. Our observations of the aggregations of abundant species basically support the hypothesis that dispersal limitation decreases as the number of reproductive trees increases. The rest species are randomly distributed, with less than 10 individuals. For most common species, spatial aggregation is weaker in larger diameter classes, and the distance between adults is larger than that between juveniles and adults, suggesting that density-dependence works on loosing aggregation and excluding conspecific juveniles away from adult trees. However, the density-dependent effect is not strong enough to eliminate all seedlings near adult trees and to result in a regular distribution of trees; thus the density dependence is usually masked by the refuge effect.     

Distribution patterns of tree species in an evergreen broadleaved forest in eastern China

Ecological assembly rules in evergreen broad-leaved forest are far from clear understanding. Spatial dispersion of individuals in a species is central in ecological theory. We analyzed the spatial patterns as well as associations between adult and juvenile of each tree species in a 5-ha subtropical evergreen broad-leaved forest plot in eastern China. Out of the 74 species occurring with more than 10 individuals, 88.4% of these species are aggregated. Most of them are aggregated from small to large scales. Spatial distributions of some species correspond with topography. Many bad dispersed species in the Baishanzu exhibit a highly aggregated distribution at small scales. These suggest that environmental heterogeneity and/or dispersal limitation may be the most important mechanisms that control the distribution patterns of these species. Our observations of the aggregations of abundant species basically support the hypothesis that dispersal limitation decreases as the number of reproductive trees increases. The rest species are randomly distributed, with less than 10 individuals. For most common species, spatial aggregation is weaker in larger diameter classes, and the distance between adults is larger than that between juveniles and adults, suggesting that density-dependence works on loosing aggregation and excluding conspecific juveniles away from adult trees. However, the density-dependent effect is not strong enough to eliminate all seedlings near adult trees and to result in a regular distribution of trees; thus the density dependence is usually masked by the refuge effect.     

Winter photosynthesis by saplings of evergreen broad-leaved trees in a deciduous temperate forest

* Here we investigated photosynthetic traits of evergreen species under a deciduous canopy in a temperate forest and revealed the importance of CO2 assimilation during winter for annual CO2 assimilation. * Saplings were shaded by the canopy trees from spring through to autumn, but were less shaded during the winter months. Photosynthetic rates at light saturation (Aarea) were lower during winter than during the growing season. Aarea was higher in Camellia, Ilex and Photinia than in Castanopsis, Cleyera and Quercus during the winter, but differed little during summer and autumn. * Estimated daily CO2 assimilation (Aday) was higher during the winter than during the growing season in Camellia, Ilex and Photinia but was higher than that during the growing season only at the beginning and end of winter in Castanopsis, Cleyera and Quercus. Aday was higher in Camellia, Ilex and Photinia than in Castanopsis, Cleyera and Quercus but differed little among them during the growing season. * These results reveal the importance of winter CO2 assimilation for the growth of Camellia, Ilex and Photinia. Furthermore, differences in annual CO2 assimilation among species are strongly modified by species-specific photosynthetic traits during the winter under deciduous canopy trees.     

The regeneration of pioneer tree species under browsing pressure of Sika deer in an evergreen oak forest

The effects of Sika deer (Cervus nippon) browsing on the regeneration of pioneer species were studied in relation to canopy gaps in a warm temperate evergreen oak forest in Kasugayama, Nara City. Four study sites, three in canopy gaps and one under a closed canopy, were selected and each divided into fenced and unfenced plots. Under the closed canopy, seedlings of all the pioneer species died irrespective of browsing pressure. However, in the canopy gap sites, seedlings of the pioneer species could establish and grow well. The seedling survival ratio in the fenced plots in the canopy gaps was >60% 1 year after germination. However, in the unfenced plots, only <20% of the seedlings survived 1 year, with all dying within 3 years after germination. Thus, the regeneration of pioneer trees in this forest was strongly inhibited by deer browsing. Successful regeneration of a pioneer,Zanthoxylum ailanthoides, occurred for several years even after two major wind disturbances during the past 90 years. This may be due to less browsing pressure from the deer.     

Shade tolerance and regeneration of some tree species of a tropical rain forest in Western Kenya

Shadetolerance and regeneration of some heavily exploited timber species(Uvariopsis congensis, Antiaris toxicaria, Funtumia africana,Prunus africana and Aningeria altissima)were studied in two adjacent sites in Kakamega tropical rain forest. The twosites have been subjected to different logging intensities. Within one of thesites, relatively intact areas were compared with adjoining more disturbedones.Seedling presence or absence in mature forest phase, sapling proportions in gapand the mature phase and diameter distributions were used to classify thespecies into shade tolerance groups. Results indicate that except forUvariopsis, the species are non-pioneer lightdemanders and could fit into three shade tolerance groups: A groupresembling pioneers but with seedling in shade, a mid-tolerant group andone showing a high shade tolerance. Amount of regeneration and disturbance wererelated: regeneration being higher in the more disturbed site except forUvariopsis. The intersite differences were mirrored whendisturbed and adjoining more intact areas within one of the sites werecompared.Species more abundant in the disturbed site also generally scored low in shadetolerance. However, the mid-tolerant Funtumia andnot Prunus or Aningeria that hadlowershade tolerance scores, dominated regeneration in the disturbed forest parts.The shade tolerance data suggest that Uvariopsis, Funtumiaand Antiaris could be amenable to natural forestmanagement. For Prunus and Aningeria,artificial regeneration in buffer plantations around the natural forest may beabetter option.     

Spatial patterns and associations of four species in an old-growth temperate forest

The spatial pattern of a tree species is an important characteristic of plant communities, providing critical information to explain species coexistence. The spatial distribution and association of four different successional species were analyzed among different life-history stages in an old-temperate forest. Significant aggregation patterns were found, and the degree of aggregation decreased with the scales and life-history stages. Significant interspecific spatial associations were detected. In comparing the relationships among the different life-history stages, positive associations were found at small scales in all of the juvenile species pairs. In the adult stage, negative associations were detected in coniferous vs. deciduous species pairs, while the deciduous species pairs, which have identical resource requirements, showed a positive association in this study. The coniferous species pairs showed a positive association at small scales. We infer that seed dispersal, competitive ability, or the requirement for specific topographic and light environments may contribute to the coexistence of these species.     

Modeling qualitative and quantitative elements of branch growth in saplings of four evergreen broad-leaved tree species growing in a temperate Japanese forest

Key message

Factors influencing branch growth differed between a qualitative element (whether a branch grows) and a quantitative element (the amount of biomass accretion).

Abstract

We measured branch growth in saplings of four evergreen broad-leaved tree species growing in a temperate Japanese forest. A hierarchical Bayesian approach was used to model two elements of branch growth: a qualitative component [whether branches produce new annual shoots (AS_n)] and a quantitative component (total AS_n mass). The two components were influenced by somewhat different factors. The probability that a branch would produce AS_n was affected by the total mass of previous-year annual shoots (total AS_n–1 mass), branch age, relative branch height, canopy openness measured at the branch tip, maximum canopy openness within a single sapling, and the interaction between canopy openness and maximum canopy openness. The total AS_n mass was influenced by the total AS_n–1 mass, relative branch height, branch inclination, and maximum canopy openness. These two components of branch growth should be considered separately when tree architecture is modeled. In addition, we detected interactive responses among branches within individual saplings. The relative importance of interactive modular response and branch autonomy may differ depending on the status of plants, such as individual age, environmental conditions, and the timing of the measurements. We found considerable similarities in the responses of branch growth to the internal and environmental factors among the species studied. We also found some among-species differences in branch growth responses to the explanatory variables measured.     

Population structure and spatial patterns for trees in a temperate old-growth evergreen broad-leaved forest in Japan

The population structure and spatial pattern of major tree species in a warm-temperate old-growth evergreen broad-leaved forest in the Tatera Forest Reserve of Japan were investigated. All stems 5 cm in diameter at breast height (DBH) were mapped on a 4 ha plot and analyses were made of population structure and the spatial distribution and spatial association of stems in different vertical layers for nine species. This was done in the context of scale dependency. The plot was located on a very gentle slope and 17.1% of its canopy layer was in gaps. It contained 45 woody plant species and 4570 living stems with a basal area of 63.9 m² ha^–1. Castanopsis cuspidata var. sieboldii, the most dominant species for the basal area, had the maximum DBH among the species present, fewer smaller stems and a lower coefficient of statistical skewness of the DBH distribution. The second most dominant species, Dystilium racemosum, had the highest stem density (410 ha^–1), more abundant smaller stems and a relatively higher coefficient of skewness. Most stems in different vertical layers showed a weakly aggregated distribution with loose colonies as basic units. Gap dependency for the occurrence of stems under the canopy layer was weak. Maximum slope degree of the plot also weakly affected the occurrence of stems. Spatial associations varied among intra- and interspecific cohorts in the different layers and spatial scales examined, and positive associations among cohorts were found more frequently as the scales examined became larger. This tendency suggests that key factors forming observed spatial associations might vary with the spatial scales.     

Establishment of the evergreen broad-leaved tree species Castanopsis cuspidata in an abandoned secondary forest in western Japan

Recently, populations of Castanopsis cuspidata have often expanded into secondary forests in western Japan. To determine the establishment processes of this species, we analyzed its spatial distribution in a secondary forest dominated by Quercus variabilis and Quercus serrata that is located adjacent to a stand dominated by C. cuspidata. Saplings, defined as ≥30 cm stem length (SL) and <5 cm diameter at breast height (DBH), were abundant and their size distribution was inversely J-shaped, indicating continuous recruitment. Although seedlings (SL < 30 cm) and small saplings (30 ≤ SL < 50 cm) of C. cuspidata were aggregated near flowering trees of this species, some were found ≥40 m from the nearest adults, suggesting that there is animal-aided dispersal of acorns. The distribution of larger-sized individuals (≥100 cm SL) of C. cuspidata was unrelated to distance from the nearest flowering C. cuspidata or dominant Quercus species (≥5 cm DBH), but was associated with dead Pinus densiflora trees, which were abundant at the site. Thus, the establishment of C. cuspidata in this forest is controlled mainly by two factors, viz. patterns of acorn dispersal by animals, and forest disturbance regime (i.e., deaths of pine trees).     

The effect of tree species diversity on fine-root production in a young temperate forest

The phenomenon of overyielding in species-diverse plant communities is mainly attributed to complementary resource use. Vertical niche differentiation belowground might be one potential mechanism for such complementarity. However, most studies that have analysed the diversity/productivity relationship and belowground niche differentiation have done so for fully occupied sites, not very young tree communities that are in the process of occupying belowground space. Here we used a 5–6 year old forest diversity experiment to analyse how fine-root (<2 mm) production in ingrowth cores (0–30 cm) was influenced by tree species identity, as well as the species diversity and richness of tree neighbourhoods. Fine-root production during the first growing season after the installation of ingrowth cores increased slightly with tree species diversity, and four-species combinations produced on average 94.8% more fine-root biomass than monocultures. During the second growing season, fine-root mortality increased with tree species diversity, indicating an increased fine-root turnover in species-rich communities. The initial overyielding was attributable to the response to mixing by the dominant species, Pseudotsuga menziesii and Picea abies, which produced more fine roots in mixtures than could be expected from monocultures. In species-rich neighbourhoods, P. abies allocated more fine roots to the upper soil layer (0–15 cm), whereas P. menziesii produced more fine roots in the deeper layer (15–30 cm) than in species-poor neighbourhoods. Our results indicate that, although there may be no lasting overyielding in the fine-root production of species-diverse tree communities, increasing species diversity can lead to substantial changes in the production, vertical distribution, and turnover of fine roots of individual species.     

Direct and indirect effects of understorey bamboo shape tree regeneration niches in a mixed temperate forest

Plant cover plays a major role in shaping the nature of recruitment microsites through direct (resource mediated) and indirect (consumer mediated) interactions. Understorey plants may differentially affect seedling establishment, thus contributing to regeneration-niche separation among canopy tree species. We examined patterns of early tree seedling survival resulting from interactive effects of understorey bamboo (Chusquea culeou) and resident consumers in a mixed temperate Patagonian forest, Argentina. Newly germinated seedlings of Nothofagus dombeyi and Austrocedrus chilensis were planted in bamboo thickets and non-bamboo patches, with or without small-vertebrate exclosures. We found species-specific patterns of seedling survival in relation to bamboo cover. Nothofagus survival was generally low but increased under bamboo, irrespective of cage treatment. Desiccation stress accounted for most Nothofagus mortality in open, non-bamboo areas. In contrast, Austrocedrus survival was highest in non-bamboo microsites, as most seedlings beneath bamboo were killed by small vertebrates through direct consumption or non-trophic physical damage. There was little evidence for a negative impact of bamboo on tree seedling survival attributable to resource competition. The balance of simultaneous positive and negative interactions implied that bamboo presence facilitated Nothofagus early establishment but inhibited Austrocedrus recruitment via apparent competition. These results illustrate the potential for dominant understorey plants to promote microsite segregation during early stages of recruitment between tree seedlings having different susceptibilities to water stress and herbivory. We recognise, however, that patterns of bamboo–seedling interactions may be conditional on moisture levels and consumer activity during establishment. Hence, both biotic and abiotic heterogeneity in understorey environments should be incorporated into conceptual models of regeneration dynamics and tree coexistence in forest communities.     

Local-scale drivers of tree survival in a temperate forest

Tree survival plays a central role in forest ecosystems. Although many factors such as tree size, abiotic and biotic neighborhoods have been proposed as being important in explaining patterns of tree survival, their contributions are still subject to debate. We used generalized linear mixed models to examine the relative importance of tree size, local abiotic conditions and the density and identity of neighbors on tree survival in an old-growth temperate forest in northeastern China at three levels (community, guild and species). Tree size and both abiotic and biotic neighborhood variables influenced tree survival under current forest conditions, but their relative importance varied dramatically within and among the community, guild and species levels. Of the variables tested, tree size was typically the most important predictor of tree survival, followed by biotic and then abiotic variables. The effect of tree size on survival varied from strongly positive for small trees (1-20 cm dbh) and medium trees (20-40 cm dbh), to slightly negative for large trees (>40 cm dbh). Among the biotic factors, we found strong evidence for negative density and frequency dependence in this temperate forest, as indicated by negative effects of both total basal area of neighbors and the frequency of conspecific neighbors. Among the abiotic factors tested, soil nutrients tended to be more important in affecting tree survival than topographic variables. Abiotic factors generally influenced survival for species with relatively high abundance, for individuals in smaller size classes and for shade-tolerant species. Our study demonstrates that the relative importance of variables driving patterns of tree survival differs greatly among size classes, species guilds and abundance classes in temperate forest, which can further understanding of forest dynamics and offer important insights into forest management.     

Drought acclimation of two deciduous tree species of different layers in a temperate forest canopy

Water-use strategies of Populus tremula and Tilia cordata, and the role of abscisic acid in these strategies, were analysed. P. tremula dominated in the overstorey and T. cordata in the lower layer of the tree canopy of the temperate deciduous forest canopy. Shoot water potential (), bulk-leaf abscisic acid concentration ([ABA]_leaf), abscisic acid concentration in xylem sap ([ABA]_xyl), and rate of stomatal closure following the supply of exogenous ABA (v) decreased acropetally through the whole tree canopy, and foliar water content per area (w), concentration of the leaf osmoticum (c), maximum leaf-specific hydraulic conductance of shoot (L), stomatal conductance (g_s), and the threshold dose per leaf area of the exogenous ABA (d_a) required to reduce stomatal conductance increased acropetally through the tree canopy (from the base of the foliage of T. cordata to the top of the foliage of P. tremula) in non-stressed trees. The threshold dose per leaf dry mass of the exogenous ABA (d_w) required to reduce stomatal conductance, was similar through the tree canopy. After a drought period (3 weeks), the , w, L, g_s, d_a and d_w had decreased, and c and v had increased in both species. Yet, the effect of the drought period was more pronounced on L, g_s, d_a, d_w and v in T. cordata, and on , w and c in P. tremula. It was concluded that the water use of the species of the lower canopy layer—T. cordata, is more conservative than that of the species of the overstorey, P. tremula. [ABA]_leaf had not been significantly changed in these trees, and [ABA]_xyl had increased during the drought period only in P. tremula. The relations between [ABA]_leaf, [ABA]_xyl and the stomatal conductance, the osmotic adjustment and the shoot hydraulic conductance are also discussed.     

长白山次生林乔木树种空间分布格局

应用RipleyK函数,分析了长白山一个5．2hm^2次生林样地内乔木树种的空间分布以及主要乔木树种的幼树、小树、大树的空间分布和空间关系。结果表明：共调查到34种乔木,其中常见种27种,少见种7种,共计15688株;在27个常见种中,有23个种在空间上主要呈聚集分布,另外4个种主要表现为随机分布;在7个少见种中,均主要表现为随机分布;簇毛槭、假色槭、水曲柳、臭松和红松随着种群发育（幼树→小树→大树）聚集强度逐渐减弱,直至趋向于随机分布,其他树种的空间分布随种群发育未表现出明显的变化规律;在10个主要树种组成的45个种对中,有11个种对表现出显著正相关,1个种对为显著负相关。     

Structure and regeneration of canopy species in an old-growth evergreen broad-leaved forest in Aya district,southwestern Japan

The population structure and regeneration of canopy species were studied in a 4 ha plot in an old-growth evergreen broad-leaved forest in the Aya district of southwestern Japan. The 200 m × 200 m plot contained 50 tree species, including 22 canopy species, 3,904 trees (dbh5 cm) and a total basal area of 48.3 m²/ha. Forty one gaps occurred within the plot, and both the average gap size (67.3 m²) and the total area of gap to plot area (6.9%) were small. Species found in the canopy in the plot were divided into three groups (A, B, C) based on size and spatial distribution patterns, and density in each tree size. Group A (typical species: Distylium racemosum, Persea japonica) showed a high density, nearly random distribution and an inverse J-shaped size distribution. Species in group B (Quercus salicina, Quercus acuta, Quercus gilva) were distributed contagiously with conspicuous concentration of small trees (<5 cm dbh) around gaps. However, the species in this group included few trees likely to reach the canopy in the near future. Group C included fast-growing pioneer and shade intolerant species (e.g. Cornus controversa, Carpinus tschonoskii, Fagara ailanthoides), which formed large clumps. Most gaps were not characterized by successful regeneration of group B and C but did appear to accelerate the growth of group A. Group B species appear to require long-lived or large gaps while group C species require large, catastrophic disturbances, such as landslides, for regeneration.     

Habitat niche specialization in an understory species in a warm temperate forest

Relationships between microhabitat variables; understory light conditions in summer and winter, altitude, slope inclination and topographic categories (valley, ridge, and slope) and the distribution of Aucuba japonica Thunb. (Cornaceae), a common understory shrub species in Japan were examined using non-contagious 66, 20 × 20 m² quadrats. The Morishita’s I _δ suggested that A. japonica distributions were strongly heterogeneous among the quadrats. Therefore positive spatial autocorrelation of A. japonica at a within-quadrat level (≤20 m) was obvious. Moran’s I statistics showed a significant positive spatial autocorrelation in A. japonica abundance within the distance shorter than 60 m. But the partial Mantel tests suggested that the mass effect from neighboring quadrats would little explain A. japonica abundance in a quadrat. The partial Mantel tests also clearly showed that A. japonica distributions were strongly structured by topography and understory light conditions. Using Monte Carlo randomization tests, we found that A. japonica was aggregately distributed in quadrats in valley which were covered by deciduous canopies. Better understory light conditions in winter together with valley edaphic conditions may increase the abundance of A. japonica there. It is concluded that habitat niche specialization is important in structuring distribution of A. japonica in this forest community.