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.     

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.     

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.     

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.     

四种常绿阔叶树种的抗寒性

以八角金盘、夹竹桃、阔叶十大功劳、樟叶槭4种常绿植物为试材,测定人工模拟低温条件下离体叶片可溶性糖、可溶性蛋白、游离脯氨酸、过氧化物酶活性、叶绿素含量和相对电导率的变化,结合自然越冬过程中叶片叶绿素指数(SPAD)、叶表面特征及解剖组织结构的变化,综合评价这4种常绿树种的抗寒能力.结果表明: 在人工低温胁迫过程中,4种常绿植物叶片叶绿素含量呈下降趋势,可溶性蛋白含量在-20 ℃出现高峰值,可溶性糖、游离脯氨酸、过氧化物酶活性和相对电导率整体呈上升趋势.八角金盘、夹竹桃、阔叶十大功劳和樟叶槭的低温半致死温度分别为-8.0、-13.4、-19.4和-14.8 ℃.自然越冬期间,4种植物叶片SPAD值变化幅度较大,显示出植物叶片中叶绿素相对含量变化对低温的适应性;同时,叶片厚度、角质层厚度、栅栏组织厚度和紧密度均有增加,并先后出现质壁分离,叶肉细胞中淀粉粒、草酸钙簇晶含量呈增高的趋势.夹竹桃叶片特有的气孔窝和窝内密被的非腺毛,阔叶十大功劳叶中的厚壁组织等在一定程度上提高了其抗寒性.叶片表面蜡质层破损、表皮毛断裂、栅栏组织形变、胞间空隙增大等现象的出现表明植物遭到了一定程度的低温伤害.     

Spatial associations of tree species in a subtropical evergreen broad-leaved forest

Aims The spatial segregation hypothesis and the low-frequency hypothesis are two important proposed mechanisms that delay or prevent competitive exclusion in ecosystems. Because tree species interact with their neighbors, the importance of these potential processes can be investigated by analyzing the spatial structures of tree species.Methods The distribution of the adults of 27 common tree species in a fully mapped 5-ha subtropical forest plot in Baishanzu, eastern China, was analyzed to investigate the community-level intra- and interspecific spatial association patterns. We first tested for the overall spatial pattern in the 5- to 40-m neighborhoods and classified first-order bivariate associations with a diametric scheme based on Ripley's K and nearest-neighbor statistic (G -function). Then heterogeneous Poisson null models were used to distinguish second-order interactions from overall spatial associations (including first-order effects). Finally, we analyzed correlations between the existence of species interactions and some attributes of the species involved.Important findings Partial overlap and segregation increased with scale, whereas mixing decreased. Nearly 70% of the species pairs occurred less than expected at random, and only 3.4% of the species pairs were well mixed; 11.0% of all species pairs showed significant small-scale interactions, which was a greater frequency than expected by chance if species are abundant or prefer the same habitat, but less frequent than expected if species are highly aggregated. This suggests that both spatial segregation and low frequency of species facilitate species coexistence by reducing the opportunity that trees of two species encounter each other. The study also revealed that positive interactions were more prevalent than negative interactions in the forest, which indicates that positive interactions may have important effects on forest species assemblies.     

东北温带森林20种乔木树种叶片干旱容忍性特征

全球范围内干旱频率和强度的增加严重影响树木生长,甚至导致森林大面积死亡。压力-容积(PV)曲线能够反映树木对干旱的容忍能力,但在局域尺度上尚未确定哪个PV曲线参数具有最优指示性。通过测定东北温带森林20种主要树种(包括16种被子植物和4种裸子植物)的PV曲线性状,包括质壁分离时的相对含水量(RWC_tlp)、失膨点叶水势(TLP)、饱和含水时的叶渗透势(π₀)、细胞弹性模量(ε)、叶水容(C_leaf)及叶结构性状(比叶面积和叶密度),研究局域尺度上叶片耐旱性的最佳指示性状,并分析叶片PV性状与结构性状间的相关性。结果表明: 被子植物的RWC_tlp 显著大于裸子植物,但其C_leaf 显著小于裸子植物,这表明用RWC_tlp和C_leaf可以指示东北温带森林不同功能型树种间耐旱性的大小。在被子植物中,TLP和π₀与叶密度呈显著负相关,且均与比叶面积呈显著正相关;而ε与比叶面积呈显著负相关。然而,裸子植物PV曲线性状与叶结构性状之间呈现与被子植物完全相反的趋势。裸子植物与被子植物树种之间PV曲线性状与叶结构性状关系的差异,可能归因于二者采取不同的干旱响应和适应策略。     

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.     

Nutrients limit photosynthesis in seedlings of a lowland tropical forest tree species

We investigated how photosynthesis by understory seedlings of the lowland tropical tree species Alseis blackiana responded to 10 years of soil nutrient fertilization with N, P and K. We ask whether nutrients are limiting to light and CO₂ acquisition in a low light understory environment. We measured foliar nutrient concentrations of N, P and K, isotopic composition of carbon (δ¹³C) and nitrogen (δ¹⁵N), and light response curves of photosynthesis and chlorophyll fluorescence. Canopy openness was measured above each study seedling and included in statistical analyses to account for variation in light availability. Foliar N concentration increased by 20% with N addition. Foliar P concentration increased by 78% with P addition and decreased by 14% with N addition. Foliar K increased by 8% with K addition. Foliar δ¹³C showed no significant responses, and foliar δ¹⁵N decreased strongly with N addition, matching the low δ¹⁵N values of applied fertilizer. Canopy openness ranged from 0.01 to 6.71% with a mean of 1.76 ± 0.14 (±1SE). Maximum photosynthetic CO₂ assimilation rate increased by 9% with N addition. Stomatal conductance increased with P addition and with P and K in combination. Chlorophyll fluorescence measurements revealed that quantum yield of photosystem II increased with K addition, maximum electron transport rate trended 9% greater with N addition (p = 0.07), and saturating photosynthetically active radiation increased with N addition. The results demonstrate that nutrient addition can enhance photosynthetic processes, even under low light availability.     

常绿阔叶树种的耐低温特性及其生态学评价

选取在南京地区已露地引种的 2 4种常绿阔叶树种 ,其自然分布北界止于亚热带不同的纬度 ,运用生理生态方法 ,测定它们在模拟低温处理下的电导率 ,并结合寒冷指数和引种后其适应性表现 ,分析这些树种的耐低温能力。结果表明 ,以某一树种自然分布北界气象资料为依据的寒冷指数与其自然植被的水温条件关系更为密切 ,寒冷指数有时不能反映树种本身耐低温能力的高低。如乐东拟单性木兰 ,其自然分布于亚热带南部 2 6°N,引种到南京耐低温表现明显好于自然分布于亚热带北部的香樟。而采用低温循环仪自动降温程序模拟冰冻处理来测定离体叶片的相对电导率 ,计算出各常绿阔叶树种叶片的半致死温度 ,能定量地反映这些树种本身的耐低温能力高低。在测试中如借助已知耐低温能力的树种作对照 ,其数据结果可作为常绿园林阔叶树种能否向更高纬度引种的重要参考依据。在上述研究基础上选出 6种 ,以其离体叶片为试材 ,测定其可溶性糖和游离脯氨酸含量 ,结果显示 ,随处理温度的降低 ,可溶性糖和游离脯氨酸的含量呈增加趋势 ,然而只有在适宜的处理下才呈现规律性变化 ,这些研究结果可作为树种耐低温能力的定性补充证据。对常绿阔叶树种的耐低温能力评价及预测有重要的参考价值     

Comparison of tree sprouting in three regeneration stages of an evergreen broadleaved forest in a karst landscape, SW China

Little is known about the role of tree sprouting in the regeneration of karst forest communities. In Shilin Stone Forest Geographical Park, southwestern China, all genets with the largest stem ?3 cm DBH (diameter at breast height) and/or stumps ?3 cm BD (basal diameter) were identified and number of sprouts counted in 10 transects (10 m × 100 m) in each of three evergreen broadleaved forest stands representing three regeneration stages (about 10, 20, and 30 years old). Species with >10 genets accounted for 72.4% of the 76 species, and all of them showed evidence of sprouting. One-third to two-thirds of the genets in the three forests were sprouting, with an average of 4.0–5.7 sprouts per sprouting genet. Sprouting capability (sprouting genets/total genets) and intensity (sprouts per sprouting genet) differed significantly among the three forest stages. More than 90% of the damaged genets were sprouting. The number of sprouts in a non-damaged genet was determined by intrinsic sprouting ability, and the number of sprouts in damaged genets was determined by stump size. As the forest developed, percentage of damaged genets increased, the portion of shoots ≥3 cm DBH co-existing in a genet decreased, and the portion of shoots <3 cm DBH sprouted from damaged genet increased. Thus the role of sprouting changed from contributing recruitment in the young stage to persistence in the later stage.     

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.     

Comparison of tree sprouting in three regeneration stages of an evergreen broadleaved forest in a karst landscape, SW China

Little is known about the role of tree sprouting in the regeneration of karst forest communities. In Shilin Stone Forest Geographical Park, southwestern China, all genets with the largest stem ?3 cm DBH (diameter at breast height) and/or stumps ?3 cm BD (basal diameter) were identified and number of sprouts counted in 10 transects (10 m × 100 m) in each of three evergreen broadleaved forest stands representing three regeneration stages (about 10, 20, and 30 years old). Species with >10 genets accounted for 72.4% of the 76 species, and all of them showed evidence of sprouting. One-third to two-thirds of the genets in the three forests were sprouting, with an average of 4.0–5.7 sprouts per sprouting genet. Sprouting capability (sprouting genets/total genets) and intensity (sprouts per sprouting genet) differed significantly among the three forest stages. More than 90% of the damaged genets were sprouting. The number of sprouts in a non-damaged genet was determined by intrinsic sprouting ability, and the number of sprouts in damaged genets was determined by stump size. As the forest developed, percentage of damaged genets increased, the portion of shoots ≥3 cm DBH co-existing in a genet decreased, and the portion of shoots <3 cm DBH sprouted from damaged genet increased. Thus the role of sprouting changed from contributing recruitment in the young stage to persistence in the later stage.     

Root ammonium transport efficiency as a determinant in forest colonization patterns: an hypothesis

Ratios of ammonium (NH₄⁺) to nitrate (NO₃^–) in soils are known to increase during forest succession. Using evidence from several previous studies, we hypothesize that a malfunction in NH₄⁺ transport at the membrane level might limit the persistence of early successional tree species in later seral stages. In those studies, ¹³N radiotracing was used to determine unidirectional fluxes and pool sizes of NH₄⁺ and NO₃^– in seedlings of the late-successional species white spruce ( Picea glauca ) and in the early successional species Douglas-fir ( Pseudotsuga menziesii var. glauca ) and trembling aspen ( Populus tremuloides ). At high external NH₄⁺, the two early successional species accumulated excessive NH₄⁺ in the root cytosol, and exhibited high-velocity, low-efficiency (15% to 22%), membrane fluxes of NH₄⁺. In sharp contrast, white spruce had low cytosolic NH₄⁺ accumulation, and lower-velocity but much higher-efficiency (65%), NH₄⁺ fluxes. Because these divergent responses parallel known differences in tolerance and toxicity to NH₄⁺ amongst these species, we propose that they constitute a significant driving force in forest succession, complementing the discrimination against NO₃^– documented in white spruce (Kronzucker et al. 1997).     

Distribution patterns of tree species in a Malaysian tropical rain forest

Abstract. Spatial patterns of tree species were studied in a 50-ha tropical rain forest plot in the Pasoh forest, Malaysia. This forest is characterized by a high diversity and very high number of rare species. Out of the 745 species occurring with > five individuals, 80.4 % had an aggregated distribution, 19.5% were randomly distributed and one species had a regular distribution. The spatial patterns of rare vs. common species, juvenile vs. adult trees, and coarse vs. fine scales were compared. Rare species are generally less aggregated than common ones and most of the randomly distributed species are rare. Spatial patterns shift from high clumping to looser intensity or random distribution when moving from juveniles to adults for the same species. No adult tree species display a regular pattern, however. Regular distributions were rarely found; this is probably due to intraspecific competition at a local scale. There is a negative correlation between per capita death rate and population density. This study suggests that the Pasoh forest and its high diversity are subjected to multiple controlling factors, e.g., topography, spacing effect, density-dependent processes and species rarity. The importance of any factor changes across spatial and temporal 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).