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.     

暖温带落叶阔叶林动态变化的模拟研究

用森林动态林窗模型 FORET1模拟了暖温带落叶阔叶林的长期变化特征。模型参数取自暖温带地区长期森林研究和经营的历史数据 ,对过去数据中缺少的参数进行了实地测定 ,并用观测的数据对模型作了检验。结果表明模型能较好地模拟暖温带落叶阔叶林的长期动态变化特征。通过模拟可以看出 ,森林的净初级生产力没有明显变化规律且极度不稳定 ,峰值出现在30 a左右 ,相似于世界上其它地区森林动态格局变化 ,生物量格局呈循环状态变化 ,循环周期大致在 110 a左右。     

Population dynamics of coniferous and broad-leaved trees in a Japanese temperate mixed forest

The growth and survival of coniferous and broad-leaved trees were followed over a 5-yr period in a temperate old-growth mixed forest in Japan, and dynamic features of the forest were studied in relation to the life history of the dominants, the coniferous Abies homolepis and the broad-leaved Fagus crenata. During this period, the gap formation rate was 31m² ha^?1yr^?1, the mortality of trees > 2m high was 1.7%/yr, and the rate of loss in basal area 1.4%/yr. These values were much higher than the recruitment, 0.3%/yr, and the total growth of surviving and new trees, 0.6%/yr, owing to the inhibition of regeneration by understorey dwarf bamboo (Sasa borealis). A transition matrix model based on DBH size classes predicts that the basal area of the forest will decrease by 14% in 50 yr, but that the DBH distribution of trees > 10 cm diameter will change little. Equilibrium DBH distributions assuming recruitment being equal to mortality, were quite different between broad-leaved and coniferous trees, reflecting different survivorship curves of the two dominants. The composition and structure of the forest may change depending on the pattern and frequency of disturbances, or episodic events, notably the synchronous death of Sasa borealis.     

Seed demography of three co-occurring Acer species in a Japanese temperate deciduous forest

Abstract. Seed demography of three co-occurring Acer species in an old-growth mixed deciduous forest in Japan was studied. Almost all of the seeds of A. mono germinated in the first spring, while those of A. palmatum var. amoenum showed a delay in germination of almost one year. A. rufinerve showed a rather opportunistic germination habit. Both A. palmatum var. amoenum and A. rufinerve form short-term persistent seed banks, but without input of newly dispersed seeds they may become extinct in about one year. The seed bank for these two species is not as significant as for a typical pioneer species, and the seedling bank is important for all three species. Only a small proportion of the dispersed propagules contained viable embryos, mainly due to pollination failure or abortion (A. mono and A. palmatum var. amoenum), and invertebrate predation (A. rufinerve). For all three species, larger seed crops had a higher percentage of viable seeds. Even for these relatively small, wind-dispersed seeds, the predation pressure was very high. A large part of the dispersed seeds was eaten by wood mice during the first winter (30–80 %). Estimation from the 5-yr average of seed dispersal and seedling emergence showed that only 7–16 % of the dispersed viable seeds succeeded in germinating.     

Pollination systems in a warm temperate evergreen broad-leaved forest on Yaku Island

Animal pollination in a warm temperate evergreen broad-leaved forest was observed on Yaku-shima Island, south of Kyushu, Japan. Three groups of plants were categorized: canopy-flowering tree species, understory-flowering tree species, and climber and epiphyte species. Each of these formed different pollination systems. The canopy-flowering tree species had shallow, dish-shaped flowers and utilized various types of opportunistic pollinators. Most of the climber and epiphyte species had deep, tube-shaped flowers and specialized pollinators, although some climber species which bloomed in the canopy especially in winter, had opportunistic pollinators. The understory-flowering tree species had large dish- or funnel-shaped flowers and endothermic pollinators able to tolerate the dark and cold conditions under the canopy. The individual trees of canopy-flowering tree species produced large numbers of flowers simultaneously (mass-flowering) and had a well synchronized flowering period. Each canopy-flowering tree species segregated its flowering time from those of the anothers. Climber and epiphyte species and most of the understory-flowering tree species produced small numbers of flowers sequentially (extended flowering) and showed a long flowering period.     

CO<Subscript>2</Subscript> exchange in a temperate Japanese cypress forest compared with that in a cool-temperate deciduous broad-leaved forest

To examine the characteristics of carbon exchange in coniferous forests, we analysed the seasonal and diurnal patterns of CO₂ exchange, as measured using the eddy covariance method, in a Japanese cypress forest in the Kiryu Experimental Watershed (KEW) in central Japan. The net CO₂ exchange data during periods of low-friction velocity conditions and during periods of missing data were interpolated. The daily CO₂ uptake was observed throughout the year, with maximum values occurring in early summer. Periods of low carbon uptake were seen in late summer owing to high respiratory CO₂ efflux. The diurnal and seasonal patterns of daytime CO₂ exchange at KEW were compared with those in a cool-temperate deciduous forest of the Tomakomai Experimental Forest (TOEF) in Japan. The environmental differences between evergreen and deciduous forests affected the seasonal patterns of carbon uptake. Although there were great differences in the mean monthly air temperatures between the sites, the mean monthly daytime carbon uptake was almost equal at both sites during the peak growing period. The carbon-uptake values at the same PAR level were greater before noon than after noon, especially at TOEF, suggesting the stomatal regulation of carbon uptake.     

Sap flux of five co-occurring tree species in a temperate broad-leaved forest during seasonal soil drought

In an old-growth forest in Central Germany, sap flux was studied in five broad-leaved tree species that were assumed to differ in drought sensitivity. Under moist soil conditions, average daily sap flux density (J _s) in the outermost xylem varied by a factor of 2.3 among the species (67–152 g cm⁻² per day, n=5 trees per species), and declined in the sequence Fagus sylvatica > Acer pseudoplatanus > Tilia cordata > Carpinus betulus > Fraxinus excelsior. Decreasing soil moisture content (Θ) resulted in linearly reduced J _s in four of the species. During a dry period, J _s was reduced by 44% in T. cordata, 39% in F. sylvatica, 37% in A. pseudoplatanus and 31% in C. betulus compared to sap flux at equal vapour pressure deficit (D) in the wet period. F. excelsior, the only ring-porous species studied, lacked a significant response in J _s to D and Θ. The relative reduction in water use during the dry period was not related to the assumed drought sensitivity of the species as inferred from their abundance in natural woodlands. J _s was positively correlated with tree diameter at breast height (DBH) in three species but decreased with DBH in two species. Dyeing experiments revealed that DBH accounted for 94% of the variation in sapwood area found in a bulk sample of all diffuse-porous trees. This suggests that DBH is a reliable estimator of sapwood area of temperate diffuse-porous species irrespective of species identity. In contrast, sap flux density was found to be greatly dependent on tree species. The estimated whole-plant water use for diffuse-porous trees of a given diameter (49 cm) ranged between 74 and 168 kg per day per species under moist soil conditions. Thus, in temperate mixed forests, species-specific differences in water use can result in a considerable spatial heterogeneity of canopy transpiration.     

温带落叶阔叶林地表鞘翅目成虫小尺度空间格局动态分析

土壤动物空间格局是格局—过程以及生物多样性维持机制研究的重要基础,目前小尺度空间土壤动物空间格局动态特征仍不清楚。基于地统计空间分析方法,以鞘翅目成虫为研究对象,研究帽儿山温带落叶阔叶林小尺度空间(5m)地表鞘翅目成虫群落及类群的空间格局动态特征。结果表明:4次调查共捕获鞘翅目成虫11科、29类、1021只个体,调查月份鞘翅目成虫群落具有较强的时空变异性;Moran'sⅠ系数表明鞘翅目成虫群落和类群具有复杂的正的空间自相关性,其空间异质性可用球状、指数、高斯和线性模型进行拟合。这种空间异质性具有一定的时间变化特征,且这种空间分异是由随机性因素单一调控或结构性因素和随机性因素共同调控的结果;类群之间在多种尺度上表现为复杂的以负相关居多的空间关联性,这种空间关联性的形成主要是结构性因素或随机性因素单一调控的结果。本实验表明地表鞘翅目成虫群落在小尺度空间具有明显的空间异质性特征,这种空间异质性时间变异性较明显。     

长白山阔叶红松林径级结构动态模拟和优化经营

以原始阔叶红松林为研究对象,采用密度依赖矩阵模型,模拟了自然生长预案下林分径级结构的动态变化,分析了一种择伐预案对林分径级结构的影响,计算了7种不同择伐强度下森林的恢复期。结果表明:原始阔叶红松林比较稳定,但也有比较缓慢的自然生长,林分株数密度呈下降趋势,符合森林的自然稀疏规律,随着时间的推移,各径阶株数的变化速度逐渐减弱,趋于稳定,验证了演替顶极理论。以生长量、收获量、保留林分结构和采伐费用为森林经营效果的评价指标,则20%的采伐强度、35年采伐周期和25%的采伐强度、45年的采伐周期的2种方案较优。     

Three-dimensional structure of a mixed broad-leaved forest in Japan

An investigation was made in a mixed broad-leaved forest in Japan to determine the three-dimensional structure of the crowns of the component species. Vertical extensions and positions of foliage above a circle with a 20-cm diamter on the forest floor were measured using a grid of 177 points in the forest. The first leaf layer above a point was defined to be the highest aggregation of the foliage above the point, and the second leaf layer to be the one below. There were generally two leaf layers above each point, the first and the second. Each of the nineteen species in the plot belonged to either the first leaf layer or the second. Because the leaf layers varied in height from place to place in the forest, the two leaf layers were not separated clearly. Mean vertical depth of each leaf layer was about 2 m irrespective of the leaf layers and component species. Since the depths were similar among the species, species mean crown volumes (volumes of the spaces occupied by foliage) per unit land area mainly depended on horizontal extensions of their crowns, or the coverage. That is, species varied in their horizontal rather than vertical crown extensions. There was an upper limit (6 m) to the sum of vertical depths of the leaf layers above a point on the forest floor. On an average, about three species occurred above a point.     

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.     

Biogeochemistry of calcium in a broad-leaved forest ecosystem

This study of the biogeochemical cycle of Ca has been carried out in an oak-hornbeam forest ecosystem on a soil derived from the alteration of Famennian (upper Devonian) shales. The physical nature of the geologic substrate made it possible to use the watershed-ecosystem approach by monitoring the loss of Ca by streamflow. Input of hydrogen ions and Ca from rain were average as well as that of mineral nitrogen of which 50% was ammonium-N. Modelling the biogeochemical cycle of Ca in this ecosystem proved to be difficult owing to the heterogeneity of the geologic substrate.The cationic denudation occured mostly with bicarbonate ion as a carrier of the base cations towards the streamflow. Seasonal effects on streamwater cationic composition were in relation to maximal soil respiration occuring in the autumn. In addition to this effect, the electrolyte concentration of streamwater was determined by rain and by changes in concentration of the soil solution either by evapotranspiration or by freezing. In spite of widely fluctuating values of flow rate, the concentration of the streamflow remained constant within a factor of two or at the most three. The mechanism underlying this buffering action of the soil on the concentration of the solution flowing through it, was shown to be related to ion-exchange processes.     

植被光合呼吸模型在长白山温带阔叶红松林的优化及验证

植被光合呼吸模型(VPRM)关键参数的确定和优化是准确计算生态系统净CO_2交换(NEE)的基础。利用中国通量观测研究联盟(China FLUX)长白山站温带阔叶红松林2005年的通量观测资料,对VPRM的4个参数(最大光能利用率ε_0、光照为半饱和条件下光合有效辐射值PAR0和呼吸参数(α、β))进行优化,并使用2006年的观测资料对参数优化前后的模拟结果进行评估。结果表明:参数优化后,VPRM能够较好地模拟长白山地区2006年植物生长季NEE的变化。对30min NEE模拟的平均误差为-1.81μmol m~(-2)s~(-1),相关系数为0.72,模拟NEE平均日变化的峰值约为观测值的91%,相关系数为0.97。但在植物非生长季模型对森林NEE的模拟效果较差。模型模拟30min NEE的平均误差为0.39μmol m~(-2)s~(-1),相关系数仅为0.10,并且模拟低估NEE平均日变化白天吸收峰值约82%,日变化模拟值与观测值的相关系数为0.50。通过分析不同天气个例,发现模型可以较好地模拟晴天条件下NEE的变化,而对阴雨天NEE的模拟误差较大。该研究有利于提高VPRM模型对温带落叶阔叶林NEE的模拟能力,对进一步改进区域陆地NEE的模拟具有重要意义。     

东灵山暖温带落叶阔叶次生林动态监测样地:物种组成与群落结构

群落结构状态是植被演替进程中的重要体现之一,同时也是下一步演替过程发生的基础.暖温带落叶阔叶次生林是中国暖温带森林植被的主要类型.为了更好地研究其演替动态、生物多样性维持机制,我们参照巴拿马Barro Colorado Island(BCI)50 ha热带雨林样地的技术规范,于2009年11月至2010年9月在北京门头沟区小龙门森林公园的暖温带落叶阔叶次生林内建立了一块20 ha的固定样地(简称DLS),对样地内胸径≥1 cm的所有木本植物进行了鉴定、调查及定位,分析了群落的组成和结构.结果表明,样地内木本植物有58种,隶属于18科33属.独立个体的总数为52,136,包括独立个体分枝的总数为103,284,全部为落叶树种.群落的区系类型以北温带成分居多,同时混有一些亚热带和热带成分,属典型的温带森林类型.群落优势种明显,个体数最多的前5个种的个体数占到总个体数的61%,前20个种占到92%,而其余38个种只占8%.群落成层现象明显,垂直结构由主林层(19个种)、次林层(18个种)和灌木层(21个种)组成.样地所有木本植物个体总径级分布呈倒"J"型,群落更新良好.主林层树种的径级结构近似于双峰或正态分布,而次林层和灌木层树种则表现出倒"J"型或"L"型.几个主要树种的空间分布表现出不同的分布格局,随着径级增大,聚集程度降低.空间分布格局显示主要优势种自身个体在其径级大小的空间分分布上互补,不同径级的个体占据了样地内不同的空间位置.     

Effects of rainfall events on soil CO2 flux in a cool temperate deciduous broad-leaved forest

The effects of rainfall events on soil CO₂ fluxes were examined in a cool temperate Quercus/Betula forest in Japan. The soil CO₂ fluxes were measured using an open-flow gas exchange system with an infrared gas analyzer in the snow-free season from August 1999 to November 2000. Soil CO₂ flux showed no significant diurnal trend on days without rain. In contrast, rainfall events caused a significant increase in soil CO₂ flux. To determine the effect of rainfall events and to evaluate more precisely the daily and annual soil carbon flux, we constructed a multiple polynomial regression model that included two variables, soil temperature and soil water content, using the soil CO₂ flux data recorded on sunny days. Daily soil carbon fluxes on sunny days calculated by the model were almost the same as those determined by the field measurements. On the contrary, the fluxes measured on rainy days were significantly higher than those calculated daily from the soil carbon fluxes by the model. Annual soil carbon fluxes in 1999 and 2000 were estimated using models that both do and do not take rainfall effects into consideration. The result indicates that post-rainfall increases in soil CO₂ flux represent approximately 16–21% of the annual soil carbon flux in this cool temperate deciduous forest.     

Shoot life span in relation to successional status in deciduous broad-leaved tree species in a temperate forest

In trees, leaf life span is closely related to successional status. Although leaves are attached to shoots, shoot life span has been insufficiently studied in the context of ecological systems. Interspecific variation in shoot survivorship was investigated over 27 months in 15 temperate hardwood tree species. Relationships between shoot architecture and shoot survival were also investigated. Shoot life span was shortest in early successional species, and longest in late successional species, in each of the families Betulaceae and Fagaceae. In Salicaceae, all of which were early successional species, shoot life span was longer in mountainous than in riparian species. Early successional or riparian species distributed longer shoots densely, even in proximal positions on mother shoots, resulting in mutual shading and consequent early and massive shoot shedding. By contrast, late successional or mountainous species concentrated shoots in distal positions, allowing shoots to receive equally favorable light, resulting in a longer life span. These results reveal close relationships between shoot life span and environmental resource availability or successional status and suggest a causal relationship between shoot shedding and shoot architecture.     

Seasonal variation in leaf properties and ecosystem carbon budget in a cool-temperate deciduous broad-leaved forest: simulation analysis at Takayama site, Japan

Seasonal changes in gross primary production (GPP) and net ecosystem production (NEP) in temperate deciduous forests are mostly driven by environmental conditions and the phenology of leaf demography. This study addresses another factor, temporal changes in leaf properties, i.e., leaf aging from emergence to senescence. A process-based model was used to link the ecosystem-scale carbon budget with leaf-level properties on the basis of field observation and scaling procedures; temporal variations in leaf thickness (leaf mass per area, LMA), photosynthetic rubisco (V_cmax) and electron-transport (J_max) capacity, and dark respiration (R_d) were empirically parameterized. The model was applied to a cool-temperate deciduous broad-leaved forest at Takayama, in central Japan, and validated with data of net ecosystem CO₂ exchange (NEE=–NEP) measured using the eddy-covariance method. NEP of the Takayama site varied seasonally from 3 g C m^–2 day^–1 net source in late winter to 5 g C m^–2 day^–1 net sink in early to mid-summer. A sensitivity experiment showed that removing the leaf-aging effect changed the seasonal CO₂ exchange pattern, and led to overestimation of annual GPP by 6% and annual NEP by 38%. We found that seasonal variation in V_cmax affected the seasonal pattern and annual budget of CO₂ exchange most strongly; LMA and R_d had moderate influences. The rapid change in V_cmax and R_d during leaf emergence and senescence was important in evaluating GPP and NEP of the temperate deciduous forest.     

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).     

Pollination systems in the cool temperate mixed coniferous and broad-leaved forest zone of Yakushima Island

Animal pollination was observed in a cool temperate mixed coniferous and broadleaved forest, and in shrubby vegetation on a mountain summit, on Yakushima Island (30.2°N, 130.3°E), to the south of Kyushu, Japan. In the mixed forest, two groups of plants were recognized: exclusively canopy-flowering species, and understory-flowering species. All of the canopy-flowering species had dish-shaped flowers or flowers without petals, were visited by opportunist insects, and most of them showed a mass-flowering pattern. Each segregated its flowering time from those of the others. On the other hand, most of the understory-flowering species had bell-or funnel-shaped flowers which were pollinated by birds or bumble bees, and showed an extended flowering pattern. Their phenological flowering series (exceptCamellia japonica that was pollinated by birds), without a break, coincided very well with the active period of a bumble bee species,Bombus ardens. In the shrubby vegetation on the mountain summit, two types of species were recognized: one type also grew in the forest, whereas the other type only grew in the shrubby vegetation. The former type of species in this vegetation was visited by a more diverse range of insects than that in the forest. In particular, species visited mainly by bumble bees in the forest attracted many opportunist insects. All but one of the species that only grew in the shrubby vegetation were visited only by opportunist insects and never by bumble bees.