Succession May Maintain High Leaf Area: Sapwood Ratios and Productivity in OldSubalpine Forests

Old forests are generally believed to exhibit low net primary productivity (NPP) and therefore to be insignificant carbon sinks. This relationship between age and NPP is based, in part, on the hypothesis that the biomass of respiratory tissues such as sapwood increases with age to a point where all photosynthate is required just to maintain existing tissue. However, this theoretical connection between respiration:assimilation ratios and forest productivity is based on age-dependent trends in the sapwood:leaf ratios of individual trees and even-aged stands; it does not take into account such processes in natural forests as disproportional increases in shade-tolerant species over time and multiple-age cohorts. Ignoring succession and structural complexity may lead to large underestimates of the productivity of old forests and inaccurate estimates of the ages at which forest productivity declines. To address this problem, we compared biomass allocation and productivity between whitebark pine, a shade-intolerant, early-successional tree species, and subalpine fir, a shade-tolerant, late-successional species, by harvesting 14 whitebark pines and nine subalpine firs that varied widely in dbh and calculating regression models for dbh vs annual productivity and biomass allocation to leaves, sapwood, and heartwood. Late-successional subalpine fir allocated almost twice as much biomass to leaves as early-successional whitebark pine. Subalpine firs also had a much lower allocation to sapwood and higher growth rates across all tree sizes. We then modeled biomass allocation and productivity for 12 natural stands in western Montana that were dominated by subalpine fir and whitebark pine varying in age from 67 to 458 years by applying the regressions to all trees in each stand. Whole-stand sapwood:leaf ratios and stand productivity increased asymptotically with age. Sapwood:leaf ratios and productivity of whitebark pine in these stands increased for approximately 200–300 years and then decreased slowly over the next 200 years. In contrast, sapwood:leaf ratios of all sizes of subalpine fir were lower than those of pine and productivity was higher. As stands shifted in dominance from pine to fir with age, subalpine fir appeared to maintain gradually increasing rates of whole-forest productivity until stands were approximately 400 years old. These results suggest that forests such as these may continue to sequester carbon for centuries. If shade-tolerant species that predominate late in succession maintain high assimilation-to-respiration ratios in other forests, we may be underestimating production in old forests, and current models may underestimate the importance of mature forests as carbon sinks for atmospheric CO₂ in the global carbon cycle. Received 16 February 1999; accepted 24 November 1999.     

Estimating tree and stand sapwood area in spatially heterogeneous southeastern Australian forests

Aims Natural and anthropogenic changes in forests can have important influences on transpiration and water production. Understanding the effects of increasing disturbances, due for example to climate change and forest harvesting, requires detailed information on how forest density and structural attributes relate to transpiration. Mean annual transpiration of eucalypt forest communities is often strongly correlated with total cross-sectional sapwood area. Our aim was to test an efficient method for estimating sapwood area at 1.3 m height (SA 1.3) in a large number of trees to understand the spatial heterogeneity of tree and stand sapwood area within and between forest communities, and develop allometric relationships that predict SA 1.3 with forest inventory data. We also apply tree competition models to determine the degree to which the relationship between SA 1.3 and tree basal area at 1.3 m height (BA 1.3) is influenced by competition.Methods We visited 25 recently harvested southeastern Australian forest sites consisting of 1379 trees and 5 Eucalyptus species to evaluate a new efficient data collection method for estimating SA 1.3 with tree taper and stump dimensions data using mixed effects models. The locations of 784 stumps within one 5-ha site were accurately mapped using an unmanned aerial vehicle (UAV), and four distance-dependent tree competition models were applied across the site to explain within-stand variation in the ratio of SA 1.3 to BA 1.3. Data from 24 additional sites, consisting of ten 15 m radial plots per site, were used to analyse within-site variation in R Ha (the ratio of stand sapwood area SA Ha to stand basal area BA Ha). The radial plots were merged within each site to evaluate between-site variations in R Ha across the landscape. For predicting SA Ha with forest inventory data, we computed the relationship between SA Ha and a new index of total stem perimeter per hectare, defined as ? B A H a N T, where N T is tree stocking density.Important findings Our 1379 measured stems represent the most comprehensive measure of sapwood area, surpassing the 757 measured stems in native eucalypt forests published in literature. The species-specific R Ha varied considerably across sites and therefore extrapolating SA Ha with spatially distributed BA Ha maps and a generalized R Ha would introduce local uncertainty. We found that the species-specific stem perimeter index was more effective at capturing variability in SA Ha across the landscape using forest composition, structure and density data (R 2 : 0.72–0.77). The strong correlation between tree SA 1.3 and BA 1.3 improved slightly using tree competition models (R 2 increased from 0.86 to 0.88). Relating SA Ha to routinely measured forest inventory attributes within permanent plots and Light Detection and Ranging (LiDAR) data may provide opportunities to map forest water use in time and space across large areas disturbed by wildfire and logging.     

Changes in forest biomass and linkage to climate and forest disturbances over Northeastern China

The forests of northeastern China store nearly half of the country's total biomass carbon stocks. In this study, we investigated the changes in forest biomass by using satellite observations and found that a significant increase in forest biomass took place between 2001 and 2010. To determine the possible reasons for this change, several statistical methods were used to analyze the correlations between forest biomass dynamics and forest disturbances (i.e. fires, insect damage, logging, and afforestation and reforestation), climatic factors, and forest development. Results showed that forest development was the most important contributor to the increasing trend of forest biomass from 2001 to 2010, and climate controls were the secondary important factor. Among the four types of forest disturbance considered in this study, forest recovery from fires, and afforestation and reforestation during the past few decades played an important role in short‐term biomass dynamics. This study provided observational evidence and valuable information for the relationships between forest biomass and climate as well as forest disturbances.     

Forest Structure and Biomass of a Tropical Seasonal Rain Forest in Xishuangbanna,Southwest China1

Xishuangbanna is a region located at the northern edge of tropical Asia. Biomass estimates of its tropical rain forest have not been published in English literature. We estimated forest biomass and its allocation patterns in five 0.185–1.0 ha plots in tropical seasonal rain forests of Xishuangbanna. Forest biomass ranged from 362.1 to 692.6 Mg/ha. Biomass of trees with diameter at 1.3 m breast height (DBH) ≥ 5 cm accounted for 98.2 percent of the rain forest biomass, followed by shrubs (0.9%), woody lianas (0.8%), and herbs (0.2%). Biomass allocation to different tree components was 68.4–70.0 percent to stems, 19.8–21.8 percent to roots, 7.4–10.6 percent to branches, and 0.7–1.3 percent to leaves. Biomass allocation to the tree sublayers was 55.3–62.2 percent to the A layer (upper layer), 30.6–37.1 percent to the B layer (middle), and 2.7–7.6 percent to the C layer (lower). Biomass of Pometia tomentosa, a dominant species, accounted for 19.7–21.1 percent of the total tree biomass. The average density of large trees (DBH ≥100 cm) was 9.4 stems/ha on two small plots and 3.5 stems/ha on two large plots, illustrating the potential to overestimate biomass on a landscape scale if only small plots are sampled. Biomass estimations are similar to typical tropical rain forests in Southeast Asia and the Neotropics.     

Forest biomass estimation at regional and global levels, with special reference to China’s forest biomass

Accurate estimation of forest biomass size and regional distribution is a prerequisite in answering a long-standing debate on the role of forest vegetation in the regional and global carbon cycle. Appropriate biomass estimation methods and available forest data sources are two key factors for this purpose. Among the estimation methods, the continuous Biomass Expansion Factor (BEF; defined as the ratio of all stand biomass to stem volume or biomass) method is considered to be the best. We applied the continuous BEF to forest inventory data of China and estimated a biomass carbon of 4.6 PgC and a biomass carbon density of 38.4 Mg ha^–1. A review of recent literature shows that forest carbon density in major temperate and boreal forest regions in the Northern Hemisphere has a narrow variance ranging from 29 Mg ha^–1 to 50 Mg ha^–1, with a global mean of 36.9 Mg ha^–1. This suggests that the forest biomass density in China is closely coincident with the global mean.     

贵州普定喀斯特次生林乔灌层地上生物量

生物量是植物群落最重要的特征之一, 也是研究生态系统基本过程和功能的重要参数。我国西南喀斯特山地次生林是退化生态系统恢复过程中的重要阶段, 其生境特殊, 乔灌层种类繁杂且生长形态多样, 物种间体积质量密度差异悬殊, 生物量测定极为困难。所以, 有关其生物量的基础资料极为缺乏。该文根据对普定喀斯特生态站的窄叶石栎(Lithocarpus confinis)+云南鼠刺(Itea yunnanensis)林、圆果化香(Platycarya longipes)+云南鼠刺林、圆果化香+槲栎(Quercus aliena)林3个主要次生森林类型的样地调查资料, 采用样木回归模型法和收获法, 研究了群落乔灌层地上生物量及其分配格局。利用172株样木数据建立了圆果化香、窄叶石栎、云南鼠刺、刺楸(Kalopanax septemlobus)、安顺润楠(Machilus cavaleriei)、槲栎、香叶树(Lindera communis)、川钓樟(Lindera pulcherrima var. hemsleyana)、异叶鼠李(Rhamnus heterophylla)、倒卵叶旌节花(Stachyurus obovatus)、薄叶鼠李(Rhamnus leptophylla)、贵州花椒(Zanthoxylum esquirolii)、竹叶椒(Zanthoxylum planispinum)、铁仔(Myrsine africana)和刺异叶花椒(Zanthoxylum dimorphophyllum var. spinifolium)15个主要树种的分种生物量回归模型。同时, 利用这些样木建立了3个不同胸径(D)组(D＜1 cm、1 cm≤D≤5 cm和D>5 cm)的生物量回归模型。通过进一步分种和分组的计算得出: 这3个次生乔木林的乔灌层地上生物量分别为85.6×10³、65.3×10³和115.2×10³ kg·hm^-2; 层次分配上, 3个样地的乔木层地上生物量占绝对优势, 分别约占乔灌层地上生物量的98.5%、96.6%和99.0%; 径级分配上, 3个样地的生物量主要集中在大径级(D≥10 cm)的个体上; 物种分配上, 3个样地的生物量排序前10位的物种分别约占乔灌层地上生物量的99.3%、97.3%和99.0%, 并集中分配在少量优势树种中。     

基于机载激光雷达的小兴安岭温带森林组分生物量反演

使用小兴安岭温带森林机载遥感-地面观测同步试验获取的机载激光雷达(light detection and ranging, Lidar)点云数据和地面实测样地数据, 估测了典型森林类型的树叶、树枝、树干、地上、树根和总生物量等组分的生物量。从激光雷达数据中提取了两组变量(树冠高度变量组和植被密度变量组)作为自变量, 并采用逐步回归方法进行自变量选择。结果表明: 激光雷达数据得到的变量与森林各组分生物量有很强的相关性; 对于针叶林、阔叶林和针阔叶混交林三种不同森林类型生物量的估测结果是: 针叶林优于阔叶林, 阔叶林优于针阔叶混交林; 不区分森林类型的各组分生物量估测与地面实测值显著相关, 模型决定系数在0.6以上; 区分森林类型进行建模可以进一步提高生物量的估测精度。     

西双版纳原始热带季节雨林生物量研究

用标准木回归分析法（ 乔木、木质藤本） 和样方收获法（ 灌木、草本） , 研究了西双版纳原始热带季节雨林生物量及其分配。雨林总生物量为６９２－５９０ ｔ／ｈｍ２ , 其分配为： 乔木层占９８－６６ ％ 、灌木层占０－７６ ％ 、木质藤本占０－５０ ％ 、草本层占０－０９ ％ , 生物量主要集中于乔木层。乔木层生物量的器官分配向树干和树根集中： 树干占６９－８０ ％ , 树根占２１－５６ ％ , 树枝占７－７７ ％ ,树叶占０－７７ ％ ; 生物量径级分配向中等径级（６０ ～７０ ｃｍ） 和最大径级（１５０ ～１６０ ｃｍ） 集中; 生物量垂直分配向上层集中; Ⅰ亚层（ 高度＞ ４０ ｍ） 占６０－５５ ％ 、Ⅱ亚层（２０ ～４０ ｍ） 占３６－７２ ％ 、Ⅲ亚层（３ ～２０ ｍ） 占２－７３ ％ ; 优势种番龙眼生物量占乔木层的２０－０７ ％ ; 乔木层叶面积指数为６－９１ 。     

基于森林调查数据的长白山天然林森林生物量相容性模型

森林生物量估算是进行陆地生态系统碳循环和碳动态分析的基础,但现有估测模型存在着总量与分量不相容的问题.本文以吉林省汪清天然林区为例,提出了基于森林调查的相容性森林生物量模型设计思想,并采用联立方程组为不同森林群落构造了一系列引入林分蓄积因子的相容性生物量模型,得到的预估精度较高.其中,针叶林、阔叶林和针阔混交林群落的森林生物量模型预估精度均在95%以上,基本上解决了森林生物量模型的相容性问题.     

Components of ecosystem evaporation in a temperate coniferous rainforest, with canopy transpiration scaled using sapwood density

Here we develop and test a method to scale sap velocity measurements from individual trees to canopy transpiration (E(c)) in a low-productivity, old-growth rainforest dominated by the conifer Dacrydium cupressinum. Further, E(c) as a component of the ecosystem water balance is quantified in relation to forest floor evaporation rates and measurements of ecosystem evaporation using eddy covariance (E(eco)) in conditions when the canopy was dry and partly wet. Thermal dissipation probes were used to measure sap velocity of individual trees, and scaled to transpiration at the canopy level by dividing trees into classes based on sapwood density and canopy position (sheltered or exposed). When compared with ecosystem eddy covariance measurements, E(c) accounted for 51% of E(eco) on dry days, and 22% of E(eco) on wet days. Low transpiration rates, and significant contributions to E(eco) from wet canopy evaporation and understorey transpiration (35%) and forest floor evaporation (25%), were attributable to the unique characteristics of the forest: in particular, high rainfall, low leaf area index, low stomatal conductance and low productivity associated with severe nutrient limitation.     

珠江三角洲地区森林生物量及其动态

利用生物量转换因子连续函数法,通过69组不同龄级的森林样地实测数据,拟合了珠江三角洲主要森林类型的生物量和蓄积量之间的回归方程,并结合3个时段森林清查资料,估算了区域森林生物量及其动态.结果表明：珠江三角洲的中幼林面积占森林总面积的80％以上,其林下植被生物量约占森林总生物量的33％,充分考虑林下植被生物量能提高区域森林生物量估算的精度.在1989—1993年、1994—1998年、1999—2003年3个研究时段,珠江三角洲森林生物量共增加了14.67×10⁶ t.其中,马尾松林、常绿阔叶林和针阔混交林的生物量约占区域总生物量的80％,是区域森林生物量的主体;而中、幼林的生物量所占比例达90％,但呈逐年下降趋势.珠江三角洲快速城市化和经济发展对区域森林生物量的积累并没有产生明显影响,区域森林面积基本保持不变,而区域森林生物量呈逐年增长趋势,年增长率为1.2％.随着珠江三角洲区域中、幼林不断发育成熟,区域森林的生物量将不断增加,其环境效应也将不断增强.     

A system-wide assessment of forest biomass production,markets, and carbon

This study examines the effects of supplying forest biomass on forest ecosystem services and goods with a dynamic systems model. This unique analysis models dynamic trade and investments in forestry, thereby capturing price changes from increased forest biomass demand on current and future flows of forest ecosystem services and natural capital stocks. Forests across the globe are interconnected through timber and forest biomass markets, which influence forest management decisions, land rents, and policy responses. Results indicate that expanding forest biomass consumption, even at relatively low levels, will have important impacts on ecosystem services, particularly the benefits of terrestrial carbon sequestration and timber outputs. Increased forest biomass production can be achieved with smaller impacts on ecosystem services through policies targeting natural forest preservation. However, policies that encourage residual biomass use for energy or discourage forest plantation expansion could potentially compromise carbon benefits.     

中国森林植被碳库的动态变化及其意义

利用1949年至1998年间7次森林资源清查资料,结合使用森林生物量实测资料,采用改良的生物量换算因子法,推算了中国50年来森林碳库和平均碳密度的变化,分析了中国森林植被的CO2源汇功能,结果表明,70年代中期以前,主要由于森林砍伐等人为作用,中国森林碳库和碳密度都是减少的,碳储量减少了0．62PgC（Pg＝10^15g),年均减少约0．024PgC。之后,呈增加趋势。在最近的20多年中,森林碳库由70年代末期的4．38PgC增加到1998年的4．75PgC,共增加0．37PgC,年平均增加0．022PgC。这种增加主要由人工造林增加所致。20多年来,由于人工林增加导致碳汇增加0．45PgC,年平均增加吸收0．021PgC／a。人工林的平均碳密度也显增加,共增加了约一倍。这除了人工成林增多外,气温上升和CO2浓度施肥也可能是促进森林生长的重要因子。     

Benchmarks and predictors of coarse woody debris in native forests of eastern Australia

Fallen coarse woody debris (CWD) is critical to forest biodiversity and function. Few studies model factors that influence CWD availability, although such investigations are critically needed to inform sustainable forest management. We assess benchmark levels of CWD in unharvested native forests and those harvested for timber, across a range of forests in north‐east New South Wales, Australia. We found timber‐harvesting was the dominant driver of CWD, with almost double the count (pieces ha^?1) and volume (m³ ha^?1) of total CWD in selectively harvested than unharvested sites. This pattern was consistent across wet and dry forest types. Harvested sites had greater counts of hollow‐bearing logs, and greater volumes of small and medium‐sized CWD (15–50 cm diameter) than unharvested sites. There was no effect of harvesting on the volume of large CWD (>51 cm diameter). Total volumes of CWD (>15 cm diameter) varied from 114 to 166 m³ ha^?1. We found few differences in CWD counts and volumes between forest types, with grassy woodlands and forests containing less CWD than other dry and shrubby forest types, reflecting lower potential input rates. The CWD levels recorded here are similar to those recorded in dry and wet sclerophyll forests elsewhere in Australia and are typical of global estimates for ‘old growth’ forests. Using general linear models we captured up to 57% of the variation in CWD across sites, and found that timber harvesting, topography and the numbers of standing hollow‐bearing and dead trees were significant predictors of CWD. Values for unharvested forest provide a benchmark that could be used to inform retention guidelines for CWD in managed forests in this region. Further assessment of the effect of repeat timber harvesting is needed to fully understand its impact on CWD dynamics, especially if forest residues resulting from timber harvesting are removed from native forests for bioenergy production.     

Relationships among the Stem, Aboveground and Total Biomass across Chinese Forests

Forest biomass plays a key role in the global carbon cycle. In the present study, a general allometric model was derived to predict the relationships among the stem biomass Ms, aboveground biomass MA and total biomass MT, based on previously developed scaling relationships for leaf, stem and root standing biomass. The model predicted complex scaling exponents for MT and/or MA with respect to Ms. Because annual biomass accumulation in the stem, root and branch far exceeded the annual increase in standing leaf biomass, we can predict that MT ∝MA ∝ Ms as a simple result of the model. Although slight variations existed in different phyletic affiliations （i.e. conifers versus angiosperms）, empirical results using Model Type Ⅱ （reduced major axis） regression supported the model＇s predictions. The predictive formulas among stem, aboveground and total biomass were obtained using Model Type I （ordinary least squares） regression to estimate forest biomass. Given the low mean percentage prediction errors for aboveground （and total biomass） based on the stem biomass, the results provided a reasonable method to estimate the biomass of forests at the individual level, which was insensitive to the variation in local environmental conditions （e.g. precipitation, temperature, etc.）.     

Forest biomass stocks and dynamics across the subtropical Andes

Forest biomass plays an important role in the global carbon cycle. Therefore, understanding the factors that control forest biomass stocks and dynamics is a key challenge in the context of global change. We analyzed data from 60 forest plots in the subtropical Andes (22–27.5° S and 300–2300 m asl) to describe patterns and identify drivers of aboveground biomass (AGB) stocks and dynamics. We found that AGB stocks remained roughly constant with elevation due to compensating changes in basal area (which increased with elevation) and plot‐mean wood specific gravity (which decreased with elevation). AGB gain and loss rates both decreased with elevation and were explained mainly by temperature and rainfall (positive effects on both AGB gains and losses). AGB gain was also correlated with forest‐use history and weakly correlated with forest structure. Mean annual temperature and rainfall showed minor effects on AGB stocks and AGB change (gains minus losses) over recent decades. Although AGB change was only weakly correlated with climate variables, increases in AGB gains and losses with increasing rainfall—together with observed increases in rainfall in the subtropical Andes—suggest that these forests may become increasingly dynamic in the future. Abstract in Spanish is available with online material     

鼎湖山黄果厚壳桂群落的生物量

用样木收获法和相关生长法测定并推算了鼎湖山黄果厚壳桂群落的生物量和净生产量.结果表明,群落的生物量是195.97(t·ha~(-1)),净生产量20.64(t·ha~(-1),·a~(-1)).     

Forest structure and biomass of mangroves in the Mgeni estuary,South Africa

Forest structure and biomass were determined in a mangrove stand dominated by Bruguiera gymnorrhiza (L.) Lam. Trees in 5 m² sample plots were harvested at ground level and then further cut into 1 m strata for separation into living wood, dead wood, leaves, reproductive material and pneumatophores. Mean above-ground living biomass was calculated at 94.49±7.83 t dry matter ha^–1, while dead wood contributed a mean mass of 7.63±0.89 t dry matter ha^–1. Excavations of roots yielded a below-ground biomass of 9.67 t dry matter ha^–1 which represented only 9.8% of the above-ground value. There was a mean density of 4700 living stems ha^–1 with plant heights ranging from 0.57 m to 5.80 m. Mean LAI was 4.95±0.80. As a basis for estimating standing biomass, regression lines were fitted to biomass values from individual trees of B. gymnorrhiza and Avicennia marina (Forssk.) Vierh. of various sizes. A comparison of these relationships with methods used by previous workers for estimating biomass suggests that most other methods cannot be applied without modification for local stands of mangroves.     

浙江省1991~2006年森林火灾释放黑碳量的估算

黑碳气溶胶不仅造成环境污染危害人的健康,同时通过吸收太阳和大气辐射产生的辐射强迫对全球和区域产生影响,成为影响全球变暖的仅次于CO2的重要成分.在全球变暖及大气中CO2浓度不断升高的趋势下,各国开始采用造林再造林来减缓全球变暖的趋势,随着森林面积和林内生物量的不断增加,以及森林火灾的频发,林火带来的黑碳排放量也不容忽视.然而在世界上大部分地区,这方面的工作却很少被人所关注.根据1991～2006年浙江省森林火灾统计资料和浙江省各种森林类型地上生物量的数据,计算出了浙江省每年的森林火灾导致的生物量损失量.同时采取释放因子法,对1991～2006浙江省每年森林火灾释放的黑碳量进行了估算.其中1995年和2000年森林火灾释放的黑碳总量分别是38.4、97.2 t,占整个浙江省黑碳释放总量中的比重分别是0.12%、0.17%,对于区域大气质量有一定的影响.     

沈阳城市森林绿量测算

借助ARC/GIS地理信息系统,以“平面量推算平面量”的方法,测算了沈阳市城市森林的绿量.结果表明:沈阳市城市森林不同类型单位面积绿量以风景游憩林最高,为3·86m2·m-2;生态公益林最低,为2·27m2·m-2;城市森林分布区单位面积绿量为2·99m2·m-2;城区单位面积绿量为0·25m2·m-2.沈阳城市森林总体绿量约为1·13×108m2·其中,附属林为4·15×107m2,占36·64%;生态公益林2·72×107m2,占23·99%;风景游憩林2·20×107m2,占19·38%;道路林1·84×107m2,占16·20%;生产经营林0·43×107m2,占3·79%.经检验,平面量推算平面量方法测算精度达到91·81%(α=0·05).