森林生态系统粗死木质残体碳储量研究进展

粗死木质残体(CWD)是森林生态系统重要的结构性与功能性单元,作为连接植被碳库与土壤碳库的关键纽带,对全球森林生态系统碳循环发挥着重要而独特的作用,越来越多的学者开始关注CWD碳储量相关研究。系统阐述了国内外CWD碳储量研究的发展历程、研究范围与基本特征等内容,总结概括了CWD体积测算、CWD碳浓度估算等碳计量相关方法的研究进展。通过梳理发现:国内外学者对CWD碳储量的研究仍处于初级阶段,主要集中于探讨不同树种、不同林龄、不同腐解等级、不同海拔、不同存在形式的森林CWD碳储量分布特征,而较少关注CWD碳库对土壤碳库和植被碳库的作用机制以及对未来气候变化的响应和反馈机制;CWD碳计量方法较为单一,普遍采用的是传统的\"生物量-碳浓度法\",而运用机器学习算法对CWD碳储量进行估算的研究尚不多见。此外,相对国外研究而言,国内研究主要局限于小尺度范围。文章据此提出未来CWD碳储量研究的发展方向:(1)拓展研究尺度,建立CWD碳储量长期观测体系;(2)深入开展不同森林生态系统CWD碳储量对气候变化的响应机制研究;(3)探索更加多元化的CWD碳储量计量方法;(4)深入探讨CWD碳库对土壤碳库与植被碳...     

鼎湖山黄果厚壳桂粗死木质残体的分解

通过建立时间序列法、测定粗死木质残体的密度变化,研究鼎湖山黄果厚壳桂粗死木质残体三个直径(5~10cm、10~20cm和20~30cm)的分解过程,探讨粗死木分解过程中C、N元素及其C/N比值与分解速率的关系。结果表明粗死木质残体的分解常数K值随直径的增加从0.2225a-1呈指数降低到0.1257a-1,由粗死木质残体在分解过程中密度变化得出三个直径从小到大分解95%所需的时间分别为13a、19a和24a;黄果厚壳桂粗死木质残体在分解过程中不存在时滞效应,其树皮和心材的分解速度也相近,用单因素数指数方程能准确反映黄果厚壳桂的分解过程;与高纬度地区比较,鼎湖山黄果厚壳桂粗死木残体的分解速率常数K值显著高于前者。还就粗死木残体基质组成和性质、气候与环境条件、生物等因素对粗死木分解速率的影响进行了详细的讨论。     

Acetylene reduction in conifer logs during early stages of decomposition

Acetylene reduction was examined periodically for as long as 68 months in the outer and inner bark, sapwood, and heartwood of decaying logs of western hemlock [Tsuga heterophylla (Raf.) Sarg.] western redcedar (Thuja plicata D. Don), Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco], and Pacific silver fir (Abies amabilis Dougl. ex Forbes) in the western Oregon Cascade Mountains. Tissues from freshly cut logs from sound trees were unable to reduce acetylene. However, after 18 months of decomposition, acetylene reduction was found in all log tissues except heartwood. Over the 68-month study period, no significant relationship between reduction rate and tissue moisture was found. Acetylene reduction rates differed significantly among tissues, log species, and time of exposure to decomposers. Although acetylene reduction generally showed a steady increase with time, tissues of some species showed a more complex, nonlinear pattern of change. Although the amount of nitrogen fixed is low compared to the total present in decaying logs, it might be an important source of readily available nitrogen for the microbiota responsible for decomposition.Paper 2587, Forest Research Laboratory, Oregon State University, Corvallis.Paper 2587, Forest Research Laboratory, Oregon State University, Corvallis.     

森林粗死木质残体的概念及其分类

森林粗死木残体 (Coarse woody debris,CWD)在不同的文献中有不同的定义 ,没有通用而确切的概念用来描述 CWD,对研究结果的比较造成了很大障碍。 2 0世纪 90年代以来 ,随着景观生态学的发展 ,以及对 CWD生态功能的深入研究 ,国外的森林管理和研究机构 (例如 USDA Forest Service和 L TER)为了把 CWD放在区域以及景观尺度上进行比较 ,对 CWD的概念等进行了统一 ,将其直径标准由原来的≥ 2 .5 cm调整到≥ 10 cm,但是我国在此方面还没有与国际接轨 ,仍采用旧标准 (≥ 2 .5 cm ) ,这样的研究结果难于和国外进行比较 ,不利于我国 CWD的长期深入研究。另外 ,有关 CWD的分类一直以来也没有形成一个完整的分类系统 ,我国也缺少 CWD分类方法的介绍。鉴于以上情况 ,综合国内外近年来在 CWD方面的研究动态 ,综述了 CWD的概念和分类情况 ,并初步提出较综合的 CWD概念及其分类系统 ,以供相关研究者讨论和参考 ,为我国的 CWD研究起到推动作用     

Concept and Classification of Coarse Woody Debris in Forest Ecosystems

Coarse woody debris (CWD) is generally considered as dead woody materials in various stages of decomposition, including sound and rotting logs, snags, and large branches. CWD is an important functional and structural component of forested ecosystems and plays an important role in nutrient cycling, long-term carbon storage, tree regeneration, and maintenance of heterogeneous environmental and biological diversity. However, the definition and classification of CWD have been the subject of a long debate in forest ecology. CWD has not been precisely defined. Recently, with the rapid development of landscape ecology in CWD, the USDA Forest Service and the Long Term Ecological Research (LTER) have provided a standardized definition and classification for CWD, which makes data comparison in landscape scale possible. Important characteristics of their definition include: (1) a minimum diameter (or an equivalent cross-section) of CWD ≥10 cm at the widest point (the woody debris with a diameter from 1 to 10 cm should be defined as fine woody debris, and the rest is litterfall); and (2) sound and rotting logs, snags, stumps, and large branches (located above the soil), and coarse root debris (larger than 1 cm in diameter). This classification has greatly facilitated CWD studies. Therefore, it has been widely applied in some countries (particularly in North America). However, this classification has long been a source of confusion for forest ecologists in China. Furthermore, different definitions and criteria are still adopted in individual studies, which makes the interpretation and generalization of their work difficult. This article reviewed recent progress in classifying CWD, with an emphasis on introducing the classification system of the USDA Forest Service and the LTER. It is expected that this review will help facilitate the development of standardized definition and classification suitable to forest ecosystems in China. Translated from Acta Ecologica Sinica, 2005, 25(1) (in Chinese)     

Concept and Classification of Coarse Woody Debris in Forest Ecosystems

Coarse woody debris (CWD) is generally considered as dead woody materials in various stages of decomposition,including sound and rotting logs,snags,and large branches.CWD is an important functional and structural component of forested ecosystems and plays an important role in nutrient cycling,long-term carbon storage,tree regeneration,and maintenance of heterogeneous environmental and biological diversity.However,the definition and classification of CWD have been the subject of a long debate in forest ecology.CWD has not been precisely defined.Recently,with the rapid development of landscape ecology in CWD,the USDA Forest Service and the Long Term Ecological Research (LTER)have provided a standardized definition and classification for CWD,which makes data comparison in landscape scale possible.Important characteristics of their definition include:(1) a minimum diameter (or an equivalent crosssection) of CWD≥10 cm at the widest point (the woody debris with a diameter from 1 to 10 cm should be defined as fine woody debris,and the rest is litterfall);and (2) sound and rotting logs,snags,stumps,and large branches (located above the soil),and coarse root debris (larger than 1 cm in diameter).This classification has greatly facilitated CWD studies.Therefore,it has been widely applied in some countries (particularly in North America).However,this classification has long been a source of confusion for forest ecologists in China.Furthermore,different definitions and criteria are still adopted in individual studies,which makes the interpretation and generalization of their work difficult.This article reviewed recent progress in classifying CWD,with an emphasis on introducing the classification system of the USDA Forest Service and the LTER.It is expected that this review will help facilitate the development of standardized definition and classification suitable to forest ecosystems in China.     

Relationships between carbon turnover and bioavailable energy fluxes in two temperate forest soils

Recent research has shown that the broad empirical relationships used in many ecosystem models to predict carbon turnover and stabilization in soils can fail to capture differences across vegetation types or climates. Theoretically, because energy flow is fundamental to the function of decomposer organisms and ecosystems, energetics could provide complimentary fundamental constraints on soil C dynamics. Often, however, C is considered as a surrogate for energy in studies of detrital decay and C turnover in soil. Bomb calorimetry has long been used to measure stored energy in organic matter, but in detritus not all of the energy is bioavailable. Here I outline an approach to quantify the flux of bioavailable energy dissipated by resident heterotrophic communities in soil organic horizons in situ. I used the principle of energy balance together with a biogeochemical process model parameterized through calorimetric analysis of field samples. I also tested relationships between C and energy across samples of forest detritus (foliar and fine root litter, well‐decayed Oea material, and woody debris), across decay stages, and between a deciduous and coniferous forest at the Harvard Forest, MA, USA. As a first approximation, energy and C concentrations were closely related (within ca. 10%), as were ratios of heterotrophic energy dissipation to C mineralization across types of detritus (within 16%). Differences in energy content and energy : C ratios were measurable in forest detritus (particularly woody vs nonwoody), but did not vary reliably enough between forest types or through detrital stages to indicate that soil C models could be improved by including energetics. Model results indicated that there are strong similarities in energy flows and storage in the O horizons of the contrasting forest types studied at this location. Future research could focus on broader patterns across climates or biomes.     

东北主要树种倒木分解释放的CO2通量

在倒木丰富的森林生态系统中,倒木分解释放的CO2通量(RCWD)是生态系统碳收支中不容忽视的一个组分。采用红外气体分析法(Li-6400IRGA)测定东北东部山区典型温带天然次生林中11个主要树种的RCWD及其相关环境因子。主要研究目标包括:比较11个树种的RCWD、倒木温度(TCWD)和倒木含水量(WCWD);量化RCWD与TCWD和WCWD的关系;量化RCWD的季节动态。研究结果表明:白桦、山杨、紫椴、胡桃楸、蒙古栎、色木槭、春榆、红松、黄菠萝、落叶松和水曲柳在测定期间RCWD的平均值分别为:10.64、8.38、7.85、6.59、6.01、4.07、3.88、2.55、2.29、1.96μmolCO.2m-.2s-1和1.90μmolCO.2m-.2s-1。软阔叶树种的RCWD最高;针叶树种的RCWD总体上低于阔叶树种的。在整个测定期间,不同树种的TCWD虽然没有显著性差异(p>0.1),但是其WCWD差异极显著(p<0.001)。树种、倒木个体、倒木所处的立地状况及其交互作用均显著地影响RCWD,但其影响程度因树种而异。所有树种的TCWD、WCWD及其交互作用显著地(p<0.01)影响RCWD;包括了这些作用的RCWD模型解释了39.9%~72.9%的RCWD变异。不同树种RCWD的季节变化呈现基本一致的单峰曲线格局,主要受TCWD的驱动;而WCWD主要影响RCWD的季节内变化和树种间的差异。     

Decomposition and Fragmentation of Coarse Woody Debris: Re-visiting a Boreal Black Spruce Chronosequence

We re-visited a seven-stand boreal chronosequence west of Thompson, Manitoba, Canada, in which coarse woody debris (CWD) and its instantaneous decomposition were measured in 2000. New CWD measurements were performed in 2007, and tree inventories updated to provide mortality and snag failure data. These data were used to model CWD changes, compare methods of estimating decomposition, and infer possible fragmentation rates. Measured CWD was between 9.7 (in both the 77- and 43-year-old stands) and 80.4 (in the 18-year-old stand) Mg ha⁻¹ in 2007. Spatial variability was high; at most stands CWD levels had not changed significantly from 2000 to 2007. Tree mortality was a significant flux only in older stands, whereas snag fall rate varied by an order of magnitude, from 2.9% y⁻¹ (0.2 Mg ha⁻¹ y⁻¹) in the 9-year-old stand to 9.8% y⁻¹ (2.3 Mg ha⁻¹ y⁻¹) in the 12-year-old stand. A one-pool model based on these inputs underestimated actual 2000–2007 CWD decomposition in the younger stands, suggesting that fragmentation could be an important part of the carbon flux exiting the CWD pool. We compared three independent measures of annual decomposition (k): direct measurements of CWD respiration, rates based on the 7-year re-sampling effort described here, and rates inferred from the chronosequence design itself. Mean k values arrived at via these techniques were 0.06 ± 0.03, 0.05 ± 0.04, and 0.05 ± 0.05 y⁻¹, respectively. The four-pool model suggested that the transition rate between decay classes was 0.14–0.19 y⁻¹; the model was most sensitive to initial CWD values. Although the computed k values implied a problem with chronosequence site selection for at least one site, the overall CWD trend was consistent with a larger number of sites surveyed in the region.     

中亚热带常绿阔叶林粗木质残体呼吸季节动态及影响因素

粗木质残体呼吸(RCWD)释放的CO2是生态系统碳收支中的一个重要组成部分。采用红外气体分析法(Li-Cor8100土壤碳通量系统连接自制腔室)对中亚热带常绿阔叶林不同分解等级粗木质残体呼吸进行测量,探讨分解等级、温度(TCWD)和含水量(WCWD)对RCWD的影响机制。结果表明:不同分解等级粗木质残体呼吸季节变化曲线均呈明显的单峰型,最大值(9.69μmolCO2·m-2·s-1)出现在8月,最小值(0.60μmol CO2·m-2·s-1)出现在2月;不同分解等级粗木质残体呼吸存在着明显差异,Ⅲ级和Ⅳ级粗木质残体呼吸显著高于Ⅰ级(P<0.05);粗木质残体呼吸与TCWD呈显著的正相关关系(P<0.01),TCWD可以解释RCWD变化的70.2%—85.6%;RCWD与WCWD相关性不显著(P>0.05);不同分解等级粗木质残体呼吸的Q10值变化范围为2.46—2.83,平均值为2.64,Q10值随分解等级升高而增大。     

Water balance of conifer logs in early stages of decomposition

Seasonal and long-term changes in the water balance of conifer logs during the first 8 years of decomposition were studied in an old-growth Pseudotsuga/Tsuga forest in the Oregon Cascade Mountains. Measurements were made of the moisture content of outer bark, inner bark, sapwood, and heartwood and of the flow of water into and out of logs of four species (Abies amabilis, Pseudotsuga menziesii, Thuja plicata, and Tsuga heterophylla). After the logs had decomposed from 1 to 2 years, 38–47% of the canopy throughfall landing upon them ran off the surface, 29–34% leached from the bottom, and 21–30% was absorbed and evaporated. After 8 years of decomposition, water entering and then leaching from logs increased 1.3 times while runoff decreased a similar amount. The proportion of water stored by and evaporated from logs in this study indicates that in old growth forests they may intercept 2–5% of the canopy throughfall to the forest floor and that, even in early stages of decomposition, they may affect the hydrological cycle of Pacific Northwest old-growth forests.This is paper 2945 of the Forest Research Laboratory, Oregon State University, Corvallis     

Current Loads of Coarse Woody Debris on Southeastern Australian Floodplains: Evaluation of Change and Implications for Restoration

We evaluated the status of coarse woody debris (CWD, fallen wood) on floodplains of the southern Murray‐Darling basin of southeastern Australia. The floodplains are dominated floristically by the river red gum Eucalyptus camaldulensis. Aerial survey techniques were used to estimate the amounts of woody debris within 200 m of the channels along 2,442 km of 11 rivers of the system, including the Murray and Darling Rivers and the Darling Anabranch. Aerially based indices were converted into wood volumes by using ground‐truthing at a selection of sites; there was a strong correlation between index values and measured wood volume densities. For thickly forested sites such as Barmah, Gunbower Island, and the Ovens floodplains, the aerial method was not useful, so ground measurements at randomly positioned sites within the forests were used. Volumes were translated into mass by using conversion factors drawn from the literature. We estimated that total tonnage on approximately 221,000 ha of floodplain forests was 4.175 ± 0.579 × 10⁶ tonne. In the larger forested blocks (>7,000 ha), mean wood densities ranged between approximately 12 tonne/ha on the lower Goulburn up to approximately 24 tonne/ha at Barmah State Forest. The area‐weighted mean for the entire area was approximately 19 tonne/ha. A main purpose of the research was to place these figures into an historical perspective to evaluate implications for restoration. A thorough search of historical documentation revealed that there are no extant data upon which to estimate pre‐European settlement levels. We used information from an apparently undisturbed “unmanaged” site in the Millewa forests of southern New South Wales as a basis. Wood density there corresponded to a mean figure of 125 tonne/ha wood‐mass density. By using this figure we estimate that CWD levels on the southern Murray‐Darling basin may be of the order of 15% of pre‐European settlement levels. Full restoration of the 221,000 ha surveyed would require 23.5 ± 0.579 × 10⁶ tonne, which is equivalent to about 600,000 mature (1 m diameter at breast height) river red gum trees or the amount of timber derived from clear felling about 115,000 ha of river red gum forest at current stocking levels. We discuss the implications of this massive deficit and possible short‐ and long‐term solutions.     

高山森林粗木质残体附生苔藓植物的重金属吸存特征

粗木质残体及其附生苔藓植物是高山森林生态系统中两个相互联系的基本组成部分,二者的相互作用可能影响森林生态系统重金属循环,但有关不同类型和不同腐解等级粗木质残体对附生苔藓植物重金属吸存特征尚不清楚。因此,于2015年7月在川西高山森林调查了岷江冷杉(Abies faxoniana)原始林内不同类型和不同腐解等级的粗木质残体附生苔藓镉、铅、铜和锌浓度与吸存特征。结果表明:高山森林粗木质残体附生苔藓植物Cd、Pb、Cu和Zn的总吸存量依次为4700mg/hm~2、21236mg/hm~2、6179mg/hm~2和2622mg/hm~2。粗木质残体附生苔藓的4种重金属吸存量都表现为倒木大枯枝枯立木根桩;倒木附生苔藓Cd、Pb、Cu和Zn吸存量分别占高山森林粗木质残体总吸存量的54.53%、66.08%、51.13%和66.30%,根桩附生苔藓重金属吸存量不足总吸存量的3%。粗木质残体的类型和腐解等级都会影响附生植物重金属吸存特征。随着腐解等级的增加,倒木和大枯枝附生苔藓中Cd和Pb呈现\"积累-释放\"的变化特征,其浓度和吸存量在第Ⅱ、Ⅲ腐解等级较高。附生苔藓Cu和Zn浓度和吸存量在不同腐解等级粗木质残体间的差异均不明显。这些结果表明,粗木质残体附生苔藓对重金属元素具有明显的吸存作用,为认识高山森林生态系统重金属元素循环及其迁移过程提供了新的思路,也为进一步了解粗木质残体在高山森林生态系统中的重要作用提供了新的角度。     

Short-term decompositional state does not influence use of wood by macroinvertebrates in subtropical, coastal plain streams

Woody debris is an important habitat component, particularly in streams that lack other hard substrates. Research suggests a general relationship between increasing invertebrate density, diversity, and taxa richness with increasing wood decay in lotic systems, with some authors observing invertebrate taxonomic succession as decay proceeds. We designed a field experiment using colonization of known-aged woody debris in two streams to examine patterns in invertebrate colonization, density, diversity, richness, and succession. After aging woody debris 0–6 weeks in laboratory tanks and then placing the debris in the two subtropical, coastal plain streams for five additional weeks, we did not detect any statistical relationship between invertebrate density, diversity, evenness, richness, or life-history pattern with increasing woody debris decay, nor did we detect any relationships between the colonization or abundance of individual taxa and the decompositional state of the wood. In this paper, we propose two non-exclusive explanations for these trends based on opportunistic colonization and evolutionary filtering. Despite the apparent unimportance of decompositional state, woody debris still supported many taxa and remains an important habitat component. Our research further supports the importance of flooding and maintenance of intact riparian and floodplain forests to the woody debris dynamics and macroinvertebrates in coastal plain lotic systems.