地形对小兴安岭阔叶红松(Pinus koraiensis)林粗木质残体分布的影响

以小兴安岭典型阔叶红松林大面积固定样地(9hm2)为对象,分析不同地形下粗木质残体(CWD)的数量、存在形式、腐烂程度,探讨其与主要地形因子的相互关系.随着坡位的上升,CWD的密度和胸高断面积逐渐增加;从阴坡、半阴、半阳到阳坡,CWD的密度、胸高断面积和体积都呈现递减的趋势;CWD的数量分布随着样方坡度的增加没有呈现出明显的趋势.对不同地形下CWD的分布进行x2检验,各地形因子对CWD密度分布均差异显著,山脊、阴坡和坡度16～25°分别是CWD易发生的地形.在不同的地形中,CWD主要以干基折断、干中折断和枯立木3种形式存在.从谷地到山脊以拔根倒存在的CWD密度、胸高断面积和体积呈逐渐减少的趋势,而干中折断、根桩、枯立木和干基折断的密度大体上表现出增加的趋势;拔根倒的密度随坡度的上升逐渐减少,而枯立木则增加;由阳坡到阴坡枯立木的密度逐渐减少.不同存在形式的倒木,其树倒方向与坡向和风向均没有显著的相关性(p>0.05).不同地形下的各腐烂等级的CWD大致呈正态分布,且主要分布在2、3腐烂等级上,说明阔叶红松林不同地形的CWD输入和输出相对稳定.     

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.     

小兴安岭谷地云冷杉林粗木质残体碳密度特征

以小兴安岭谷地云冷杉林9.12 hm~2固定样地为研究对象,分析粗木质残体(CWD)碳密度的基础特征,揭示其与林分因子和物种多样性的关系。结果表明:(1)谷地云冷杉林CWD碳密度为13.25 t C/hm~2,其中云杉(Picea spp.)、冷杉(Abies nephrolepis)、兴安落叶松(Larix gmelinii)和未知种的CWD碳密度分别为3.59、2.61、3.06和2.85 t C/hm~2。(2)不同腐烂等级下CWD碳密度呈近正态分布,多集中在Ⅱ和Ⅲ等级,分别占总量的42.7%和35.4%。不同径级的CWD碳密度也呈近正态分布,主要分布在30—40 cm和40—50 cm径级上。干中折断、拔根倒、枯立木和干基折断为谷地云冷杉林CWD碳密度的主要存在方式。腐烂等级为Ⅰ和Ⅴ的CWD中,拔根倒的碳密度最高,其他腐烂等级中均为干中折断的碳密度最高。(3)CWD碳密度表现出较强的空间异质性,其随着林分平均胸径、最大胸径和胸高断面积的增加而下降,呈显著负相关关系(P0.05);而与林分密度、多样性指数和均匀度指数均无显著相关性(P0.05)。     

溪流粗木质残体的生态学研究进展

粗木质残体（CWD）是森林或溪流生态系统中残存的超过一定直径大小的站杆、倒木、枝桠及根系等死木质物的总称,溪流CWD对于溪流生态系统的稳定,水生生物多样性,河槽形态及其变化过程有着重要的作用。对溪流CWD的产生和分类,溪流CWD对于溪流生态系统的稳定,水生生物多样性,河槽形态及其变化过程有着重要的作用。对溪流CWD的产生和分类,溪流CWD贮量,分布和动态,以及溪流CWD的功能和管理分别进行了总结,并指出应尽快在国内开展溪流CWD的研究和管理。     

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.     

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.     

小兴安岭阔叶红松林粗木质残体空间分布的点格局分析

采用点格局分析方法对小兴安岭典型阔叶红松林9hm2(300m×300m)固定样地内粗木质残体(CWD)的空间分布格局进行了研究。结果表明:固定样地内CWD的总密度为368.8株/hm2,径级结构呈现明显的正态分布,各径级密度差别较大。花楷槭(27.8株/hm2)和枫桦(26.1株/hm2)是阔叶CWD的主要组成树种。红松(41.6株/hm2)是针叶CWD的主要组成树种,针叶树种CWD的数量随着径级的增大而增加,呈典型的J型分布。在150m的空间尺度内,CWD总体在较小尺度上表现为集群分布,在40m尺度上聚集强度最大(0.40),随着尺度的增加,CWD趋于均匀分布。不同径级组在不同规模尺度聚集,随着径级的增加,聚集强度呈下降趋势。不同物种CWD在各径级下的空间分布格局有所不同,具体表现为集群分布和由集群分布向随机分布发展两种形式。不同存在形式的CWD在研究尺度内随着尺度的增加,由集群分布向随机分布发展。随着CWD腐烂等级的增高,其聚集强度和聚集尺度均增大。CWD的空间分布格局是阔叶红松林群落与其自然环境长期作用的结果,从某种程度上也反映了该林型天然更新的格局和机制。     

Coarse woody debris in undisturbed and logged forests in the eastern Brazilian Amazon

Coarse woody debris (CWD) is an important component of the carbon cycle in tropical forests. We measured the volume and density of fallen CWD at two sites, Cauaxi and Tapajós in the Eastern Amazon. At both sites we studied undisturbed forests (UFs) and logged forests 1 year after harvest. Conventional logging (CL) and reduced impact logging (RIL) were used for management on areas where the geometric volumes of logs harvested was about 25–30 m³ ha^?1. Density for five classes of fallen CWD for large material (>10 cm diameter) ranged from 0.71 to 0.28 Mg m^?3 depending upon the degree of decomposition. Density of wood within large fallen logs varied with position relative to the ground and with distance from the center of the log. Densities for materials with diameters from 2 to 5 and 5 to 10 cm were 0.36 and 0.45 Mg m^?3, respectively. The average mass (±SE) of fallen CWD at Cauaxi was 55.2 (4.7), 74.7 (0.6), and 107.8 (10.5) Mg ha^?1 for duplicate UF, RIL, and CL sites, respectively. At Tapajós, the average mass of fallen CWD was 50.7 (1.1) Mg ha^?1 for UF and 76.2 (10.2) Mg ha^?1 for RIL for duplicate sites compared with 282 Mg ha^?1 for live aboveground biomass. Small‐ and medium‐sized material (<10 cm dia.) accounted for 8–18% of the total fallen CWD mass. The large amount of fallen CWD at these UF sites relative to standing aboveground biomass suggests either that the forests have recently been subjected to a pulse of high mortality or that they normally suffer a high mortality rate in the range of 0.03 per year. Accounting for background CWD in UF, CL management produced 2.7 times as much CWD as RIL management. Excess CWD at logging sites would generate a substantial CO₂ emission given the high rates of decay in moist tropical forests.     

喀斯特常绿落叶阔叶混交林粗木质残体的空间分布格局

粗木质残体(coarse woody debris,CWD)是森林生态系统中重要的结构性和功能性组成要素,是维护系统完整性和稳定性的关键。对CWD空间格局的研究将有助于深入探索种群格局的形成和森林生态系统的维持机制。采用g(r)函数对茂兰喀斯特常绿落叶阔叶混交林1.28 hm~2固定样地内不同径级、不同腐烂等级、不同存在形式的CWD的空间分布格局及空间关联性进行了研究。结果表明:1)在40 m的空间尺度内,CWD总体在0—12 m尺度上表现为集群分布,随着尺度的增加格局强度降低,趋于随机分布,剔除生境异质性后,格局尺度降低至7 m。2)CWD径级格局表现为:小径级中径级大径级。拔根倒和干中折断在整个研究尺度上为随机分布,其他不同径级、不同腐烂等级、不同存在形式的CWD均在小规模尺度(2—8 m)表现为集群分布,随着尺度的增加聚集强度急剧变小,趋于随机分布或均匀分布。3)除了干中折断与树段之间、大径级与小径级之间的CWD在空间上相互独立,其他不同径级、不同腐烂等级或不同存在形式的CWD之间均在小规模尺度(2—8 m)上表现为显著的正相关,随着尺度的增加空间关联性降低。喀斯特常绿落叶阔叶混交林CWD的分布格局可能是在小尺度内由密度制约、在大尺度内由生境过滤和个体自然衰老等生态学过程所形成,大径级对临近的小径级、先死亡对后死亡、站杆对倒下的个体具有一定的正向影响作用,在一定程度上揭示了该林型天然更新的作用和机制。