Linking wood-decay fungal communities to decay rates: Using a long-term experimental manipulation of deadwood and canopy gaps

Decomposition transfers carbon (C) from detrital organic matter to soil and atmospheric pools. In forested ecosystems, deadwood accounts for a large proportion of the detrital C pool and is primarily decomposed by wood-inhabiting fungi (WIF). Deadwood reductions linked to forest harvesting may alter WIF richness and composition, thus indirectly influencing the persistence of deadwood and its contribution to C and nutrient cycling. Forest structure was enhanced via canopy gap creation and coarse woody debris (CWD) addition that mimic natural disturbance by windfall within a deciduous northern hardwood forest (Wisconsin, USA) to examine its effect on deadwood-associated biodiversity and function. Experimental sugar maple (Acer saccharum) logs were sampled, for DNA extraction, ten years after placement to determine the assembly of fungal community composition and its relationship to wood decay rates.Our findings suggest that the WIF community responded to gap disturbance by favoring species able to persist under more extreme microclimates caused by gaps. CWD addition under closed canopy tended to favor a different species assemblage from gap creation treatments and the control, where canopy was undisturbed and CWD was not added. This was presumably due to consistent microclimatic conditions and the abundance of CWD substrates for host specialists. Fungal OTU richness was significantly and inversely related to CWD decay rates, likely due to competition for resources. In contrast, fungal OTU composition was not significantly related to CWD decay rates, canopy openness or CWD addition amounts. Our study site represents a diverse fungal community in which complex interactions among wood-inhabiting organisms and abiotic factors are likely to slow CWD decomposition, which suggests that maintaining a biodiverse and microsite-rich ecosystem may enhance the capacity for C storage within temperate forests.     

Low stocks of coarse woody debris in a southwest Amazonian forest

The stocks and dynamics of coarse woody debris (CWD) are significant components of the carbon cycle within tropical forests. However, to date, there have been no reports of CWD stocks and fluxes from the approximately 1.3 million km² of lowland western Amazonian forests. Here, we present estimates of CWD stocks and annual CWD inputs from forests in southern Peru. Total stocks were low compared to other tropical forest sites, whether estimated by line-intercept sampling (24.4 ± 5.3 Mg ha⁻¹) or by complete inventories within 11 permanent plots (17.7 ± 2.4 Mg ha⁻¹). However, annual inputs, estimated from long-term data on tree mortality rates in the same plots, were similar to other studies (3.8 ± 0.2 or 2.9 ± 0.2 Mg ha⁻¹ year⁻¹, depending on the equation used to estimate biomass). Assuming the CWD pool is at steady state, the turnover time of coarse woody debris is low (4.7 ± 2.6 or 6.1 ± 2.6 years). These results indicate that these sites have not experienced a recent, large-scale disturbance event and emphasise the distinctive, rapid nature of carbon cycling in these western Amazonian forests.     

Woody debris stocks in different secondary and primary forests in the subtropical Ailao Mountains, southwest China

Woody debris (WD), including coarse woody debris (CWD) and fine woody debris (FWD), is an essential structural and functional component of many ecosystems, particularly in montane forests. CWD is considered to be the major part in forest WD and it is primarily composed of logs, snags, stumps and large branches, while FWD mainly consists of small twigs. Attributes of dead woody material may change in accordance with trends in stand dynamics. The primary forest (primary montane moist evergreen broad-leaved forest) in Ailao Mountain National Nature Reserve (NNR) preserves the largest tract of natural vegetation in China. The Alnus nepalensis (D. Don) association, Populus bonatii (Levl.) association and secondary Lithocarpus association represent the secondary and chronological types following human disturbance by fires and logging under different intensity. The mass and composition of coarse woody debris (CWD, ≥10 cm in diameter) and fine woody debris (FWD, 2.5–10 cm in diameter) were inventoried in a primary forest and its three secondary counterparts. Estimates of total mass of woody debris across secondary types to primary forest ranged from 2.4 to 74.9 Mg ha⁻¹. The lowest value was found in the A. nepalensis association and the highest values were in the primary forest of which logs are the considerable differences. The ratios of CWD to FWD were low in the secondary types (about 1–4) but high in the primary forest (above 15). Our results suggested that for the recovery of woody debris in the secondary forest, it might last longer than the age of the oldest successional stage studied. Yang Lipan and Ma Wenzhang contributed equally to this work.     

The Imprint of Land-use History: Patterns of Carbon and Nitrogen in Downed Woody Debris at the Harvard Forest

Few data sets have characterized carbon (C) and nitrogen (N) pools in woody debris at sites where other aspects of C and N cycling are studied and histories of land use and disturbance are well documented. We quantified pools of mass, C, and N in fine and coarse woody debris (CWD) in two contrasting stands: a 73-year-old red pine plantation on abandoned agricultural land and a naturally regenerated deciduous forest that has experienced several disturbances in the past 150 years. Masses of downed woody debris amounted to 40.0 Mg ha⁻¹ in the coniferous stand and 26.9 Mg ha⁻¹ in the deciduous forest (20.4 and 13.8 Mg C ha⁻¹, respectively). Concentrations of N were higher and C:N ratios were lower in the deciduous forest compared to the coniferous. Pools of N amounted to 146 kg N ha⁻¹ in the coniferous stand and 155 kg N ha⁻¹ in the deciduous forest; both are larger than previously published pools of N in woody debris of temperate forests. Woody detritus buried in O horizons was minimal in these forests, contrary to previous findings in forests of New England. Differences in the patterns of mass, C, and N in size and decay classes of woody debris were related to stand histories. In the naturally regenerated deciduous forest, detritus was distributed across all size categories, and most CWD mass and N was present in the most advanced decay stages. In the coniferous plantation, nearly all of the CWD mass was present in the smallest size class (less than 25 cm diameter), and a recognizable cohort of decayed stems was evident from the stem-exclusion phase of this even-aged stand. These results indicate that heterogeneities in site histories should be explicitly included when biogeochemical process models are used to scale C and N stocks in woody debris to landscapes and regions. Received 27 April 2001; accepted 4 January 2002.     

Woody debris contribution to the carbon budget of selectively logged and maturing mid-latitude forests

Woody debris (WD) is an important component of forest C budgets, both as a C reservoir and source of CO₂ to the atmosphere. We used an infrared gas analyzer and closed dynamic chamber to measure CO₂ efflux from downed coarse WD (CWD; diameter≥7.5 cm) and fine WD (FWD; 7.5 cm>diameter≥2 cm) to assess respiration in a selectively logged forest and a maturing forest (control site) in the northeastern USA. We developed two linear regression models to predict WD respiration: one based on WD temperature, moisture, and size (R ²=0.57), and the other on decay class and air temperature (R ²=0.32). WD respiration (0.28±0.09 Mg C ha⁻¹ year⁻¹) contributed only ≈2% of total ecosystem respiration (12.3±0.7 Mg C ha⁻¹ year⁻¹, 1999–2003), but net C flux from CWD accounted for up to 30% of net ecosystem exchange in the maturing forest. C flux from CWD on the logged site increased modestly, from 0.61±0.29 Mg C ha⁻¹ year⁻¹ prior to logging to 0.77±0.23 Mg C ha⁻¹ year⁻¹ after logging, reflecting increased CWD stocks. FWD biomass and associated respiration flux were ≈7 times and ≈5 times greater, respectively, in the logged site than the control site. The net C flux associated with CWD, including inputs and respiratory outputs, was 0.35±0.19 Mg C ha⁻¹ year⁻¹ (net C sink) in the control site and −0.30±0.30 Mg C ha⁻¹ year⁻¹ (net C source) in the logged site. We infer that accumulation of WD may represent a small net C sink in maturing northern hardwood forests. Disturbance, such as selective logging, can enlarge the WD pool, increasing the net C flux from the WD pool to the atmosphere and potentially causing it to become a net C source.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

广州三种森林粗死木质残体（CWD）的储量与分解特征

粗死木质残体(Coarse Woody Debris, CWD)对森林生态系统的稳定性具有不可忽视的贡献.对广州3种森林CWD的储量与分解特征进行了调查分析,结果表明:(1)CWD储量及其与相应森林总生物量比值均表现为常绿阔叶林>针阔混交林>针叶林;枯立木与倒木为CWD的主要成分,其中,针叶CWD主要物种为马尾松(Pinus massonianai),阔叶CWD物种主要为荷木(Schima superba)与黄杞(Engelhardtia chrysolepis)等.(2)CWD径级主要集中在<10cm的范围内,存在状态主要为中级腐烂状态,干扰与竞争是3种森林CWD产生的主要因素.(3)针叶林、针阔混交林与常绿阔叶林CWD的分解常数k分别为0 0244、0.0407和0.0487,即分解速率为常绿阔叶林>混交林>针叶林;随着CWD的分解,N、P与木质素的含量逐渐升高,C、C/N与木质素/N呈降低趋势.     

Diversity of wood-inhabiting Agaricomycotina on wood of different size classes in riparian forests of Uruguay

Many Agaricomycotina species are saprobes, playing a fundamental role in nutrient cycling in forest ecosystems by decomposing wood. Little is known about factors affecting diversity of wood-inhabiting fungi in the neotropical, warm temperate native forests of Uruguay. Most of these native forests are riparian harboring about 300 tree species. In this study, we assessed the diversity of wood-inhabiting fungi on wood of different size classes in riparian forests of Uruguay. We recovered 186 species of Agaricomycotina, including 113 corticioid and 58 polyporoid taxa. Eleven taxa accounted for 38% of the all the samples. The highest number of species was found on fine woody debris (FWD, 2–10 cm diam) than coarse woody debris (CWD, >10 cm diam) and very fine woody debris (VFWD, <2 cm diam). Species-accumulation curves did not reach an asymptote for any of the groups or wood diameter classes studied. Polyporoids were more frequently recorded on CWD (61% of collections) and corticioids on VFWD (77%). Species richness estimated by non-parametric estimators indicates an Agaricomycotina species richness between 450 and 700 taxa. Our results show that Uruguayan riparian forests, despite its limited area and fragmentation, support a wood-inhabiting Agaricomycotina diversity comparable to less fragmented forests with more plant diversity.     

川西亚高山森林木质残体及其附生苔藓持水特性

木质残体及其附生苔藓是森林生态系统的基本结构,在水源涵养、水土保持、生物多样性保育、碳和养分循环等方面具有重要作用。然而,有关亚高山森林木质残体及其附生苔藓的水源涵养功能研究尚未见报道。因此,以川西亚高山针叶林区紫果云杉原始林、岷江冷杉原始林、方枝柏原始林等8种典型森林类型为研究对象,调查研究了木质残体及其附生苔藓持水能力随林型、腐烂等级和径级的变化特征。结果表明：（1）亚高山针叶林木质残体饱和储水量在林型之间变化显著。其中,紫果云杉原始林具有最大的饱和储水量（22.06 mm）,柳树次生林木质残体饱和储水量最小（4.55 mm）,但林型之间木质残体饱和持水率的差异不显著。（2）不同林型的附生苔藓持水能力具有显著差异。其中,紫果云杉原始林木质残体附生苔藓饱和储水量最高（7.01 mm）,方枝柏原始林最低（0.21 mm）。（3）腐烂等级显著影响木质残体饱和储水量,IV腐烂等级的饱和储水量较高（3.77 mm）,II腐烂等级较低（0.75 mm）;木质残体饱和持水率和附生苔藓饱和储水量与腐解程度显著正相关,均符合Q（Q''）=ex²+fx+g的函数关系式。（4）木质残体和附生苔藓的饱和储水量均随径级增大而升高,大径级（D5）的饱和储水量占比分别超过60%和40%。可见,林型和木质残体腐解程度及径级大小是决定亚高山针叶林区木质残体及其附生苔藓持水性能的关键因子,特别是附生苔藓对于提高亚高山针叶林水源涵养具有重要作用。     

Coarse woody debris in Australian forest ecosystems: A review

Abstract Coarse woody debris (CWD) is the standing and fallen dead wood in a forest and serves an important role in ecosystem functioning. There have been several studies that include estimates of CWD in Australian forests but little synthesis of these results. This paper presents findings from a literature review of CWD and fine litter quantities. Estimates of forest‐floor CWD, snags and litter from the literature are presented for woodland, rainforest, open forest and tall open forest, pine plantation and native hardwood plantation. Mean mass of forest floor CWD in Australian native forests ranged from 19 t ha^?1 in woodland to 134 t ha^?1 in tall open forest. These values were generally within the range of those observed for similar ecosystems in other parts of the world. Quantities in tall open forests were found to be considerably higher than those observed for hardwood forests in North America, and more similar to the amounts reported for coniferous forests with large sized trees on the west coast of the USA and Canada. Mean proportion of total above‐ground biomass as forest floor CWD was approximately 18% in open forests, 16% in tall open forests, 13% in rainforests, and 4% in eucalypt plantations. CWD can be high in exotic pine plantations when there are considerable quantities of residue from previous native forest stands. Mean snag biomass in Australian forests was generally lower than the US mean for snags in conifer forests and higher than hardwood forest. These results are of value for studies of carbon and nutrient stocks and dynamics, habitat values and fire hazards.     

Influence of fine woody debris on spider diversity and community structure in forest leaf litter

Exploitation of forests for biofuel results in variable amounts of fine woody debris (FWD) being left on the forest floor. Such practices have the potential to affect the diversity and taxonomic composition of litter-dwelling organisms. The ability to predict the consequences of harvesting biofuel will be strengthened by knowledge of how variation in FWD affects litter organisms in non-harvested forests. A field experiment with three treatments [Control (no manipulation), FWD Removal, and FWD Addition] was carried out in replicated open plots in an unexploited forest in order to determine how the amount of fallen FWD affects the abundance, diversity and community composition of spiders, selected because of their importance as predators in the leaf-litter food web. Manipulation of FWD started in August, and by the following July total spider density in the FWD Addition treatment was 1.6× that of FWD Removal plots. Spider density in FWD Removal plots was 30% lower than the Control treatment, with no statistically significant difference between Control and FWD Addition. Manipulating FWD had no statistically significant effects on standard indices of spider diversity. However, multivariate community-level analyses revealed statistically significant differences in spider community structure between the FWD Removal and FWD Addition treatments. Two dominant genera of web builders contributed the most to this effect of manipulating FWD. Our results, and the findings of studies of woody debris in younger forests, suggest that the relatively small effects of FWD in our field experiment may reflect the age of the forest, with effects on the fauna likely being larger in forests younger than the one we studied.     

From Models to Measurements: Comparing Downed Dead Wood Carbon Stock Estimates in the U.S. Forest Inventory

The inventory and monitoring of coarse woody debris (CWD) carbon (C) stocks is an essential component of any comprehensive National Greenhouse Gas Inventory (NGHGI). Due to the expense and difficulty associated with conducting field inventories of CWD pools, CWD C stocks are often modeled as a function of more commonly measured stand attributes such as live tree C density. In order to assess potential benefits of adopting a field-based inventory of CWD C stocks in lieu of the current model-based approach, a national inventory of downed dead wood C across the U.S. was compared to estimates calculated from models associated with the U.S.’s NGHGI and used in the USDA Forest Service, Forest Inventory and Analysis program. The model-based population estimate of C stocks for CWD (i.e., pieces and slash piles) in the conterminous U.S. was 9 percent (145.1 Tg) greater than the field-based estimate. The relatively small absolute difference was driven by contrasting results for each CWD component. The model-based population estimate of C stocks from CWD pieces was 17 percent (230.3 Tg) greater than the field-based estimate, while the model-based estimate of C stocks from CWD slash piles was 27 percent (85.2 Tg) smaller than the field-based estimate. In general, models overestimated the C density per-unit-area from slash piles early in stand development and underestimated the C density from CWD pieces in young stands. This resulted in significant differences in CWD C stocks by region and ownership. The disparity in estimates across spatial scales illustrates the complexity in estimating CWD C in a NGHGI. Based on the results of this study, it is suggested that the U.S. adopt field-based estimates of CWD C stocks as a component of its NGHGI to both reduce the uncertainty within the inventory and improve the sensitivity to potential management and climate change events.     

湘南典型次生林粗木质残体生物量、碳储量和养分特征

对九龙江森林公园次生常绿阔叶林粗木质残体(CWD)进行量化研究,了解亚热带典型次生林CWD的本底数据。以九龙江森林公园典型天然次生林中的6个20m×20m的标准样地为对象,调查并分析样地内不同分解等级CWD的生物量、碳储量和养分特征变化。九龙江森林公园亚热带典型次生林的CWD储量在2.8—30.4t/hm~2之间,碳储量在6.10—6.75t/hm~2之间,大量营养元素(N、P、K、 Ca、 Mg)含量相对稳定,化学计量变化不显著,微量营养元素中Pb与Cd元素含量随着分解等级的增加而增加。九龙江森林公园亚热带典型次生林的CWD储量在亚热带常绿阔叶林中处于中等水平,CWD主要以分解中后期的倒木为主,反映出该次生林处于中幼龄林阶段,具有较高的碳储量,养分含量相对稳定,并表现出Pb与Cd在CWD中积累的现象,在中度分解和重度分解的CWD中Pb含量分别增加了62.65%和69.88%,Cd含量则分别增加了33.33%和100%,其内在机理有待进一步深入研究。研究结果有助于进一步了解CWD如何参与森林生态系统养分循环、重金属积累等生态过程,比较不同林分干扰历史下CWD储量、分布及养分特征的异质性...     

Understorey plant community structure in lower montane and subalpine forests, Grand Canyon National Park, USA

Aim Our objectives were to compare understorey plant community structure among forest types, and to test hypotheses relating understorey community structure within lower montane and subalpine forests to fire history, forest structure, fuel loads and topography. Location Forests on the North Rim of Grand Canyon National Park, Arizona, USA. Methods We measured understorey (< 1.4 m) plant community structure in 0.1‐ha plots. We examined differences in univariate response variables among forest types, used permutational manova to assess compositional differences between forest types, and used indicator species analysis to identify species driving the differences between forest types. We then compiled sets of proposed models for predicting plant community structure, and used Akaike's information criterion (AIC_C) to determine the support for each model. Model averaging was used to make multi‐model inferences if no single model was supported. Results Within the lower montane zone, pine–oak forests had greater understorey plant cover, richness and diversity than pure stands of ponderosa pine (Pinus ponderosa P. & C. Lawson var. scopulorum Engelm.). Plant cover was negatively related to time since fire and to ponderosa pine basal area, and was highest on northern slopes and where Gambel oak (Quercus gambelii Nutt.) was present. Species richness was negatively related to time since fire and to ponderosa pine basal area, and was highest on southern slopes and where Gambel oak was present. Annual forb species richness was negatively related to time since fire. Community composition was related to time since fire, pine and oak basal area, and topography. Within subalpine forests, plant cover was negatively related to subalpine fir basal area and amounts of coarse woody debris (CWD), and positively related to Engelmann spruce basal area. Species richness was negatively related to subalpine fir basal area and amounts of CWD, was positively related to Engelmann spruce basal area, and was highest on southern slopes. Community composition was related to spruce, fir and aspen basal areas, amounts of CWD, and topography. Main conclusions In montane forests, low‐intensity surface fire is an important ecological process that maintains understorey communities within the range of natural variability and appears to promote landscape heterogeneity. The presence of Gambel oak was positively associated with high floristic diversity. Therefore management that encourages lightning‐initiated wildfires and Gambel oak production may promote floristic diversity. In subalpine forests, warm southern slopes and areas with low amounts of subalpine fir and CWD were positively associated with high floristic diversity. Therefore the reduction of CWD and forest densities through managed wildfire may promote floristic diversity, although fire use in subalpine forests is inherently more difficult due to intense fire behaviour in dense spruce–fir forests.     

Structures determining bryophyte species richness in a managed forest landscape in boreo-nemoral Europe

Forest patches with high biological value are protected as woodland key habitats (WKH), which are identified by the presence of forest structures and indicator species. However, management for conservation needs to consider also managed forests as habitats for species. In this respect, there is a need to set quantitative targets for species and structures at different landscape scales. Due to non-intensive methods of forest management used prior to 1940 in Latvia, it might be expected that large areas of forest have developed structures that can support many species characteristic of natural forests. The aim of the study was to create a model that best described the richness of bryophyte species that are characteristic of natural forests, using forest structures as explanatory factors. The structures and bryophyte communities on living trees and coarse woody debris (CWD) were described in plots along transects blindly placed in areas dominated by State forests under commercial management. Explanatory variables related to tree species composition and tree size explained 54% of the variation in WKH indicator species richness on living trees. The best explanatory factors were maximum diameter of deciduous tree species and CWD. Low richness of total bryophyte and indicator species was found on dead wood, and the amount of variation in bryophyte species richness on CWD explained by explanatory variables was low. The study indicates the importance of deciduous tree substrate in managed forests in maintaining the spatial continuity of epiphytic species diversity. However, the forests in the managed forest landscape did not support high diversity of epixylic species, even in the WKHs, due to low diversity of suitable dead wood substrate.     

东北东部山区11种温带树种粗木质残体分解与碳氮释放

采用长期定位跟踪实测方法,比较分析了我国东北温带森林4个水热状况不同的立地条件(红松(Pinus koraiensis)人工林、硬阔叶林、蒙古栎(Quercus mongolica)林和林外空旷地)下11个温带树种粗木质残体(CWD)分解初期3年中的碳氮动态及其影响因子。测定树种包括:白桦(Betula platyphylla)、山杨(Populus davidiana)、紫椴(Tilia amurensis)、胡桃楸(Juglans mandshurica)、蒙古栎、色木槭(Acer mono)、春榆(Ulmus japonica)、红松、黄檗(Phellodendron amurense)、兴安落叶松(Larix gmelinii)和水曲柳(Fraxinus mandshurica)。结果表明:在分解过程中,所有树种CWD的碳浓度没有明显变化(p0.05),但其干重、碳密度、氮浓度和氮密度均随分解进程不同程度地减小,碳氮比(C/N)则增大,而且树种间差异显著(p0.001)。针叶树种的CWD分解速率显著地低于阔叶树种,其中白桦的3年CWD干重损失率(65%)约为兴安落叶松(22%)的3倍。径级大的CWD分解较慢。CWD分解与碳氮释放均与CWD的初始N含量呈正相关,而与初始C/N呈负相关。4个立地条件下CWD的干重和碳氮含量的变化差异不显著,均表现出一致的变化趋势。该研究指出,在分解初期的前3年中,CWD基本上是一个碳源和氮源。     

Long-term movement of 15N tracers into fine woody debris under chronically elevated N inputs

Two key questions in the study of large-scale C (carbon) and N (nitrogen) cycling in temperate forests are how N cycling in soil detritus controls ecosystem-level retention of elevated N deposition, and whether elevated N deposition is likely to cause increases in C pools. The large C:N ratios in woody detritus make it a potentially important contributor to N retention, if N immobilization increases, and a potentially important contributor to C sequestration, if pool sizes increase. We studied N concentrations, C:N ratios, and pool sizes of N and biomass in fine woody debris (FWD < 5 cm diam.) 12 years into a long-term N-amendment study in two contrasting forests, a naturally-regenerated forest dominated by Quercus spp., and a 63-yr old plantation of Pinus resinosa. We also quantitatively recovered ¹⁵N tracers (originally applied as ¹⁵NH₄ and ¹⁵NO₃) in FWD, eight years following their application in the same study, in both ambient and N-amended plots. We used these data to test predictions of tracer redistributions made by a biogeochemical process model that included ¹⁵N. Results from the N pool-size analysis and the ¹⁵N tracer-recovery analysis indicated that under elevated N inputs of 5 g N m^–2 yr^–1 (as NH₄NO₃) over the decadal time period, only 0.15%–0.76% of the elevated N inputs were recovered in FWD of N-amended plots relative to ambient. Any increase in N immobilization in wood appeared to be minimal, in agreement with model predictions. Under N amendments, pool sizes of C in FWD were not significantly different from ambient, whereas pool sizes of N were marginally higher. Patterns of ¹⁵NH₄ vs. ¹⁵NO₃ recovery, treatment differences, and forest-type differences suggested that plant uptake, rather than detrital immobilization, was the dominant mechanism of ¹⁵N tracer movement into FWD. This result indicates that plant-soil cycling operating over a decadal time scale or longer controls C:N ratios and N pool sizes in woody debris.     

Guild structure of wood-rotting fungi based on volume and decay stage of coarse woody debris

Wood-rotting fungi are major organisms exploiting coarse woody debris (CWD) in forests. Here, guild structure of wood-rotting fungi was investigated in cool temperate (Chichibu) and warm temperate (Chiba) forests in central Japan, based on their occurrence on CWD of different volumes and decay stages. Analysis with the program partitioning around medoids (PAM) recognized two clusters in Chichibu and four in Chiba, and their silhouette coefficients (an index for reliability of clustering) were adequately high, suggesting the existence of non-random clustering structure. To examine whether the clustering structure observed in this study was based on fungal preference for CWD or discontinuities in the distribution of CWD characteristics (decay stage and volume), null model analyses were made in which all fungal species were randomly redistributed among CWD. Silhouette coefficients based on the null model were reasonably high in both localities, indicating the observed clustering structure was at least partly attributable to the discontinuity in the distribution of CWD characteristics. In addition, the silhouette coefficient of the observed clustering structure was significantly higher than that of the null model in both localities. This result suggests that guild structure was present in the wood-rotting fungal communities, and the difference in CWD preference among wood-rotting fungi contributed to the structuralization of the communities.     

鼎湖山锥栗粗木质残体的分解和元素动态

选取锥栗(Castanopsischinensis)粗木质残体(coarsewoodydebris,CWD)的3个径级(径级1-3分别为5-10cm,10-20cm和20-30cm),并且将每个径级的锥栗粗木质残体分为Ⅰ、Ⅱ、Ⅲ、Ⅳ、Ⅴ5个分解阶段。通过测定锥栗粗木质残体3个径级的Ⅰ-Ⅲ阶段的7种化学元素(C、N、P、K、Ca、Na、Mg)的浓度和CWD密度变化来研究其分解过程中的元素动态、分解速率及其分解过程中基质质量的变化。到分解阶段Ⅲ时,3个径级的重量与原来相比分别损失了36%、48%和43%。元素N、P、Mg、Ca、Na的浓度升高。元素N的累积可能和锥栗粗木质残体中寄生着固氮细菌和真菌有关。元素P、Mg、Ca、Na的浓度升高则可能是由于这4个元素因淋溶流失的速度小于锥栗粗木质残体质量的损失速度,造成元素积聚,元素C、K的浓度降低。N/P比值是较好的分解指示指标。虽然存在元素的净释放,但是由于C和其它主要元素的释放速率较慢,因而锥栗粗木质残体是森林中重要的C库和长期的元素来源。     

Coarse Woody Debris Stimulates Soil Enzymatic Activity and Litter Decomposition in an Old-Growth Temperate Forest of Patagonia, Argentina

In most temperate forest ecosystems, tree mortality over time generates downed logs that accumulate as coarse woody debris (CWD) on the forest floor. These downed logs and trunks have important recognized ecosystem functions including habitat for different organisms and long-term organic C storage. Due to its recalcitrant chemical composition and slow decomposition, CWD can also have direct effects on ecosystem carbon and nutrient turnover. CWD could also cause changes indirectly through the physical and chemical alterations that it generates, although it is not well-understood how important these indirect effects could be for ecosystem processes and soil biogeochemistry. We hypothesized that in an old-growth mature forest, CWD affects carbon and nutrient cycles through its “proximity effects”, meaning that the forest floor near CWD would have altered soil biotic activity due to the environmental and biogeochemical effects of the presence of CWD. We conducted our study in an old-growth southern beech temperate forest in Patagonia, Argentina, where we estimated and classified the distribution and mass, nutrient pools and decay stage of CWD on the forest floor, and evaluated its impact on litter decomposition, soil mites and soil enzymatic activity of carbon and phosphorus-degrading enzymes. We demonstrate here that CWD in this ecosystem represents an important organic carbon reservoir (85 Mg ha^?1) and nitrogen pool (0.42 Mg ha^?1), similar in magnitude to other old-growth forests of the Northern Hemisphere. In addition, we found significant proximity effects of CWD, with increased C-degrading soil enzyme activity, decreased mite abundance, and more rapid litter decomposition beneath highly decayed CWD. Considered at the ecosystem scale in this forest, the removal of CWD could cause a decrease of 6% in soil enzyme activity, particularly in the summer dry season, and nearly 15% in annual litter decomposition. We conclude that beyond the established importance of CWD as a long-term carbon reservoir and habitat, CWD contributes functionally to the forest floor by influencing the spatial heterogeneity of microbial activity and carbon and nutrient turnover. These proximity effects demonstrate the importance of maintenance of this ecosystem component and should be taken into consideration for management decisions pertaining to carbon sequestration and functional diversity in natural forest ecosystems.     

Accumulation and Connectivity of Coarse Woody Debris in Partial Harvest and Unmanaged Relict Forests

When a tree dies, it continues to play an important ecological role within forests. Coarse woody debris (CWD), including standing deadwood (SDW) and downed deadwood (DDW), is an important functional component of forest ecosystems, particularly for many dispersal-limited saproxylic taxa and for metapopulation dynamics across landscapes. Processes, such as natural disturbance or management, modify forest composition and structure, thereby influencing CWD abundance and distribution. Many studies have compared older forests to forests managed with even-aged silvicultural systems and observed a prolonged period of low CWD occurrence after harvesting. With fine-scale spatial data, our study compares the long-term impacts of light partial harvesting on the CWD structure of eastern deciduous hardwood forests. We mapped and inventoried DDW and SDW using variable radius plots based on a 10 m×10 m grid throughout an unmanaged, structurally-complex relict forest and two nearby forests that were partially harvested over 46 years ago. The relict stand had significantly larger individual pieces and higher accumulations of DDW and SDW than both of the partially harvested stands. Connectivity of CWD was much higher in the relict stand, which had fewer, larger patches. Larger pieces and higher proportion of decay-resistant species (e.g. Quercus spp.) in the relict forest resulted in slower decomposition, greater accumulation and increased connectivity of CWD. Partial harvests, such that occur with selection forestry, are generally considered less disruptive of ecosystem services, but this study highlights the long-term impacts of even light partial harvests on CWD stocks and distribution. When planning harvesting events, forest managers should also consider alternative methods to ensure the sustainability of deadwood resources and function.