Wood-decay type and fungal guild dominance across a North American log transplant experiment

We incubated 196 large-diameter aspen (Populus tremuloides), birch (Betula papyrifera), and pine (Pinus taeda) logs on the FACE Wood Decomposition Experiment encompassing eight climatically-distinct forest sites in the United States. We sampled dead wood from these large-diameter logs after 2 to 6 y of decomposition and determined wood rot type as a continuous variable using the lignin loss/density loss ratio (L/D) and assessed wood-rotting fungal guilds using high-throughput amplicon sequencing (HTAS) of the ITS-2 marker. We found L/D values in line with a white rot dominance in all three tree species, with pine having lower L/D values than aspen and birch. Based on HTAS data, white rot fungi were the most abundant and diverse wood-rotting fungal guild, and soft rot fungi were more abundant and diverse than brown rot fungi in logs with low L/D values. For aspen and birch logs, decay type was related to the wood density at sampling. For the pine logs, decay type was associated with the balance between white and brown/soft rot fungi abundance and OTU richness. Our results demonstrate that decay type is governed by biotic and abiotic factors, which vary by tree species.     

腐烂等级、径级对典型阔叶红松林红松倒木物理化学性质的影响

倒木是森林生态系统的重要组成部分, 在地力维护、生物多样性保持以及碳(C)和养分循环等方面具有重要意义, 但倒木物理化学性质随其腐烂等级和径级而变化。为了深入理解腐烂等级和径级对倒木物理化学性质的影响, 该研究以典型阔叶红松林的建群种——红松(Pinus koraiensis)的倒木为研究对象, 将其每个腐烂等级(I-V)下的倒木分为4个径级(径级i ≤ 10.0 cm、径级ii 10.1-30.0 cm、径级iii 30.1-50.0 cm、径级iv >50.0 cm), 研究了不同腐烂等级、径级及两者交互作用对倒木心材和边材物理化学性质的影响。结果表明: 心材和边材具有相似的变化规律。倒木心材和边材含水率随着腐烂等级增加而增加, 而木材密度随腐烂等级和径级的增加均呈下降趋势; 边材C含量以及心材和边材的氮(N)、磷(P)含量随腐烂等级增加呈上升趋势, 心材N、P含量随径级增加呈先增加后减少的趋势; 纤维素含量随腐烂等级增加呈下降趋势, 而木质素含量呈上升趋势, 纤维素和木质素含量随径级增加没有明显变化规律。倒木含水率与C、N、P、木质素含量(除心材P含量)显著正相关, 与纤维素含量显著负相关; 木材密度与C、N、P、木质素含量显著负相关, 与纤维素含量显著正相关。由此可见, 倒木物理化学性质受不同腐烂等级和径级的影响有各自的变化规律, 且倒木的物理性质(含水率和木材密度)是影响化学含量变化的重要因素。     

Effects of decay classes and diameter classes on physico-chemical properties of Pinus koraiensis log in a typical mixed broadleaved-Korean pine forest

AimsLog is an important component for most of forest ecosystems. It plays crucial roles in maintaining soil fertility, sustaining biodiversity and cycling of carbon (C) and nutrient. However, physico-chemical properties of logs vary with decay classes and diameter classes. Our objective was to study effects of decay classes and diameter classes on physico-chemical properties of logs in a typical mixed broadleaved-Korean pine forest in northern China.MethodsIn this study, logs of Pinus koraiensis were chosen as it was the constructive species in the typical mixed broadleaved-Korean pine forest. Logs of P. koraiensis at each decay classes (I-V) were divided into four diameter classes, including diameter class i ≤ 10.0 cm, diameter class ii: 10.1-30.0 cm, diameter class iii: 30.1-50.0 cm, and diameter class iv > 50.0 cm. Then, we explored effects of different decay classes, diameter classes and their interactions on the physico-chemical properties of logs for both the heartwood and sapwood.Important findings The results showed that the physico-chemical properties of heartwood and sapwood generally exhibited similar variations. Their moisture content both increased with an increasing decay class, whereas wood density both decreased with an increased decay class and diameter class. The carbon concentrations of the sapwood showed a trend of gradual increasing during decomposition, and there was an accumulation in nitrogen (N) and phosphorus (P) concentrations of the heartwood and sapwood with an increased decay class, simultaneously. Only N and P concentrations of the heartwood increased and then decreased with an increasing diameter class. The cellulose content decreased with an increasing decay class. In contrast to the cellulose, the lignin content increased with an increased decay class. However, cellulose and lignin contents exhibited no distinct trend among diameter classes. The moisture content of logs had a significant positive correlation with C, N, P concentrations and lignin content (except P concentrations of the heartwood), but had a significant negative correlation with the cellulose content (p< 0.05). The wood density was negatively correlated with C, N, P concentrations and the lignin content, but it was positively correlated with the cellulose content (p< 0.05). Therefore, physico-chemical properties of logs had unique patterns along both decay classes and diameter classes, and the physical properties of logs (the moisture content and wood density) were important factor affecting the variations of their chemical contents.     

Decomposition of Japanese beech wood by diverse fungi isolated from a cool temperate deciduous forest

We assessed 62 fungal strains in 31 species of wood decay fungi in the ability to decompose wood blocks of Japanese beech (Fagus crenata) under a pure culture condition. Fungi were collected in a cool temperate beech forest in Japan and isolated from the inside of beech logs and from sporocarps fruiting on logs and snags of beech that were different in diameter and decay class. Fungi in Holobasidiomycetidae showed marked decomposition of lignin and carbohydrate. These fungi were divided into three groups according to the pattern of lignin and carbohydrate utilization. Phanerochaete filamentosa decomposed lignin selectively. Lampteromyces japonicus, Steccherinum rhois, Trichaptum biforme, Stereum ostrea, Mycena haematopoda, Antrodiella albocinnamomea, Daedalea dickinsii, Daedaleopsis tricolor, Ganoderma tsunodae, and Trametes versicolor decomposed lignin and carbohydrates simultaneously. Psathyrella candolleana, Lenzites betulinus, and Trametes hirsuta decomposed carbohydrates selectively. Species in the Phragmobasidiomycetidae and in the Ascomycota caused low mass loss of wood.     

Wood-inhabiting fungi on fallen logs of Norway spruce: relations to forest management and substrate quality

A survey of the patterns of wood decaying fungi as to occurrence of sporocarps on naturally fallen logs of Norway spruce ( Picea abies ) was undertaken in two nearby forest stands with different histories of management. One stand was an old-growth forest with few signs of logging, and the other stand was selectively logged 60–80 years ago. Altogether 118 species were found. Forest management had a negative impact on the species diversity. Newly fallen and weakly decayed logs in a natural forest had a higher species richness, more red-listed species, as well as more indicator species compared to similar logs in a managed forest. The importance of dead wood for species diversity of wood inhabiting fungi was clearly demonstrated. Presence of logs in later stages of decomposition increased the total species number in a natural forest stand with 42 (63 %), compared to a survey of only newly fallen and weakly decayed logs. Presence of logs in later stages of decomposition also increased the diversity of the species pool colonising newly fallen and weakly decayed logs. The highest number of fruiting species was found on intermediately decayed logs and on logs lying in contact with the ground. The fungal gradient as revealed in a DCA ordination was primarily related to decay. A successional pathway based on the primary decayer Fomitopsis pinicola was not detected.     

Spatial Ecology of the Fungal Genus Xylaria in a Tropical Cloud Forest

Fungal symbioses with plants are ubiquitous, ancient, and vital to both ecosystem function and plant health. However, benefits to fungal symbionts are not well explored, especially in non‐mycorrhizal fungi. The Foraging Ascomycete hypothesis proposes that some wood‐decomposing fungi may shift life‐history strategies to endophytism to bridge gaps in time and space between suitable substrates. To test this hypothesis we examine spatial relationships of Xylaria endophytic fungi in the forest canopy with Xylaria decomposer fungi on the forest floor. We sampled for fungi of the genus Xylaria using a spatially explicit sampling scheme in a remote Ecuadorian cloud forest, and concurrently carried out an extensive culture‐based sampling of fungal foliar endophytes. We found 36 species of Xylaria in our 0.5 ha plot, 31 of which were found to only occur as fruiting bodies. All five species of Xylaria found as endophytes were also found as fruiting bodies. We also tested the relationships of both stages of these fungi to environmental variables. Decomposer fungi were differentiated by species‐specific habitat preferences, with three species being found closer to water than expected by chance. In contrast, endophytes displayed no sensitivity to environmental conditions, such as host, moisture, or canopy cover. We found evidence of spatial linkage between life stages in two species. We also demonstrate that direct transmission of endophytes from leaves to woody substrates is possible. These results indicate that endophytism may represent one way for decomposer fungi to escape moisture limitation, and that endophytic fungi may act as sources of dispersal for decomposer fungi consistent with predictions of the Foraging Ascomycete hypothesis.     

丰林国家级自然保护区木腐真菌多样性与寄主倒木的关系

木腐真菌是一类以木材为生长基质的大型真菌, 通过分泌各种水解酶全部或部分降解木材中的木质素、纤维素和半纤维素, 促进森林生态系统的物质循环, 具有重要的生态功能。本研究调查了丰林国家级自然保护区固定样地中木腐真菌的多样性和倒木特征, 并进行了木腐真菌的物种多样性和数量与倒木的种类、数量、腐朽程度、直径大小等的相关性分析。结果显示: 在样地内共采集木腐真菌标本295份, 经鉴定为93种, Shannon多样性指数为3.86, Simpson指数为0.96。相关性分析发现木腐真菌的数量和种类与直径为2-5 cm和5-10 cm的倒木、2级腐烂的倒木和红松倒木均显著相关。样地中优势倒木寄主分别为槭属(Acer)、榛属(Corylus)、云杉属(Picea)和松属(Pinus), 这4类倒木上生长的木腐真菌种类组成具有明显的差异, 槭属和榛属倒木上的共有优势种主要是三色拟迷孔菌(Daedaleopsis tricolor)、云芝(Trametes versicolor)和桦附毛孔菌(Trichaptum pargamenum), 而松属和云杉属的共有优势种主要有白囊耙齿菌(Irpex lacteus)、云芝、冷杉附毛孔菌(Trichaptum abietinum)和褐紫附毛孔菌(T. fuscoviolaceum)。倒木产生真菌子实体的概率研究表明, 同一类寄主倒木上发生木腐真菌子实体的概率在调查面积增加到0.36 ha后趋于一个定值, 松属倒木中仅有10.2%产生真菌子实体, 槭属和云杉属分别是12.9%和13.4%, 榛属最高, 达到53.7%。本研究结果对于预测森林生态系统中木腐真菌的发生具有重要理论意义。     

A new source of methane in boreal forests

Methane was found among the gases evolved during natural wood decay caused by bracket fungi in boreal forests. Methane was detected both in decaying wood and fungal fruiting bodies. A scheme of symbiotic association of wood-degrading fungi and anaerobic microorganisms providing the methanogenesis in the wood was proposed. The scale of mycogenic methane emission has to be consistent with the huge volume of decaying wood in boreal forest ecosystems.     

A new source of methane in boreal forests

Methane was found among the gases evolved during natural wood decay caused by bracket fungi in boreal forests. Methane was detected both in decaying wood and fungal fruiting bodies. A scheme of symbiotic association of wood-degrading fungi and anaerobic microorganisms providing the methanogenesis in the wood was proposed. The scale of mycogenic methane emission has to be consistent with the huge volume of decaying wood in boreal forest ecosystems.     

Isolation of wood-inhabiting fungi from Canadian hardwood logs

Wood-inhabiting fungi include many molds, wood-staining fungi, and decay fungi. Most of these fungal species can result in economic losses to wood users. Studies on molds, staining fungi, and decay fungi are necessary to be able to control their growth on wood and wood products. In this study, wood-inhabiting fungi were isolated from logs of 3 major Canadian hardwood species: sugar maple, white birch, and yellow birch. Two media were used for isolation. From these 3 wood species, a total of 1198 fungal cultures were obtained from summer- and winter-harvested logs in dry storage and under water sprinkling. The results showed that most fungal species were not host specific and affected all of the wood species tested. Frequently isolated molds were Alternaria alternata, Trichoderma species, and Mucor/Rhizopus (Zygomycota) species, frequently isolated staining fungi were Ophiostoma piceae and Ophiostoma piliferum, a frequently isolated bark saprophyte was Nectria cinnabarina, and frequently isolated decay fungi were taxa of the phylum Basidiomycota. More fungal species were isolated from summer-harvested logs than from winter-harvested logs. Fewer fungal cultures, especially decay fungi, were isolated from logs in early storage than from logs in late storage.     

Fine‐scale spatiotemporal dynamics of fungal fruiting: prevalence,amplitude, range and continuity

Despite the critical importance of fungi as symbionts with plants, resources for animals, and drivers of ecosystem function, the spatiotemporal distributions of fungi remain poorly understood. The belowground life cycle of fungi makes it difficult to assess spatial patterns and dynamic processes even with recent molecular techniques. Here we offer an explicit spatiotemporal Bayesian inference of the drivers behind spatial distributions from investigation of a Swiss inventory of fungal fruit bodies. The unique inventory includes three temperate forest sites in which a total of 73 952 fungal fruit bodies were recorded systematically in a spatially explicit design between 1992 and 2006. Our motivation is to understand how broad‐scale climate factors may influence spatiotemporal dynamics of fungal fruiting within forests, and if any such effects vary between two functional groups, ectomycorrhizal (ECM) and saprotrophic fungi. For both groups we asked: 1) how consistent are the locations of fruiting patches, the sizes of patches, the quantities of fruit bodies, and of prevalence (occupancy)? 2) Do the annual spatial characteristics of fungal fruiting change systematically over time? 3) Are spatial characteristics of fungal fruiting driven by climatic variation? We found high inter‐annual continuity in fruiting for both functional groups. The saprotrophic species were characterised by small patches with variable fruit body counts. In contrast, ECM species were present in larger, but more distinctly delimited patches. The spatial characteristics of the fungal community were only indirectly influenced by climate. However, climate variability influenced overall yields and prevalence, which again links to spatial structure of fruit bodies. Both yield and prevalence were correlated with the amplitudes of occurrence and of fruit body counts, but only prevalence influenced the spatial range. Summarizing, climatic variability affects forest‐stand fungal distributions via its influence on yield (amount) and prevalence (occupancy), whereas fungal life‐history strategies dictate fine‐scale spatial characteristics.     

古田山国家级自然保护区木腐真菌物种多样性及分布

木腐真菌是微生物的一个重要类群, 主要以倒木为生长基质, 通过产生各种水解酶将倒木的纤维素、木质素和半纤维素分解为小分子物质, 对促进森林生态系统中的营养物质循环发挥着重要的生态功能。于2016年8月在浙江古田山国家级自然保护区开展的木腐真菌野外调查, 利用形态学和DNA序列分析对采集的标本进行了物种鉴定, 并分析了木腐真菌的物种组成和地理成分。在采集的158份标本中鉴定木腐真菌45属92种, 其中白腐真菌78种, 褐腐真菌14种。古田山的木腐真菌物种区系组成中, 热带-亚热带成分比例最高。在158份木腐真菌标本中, 97份标本采自直径大于10 cm的倒木或树桩上, 分属于76个种, 是木腐真菌生长的主要基质大小类型; 48份标本采自直径为2-10 cm的枝干上, 分属38个种; 13份标本采自直径小于2 cm的枝干上, 分属12种。不同腐烂等级倒木上生长的真菌数量和种类差异明显, 其中一级腐烂倒木上采集到9份标本(7种), 二级腐烂倒木上采集到86份标本(45种), 三级腐烂倒木上49份标本(29种), 四级腐烂倒木上14份标本(14种)。结果表明, 林分中倒木直径大小和腐烂程度是影响木腐真菌生长与分布的重要因子。     

Fungal wood decomposer activities influence community structures of myxomycetes and bryophytes on coarse woody debris

Dead wood is an important habitat for forest organisms, and wood decay fungi are the principal agents determining the dead wood properties that influence the communities of organisms inhabiting dead wood. In this study, we investigated the effects of wood decomposer fungi on the communities of myxomycetes and bryophytes inhabiting decayed logs. On 196 pine logs, 72 species of fungi, 34 species and seven varieties of myxomycetes, and 16 species of bryophytes were identified. Although white rot was the dominant decay type in sapwood and heartwood, brown and soft rots were also prevalent, particularly in sapwood. Moreover, white rot and soft rot were positively and brown rot negatively correlated with wood pH. Ordination analyses clearly showed a succession of cryptogam species during log decomposition and showed significant correlations of communities with the pH, water content, and decay type of wood. These analyses indicate that fungal wood decomposer activities strongly influence the cryptogam communities on dead wood.     

Species richness of tropical wood-inhabiting macrofungi provides support for species-energy theory

A study was undertaken at the El Verde Field Station in Puerto Rico to determine the effect of energy available from newly dead trees on the species richness of macrofungal communities that inhabit them. It is hypothesized that there is a positive relationship between available energy and species richness. Energy was measured using the volume of the dead trees and the wood density of living trees of the same species. One hundred ninety-four logs of known tree species were surveyed 1 y for fruiting bodies of macrofungi at monthly intervals. For individual logs, log volume had a significant positive effect on macrofungal species richness. Younger logs had significantly higher species richness than older logs, and those with less apparent decay had more species than those with more decay. When logs were grouped by tree species, total wood volume and density of live wood had a significant positive effect and average log diameter had a negative effect on total species richness and abundance of the wood-inhabiting macrofungi. Macrofungal richness and abundance constantly increased with initial wood density; there was no evidence for a unimodal relationship. These results support the proposed relationship between species richness and energy.     

Wood ash application in a managed Norway spruce plantation did not affect ectomycorrhizal diversity or N retention capacity

Ectomycorrhizal (ECM) fungi are key players in N cycling in coniferous forests, and forest management such as application of wood ash can affect their functionality. The aim of this study was to determine the effects of wood ash application on ECM fungal mycelial production, capacity to retain N, diversity and community composition. In-growth mesh bags were installed in control and treated plots. After 6 months, ¹⁵N labeled ammonium and nitrate were applied into the mesh bags, and 24 h later extramatrical mycelium (EMM) was extracted and analyzed. Wood ash had no effects on EMM in-growth, N retention capacity, diversity or community composition. In contrast, there were significant seasonal differences in the amount of EMM produced. These results demonstrate that applying up to 6 t ha⁻¹ of wood ash in this type of plantation forest is a safe management practice that does not increase N leaching or negatively affect ECM fungi.     

Successional development of wood-inhabiting fungi associated with dominant tree species in a natural temperate floodplain forest

Fungi play a crucial role in dead wood decay, being the major decomposers of wood and affecting microbiota associated with dead wood. We sampled dead wood from five deciduous tree species over more than forty years of decay in a natural European floodplain forest with high tree species diversity. While the assembly of dead wood fungal communities shows a high level of stochasticity, it also indicates clear successional patterns, with fungal taxa either specific for early or late stages of wood decay. No clear patterns of fungal biomass content over time were observed. Out of 220 major fungal operational taxonomic units, less than 8% were associated with a single tree species, most of them with Quercus robur. Tree species and wood chemistry, particularly pH, were the most important drivers of fungal community composition. This study highlights the importance of dead wood and tree species diversity for preserving the biodiversity of fungi.     

Wood Decomposition of Cyrilla racemiflora (Cyrillaceae) in Puerto Rican Dry and Wet Forests: A 13‐year Case Study1

We studied the decomposition of Cyrilla racemiflora logs over a 13‐yr period in tropical dry and wet forests in Puerto Rico. The mean mass loss, ratio of soft to hard wood, nutrient concentrations, and the diversity of wood‐inhabiting organisms were greater in logs decomposing in the dry forest than in the wet forest. Termites were also more abundant in the logs collected from the tropical dry forest than the tropical wet forest. High moisture content and a low animal diversity on the logs in the wet forest seem to retard wood decay in this habitat. Wood decay rates in the tropical dry forest can be related to the high diversity of species and functional groups of wood‐inhabiting organisms.     

Soil-wood interactions: Influence of decaying coniferous and broadleaf logs on composition of soil fungal communities

Wood-inhabiting fungi may affect soil fungal communities directly underneath decaying wood via their exploratory hyphae. In addition, differences in wood leachates between decaying tree species may influence soil fungal communities. We determined the composition of fungi in 4-yr old decaying logs of Larix kaempferi and Quercus rubra as well as in soil directly underneath and next to logs. Fungal community composition in soil covered by logs was different from that in wood and uncovered soil and was clearly influenced by the tree species. Soil fungal species richness under logs was lower than in uncovered soil but higher than in decaying wood. The amount of exploratory hyphae of log-inhabiting fungi was only high close to decaying logs. In conclusion, there is a small but significant effect of decaying coniferous and broadleaf logs on soil fungal communities directly underneath logs, likely affected by differences in wood chemistry and fungal preference between tree species.     

Forest microsite effects on community composition of ectomycorrhizal fungi on seedlings of Picea abies and Betula pendula

Niche differentiation in soil horizons, host species and natural nutrient gradients contribute to the high diversity of ectomycorrhizal fungi in boreal forests. This study aims at documenting the diversity and community composition of ectomycorrhizal fungi of Norway spruce ( Picea abies ) and silver birch ( Betula pendula ) seedlings in five most abundant microsites in three Estonian old-growth forests. Undisturbed forest floor, windthrow mounds and pits harboured more species than brown- and white-rotted wood. Several species of ectomycorrhizal fungi were differentially represented on either hosts, microsites and sites. Generally, the most frequent species in dead wood were also common in forest floor soil. Ordination analyses suggested that decay type determined the composition of EcM fungal community in dead wood. Root connections with in-growing mature tree roots from below affected the occurrence of certain fungal species on seedling roots systems in dead wood. This study demonstrates that ectomycorrhizal fungi differentially establish in certain forest microsites that is attributable to their dispersal and competitive abilities. Elevated microsites, especially decayed wood, act as seed beds for both ectomycorrhizal forest trees and fungi, thus affecting the succession of boreal forest ecosystems.     

森林生态系统中木腐真菌群落形成机理及生态功能

在森林生态系统中,枯死木是一个重要的组成部分,为很多生物提供栖息地,有助于养分循环以及碳和水的储存。木材分解是森林生态系统养分循环、土壤形成和碳收支的决定性过程,越来越受到森林生态学家、病理学家和管理者的重视。在此过程中,木腐真菌通过分泌多种酶降解木材主要成分,实现生态系统中的物质循环,具有极为关键和重要的作用。木腐真菌群落的形成受多种因素的影响,环境因子(降水、纬度、光照、温度、湿度等)和寄主倒木特征(化学成分、物理结构、体积大小以及腐烂程度等)通过影响不同真菌物种的分布和生长,进而影响真菌群落的物种组成;木腐真菌在一定程度上对其周围的微生物群落具有较强的控制力,其中能形成菌索的真菌更擅长控制细菌群落,而细菌通过消耗真菌降解酶的活性分解产物间接影响分解速率,也影响着真菌群落的动态变化;木腐真菌和腐生无脊椎动物作为倒木分解的两类主要群体,它们彼此之间相互影响,既有抑制作用也存在互惠互利;一些土壤真菌类群通过菌丝扩散定殖于倒木上进行生长。木腐真菌群落组成的最大干扰来自人类对森林的砍伐和利用,倒木的移除直接造成木腐真菌生长基质和环境的丧失,许多物种濒临灭绝,使得真菌群落组成单一、功能相对简单化。在倒木降解过程中,白腐真菌主要分泌过氧化物酶和漆酶降解倒木木质素,褐腐真菌可以有效地降解半纤维素和纤维素,降解酶的活性和种类直接影响木材腐烂速率和营养元素含量,而真菌菌丝对木材中C、N等营养元素具有重要的吸收和转移作用。木腐真菌的功能性状能够反映物种对环境条件的适应,有助于理解在不同环境中影响真菌群落聚集的机制。揭示森林生态系统中木腐真菌群落的形成机理及其生态功能,对于研究森林生态系统物质循环过程和生物多样性维持具有重要的理论意义和参考价值。