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.     

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     

丰林自然保护区阔叶红松林粗木质残体研究

粗木质残体(CWD)是森林生态系统的重要组成部分,对维持森林生态系统健康具有不可忽视的贡献.采用典型取样法在黑龙江丰林国家级自然保护区设置了17块20×20m的标准样地,拟对保护区内阔叶红松林中粗木质残体的贮量、形态组成、直径、长度及腐烂度等特征进行研究.研究结果表明:(1)CWD总贮量为75.1m³·hm^-2,其中倒木,枯立木和树桩的贮量分别为.24 m³·hm^-2,2.03m³·hm^-2和7.34m³ ·hm^-2.(2)倒木和枯立木的优势径级范围分别为11cm～20cm(占CWD总数的4%)和0-10cm(45%);倒木和枯立木的优势长度范围分别为0-5m (%)和m-10m(49%).(3)CWD腐烂度呈近正态分布,且主要分布在Ⅱ(28%),Ⅲ(35%)和Ⅳ(2%)腐烂等级上.     

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.     

人为干扰对大兴安岭北坡兴安落叶松林粗木质残体的影响

比较了兴安落叶松天然林和两种不同干扰类型兴安落叶松林(一次干扰林、二次干扰林)之间活立木蓄积、粗木质残体(CWD)蓄积和组成的差异.结果表明：天然林、一次干扰林和二次干扰林的活立木蓄积量分别为161.6、138.3和114.8 m³·hm^-2,粗木质残体的蓄积量分别为69.77、36.64和32.61 m³·hm^-2.天然林粗木质残体大部分径级在20～40 cm,其中倒木、枯立木分别占总材积的72%和28%;一次干扰林和二次干扰林粗木质残体大部分径级在10～30 cm,其中倒木、枯立木和伐桩分别占各自总材积的70%、14%、16%和57%、15%、28%.人为干扰造成兴安落叶松林粗木质残体蓄积减少,改变了粗木质残体的组成.     

人为干扰对大兴安岭北坡兴安落叶松林粗木质残体的影响

比较了兴安落叶松天然林和两种不同干扰类型兴安落叶松林(一次干扰林、二次干扰林)之间活立木蓄积、粗木质残体(CWD)蓄积和组成的差异.结果表明：天然林、一次干扰林和二次干扰林的活立木蓄积量分别为161.6、138.3和114.8 m³·hm^-2,粗木质残体的蓄积量分别为69.77、36.64和32.61 m³·hm^-2.天然林粗木质残体大部分径级在20～40 cm,其中倒木、枯立木分别占总材积的72%和28%;一次干扰林和二次干扰林粗木质残体大部分径级在10～30 cm,其中倒木、枯立木和伐桩分别占各自总材积的70%、14%、16%和57%、15%、28%.人为干扰造成兴安落叶松林粗木质残体蓄积减少,改变了粗木质残体的组成.     

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。     

Sampling coarse woody debris for multiple attributes in extensive resource inventories

Information on the amount, distribution, and characteristics of coarse woody debris (CWD) in forest ecosystems is in high demand by wildlife biologists, fire specialists, and ecologists. In its important role in wildlife habitat, fuel loading, forest productivity, and carbon sequestration, CWD is an indicator of forest health. Because of this, the USDA Forest Service Pacific Northwest Research Station’s Forest Inventory and Analysis (FIA) program recognized the need to collect data on CWD in their extensive resource inventories. This paper describes a sampling method, measurement protocols, and estimation procedures to collect and compile data on CWD attributes within FIA’s forest inventory. The line-intersect method was used to sample CWD inside the boundaries of the standard inventory field plot. Previously published equations were customized to allow for easy calculation of per-unit-area values, such as biomass and carbon per hectare, log density per hectare, or volume per hectare, for each plot. These estimates are associated with all other information recorded or calculated for an inventory plot. This allows for indepth analysis of CWD data in relation to stand level characteristics. The data on CWD can be used to address current, relevant issues such as criteria no. 5 outlined in the 1994 Montreal process and the 1995 Santiago declaration. This criteria assesses the contribution of forests to the global carbon cycle by measuring such indicators as CWD, live plant biomass, and soil carbon.     

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.     

Common Factors Drive Disease and Coarse Woody Debris Dynamics in Forests Impacted by Sudden Oak Death

Disease ecology has made important steps in describing how epidemiological processes control the impact of pathogens on populations and communities but fewer field or theoretical studies address disease effects at the ecosystem level. We demonstrate that the same epidemiological mechanisms drive disease intensity and coarse woody debris (CWD) dynamics in natural forest ecosystems impacted by an emerging disease. Sudden oak death (causal agent, Phytophthora ramorum) has caused mortality of tanoak (Notholithocarpus densiflorus) on a spatial scale and rate comparable to other major North American forest diseases caused by invasive pathogens. In pathogen invaded stands, mean CWD masses were 22.4 Mg ha⁻¹ of standing dead tanoak (snags) and 11.5 Mg ha⁻¹ in logs compared to 0.27 and 1.16 Mg ha⁻¹ of snags and logs in an uninvaded stand. Within invaded stands variation in CWD mass and accumulation rates were largely driven by the distribution of pre-disease tanoak biomass and the densities of infected tanoak and California bay laurel (Umbellularia californica) which jointly determine P. ramorum sporulation and disease emergence rates. In a narrow range of community and host characteristics sudden oak death can result in woody debris dynamics similar to discrete disturbances such as fire and forest harvest but it is more common to have lower maximum amounts with slower rates of accumulation than these better studied disturbances. Our results indicate that models of CWD dynamics need to integrate epidemiological processes to predict realistic ecosystem impacts and lead to management applications for forest pathogens.     

Natural and anthropogenic influences on coarse woody debris stocks in Nothofagus–Araucaria forests of northern Patagonia,Argentina

Coarse woody debris (CWD) is an important element driving ecological processes, strengthening ecosystem resilience and for biodiversity within forest ecosystems. However, the abundance and distribution of CWD and their relation to natural and human factors are poorly known in southern South America. In this work we studied the density and volume of CWD types in Nothofagus–Araucaria stands in northern Patagonia (Neuquén – Argentina) and relationships with forest composition and structure. We also studied their relationships with fire history, topography and human‐related variables. Twenty‐three stands with Nothofagus pumilio, Nothofagus antarctica and/or Araucaria araucana were sampled to estimate quantities of logs, snags and dead branches using the planar‐intersect method. CWD density and volume in these forests were moderate and varied across the landscape with a spatial pattern determined by biotic, abiotic and human use–related variables. Mean CWD volume was 52.9 m³ ha^?1 (range: 1.6–143.7) and significantly varied among forest types and watersheds. CWD was positively related to dbh, tree height and slope, but negatively related to tree density. CWD was clearly influenced by composition and structural characteristics of stands, where the tree species traits had an important role. As well, the observed amount and type of CWD, whereby most of the stands showed low levels of old (pre‐disturbance) logs/snags and poor new inputs of deadwood, may be explained by fire frequency. Firewood gathering and livestock grazing negatively affected deadwood stocks and topography counteracts this effect by limiting human access. Fire disturbance history, windthrow and dieback pulses produced by insect outbreaks and human access seemed to be the main causes that best explained CWD spatial distribution and abundance patterns in north‐western Patagonian forests.     

Coarse Woody Debris following Fire and Logging in Wyoming Lodgepole Pine Forests

The accumulation and decomposition of coarse woody debris (CWD) are processes that affect habitat, soil structure and organic matter inputs, and energy and nutrient flows in forest ecosystems. Natural disturbances such as fires typically produce large quantities of CWD as trees fall and break, whereas human disturbances such as timber harvesting remove much of the CWD. Our objective was to compare the amount of CWD removed and left behind after clear-cutting to the amount consumed and left behind after natural fires in Rocky Mountain lodgepole pine. The masses of fallen logs, dead-standing trees, stumps, and root crowns more than 7.5 cm in diameter were estimated in clear-cut and intact lodgepole pine forests in Wyoming and compared to estimates made in burned and unburned stands in Yellowstone National Park (YNP), where no timber harvesting has occurred. Estimates of downed CWD consumed or converted to charcoal during an intense crown fire were also made in YNP. No significant differences in biomass of downed CWD more than 7.5 cm in diameter were detected between burned stands and those following a single clear-cut. However, the total mass of downed CWD plus the mass of snags that will become CWD was nearly twice as high in burned stands than in clear-cuts. In YNP, approximately 8% of the downed CWD was consumed by fire and an additional 8% was converted to charcoal, for an estimated loss of about 16%. In contrast, approximately four times more wood (70%) was removed by clear-cutting. Considering all CWD more than 7.5 cm in diameter that was either still present in the stand or removed by harvesting, slash treatment, or burning, clear-cut stands lost an average of 80 Mg ha⁻¹ whereas stands that burned gained an average of 95 Mg ha⁻¹. Some CWD remains as slash and stumps left behind after harvesting, but stands subjected to repeated harvesting will have forest floor and surface soil characteristics that are beyond the historic range of variability of naturally developing stands. Received 16 November 1999; Accepted 31 May 2000.     

Availability of coarse woody debris in a boreal old‐growth Picea abies forest

Abstract. This study reports temporal (based on cross‐dated dead trees) and spatial patterns of availability of coarse woody debris (CWD) from Picea abies in a Swedish boreal landscape with discrete old‐growth forest patches in a wetland matrix. Data were collected from 29 patches ranging in size from 0.3 to 28 ha. A total of 897 dead trees with a minimum diameter of > 15 cm occurred on the 7.2 ha area analysed. The year of death was established for 50% of these trees. CWD volume ranged from 17 to 65 m³/ha for downed logs and from 0.5 to 13 m³/ha for standing snags. CWD of all decay stages and diameter classes occurred abundantly and the probability of finding logs of all decay stages and sizes was very high at the scale of single hectares. Tree mortality differed among 5 yr periods. However, during the last 50 yr no 5 yr period produced less than 3 logs/ha. Decay rates were highly variable among different logs. Logs with soft wood and some wood pieces lost (decay stage 5) died ca. 34 years ago. This suggests a fairly rapid decay in this northern forest. The data indicate a high and continuous availability of CWD of all types. It is likely, therefore, that selection pressures for efficient dispersal among CWD dependent species may not be very high. Consequently, species with narrow habitat demands and/or low dispersal ability may have evolved and this may contribute to the decrease of certain species in the managed landscape.     

呼中林区与呼中自然保护区森林粗木质残体储量的比较

采用样线法研究了人为干扰对大兴安岭呼中林区粗木质残体(Coarse woody debris, CWD)储量的影响。结果表明, 未受人为干扰的呼中自然保护区中CWD变化范围集中在5.92~43.53 m³·hm^-2, 受到人为干扰的呼中林区中CWD变化范围集中在12.70~47.59 m³·hm^-2之间。人为干扰对混交林和针叶林的CWD储量影响显著, 而对阔叶林CWD储量影响不显著。人为干扰增加了混交林CWD储量, 减少了针叶林CWD储量, 同时减少了枯立木的数量, 增加了高度腐烂的CWD的比例。对兴安落叶松各林型的分析表明, 人为干扰条件下CWD储量(8.20~25.60 m³·hm^-2)是未受干扰条件下CWD储量(18.70~36.99 m³·hm^-2)的1/4~1/3。为促使采伐后的森林能维持原始林粗木质残体的结构特征, 建议保留采伐剩余物以维持一定储量的CWD。     

胜山阔叶红松林粗木质残体空间分布格局

为探究胜山阔叶红松(Pinus koraiensis)林粗木质残体(Coarse woody debris, CWD)的形成过程及机制,采用点格局分析法对10.4 hm²大型样地内不同分类的CWD 的空间格局、空间关联性进行了研究。结果表明:(1) 总CWD 在0-7 m尺度聚集分布,7-50 m尺度随机分布。(2) 腐烂等级5级CWD 、倒木、伐根在所有尺度随机分布;其他CWD 随尺度增大由聚集分布转为随机分布,但格局尺度不同(在2-10 m之间)。(3) 大、中径级与小径级CWD 在0-4 m、0-6 m尺度正相关;高腐烂等级与低腐烂等级CWD 在0-5 m尺度以内正相关;枯立木与倒木在0-4 m尺度正相关;落叶松(Larix gmelinii)CWD与小径级、腐烂等级2级的红皮云杉(Picea koraiensis)CWD 在0-3 m尺度正相关,与中径级、腐烂等级1、2级在13-16 m尺度负相关。胜山阔叶红松林CWD 的形成是由树种特性、小尺度的个体竞争、大尺度的自然衰老、外界干扰、生境异质性等共同决定的;大、中径级对小径级CWD、先形成对后形成CWD、枯立木对倒木具有一定的正向影响,落叶松对红皮云杉CWD 在小尺度有正向影响,而在稍大尺度有负面影响;在一定程度上揭示了该林型CWD 的形成过程和机制。     

Preliminary Studies on the Litterfall and Coarse Woody Debris in Mid-Mountain Humid Evergreen Broad-Leaved Forest in Ailao Mountains

The composition and amount of the litterfall and coarse woody debris (CWD), their time and space distribution and the element return were studied in mid-mountain humid ev-ergreen broad-leaved forest in Ailao Mountain in Yunnan, China. The average litterfall (leaves, wood (< 2.5 cm diameter), flower, fruit and mixed matter) was (6.77± 1.43) t/(hm2 · a) while the large wood litterfall () 2.5 cm diameter) was (0.45±0.18) t/(hm2 · a). There were two marked peaks of annual litterfalh the main one from April to May and the second one from October to November. The standing crop of CWD in the forest was 98.46 t/hm2, in which logs made up to 85.8% of the total. As the CWD was not well-distributed in the forest, it increased the diversities of structure and site in the forest ecosystem. The total storage of C, N, P, K, Ca, Mg, Mn, A1 and Fe in the CWD was 48699 t/hm2, but return of elements to the forest ground in the litterfall only made up 7.77% of the nutrient storage of the CWD.     

Diameter of downed woody debris does matter for saproxylic beetle assemblages in temperate oak and pine forests

Deadwood is widely recognized to be an important issue for biodiversity conservation in forest ecosystems. Establishing guidelines for its management requires a better understanding of relationships between woody debris characteristics and associated species assemblages. Although deadwood diameter has been identified as an important factor predicting occurrence of many saproxylic species, the boundary between small and large diameter has not yet been precisely defined. In commercial forests, it is also of critical importance to know which diameter is large enough to host the beetle species associated with large logs in order to ensure cost-effectiveness of biodiversity conservation measures. We investigated the differences in saproxylic beetle assemblages among four different diameter classes of downed woody oak and maritime pine debris, in France. Beetles were sampled using in situ emergence traps. The diameter of deadwood pieces ranged from 1 to 40 cm. No patterns of nestedness associated with the gradient of diameter size were identified for either tree species. More indicator saproxylic species were observed in large logs and branches than in small logs. A clear distinction appeared in assemblage composition around the 5-cm diameter threshold whereas no similar pattern occurred around the 10 cm value, i.e. the classical threshold used in forestry to distinguish fine woody debris from coarse woody debris. For both tree species, the mean body length of beetles increased with the diameter of deadwood suggesting that the quantity of available resources per piece may constitute a limiting factor for large beetle species. This study confirms that not only large deadwood pieces are relevant for saproxylic biodiversity conservation but also the smallest pieces. Therefore, forest managers would be well advised to maintain a high diversity of deadwoods to maintain saproxylic biodiversity.     

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.     

The longevity of constructed log pile fauna habitats in restored bauxite mines in relation to recurrent wildfire in the jarrah forest of Western Australia

Coarse woody debris (CWD) is often returned to rehabilitated areas after mining to provide specific habitat features for fauna that might otherwise take many decades to develop naturally. The CWD, typically constructed into log piles, is assumed to persist for comparable periods of time to fulfil their role. In the fire‐prone jarrah forest in Western Australia, these structures will be subject to recurrent wildfire, but there is little information on their survival. In January 2006, a wildfire burnt through 310 ha of rehabilitation at Alcoa’s Willowdale bauxite mine, 140 km south east of Perth, enabling the first estimates of the longevity of log pile fauna habitats to be made. Reductions in numbers of logs were greatest for small (<30 cm diameter) logs (48%), intermediate for large (>30 cm diameter) logs (37%) and least for stumps (17%). Assuming an exponential decay pattern, half‐lives of 1.1, 1.5 and 3.7 wildfires were calculated for small logs, large logs and stumps respectively, from which we estimate a useful ‘life’ of a typical log pile fauna habitat of 57–73 years. Increasing fire intensity was associated with greater reductions in the number of large logs, significantly greater surface charring and, at the highest intensity, significantly fewer crevices compared to unburnt log piles. Implications of these findings for the management of log piles in rehabilitation and CWD in the adjacent unmined forest, and of the role of prescribed fire, are discussed.     

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.