The ectomycorrhizal flora of primary and secondary stands of Pinus sylvestris in relation to soil conditions and ectomycorrhizal succession

Abstract. Ectomycorrhizal species composition and sporocarp abundance in two 15–20-yr old primary stands of Pinus sylvestris in the central part of the Netherlands was compared with those in two 16- and 27-yr old secondary stands of P. sylvestris in the Northeast of the country. The trees of the primary stands were spontaneously seeded in a drifting sand area. Only thin litter and humus layers were present. The trees of the 16-yr old secondary stand were planted on podzolic sandy soil and those of the 27-yr old stand on non-podzolic sandy soil. In both secondary stands thick litter and humus layers had developed. The litter and humus partly originated from former stands at the same sites and partly from the present stands. In the secondary stands the thick litter and humus layers and herb vegetation were removed (‘sod-cutting’) in order to simulate the thin litter and humus layers in the primary stands. Control treatments were present. Surveys in 1991, 1992 and 1993 showed that sod-cutting enhanced both abundance and diversity of ectomycorrhizal fungi. However, species richness and diversity were higher in the primary stands than in the secondary ones, also even in the sod-cut plots. High species richness and diversity were associated with low concentrations of nitrogen and relatively high pH in the litter and humus layers, and in the mineral soil.     

云南松(Pinus yunnanensis)林外生菌根真菌的时空分布

2000年至2005年,调查了滇中及其附近云南松林下外生菌根真菌的生态分布,共采集、鉴定标本834号,计有27科39属211种（含变种、变型）。结果表明,红菇属（Russula）、牛肝菌属（Boletus）、乳菇属（Lactarius）、乳牛肝菌属（Suillus）、口蘑属（Tricholoma）、鸡油菌属（Cantharellus）和革菌属（Thelephora）等为云南松林下的主要外生菌根菌类群。它们的发生与分布受到气候（如：气温和降水）、植被（如：林龄、林地郁闭度和草本植被）、地形特征（如：海拔、坡向和坡度）、土壤条件（如：pH值、地表腐殖质和枯枝落叶层等）和人为干扰（比如：商业化采集、林木采伐、火烧和地表物清理）诸多因素的影响。总结为如下：（1）5a的调查结果显示,云南松外生菌根菌的分布表现出季节性变化的规律;其中以每年1、2、3月份的物种多样性为最低,雨季期间急剧增加,至中夏和秋末达到顶峰,种类最为繁多。（2）在海拔1500-2100m,云南松外生菌根菌种类随着海拔的升高而逐渐增加,至顶峰后,又呈缓慢下降趋势。海拔因素不但对其物种多样性,而且对于类群的组成也具有重要的影响。特定的类群往往发生在特定海拔范围。（3）随着云南松林龄的增加,外生菌根菌呈现由少至多的演替过程。外生菌根菌多样性随云南松林生长而逐渐增加的演替方式,可能与宿主光合作用产物、根部分泌物和土壤条件的逐渐变化有关。（4）人类干扰是云南松外生菌根菌物种多样性和类群组成的主要负影响因子。大规模的商业化采集可破坏或枯竭地下菌丝体,打破各物种之间的竞争平衡,减少孢子释放影响资源再生能力,进而直接影响到子实体的产生。外生菌根菌物种多样性的减少趋势会随林木砍伐和火烧强度的增加而加剧。地表枯枝落叶层与杂草密度也会影响子实体的产生,其中枯枝落叶层的厚度与云南松外生菌根菌子实体的发生呈负相关性,而被紫茎泽兰覆盖的云南松林地内也很少会发现相应的子实体。     

Effects of manipulation of litter and humus layers on ectomycorrhizal colonization potential in Scots pine stands of different age

Effects of manipulation of litter and humus layers (removal, doubling and control treatments) on the colonization potential of ectomycorrhizal fungi were studied in two secondary stands of Pinus sylvestris (5 and 18 years old) in The Netherlands. Five-mont-hold, sterile-grown Scots pine seedlings, inoculated with Laccaria bicolor, Paxillus involutus or Rhizopogon luteolus and noninoculated seedlings were used as baits. The seedlings were harvested after one growing season. For comparison, sporocarps of ectomycorrhizal fungi were also investigated. Genus composition on the seedlings was independent of initial inoculum, but determined by both treatment and age of the stands. In both stands, removal of litter and humus layers increased, and addition of organic material decreased the number of ectomycorrhizal types on the seedlings. Not all indigenous genera were observed by either outplanting seedlings or sporocarp surveys.     

云南松（Pinus yunnanensis）林外生菌根真菌的时空分布

2000年至2005年,调查了滇中及其附近云南松林下外生菌根真菌的生态分布,共采集、鉴定标本834号,计有27科39属211种（含变种、变型）。结果表明,红菇属（Russula）、牛肝菌属（Boletus）、乳菇属（Lactarius）、乳牛肝菌属（Suillus）、口蘑属（Tricholoma）、鸡油菌属（Cantharellus）和革菌属（Thelephora）等为云南松林下的主要外生菌根菌类群。它们的发生与分布受到气候（如：气温和降水）、植被（如：林龄、林地郁闭度和草本植被）、地形特征（如：海拔、坡向和坡度）、土壤条件（如：pH值、地表腐殖质和枯枝落叶层等）和人为干扰（比如：商业化采集、林木采伐、火烧和地表物清理）诸多因素的影响。总结为如下：(1）5a的调查结果显示,云南松外生菌根菌的分布表现出季节性变化的规律;其中以每年1、2、3月份的物种多样性为最低,雨季期间急剧增加,至中夏和秋末达到顶峰,种类最为繁多。(2）在海拔1500～2100 m,云南松外生菌根菌种类随着海拔的升高而逐渐增加,至顶峰后,又呈缓慢下降趋势。海拔因素不但对其物种多样性,而且对于类群的组成也具有重要的影响。特定的类群往往发生在特定海拔范围。(3）随着云南松林龄的增加,外生菌根菌呈现由少至多的演替过程。外生菌根菌多样性随云南松林生长而逐渐增加的演替方式,可能与宿主光合作用产物、根部分泌物和土壤条件的逐渐变化有关。(4）人类干扰是云南松外生菌根菌物种多样性和类群组成的主要负影响因子。大规模的商业化采集可破坏或枯竭地下菌丝体,打破各物种之间的竞争平衡,减少孢子释放影响资源再生能力,进而直接影响到子实体的产生。外生菌根菌物种多样性的减少趋势会随林木砍伐和火烧强度的增加而加剧。地表枯枝落叶层与杂草密度也会影响子实体的产生,其中枯枝落叶层的厚度与云南松外生菌根菌子实体的发生呈负相关性,而被紫茎泽兰覆盖的云南松林地内也很少会发现相应的子实体。     

Partial restoration of fungal and plant species diversity by removal of litter and humus layers in stands of Scots pine in the Netherlands

A field experiment was set up in five mature Scots pine (Pinus sylvestris) stands in the Netherlands in order to study the effects of removal of litter and humus on ectomycorrhizal fungi and vegetation of higher plants, bryophytes, and lichens. Lime was also applied as a countermeasure to the removal of surplus cations. Application of lime did not have significant effects on mycorrhizal fungi. Removal of ectorganic material increased species richness of ectomycorrhizal fungi. The number of fruit bodies was also higher in the sod-cut plots, but the difference compared with the controls was not significant in all years. A major effect of removal of the ectorganic layer was a reversion of ectomycorrhizal succession to earlier stages. Liming increased the number and cover of ruderal, nitrophilous plant species. Sod-cutting caused a reversion of the green vegetation to earlier successional stages.     

抚育间伐对油松人工林下大型真菌的影响

真菌在森林凋落物分解过程中起重要作用,研究间伐如何影响真菌进而影响凋落物分解,对深入了解间伐调控人工林凋落物分解有重要意义。本文以抚育间伐后的中龄油松人工林为研究对象,设立对照（I）、轻度（II）、中度（III）和强度（IV）4种间伐强度,于2011年对间伐后林下大型真菌进行两次调查,分析了不同间伐强度下大型真菌的科的分布、优势种组成和生态指标（包括丰富度指数、多样性指数和均匀度指数）。结果表明：（1）8月、9月采集到的大型真菌分别为35种和25种,分属13个科和10个科;（2）在大型真菌出菇期,间伐改变了大型真菌的优势种组成,对照林下大型真菌优势种最初为外生菌根菌（粘盖乳牛肝菌 血红铆钉菇）后变为腐生菌（大盖小皮伞和脐顶小皮伞）,而间伐后林下优势种始终为腐生菌;（3）间伐影响大型真菌的生态指标,中度间伐林下大型真菌丰富度和多样性指数最高。总之,适度间伐不仅有利于提高林下大型真菌的丰富度与多样性,同时使其群落结构发生改变,群落优势种由外生菌根菌变为以分解凋落物为主的腐生菌,可促进凋落物的分解和养分循环。     

Ectomycorrhizal and wood-decay macromycete communities along development stages of managed Pinus patula stands in Southwest Mexico

This study used fruit body surveys to evaluate the diversity and distribution of ectomycorrhizal and wood-decay macromycetes along developing stages of stands artificially reforested with Pinus patula in Southern Mexico. Three stand ages were selected: 1, 11, and 60 y old. Macromycetes were collected from June to October 2016 in 0.1 ha plots per stand age. A total of 106 taxa were recorded, 63 ectomycorrhizal and 43 wood-decay fungi. The highest diversity of ectomycorrhizal fungi was in 11-y stands, and 11- and 60-y stands had similar species composition. Wood-decay fungi showed a trend towards higher diversity in the 1- and 11-y stands, and species composition in these sites corresponded. Patterns of diversity and distribution were mainly determined by factors related to substratum availability and vegetation structure. Knowledge about the influence of artificially reforested stands on macrofungal communities is relevant to improve management plans and achieve a more sustainable use of forest timber and non-timber products.     

15N natural abundance in fruit bodies of different functional groups of fungi in relation to substrate utilization

Natural abundances of ¹⁵N and N concentrations of 34 fruit bodies from 24 species of ectomycorrhizal and saprophytic fungi were measured in a temperate Central European mixed forest stand. The fungi of the two life forms are known to be capable of utilizing different types of N sources (organic N compounds from the humus, inorganic N from the soil and N from litter or wood) differing by their ¹⁵N natural abundance values. Based on the two life forms and the three different N sources, four functional groups of fungi were distinguished: (1) ectomycorrhizal fungi capable of utilizing organic N from the humus; (2) ectomycorrhizal fungi known to depend on inorganic N compounds in the soil; (3) saprophytes capable of utilizing organic N from the humus; and (4) saprophytes utilizing N from dead wood or litter. Large differences were found between species in the δ¹⁵N values (−3.0 to 3.3‰) and in the N concentrations (0.84 to 6.61 mmol eq N g dw⁻¹) of the fruit bodies. In most cases fungi were more enriched in ¹⁵N than their respective bulk N source was. Fungi living in humus, and presumably having access to organic N compounds (groups 1 and 3), were significantly more enriched in ¹⁵N than fungi which are known to depend on inorganic N (e.g. Laccaria , group 2), or fungi living on litter or wood (group 4), irrespective of whether they were ectomycorrhizal or saprophytic species. Fungi living in humus had significantly higher N concentrations than fungi living on litter or wood.     

Ectomycorrhizal fungal communities in late-successional Swedish boreal forests, and their composition following wildfire

This study was conducted to evaluate the effects of wildfires on ectomycorrhizal (EM) fungal communities in Scots pine ( Pinus sylvestris ) stands. Below- and above-ground communities were analysed in terms of species richness and evenness by examining mycorrhizas and sporocarps in a chronosequence of burned stands in comparison with adjacent unburned late-successional stands. The internal transcribed spacer (ITS)-region (rDNA) of mycobionts from single mycorrhizas was digested with three restriction enzymes and compared with an ITS–restriction fragment length polymorphism (RFLP) reference database of EM sporocarps. Spatial variation seemed to be more prominent than the effects of fire on the EM fungal species composition. Most of the common species tended to be found in all sites, suggesting that EM fungal communities show a high degree of continuity following low-intensity wildfires. Species richness was not affected by fire, whereas the evenness of species distributions of mycorrhizas was lower in the burned stands. The diversity of EM fungi was relatively high considering that there were only three EM tree species present in the stands. In total, 135 EM taxa were identified on the basis of their RFLP patterns; 66 species were recorded as sporocarps, but only 11 of these were also recorded as mycorrhizas. The species composition of the below-ground community of EM fungi did not reflect that of the sporocarps produced. EM fungal species present in our ITS–RFLP reference database accounted for 54–99% of the total sporocarp production in the stands, but only 0–32% of the mycorrhizal abundance.     

毛乌素沙地樟子松外生菌根真菌多样性特征

【背景】外生菌根真菌(ectomycorrhizal fungi,ECMF)侵入林木根系形成外生菌根,是樟子松(Pinus sylvestris var. mongholica)适应环境变化和环境胁迫的重要途径。【目的】深入理解沙地樟子松人工林外生菌根真菌群落组成和多样性。【方法】以毛乌素沙地3个林龄(27、33和44 a)沙地樟子松人工林为研究对象,采用rDNA ITS区段PCR扩增测序方法鉴定分析沙地樟子松外生菌根真菌群落组成并计算多样性指数。【结果】毛乌素沙地樟子松共获得56个外生菌根真菌OTU,隶属于2门3纲8目15科21属,优势属为棉革菌属(Tomentella)、地孔菌属(Geopora)和阿太菌属(Amphinema)。27 a人工林外生菌根真菌丰富度指数最高,33 a人工林ECMF的α多样性指数最高,但不同林龄之间不存在显著差异(P>0.05)。不同林龄沙地樟子松ECMF群落组成存在差异,地孔菌属、棉革菌属、Delastria和Mallocybe的动态变化是造成群落组成差异的主要原因。【结论】毛乌素沙地樟子松人工林外生菌根真菌群落组成较为丰富,该研究区的群落组成可能比多样性更能体现外生菌根真菌群落随林龄的动态变化,伴随着林木的生长成熟,部分功能性外生菌根真菌的动态变化可能与沙地樟子松人工林退化存在密切联系,有待进一步研究。     

Ectomycorrhizal root growth in scots pine stands in response to manipulation of litter and humus layers

 The effect on ectomycorrhizal root growth in a nitrogen-enriched planted stand of Scots pine (Pinus sylvestris L.) on podzolic sandy soil to manipulation of litter and humus layers (removal, doubling and control treatments) was examined, and compared to ectomycorrhizal root growth in an untreated naturally established Scots pine stand on nutrient-poor non-podzolic sandy soil. Half a year after manipulation of litter and humus layers in the planted stand, ingrowth-cores to a depth of 60 cm were installed in both stands. Scots pine roots were sampled four times during two growing seasons. Ectomycorrhizal roots were found at all sampled soil depths to 60 cm in all plots. Root growth and ectomycorrhizal development were greater in the naturally established stand than in all plots in the planted stand. Numbers of ectomycorrhizal root tips in the litter and humus removal plots were generally higher than in the control plots in the planted stand until May 1992. Doubling litter and humus did not significantly affect root length or the numbers of ectomycorrhizal root tips. The N _dissolved , NH₄ ⁺ and NO₃ ^– concentrations and the organic matter content in the upper 5 cm of the mineral soil in the planted stand on podzolic sandy soil were generally higher and the pH significantly lower than in the naturally established stand on non-podzolic sandy soil. Root growth and ectomycorrhizal development in the secondary stand may have been negatively affected by the chemical composition of the podzolic sandy soil. Accepted: 19 March 1997     

Spatial structure and diversity of woody plants and ectomycorrhizal fungus sporocarps in a natural subtropical forest

Spatial patterns of ectomycorrhizal fungi, ectomycorrhizal plants, and non-ectomycorrhizal plants were investigated in a natural subtropical forest using second-order analysis. The results of spatial pattern analysis showed that the degree of clumping of woody plants and ectomycorrhizal sporocarps were correlated. There was a significantly positive correlation of relative aggregation indices between ectomycorrhizal fungi and both non-ectomycorrhizal trees and ectomycorrhizal saplings. Correlations between percentage of ectomycorrhizal trees and sporocarp occurrence of ectomycorrhizal fungi and between diversities of woody plants and ectomycorrhizal fungi were distance-dependent or scale-related. A significantly high percentage of ectomycorrhizal trees was found only at relatively short distance from ectomycorrhizal fungal sporocarps, and significantly positive correlation of the diversity between woody plants and ectomycorrhizal fungi was found only at relative long distance, which implied that ectomycorrhizal sporocarps prefer ectomycorrhizal-tree-dominant micro-sites at near distances and at relatively large scales, diverse ectomycorrhizal sporocarps could be found in woodlands with high diversity of woody plants. Important factors affecting the spatial distribution, occurrence, and diversity of ectomycorrhizal fungi include spatial pattern of ectomycorrhizal plants and non-ectomycorrhizal plants, percentage of ectomycorrhizal plants, and plant diversity in a natural forest.     

外生菌根对干旱胁迫的响应

从外生菌根真菌、外生菌根共生体以及外生菌根的间接作用等方面阐述外生菌根如何抵制干旱胁迫,并对未来我国外生菌根的研究提出了建议。干旱可以抑制外生菌根真菌的生长并降低其群落中真菌的多样性,干旱胁迫下外生菌根真菌子实体可以利用深度30cm以下的土壤水,子实体的表面积和体积比可作为筛选抗旱真菌的一个重要因子;在遭受干旱胁迫时,外生菌根共生体可以发生形态变化来应对干旱,同时增加了植株水分的吸收并改善了植物的光合作用、活性氧以及激素等相关代谢;外生菌根对植物生长的促进作用、增加土壤碳汇以及对其他根际微生物生长的促进作用等对宿主植物应对干旱胁迫有利。未来我国外生菌根研究应加强对干旱区优良菌-树组合的筛选工作,同时加大对乡土外生菌根真菌资源的调查力度,未来研究应重点向分子生物学领域推进。     

川中丘陵区柏木低效林改造模式植物多样性对土壤有机碳的影响

以川中丘陵区柏木低效林改造10a后的4种模式:纯杂交竹模式(Bambusa pervariabilis×Dendrocalamopsis daii)(CZ)、柏木(Cupressus funebris Endl.)+桤木(Alnus cremastogyne Burk.)+杂交竹模式(BZQ)、柏木+麻栎(Quercus acutissima Carruth)模式(BL)、柏木+杂交竹模式(BZ)为研究对象,纯柏(CB)为对照,对土壤有机碳含量、碳密度和土壤活性有机碳(土壤易氧化碳、水溶性碳和微生物量碳)含量及植物多样性进行了研究。结果表明:(1)随着土层深度的增加,土壤有机碳含量、碳密度及其组分均下降,其中土壤有机碳含量下降幅度最大,易氧化碳含量下降幅度最小;0—10 cm土壤有机碳密度占整个取样剖面的45.7%—64.4%,具有明显的表聚性。(2)不同模式0—40 cm土层土壤有机碳含量、碳密度和土壤易氧化碳、水溶性碳和微生物量碳含量均为BZQBZCBBLCZ,差异均达到显著水平(P0.05)。几种模式中,BZQ与CB相比,土壤有机碳含量、碳密度、土壤易氧化碳、水溶性碳和微生物量碳含量增幅分别为37.8%、33.5%、36.6%、52.5%和23.2%,表明BZQ模式在提高土壤有机碳方面作用最明显。(3)植物Simpson优势度指数、Shannon-Wiener多样性指数、Pielou均匀度指数和丰富度指数大致表现为BZQBZCBBLCZ;相关分析表明,土壤有机碳、碳密度、易氧化碳、水溶性碳和微生物量碳与灌木层和草本层的植物多样性指数均呈显著正相关,而与乔木层植物多样性不相关。     

Difference of pine ectomycorrhizal biomass in relation to forest conditions

In this study, two plots in a secondary and another two in planted Pinus densiflora stands were exposed to different forest treatments, and the ectomycorrhizal (EM) biomass and its ergosterol content was measured for a year. The unmanaged plot in the secondary stand had greater EM biomass than those in any other plots. Whereas understory cutting had less effect on EM biomass, litter and humus removal decreased pine EM biomass and its ergosterol content, suggesting that such forest treatment alters EM biomass and its structure.     

Epigean communities of harvestmen (Opiliones) in Pannonian Basin floodplain forests: an interaction with environmental parameters

During the years 2015–2016, we studied the fauna of epigeon in eight similar habitats in Slovakia and Serbia. The study material of Opiliones was obtained by a pitfall trap method. We focused on the comparison of the incidence of harvestmen in floodplain forests and ecotones alongside the Danube River in Slovakia and along the Tisa and Begej River in Serbia. We examined their dependence on anthropogenic impact (frequency of visits), cover of vegetation layers (herbs, shrubs and trees), plant diversity of vegetation layers, area of forest stands, distance to the forest edge, circumference of forest stands, age of forest stands, thickness of litter layer and physico-chemical properties of soil and litter (pH, conductivity, P, N, C, H). Total epigeic activity of harvestman showed significant positive relationship with proportion of N in litter. Community equitability was positively related with species richness of plant communities in shrub layer and negatively related with H content of soil. We did not find any significant correlations between diversity of harvestman communities and measured environmental characteristics.     

辽宁章古台樟子松人工林的生物量与营养元素分布的初步研究

樟子松(Pinus sylvestris var.mongolica)人工林是章古台沙地生物生产量较高的林分,其生长主要受沙丘部位和造林类型所影响。樟子松林乔木层与凋落物中贮存了大量营养元素,土壤上层的有机质与N、P元素含量较高,下层含量较低,出现一定的降低趋势。樟子松林混以小青杨可提高生物产量,增加土壤腐殖质,且改善土壤腐殖酸组成成分。     

Dynamics of vesicular-arbuscular mycorrhizae during old field succession

Summary The species composition of vesicular-arbuscular mycorrhizal (VAM) fungal communities changed during secondary succession of abandoned fields based on a field to forest chronosequence. Twenty-five VAM fungal species were identified. Seven species were clearly early successional and five species were clearly late successional. The total number of VAM fungal species did not increase with successional time, but diversity as measured by the Shannon-Wiener index tended to increase, primarily because the community became more even as a single species, Glomus aggregatum, became less dominant in the older sites. Diversity of the VAM fungal community was positively correlated with soil C and N. The density of VAM fungi, as measured by infectivity and total spore count, first increased with time since abandonment and then decreased in the late successional forest sites. Within 12 abandoned fields, VAM fungal density increased with increasing soil pH, H₂O soluble soil C, and root biomass, but was inversely related to extractable soil P and percent cover of non-host plant species. The lower abundance of VAM fungi in the forest sites compared with the field sites agrees with the findings of other workers and corresponds with a shift in the dominant vegetation from herbaceous VAM hosts to woody ectomycorrhizal hosts.     

Strategies of carbon and nitrogen acquisition by saprotrophic and ectomycorrhizal fungi in Finnish boreal Picea abies-dominated forests

We compared the δ¹³C and δ¹⁵N of forest material with an extensive sporocarp collection to elucidate the role of litter, wood and soil as fungal carbon and nitrogen sources in Finnish boreal Picea abies-dominated forests. Ectomycorrhizal Hydnum and Cortinarius had higher δ¹⁵N than other ectomycorrhizal fungi, suggesting use of ¹⁵N-enriched, deeper nitrogen. Russula had lower δ¹⁵N than other ectomycorrhizal fungi and resembled some litter decay genera, suggesting use of litter-derived nitrogen. There was little variation in δ¹⁵N among other genera of ectomycorrhizal fungi, indicating limited functional diversity in nitrogen use. Saprotrophic Leotia, Gymnopus, Hypholoma, Pholiota, Rhodocollybia and Calocera had δ¹⁵N values similar to ectomycorrhizal fungi, indicating overlap in use of older nitrogen from soil or roots or use of newly fixed nitrogen. Genera of litter and wood decay fungi varied up to 6‰ in δ¹³C and 10‰ in δ¹⁵N, suggesting large differences in carbon and nitrogen sources and processing. Similar δ¹³C between white and brown rot wood decay fungi also suggest that white rot fungi do not use lignin-derived carbon. Together, these δ¹³C and δ¹⁵N patterns of fungi from Finnish boreal forests enhance our knowledge of fungal functional diversity and indicate broad use of litter, wood and soil resources.     

Reed cut,habitat diversity and productivity in wetlands

There is a conflict between nature conservation and thatching industry regarding the management of reedbeds. On one hand, reedbeds are of an economical importance by providing thatching material, on the other hand, they harbour several endangered species. Reedbeds are typically managed by winter cutting, but its impacts on biodiversity are poorly understood. Our aim was to study the effects of winter cutting on the habitat diversity and structural heterogeneity of wetlands in a lowland alkali landscape (East-Hungary). We tested the following hypotheses: (i) Both diversity of plant species and habitat diversity are lower in winter cut wetlands compared to unmanaged stands. (ii) The distribution of biomass (green biomass, litter and standing dead biomass) is more homogeneous in winter cut wetlands compared to unmanaged ones. We found that winter cutting decreased habitat diversity and structural heterogeneity at multiple scales. Number of plant species and all measures of habitat diversity (number of patches, vegetation types and the length of vegetation margins) had lower scores in cut wetlands than in unmanaged ones. We found that unmanaged wetlands harboured high amount of accumulated biomass and they also maintained high habitat diversity likely due to the heterogeneous distribution of the biomass. In unmanaged wetlands, biomass accumulation did not decrease habitat diversity and also contributed to a higher structural heterogeneity. In cut wetlands, expansion of reed was an important driver of the decrease in habitat diversity and structural heterogeneity. Reed expansion likely overrode fine-scale edaphic conditions (hydrology and salinity) in shaping vegetation patterns; thus we suggest to avoid intensive winter cutting.