Differential response of tellimagrandin II and total bioactive hydrolysable tannins in an aquatic angiosperm to changes in light and nitrogen

Ectomycorrhizal (ECM) diversity was measured in 12 mixed-wood stands in the Abitibi region of north-western Québec. Stands were of similar age and were situated on similar mineral soil deposits, but supported varying proportions of ECM host trees. Host roots were sampled in a manner that enabled their separation into species on the basis of wood anatomy. Shannon diversity indices for the ECM colonizing each host species were determined on the basis of ECM anatomy. The diversity of overstory trees, understory plants and host roots, as well as overstory tree composition, root density and pertinent abiotic factors were measured and used as independent variables in multiple regressions against ECM diversity. We found a positive relationship between overstory tree diversity and ECM diversity, which appears related to fungal host specificity. Although no direct relationship was seen between ECM diversity and soil factors, levels of exchangeable base cations were related to ECM fungal species composition which correlated with ECM diversity at the scale sampled.     

Determination of Diversity,Distribution and Host Specificity of Korean Laccaria Using Four Approaches

The genus Laccaria (Hydnangiaceae, Agaricales) plays an important role in forest ecosystems as an ectomycorrhizal fungus, contributing to nutrient cycles through symbiosis with many types of trees. Though understanding Laccaria diversity and distribution patterns, as well as its association with host plants, is fundamental to constructing a balanced plant diversity and conducting effective forest management, previous studies have not been effective in accurately investigating, as they relied heavily on specimen collection alone. To investigate the true diversity and distribution pattern of Laccaria species and determine their host types, we used four different approaches: specimen-based analysis, open database search (ODS), NGS analysis, and species-specific PCR (SSP). As a result, 14 Laccaria species have been confirmed in Korea. Results regarding the species distribution pattern were different between specimen-based analysis and SSP. However, when both were integrated, the exact distribution pattern of each Laccaria species was determined. In addition, the SSP revealed that many Laccaria species have a wide range of host types. This study shows that using these four different approaches is useful in determining the diversity, distribution, and host of ECM fungi. Furthermore, results obtained for Laccaria will serve as a baseline to help understand the role of ECM fungi in forest management in response to climate change.     

Spatial analysis of ectomycorrhizal fungi reveals that root tip communities are structured by competitive interactions

Microbial ecology has made large advances over the last decade, mostly because of improvements in molecular analysis techniques that have enabled the detection and identification of progressively larger numbers of microbial species. However, determining the ecological patterns and processes taking place in communities of microbes remains a significant challenge. Are communities randomly assembled through dispersal and priority effects, or do species interact with each other leading to positive and negative associations? For mycorrhizal fungi, evidence is accumulating that stochastic and competitive interactions between species may both have a role in shaping community structure. Could the methodological approach, which is often incidence based, impact the outcomes detected? Here, we applied an incidence‐based Terminal Restriction Fragment Length Polymorphism (T‐RFLP) database approach to examine species diversity and ecological interactions within a community of ectomycorrhizal (ECM) fungi. Co‐occurrence analysis revealed that the ECM community colonizing root tips was strongly structured by competitive interactions, or ecological processes generating a similar spatial pattern, rather than neutral processes. Analysis of β‐diversity indicated that community structure was significantly more similar (spatially autocorrelated) at distances equal to or <3.41 m. The eight most frequently encountered species in the root tip community of ECM fungi displayed significant competitive interactions with at least one other species, showing that the incidence‐based approach was capable of detecting this sort of ecological information.     

Genetic diversity of ectomycorrhizal Basidiomycetes from African and Indian tropical rain forests

Ectomycorrhizal (ECM) fungi have a worldwide distribution. However, the ecology of tropical ECM fungi is poorly documented, limiting our understanding of the symbiotic associations between tropical plants and fungi. ECM Basidiomycete diversity was investigated for the first time in two tropical rain forests in Africa (Western Upper Guinea) and in Asia (Western Ghats, India), using a fragment of the mitochondrial large subunit rRNA gene to type 140 sporocarps and 54 ectomycorrhizas. To evaluate taxonomic diversity, phylogenetic analyses were performed, and 40 sequences included from identified European specimens were used as taxonomic benchmarks. Five clades were recovered corresponding to six taxonomic groups: boletoids, sclerodermatoids, russuloids, thelephoroids, and a clade grouping the Amanitaceae and Tricholomataceae families. Our results revealed that the Russulaceae species display a great diversity with several putative new species, especially in Guinea. Other taxonomic issues at family/section levels are also briefly discussed. This study provides preliminary insights into taxonomic diversity, ECM status, and biogeographic patterns of ECM fungi in tropical two rain forest ecosystems, which appear to be as diverse as in temperate and boreal forests.     

Fungal diversity in ectomycorrhizal communities: sampling effort and species detection

A number of recent review articles on ectomycorrhizal (ECM) fungal community diversity have highlighted the unprecedented increase in the number of publications on this ecologically important but neglected area. The general features of these species-rich, highly dynamic and complex communities have been comprehensively covered but one aspect crucial to our assessment of diversity, namely the sampling of ECM communities has received less attention. This is a complex issue with two principal components, the physical sampling strategy employed and the life cycle traits of the ECM fungi being examined. Combined, these two components provide the image that we perceive as ECM diversity. This contribution will focus primarily on the former of these components using a recent study from a pine forest in central Sweden to highlight some sampling problems and also to discuss some features common to ECM communities. The two commonly used elements of diversity, species richness and community evenness, present rather different problems in the assessment of ECM diversity. The applicability of using current measures of abundance (number or percentage of root tips colonised) to determine community evenness is discussed in relation to our lack of knowledge on the size of individual genets of ECM fungi. The inherent structure of most ECM communities, with a few common species and a large number of rare species, severely limits our ability to accurately assess species richness. A discussion of theoretical detection limits is included that demonstrates the importance of the sampling effort (no. of samples or tips) involved in assessing species richness. Species area abundance plots are also discussed in this context. It is suggested that sampling strategy (bulk samples versus multiple collections of single tips) may have important consequences when sampling from communities where root tip densities differ. Finally, the need for studies of the spatial distribution of ECM on roots in relation to small-scale soil heterogeneity and of the temporal aspects of ECM community dynamics is raised.     

Diversity and specificity of ectomycorrhizal fungi retrieved from an old-growth Mediterranean forest dominated by Quercus ilex

We analysed the ectomycorrhizal (ECM) fungal diversity in a Mediterranean old-growth Quercus ilex forest stand from Corsica (France), where Arbutus unedo was the only other ECM host. On a 6400 m2 stand, we investigated whether oak age and host species shaped below-ground ECM diversity. Ectomycorrhizas were collected under Q. ilex individuals of various ages (1 yr seedlings; 3-10 yr saplings; old trees) and A. unedo. They were typed by ITS-RFLP analysis and identified by match to RFLP patterns of fruitbodies, or by sequencing. A diversity of 140 taxa was found among 558 ectomycorrhizas, with many rare taxa. Cenococcum geophilum dominated (35% of ECMs), as well as Russulaceae, Cortinariaceae and Thelephoraceae. Fungal species richness was comparable above and below ground, but the two levels exhibited < 20% overlap in fungal species composition. Quercus ilex age did not strongly shape ECM diversity. The two ECM hosts, A. unedo and Q. ilex, tended to share few ECM species (< 15% of the ECM diversity). Implications for oak forest dynamics are discussed.     

Alpine bistort (<Emphasis Type="Italic">Bistorta vivipara</Emphasis>) in edge habitat associates with fewer but distinct ectomycorrhizal fungal species: a comparative study of three contrasting soil environments in Svalbard

Bistorta vivipara is a widespread arctic-alpine ectomycorrhizal (ECM) plant species. Recent findings suggest that fungal communities associated with B. vivipara roots appear random over short distances, but at larger scales, environmental filtering structure fungal communities. Habitats in highly stressful environments where specialist species with narrower niches may have an advantage represent unique opportunity to test the effect of environmental filtering. We utilised high-throughput amplicon sequencing to identify ECM communities associated with B. vivipara in Svalbard. We compared ECM communities in a core habitat where B. vivipara is frequent (Dryas-heath) with edge habitats representing extremes in terms of nutrient availability where B. vivipara is less frequent (bird-manured meadow and a nutrient-depleted mine tilling). Our analysis revealed that soil conditions in edge habitats favour less diverse but more distinct ECM fungal communities with functional traits adapted to local conditions. ECM richness was overall lower in both edge habitats, and the taxonomic compositions of ECM fungi were in line with our functional expectations. Stress-tolerant genera such as Laccaria and Hebeloma were abundant in nutrient-poor mine site whereas functional competitors genera such as Lactarius and Russula were dominant in the nutrient-rich bird-cliff site. Our results suggest that ECM communities in rare edge habitats are most likely not subsets of the larger pool of ECM fungi found in natural tundra, and they may represent a significant contribution to the overall diversity of ECM fungi in the Arctic.     

Similar taxonomic richness but different communities of ectomycorrhizas in native forests and non-native plantation forests

This investigation sought to examine if there was a difference between the ectomycorrhizal (ECM) communities in plots of native oak and introduced Scots pine and Sitka spruce forest. The ECM communities in four plots of each forest type were described, from five soil cores collected in each plot, by morphotyping, internal transcribed spacer (ITS)-restriction fragment length polymorphism matching of mycorrhizas and sporocarps and ITS sequencing. Fifty-one distinct taxa were distinguished; 25 were identified to species level, 11 to genus and 15 remained unidentified. Seventy-one ECM species were recorded as sporocarps from the forest plots; most (43 species) were found in the Sitka spruce plots. The below-ground ECM communities of the different forest types did not differ significantly with respect to species richness of taxa on roots, but differed in species composition. Multivariate analysis produced a clear separation of the communities of the different forest types using below-ground data, but the above-ground sporocarp data did not separate the forest types. Moreover, results of a Mantel test found no relationship between the above- and below-ground similarity matrices. The oak plots had the most distinctive ECM community, with Laccaria amethystina and Elaphomyces granulatus being frequent. The Sitka spruce plots showed the lowest intra-forest type similarity and were often dominated by "nursery type" ectomycorrhizas. There was only 10% similarity between the above- and below-ground ECM species in these plots, different colonisation methods of ectomycorrhizal taxa and insufficient below-ground sampling being possible reasons for this disparity. Our results indicate that plantations of non-native Sitka spruce can support similar levels of ECM diversity as native forests.     

Asynchronous origins of ectomycorrhizal clades of Agaricales

The ectomycorrhizal (ECM) symbiosis is the most widespread biotrophic nutritional mode in mushroom-forming fungi. ECM fungi include, though are not limited to, about 5000 described species of Agaricales from numerous, independently evolved lineages. Two central hypotheses suggest different explanations for the origin of ECM fungal diversity: (i) dual origins, initially with the Pinaceae in the Jurassic and later with angiosperms during the Late Cretaceous, and (ii) a simultaneous and convergent radiation of ECM lineages in response to cooling climate during the Palaeogene and advancing temperate ECM plant communities. Neither of these hypotheses is supported here. While we demonstrate support for asynchronous origins of ECM Agaricales, the timing of such events appears to have occurred more recently than suggested by the first hypothesis, first during the Cretaceous and later during the Palaeogene. We are also unable to reject models of rate constancy, which suggests that the diversity of ECM Agaricales is not a consequence of convergent rapid radiations following evolutionary transitions from saprotrophic to ECM habits. ECM lineages of Agaricales differ not only in age, but also in rates of diversification and rate of substitution at nuclear ribosomal RNA loci. These results question the biological uniformity of the ECM guild.     

Ectomycorrhizal community structure varies among Norway spruce (Picea abies) clones

In northern boreal forests, the diversity of ectomycorrhizal (ECM) species is much greater than that of their host trees. This field study investigated the role of individual trees in shaping the ECM community. We compared ECM communities of eight Norway spruce (Picea abies) clones planted in a clear-cut area in 1994 with a randomized block design. In 2003, the ECM fungi were identified from randomly sampled root tips using denaturing gradient gel electrophoresis (DGGE) and rDNA internal transcribed spacer (ITS) sequence similarity. ECM diversity varied among clone groups, showing twofold growth differences. Moreover, according to detrended correspondence analysis (DCA), ECM community structure varied not only among but also within slow-growing or fast-growing clones. Results suggest that ECM diversity and community structure are related to the growth rate or size of the host. A direct or indirect influence of host genotype was also observed, and we therefore suggest that individual trees are partly responsible for the high diversity and patchy distribution of ECM communities in boreal forests.     

Successional changes in ectomycorrhizal fungi associated with the polar willow Salix polaris in a deglaciated area in the High Arctic, Svalbard

Polar willow (Salix polaris Wahlenb.), a mycorrhizal dwarf shrub, colonizes recently deglaciated areas in the High Arctic, Svalbard. To clarify successional changes in ECM fungi associated with S. polaris after glacier retreat, we examined the diversity and density of ECM fungi in culture and field conditions. Plant and soil samples were collected from three sites of different successional stages in the deglaciated area of Austre Br?ggerbreen, near Ny-?lesund, Svalbard. The successional stages were early stage with newly exposed bare ground (site I), transient stage with scattered colonization of Salix (sites IIa and IIb), and late stage with well-developed vegetation (site III). No ECM colonization on Salix was observed in soils collected from bare ground in early and transient stages (sites I and IIa). However, most Salix individuals showed ECM colonization in soils collected from sites close to Salix colonies in transient and late stages (sites IIb and III). Based on molecular analyses and operational taxonomic unit (OTU: >95% ITS sequence similarity) delimitations, we identified 15 OTUs/species in eight genera. The dominant OTU/species of ECM fungi identified in the transient and late stages was Geopora sp.1 and Cenococcum sp.1, respectively. In the culture experiment, ECM diversity was greater in late stage (eight OTUs/species) than in transient stage (three OTUs/species). This pattern was consistent with field observations, i.e., late-stage sites contained more OTUs/species of ECM fungi. These results indicate that species diversity of ECM fungi increases and the dominant species changes with the progress of succession after glacier retreat in the High Arctic.     

Ectomycorrhizal fungal sporocarp diversity and discovery of new taxa in Dicymbe monodominant forests of the Guiana Shield

Ectomycorrhizal (ECM) fungi historically were considered poorly represented in Neotropical forests but in the central Guiana Shield substantial areas are dominated by leguminous ECM trees. In the Upper Potaro Basin of Western Guyana, ECM fungi were sampled for 7?years during the rainy seasons of 2000–2008 in three 1-ha plots in primary monodominant forests of the ECM canopy tree Dicymbe corymbosa (Fabaceae subfam. Caesalpinioideae). Over the plot sampling period sporocarps of 126 species of putative or confirmed ECM fungi were recovered. These taxa represented 13 families and 25 genera of primarily Agaricomycetes, but also Ascomycota (Elaphomycetaceae), the majority of which are new to science. Russulaceae contained the most species (20 Russula; 9 Lactarius), followed by Boletaceae (8 genera, 25 spp.), Clavulinaceae (17 Clavulina), and Amanitaceae (16 Amanita). An additional 46 species of ECM fungi were collected in forests of the Upper Potaro Basin outside the study plots between 2000 and 2010, bringing the regional number of ECM species known from sporocarps to 172. This is the first long-term ECM macrofungal dataset from an ECM-dominated Neotropical forest, and sporocarp diversity is comparable to that recorded for ECM-diverse temperate and boreal forests. While a species accumulation curve indicated that ECM sporocarp diversity was not fully recovered inside of the plots,?~80% of the total species were recovered in the first year. Sequence data from ECM roots have confirmed the ECM status of 56 taxa represented by corresponding sporocarp data. However,?>50% of ECM fungal species from roots remain undiscovered as sporocarps, leading to a conservative estimate of?>?250 ECM species at the Potaro site. Dicymbe forests in Guyana are a hotspot for ECM fungal diversity in the Neotropics.     

Diversity of ectomycorrhizal fungi in Britain: a test of the species–area relationship, and the role of host specificity

The host range of ectomycorrhizal (ECM) fungi in Britain was examined by compilation of a data matrix from published literature sources, based primarily on accounts of sporocarp associations with particular host genera. Information was gathered for 577 species of ECM fungi belonging to 51 genera, and 25 genera of host trees, representing the majority of ECM fungal species and host genera recorded in Britain.
Pronounced variation was recorded in the number of ECM fungal species associated with different host genera, with over 200 species recorded with Betula , Fagus , Pinus and Quercus . There was a positive linear relationship ( r ²=0·47, P =0·007) between the number of species of ECM fungi associated with different host genera and the total area occupied by each tree genus in Britain (both values log-transformed). There was also variation in the number of species of ECM fungi which were apparently specific to particular host genera, values ranging from zero (in 15 genera) to >40 in the case of Betula and Fagus . In total, 233 fungal species appeared to be specific to a single host genus (i.e. 40% of those surveyed). Comparison of the ECM mycota associated with different host genera by PCA accounted for 17% of the total variation, with genera belonging to the Fagaceae ( Quercus , Fagus and Castanea ) tending to cluster together, indicating a degree of overlap in their ECM associates. Exotic conifer species, which displayed a lower ECM diversity than would be expected from their distributional areas, were characterized by a high degree of overlap with the ECM associates of Pinus and Betula .
These results indicate that the abundance of different genera of host trees and variation in host specificity could provide a basis for understanding patterns of diversity in ECM fungi within Britain.     

Strong host preference of ectomycorrhizal fungi in a Tasmanian wet sclerophyll forest as revealed by DNA barcoding and taxon-specific primers

Ectomycorrhizal (ECM) symbiosis is a widespread plant nutrition strategy in Australia, especially in semiarid regions. This study aims to determine the diversity, community structure and host preference of ECM fungi in a Tasmanian wet sclerophyll forest. Ectomycorrhizal fungi were identified based on anatomotyping and rDNA internal transcribed spacer (ITS)-large subunit (LSU) sequence analysis using taxon-specific primers. Host tree roots were identified based on root morphology and length differences of the chloroplast trnL region. A total of 123 species of ECM fungi were recovered from root tips of Eucalyptus regnans (Myrtaceae), Pomaderris apetala (Rhamnaceae) and Nothofagus cunninghamii (Nothofagaceae). The frequency of two thirds of the most common ECM fungi from several lineages was significantly influenced by host species. The lineages of Cortinarius, Tomentella-Thelephora, Russula-Lactarius, Clavulina, Descolea and Laccaria prevailed in the total community and their species richness and relative abundance did not differ by host species. This study demonstrates that strongly host-preferring, though not directly specific, ECM fungi may dominate the below-ground community. Apart from the richness of Descolea, Tulasnella and Helotiales and the lack of Suillus-Rhizopogon and Amphinema-Tylospora, the ECM fungal diversity and phylogenetic community structure is similar to that in the Holarctic realm.     

Ectomycorrhiza communities of red oak (<Emphasis Type="BoldItalic">Quercus rubra</Emphasis> L.) of different age in the Lusatian lignite mining district,East Germany

Ectomycorrhizal (ECM) communities were assessed on a 720 m² plot along a chronosequence of red oak (Quercus rubra) stands on a forest reclamation site with disturbed soil in the lignite mining area of Lower Lusatia (Brandenburg, Germany). Adjacent to the mining area, a red oak reference stand with undisturbed soil was investigated reflecting mycorrhiza diversity of the intact landscape. Aboveground, sporocarp surveys were carried out during the fruiting season in a 2-week interval in the years 2002 and 2003. Belowground, ECM morphotypes were identified by comparing sequences of the internal transcribed spacer regions from nuclear rDNA with sequences from the GenBank database. Fifteen ECM fungal species were identified as sporocarps and 61 belowground as determined by morphological/anatomical and molecular analysis of their ectomycorrhizas. The number of ECM morphotypes increased with stand age along the chronosequence. However, the number of morphotypes was lower in stands with disturbed soil than with undisturbed soil. All stands showed site-specific ECM communities with low similarity between the chronosequence stands. The dominant ECM species in nearly all stands was Cenococcum geophilum, which reached an abundance approaching 80% in the 21-year-old chronosequence stand. Colonization rate of red oak was high (>95%) at all stands besides the youngest chronosequence stand where colonization rate was only 15%. This supports our idea that artificial inoculation with site-adapted mycorrhizal fungi would enhance colonization rate of red oak and thus plant growth and survival in the first years after outplanting.     

Ectomycorrhizal fungal diversity and community structure on three co-occurring leguminous canopy tree species in a Neotropical rainforest

? The ectomycorrhizal (ECM) symbiosis was historically considered restricted to the temperate zones, but recent studies have shown the importance of this symbiosis across the tropics. We examined ECM fungal diversity, host plant phylogeny and ECM host preferences in a rainforest dominated by the leguminous host plants Dicymbe corymbosa, Dicymbe altsonii and Aldina insignis. ? Ectomycorrhizal fungi were identified by internal transcribed spacer rDNA sequencing and host species were verified with chloroplast trnL sequencing. To test whether Dicymbe and Aldina represent independent gains of the ECM symbiosis, we constructed a Fabaceae phylogeny using MatK and trnL. We identified four independent ECM lineages within the Fabaceae. ? We detected a diverse community of 118 ECM species dominated by the /clavulina, /russula-lactarius, /boletus, and /tomentella-thelephora lineages. Ectomycorrhizal species in Agaricales, Atheliales and Polyporales may represent previously unrecognized tropical-endemic ECM lineages. Previous studies suggested that ECM fungi did not diversify in the tropics, but the /clavulina lineage appears to have a center of diversity in tropical South America. ? Dicymbe and Aldina represent independent gains of the ECM symbiosis in Fabaceae but their fungal symbionts showed no host preferences. Spatial factors are more important than hosts in structuring the ECM fungal community in this ecosystem.     

Ectomycorrhizal fungi of the Seychelles: diversity patterns and host shifts from the native Vateriopsis seychellarum (Dipterocarpaceae) and Intsia bijuga (Caesalpiniaceae) to the introduced Eucalyptus robusta (Myrtaceae), but not Pinus caribea (Pinaceae)

Ectomycorrhizal (ECM) fungi form highly diverse communities in temperate forests, but little is known about their community ecology in tropical ecosystems. Using anatomotyping and rDNA sequencing, ECM fungi were identified on root tips of the introduced Eucalyptus robusta and Pinus caribea as well as the endemic Vateriopsis seychellarum and indigenous Intsia bijuga in the Seychelles. Sequencing revealed 30 species of ECM fungi on root tips of V. seychellarum and I. bijuga, with three species overlapping. Eucalyptus robusta shared five of these taxa, whereas P. caribea hosted three unique species of ECM fungi that were likely cointroduced with containerized seedlings. The thelephoroid (including the anamorphic genus Riessiella), euagaric, boletoid and hymenochaetoid clades of basidiomycetes dominated the ECM fungal community of native trees. Two species of Annulatascaceae (Sordariales, Ascomycota) were identified and described as ECM symbionts of V. seychellarum. The low diversity of native ECM fungi is attributed to deforestation and long-term isolation of the Seychelles. Native ECM fungi associate with exotic eucalypts, whereas cointroduced ECM fungi persist in pine plantations for decades.     

Common environmental factors explain both ectomycorrhizal species diversity and pine regeneration variability in a post-fire Mediterranean forest

Natural seedling regeneration and establishment after stand replacing wildfires is influenced by a series of environmental and biological constraints. In this study, we characterized the diversity and structure of the ectomycorrhizal (ECM) fungal community associated with post-fire naturally regenerated maritime pine saplings, and individuate the environmental factors responsible for fungal species distribution. We also identify the main environmental factors responsible for maritime pine regeneration variability and assessed the relation between saplings performance and ECM fungal diversity indices. Fungal species were identified by direct sequencing of internal transcribed spacer regions. Five years after the disturbance event, a total of 30 taxa colonized the pine saplings. The ECM fungal community was dominated by ruderal species of the genus Rhizopogon (present in almost half of the samples). Almost one third of the identified ECM fungal species belonged to the family Thelephoraceae. Typical k-selected species like Amanita pantherina, Boletus aestivalis, Lactarius chrysorrheus, and Russula densifolia were found on pine saplings collected in proximity of unburnt pine trees, in correspondence with low erosion extents. Pine regeneration varied throughout the study areas and was enhanced at higher elevations, in correspondence with moderate slopes, shallower soils, and a reduced cover of ericaceous shrubs and bare ground. These conditions were found in close proximity to patches of pine trees that survived the disturbance event and were previously characterized by a higher pre-fire pine biomass. Even though no correlations were found between saplings performance and ECM fungal diversity indices, common environmental factors (i.e., ericaceous shrub cover, extent of erosion, slope, and soil depth) were responsible for shaping the ECM fungal distribution and for describing most of the explained regeneration variability.     

Vegetation and Soil Environment Influence the Spatial Distribution of Root-Associated Fungi in a Mature Beech-Maple Forest

Although the level of diversity of root-associated fungi can be quite high, the effect of plant distribution and soil environment on root-associated fungal communities at fine spatial scales has received little attention. Here, we examine how soil environment and plant distribution affect the occurrence, diversity, and community structure of root-associated fungi at local patch scales within a mature forest. We used terminal restriction fragment length polymorphism and sequence analysis to detect 63 fungal species representing 28 different genera colonizing tree root tips. At least 32 species matched previously identified mycorrhizal fungi, with the remaining fungi including both saprotrophic and parasitic species. Root fungal communities were significantly different between June and September, suggesting a rapid temporal change in root fungal communities. Plant distribution affected root fungal communities, with some root fungi positively correlated with tree diameter and herbaceous-plant coverage. Some aspects of the soil environment were correlated with root fungal community structure, with the abundance of some root fungi positively correlated with soil pH and moisture content in June and with soil phosphorous (P) in September. Fungal distribution and community structure may be governed by plant-soil interactions at fine spatial scales within a mature forest. Soil P may play a role in structuring root fungal communities at certain times of the year.In temperate forests, most trees form relationships with ectomycorrhizal (ECM) fungi, and the diversity of this fungal group alone can approach 100 species within a forest stand (17, 20, 60). The ECM mutualism may be necessary for the success of some native plant species, as approximately 90% of roots of some tree species are colonized by ECM fungi (65). Nevertheless, we still know surprisingly little about what controls the community structure and distribution of root-associated fungi in forest systems (44, 46). The occurrence of root-associated fungi may broadly reflect soil environmental conditions and the presence of preferred plant hosts (28, 61), but how these factors interact to influence the diversity, distribution, and community structure of these fungi within forest habitat patches at a local scale is uncertain.The distribution of root-associated fungi may be primarily a species response to local soil environmental conditions. For example, both the quality (i.e., nutrient content) and the quantity of soil organic matter are known to influence the diversity of ECM communities (18, 20, 32). ECM fungi also vary in drought tolerance (14, 36), resistance to fire (61, 65), and tolerance to soil acidity (19) and temperature (56). Changes in soil chemistry, especially as they relate to pH and the availability of nitrogen (N) and phosphorous (P), might favor selection of fungi most capable of tolerating environmental extremes (2, 28, 29).Plant distribution and identity may, however, play the strongest role in structuring the below-ground diversity of root-associated fungi. Many ECM fungi can colonize a wide range of plant species, and plant species can be host to a large number of ECM fungi (63), especially those in the families Russulaceae and Thelephoraceae (34, 35, 62). Moreover, some ECM fungi are also specific to certain tree species (e.g., Suillus and Rhizopogon species are specific to species in the family Pinaceae [38, 39]). At the local scale, fungal distribution and richness might be influenced by differences in root growth and architecture (30, 42), by the distance to the bole of the tree (11, 42, 49), or by the presence of neighboring trees (29, 64). Temporal changes in ECM communities could be associated with seasonal changes in plant physiology and phenology (3, 8, 17).An often overlooked factor influencing root-associated fungi of tree roots is the occurrence of herbaceous plant species within forest stands. Many species of parasitic, achlorophyllous angiosperms obtain carbon (C) from ECM fungi that colonize tree roots (43), and some autotrophic plants could also obtain C from ECM fungi during certain times of the year (58). Herbaceous plants also influence the cycling of nutrients, including N, P, and K (potassium) (31, 50), within forests, which could affect the distribution of root-associated fungi. Herbaceous plants can also produce secondary compounds that inhibit colonization of tree roots (68).In this study, we examine the effect of soil environment and plant distribution on root-associated fungi of tree roots in a mature beech-maple forest at two points in the growing season. We predict that plant distribution, both the distribution of host trees and that of herbaceous plants, influences fungi associated with tree roots in terms of both community structure and diversity. Molecular typing protocols, including a site-specific database of fungal sequences and fingerprints, were used to identify fungi on tree roots (i.e., beech or maple trees) to the species level.     

欧亚大陆东部毛茛科植物多样性格局及主导因子

毛茛科是真双子叶植物的基部类群之一, 包含多种药用植物, 具有较高的保护价值, 但关于毛茛科物种多样性和谱系多样性大尺度格局及其影响因子的研究还比较匮乏, 特别是以较高分辨率分布数据为基础的物种多样性格局研究尚未见报道。本文旨在: (1)建立欧亚大陆东部毛茛科植物分布数据库, 估算不同生活型物种多样性和谱系多样性格局, 并探究格局的形成机制。(2)分析毛茛科物种多样性和谱系多样性的相关关系, 确定多样性热点地区, 为毛茛科保护规划提供依据。根据中国、哈萨克斯坦、吉尔吉斯斯坦、塔吉克斯坦、土库曼斯坦、乌兹别克斯坦、蒙古和俄罗斯等国家的区域和地方植物志, 建立了“欧亚大陆东部地区毛茛科物种分布数据库”。该数据库包含了欧亚大陆东部地区1,688种毛茛科物种的分布数据, 空间分辨率为100 km × 100 km。在此基础上, 估算了毛茛科全部及不同生活型的物种多样性和谱系多样性格局, 并利用广义线性模型和等级方差分离方法分析了毛茛科物种和谱系多样性格局与环境因子的关系。最后比较了物种多样性和谱系多样性的相关关系, 确定了毛茛科的古热点地区。结果显示: (1)欧亚大陆东部毛茛科植物物种和谱系多样性均呈明显的纬度格局, 且在山区具有较高的多样性。(2)毛茛科植物物种和谱系多样性受现代气候、地形异质性和末次冰期以来的气候变化的共同影响, 但不同影响因子的相对贡献率在物种和谱系多样性及不同生活型之间差异显著。(3)中高纬度地区的谱系多样性高于给定物种数的预期, 是毛茛科的古热点地区, 在毛茛科保护规划中应受到重视。