Host effects in high ectomycorrhizal diversity tropical rainforests on ultramafic soils in New Caledonia

Tropical rainforests have been thought to have low prevalence and diversity of ectomycorrhizal symbioses. However, to date, tropical regions have been poorly sampled for ectomycorrhizal fungi. Here, we investigated ectomycorrhizal fungal community diversity and the role of host plants in shaping this diversity in three main ultramafic rainforests in New Caledonia, an archipelago renowned for its exceptional plant diversity and recognized as a biodiversity hotspot. Sampling of ectomycorrhizal root tips and fruit bodies in Nothofagus aequilateralis-dominated, Arillastrum gummiferum-dominated and mixed rainforests showed high fungal diversity with, in total, 28 lineages and 311 operational taxonomic units (OTUs), of which 95% might be endemic. We also found that host preference and host density influenced ectomycorrhizal community composition and contributed to the high fungal diversity of New Caledonian rainforests. Finally, the /cortinarius lineage dominated the below- and above-ground communities, which suggests that this lineage plays a central role in ultramafic ecosystems functioning.     

Regional scale gradients of climate and nitrogen deposition drive variation in ectomycorrhizal fungal communities associated with native Scots pine

Ectomycorrhizal fungi commonly associate with the roots of forest trees where they enhance nutrient and water uptake, promote seedling establishment and have an important role in forest nutrient cycling. Predicting the response of ectomycorrhizal fungi to environmental change is an important step to maintaining forest productivity in the future. These predictions are currently limited by an incomplete understanding of the relative significance of environmental drivers in determining the community composition of ectomycorrhizal (ECM) fungi at large spatial scales. To identify patterns of community composition in ECM fungi along regional scale gradients of climate and nitrogen deposition in Scotland, fungal communities were analysed from 15 seminatural Scots pine (Pinus sylvestris L.) forests. Fungal taxa were identified by sequencing of the ITS rDNA region using fungal‐specific primers. Nonmetric multidimensional scaling was used to assess the significance of 16 climatic, pollutant and edaphic variables on community composition. Vector fitting showed that there was a strong influence of rainfall and soil moisture on community composition at the species level, and a smaller impact of temperature on the abundance of ectomycorrhizal exploration types. Nitrogen deposition was also found to be important in determining community composition, but only when the forest experiencing the highest deposition (9.8 kg N ha^?1 yr^?1) was included in the analysis. This finding supports previously published critical load estimates for ectomycorrhizal fungi of 5–10 kg N ha^?1 yr^?1. This work demonstrates that both climate and nitrogen deposition can drive gradients of fungal community composition at a regional scale.     

Distribution of ectomycorrhizal fungi in a Chamaecyparis obtusa stand at different distances from a mature Quercus serrata tree

Ectomycorrhizal fungal distributions at different distances from one mature Quercus serrata tree were demonstrated in a Japanese cypress stand. After 4 months for field-planted and 3 months for laboratory-incubated Q. serrata seedlings, they were evaluated ectomycorrhiza formations. For the field-planted seedlings, molecular discrimination was conducted to clarify the species diversity of ectomycorrhizal fungi. Ectomycorrhiza formation was confirmed at all distances in both methods. However, the formation and associating fungal species were decreased with increasing distance. These results indicate that the density and diversity of ectomycorrhizal fungi decrease rapidly within several tens of meters from arbuscular mycorrhizal forest edges.     

Ectomycorrhizal fungal communities associated with Masson pine (Pinus massoniana Lamb.) in Pb–Zn mine sites of central south China

To advance our understanding of ectomycorrhizal fungal communities in mining areas, the diversity and composition of ectomycorrhizal fungi associated with Masson pine (Pinus massoniana Lamb.) and soil chemistry were investigated in Taolin lead–zinc (Pb–Zn) mine tailings (TLT), two fragmented forest patches in a Huayuan Pb–Zn mineland (HY1 and HY2), and a non-polluted forest in Taolin in central south China. Ectomycorrhizal fungal species were identified by morphotyping and sequence analyses of the internally transcribed spacer regions of ribosomal DNA. The two study sites in the Huayuan mineland (HY1 and HY2) were significantly different in soil Pb, Zn, and cadmium (Cd) concentrations, but no significant difference was observed in ectomycorrhizal colonization, ectomycorrhizal fungal richness, diversity, or rank–abundance. In addition, the similarity of ectomycorrhizal fungal communities between HY1 and HY2 was quite high (S?rensen similarity index?=?0.47). Thus, the concentration of heavy metals may not be determining factors in the structure of these communities. In the tailings, however, significantly lower ectomycorrhizal colonization and ectomycorrhizal fungal richness were observed. The amounts of Pb and Zn in the tailing sand were higher than the non-polluted forest but far lower than in HY1. Thus, these heavy metals did not account for the reduced colonization and ectomycorrhizal fungal richness in TLT. The ectomycorrhizal fungal community in TLT was dominated by four pioneer species (Rhizopogon buenoi, Tomentella ellisii, Inocybe curvipes, and Suillus granulatus), which collectively accounted for 93.2?% of root tip colonization. The immature soil conditions in tailing (low N and P, sand texture, and lack of organic matter) may only allow certain pioneer ectomycorrhizal fungal species to colonize the site. When soil samples from four sites were combined, we found that the occurrences of major ectomycorrhizal fungal taxa were not clearly related to the concentrations of Pb, Zn, and Cd. In conclusion, our results suggest that ectomycorrhizal fungal communities in mining areas are not necessarily affected by heavy metals themselves but could be largely determined by soil maturity.     

Fungal communities associated with roots of two closely related Juglandaceae species with a disjunct distribution in the tropics

We studied the biogeography and community structure of root-associated and ectomycorrhizal fungal communities in two related species of tropical Juglandaceae that have disjunct distributions in Asia and Mesoamerica. We tested the effects of environmental and dispersal factors in structuring root-associated fungi at a regional scale. We used Illumina sequencing to document fungi on the roots of Oreomunnea mexicana in Panama and Mexico and Alfaropsis roxburghiana in China. Ectomycorrhizal fungi dominated the communities with both hosts but we detected a more diverse root-associated fungal community in Alfaropsis but higher ectomycorrhizal fungi richness in Oreomunnea. Geographic distance was the best predictor of variation in fungal species composition, when including both hosts and when analyzing each host independently. However, our results showed a high correlation between geographic distance and abiotic variables, and therefore we were not able to determine if the observed changes in fungal community composition were explained also by spatially structured environmental or phylogenetic factors.     

连续三年夜间增温和施氮对云杉外生菌根及菌根真菌多样性的影响

以西南亚高山针叶林建群种粗枝云杉(Picea asperata)为研究对象,采用红外加热模拟增温结合外施氮肥(NH₄NO₃ 25 g N m^-2 a^-1)的方法,研究连续3a夜间增温和施肥对云杉幼苗外生菌根侵染率、土壤外生菌根真菌生物量及其群落多样性的影响。结果表明:夜间增温对云杉外生菌根侵染率的影响具有季节性及根级差异。夜间增温对春季(2011年5月)云杉1级根,夏季(2011年7月)和秋季(2010年10月)云杉2级根侵染率影响显著。除2011年7月1级根外,施氮对云杉1、2级根侵染率无显著影响。夜间增温对土壤中外生菌根真菌的生物量和群落多样性无显著影响,施氮及增温与施氮联合处理使土壤中外生菌根真菌生物量显著降低,但却提高了外生菌根真菌群落的多样性。这说明云杉幼苗外生菌根侵染率对温度较敏感,土壤外生菌根真菌生物量及其群落多样性对施氮较敏感。这为进一步研究该区域亚高山针叶林地下过程对全球气候变化的响应机制提供了科学依据。     

Ectomycorrhizal fungi are shared between seedlings and adults in a monodominant Gilbertiodendron dewevrei rain forest in Cameroon

Ectomycorrhizal networks may facilitate the establishment and survival of seedlings regenerating under the canopies of tropical forests and are often invoked as a potential contributor to monodominance. We identified ectomycorrhizal fungi in a monodominant Gilbertiodendron dewevrei (Fabaceae) rain forest in Cameroon, using sporocarps and ectomycorrhizae of three age categories (seedlings, intermediate trees, and large trees) and tentatively revealed nutrient transfer through ectomycorrhizal networks by measuring spontaneous isotopic (¹³C and ¹⁵N) abundances in seedlings. Sporocarp surveys revealed fewer ectomycorrhizal fungal taxa (59 species from 1030 sporocarps) than molecular barcoding of ectomycorrhizal roots (75 operational taxonomic units from 828 ectomycorrhizae). Our observations suggest that ectomycorrhizal fungal diversity is similar to that in other mixed tropical forests and provide the first report of the Tuber‐Helvella lineage in a tropical forest. Despite some differences, all age categories of G. dewevrei had overlapping ectomycorrhizal fungal communities, with families belonging to Thelephoraceae, Russulaceae, Sebacinaceae, Boletaceae, and Clavulinaceae. Of the 49 operational taxonomic units shared by the three age categories (65.3% of the ectomycorrhizal fungal community), 19 were the most abundant on root tips of all categories (38.7% of the shared taxa), supporting the likelihood of ectomycorrhizal networks. However, we obtained no evidence for nutrient transfer from trees to seedlings. We discuss the composition of the ectomycorrhizal fungal community among the G. dewevrei age categories and the possible role of common ectomycorrhizal networks in this rain forest.     

In vitro mycorrhization and acclimatization of Amanita caesareoides and its relatives on Pinus densiflora

Amanita caesareoides is a sister species of Amanita caesarea, also known as Caesar’s mushroom and one of the most desirable edible mycorrhizal mushrooms. However, cultivation of Caesar’s mushrooms has not yet been successful due to the difficulties involved in establishing pure cultures. In this study, we established pure cultures of four Asian Caesar’s mushroom species, i.e., A. caesareoides, Amanita javanica, Amanita esculenta, and Amanita similis, which were identified by sequence analysis of their rDNA internal transcribed spacer (ITS) region. Five selected isolates in A. caesareoides, A. javanica, and A. esculenta were tested for ectomycorrhizal syntheses with axenic Pinus densiflora seedlings in vitro. Ectomycorrhizal tips of each fungal isolate tested were observed on pine lateral roots within 5 months of inoculation. Seventeen pine seedlings that formed ectomycorrhizas in vitro with these three Amanita species were acclimatized under non-sterile conditions. Seven months following acclimatization, ectomycorrhizal colonization by A. caesareoides was observed on newly grown root tips, which was confirmed by polymerase chain reaction restriction fragment length polymorphism analysis of the fungal rDNA ITS region. Two other Amanita species also survived during ectomycorrhizal acclimatization. These results suggest that the cultivation of A. caesareoides and its relatives can be attempted through mycorrhizal synthesis using P. densiflora as a host. This is the first report of in vitro mycorrhization of Asian Caesar’s mushrooms and their acclimatization under non-sterile conditions.     

Forest Age and Plant Species Composition Determine the Soil Fungal Community Composition in a Chinese Subtropical Forest

Fungal diversity and community composition are mainly related to soil and vegetation factors. However, the relative contribution of the different drivers remains largely unexplored, especially in subtropical forest ecosystems. We studied the fungal diversity and community composition of soils sampled from 12 comparative study plots representing three forest age classes (Young: 10–40 yrs; Medium: 40–80 yrs; Old: ≥80 yrs) in Gutianshan National Nature Reserve in South-eastern China. Soil fungal communities were assessed employing ITS rDNA pyrotag sequencing. Members of Basidiomycota and Ascomycota dominated the fungal community, with 22 putative ectomycorrhizal fungal families, where Russulaceae and Thelephoraceae were the most abundant taxa. Analysis of similarity showed that the fungal community composition significantly differed among the three forest age classes. Forest age class, elevation of the study plots, and soil organic carbon (SOC) were the most important factors shaping the fungal community composition. We found a significant correlation between plant and fungal communities at different taxonomic and functional group levels, including a strong relationship between ectomycorrhizal fungal and non-ectomycorrhizal plant communities. Our results suggest that in subtropical forests, plant species community composition is the main driver of the soil fungal diversity and community composition.     

New evidence of ectomycorrhizal fungi in the Hawaiian Islands associated with the endemic host Pisonia sandwicensis (Nyctaginaceae)

Ectomycorrhizal plants and fungi are ubiquitous in mainland forests, but because of dispersal limitations are predicted to be less common on isolated islands. For instance, no native ectomycorrhizal plants or fungi have ever been reported from Hawaii, one of the most remote archipelagos on Earth. Members of the plant tribe Pisonieae are common on many islands, and prior evidence shows that some species associate with ectomycorrhizal fungi. However, until now, the Pisonieae species of Hawaii had yet to be examined for their mycorrhizal status. Here we sampled roots from members of the genus Pisonia growing on the Hawaiian islands of Oahu, Maui and Hawaii. We used molecular and microscopic techniques to categorize trees with respect to their mycorrhizal associations. We report that the Hawaiian endemic Pisonia sandwicensis forms ectomycorrhizas with at least five fungal operational taxonomic units (corresponding closely to species) belonging to four genera. We also report that this tree species is monophyletic with other ectomycorrhizal Pisonia species. We suggest that in light of the newly discovered Hawaiian ectomycorrhizal fungal community and other island ectomycorrhizal communities, dispersal limitations do not prevent the colonization of remote islands by at least some ectomycorrhizal fungi.     

Fungal content of ectomycorrhizal tips: comparison among 13 tree species

To better understand soil carbon cycling in forest ecosystems, we studied the proportion of fungal sheath area (FSA) in the cross-sectional ectomycorrhizal area in 13 tree species. Ectomycorrhizal samples were collected from subalpine and temperate forests in Japan. The FSA values were in the range of 12% to 56% across all tree species, tree ages, and fungal species. In Abies firma and Quercus serrata, the FSA values were larger in mature trees than in seedlings, whereas no such differences were found in Pinus densiflora and Fagus crenata. In broad-leaved trees, because the plant tissue radii lay within a narrow range, the FSA was affected mainly by the fungal sheath thickness. In conifers, however, the plant tissue radii varied widely among genera, so the FSA was affected by both the plant tissue radius and the fungal sheath thickness. Our findings suggest that the fungal content of ectomycorrhizal tips differs among tree species and fungal species, so that both parameters must be considered in studies of forest carbon cycling. The estimates revealed that data gathering in each type of forest leads to more accurate estimates of the biomass of fungi in ectomycorrhizal tips.     

Evidence of species interactions within an ectomycorrhizal fungal community

Ectomycorrhizal fungal communities can be structured by abiotic and biotic factors. Here, we present evidence for community structuring by species interactions. We sampled ectomycorrhizas and forest floor seven times during a 13-month period. The presence of various ectomycorrhizal fungal species was determined for each sample, and species co-occurrence analyses were performed. For both ectomycorrhizas and forest floor samples there was significantly less co-occurrence among species within the community than expected by chance, mostly because of negative associations involving Cenococcum geophilum or Clavulina cinerea. For some species pairs, there was significantly more co-occurrence than expected by chance. Both nitrogen and tannin additions to the forest floor altered some interactions among species. The causes of these nonrandom distributions are currently unknown. Future investigations on competition, antibiosis, parasitism and facilitation among ectomycorrhizal fungal species appear to be warranted.     

Ectomycorrhizal Specificity Patterns in a Mixed Pinus contorta and Picea engelmannii Forest in Yellowstone National Park

We used molecular genetic methods to test two hypotheses, (i) that host plant specificity among ectomycorrhizal fungi would be common in a closed-canopy, mixed Pinus contorta-Picea engelmannii forest in Yellowstone National Park and (ii) that specificity would be more common in the early successional tree species, P. contorta, than in the invader, P. engelmannii. We identified 28 ectomycorrhizal fungal species collected from 27 soil cores. The proportion of P. engelmannii to P. contorta ectomycorrhizae was nearly equal (52 and 48%, respectively). Of the 28 fungal species, 18 composed greater than 95% of the fungal community. No species was associated exclusively with P. contorta, but four species, each found in only one core, and one species found in two cores were associated exclusively with P. engelmannii. These fungi composed less than 5% of the total ectomycorrhizae. Thus, neither hypothesis was supported, and hypothesized benefits of ectomycorrhizal specificity to both trees and fungi probably do not exist in this system.     

Host plants and edaphic factors influence the distribution and diversity of ectomycorrhizal fungal fruiting bodies within rainforests from Tshopo,Democratic Republic of the Congo

Ectomycorrhizal fungi constitute an important component of forest ecosystems that enhances plant nutrition and resistance against stresses. Diversity of ectomycorrhizal (EcM) fungi is, however, affected by host plant diversity and soil heterogeneity. This study provides information about the influence of host plants and soil resources on the diversity of ectomycorrhizal fungal fruiting bodies from rainforests of the Democratic Republic of the Congo. Based on the presence of fungal fruiting bodies, significant differences in the number of ectomycorrhizal fungi species existed between forest stand types (p < 0.001). The most ectomycorrhizal species‐rich forest was the Gilbertiodendron dewevrei‐dominated forest (61 species). Of all 93 species of ectomycorrhizal fungi, 19 demonstrated a significant indicator value for particular forest stand types. Of all analysed edaphic factors, the percentage of silt particles was the most important parameter influencing EcM fungi host plant tree distribution. Both host trees and edaphic factors strongly affected the distribution and diversity of EcM fungi. EcM fungi may have developed differently their ability to successfully colonise root systems in relation to the availability of nutrients.     

Establishment of ectomycorrhizal fungal community on isolated Nothofagus cunninghamii seedlings regenerating on dead wood in Australian wet temperate forests: does fruit-body type matter?

Decaying wood provides an important habitat for animals and forms a seed bed for many shade-intolerant, small-seeded plants, particularly Nothofagus. Using morphotyping and rDNA sequence analysis, we compared the ectomycorrhizal fungal community of isolated N. cunninghamii seedlings regenerating in decayed wood against that of mature tree roots in the forest floor soil. The /cortinarius, /russula-lactarius, and /laccaria were the most species-rich and abundant lineages in forest floor soil in Australian sites at Yarra, Victoria and Warra, Tasmania. On root tips of seedlings in dead wood, a subset of the forest floor taxa were prevalent among them species of /laccaria, /tomentella-thelephora, and /descolea, but other forest floor dominants were rare. Statistical analyses suggested that the fungal community differs between forest floor soil and dead wood at the level of both species and phylogenetic lineage. The fungal species colonizing isolated seedlings on decayed wood in austral forests were taxonomically dissimilar to the species dominating in similar habitats in Europe. We conclude that formation of a resupinate fruit body type on the underside of decayed wood is not necessarily related to preferential root colonization in decayed wood. Rather, biogeographic factors as well as differential dispersal and competitive abilities of fungal taxa are likely to play a key role in structuring the ectomycorrhizal fungal community on isolated seedlings in decaying wood.     

Contrasting below-ground views of an ectomycorrhizal fungal community

Ectomycorrhizal fungal communities have been characterized in a number of ways. Here we compare colonized root-tip and mycelia views of an ectomycorrhizal fungal community. Ectomycorrhizal fungi, both as mycelia and colonized root tips, were identified in soil samples taken from a pine plantation. We determined that for some ectomycorrhizal fungal species multiple root tips from a single soil sample were not independent. Therefore in the comparison of root-tip and mycelia views, we considered species to be present or absent from each soil sample irrespective of the number of root tips colonized by the species. We observed 39 ectomycorrhizal fungal species in total, but 12 were observed exclusively as mycelia and 11 exclusively colonizing root tips. The relative frequencies of 10 species occurring as both mycelia and root tips were not independent of the method of observation. Our results suggest that ectomycorrhizal fungal species differ in their spatial distributions on root tips, and that root-tip and mycelia views of the community are different.     

Plant identity strongly structures the root-associated fungal community in a diverse subtropical forest

Revealing the relationship between plants and root-associated fungi is very important in understanding diversity maintenance and community assembly in ecosystems. However, the community assembly of root-associated fungi of focal plant species along a subtropical plant species diversity gradient is less documented. Here, we examined root-associated fungal communities associated with five ectomycorrhizal (EM) plant species (Betula luminifera, Castanea henryi, Castanopsis fargesii, C. sclerophylla, and Quercus serrate) in a Chinese subtropical woody plant species diversity (1, 2, 4, 8, 16 and 24 species) experiment, using paired-end Illumina MiSeq sequencing of the ITS2 region. In total, we detected 1933 root-associated fungal operational taxonomic units (OTUs) at a 97% sequence similarity level. Plant identity had a significant effect on total and saprotrophic fungal OTU richness, but plant species diversity level had a significant effect on saprotrophic and pathogenic fungal OTU richness. The community composition of total, saprotrophic and EM fungi was structured by plant identity and plant species diversity level. However, the community composition of pathogenic fungi was only shaped by plant identity. This study highlights that plant identity has a stronger effect on the root-associated fungal community than plant species diversity level in a diverse subtropical forest ecosystem.     

Mycorrhizal networks mediate overstorey-understorey competition in a temperate forest

In forests, common mycorrhizal networks (CMNs) often connect the roots of neighbouring plants. Observations of material flows between hosts connected by CMNs have given rise to the hypothesis that CMNs limit the negative effects of competition by overstorey trees on seedlings recruiting underneath them. I conducted an experiment in a temperate forest dominated by ectomycorrhizal conifers and hardwoods to isolate the effects of CMNs on the growth and survival of four tree species that co‐occur in the understorey. Ectomycorrhizal networks had strong negative effects on the survival of an arbuscular mycorrhizal species, Acer rubrum, and neutral effects on the survival of three ectomycorrhizal species, Betula allegheniensis, Pinus strobus, and Tsuga canadensis. CMNs had positive effects on the growth of at least one ectomycorrhizal species, P. strobus. Interspecific differences in juvenile responses to CMNs may influence forest community development, promoting coexistence of some tree species while limiting the success of others.     

Diversity and Spatial Structure of Belowground Plant–Fungal Symbiosis in a Mixed Subtropical Forest of Ectomycorrhizal and Arbuscular Mycorrhizal Plants

Plant–mycorrhizal fungal interactions are ubiquitous in forest ecosystems. While ectomycorrhizal plants and their fungi generally dominate temperate forests, arbuscular mycorrhizal symbiosis is common in the tropics. In subtropical regions, however, ectomycorrhizal and arbuscular mycorrhizal plants co-occur at comparable abundances in single forests, presumably generating complex community structures of root-associated fungi. To reveal root-associated fungal community structure in a mixed forest of ectomycorrhizal and arbuscular mycorrhizal plants, we conducted a massively-parallel pyrosequencing analysis, targeting fungi in the roots of 36 plant species that co-occur in a subtropical forest. In total, 580 fungal operational taxonomic units were detected, of which 132 and 58 were probably ectomycorrhizal and arbuscular mycorrhizal, respectively. As expected, the composition of fungal symbionts differed between fagaceous (ectomycorrhizal) and non-fagaceous (possibly arbuscular mycorrhizal) plants. However, non-fagaceous plants were associated with not only arbuscular mycorrhizal fungi but also several clades of ectomycorrhizal (e.g., Russula) and root-endophytic ascomycete fungi. Many of the ectomycorrhizal and root-endophytic fungi were detected from both fagaceous and non-fagaceous plants in the community. Interestingly, ectomycorrhizal and arbuscular mycorrhizal fungi were concurrently detected from tiny root fragments of non-fagaceous plants. The plant–fungal associations in the forest were spatially structured, and non-fagaceous plant roots hosted ectomycorrhizal fungi more often in the proximity of ectomycorrhizal plant roots. Overall, this study suggests that belowground plant–fungal symbiosis in subtropical forests is complex in that it includes “non-typical” plant–fungal combinations (e.g., ectomycorrhizal fungi on possibly arbuscular mycorrhizal plants) that do not fall within the conventional classification of mycorrhizal symbioses, and in that associations with multiple functional (or phylogenetic) groups of fungi are ubiquitous among plants. Moreover, ectomycorrhizal fungal symbionts of fagaceous plants may “invade” the roots of neighboring non-fagaceous plants, potentially influencing the interactions between non-fagaceous plants and their arbuscular-mycorrhizal fungal symbionts at a fine spatial scale.     

Disproportionate abundance between ectomycorrhizal root tips and their associated mycelia

Extensive knowledge of various ectomycorrhizal fungal communities has been obtained over the past 10 years based on molecular identification of the fungi colonizing fine roots. In contrast, only limited information exists about the species composition of ectomycorrhizal hyphae in soil. This study compared the ectomycorrhizal external mycelial community with the adjacent root-tip community in a Danish beech forest. Sand-filled in-growth mesh bags were used to trap external mycelia by incubating the mesh bags in the soil for 70 days. The adjacent ectomycorrhizal root-tip communities were recorded at the times of insertion and retrieval of the mesh bags. Ectomycorrhizal fungi were identified by sequencing the internal transcribed spacer region. In total, 20, 31 and 24 ectomycorrhizal species were recorded from the two root-tip harvests and from the mesh bags, respectively. Boletoid species were significantly more frequent as mycelia than as root tips, while russuloid and Cortinarius species appeared to be less dominant as mycelia than as root tips. Tomentella species were equally frequent as root tips and as mycelia. These discrepancies between the root-tip and the mycelial view of the ectomycorrhizal fungal community are discussed within the framework of ectomycorrrhizal exploration types.