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Summary• The role of mycorrhizal networks in forest dynamics is poorly understood because of the elusiveness of their spatial structure. We mapped the belowground distribution of the fungi Rhizopogon vesiculosus and Rhizopogon vinicolor and interior Douglas-fir trees (Pseudotsuga menziesii var. glauca) to determine the architecture of a mycorrhizal network in a multi-aged old-growth forest.• Rhizopogon spp. mycorrhizas were collected within a 30 · 30 m plot. Trees and fungal genets were identified using multi-locus microsatellite DNA analysis. Tree genotypes from mycorrhizas were matched to reference trees aboveground. Two trees were considered linked if they shared the same fungal genet(s).• The two Rhizopogon species each formed 13-14 genets, each colonizing up to 19 trees in the plot. Rhizopogon vesiculosus genets were larger, occurred at greater depths, and linked more trees than genets of R. vinicolor. Multiple tree cohorts were linked, with young saplings established within the mycorrhizal network of Douglas-fir veterans. A strong positive relationship was found between tree size and connectivity, resulting in a scale-free network architecture with smallworld properties.• This mycorrhizal network architecture suggests an efficient and robust network, where large trees play a foundational role in facilitating conspecific regeneration and stabilizing the ecosystem.

1. From the phytocentric perspective, a mycorrhizal network (MN) is formed when the roots of two or more plants are colonized by the same fungal genet. MNs can be modelled as interaction networks with plants as nodes and fungal genets as links. The potential effects of MNs on facilitation or competition between plants are increasingly recognized, but their network topologies remain largely unknown. This information is needed to understand the ecological significance of MN functional traits. 2. The objectives of this study were to describe the interaction network topologies of MNs formed between two ectomycorrhizal fungal species, Rhizopogon vesiculosus and R. vinicolor, and interior Douglas-fir trees at the forest stand scale, identify factors leading to this structure and to contrast MN structures between forest plots with xeric versus mesic soil moisture regimes. 3. Tuberculate mycorrhizas were sampled in six 10 9 10 m plots with either xeric or mesic soil moisture regimes. Microsatellite DNA markers were used to identify tree and fungal genotypes isolated from mycorrhizas and for comparison with reference tree boles above-ground. 4. In all six plots, trees and fungal genets were highly interconnected. Size asymmetries between different tree cohorts led to non-random MN topologies, while differences in size and connectivity between Rhizopogon species-specific subnetwork components contributed towards MN nestedness. Large mature trees acted as network hubs with a significantly higher node degree compared to smaller trees. MNs representing trees linked by R. vinicolor genets were mostly nested within larger, more highly connected R. vesiculosus-linked MNs. 5. Attributes of network nodes showed that hub trees were more important to MN topology on xeric than mesic sites, but the emergent structures of MNs were similar in the two soil moisture regimes. 6. Synthesis. This study suggests MNs formed between interior Douglas-fir trees and R. vesiculosus and R. vinicolor genets are resilient to the random loss of participants, and to soil water stress, but may be susceptible to the loss of large trees or fungal genets. Our results regarding the topology of MNs contribute to the understanding of forest stand dynamics and the resilience of forests to stress or disturbance.

Understanding ectomycorrhizal fungal (EMF) community structure is limited by a lack of taxonomic resolution and autecological information. Rhizopogon vesiculosus and Rhizopogon vinicolor (Basidiomycota) are morphologically and genetically related species. They are dominant members of interior Douglas-fir (Pseudotsuga menziesii var. glauca) EMF communities, but mechanisms leading to their coexistence are unknown. We investigated the microsite associations and foraging strategy of individual R. vesiculosus and R. vinicolor genets. Mycelia spatial patterns, pervasiveness and root colonization patterns of fungal genets were compared between Rhizopogon species and between xeric and mesic soil moisture regimes. Rhizopogon spp. mycelia were systematically excavated from the soil and identified using microsatellite DNA markers. Rhizopogon vesiculosus mycelia occurred at greater depth, were more spatially pervasive, and colonized more tree roots than R. vinicolor mycelia. Both species were frequently encountered in organic layers and between the interface of organic and mineral horizons. They were particularly abundant within microsites associated with soil moisture retention. The occurrence of R. vesiculosus shifted in the presence of R. vinicolor towards mineral soil horizons, where R. vinicolor was mostly absent. This suggests that competition and foraging strategy may contribute towards the vertical partitioning observed between these species. Rhizopogon vesiculosus and R. vinicolor mycelia systems occurred at greater mean depths and were more pervasive in mesic plots compared with xeric plots. The spatial continuity and number of trees colonized by genets of each species did not significantly differ between soil moisture regimes.

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Mycorrhizal networks are conduits for the transfer of resources between hosts. While ectomycorrhizal networks (EMN) are known to influence seedlings, their effect on adult tree growth remains unknown and may have important implications for forest responses to future climates. We used annual basal area increment of trees and previously described Rhizopogon vesiculosus and Rhizopogon vinicolor EMNs to examine an association between the number of connections between trees through an EMN and the growth of adult interior Douglas‐fir. We compared this relationship for the year the networks were mapped, in 2008, with 8 years previous and 8 years afterward. We also compared the variation in standardized growth (2000–2016) to examine the association between growth variability and EMN variables. Greater growth was positively associated with (a) the number of connections to other trees via a R. vinicolor EMN and (b) the number of genets of Rhizopogon vesiculosis by which a tree was colonized. Variation of growth (2000–2016) was negatively associated with increasing number of connections to other trees via R. vinicolor. Synthesis. These findings, for the first time, indicate that EMNs may positively influence the growth of adult trees. The difference in tree growth response between the sister fungal species highlights a novel avenue to identify interspecific and intraspecific differences between fungi occurring at different depths in the soil. Our study has important implications when considering the role of EMNs in influencing forest health and mitigating stress from environmental conditions.

Rhizopogon vesiculosus and Rhizopogon vinicolor are sister species of ectomycorrhizal fungi that associate exclusively with Douglas-fir (DF). They form tuberculate mycorrhizas and they can be easily distinguished using molecular tools. We are not aware of studies relating their relative abundance in forests with different age classes. Our objective was to determine whether a change in the number or relative abundance of R. vesiculosus and R. vinicolor tubercules and genotypes was related to a change in the percent of DF in a regenerating phase (<50 years old). R. vesiculosus and R. vinicolor were located by excavating tuberculate mycorrhizas from the forest floor. A DNA Alu1 digest was used to distinguish between the two species. Microsatellite markers were used to identify genotypes. The number of R. vesiculosus tubercules correlated positively with an increasing proportion of DF in a regenerating phase, while the number of R. vinicolor tubercules was similar across all forest age structures. The number of R. vesiculosus genotypes did not correlate with forest age structure, whereas the number of R. vinicolor genotypes showed a negative relationship with an increasing proportion of DF in a regenerating phase. When the numbers of R. vesiculosus tubercules and genotypes were expressed as a relative abundance of the two species, there was a positive correlation with an increasing proportion of DF in a regenerating phase for both genotypes and tubercules. Our results suggest that the degree of DF regeneration or ecosystem factors related to DF regeneration affect the population dynamics of R. vesiculosus and R. vinicolor differently.

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