Diversity of coleoptera of the bracket fungus Fomitopsis pinicola in a Norwegian spruce forest

The beetle fauna of 299 sporocarps of the bracket fungus Fomitopsis pinicola in a 200 ha spruce forest in southeastern Norway was investigated in relation to sporocarp, tree and forest variables. The sporocarps contained 36 species of beetles, of which six species are on the Norwegian Red List. Of 12,373 individual beetles collected, 91% were Cis glabratus. Plots of species accumulation curves suggested that there may be more than 60 beetle species present in F. pinicola in the area, but that probably all the specialist Ciidae were found. The major factor influencing beetle diversity turned out to be the level of dead wood at and in the vicinity of the sampling site, with a higher number of species per unit volume of sporocarps in areas with high levels of dead wood. There were also significantly more red-listed species in those areas. Analysing the species occurrence with stepwise logistic regression, we show preferential habitat selection of the six most abundant species of Ciidae. Conservation of beetles associated with bracket fungi using amount of dead wood as a surrogate measurement is discussed.     

The Red-belted Bracket (Fomitopsis pinicola) colonizes spruce trees early after bark beetle attack and persists

The Red-belted Bracket (Fomitopsis pinicola) is one of the major decomposers of coniferous wood in Europe and can reach high densities after outbreaks of bark beetles. However, factors of dead wood type and decay stage, which determine the growth of reproductive biomass, i.e. basidiomes, remain unclear. In 2013, we surveyed 1280 dead wood objects and vital trees in spruce stands killed by the bark beetle Ips typographus in 2012, 2002, 1992 and in undisturbed stands for the presence, number, mean basidiome size and total volume of basidiomes. Living basidiomes were equally abundant on dead wood 1, 11, and 21 y after bark beetle outbreak, but were lacking on living trees. Our results indicate that F. pinicola is an effective early colonizer of the huge resource pulse of dead wood caused by the outbreak of bark beetles and basidiomes can persist for 21 y.     

Saproxylic beetle assemblages related to silvicultural management intensity and stand structures in a beech forest in Southern Germany

Compared to agricultural land and spruce plantations, central European beech-oak forests are often relatively close to natural conditions. However, forest management may alter these conditions. In Steigerwald, southern Germany, a large beech-dominated forest area, three management intensities were applied during the past 30–70 years. Here, we examined the influence of management intensity on saproxylic beetles in >100-year old mature stands at 69 sampling plots in 2004. We sampled beetles using flight-window traps and time standard direct searches. The community structure based on presence/absence data changed remarkably along the gradient from unmanaged to low-intensity to high-intensity management, but these differences were not evident using abundance data from flight interception traps. Saproxylic species richness decreased in intensively managed forests. Elateridae and threatened species richness peaked in unmanaged forests and in forests under low-intensity management. Saproxylic species richness was dependent on certain micro-habitat factors. These factors were (1) the amount of dead wood for Elateridae, overall and threatened saproxylic beetle richness; (2) the amount of flowering plants for Cerambycidae; (3) the richness of wood-inhabiting fungi for Staphylinidae, Melandryidae and overall saproxylic beetle richness; and (4) the frequency of Fomes fomentarius for threatened species. Species richness was better explained by plot factors, such as dead wood or fungi, than by management intensity. These results suggest that the natural variation of dead wood niches (decay stages, snag sizes, tree cavities and wood-inhabiting fungi species) must be maintained to efficiently conserve the whole saproxylic beetle fauna of beech forests. Also, intensive management may alter the specialised saproxylic beetle community even if the initial tree-species composition is maintained, which was the case in our study. For monitoring the ecological sustainability of forest management we must focus on threatened species. If structures alone are sampled then the amount of dead wood is the best indicator for a rich saproxylic beetle fauna.     

Fast but ephemeral effects of ecological restoration on forest beetle community

Many protected areas have a long history of human intervention before being protected. In protected forests, the past land use has reduced the amount of natural structures, which are crucial substrates for thousands of species. We evaluate the short-term ecological effect of forest restoration (dead wood creation) on conifer-associated saproxylic (dead-wood dependent) beetles. More specifically, we analyze the effect of dead wood creation on the number of beetle species and individuals 1 and 5 years after restoration in spruce and pine forests, using a large-scale monitoring network over Finland. The number of saproxylic beetle species and individuals was larger at restored than at control plots both 1 and 5 years after restoration in both spruce and pine forests. Community composition in restored plots was different from control plots 1 year after restoration, but had returned towards the control plot composition 5 years after restoration, while control plots remained largely unchanged. Both in spruce and pine forests, there were more red-listed and rare saproxylic beetles in restored than in control plots 1 and 5 years after restoration. Our results indicate that restoration has an overall positive influence on saproxylic beetle diversity immediately after dead wood creation, but this effect is rather short-lived. Long term monitoring of restored dead wood is crucial in investigating successional pathways as well as biotic communities in advanced decay stages, and in fully evaluating the ecological effect of dead wood creation as a forest restoration measure.     

Wood-inhabiting insects can function as targeted vectors for decomposer fungi

Most wood-inhabiting fungi are assumed to be dispersed primarily by wind, with the exception of a few species involved in mutualistic relationships with insects. In this study we tested whether several species of wood-inhabiting insects can function as dispersal vectors for non-mutualistic fungi, which would indicate that wood-inhabiting fungi can benefit from targeted animal-mediated dispersal. We sampled wood-inhabiting beetles (Coleoptera) from freshly felled wood experimentally added to forests and used DNA metabarcoding to investigate the fungal DNA carried by these insects. Staphylinid beetles rarely contained fungal DNA, while Endomychus coccineus, Glischrochilus hortensis and Glischrochilus quadripunctatus frequently carried fungal DNA with a composition specific to the insect taxon. A large proportion of the obtained fungal sequences (34%) represented decomposer fungi, including well-known wood-decay fungi such as Fomitopsis pinicola, Fomes fomentarius, Trichaptum abietinum and Trametes versicolor. Scanning electron microscopy further showed that some of the fungal material was carried as spores or yeast cells on the insect exoskeletons. Our results suggest that insect-vectored dispersal is of broader importance to wood-inhabiting fungi than previously assumed.     

Fungi associated with beetles dispersing from dead wood – Let's take the beetle bus!

Spore characteristics of wood-inhabiting fungi suggest that wind is their predominant dispersal vector. However, since they are restricted to ephemeral habitats, colonizing new patches should benefit from dispersal by animals with similar habitat preferences because the directed, resource-searching movement of animals increases the likelihood of reaching suitable habitats. Here we determine which fungal guilds are carried by wood-inhabiting beetles and what influences beetle-associated fungal communities. High-throughput sequencing identified >1800 fungal taxa from beetle communities that emerged from 64 experimental logs. Beetle-associated fungi included mutualistic, decomposing, pathogenic and mycorrhizal fungi; decomposers were the most diverse. Partial-procrustes analysis revealed that the total beetle-associated community and mutualists were correlated (p ≤ 0.05) with beetle community composition and decomposers were marginally correlated (p ≤ 0.10) with beetle community composition. All three groups were marginally correlated with the total fungal communities that inhabit the dead wood. Our results show that beetles carry a broad range of wood-inhabiting fungi and beetle-associated fungal communities are determined by environmental factors and the vectoring beetle community and to some degree by the fungal source community. This suggests that wood-inhabiting beetles contribute to fungal dispersal, including directed dispersal, which could affect fungal community assembly and ecosystem processes like wood decomposition.     

Can 'continuity indicator species' predict species richness or red-listed species of saproxylic beetles?

This study investigates the relationship between the abundance of wood-rotting fungus suggested as 'continuity indicator species' and environmental variables for the assemblage of saproxylic (wood-living) beetles associated with Fomitopsis pinicola fruiting bodies in a mature spruce forest in southeastern Norway. The presence of species thought to indicate continuity in old growth is one of the criteria used when finding and delineating small protected areas ('woodland key habitats') in Scandinavian forestry. Although it is clear that remnants of old-growth forest are important for many taxa, documentation as to which entities or species the indicator species indeed indicate is scarce. If stands with a continuous and unbroken input of dead wood have a unique assemblage of wood-rotting fungi, it seems relevant to ask if these stands also have a unique assemblage of rare saproxylic beetles. I find that the indicator species exhibit no significant correlations with beetle species richness or with the presence of red-listed saproxylic beetles as a group. The different characteristics of dead wood conditions are the most important environmental variables that explain both the species richness and the presence of red-listed beetles. Single-species analyses reveal contrasting relationships. The red-listed beetle Atomaria alpina shows a significant and positive association to the abundance of indicator species. Contrary, a group of three red-listed species with similar ecology in the family Cisidae exhibits a significant and negative association to indicator species abundance. This indicates that important patterns are concealed when considering general measures such as overall presence of red-listed beetles. Single-species studies are necessary in order to correctly understand how rare beetles respond to forestry activities and to develop a policy that can secure their continuing existence in the boreal forest.     

Wood-inhabiting fungi on fallen logs of Norway spruce: relations to forest management and substrate quality

A survey of the patterns of wood decaying fungi as to occurrence of sporocarps on naturally fallen logs of Norway spruce ( Picea abies ) was undertaken in two nearby forest stands with different histories of management. One stand was an old-growth forest with few signs of logging, and the other stand was selectively logged 60–80 years ago. Altogether 118 species were found. Forest management had a negative impact on the species diversity. Newly fallen and weakly decayed logs in a natural forest had a higher species richness, more red-listed species, as well as more indicator species compared to similar logs in a managed forest. The importance of dead wood for species diversity of wood inhabiting fungi was clearly demonstrated. Presence of logs in later stages of decomposition increased the total species number in a natural forest stand with 42 (63 %), compared to a survey of only newly fallen and weakly decayed logs. Presence of logs in later stages of decomposition also increased the diversity of the species pool colonising newly fallen and weakly decayed logs. The highest number of fruiting species was found on intermediately decayed logs and on logs lying in contact with the ground. The fungal gradient as revealed in a DCA ordination was primarily related to decay. A successional pathway based on the primary decayer Fomitopsis pinicola was not detected.     

The saproxylic beetle assemblage associated with different host trees in Southwest China

Dead wood is a habitat for many insects and other small animals, some of which may be rare or endangered and in need of effective protection. In this paper, saproxylic beetle assemblages associated with different host trees in the subtropical forests in southwestern China were investigated. A total of 277 species (1 439 specimens) in 36 beetle families were collected from 117 dead wood samples, of which 101 samples were identified and respectively belonged to 12 tree genera. The number of saproxylic beetle species varied greatly among logs of different tree genera, with the highest diversity on logs of Juglans. Generally, broad‐leaved trees had a higher richness and abundance of saproxylic species than coniferous trees. Cluster analysis revealed that assemblages from broad‐leaved tree genera were generally similar (except for Betula) and assemblages from coniferous trees formed another distinct cluster. The subsequent indicator analysis proposed that there are different characteristic species for different cluster groups of host tree genera. In our study, log diameter has no positive influence on beetle species density. Conversely, comparisons of individual‐based rarefaction curves suggested that beetle species richness was highest in the small diameter class both in coniferous and broad‐leaved tree genera. With increased wood decay, proportion of habitat specialists (saproxylic beetles living on one tree genus) decreased, whereas proportion of habitat generalists (living on more than three tree genera) increased. The beetle species density was found to be higher in early stages, and decreased in later stages as well. A negative influence of altitude on saproxylic beetle species richness and abundance was detected. It was indicated that different tree genera and altitudes possibly display cross effects in modulating the altitudinal distribution and host preference of the beetles.     

Abundance and viability of fungal spores along a forestry gradient – responses to habitat loss and isolation?

Regional variation in spore deposition and viability was studied for two fungi, Fomitopsis rosea (Alb. & Schwein.: Fr.) P. Karst. and Phlebia centrifuga P. Karst., both confined to old‐growth spruce forests in the boreal zone. Seven regions in Sweden were studied along a north‐south transect in which the historical impact from forestry increases and the amount old forests decreases towards the south. The two southernmost regions were located outside the distribution border of the species. Spore deposition was measured species specifically as heterokaryotisation of homokaryotic mycelia growing on wood discs. There was a significant decline in spore deposition towards the south for both species. F. rosea deposited an average amount of 111 spores m^?2 24 h^?1 in the northernmost region compared to less than 1 spore in the four southernmost regions. The corresponding values for P. centrifuga were 27 spores m^?2 24 h^?1 in the north compared to less than 2 spores in the 4 southernmost regions. No deposition was found south of the distribution borders. The viability of spores from local populations within each region was measured as germination success on nutrient media. Individual fruiting bodies from large populations in the north generally produced spores with higher germinability than fruiting bodies from geographically isolated populations in the central and southern regions. However, there was a high variation among the southern populations. Our data suggest that some populations in mid‐ and south Sweden may suffer from negative genetic effects, possibly associated with fragmentation and loss of habitat. Thus, the combination of low spore deposition and low germinability of spores may be a threat to the long‐term persistence of F. rosea and P. centrifuga in southern Sweden. Several other species may experience the same situation, especially when considering the severe decline of dead wood in Swedish forests.     

Habitat models of wood-inhabiting fungi along a decay gradient of Norway spruce logs

Information on the habitat requirements of wood-inhabiting fungi is needed to understand the factors that affect their diversity. We applied culture-free DNA extraction and 454-pyrosequencing to study the mycobiota of decaying Norway spruce (Picea abies) logs in five unmanaged boreal forests. Fungal habitat preferences in respect of wood density gradient were then estimated with generalized additive mixed models. Fungal diversity and wood density were inversely related, i.e., OTU richness generally increased as the log became increasingly decomposed. White-rot fungi (e.g., Phellinus nigrolimitatus) and members of Hyphodontia did not show a clear response to the wood-density gradient, whereas abundance of Phellinus viticola and brown-rot fungi (e.g., Fomitopsis pinicola, Antrodia serialis, Coniophora olivaceae) peaked during intermediate decay and mycorrhizal fungi (e.g., Piloderma, Tylospora, Russula) increased in the later stages. This information on fungal habitat requirements facilitates the development of management practices that preserve fungal diversity in managed forests.     

The importance of substrate type, shading and scorching for the attractiveness of dead wood to saproxylic beetles

Modern forestry management has reduced the amount of dead wood in forest ecosystems and this has become a serious threat to flora and fauna. Efforts are therefore being made to reverse this trend but one problem is that we still lack detailed knowledge regarding the substrate requirements of many saproxylic species. In a field experiment, conducted in three forest types (forest reserve, mature managed forest and clear-cut), we evaluated the value, from a conservation perspective, of different substrate types (logs, snags and tops) of Norway Spruce, Picea abies, and if the quality of spruce logs as saproxylic habitats can be improved by simple log treatments (scorching and shading). We collected 9982 individuals representing 262 saproxylic beetle species in window traps. Both substrate type and, to a lesser extent, log treatment had a significant effect on the abundance and species richness of saproxylic beetles attracted to the different dead wood substrates. However, more importantly, the composition of the beetle assemblages differed significantly between both substrates and log treatments. Snags, logs and tops all attracted significantly different beetle assemblages and scorched logs differed from untreated control logs. Sixteen red listed species were trapped, with the highest number (11 species) being found on scorched logs. We found strong evidence that some species preferred a specific substrate type, mainly logs, in some cases treated logs (scorched or shaded), but not snags, the substrate commonly provided for conservation purposes on e.g. clear-cuts. This stresses the importance of conducting forestry in such a way that a multitude of both forest habitats and dead wood substrates are available continuously in the forest landscape to maintain biodiversity.     

Spore dispersal of a resupinate ectomycorrhizal fungus, Tomentella sublilacina, via soil food webs

Patterns of fungal spore dispersal affect gene flow, population structure and fungal community structure. Many Basidiomycota produce resupinate (crust-like) basidiocarps buried in the soil. Although spores are actively discharged, they often do not appear to be well positioned for aerial dispersal. We investigated the potential spore dispersal mechanisms of one exemplar of this growth form, Tomentella sublilacina. It is a widespread ectomycorrhizal fungus that sporulates in the soil organic horizon, can establish from the spore bank shortly after disturbance, but also can be a dominant species in mature forest stands. We investigated whether its spores could be dispersed via spore-based food webs. We examined external surfaces, gut contents and feces from arthropod fungivores (mites, springtails, millipedes, beetles, fly larvae) and arthropod and vertebrate predators (centipedes, salamanders) from on and around T. sublilacina sporocarps. Spore densities were high in the guts of many individuals from all fungivore groups. Centipede gut contents, centipede feces and salamander feces contained undigested invertebrate exoskeletons and many apparently intact spores. DAPI staining of spores from feces of fungivores indicated that 7-73% of spores contained intact nuclei, whereas spores from predators had lower percentages of intact nuclei. The spiny spores often were lodged on invertebrate exoskeletons. To test the viability of spores that had passed through invertebrate guts we used fecal droppings of the millipede Harpaphe haydeniana to successfully inoculate seedlings of Pinus muricata (Bishop pine). These results indicate the potential for T. sublilacina spore dispersal via invertebrates and their predators in soil food webs and might help to explain the widespread distribution of this species. It is likely that this is a general mechanism of dispersal for fungi producing resupinate sporocarps, indicating a need to develop a fuller understanding of the linkages of soil food webs and spore dispersal.     

Colonisation of ephemeral forest habitats by specialised species: beetles and bugs associated with recently dead aspen wood

The most appropriate strategy for preserving fragmented populations depends on a species’ ability to colonise distant habitat patches. Insects associated with early decay stages of dead wood are expected to have a high capacity to colonise new habitat patches. To study the dispersal ranges of beetles (Coleoptera) and flat bugs (Hemiptera: Aradidae) dependent on recently dead aspen (Populus tremula) wood in Finland, we set out 58 piles of recently cut aspen logs at various distances up to 1.6 km from forests that contained a high density of old aspen trees. We captured insects by trunk window-traps, and counted beetles’ exit holes. Habitat connectivity was measured in terms of the amount of suitable aspen-wood in the surrounding environment, with the closest dead wood items up-weighted by a negative-exponential function. The log-piles attracted many saproxylic insects including four red-listed aspen-specialist species. The exposure of log-piles to the sun, and high levels of habitat connectivity increased the species richness of aspen-specialists, whereas bark peeling by moose decreased richness. The spatial scale at which species richness had its strongest response to habitat was 93 m. Among individual species there was a wide variability in spatial scale of response. This study supports the view that conservation efforts in boreal forests should be concentrated on sites where colonisation by target species is most likely. Restoration of habitat by re-locating logs may be useful at localities with a rich and specialised fauna but which have too low rate of formation of dead wood by natural processes.     

Metabolic activities and ultrastructure imaging at late-stage of wood decomposition in interactive brown rot - white rot fungal combinations

Basidiomycota brown rot fungus (Fomitopsis pinicola) and two white rot fungi (Phlebia radiata, Trichaptum abietinum) were cultivated on thin slices of spruce wood individually and in interspecies combinations. Within 12 months, F. pinicola substantially decomposed spruce wood observed as mass loss, also in three-species combinations. However, white rot fungi through hyphal interactions negatively affected the brown-rot indicative iron reduction capacity of F. pinicola. Decay-signature gene expression in mycelial interaction zones indicated suppression of brown rot mechanism but stimulation of enzymatic white-rot lignin attack by P. radiata. Wood ultrastructure imaging showed white-rot dominance in the fungal combinations, whereas destructive brown-rot was evident with F. pinicola alone. Our results confirm the dynamic pattern of enzyme production in fungal combinations, and transition from brown to white rot decomposition metabolism during the late stage of wood decay after one year of interspecific interactions.     

Forest microsite effects on community composition of ectomycorrhizal fungi on seedlings of Picea abies and Betula pendula

Niche differentiation in soil horizons, host species and natural nutrient gradients contribute to the high diversity of ectomycorrhizal fungi in boreal forests. This study aims at documenting the diversity and community composition of ectomycorrhizal fungi of Norway spruce ( Picea abies ) and silver birch ( Betula pendula ) seedlings in five most abundant microsites in three Estonian old-growth forests. Undisturbed forest floor, windthrow mounds and pits harboured more species than brown- and white-rotted wood. Several species of ectomycorrhizal fungi were differentially represented on either hosts, microsites and sites. Generally, the most frequent species in dead wood were also common in forest floor soil. Ordination analyses suggested that decay type determined the composition of EcM fungal community in dead wood. Root connections with in-growing mature tree roots from below affected the occurrence of certain fungal species on seedling roots systems in dead wood. This study demonstrates that ectomycorrhizal fungi differentially establish in certain forest microsites that is attributable to their dispersal and competitive abilities. Elevated microsites, especially decayed wood, act as seed beds for both ectomycorrhizal forest trees and fungi, thus affecting the succession of boreal forest ecosystems.     

Diversity and ecology of aphyllophoroid fungi on driftwood logs on the shores of the Baltic Sea

Driftwood is woody debris that is floating on the sea or brought onto the shore by the sea. It can have a natural origin but often it originates from human activities. Driftwood has a significant ecological role as a microhabitat for a large range of species. Dead-wood-associated aphyllophoroid fungi on driftwood have been studied rather little globally, and there are hardly any studies conducted in the Baltic Sea. We studied the diversity and ecology of polypores and corticioids growing on driftwood (man-made logs) on the shores of the Finnish SW-archipelago. In total, 394 driftwood logs were surveyed for visible fungal sporocarps. We found altogether 145 species (1023 records) of which approximately three-fourths were corticioids. The driftwood logs hosted several rare and noteworthy species, such as one new corticioid species, 16 nationally rare species, and 40 new species to the region. The five most common species accounted for one-third of all observations, whereas 82 species (11% of all records) were recorded only once or twice. Larger logs hosted, on average, more species compared to small-diameter logs. The mean number of species increased significantly when comparing the logs in the first and the middle stages of decay. Our results show that driftwood logs can host a wide variety of species and they provide an important substrate for many dead-wood-associated fungi, including species of conservation concern and species with restricted ecological requirements. Driftwood hosted several species that have not been previously found in the archipelago forests, and the logs clearly enrich the fungal diversity in the area. Our results encourage to increase dead wood for fungi in coastal forests where it has been dramatically decreased as a result of strong hemeroby.     

Linking taxonomical and functional biodiversity of saproxylic fungi and beetles in broad‐leaved forests in southern Italy with varying management histories

Abstract

The fundamental ecological significance of deadwood decomposition in forests has been highlighted in several reviews, some conclusions regarding silviculture being drawn. Old‐growth forests are natural centres of biodiversity. Saproxylic fungi and beetles, which are vital components of these ecosystems, occupy a variety of spatial and trophic niches. Fungal and beetle diversity on coarse woody debris (CWD) was analysed in 36 forest sites in the Cilento and Vallo di Diano National Park, Italy. The data were analysed by DCA and Spearman’s rank correlation. The results provide empirical evidence of the existence of a pattern of joint colonization of the woody substrate by fungi and beetles, which includes an assemblage of reciprocal trophic roles within fungal/beetle communities. These organisms act together to form a dynamic taxonomical and functional ecosystem component within the complex set of processes involved in wood decay. The variables most predictive of correlations between management‐related structural attributes and fungal/beetle species richness and their trophic roles for old‐growth forest are: number of logs, number of decay classes and CWD total volume. Deadwood spatio‐temporal continuity should be the main objective of forest planning to stop the loss of saproxylic fungal and insect biodiversity.     

Dispersal may limit the occurrence of specialist wood decay fungi already at small spatial scales

We used species‐specific spore traps to measure airborne dispersal of the wood decay fungus Phlebia centrifuga (spore size 6.5–9 × 2.5–3 μm) up to 1000 m distance from a point source. We fitted two simple dispersal models, an empirical power law model and a semi‐mechanistic diffusion model to the data using the Bayesian approach. The diffusion model provided a better fit than the power law model which underestimated deposition at 3–55 m and overestimated deposition at longer and shorter distances. Model fit improved by allowing overdispersion, suggesting that spores are not dispersed independently but wind can transport spores in groups inside discrete air packages up to considerable distances. Using the fitted diffusion model and available information on the establishment rates of wood‐decay fungi, we examine the distance up to which colonisation from a single fruit body is likely to occur. We conclude that the diluting effect of distance and low establishment success make the occurrence of P. centrifuga dispersal limited possibly already at the distance of tens of metres and very probably at a few hundred metres from the nearest fruit body, despite the fact that under favourable conditions a high proportion of the spores can disperse considerably further. This conclusion is likely to hold generally for those fungal species that inhabit fragmented landscapes, have specialised resource and habitat requirements, and have similar spore size and other dispersal traits as P. centrifuga.     

Trichostomopsis aaronis (Lor.) Agnew & Townsend new to the U.S.S.R.

Abstract

Occurrence of Ptilidium pulcherrimum in transects and spore dispersal from a single colony have been studied in a coastal spruce forest in northern Sweden. The main substrate type was rotting wood with 75% of all occurrences. Annual spore production was 68,500 spores/m² forest, 640,000 spores/m² substrate and 44,000,000 spores/m² colony. Almost 50% of the spores were deposited within 2.5 m of the colony. Annual spore deposition between colonies was estimated to be between 24,000–39,000 and deposition on the main substrate, decaying logs, was about 340–600 spores/m² forest. P. pulcherrimum showed a clumped distribution pattern up to about a 15 m neighbourhood distance. This pattern could not be explained by a similar clumping of the substrate. Instead a limitation by distance in establishment due to a deficit of spores is assumed.