Variation in terricolous bryophyte and macrolichen vegetation along primary gradients in Canadian boreal forests

Abstract. Compositional variation among 172 boreal forests stands, based upon the understorey terricolous bryophyte and macrolichen vegetation, is compared with that based upon vascular vegetation. Detrended correspondence analysis (DCA) of the two datasets yielded stand ordinations each indicating a single dominant upland-bottomland gradient. This was most clear among 138 stands in the cryptogam dataset. Canonical correspondence analysis (CCA) of the 138 stand subset yielded a pronounced first axis gradient from dry, nutrient poor pine dominated forest sites to moist, nutrient rich bottomland sites dominated by balsam poplar woodland. Individual species response curves, by the method of log-linear least squares regression, yielded three different respective patterns among the most abundant tree, understorey vascular and cryptogamic plant species. Whereas curves for the tree species were ‘Gaussian’ in shape and displaced at somewhat regular intervals along the gradient, the abundant understorey vascular plant species curves clustered in the mesic, mixed wood region two thirds of the way along. Curves for the abundant cryptogams were mostly linear in shape and absent from the mixed wood zone, showing concentrations at one end of the gradient or the other. Two interpretations of the understorey pattern are considered. One proposes competitive exclusion of cryptogams from the mesic mixed wood region by the vascular understorey plants. Another proposes that the cryptogams are adapted to the sharply contrasting ecosystems encountered at one end of the gradient or another but that none can effectively cope with the intermediate, mixed wood ecosystems. Evidence from the response curves favours this second interpretation.     

Diversity and species distribution of ectomycorrhizal fungi along productivity gradients of a southern boreal forest

Coniferous forests with diverse ectomycorrhizal fungus (EMF) communities are associated with nutrient-poor, acidic soils but there is some debate whether EMF can be equally adapted to more productive, nitrogen-rich sites. We compared EMF species distribution and diversity along a replicated productivity gradient in a southern boreal forest of British Columbia (Canada). Roots from subalpine fir (Abies lasiocarpa) saplings of the understory were sampled and EMF species were identified by morphotypes supplemented with ITS rDNA analysis. There were significant changes in the distribution and abundance of 74 EMF species along the productivity gradient, with as little as 24% community similarity among contrasting sites. Species richness per plot increased asymptotically with foliar nitrogen concentrations of subalpine fir, demonstrating that many EMF species were well suited to soils with high rates of nitrogen mineralization. EMF species abundance in relation to site productivity included parabolic, negative linear, and positive exponential curves. Both multi-site and more narrowly distributed EMF were documented, and a diverse mix of mantle exploration types was present across the entire productivity gradient. The results demonstrate strong associations of EMF fungal species with edaphic characteristics, especially nitrogen availability, and a specialization in EMF communities that may contribute to the successful exploitation of such contrasting extremes in soil fertility by a single tree host.     

Deadwood-rich managed forests provide insights into the old-forest association of wood-inhabiting fungi

A major question in fungal conservation is why many species are confined to old forests, and how they could be supported by contemporary landscape matrix. Specifically, forestry that retains large biological legacies across landscape could reduce old-forest dependencies to species that require unusual substrate conditions. We sampled polypores in 112 2 ha plots (both old and harvested stands) in a semi-natural forestry context in Estonia and modelled the habitat factors of species confined to old growth. The results confirmed that old-growth assemblages emerged mostly due to diverse and abundant substrate supply (notably downed CWD). Only 10 species (five spruce-dwellers) were confined to old growth; of these, only Fomitopsis rosea and Oxyporus corticola were additionally affected by forest connectivity. The forestry system studied appeared particularly favourable for the species inhabiting deciduous wood. To better address habitat degradation in conservation, expert lists of ‘old-forest (indicator) fungi’ should be replaced with evidence-based focal taxa.     

Saprotrophic and ectomycorrhizal fungi exhibit contrasting richness patterns along elevational gradients in cool-temperate montane forests

Increasing evidence suggests that elevational gradients of soil fungal richness are highly variable, but few studies have examined how diversity components of each guild contribute to overall fungal diversity. Here, we aimed to disentangle the relationships between total, saprotrophic, and ectomycorrhizal fungal richness, and environmental factors across elevational gradients in cool-temperate montane forests. We observed that total and saprotrophic richness decreased but ectomycorrhizal richness increased with increase in elevation. Elevational range size and nestedness analyses illustrated that saprotrophic and ectomycorrhizal source communities were located at lower and higher elevations, respectively. The observed total and saprotrophic richness were directly influenced by soil properties and indirectly influenced by climate and plant communities. Ectomycorrhizal fungal richness was affected by climate and the dominance of ectomycorrhizal trees. We highlight that two directional source–sink dynamics lead to opposite elevational patterns between saprotrophic and ectomycorrhizal fungal richness, shaping the variation in elevational richness gradients.     

Biodiversity of wood-inhabiting fungi in a boreal coniferous forest in Ser-Trendelag County, Central Norway

A survey of the patterns of lignicolous fungi to record occurrence of fruitbodies on fallen logs of Picea abies . has been performed. Ninetythree forest stands with different ages. ecology, and histories of management in Ser-Trendelag County, Central Norway were investigated. In each stand an investigation plot of 40 m diameter was established, in which five logs were randomly selected. Environmental variables of the logs and plots and geographical positions of the plots were recorded. The numerical methods used include detrended correspondence analysis (DCA) and canonical correspondence analysis (CCA).
A total of 140 species of Aphyllophorales, Agaricales with eccentrically attached caps, heterobasiodomycetes, and prominent ascomycetes and deuteromycetes were registered. More than one third of the species were only found once. The main gradient in the data set was shown to be related to decay. The second gradient was related to size of the logs and some forest management variables (in addition to decay). Of the variation in composition of the species, 13.8 % can be ascribed to solely environmental variance, 6 % to solely spatial structure, and only 0.8 % to the combination of the environment and spatial component of variation. The results are discussed in the light of wood decay, forest management, and landscape fragmentation.     

Impact of cervid browsing and grazing on the terrestrial gastropod fauna in the boreal forests of Fennoscandia

Selective foraging by cervids can affect vegetation, and that in turn can have far-reaching effects on various components of the boreal forest ecosystem. Since terrestrial gastropods are sensitive to habitat alterations, they can be expected to respond to changes caused by grazing. This study is based on gastropod specimens from two data sets that were collected with pitfall traps in order to study the effects of moose and reindeer on ground-layer invertebrates. Invertebrates were trapped from 23 pairs of plots, where one plot was open to all animals while the other one was fenced to exclude large mammals. Half of the sites were in Finnish Lapland, where reindeer grazing was the main cause of differences between the plots; the rest were located in central Sweden and southern Finland, where moose was the most important cervid grazer. The composition of the gastropod fauna differed between grazed and ungrazed plots in both areas, and the abundance of gastropods in general and that of many species was higher in ungrazed plots (the number of gastropods caught was 17% lower in grazed than in ungrazed plots in moose sites, and 24% smaller in reindeer sites). Species richness differed significantly between grazed and ungrazed plots in the combined data of both site types, and was 15% higher in ungrazed plots. In moose sites the relative diversity (H′/H′_max) of gastropods was 17% higher in grazed plots. In reindeer sites, the means of richness and diversity (H′) were considerably higher in ungrazed plots (35% and 40% respectively) but the differences were only marginally significant. By indirectly modifying the microclimate within the forest, cervids fulfil the criteria for a ‘physical ecosystem engineer’. Since the population density and distribution of terrestrial gastropods depend largely on the physical conditions of the microhabitat, the engineering effects of cervids are the most likely cause of the changes observed in the density and composition of the gastropod fauna in the boreal forest.     

Communities of wood-inhabiting fungi in dead pine logs along a geographical gradient in Japan

Fungi are the main agents of coarse woody debris decomposition in forest ecosystems. We examined the associations of environmental variables with fungal community structures in dead pine logs at 12 geographically distant sites using amplicon pyrosequencing of fungal ITS rDNA. A total of 575 operational taxonomic units (OTUs) were identified based on clustering at 97% similarity. Among the known fungal ecological groups, saprotrophic fungi generally showed highest frequency of occurrence and were positively associated with mean annual temperature (MAT) and log diameter. Wood decay fungi with unknown decay type were positively associated with pine wilt disease and negatively associated with log diameter. Ordination analysis of the 42 most prevalent OTUs showed that MAT and annual precipitation significantly explained the observed fungal community structure. These results suggested that climate conditions and site history differentially effect structure fungal communities in pine logs among different ecological groups.     

Stand-replacing wildfires alter the community structure of wood-inhabiting fungi in southwestern ponderosa pine forests of the USA

Increases in stand-replacing wildfires in the western USA have widespread implications for ecosystem carbon (C) cycling, in part because the decomposition of trees killed by fire can be a long-term source of CO₂ to the atmosphere. Knowledge of the composition and function of decay fungi communities may be important to understanding how wildfire alters C cycles. We assessed the effects of stand-replacing wildfires on the community structure of wood-inhabiting fungi along a 32-yr wildfire chronosequence. Fire was associated with low species richness for up to 4 yr and altered species composition relative to unburned forest for the length of the chronosequence. A laboratory incubation demonstrated that species varied in their capacity to decompose wood; Hypocrea lixii, an indicator of the most recent burn, caused the lowest decomposition rate. Our results show that stand-replacing wildfires have long-term effects on fungal communities, which may have consequences for wood decomposition and C cycling.     

Simulating forest succession along ecological gradients in southern Central Europe

Forest stand development was simulated using a forest succession model of the JABOWA/FORET type. The environmental conditions are representative for a wide spectrum of Swiss forest sites ranging from 220 m to 1 700 m a.s.l. Each model run covers a period of 1 200 yr and is based on the averaged successional characteristics of 50 forest plots with an individual size of 1/12 ha. These small forest plots serve as basic units to simulate establishment, growth, and death of individual trees of 29 species. Existing light in the forest stand, climatic conditions, soil properties, and other environmental factors control the growth of each individual tree. Compared with previous simulation studies, some major modifications were made, including the incorporation of the indicator values of Ellenberg (1979) to describe the ecophysiological behaviour of the species considered. As a test, the simulated species composition through time was compared with the actual vegetation and the potentially natural species composition on the corresponding site types. The extensive comparison revealed that approximately 80% of the simulations match the expected species configurations. Thus, it was concluded that the model is valid for the purpose of evaluating impacts of natural and human disturbances on forest communities.     

Belowground interspecific competition in mixed boreal forests: fine root and ectomycorrhiza characteristics along stand developmental stage and soil fertility gradients

We studied fine roots and ectomycorrhizas in relation to aboveground tree and stand characteristics in five mixed Betula pendula Roth, Picea abies (L.) H. Karst., and Pinus sylvestris L. stands in Southern Finland. The stands formed gradients of developmental stage (15-, 30-, and 50-year-old stands) in the stands of medium fertility, and of site fertility in the young stands (30-year-old fertile, medium fertile, and least fertile stands). The biomass of the external hyphae of ectomycorrhizas (ECM) was the highest, and the diversity of the fungal community the lowest, in the most fertile stand. The vertical distributions of fine roots of the three tree species were mostly overlapping, indicating high inter-specific belowground competition in the stands. We did not find any clear trends in the fine root biomass (FRB) or length across the stand developmental stages. The FRB of the conifers varied with site fertility, whereas in B. pendula it was almost constant. In contrast to the conifers, the specific root length (SRL) of B. pendula clearly increased from the most fertile to the least fertile stand. This indicates differences in the primary nutrient acquisition strategy between conifers and B. pendula.     

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.     

Species associations during the succession of wood-inhabiting fungal communities

We studied fungal succession in decaying wood by compiling time-series data of fruit body observations. We tested the hypothesis that the presence of a primary species affects the probability of a succeeding species occurring later on the same log. Significant associations were detected for 15 species pairs; these were consistent with earlier findings on co-occurrence patterns in single time surveys. We used enrichment analysis to test if species with particular life-history attributes were more often associated with the occurrence of a succeeding species, or vice versa. White rot fungi and fungi abundant as mycelia were more often associated with the occurrence of succeeding species, compared to brown rot fungi and species with low mycelial abundance. Our results indicate that certain primary species cause priority effects and non-random co-occurrence patterns in the field. These successional patterns are likely to be connected both with substrate modification and species interactions.     

Plant diversity changes and succession along resource availability and disturbance gradients in Kamchatka

Changes in plant species richness across environmental and temporal gradients have often been explained by the intermediate disturbance hypothesis and a unimodal diversity–productivity relationship. We tested these predictions using two sets of mountain plant communities assembled along postglacial successional and snow depth (disturbance and stress) gradients in maritime Kamchatka. In each community, we counted the number of species in plots of increasing sizes (0.0025–100 m²) and analyzed them using species–area curves fitted by the Arrhenius power function and the Gleason logarithmic function. A comparison of successional communities along a 270-year-old moraine chronosequence behind the receding Koryto Glacier—representing gradients of increasing productivity and resource competition—confirmed the unimodal species richness pattern. The plant diversity peaked in a 60–80-year-old Salix–Alnus stand where light availability was sufficient to sustain a rich understory combining pioneer and late successional herbs. The closed Alnus canopy on older moraines caused a pronounced decrease in species richness for all plot sizes (interactive stage 80–120 years since deglaciation). A slight increase in species richness in the oldest assortative stages (120–270 years), when Alnus stands are mature, was found only at the smaller spatial scales. This reflects (i) the consolidation of clonal understory dominants and (ii) the absence of other woody species such as Betula ermanii whose invasion would eliminate Alnus and increase diversity at larger spatial scales. A comparative study of major mountain plant communities distributed above the Koryto Glacier foreland did not confirm the highest species richness at intermediate levels of disturbance and stress. Contrary to our expectation, the species richness was highest in alpine tundra and snowbed communities, which are subjected to severe winter frost and a short summer season, while less disturbed communities of subalpine meadows, heaths, and Betula ermanii woods were less species-rich. We attribute this pattern to differences in habitat area and species pool size.     

Variations of root and heterotrophic respiration along environmental gradients in China's forests

Aims Root and heterotrophic respiration may respond differently to environmental variability, but little evidence is available from large-scale observations. Here we aimed to examine variations of root and heterotrophic respiration across broad geographic, climatic, soil and biotic gradients.Methods We conducted a synthesis of 59 field measurements on root and heterotrophic respiration across China's forests.Important findings Root and heterotrophic respiration varied differently with forest types, of which evergreen broadleaf forest was significantly different from those in other forest types on heterotrophic respiration but without statistically significant differences on root respiration. The results also indicated that root and heterotrophic respiration exhibited similar trends along gradients of precipitation, soil organic carbon and satellite-indicated vegetation growth. However, they exhibited different relationships with temperature: root respiration exhibited bimodal patterns along the temperature gradient, while heterotrophic respiration increased monotonically with temperature. Moreover, they showed different relationships with MOD17 GPP, with increasing trend observed for root respiration whereas insignificant change for heterotrophic respiration. In addition, root and heterotrophic respiration exhibited different changes along the age sequence, with insignificant change for root respiration and decreasing trend for heterotrophic respiration. Overall, these results suggest that root and heterotrophic respiration may respond differently to environmental variability. Our findings could advance our understanding on the different environmental controls of root and heterotrophic respiration and also improve our ability to predict soil CO₂ flux under a changing environment.     

The effects of green tree retention and subsequent prescribed burning on ground beetles (Coleoptera: Carabidae) in boreal pine-dominated forests

We studied how two methods to promote biodiversity in managed forests, i.e. green tree retention and prescribed fire, affect the assemblages of carabid beetles. Our experiment consisted of 24 study sites, each 3–5 ha in size, which had been prepared according to factorial design. Each of the eight treatment combinations determined by the two factors explored – tree retention level (0, 10, 50 m³/ha^?1 and uncut controls) and prescribed use of fire (yes/no) – was replicated three times. We sampled carabids using pitfall traps one year after the treatments. Significantly more individuals were caught in most of the burned sites, but this difference was partially reflective of the trap‐catches of Pterostichus adstrictus. The fire did not increase no. of P. adstrictus in the uncut sites as much as in the other sites. Species richness was significantly affected by both factors, being higher in the burned than in the unburned sites and in the harvested than in the unharvested sites. Many species were concentrated in the groups of retention trees in the burned sites, but only a few were in the unburned sites. The species turnover was greater in the burned than in the unburned sites, as indicated by the NMDS ordinations. Greater numbers of smaller sized species and proportion of brachypterous species were present in the burned sites. Fire‐favored species, and also the majority of other species that prefer open habitats were more abundantly caught in the burned sites than in the unburned sites. Dead wood or logging waste around the traps did not correlate with the occurrence of species. We conclude that carabids are well adapted to disturbances, and that frequent use of prescribed fire is essential for the maintenance of natural assemblages of carabid beetles in the boreal forest. Small retention tree groups can not maintain assemblages of uncut forest, but they can be important by providing food, shelter and breeding sites for many species, particularly in the burned sites.     

Diversity and abundance of resupinate thelephoroid fungi as ectomycorrhizal symbionts in Swedish boreal forests

Resupinate thelephoroid fungi (hereafter called tomentelloid fungi) have a world-wide distribution and comprise approximately 70 basidiomycete species with inconspicuous, resupinate sporocarps. It is only recently that their ability to form ectomycorrhizas (EM) has been realized, so their distribution, abundance and significance as mycobionts in forest ecosystems is still largely unexplored. In order to provide baseline data for future ecological studies of tomentelloid fungi, we explored their presence and abundance in nine Swedish boreal forests in which the EM communities had been analysed. Phylogenetic analyses were used to compare the internal transcribed spacer of nuclear ribosomal DNA (ITS rDNA) sequence data obtained from mycobionts on single ectomycorrhizal tips with that obtained from sporocarps of identified tomentelloid fungi. Five species of Tomentella and one species of Pseudotomentella were identified as ectomycorrhizal fungi. The symbiotic nature of Tomentella bryophila, T. stuposa, T. badia and T. atramentaria is demonstrated for the first time. T. stuposa and Pseudotomentella tristis were the most commonly encountered tomentelloid fungi, with the other species, including T. sublilacina, only being recorded from single stands. Overall, tomentelloid fungi were found in five of the studies, colonizing between 1 and 8% of the mycorrhizal root tips. Two of the five sites supported several tomentelloid species. Tomentelloid fungi appear to be relatively common ectomycorrhizal symbionts with a wide distribution in Swedish coniferous forests. The results are in accordance with accumulating data that fungal species which lack conspicuous sporocarps may be of considerable importance in EM communities.     

Niche modelling predicts that soil fungi occupy a precarious climate in boreal forests

Aim

Efforts to predict the responses of soil fungal communities to climate change are hindered by limited information on how fungal niches are distributed across environmental hyperspace. We predict the climate sensitivity of North American soil fungal assemblage composition by modelling the ecological niches of several thousand fungal species.

Location

One hundred and thirteen sites in the United States and Canada spanning all biomes except tropical rain forest.

Major Taxa Studied

Fungi.

Time Period

2011–2018.

Methods

We combine internal transcribed spacer (ITS) sequences from two continental-scale sampling networks in North America and cluster them into operational taxonomic units (OTUs) at 97% similarity. Using climate and soil data, we fit ecological niche models (ENMs) based on logistic ridge regression for all OTUs present in at least 10 sites (n = 8597). To describe the compositional turnover of soil fungal assemblages over climatic gradients, we introduce a novel niche-based metric of climate sensitivity, the Sørensen climate sensitivity index. Finally, we map climate sensitivity across North America.

Results

ENMs have a mean out-of-sample predictive accuracy of 73.8%, with temperature variables being strong predictors of fungal distributions. Soil fungal climate niches clump together across environmental space, which suggests common physiological limits and predicts abrupt changes in composition with respect to changes in climate. Soil fungi in North American climates are more likely to be limited by cold and dry conditions than by warm and wet conditions, and ectomycorrhizal fungi generally tolerate colder temperatures than saprotrophic fungi. Sørensen climate sensitivity exhibits a multimodal distribution across environmental space, with a peak in climates corresponding to boreal forests.

Main Conclusions

The boreal forest occupies an especially precarious region of environmental space for the composition of soil fungal assemblages in North America, as even small degrees of warming could trigger large compositional changes characterized mainly by an influx of warm-adapted species.     

木材腐朽菌在森林生态系统中的功能

木材腐朽菌是森林生态系统的重要组成部分,在森林生态系统中起着极为重要的降解还原作用,主要包括担子菌门非褶菌目、子囊菌门盘菌纲和半知菌类的部分真菌,能全部或部分降解木材中的木质素、纤维素和半纤维素,其降解机制有3种：白色腐朽、褐色腐朽和软腐朽．木材腐朽菌与生态系统中其它生物关系密切,为很多昆虫、鸟类提供营养,有些昆虫也能使木腐菌得到传播．保护木材腐朽菌的生物多样性是保护森林生态系统、维护生态系统健康的重要因素．