Phylogenetic diversity of macromycetes and woody plants along an elevational gradient in Eastern Mexico

Phylogenetic information provides insight into the ecological and evolutionary processes that organize species assemblages. We compared patterns of phylogenetic diversity among macromycete and woody plant communities along a steep elevational gradient in eastern Mexico to better understand the evolutionary processes that structure their communities. Macrofungi and trees were counted and identified in eight sites from 100 to 3500 m asl, and sequence data retrieved from GenBank for the same or closely related species were used to reconstruct their phylogenies. Patterns of species richness and phylogenetic diversity were similar for both macrofungi and trees, but macromycete richness and diversity peaked at mid‐elevations, whereas woody plant richness and diversity did not show significant trends with elevation. Phylogenetic similarity among sites was low for both groups and decreased as elevational distance between sites increased. Macromycete communities displayed phylogenetic overdispersion at low elevations and phylogenetic clustering at high elevations; the latter is consistent with environmental filtering at high elevation sites. Woody plants generally exhibited phylogenetic clustering, consistent with the potential importance of environmental filtering throughout the elevational gradient.     

Fungal community composition in neotropical rain forests: the influence of tree diversity and precipitation

Plant diversity is considered one factor structuring soil fungal communities because the diversity of compounds in leaf litter might determine the extent of resource heterogeneity for decomposer communities. Lowland tropical rain forests have the highest plant diversity per area of any biome. Since fungi are responsible for much of the decomposition occurring in forest soils, understanding the factors that structure fungi in tropical forests may provide valuable insight for predicting changes in global carbon and nitrogen fluxes. To test the role of plant diversity in shaping fungal community structure and function, soil (0-20?cm) and leaf litter (O horizons) were collected from six established 1-ha forest census plots across a natural plant diversity gradient on the Isthmus of Panama. We used 454 pyrosequencing and phospholipid fatty acid analysis to evaluate correlations between microbial community composition, precipitation, soil nutrients, and plant richness. In soil, the number of fungal taxa increased significantly with increasing mean annual precipitation, but not with plant richness. There were no correlations between fungal communities in leaf litter and plant diversity or precipitation, and fungal communities were found to be compositionally distinct between soil and leaf litter. To directly test for effects of plant species richness on fungal diversity and function, we experimentally re-created litter diversity gradients in litter bags with 1, 25, and 50 species of litter. After 6?months, we found a significant effect of litter diversity on decomposition rate between one and 25 species of leaf litter. However, fungal richness did not track plant species richness. Although studies in a broader range of sites is required, these results suggest that precipitation may be a more important factor than plant diversity or soil nutrient status in structuring tropical forest soil fungal communities.     

Differences in epiphyte biomass and community composition along landscape and within-crown spatial scales

Vascular epiphytes contribute to the structural, compositional, and functional complexity of tropical montane cloud forests because of their high biomass, diversity, and ability to intercept and retain water and nutrients from atmospheric sources. However, human-caused climate change and forest-to-pasture conversion are rapidly altering tropical montane cloud forests. Epiphyte communities may be particularly vulnerable to these changes because of their dependence on direct atmospheric inputs and host trees for survival. In Monteverde, Costa Rica, we measured vascular epiphyte biomass, community composition, and richness at two spatial scales: (1) along an elevation gradient spanning premontane forests to montane cloud forests and (2) within trees along branches from inner to outer crown positions. We also compared epiphyte biomass and distribution at these scales between two different land-cover types, comparing trees in closed canopy forest to isolated trees in pastures. An ordination of epiphyte communities at the level of trees grouped forested sites above versus below the cloud base, and separated forest versus pasture trees. Species richness increased with increasing elevation and decreased from inner to outer branch positions. Although richness did not differ between land-cover types, there were significant differences in community composition. The variability in epiphyte community organization between the two spatial scales and between land-cover types underscores the potential complexity of epiphyte responses to climate and land-cover changes.     

Richness,species composition and functional groups in Agaricomycetes communities along a vegetation and elevational gradient in the Andean Yungas of Argentina

The Neotropics are among the least explored regions from a mycological perspective. A few recent molecular studies in South America have shown high fungal diversity as well as numerous groups of mostly undescribed taxa. Through soil metabarcoding analysis we compared richness and species composition among macrofungal communities, belonging to Agaricales, Russulales, Boletales and Phallomycetidae groups, in three elevational forests types in the subtropical Yungas of Northwestern Argentina (Piedmont forest; Montane forest, Montane cloud forest). The aims of this study were to assess richness of taxonomic and functional groups along the elevation gradient and to assess the relationships between environmental variables and species composition in the studied fungal communities. The results have shown rich Agaricomycetes communities, diversely structured among forests habitats. The elevation gradient differentially affected the richness and distribution of Agaricales, Russulales, Boletales and Phallomycetidae. Based on fungal trophic modes and guilds, the gradient also affected the ectomycorrhizal taxa distribution. When considering the basidiomata growth forms (agaricoid, boletoid, gasteroid, etc.), only the secotioid type showed significant elevational differences. Additional analyses indicated that saprotrophic nutritional mode was dominant along the entire gradient, being partially replaced by biotrophic modes at higher elevations. Fungal communities in the Montane cloud forests are most dissimilar when compared with communities at the Piedmont forest and Montane forest, which is consistent with the different biogeographic origins of these forests. DNA metabarcoding sequence analysis provided detailed information on the diversity and taxonomic and functional composition of macrofungal communities.     

Soil pH determines fungal diversity along an elevation gradient in Southwestern China

Fungi play important roles in ecosystem processes, and the elevational pattern of fungal diversity is still unclear. Here, we examined the diversity of fungi along a 1,000 m elevation gradient on Mount Nadu, Southwestern China. We used MiSeq sequencing to obtain fungal sequences that were clustered into operational taxonomic units(OTUs) and to measure the fungal composition and diversity. Though the species richness and phylogenetic diversity of the fungal community did not exhibit significant trends with increasing altitude, they were significantly lower at mid-altitudinal sites than at the base. The Bray-Curtis distance clustering also showed that the fungal communities varied significantly with altitude. A distance-based linear model multivariate analysis(DistLM) identified that soil pH dominated the explanatory power of the species richness(23.72%),phylogenetic diversity(24.25%) and beta diversity(28.10%) of the fungal community. Moreover, the species richness and phylogenetic diversity of the fungal community increased linearly with increasing soil pH(P0.05). Our study provides evidence that pH is an important predictor of soil fungal diversity along elevation gradients in Southwestern China.     

Amplicon‐pyrosequencing‐based detection of compositional shifts in bryophyte‐associated fungal communities along an elevation gradient

Although bryophytes are a dominant vegetation component of boreal and alpine ecosystems, little is known about their associated fungal communities. HPLC assays of ergosterol (fungal biomass) and amplicon pyrosequencing of the ITS2 region of rDNA were used to investigate how the fungal communities associated with four bryophyte species changed across an elevational gradient transitioning from conifer forest to the low‐alpine. Fungal biomass and OTU richness associated with the four moss hosts did not vary significantly across the gradient (P > 0.05), and both were more strongly affected by host and tissue type. Despite largely constant levels of fungal biomass, distinct shifts in community composition of fungi associated with Hylocomium, Pleurozium and Polytrichum occurred between the elevation zones of the gradient. This likely is a result of influence on fungal communities by major environmental factors such as temperature, directly or indirectly mediated by, or interacting with, the response of other components of the vegetation (i.e. the dominant trees). Fungal communities associated with Dicranum were an exception, exhibiting spatial autocorrelation between plots, and no significant structuring by elevation. Nevertheless, the detection of distinct fungal assemblages associated with a single host growing in different elevation zones along an elevational gradient is of particular relevance in the light of the ongoing changes in vegetation patterns in boreal and alpine systems due to global climate warming.     

The mid-domain effect in ectomycorrhizal fungi: range overlap along an elevation gradient on Mount Fuji,Japan

Mid-domain effect (MDE) models predict that the random placement of species'' ranges within a bounded geographical area leads to increased range overlap and species richness in the center of the bounded area. These models are frequently applied to study species-richness patterns of macroorganisms, but the MDE in relation to microorganisms is poorly understood. In this study, we examined the characteristics of the MDE in richness patterns of ectomycorrhizal (EM) fungi, an ecologically important group of soil symbionts. We conducted intensive soil sampling to investigate overlap among species ranges and the applicability of the MDE to EM fungi in four temperate forest stands along an elevation gradient on Mount Fuji, Japan. Molecular analyses using direct sequencing revealed 302 EM fungal species. Of 73 EM fungal species found in multiple stands, 72 inhabited a continuous range along the elevation gradient. The maximum overlap in species range and the highest species richness occurred at elevations in the middle of the gradient. The observed richness pattern also fit within the 95% confidence interval of the mid-domain null model, supporting the role of the MDE in EM fungal richness. Deviation in observed richness from the mean of the mid-domain null estimation was negatively correlated with some environmental factors, including precipitation and soil C/N, indicating that unexplained richness patterns could be driven by these environmental factors. Our results clearly support the existence of microbial species'' ranges along environmental gradients and the potential applicability of the MDE to better understand microbial diversity patterns.     

Soil propagule banks of ectomycorrhizal fungi share many common species along an elevation gradient

We conducted bioassay experiments to investigate the soil propagule banks of ectomycorrhizal (EM) fungi in old-growth forests along an elevation gradient and compared the elevation pattern with the composition of EM fungi on existing roots in the field. In total, 150 soil cores were collected from three forests on Mt. Ishizuchi, western Japan, and subjected to bioassays using Pinus densiflora and Betula maximowicziana. Using molecular analyses, we recorded 23 EM fungal species in the assayed propagule banks. Eight species (34.8 %) were shared across the three sites, which ranged from a warm–temperate evergreen mixed forest to a subalpine conifer forest. The elevation pattern of the assayed propagule banks differed dramatically from that of EM fungi on existing roots along the same gradient, where only a small proportion of EM fungal species (3.5 %) were shared across sites. The EM fungal species found in the assayed propagule banks included many pioneer fungal species and composition differed significantly from that on existing roots. Furthermore, only 4 of 23 species were shared between the two host species, indicating a strong effect of bioassay host identity in determining the propagule banks of EM fungi. These results imply that the assayed propagule bank is less affected by climate compared to EM fungal communities on existing roots. The dominance of disturbance-dependent fungal species in the assayed propagule banks may result in higher ecosystem resilience to disturbance even in old-growth temperate forests.     

Strong coupling of plant and fungal community structure across western Amazonian rainforests

The Amazon basin harbors a diverse ecological community that has a critical role in the maintenance of the biosphere. Although plant and animal communities have received much attention, basic information is lacking for fungal or prokaryotic communities. This is despite the fact that recent ecological studies have suggested a prominent role for interactions with soil fungi in structuring the diversity and abundance of tropical rainforest trees. In this study, we characterize soil fungal communities across three major tropical forest types in the western Amazon basin (terra firme, seasonally flooded and white sand) using 454 pyrosequencing. Using these data, we examine the relationship between fungal diversity and tree species richness, and between fungal community composition and tree species composition, soil environment and spatial proximity. We find that the fungal community in these ecosystems is diverse, with high degrees of spatial variability related to forest type. We also find strong correlations between α- and β-diversity of soil fungi and trees. Both fungal and plant community β-diversity were also correlated with differences in environmental conditions. The correlation between plant and fungal richness was stronger in fungal lineages known for biotrophic strategies (for example, pathogens, mycorrhizas) compared with a lineage known primarily for saprotrophy (yeasts), suggesting that this coupling is, at least in part, due to direct plant–fungal interactions. These data provide a much-needed look at an understudied dimension of the biota in an important ecosystem and supports the hypothesis that fungal communities are involved in the regulation of tropical tree diversity.     

Fruiting patterns of macrofungi in tropical and temperate land use types in Yunnan Province,China

Despite the important contribution of fungi to forest health, biomass turnover and carbon cycling, little is known about the factors that influence fungal phenology. Therefore, in order to further our understanding on how macrofungal fruiting patterns change along a gradient from temperate to tropical climate zones, we investigated the phenological patterns of macrofungal fruiting at five sites along a combined altitudinal and latitudinal gradient in SW China and NW Laos, ranging from temperate to tropical climates. Observations were conducted in the dominant land use types at these study sites: mixed forest (all sites), coniferous forest (temperate sites) and grassland (temperate sites). In total, 2866 specimens were collected, belonging to 791 morpho species, 162 genera, and 71 families. At the site level, the fruiting of ectomycorrhizal (EcMF) and saprotrophic fungi (SapF) occurred at the same time among all land use types. The fruiting season of fungi in the tropical sites began earlier and ended later compared to that of fungi in the temperate sites, which we attribute mainly to the higher temperature and more abundant rainfall of the tropical areas. EcMF taxa richness in temperate forests (both coniferous and mixed forest) showed a distinct peak at the end of the rainy season in August and September, while no significant peak was observed for SapF taxa richness. Neither functional fungal groups showed significant seasonal fluctuations in tropical areas. The temporal turnover of fungal fruiting significantly increased with the shift from tropical to temperate forests along the elevation gradient. In the grasslands, macrofungal abundance was less than 22% of that of corresponding forest sites, and taxa richness was 42% of that of corresponding forest sites. Fungal fruiting showed no significant fluctuations across the rainy season. This work represents a case study carried out over one year, and further measurements will be needed to test if these results hold true in the longer term.     

Belowground responses to elevation in a changing cloud forest

Few studies have investigated how soil fungal communities respond to elevation, especially within TMCF (tropical montane cloud forests). We used an elevation gradient in a TMCF in Costa Rica to determine how soil properties, processes, and community composition of fungi change in response to elevation and across seasons. As elevation increased, soil temperature and soil pH decreased, while soil moisture and soil C:N ratios increased with elevation. Responses of these properties varied seasonally. Fungal abundance increased with elevation during wet and dry seasons. Fungal community composition shifted in response to elevation, and to a lesser extent by season. These shifts were accompanied by varying responses of important fungal functional groups during the wet season and the relative abundance of certain fungal phyla. We suggest that elevation and the responses of certain fungal functional groups may be structuring fungal communities along this elevation gradient. TMCF are ecosystems that are rapidly changing due to climate change. Our study suggests that these changes may affect how fungal communities are structured.     

Responses of bacterial and fungal communities to an elevation gradient in a subtropical montane forest of China

Bacteria and fungi are ecologically important contributors to various functioning of forest ecosystems. In this study, we examined simultaneously the bacterial and fungal distributions in response to elevation changes of a forest. By using clone library analysis from genomic DNA extracted from forest humic clay soils, the composition and diversity of bacterial and fungal communities were determined across an elevation gradient from low via medium to high, in a subtropical forest in the Mountain Lushan, China. Our results showed that soil water content and nutrient availability, specifically total carbon, differed significantly with elevation changes. Although the soil acidity did not differ significantly among the three sites, low pH (around 4) could be an important selection factor selecting for acidophilic Acidobacteria and Alphaproteobacteria, which were the most abundant bacterial clones. As the majority of the fungi recovered, both Basidiomycota and Ascomycota, and their relative abundance were most closely associated with the total carbon. Based on the Shannon–Weaver diversity index and ∫-libshuff analysis, the soil at medium elevation contained the highest diversity of bacteria compared with those at high and low elevations. However, it is difficult to predict overall fungal diversity along elevation. The extreme high soil moisture content which may lead to the formation of anaerobic microhabitats in the forest soils potentially reduces the overall bacterial and fungal diversity.     

Tree diversity patterns in successive vegetation types along an elevation gradient in the Mountains of Eastern Mexico

Tree species richness changes along elevation gradients in response to underlying environmental conditions. Our hypothesis was that richness is associated with climatic variables and decreases with elevation. The objective was to identify trends in species, genus and family richness, diversity and vegetation structure in relation to climate variables along an elevation gradient with successive types of forest in Veracruz, Mexico. Trees were identified and measured in 0.1 ha at 15 sites located from 140 to 4000 m a.s.l. Generalized linear models were used to fit richness, diversity, basal area and density as a function of elevation; the best model was selected using Akaike’s Information Criterion. Multivariate analyses were used to explore climatic variables associated to composition of groups of sites along the gradient. Along the entire elevation gradient, species, genus and family richness decreased unimodally, and diversity decreased monotonically. Richness was positively correlated with temperature but not with precipitation. Basal area increased monotonically and highest basal area was associated with high humidity and certain tree species (Quercus and Abies). Ordinations indicated three groups of sites: lower elevation dry forest associated with temperature seasonality, mid-elevation cloud forest associated with precipitation-related variables, and coniferous forest at the top of the gradient associated with elevation. Our study shows that different plant communities are associated with certain climatic conditions and harbour different tree species, genera and families. The results support the hypothesis that species richness is associated with climate, and decreases with elevation.     

The nutrient status of epiphytes and their host trees along an elevational gradient in Costa Rica

Vascular epiphytes are a conspicuous and highly diverse group in tropical wet forests; yet, we understand little about their mineral nutrition across sites. In this study, we examined the mineral nutrition of three dominant vascular epiphyte groups: ferns, orchids, and bromeliads, and their host trees from samples collected along a 2600 m elevational gradient in the tropical wet forests of Costa Rica. We predicted that the mineral nutrition of ferns, orchids, and bromeliads would differ because of their putative differences in nutrient acquisition mechanisms and nutrient sources—atmospherically dependent, foliar feeding bromeliads would have lower nitrogen (N) and phosphorous (P) concentrations and more depleted δ¹⁵N values than those in canopy soil-rooted ferns because canopy soil is higher in available N, and more enriched in δ¹⁵N than the atmospheric sources of precipitation and throughfall. We also predicted that epiphyte foliar chemistry would mirror that of host trees because of the likely contribution of host trees to the nutrient cycle of epiphytes via foliar leaching and litter contributions to canopy soil. In the same vein, we predicted that epiphyte and host tree foliar chemistry would vary with elevation reflecting ecosystem-level nutrients—soil N availability increases and P availability decreases with increasing elevation. Our results confirmed that canopy soil-rooted epiphytes had higher N concentrations than atmospheric epiphytes; however, our predictions were not confirmed with respect to P which did not vary among groups indicating fixed P availability within sites. In addition, foliar δ¹⁵N values did not match our prediction in that canopy soil-rooted as well as atmospheric epiphytes had variable signatures. Discriminant function analysis (DFA) on foliar measurements determined that ferns, orchids, and bromeliads are statistically distinct in mineral nutrition. We also found that P concentrations of ferns and orchids, but not bromeliads, were significantly correlated with those of host trees indicating a possible link in their mineral nutrition’s via canopy soil. Interestingly, we did not find any patterns of epiphyte foliar chemistry with elevation. These data indicate that the mineral nutrition of the studied epiphyte groups are distinct and highly variable within sites and the diverse uptake mechanisms of these epiphyte groups enhance resource partitioning which may be a mechanism for species richness maintenance in tropical forest canopies.     

Bacterial functional redundancy along a soil reclamation gradient

A strategy to measure bacterial functional redundancy was developed and tested with soils collected along a soil reclamation gradient by determining the richness and diversity of bacterial groups capable of in situ growth on selected carbon substrates. Soil cores were collected from four sites along a transect from the Jamari tin mine site in the Jamari National Forest, Rondonia, RO, Brazil: denuded mine spoil, soil from below the canopy of invading pioneer trees, revegetated soil under new growth on the forest edge, and the forest floor of an adjacent preserved forest. Bacterial population responses were analyzed by amending these soil samples with individual carbon substrates in the presence of bromodeoxyuridine (BrdU). BrdU-labeled DNA was then subjected to a 16S-23S rRNA intergenic analysis to depict the actively growing bacteria from each site. The number and diversity of bacterial groups responding to four carbon substrates (L-serine, L-threonine, sodium citrate, and alpha-lactose hydrate) increased along the reclamation-vegetation gradient such that the preserved forest soil samples contained the highest functional redundancy for each substrate. These data suggest that bacterial functional redundancy increases in relation to the regrowth of plant communities and may therefore represent an important aspect of the restoration of soil biological functionality to reclaimed mine spoils. They also suggest that bacterial functional redundancy may be a useful indicator of soil quality and ecosystem functioning.     

Richness of cultivable endophytic fungi along an altitudinal gradient in wet forests of Costa Rica

We collected various plant species along an altitudinal gradient ranging from 400 to 2900 m in tropical wet forests of Costa Rica, isolated the associated endophytic fungi, and performed bioinformatic analyses to determine whether changes in altitude are related to changes in their richness and community structure. We showed that the richness of endophytic fungi varied along the altitudinal gradient, being higher in the lowest stratum and decreasing as elevation increases. Each stratum presented a particular composition and diversity of endophytes, although the whole population was characterized by the presence of a few dominant and apparently ubiquitous species, coexisting with a number of less abundant species that presented a more limited host range. These results have important implications for better understanding the role of altitude on the distribution and composition of endophytic fungal populations in tropical forests, but also for maximizing the number and diversity of endophytic isolates in bioprospecting campaigns.     

Lack of host specificity leads to independent assortment of dipterocarps and ectomycorrhizal fungi across a soil fertility gradient

Plants interact with a diversity of microorganisms, and there is often concordance in their community structures. Because most community‐level studies are observational, it is unclear if such concordance arises because of host specificity, in which microorganisms or plants limit each other's occurrence. Using a reciprocal transplant experiment, we tested the hypothesis that host specificity between trees and ectomycorrhizal fungi determines patterns of tree and fungal soil specialisation. Seedlings of 13 dipterocarp species with contrasting soil specialisations were seeded into plots crossing soil type and canopy openness. Ectomycorrhizal colonists were identified by DNA sequencing. After 2.5 years, we found no evidence of host specificity. Rather, soil environment was the primary determinant of ectomycorrhizal diversity and composition on seedlings. Despite their close symbiosis, our results show that ectomycorrhizal fungi and tree communities in this Bornean rain forest assemble independently of host‐specific interactions, raising questions about how mutualism shapes the realised niche.     

Abrupt changes in the composition and function of fungal communities along an environmental gradient in the high Arctic

Fungi play a key role in soil–plant interactions, nutrient cycling and carbon flow and are essential for the functioning of arctic terrestrial ecosystems. Some studies have shown that the composition of fungal communities is highly sensitive to variations in environmental conditions, but little is known about how the conditions control the role of fungal communities (i.e., their ecosystem function). We used DNA metabarcoding to compare taxonomic and functional composition of fungal communities along a gradient of environmental severity in Northeast Greenland. We analysed soil samples from fell fields, heaths and snowbeds, three habitats with very contrasting abiotic conditions. We also assessed within‐habitat differences by comparing three widespread microhabitats (patches with high cover of Dryas, Salix, or bare soil). The data suggest that, along the sampled mesotopographic gradient, the greatest differences in both fungal richness and community composition are observed amongst habitats, while the effect of microhabitat is weaker, although still significant. Furthermore, we found that richness and community composition of fungi are shaped primarily by abiotic factors and to a lesser, though still significant extent, by floristic composition. Along this mesotopographic gradient, environmental severity is strongly correlated with richness in all fungal functional groups: positively in saprotrophic, pathogenic and lichenised fungi, and negatively in ectomycorrhizal and root endophytic fungi. Our results suggest complex interactions amongst functional groups, possibly due to nutrient limitation or competitive exclusion, with potential implications on soil carbon stocks. These findings are important in the light of the environmental changes predicted for the Arctic.     

Plant species richness and diversity along an altitudinal gradient in the Sierra Nevada, Mexico

This article presents an analysis of plant species richness and diversity and its association with climatic and soil variables along a 1300‐m elevation gradient on the Cerro Tláloc Mountain in the northern Sierra Nevada in Mexico. Two 1000‐m² tree sampling plots were created at each of 21 selected sampling sites, as well as two 250‐m² plots for shrubs and six 9‐m² plots for herbaceous plants. Species richness and diversity were estimated for each plant life form, and beta diversity between sites was estimated along the gradient. The relationship between species richness and diversity and environmental variables was modelled using simple linear correlation and regression trees. Species richness and diversity showed a unimodal pattern with a bias towards high values in the lower half of the elevation gradient under study. This response was consistent for all three life forms. Beta diversity increased steadily along the elevation gradient, being lower between contiguous sites at intermediate elevations and high – the species replacement rate was nearly 100%– between sites at the extremes of the gradient. Few species were adapted to the full spectrum of environmental variation along the elevation gradient studied. The regression tree suggests that differences in species richness are mainly influenced by elevation (temperature and humidity) and soil variables, namely A₂ permanent wilting point, organic matter and horizon field capacity and A₁ horizon Mg²⁺.