Large‐scale fungal diversity assessment in the Andean Yungas forests reveals strong community turnover among forest types along an altitudinal gradient

The Yungas, a system of tropical and subtropical montane forests on the eastern slopes of the Andes, are extremely diverse and severely threatened by anthropogenic pressure and climate change. Previous mycological works focused on macrofungi (e.g. agarics, polypores) and mycorrhizae in Alnus acuminata forests, while fungal diversity in other parts of the Yungas has remained mostly unexplored. We carried out Ion Torrent sequencing of ITS2 rDNA from soil samples taken at 24 sites along the entire latitudinal extent of the Yungas in Argentina. The sampled sites represent the three altitudinal forest types: the piedmont (400–700 m a.s.l.), montane (700–1500 m a.s.l.) and montane cloud (1500–3000 m a.s.l.) forests. The deep sequence data presented here (i.e. 4 108 126 quality‐filtered sequences) indicate that fungal community composition correlates most strongly with elevation, with many fungi showing preference for a certain altitudinal forest type. For example, ectomycorrhizal and root endophytic fungi were most diverse in the montane cloud forests, particularly at sites dominated by Alnus acuminata, while the diversity values of various saprobic groups were highest at lower elevations. Despite the strong altitudinal community turnover, fungal diversity was comparable across the different zonal forest types. Besides elevation, soil pH, N, P, and organic matter contents correlated with fungal community structure as well, although most of these variables were co‐correlated with elevation. Our data provide an unprecedented insight into the high diversity and spatial distribution of fungi in the Yungas forests.     

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

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

Fungal colonization of shrub willow roots at the forefront of a receding glacier

Shrub willows (Salix spp.) form associations with arbuscular mycorrhizal (AM), ectomycorrhizal (EM) and dark septate endophytic (DSE) fungi. Willow root colonization by these three types of fungi was studied on a deglaciated forefront of Lyman Glacier, Washington, USA. Root colonization was low; less than 1% of the root length was colonized by AM and 25.6% by DSE. EM colonized 25% of the root tips and 19.4% of the root length. AM and DSE colonization were not related to distance from the present glacier terminus or to canopy cover. EM colonization increased with distance from the glacier terminus based on gridline intercept data but not on root tip frequency data. Availability of propagules in the substrate was low, but numbers of propagules increased with distance from the glacier terminus. The EM communities were dominated by three ascomycetes showing affinity to Sordariaceae in BLAST analyses. Other frequent taxa on the glacier forefront included species of Cortinariaceae, Pezizaceae, Russulaceae, Thelephoraceae and Tricholomataceae. When occurrence of individual taxa was used as a response variable to canopy cover, distance from the glacier terminus, and their interaction, four different fungal guilds were identified: 1) fungi that did not respond to these environmental variables; 2) fungi that occurred mainly in intercanopy areas and decreased with distance from the glacier terminus; 3) fungi that were insensitive to canopy cover but increased with distance from the glacier terminus; 4) fungi that occurred mainly under willow canopies and increased with distance from the glacier terminus. We suggest that fungal colonization is mainly limited by fungal propagule availability. Environmental conditions may also limit successful establishment of plant-fungus associations. We propose that the four EM guilds partly explain successional dynamics. The initial EM community comprises fungi that tolerate low organic matter and nitrogen environment (first and second guilds above). During later community development, these fungi are replaced by those that benefit from an increased organic matter and nitrogen environment (third and fourth guilds above).     

Dual mycorrhizal colonization of forest-dominating tropical trees and the mycorrhizal status of non-dominant tree and liana species

The contribution of mycorrhizal associations to maintaining tree diversity patterns in tropical rain forests is poorly known. Many tropical monodominant trees form ectomycorrhizal (EM) associations, and there is evidence that the EM mutualism contributes to the maintenance of monodominance. It is assumed that most other tropical tree species form arbuscular mycorrhizal (AM) associations, and while many mycorrhizal surveys have been done, the mycorrhizal status of numerous tropical tree taxa remains undocumented. In this study, we tested the assumption that most tropical trees form AM associations by sampling root vouchers from tree and liana species in monodominant Dicymbe corymbosa forest and an adjacent mixed rain forest in Guyana. Roots were assessed for the presence/absence of AM and EM structures. Of the 142 species of trees and lianas surveyed, three tree species (the monodominant D. corymbosa, the grove-forming D. altsonii, and the non-dominant Aldina insignis) were EM, 137 were exclusively AM, and two were non-mycorrhizal. Both EM and AM structures were observed in D. corymbosa and D. altsonii. These results provide empirical data supporting the assumption that most tropical trees form AM associations for this region in the Guiana Shield and provide the first report of dual EM/AM colonization in Dicymbe species. Dual colonization of the Dicymbe species should be further explored to determine if this ability contributes to the establishment and maintenance of site dominance. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Species richness and community composition of mat-forming ectomycorrhizal fungi in old- and second-growth Douglas-fir forests of the HJ Andrews Experimental Forest,Oregon, USA

We investigated the species identity of mat-forming ectomycorrhizal (EM) fungi associated with old- and second-growth Douglas-fir stands. Using molecular analyses of rhizomorphs and EM root tips, we characterized 28 unique internal transcribed spacer sequences and considered them proxies for mat-forming EM species. In both stand age classes, one Athelioid species in the genus Piloderma dominated our sample of the mat-forming fungal community. In second-growth stands, the second most frequently encountered mat-forming EM species belonged to the genus Hysterangium. In old-growth stands, several Ramaria species were associated with a frequently encountered mat morphology but no species dominated the community. After using rarefaction analysis to standardize sampling effort, the total species richness did not differ statistically between old- and second-growth habitats. Both an abundance of infrequently encountered species and incomplete sampling of the mat-forming EM community may have limited our ability to detect potential differences in species richness. Several frequently encountered Piloderma species appear to have broad (holarctic) distributions and diverse host associations and their potential importance in forest ecosystems warrants further study.     

Long-Term Impacts of Fuelwood Extraction on a Tropical Montane Cloud Forest

Fuelwood extracted from natural forests serves as a principal energy source in rural regions of many tropical countries. Although fuelwood extraction (even low intensities) might strongly impact the structure and species composition of natural forests, long-term studies remain scarce. Here, we estimate the potential long-term impacts (over several hundred years) of such repeated harvesting of single trees on tropical montane cloud forest in central Veracruz, Mexico, by applying a process-based forest growth model. We simulate a wide range of possible harvesting scenarios differing in wood volume harvested and preferred tree species and sizes, and use a set of indicators to compare their impacts on forest size structure and community composition. Results showed that the overall impact on forest structure and community composition increased linearly with the amount of harvested wood volume. Even at low levels of harvesting, forest size structure became more homogeneous in the long term because large old trees disappeared from the forest, but these changes might take decades or even centuries. Although recruitment of harvested species benefited from harvesting, species composition shifted to tree species that are not used for fuelwood. Our results demonstrate that fuelwood extraction can have marked long-term impacts on tropical montane cloud forests. The results also offer the possibility to support the design of management strategies for the natural species-rich forests that achieve a balance between economic needs and ecological goals of the stakeholders. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Diversity and community composition of ectomycorrhizal fungi in a dry deciduous dipterocarp forest in Thailand

Large forest areas of South-East Asia, are dominated by the Dipterocarpaceae tree family, which contains many important timber species. Unlike many other tropical trees, Dipterocarpaceae rely on ectomycorrhizal (ECM) root symbiosis for their mineral nutrition. This study aims to document the richness and community composition of ECM fungi in a dry deciduous forest in Thailand. Combining morphological and molecular identification methods revealed 69 species of ECM fungi that belong to 17 phylogenetic lineages. The /russula-lactarius, /tomentella-thelephora, /sordariales, /sebacina and /cantharellus lineages were the most species-rich. The fungal richness is comparable to other tropical rain forest sites, but the phylogenetic community structure has elements of both tropical and temperate ecosystems. Unlike tropical rain forests, the Cenococcum geophilum complex was one of the most frequent fungal taxa that had a relatively high ITS genetic diversity over the small sampling area. This study provides the first snapshot insight into the fungal community of dry dipterocarp forests. However, it is necessary to broaden the spatial and temporal scales of sampling to improve our understanding of the below-ground relations of dry and humid tropical forests.     

Availability and function of arbuscular mycorrhizal and ectomycorrhizal fungi during revegetation of dewatered reservoirs left after dam removal

Revegetation following dam removal projects may depend on recovery of arbuscular mycorrhizal (AM) and ectomycorrhizal (EM) fungal communities, which perform valuable ecosystem functions. This study assessed the availability and function of AM and EM fungi for plants colonizing dewatered reservoirs following a dam removal project on the Elwha River, Olympic Peninsula, Washington, United States. Availability was assessed via AM fungal spore density in soils and EM root tip colonization of Salix sitchensis (Sitka willow) in an observational field study. The effect of mycorrhizal fungi from 4 sources (reservoir soils, commercial inoculum, and 2 mature plant community soils) on growth and nutrient status of S. sitchensis was quantified in a greenhouse study. AM fungal spores and EM root tips were present in all field samples. In the greenhouse, plants receiving reservoir soil inoculum had only incipient mantle formation, while plants receiving inoculum from mature plant communities had fully formed EM root tips. EM formation corresponded with alleviation of phosphorus stress in plants (lower shoot nitrogen:phosphorus). Thus, revegetating plants have access to AM and EM fungi following dam removal, and EM formation may be especially important for plant P uptake in reservoir soils. However, availability of mycorrhizal fungi declines with distance from established plant communities. Furthermore, EM fungal communities in recently dewatered reservoirs may not be as effective at forming beneficial mycorrhizae as those from mature plant communities. Whole soil inoculum from mature plant communities may be important for the success of revegetating plants and recovery of mycorrhizal fungal communities.     

Ectomycorrhizal fungi of the Seychelles: diversity patterns and host shifts from the native Vateriopsis seychellarum (Dipterocarpaceae) and Intsia bijuga (Caesalpiniaceae) to the introduced Eucalyptus robusta (Myrtaceae), but not Pinus caribea (Pinaceae)

Ectomycorrhizal (ECM) fungi form highly diverse communities in temperate forests, but little is known about their community ecology in tropical ecosystems. Using anatomotyping and rDNA sequencing, ECM fungi were identified on root tips of the introduced Eucalyptus robusta and Pinus caribea as well as the endemic Vateriopsis seychellarum and indigenous Intsia bijuga in the Seychelles. Sequencing revealed 30 species of ECM fungi on root tips of V. seychellarum and I. bijuga, with three species overlapping. Eucalyptus robusta shared five of these taxa, whereas P. caribea hosted three unique species of ECM fungi that were likely cointroduced with containerized seedlings. The thelephoroid (including the anamorphic genus Riessiella), euagaric, boletoid and hymenochaetoid clades of basidiomycetes dominated the ECM fungal community of native trees. Two species of Annulatascaceae (Sordariales, Ascomycota) were identified and described as ECM symbionts of V. seychellarum. The low diversity of native ECM fungi is attributed to deforestation and long-term isolation of the Seychelles. Native ECM fungi associate with exotic eucalypts, whereas cointroduced ECM fungi persist in pine plantations for decades.     

Influence of host species on ectomycorrhizal communities associated with two co-occurring oaks (Quercus spp.) in a tropical cloud forest

Interactions between host tree species and ectomycorrhizal fungi are important in structuring ectomycorrhizal communities, but there are only a few studies on host influence of congeneric trees. We investigated ectomycorrhizal community assemblages on roots of deciduous Quercus crassifolia and evergreen Quercus laurina in a tropical montane cloud forest, one of the most endangered tropical forest ecosystems. Ectomycorrhizal fungi were identified by sequencing internal transcribed spacer and partial 28S rRNA gene. We sampled 80 soil cores and documented high ectomycorrhizal diversity with a total of 154 taxa. Canonical correspondence analysis indicated that oak host was significant in explaining some of the variation in ectomycorrhizal communities, despite the fact that the two Quercus species belong to the same red oak lineage (section Lobatae ). A Tuber species, found in 23% of the soil cores, was the most frequent taxon. Similar to oak-dominated ectomycorrhizal communities in temperate forests, Thelephoraceae, Russulaceae and Sebacinales were diverse and dominant.     

Diversity of an ectomycorrhizal fungal community studied by a root tip and total soil DNA approach

Molecular methods based on soil DNA extracts are increasingly being used to study the fungal diversity of ectomycorrhizal (EM) fungal communities in soil. Contrary to EM root tip identification, the use of molecular methods enables identification of extramatrical mycelia in soil. To compare fungal diversity as determined by root tip identification and mycelial identification, six soil samples were analysed. Root tips were extracted from the six samples and after amplification, the basidiomycete diversity on the root tips was analysed by denaturing gradient gel electrophoresis (DGGE). The soil from the six samples was sieved, total soil DNA was extracted and after amplification, the basidiomycete diversity in the soil fractions was analysed by DGGE. Fourteen different bands were excised from the DGGE gel and sequenced; fungal taxon names could be assigned to eight bands. Out of a total of 14 fungal taxa detected in soil, 11 fungal taxa were found on root tips, of which seven were EM fungal taxa. To examine whether the sieving treatment would affect EM species diversity, two different sieve mesh sizes were used and in addition, the organic soil fraction was analysed separately. DGGE analysis showed no differences in banding pattern for the different soil fractions. The organic fraction gave the highest DGGE band intensities. This work demonstrates that there is a high correspondence between basidiomycete diversity detected by molecular analysis of root tips and soil samples, irrespective of the soil fraction being analysed.     

Seasonal consumption of mycorrhizal fungi by a marsupial-dominated mammal community

The temperate forests of Australia support a high diversity of hypogeous fungi and a wide variety of mycophagous mammals, yet many mammal-fungal relationships are still poorly understood. We studied the seasonal fungal diets of eight sympatric mammals (seven marsupials and one rodent) in a remnant montane eucalypt forest. Fifty-five different fungal taxa were identified from 305 scat samples. Swamp wallabies (Wallabia bicolor), yellow-footed antechinus (Antechinus flavipes) and brown antechinus (A. stuartii) were the primary mycophagists in this community, but all mammals consumed fungi, including three species not previously recorded as mycophagous (eastern grey kangaroo, Macropus giganteus;common wallaroo, Osphranter robustus; and common dunnart, Sminthopsis murina). Winter was the peak season for fungal consumption and dietary diversity of fungi, however, the diversity of taxa ingested varied between species and season. Our work supports the idea that a diverse mycophagous mammal community is important for maintaining natural variation in fungal community composition.     

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

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

Moisture status during a strong El Niño explains a tropical montane cloud forest’s upper limit

Growing evidence suggests short-duration climate events may drive community structure and composition more directly than long-term climate means, particularly at ecotones where taxa are close to their physiological limits. Here we use an empirical habitat model to evaluate the role of microclimate during a strong El Niño in structuring a tropical montane cloud forest’s upper limit and composition in Hawai‘i. We interpolate climate surfaces, derived from a high-density network of climate stations, to permanent vegetation plots. Climatic predictor variables include (1) total rainfall, (2) mean relative humidity, and (3) mean temperature representing non-El Niño periods and a strong El Niño drought. Habitat models explained species composition within the cloud forest with non-El Niño rainfall; however, the ecotone at the cloud forest’s upper limit was modeled with relative humidity during a strong El Niño drought and secondarily with non-El Niño rainfall. This forest ecotone may be particularly responsive to strong, short-duration climate variability because taxa here, particularly the isohydric dominant Metrosideros polymorpha, are near their physiological limits. Overall, this study demonstrates moisture’s overarching influence on a tropical montane ecosystem, and suggests that short-term climate events affecting moisture status are particularly relevant at tropical ecotones. This study further suggests that predicting the consequences of climate change here, and perhaps in other tropical montane settings, will rely on the skill and certainty around future climate models of regional rainfall, relative humidity, and El Niño.     

Ectomycorrhizal fungal response to warming is linked to poor host performance at the boreal‐temperate ecotone

Rising temperatures associated with climate change have been shown to negatively affect the photosynthetic rates of boreal forest tree saplings at their southern range limits. To quantify the responses of ectomycorrhizal (EM) fungal communities associated with poorly performing hosts, we sampled the roots of Betula papyrifera and Abies balsamea saplings growing in the B4Warmed (Boreal Forest Warming at an Ecotone in Danger) experiment. EM fungi on the root systems of both hosts were compared from ambient and +3.4 °C air and soil warmed plots at two sites in northern Minnesota. EM fungal communities were assessed with high‐throughput sequencing along with measures of plant photosynthesis, soil temperature, moisture, and nitrogen. Warming selectively altered EM fungal community composition at both the phylum and genus levels, but had no significant effect on EM fungal operational taxonomic unit (OTU) diversity. Notably, warming strongly favored EM Ascomycetes and EM fungi with short‐contact hyphal exploration types. Declining host photosynthetic rates were also significantly inversely correlated with EM Ascomycete and EM short‐contact exploration type abundance, which may reflect a shift to less carbon demanding fungi due to lower photosynthetic capacity. Given the variation in EM host responses to warming, both within and between ecosystems, better understanding the link between host performance and EM fungal community structure will to clarify how climate change effects cascade belowground.     

Slowed decomposition is biotically mediated in an ectomycorrhizal, tropical rain forest

Bacteria and fungi drive the cycling of plant litter in forests, but little is known about their role in tropical rain forest nutrient cycling, despite the high rates of litter decay observed in these ecosystems. However, litter decay rates are not uniform across tropical rain forests. For example, decomposition can differ dramatically over small spatial scales between low-diversity, monodominant rain forests, and species-rich, mixed forests. Because the climatic patterns and soil parent material are identical in co-occurring mixed and monodominant forests, differences in forest floor accumulation, litter production, and decomposition between these forests may be biotically mediated. To test this hypothesis, we conducted field and laboratory studies in a monodominant rain forest in which the ectomycorrhizal tree Dicymbe corymbosa forms >80% of the canopy, and a diverse, mixed forest dominated by arbuscular mycorrhizal trees. After 2 years, decomposition was significantly slower in the monodominant forest (P < 0.001), but litter production was significantly greater in the mixed forest (P < 0.001). In the laboratory, we found microbial community biomass was greater in the mixed forest (P = 0.02), and the composition of fungal communities was distinct between the two rain forest types (P = 0.001). Sequencing of fungal rDNA revealed a significantly lower richness of saprotrophic fungi in the monodominant forest (19 species) relative to the species-rich forest (84 species); moreover, only 4% percent of fungal sequences occurred in both forests. These results show that nutrient cycling patterns in tropical forests can vary dramatically over small spatial scales, and that changes in microbial community structure likely drive the observed differences in decomposition.     

南岭典型植被土壤真菌群落结构与影响因素

南岭是世界生物多样性关键地区之一。土壤真菌多样性是生态系统的重要构成与系统健康稳定的重要指标。本研究采集南岭典型植被类型土壤样本,基于Illumina Novaseq测序平台分析土壤真菌群落结构,结合土壤理化性质和环境因子探讨不同植被类型下真菌群落组成的差异性和影响因子。结果显示,相对丰度最高的真菌门为担子菌门Basidiomycota,相对丰度从高海拔到低海拔变化介于28.54%–59.93%,在沟谷常绿阔叶林中比例较高,呈现出随着海拔的升高逐渐降低的变化格局;子囊菌门Ascomycota次之,相对丰度为19.52%–56.98%,随着海拔的上升呈现出先降低后升高的变化趋势;被孢霉门Mortierellomycota,相对丰度为1.03%–25.72%,随着海拔的上升,表现出先升后降又升的N型变化模式。不同真菌目在各植被类型土壤中的丰度不同,红菇目Russulales在沟谷常绿阔叶林和山地常绿阔叶林土壤中较为丰富;蘑菇目Agaricales在针阔混交林和高山草地土壤中较为丰富;被孢霉目Mortierellales在低海拔的山地常绿阔叶林和高海拔的山顶矮林土壤中较为丰富。土壤pH、全钾...     

Ectomycorrhizal fungal community structure across a bog-forest ecotone in southeastern Alaska

We examined the ectomycorrhizal (ECM) fungal community across a bog-forest ecotone in southeastern Alaska. The bog and edge were both characterized by poorly drained Histosols and a continuous layer of Sphagnum species, ericaceous shrubs, Carex species, and shore pine [Pinus contorta Dougl. ex Loud. var. contorta]. The forest had better-drained Inceptisols and Spodosols, a tree community comprised of western hemlock [Tsuga heterophylla (Raf.) Sarg.], yellow cedar (Thuja plicata Donn ex D. Don.), Sitka spruce [Picea sitchensis (Bong.) Carr.] and shore pine, and an understorey of ericaceous shrubs and herbs. ECM root tip density (tips cm^–3 soil) was significantly greater in the forest than the edge or bog and ECM colonization was significantly different in all three plant communities. The below ground ECM fungal taxa were analyzed using molecular techniques (PCR-RFLP and DNA sequencing). Three ECM fungal taxa, Suillus tomentosus (Kauffman) Singer, Cenococcum geophilum Fr.:Fr, and a Russula species, differed in relative frequency, yet were among the four most frequent in all three plant communities. Although differences in ECM fungal richness were observed across plant communities, unequal sampling of ECM roots due to root density and colonization differences confounded richness comparisons. Using resampling procedures for creating taxon-accumulation curves as a function of sampled ECM roots revealed similarities in cumulative ECM fungal taxa richness across the ecotone.