Connecting the multiple dimensions of global soil fungal diversity.

How the multiple facets of soil fungal diversity vary worldwide remains virtually unknown, hindering the management of this essential species-rich group. By sequencing high-resolution DNA markers in over 4000 topsoil samples from natural and human-altered ecosystems across all continents, we illustrate the distributions and drivers of different levels of taxonomic and phylogenetic diversity of fungi and their ecological groups. We show the impact of precipitation and temperature interactions on local fungal species richness (alpha diversity) across different climates. Our findings reveal how temperature drives fungal compositional turnover (beta diversity) and phylogenetic diversity, linking them with regional species richness (gamma diversity). We integrate fungi into the principles of global biodiversity distribution and present detailed maps for biodiversity conservation and modeling of global ecological processes.

Five new species of Gloeandromyces (Fungi, Laboulbeniales) from tropical American bat flies (Diptera, Streblidae), revealed by morphology and phylogenetic reconstruction.

This paper describes and illustrates five new species of Gloeandromyces (Ascomycota, Laboulbeniales) associated with tropical American bat flies (Diptera, Streblidae). These are Gloeandromyces cusucoensis sp. nov. from Trichobius uniformis in Costa Rica and Honduras, G. diversiformis sp. nov. from Strebla wiedemanni in Costa Rica, G. plesiosaurus sp. nov. from Trichobius yunkeri in Panama, G. pseudodickii sp. nov. from Trichobius longipes in Ecuador and Panama, and G. verbekeniae sp. nov. from Strebla galindoi in Ecuador and Panama. The description of these five species doubles the number of known species in the genus. Morphological characteristics, host association, and a three-locus (18S nuc rDNA, 28S nuc rDNA, TEF1) phylogenetic reconstruction support placement of these taxa in the genus Gloeandromyces. Three of the new species are polymorphic; they have multiple morphotypes that grow in specific positions on the host integument: G. diversiformis f. diversiformis, f. musiformis, and f. vanillicarpiformis; G. plesiosaurus f. asymmetricus and f. plesiosaurus; and G. verbekeniae f. verbekeniae and f. inflexus. Finally, a dichotomous key to all species and morphotypes is presented.

Mycophagy: A Global Review of Interactions between Invertebrates and Fungi.

Fungi are diverse organisms that occupy important niches in natural settings and agricultural settings, acting as decomposers, mutualists, and parasites and pathogens. Interactions between fungi and other organisms, specifically invertebrates, are understudied. Their numbers are also severely underestimated. Invertebrates exist in many of the same spaces as fungi and are known to engage in fungal feeding or mycophagy. This review aims to provide a comprehensive, global view of mycophagy in invertebrates to bring attention to areas that need more research, by prospecting the existing literature. Separate searches on the Web of Science were performed using the terms "mycophagy" and "fungivore". Invertebrate species and corresponding fungal species were extracted from the articles retrieved, whether the research was field- or laboratory-based, and the location of the observation if field-based. Articles were excluded if they did not list at least a genus identification for both the fungi and invertebrates. The search yielded 209 papers covering seven fungal phyla and 19 invertebrate orders. Ascomycota and Basidiomycota are the most represented fungal phyla whereas Coleoptera and Diptera make up most of the invertebrate observations. Most field-based observations originated from North America and Europe. Research on invertebrate mycophagy is lacking in some important fungal phyla, invertebrate orders, and geographic regions.

Russulaceae of the Pakaraima Mountains of Guyana. IV. New species forming a distinct lineage of Lactarius subg. Plinthogalus.

The Neotropics have recently emerged as an important region for studies of tropical ectomycorrhizal (ECM) fungi. Specific neotropical areas with high ECM host tree densities have ECM fungal diversities rivaling those of higher-latitude forests. Some forests of the Guiana Shield are dominated by endemic ECM trees of the Fabaceae, including species of Dicymbe (subfam. Detarioideae), Aldina (subfam. Papilionoideae), and Pakaraimaea (Cistaceae). One of the most species-rich ECM fungal families present in each of these systems is Russulaceae. Long-term sampling in forests in Guyana's Pakaraima Mountains has revealed a number of species of the Russulaceae genera Lactarius, Lactifluus, and Russula. In this study, we document a previously unknown, distinct lineage of Lactarius subg. Plinthogalus containing eight species from the Guiana Shield. Here, we describe five of these species from Guyana as new to science: Lactarius humiphilus, Lactarius mycenoides, Lactarius guyanensis, Lactarius dicymbophilus, and Lactarius aurantiolamellatus. Morphological descriptions, habit, habitat, and known distribution are provided for each new species. Sequence data for the barcode internal transcribed spacer (ITS) locus are provided for types and most other collections of the new taxa, and a molecular phylogenetic analysis based on the ITS, 28S, and RPB2 (second-largest subunit of the RNA polymerase II) loci across the genus Lactarius corroborates their morphology-based infrageneric placement. The discovery of this lineage changes our insights into the biogeography and evolutionary history of Lactarius subg. Plinthogalus.

Global patterns in endemicity and vulnerability of soil fungi.

Fungi are highly diverse organisms, which provide multiple ecosystem services. However, compared with charismatic animals and plants, the distribution patterns and conservation needs of fungi have been little explored. Here, we examined endemicity patterns, global change vulnerability and conservation priority areas for functional groups of soil fungi based on six global surveys using a high-resolution, long-read metabarcoding approach. We found that the endemicity of all fungi and most functional groups peaks in tropical habitats, including Amazonia, Yucatan, West-Central Africa, Sri Lanka, and New Caledonia, with a negligible island effect compared with plants and animals. We also found that fungi are predominantly vulnerable to drought, heat and land-cover change, particularly in dry tropical regions with high human population density. Fungal conservation areas of highest priority include herbaceous wetlands, tropical forests, and woodlands. We stress that more attention should be focused on the conservation of fungi, especially root symbiotic arbuscular mycorrhizal and ectomycorrhizal fungi in tropical regions as well as unicellular early-diverging groups and macrofungi in general. Given the low overlap between the endemicity of fungi and macroorganisms, but high conservation needs in both groups, detailed analyses on distribution and conservation requirements are warranted for other microorganisms and soil organisms.

Quantitative mapping of mercury and selenium in mushroom fruit bodies with laser ablation-inductively coupled plasma-mass spectrometry.

This work describes the development of a novel method for quantitative mapping of Hg and Se in mushroom fruit body tissues with laser ablation coupled to inductively coupled plasma-mass spectrometry (LA-ICP-MS). Different parameters of the protocol for preparation of the standards used for quantification via external calibration were assessed, e.g., the dissolution temperature of gelatin standards and the addition of chitosan and L-cysteine as additives to the gelatin-based calibration droplets to better match the sample matrix. While chitosan was not suited for this purpose, the presence of L-cysteine considerably improved the figures of merit of the calibration, leading to limits of detection of 0.006 and 0.3 µg g-1 for Hg and Se, respectively, at a pixel size of 20 × 20 µm. Further, an in-house reference material, ideally suited for the validation of the method for application to mushroom samples, was successfully prepared from a paste of Boletus edulis. The newly developed method was used to investigate the distribution of Hg and Se in tissue sections of five porcini mushroom individuals of three different species (Boletus edulis, Boletus aereus, and Boletus pinophilus) and one sample of a parasol mushroom (Macrolepiota procera). For one sample, additional areas were ablated at higher spatial resolution, with a laser spot size down to 5 µm, which allows a detailed investigation of the spatial distribution of Hg and Se in mushrooms.

Biogeographic history of a large clade of ectomycorrhizal fungi, the Russulaceae, in the Neotropics and adjacent regions.

The biogeography of neotropical fungi remains poorly understood. Here, we reconstruct the origins and diversification of neotropical lineages in one of the largest clades of ectomycorrhizal fungi in the globally widespread family Russulaceae. We inferred a supertree of 3285 operational taxonomic units, representing worldwide internal transcribed spacer sequences. We reconstructed biogeographic history and diversification and identified lineages in the Neotropics and adjacent Patagonia. The ectomycorrhizal Russulaceae have a tropical African origin. The oldest lineages in tropical South America, most with African sister groups, date to the mid-Eocene, possibly coinciding with a boreotropical migration corridor. There were several transatlantic dispersal events from Africa more recently. Andean and Central American lineages mostly have north-temperate origins and are associated with North Andean uplift and the general north-south biotic interchange across the Panama isthmus, respectively. Patagonian lineages have Australasian affinities. Diversification rates in tropical South America and other tropical areas are lower than in temperate areas. Neotropical Russulaceae have multiple biogeographic origins since the mid-Eocene involving dispersal and co-migration. Discontinuous distributions of host plants may explain low diversification rates of tropical lowland ectomycorrhizal fungi. Deeply diverging neotropical fungal lineages need to be better documented.

The Diversity of Lichenized Trentepohlioid Algal (Ulvophyceae) Communities is Driven by Fungal Taxonomy and Ecological Factors.

Trentepohliales are a group of both free-living and lichenized algae, with most diversity occurring in tropical regions. Recent studies showed that the abundance of lichens with a trentepohlioid photobiont has been increasing in temperate habitats, probably because of global warming, which makes them an interesting study case. A detailed molecular study of the diversity of lichenized Trentepohliales, epiphytic as well as epilithic, was performed in three forests of north-western Europe. Additional samples of lichens of the Arthoniales order (associating essentially with a trentepohlioid photobiont) from other European regions and from other continents were also sequenced. A total of 195 algal sequences were obtained. Phylogenetic analyses with rbcL and ITS loci were performed and associations between phylogenetic distances of photobionts and ecological factors (substratum, climate or Wirth indices, mycobiont taxonomy, and geographic location) were tested by variation partitioning and phylogenetic signal analyses. The high number of rbcL algal haplotypes found in some lichens or on different substrata revealed that the Trentepohliales diversity in extratropical regions was underestimated. The phylogenetic patterns showed selectivity of some photobionts in their fungal partner choice and vice-versa, while others were linked with several haplotypes. Photobionts seemed to be less selective than mycobionts. The main factors influencing lichenized algal community were climate and mycobiont species. Coevolution between mycobionts and photobionts as well as switching between free living and lichenized lifestyles appeared to drive the evolution of Trentepohliales and might explain the high cryptic diversity observed, which might be changing in some regions due to climate change.

Two new Russula species (fungi) from dry dipterocarp forest in Thailand suggest niche specialization to this habitat type.

Dry dipterocarp forests are among the most common habitat types in Thailand. Russulaceae are known as common ectomycorrhizal symbionts of Dipterocarpaceae trees in this type of habitat. The present study aims to identify collections of Russula subsection Amoeninae Buyck from dry dipterocarp forests in Thailand. A multi-locus phylogenetic analysis placed Thai Amoeninae collections in two novel lineages, and they are described here as R. bellissima sp. nov. and R. luteonana sp. nov. The closest identified relatives of both species were sequestrate species suggesting that they may belong to drought-adapted lineages. An analysis of publicly available ITS sequences in R. subsect. Amoeninae did not confirm evidence of any of the new species occurring in other Asian regions, indicating that dry dipterocarp forests might harbor a novel community of ectomycorrhizal fungi. Macromorphological characters are variable and are not totally reliable for distinguishing the new species from other previously described Asian Amoeninae species. Both new species are defined by a combination of differentiated micromorphological characteristics in spore ornamentation, hymenial cystidia and hyphal terminations in the pileipellis. The new Amoeninae species may correspond to some Russula species collected for consumption in Thailand, and the detailed description of the new species can be used for better identification of edible species and food safety in the region.

Taxonomic Revision of the Genus Lactifluus (Russulales, Basidiomycota) of South Korea.

Lactifluus (Pers.) Roussel is an ectomycorrhizal genus that was recently recognized to be distinct from the genus Lactarius. To date, 226 Lactifluus species have been reported worldwide. Misidentification of Lactifluus species is common because of intraspecific morphological variation, cryptic diversity, and the limited number of taxonomic keys available. Molecular data are indispensable for species delimitation; a multilocus phylogenetic analysis showed that most Asian Lactifluus species are not conspecific with morphologically similar species present on other continents. In particular, Korea has misused European and North American Lactifluus names. In this study, we evaluated the taxonomy of Lactifluus in Korea using both morphological and multilocus molecular (ITS, nrLSU, rpb1, and rpb2) data. We examined 199 Lactifluus specimens collected between 1980 and 2016, and a total of 24 species across the four Lactifluus subgenera were identified. All Korean species are distinct and clearly separated from European and North American species. Five taxa corresponded to previously described species from Asia and the remaining 19 taxa are confirmed as new species. Herein, we provide keys to the Korean Lactifluus species within their subgenera, molecular phylogenies, a summary of diversity, and detailed description of the new species.

Enlightening the black and white: species delimitation and UNITE species hypothesis testing in the Russula albonigra species complex.

Russula albonigra is considered a well-known species, morphologically delimited by the context of the basidiomata blackening without intermediate reddening, and the menthol-cooling taste of the lamellae. It is supposed to have a broad ecological range and a large distribution area. A thorough molecular analysis based on four nuclear markers (ITS, LSU, RPB2 and TEF1-α) shows this traditional concept of R. albonigra s. lat. represents a species complex consisting of at least five European, three North American, and one Chinese species. Morphological study shows traditional characters used to delimit R. albonigra are not always reliable. Therefore, a new delimitation of the R. albonigra complex is proposed and a key to the described European species of R. subgen. Compactae is presented. A lectotype and an epitype are designated for R. albonigra and three new European species are described: R. ambusta, R. nigrifacta, and R. ustulata. Different thresholds of UNITE species hypotheses were tested against the taxonomic data. The distance threshold of 0.5% gives a perfect match to the phylogenetically defined species within the R. albonigra complex. Publicly available sequence data can contribute to species delimitation and increase our knowledge on ecology and distribution, but the pitfalls are short and low quality sequences.

The Effect of Surrounding Vegetation on the Mycorrhizal Fungal Communities of the Temperate Tree Crataegus monogyna Jacq.

About 90% of all land plants form mycorrhiza to facilitate the acquisition of essential nutrients such as phosphorus, nitrogen, and sometimes carbon. Based on the morphology of the interaction and the identity of the interacting plants and fungi, four major mycorrhizal types have been distinguished: arbuscular mycorrhiza (AM), ectomycorrhizal (EcM), ericoid mycorrhiza, and orchid mycorrhiza. Although most plants are assumed to form only one type of mycorrhiza, some species simultaneously form associations with two mycorrhizal types within a single root system. However, the dual-mycorrhizal status of many species is under discussion and in some plant species the simultaneous association with two mycorrhizal types varies in space or time or depends on the ecological context. Here, we assessed the mycorrhizal communities associating with common hawthorn (Crataegus monogyna), a small tree that commonly associates with AM fungi, and investigated the potential factors that underlie variation in mycorrhizal community composition. Histological staining of C. monogyna roots showed the presence of a Hartig net and hyphal sheaths in and around the roots, demonstrating the capacity of C. monogyna to form EcM. Meta-barcoding of soil and root samples of C. monogyna collected in AM-dominated grassland vegetation and in mixed AM + EcM forest vegetation showed a much higher number of EcM sequences and OTUs in root and soil samples from mixed AM + EcM vegetation than in samples from pure AM vegetation. We conclude that C. monogyna is able to form both AM and EcM, but that the extent to which it does depends on the environmental context, i.e., the mycorrhizal type of the surrounding vegetation.

On the Fly: Tritrophic Associations of Bats, Bat Flies, and Fungi.

Parasitism is one of the most diverse and abundant modes of life, and of great ecological and evolutionary importance. Notwithstanding, large groups of parasites remain relatively understudied. One particularly unique form of parasitism is hyperparasitism, where a parasite is parasitized itself. Bats (Chiroptera) may be parasitized by bat flies (Diptera: Hippoboscoidea), obligate blood-sucking parasites, which in turn may be parasitized by hyperparasitic fungi, Laboulbeniales (Ascomycota: Laboulbeniomycetes). In this study, we present the global tritrophic associations among species within these groups and analyze their host specificity patterns. Bats, bat flies, and Laboulbeniales fungi are shown to form complex networks, and sixteen new associations are revealed. Bat flies are highly host-specific compared to Laboulbeniales. We discuss possible future avenues of study with regard to the dispersal of the fungi, abiotic factors influencing the parasite prevalence, and ecomorphology of the bat fly parasites.

Updated taxonomy of LactifluussectionLuteoli: L.russulisporus from Australia and L.caliendrifer from Thailand.

Lactifluusrussulisporus Dierickx & De Crop and Lactifluuscaliendrifer Froyen & De Crop are described from eucalypt forests in Queensland, Australia and different forest types in Thailand, respectively. Both species have recently been published on Index Fungorum and fit morphologically and molecularly in L.sect.Luteoli, a section within L.subg.Gymnocarpi that encompasses species with alboochraceous basidiomes, white latex that stains brown and typical capitate elements in the pileipellis and/or marginal cells.

Lactifluusbicapillus (Russulales, Russulaceae), a new species from the Guineo-Congolian rainforest.

The milkcap genus Lactifluus is one of the most common ectomycorrhizal genera within Central African rainforests. During a field trip to the Dja Biosphere Reserve in Cameroon, a new Lactifluus species was found. Molecular and morphological analyses indicate that the species belongs to LactifluussectionXerampelini and we formally describe it here as Lactifluusbicapillus sp. nov.

Looks can be deceiving: the deceptive milkcaps (Lactifluus, Russulaceae) exhibit low morphological variance but harbour high genetic diversity.

The ectomycorrhizal genus Lactifluus is known to contain many species complexes, consisting of morphologically very similar species, which can be considered cryptic or pseudocryptic. In this paper, a thorough molecular study is performed of the clade around Lactifluus deceptivus (originally described by Peck from North America) or the deceptive milkcaps. Even though most collections were identified as L. deceptivus, the clade is shown to contain at least 15 species, distributed across Asia and America, indicating that the L. deceptivus clade represents a species complex. These species are morphologically very similar and are characterized by a tomentose pileus with thin-walled hyphae and a velvety stipe with thick-walled hyphae. An ITS1 sequence was obtained through Illumina sequencing for the lectotype of L. deceptivus, dating from 1885, revealing which clade represents the true L. deceptivus. In addition, it is shown that three other described species also belong to the L. deceptivus clade: L. arcuatus, L. caeruleitinctus and L. mordax, and molecularly confirmed that L. tomentoso-marginatus represents a synonym of L. deceptivus. Furthermore, two new Neotropical species are described: Lactifluus hallingii and L. domingensis.

Novel diversity in Lactifluus section Gerardii from Asia: five new species with pleurotoid or small agaricoid basidiocarps.

The ectomycorrhizal milkcap genus Lactifluus (Russulaceae) is commonly found in tropical and subtropical forests of Southeast Asia. During several sampling expeditions in Thailand, multiple collections of Lactifluus (Lf.) species with pleurotoid or small agaricoid basidiocarps were found. A molecular study was combined with a morphological study, in which the collections were compared with herbarium material. The molecular study indicated that four Thai collections belonged to undescribed species within Lactifluus section Gerardii, as was also the case for herbarium collections of a pleurotoid species from Nepal. One other collection from Thailand appeared closely related to Lf. uyedae, known only from Japan. Five species are described as new in the genus: Lf. auriculiformis, Lf. gerardiellus, Lf. bhandaryi, Lf. pulchrellus, and Lf. raspei.

Lactarius subgenus Russularia (Basidiomycota, Russulales): novel Asian species, worldwide phylogeny and evolutionary relationships.

Lactarius subg. Russularia is a large group of milkcaps occurring almost worldwide and dominant in many ecosystems. In this study we focus on new diversity, evolutionary relationships, divergence time, and origin of the subgenus. Six conifer symbionts are described as new to science: Lactarius atrii, L. aurantionitidus, L. dombangensis, L. flavigalactus, L. lachungensis, and L. sikkimensis. Species delimitation is assessed based on the concordance between morphological characteristics and an ITS phylogeny. Infrageneric relationships were studied using a phylogeny constructed from concatenated ITS-rpb2 data using Maximum Likelihood and Bayesian inference. Results show that species in this subgenus do not cluster together according to their geographic origin. Intercontinental sister relationships between Europe/Asia/North America are common but actual conspecificity is rare. This result suggests that allopatric speciation has played an important role within this subgenus. Only few morphological characteristics tend to be phylogenetically informative, with the most important being presence or absence of true cystidia and the pileipellis structure. Two datasets were generated in order to estimate the age of L. subg. Russularia. The results suggest the origin of L. subg. Russularia to be in the Mid Miocene period.

Diversity of Edible and Medicinal Mushrooms Used in the Noun Division of the West Region of Cameroon.

This article presents discussions of mushrooms as a source of food, income, as well as medicine among the Bamoun people of the highlands of West Cameroon, where the vegetation is mainly savannah mixed with forest galleries. Like most tribes in tropical Africa, the Bamoun people use a wide range of natural products as mushrooms. This study attempts to identify the various mushrooms exploited by the Bamoun. Ethnomycological surveys and field trips were conducted over 4 years in several villages in the Noun Division. Samples of wild mushrooms were collected from both the savannah and the forest galleries. These were described, preserved, and identified. The study shows that the Bamoun people use at least 40 species of mushrooms for either food or medicine. These species belong to 8 genera: Auricularia, Cantharellus, Ganoderma, Pleurotus, Lactarius, Lactifluus, Russula, and Termitomyces. Species of genera Lactarius, Lactifluus, Russula, and Termitomyces are most often used for food, whereas Ganoderma spp. and Pleurotus tuber-regium are mainly exploited for medicinal purposes. This survey provides an overview of the diversity of mushrooms and their importance to the local people of this area. Since some of the species mentioned by the local population were not fruiting at the time of our field trips, additional investigations are needed to further clarify the diversity and the usage of mushrooms in this region.