Systematic revision of species of Atractilina and Spiropes hyperparasitic on Meliolales (Ascomycota) in the tropics.

Atractilina Dearn. & Barthol. and Spiropes Cif. are genera of asexual fungi that comprise species mainly hyperparasitic on black mildews (Meliolales, Ascomycota). Although a common group of anamorphic fungi, they have been described up to now only by morphology and their systematic position is unknown. The present study provides a morphological treatise of all known species of Atractilina and Spiropes hyperparasitic on Meliolales, including insights into their systematic position, based on DNA sequences generated here for the first time. The study was conducted, based on 33 herbarium specimens and 23 specimens recently collected in Benin and Panama. The obtained DNA sequence data (28S rDNA and ITS rDNA) of A.parasitica and of two species of Spiropes show systematic placements in the Dothideomycetes and Leotiomycetes, respectively. The sequence data of the two Spiropes spp. do not group together. Moreover, the anamorph-teleomorph connection between Atractilinaparasitica and Malacariameliolicola, a pseudothecioid fungus, is confirmed. Three species in the genus Spiropes are proposed as new to science, namely S.angylocalycis, S.carpolobiae and S.croissantiformis. Four species are reported for Benin for the first time, three species for Panama and one species for mainland America. Atractilina and Spiropes are currently two genera with highly heterogeneous species and they might have to be split in the future, once the taxonomic concepts are validated by morphology and molecular sequence data.

A new species of Inosperma, and first record of I. afromelliolens (Inocybaceae, Fungi) from West Africa.

Here, we present the newly identified Inosperma macrocarpa and the first record of I. afromelliolens from West Africa. Inosperma macrocarpa is nested in an Old World Tropical clade, based on a molecular phylogeny inferred from the sequences of ITS, LSU, RPB2, and TEF1. Complete descriptions and illustrations, including photographs and line drawings, of the new species are presented. Morphological and molecular analyses based on collections from Benin confirmed the presence of I. afromelliolens in West Africa. Toxicity analysis showed that neither species contained muscarine, which further supports the hypothesis that the ability to produce muscarine is a derived trait of Inosperma.

Historical biogeography and diversification of ringless Amanita (section Vaginatae) support an African origin and suggest niche conservatism in the Americas.

Ectomycorrhizal fungi (ECM) sustain nutrient recycling in most terrestrial ecosystems, yet we know little about what major biogeographical events gave rise to present-day diversity and distribution patterns. Given the strict relationship between some ECM lineages and their hosts, geographically well-sampled phylogenies are central to understanding major evolutionary processes of fungal biodiversity patterns. Here, we focus on Amanita sect. Vaginatae to address global diversity and distribution patterns. Ancestral-state-reconstruction based on a 4-gene timetree with over 200 species supports an African origin between the late Paleocene and the early Eocene (ca. 56 Ma). Major biogeographic "out-of-Africa" events include multiple dispersal events to Southeast Asia (ca. 45-21 Ma), Madagascar (ca. 18 Ma), and the current Amazonian basin (ca. 45-36 Ma), the last two likely trans-oceanic. Later events originating in Southeast Asia involve Nearctic dispersal to North America (ca. 20-5 Ma), Oceania (Australia and New Zealand; ca. 15 Ma), and Europe (ca. 10-5 Ma). Subsequent dispersals were also inferred from Southeast Asia to East Asia (ca. 4 Ma); from North America to East Asia (ca. 11-8 Ma), Southeast Asia (ca. 19-2 Ma), Northern Andes (ca. 15 Ma), and Europe (ca. 15-2 Ma), respectively; and from the Amazon to the Caribbean region (ca. 25-20 Ma). Finally, we detected a significant increase in the net diversification rates in the branch leading to most northern temperate species in addition to higher state-dependent diversification rates in temperate lineages, consistent with previous findings. These results suggest that species of sect. Vaginatae likely have higher dispersal ability and higher adaptability to new environments, in particular compared to those of its sister clade, sect. Caesareae. Overall, the much wider distribution of A. sect. Vaginatae, from pan-tropical to pan-arctic, provides a unique window to understanding niche conservatism across a species-rich clade of ECM fungi.

Soil fungal communities of ectomycorrhizal dominated woodlands across West Africa.

Forests and woodlands in the West African Guineo-Sudanian transition zone contain many tree species that form symbiotic interactions with ectomycorrhizal (ECM) fungi. These fungi facilitate plant growth by increasing nutrient and water uptake and include many fruiting body-forming fungi, including some edible mushrooms. Despite their importance for ecosystem functioning and anthropogenic use, diversity and distribution of ECM fungi is severely under-documented in West Africa. We conducted a broad regional sampling across five West African countries using soil eDNA to characterize the ECM as well as the total soil fungal community in gallery forests and savanna woodlands dominated by ECM host tree species. We subsequently sequenced the entire ITS region and much of the LSU region to infer a phylogeny for all detected soil fungal species. Utilizing a long read sequencing approach allows for higher taxonomic resolution by using the full ITS region, while the highly conserved LSU gene allows for a more accurate higher-level assignment of species hypotheses, including species without ITS-based taxonomy assignments. We detect no overall difference in species richness between gallery forests and woodlands. However, additional gallery forest plots and more samples per plot would have been needed to firmly conclude this pattern. Based on both abundance and richness, species from the families Russulaceae and Inocybaceae dominate the ECM fungal soil communities across both vegetation types. The community structure of both total soil fungi and ECM fungi was significantly influenced by vegetation types and showed strong correlation within plots. However, we found no significant difference in fungal community structure between samples collected adjacent to different host tree species within each plot. We conclude that within plots, the fungal community is structured more by the overall ECM host plant community than by the species of the individual host tree that each sample was collected from.

Unravelling unexplored diversity of cercosporoid fungi (Mycosphaerellaceae, Mycosphaerellales, Ascomycota) in tropical Africa.

Cercosporoid fungi (Mycosphaerellaceae, Mycosphaerellales, Ascomycota) are one of the largest and most diverse groups of hyphomycetes causing a wide range of diseases of economically important plants as well as of plants in the wild. Although more than 6000 species are known for this group, the documentation of this fungal group is far from complete. Especially in the tropics, the diversity of cercosporoid fungi is poorly known. The present study aims to identify and characterise cercosporoid fungi collected on host plants belonging to Fabaceae in Benin, West Africa. Information on their morphology, host species and DNA sequence data (18S rDNA, 28S rDNA, ITS and tef1) is provided. DNA sequence data were obtained by a simple and non-culture-based method for DNA isolation which has been applied for cercosporoid fungi for the first time in the context of the present study. Among the loci used for the phylogenetic analysis, tef1 provided the best resolution together with the multigene dataset. Species delimitation in many cases, however, was only possible by combining molecular sequence data with morphological characteristics. Based on forty specimens recently collected in Benin, 18 species are presented with morphological descriptions, illustrations and sequence data. Among these, six species in the genus Cercospora and two species in Pseudocercospora are proposed as species new to science. The newly described species are Cercospora (C.) beninensis on Crotalaria macrocalyx, C. parakouensis on Desmodium tortuosum, C. rhynchophora on Vigna unguiculata, C. vignae-subterraneae on Vigna subterranea, C. tentaculifera on Vigna unguiculata, C. zorniicola on Zornia glochidiata, Pseudocercospora sennicola on Senna occidentalis and Pseudocercospora tabei on Vigna unguiculata. Eight species of cercosporoid fungi are reported for Benin for the first time, three of them, namely C. cf. canscorina, C. cf. fagopyri and C. phaseoli-lunati are new for West Africa. The presence of two species of cercosporoid fungi on Fabaceae previously reported from Benin, namely Nothopassalora personata and Passalora arachidicola, is confirmed.

Long- and short-read metabarcoding technologies reveal similar spatiotemporal structures in fungal communities.

Fungi form diverse communities and play essential roles in many terrestrial ecosystems, yet there are methodological challenges in taxonomic and phylogenetic placement of fungi from environmental sequences. To address such challenges, we investigated spatiotemporal structure of a fungal community using soil metabarcoding with four different sequencing strategies: short-amplicon sequencing of the ITS2 region (300-400 bp) with Illumina MiSeq, Ion Torrent Ion S5 and PacBio RS II, all from the same PCR library, as well as long-amplicon sequencing of the full ITS and partial LSU regions (1200-1600 bp) with PacBio RS II. Resulting community structure and diversity depended more on statistical method than sequencing technology. The use of long-amplicon sequencing enables construction of a phylogenetic tree from metabarcoding reads, which facilitates taxonomic identification of sequences. However, long reads present issues for denoising algorithms in diverse communities. We present a solution that splits the reads into shorter homologous regions prior to denoising, and then reconstructs the full denoised reads. In the choice between short and long amplicons, we suggest a hybrid approach using short amplicons for sampling breadth and depth, and long amplicons to characterize the local species pool for improved identification and phylogenetic analyses.

Three new species of Inosperma (Agaricales, Inocybaceae) from Tropical Africa.

Here, we describe three new species of Inosperma from Tropical Africa: Inosperma africanum, I. bulbomarginatum and I. flavobrunneum. Morphological and molecular data show that these species have not been described before, hence need to be described as new. The phylogenetic placements of these species were inferred, based on molecular evidence from sequences of 28S and RPB2. Additional analysis using ITS dataset shows interspecific variation between each species. Phylogenetic analyses resolve I. flavobrunneum in Old World Tropical clade 1 with weak support, I. bulbomarginatum is sister of Old World Tropical clade 1 and I. africanum is indicated as sister to the rest of Inosperma. Complete description and illustrations, including photographs and line drawings, are presented for each species. A new combination of Inocybe shawarensis into Inosperma is also proposed.

Morphology, Multilocus Phylogeny, and Toxin Analysis Reveal Amanita albolimbata, the First Lethal Amanita Species From Benin, West Africa.

Many species of Amanita sect. Phalloideae (Fr.) Quél. cause death of people after consumption around the world. Amanita albolimbata, a new species of A. sect. Phalloideae from Benin, is described here. The taxon represents the first lethal species of A. sect. Phalloideae known from Benin. Morphology and molecular phylogenetic analyses based on five genes (ITS, nrLSU, rpb2, tef1-α, and ß-tubulin) revealed that A. albolimbata is a distinct species. The species is characterized by its smooth, white pileus sometimes covered by a patchy volval remnant, a bulbous stipe with a white limbate volva, broadly ellipsoid to ellipsoid, amyloid basidiospores, and abundant inflated cells in the volva. Screening for the most notorious toxins by liquid chromatography-high-resolution mass spectrometry revealed the presence of α-amanitin, ß-amanitin, and phallacidin in A. albolimbata.

Rhizoglomus dalpeae, R. maiae, and R. silesianum, new species.

We examined three arbuscular mycorrhizal fungi (AMF; phylum Glomeromycota) producing glomoid spores. The mode of formation and morphology of these spores suggested that they represent undescribed species in the genus Rhizoglomus of the family Glomeraceae. Subsequent morphological studies of the spores and molecular phylogenetic analyses of sequences of the nuc rDNA small subunit (18S), internal transcribed spacer (ITS1-5.8S-ITS2 = ITS), and large subunit (28S) region (= 18S-ITS-28S) confirmed the suggestion and indicated that the fungi strongly differ from all previously described Rhizoglomus species with known DNA barcodes. Consequently, the fungi were described here as new species: R. dalpeae, R. maiae, and R. silesianum. Two of these species lived hypogeously in the field in habitats subjected to strong environmental stresses. Rhizoglomus dalpeae originated from an inselberg located within Guineo-Sudanian transition savanna zone in Benin, West Africa, where the temperature of the inselberg rock during a 5-mo drought ranges from 40 to 60 C. Rhizoglomus silesianum originated from a coal mine spoil heap in Poland, whose substrate is extremely poor in nutrients, has unfavorable texture, and may heat up to 50 C. By contrast, R. maiae was found in more favorable habitat conditions. It produced an epigeous cluster of spores among shrubs growing in a tropical humid reserve in Brazil. Moreover, the compatibility of phylogenies of species of the family Glomeraceae reconstructed from analyses of sequences of 18S-ITS-28S and the largest subunit of RNA polymerase II (RPB1) gene was discussed.

Two new African siblings of Pulveroboletusravenelii (Boletaceae).

This paper sorts out the taxonomy of species affiliated with Pulveroboletusravenelii in the Guineo-soudanian and Zambezian woodlands of Africa. Morphological and genetic characters of African Pulveroboletus collections were studied and compared to those of North American and Asian species. A phylogenetic analysis showed that the African specimens form a subclade, sister to the Asian and American taxa. Although clamp connections have previously never been reported from Pulveroboletus, all specimens of the African subclade show very small clamp connections. Two new African species, Pulveroboletusafricanus sp. nov. and P.sokponianus sp. nov., are described and illustrated. Comments concerning morphology and identification, as well as distribution and ecology, are given for both species.

Phylogenetic and microscopic studies in the genus Lactifluus (Basidiomycota, Russulales) in West Africa, including the description of four new species.

Despite the crucial ecological role of lactarioid taxa (Lactifluus, Lactarius) as common ectomycorrhiza formers in tropical African seasonal forests, their current diversity is not yet adequately assessed. During the last few years, numerous lactarioid specimens have been sampled in various ecosystems from Togo (West Africa). We generated 48 ITS sequences and aligned them against lactarioid taxa from other tropical African ecozones (Guineo-Congolean evergreen forests, Zambezian miombo). A Maximum Likelihood phylogenetic tree was inferred from a dataset of 109 sequences. The phylogenetic placement of the specimens, combined with morpho-anatomical data, supported the description of four new species from Togo within the monophyletic genus Lactifluus: within subgen. Lactifluus (L. flavellus), subgen. Russulopsis (L. longibasidius and L. pectinatus), and subgen. Edules (L. melleus). This demonstrates that the current species richness of the genus is considerably higher than hitherto estimated for African species and, in addition, a need to redefine the subgenera and sections within it.

Fungal biogeography. Global diversity and geography of soil fungi.

Fungi play major roles in ecosystem processes, but the determinants of fungal diversity and biogeographic patterns remain poorly understood. Using DNA metabarcoding data from hundreds of globally distributed soil samples, we demonstrate that fungal richness is decoupled from plant diversity. The plant-to-fungus richness ratio declines exponentially toward the poles. Climatic factors, followed by edaphic and spatial variables, constitute the best predictors of fungal richness and community composition at the global scale. Fungi show similar latitudinal diversity gradients to other organisms, with several notable exceptions. These findings advance our understanding of global fungal diversity patterns and permit integration of fungi into a general macroecological framework.

The genus Lactarius s. str. (Basidiomycota, Russulales) in Togo (West Africa): phylogeny and a new species described.

Lactarius s. str. represents a monophyletic group of about 40 species in tropical Africa, although the delimitation of the genus from Lactifluus is still in progress. Recent molecular phylogenetic and taxonomic revisions have led to numerous changes in names of tropical species formerly referred to Lactarius. To better circumscribe the genus Lactarius in Togo, we combined morphological data with sequence analyses and phylogeny inference of rDNA ITS sequences. Morphological and molecular data were generated from specimens sampled in various native woodlands and riverside forests; Lactarioid- and Russula sequences from public GenBank NCBI, and UNITE are included for phylogenetic analysis. The Maximum likelihood phylogeny tree inferred from aligned sequences supports the phylogenetic position of the studied samples from Togo within the subgenera Piperites, and Plinthogali. Lactarius s. str. includes about 13 species described from West Africa, of which eight were not previously known from Togo, including one new species: Lactarius subbaliophaeus identifiable by the presence of winged basidiospores, a pallisadic pileipellis with a uprapellis composed of cylindrical cells, inconspicuous pleurocystidia, and fusiform or tortuous, often tapering apex marginal cells. It can also be recognised by a transparent white latex that turns pinkish and then blackish, and a bluish reaction of the flesh context with FeSO4. These features mentioned do not match any of the morpho-anatomically most similar species, notably L. baliophaeus and L. griseogalus.

The status of mycology in Africa: A document to promote awareness.

The African Mycological Association (AMA) promotes mycology amongst members in Africa and globally. The AMA has about 200 members, mostly from African states but also with strong representation from Europe and USA, amongst others. Recent efforts by members of the AMA focused on reviving and developing mycological research and networking in Africa. A great deal must, however, still be done to promote the AMA under African mycologists, and those elsewhere with interests in Africa. African mycologists also experience challenges typical of the developing world and a great deal of fungi still needs to be discovered. This can also be seen as representing great opportunities for research and collaboration. Several issues pertinent to mycology in Africa were discussed during Special Interest Group sessions of the 9th International Mycological Congress in 2010, and through several opinion pieces contributed by AMA members in the AMA newsletter, MycoAfrica. This contribution serves as a document to summarise these in a form that can be presented to fellow mycologists, biologists and other scientists, relevant government departments, funding bodies and Non-Governmental Organizations and that pins down the importance of mycology, the status thereof in Africa and the need to promote it more.