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A-E: Siderophilous granulation during maturation of basidia from Ossicaulis lignatilis stained with iron-acetocarmine. A. Mature basidium (2N), B. Basidium after first nuclear division. C., D. and E. Basidia after second nuclear division (phase contrast microscopy, photo: Heinz Clémençon).
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We explored evolutionary relationships within the Lyophyllaceae by combining sequence data from six loci. The most likely phylogram led us to reconsider the Lyophyllaceae classification with the recognition of two new genera (Myochromella and Sagaranella) based on ecological and/or morphological distinctiveness. Lyophyllaceae are ecologically highl...
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... the basidia of Clitocybe candicans and C. subditopoda with iron-acetocarmine (Clémençon, 1978), granules are absent from the basidia of these two species. Performing the same coloration on basidia of O. lignatilis, a few, very small granules are seen in phase contrast that can easily escape attention when observed in bright field microscopy ( Fig. 2). ...
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... This solution would be systematically advantageous if Lyophyllaceae sensu Matheny et al. (2006) only included taxa exhibiting granulation of the macro-type, character largely viewed as the key taxonomic feature for this family. However, Lyophyllaceae as delimited by Matheny et al. (2006) includes Ossicaulis with granulation of the oligo-type (Fig. 2) and Hypsizygus tessulatus (analyses not shown) lacking granulation (Singer, 1947). Also the "hemilyophylloid" clade is clearly separated from the Tricholomataceae and supported to be monophyletic with the Lyophyllaceae (ML-BS: 71%; PP: 1.0). We refrain from proposing a new genus(era) for species of this clade considering our poor ...
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... only S. paluster (Peck) Redhead & V. Hofstetter] and Ossicaulis lachnopus ) and ruderal species (in a new genus Sagaranella represented by Tephrocybe tylicolor and T. gibberosa , and Tricholomella constricta ). Within this clade only terminal relationships are well supported ( Asterophora [ML-BS100%; PP: 1.0]; Asterophora + Tricholomella [ML-BS: 98%; PP: 1.0]; Sagaranella [ML- BS: 100%; PP: 1.0]). The two species, O. lachnopus (primarily saprotrophic) and Sphagnurus paluster (a necroparasite), remain unresolved within that clade. (4) A strongly supported “lyophylloid” clade (ML-BS: 100%; PP: 1.0) that includes Lyophyllum sensu Redhead et al. (2006), with its conserved type. Finally the monophyly of the “termitomycetoid” and “calocyboid” clades (ML-BS = –; PP: 0.95) is weakly supported. The present phylogeny (Fig. 1) suggests, however without significant support, a sister relationship between the Lyophyllaceae sensu Matheny et al. (2006) and the Tricholomataceae. This topology is inferred only by ML analysis of our 6-locus 51 taxa dataset. When using Bayesian inference, Entolomataceae are resolved and supported (PP = 0.95) as sister to Lyophyllaceae with Tricholomataceae occupying a more basal position (not shown). This last topology was also inferred by Matheny et al. (2006) but with C. candicans and C. subditipoda supported as basal to a monophyletic Entolomataceae- Lyophyllaceae clade based on maximum parsimony bootstrap value. Sánchez- García et al. (2014) retreived the Lyophyllaceae basal to Entolomataceae and Tricholomataceae sensu strictus. Their topoplogy suggests, however without support, the monophyly of part of Clitocybe , including C. candicans and C. subditipoda, with Lepista, Collybia and Lyophyllaceae. The present study (Fig. 1) also resolves C. candicans and C. subditipoda at the base of Lyophyllaceae but infer their monophyly with two species that have been previously classified in Lyophyllaceae sensu auct. : Clitocybe connata (Schum.: Fr.) Gillet ( = Lyophyllum connatum (Schum.: Fr.) Singer) and Hypsizygus ulmarius (Bull.: Fr.) Redhead (= Lyophyllum ulmarium [Bull.: Fr.] Kühner). The monophyly of Clitocybe connata and C. candicans has also been inferred in a very recent study (Alvarado et al ., in press) and led the authors to propose genus Leucocybe Vizzini, P. Alvarado, G. Moreno & Consiglio for these two species. Three other recent publications have examined elements of the “Tricholomatoid” clade (Vizzini, Musumeci and Murat, 2010; Vizzini and Ercole, 2012; Yu, Deng and Yao, 2011). These studies inferred a sister relationship between Hypsizygus and Ossicaulis with significant support. The placement of Hypsizygus appears therefore incongruent between previous studies and the phylogeny depicted here (Fig. 1). However, these previous studies only sampled Hypsizygus tessulatus , type species of this genus, while the present study only sampled H. ulmarius . On the other hand, Holec and Kolarík (2013) found H. ulmarius to be monophyletic with maximum support with H. marmoreus . To check for the phylogenetic placement of H. tessulatus , we introduced the sequence data available in GenBank for this taxon in our dataset (AFTOL-ID 1898; RPB 1: DQ917665, nucLSU: DQ917664, ITS: DQ917653) and ran maximum parsimony (MP) analyses. The best MP topology (a single MP tree: length = 12793 steps, CI = 0.228, HI = 0.772, RC = 0.1025; tree not shown) places H. tessulatus sister to Ossicaulis lachnopus with significant support (MP-BS: 98%). Consequently the genus Hypsizygus, as presently delimited, appears to be polyphyletic but the identities of the source materials should be confirmed. The “hemilyophylloid” clade (Fig. 1) includes two species traditionally classified in Lyophyllaceae ( Hypsizygus ulmarius and Clitocybe connata (= Lyophyllum connatum (Schum.: Fr.) Singer), which questions the delimitation of Lyophyllaceae sensu Matheny et al. (2006). These two species have both been included in tribe Lyophylleae (Kühner 1953, Moser 1978, Singer 1986) or family Lyophyllaceae (Bon, 1999) because they exhibit granulation in their basidia, a key character for that tribe or family. However these two species exhibit a granulation of the oligo-type (Clémençon, 1978) compared to the typical Lyophyllaceae, which all exhibit a macro-type granulation except for Ossicaulis in which granulation is absent according to Singer (1947). As Clitocybe candicans and C. subditipoda cluster in the same clade as C. connata , we therefore checked for the presence or absence of granulation in the basidia of these two species and in the basidia of Ossicaulis sp. , O. lignatilis. Staining the basidia of Clitocybe candicans and C. subditopoda with iron-acetocarmine (Clémençon, 1978), granules are absent from the basidia of these two species. Performing the same coloration on basidia of O. lignatilis , a few, very small granules are seen in phase contrast that can easily escape attention when observed in bright field microscopy (Fig. 2). Two possible systematic solutions can be considered: a first solution would be to keep the delimitation of Lyophyllaceae as recovered in Matheny et al . (2006) and Sánchez-García et al. (2014), which would leave the clade including H. ulmarius and part of Clitocybe including C. connata unnamed . This solution would be systematically advantageous if Lyophyllaceae sensu Matheny et al. (2006) only included taxa exhibiting granulation of the macro-type, character largely viewed as the key taxonomic feature for this family. However, Lyophyllaceae as delimited by Matheny et al . (2006) includes Ossicaulis with granulation of the oligo-type (Fig. 2) and Hypsizygus tessulatus (analyses not shown) lacking granulation (Singer, 1947). Also the “hemilyophylloid” clade is clearly separated from the Tricholomataceae and supported to be monophyletic with the Lyophyllaceae (ML-BS: 71%; PP: 1.0). We refrain from proposing a new genus(era) for species of this clade considering our poor sampling of genera Clitocybe and Hypsisygus , the apparent polyphyly of genus Hypsizygus , and the current lack of characters to predict delimitation of the “hemilyophylloid” clade. More extensive taxon and gene sampling for the Lyophyllaceae- Tricholomataceae-Entolomataceae clade are needed to clarify the delimitation of Lyophyllaceae and its relationship with closely related families. Previous studies (Hofstetter et al. 2002; Redhead et al. , 2006) resulted in maintaining the name Lyophyllum with a conserved type, L. semitale , and to place Lyophyllum leucophaeatum in Calocybe as C. gangraenosa (Redhead et al. , 2012). Lyophyllum sensu stricto , once segregated from Lyophyllum sensu lato , forms a monophyletic lineage, the “lyophylloid clade” (Fig. 1), composed only of staining species and more or less characterized by brown or dusky pigments and presence of siderophilous lysosomes of the macro-type (Clémençon, 1978). However Lyophyllum s.s. includes some species from section Tephrophana (Fr.) Singer (now frequently classified in Tephrocybe: i.e. Lyophyllum ambustum, L. anthracophilum and L. atratum; Fig. 1). Tephrocybe , as defined by Donk (1962), was composed of small, more or less hygrophanous, collybioid taxa, several of which have since been transferred back into Lyophyllum . One taxonomic and nomenclatural solution to resolve the conflict between named taxa and results from inferred phylogenies would be to follow Singer (1986) and to lump all Lyophyllaceae species, except the ones that are part of the “hemilyophylloid” clade, under a single generic name. However, a single genus Lyophyllum would not be a reasonable choice based on the following considerations. (1) Several biologically well-characterized taxa are recognized phylogenetically in the complex of siderophilous granule producing agarics (Hofstetter et al., 2002; Moncalvo et al. , 2002), the most well-known, conspicuous genus being Termitomyces (Clémençon, 1984). Termitomyces is the famous agaric genus assiduously cultivated by termites, about which considerable literature has been written (Aanen et al. , 2002; Aanen and Eggleton, 2005; Heim, 1977; Johnson et al. 1981; Nobre and Aanen, 2010; Nobre et al. , 2010). The type for the generic name Podabrella , T. microcarpus (Berk and Broome) R. Heim, is nested by molecular phylogenetic analysis within the greater Termitomyces lineage (Frøslev et al. , 2003), and therefore we follow Heim (1977) and Pegler (1977) who do not recognize Podabrella as a distinct genus (Table 1). The closest ally to the Termitomyces clade is Tephrocybe rancida (Fr.) Donk (Fig. 1), type species of Tephrocybe . This species, which occurs in soil in coniferous forests, is characterized by a long pseudorrhiza and virtually free lamellae (as in Termitomyces ), but unlike Termitomyces has clamp connections. We restrict Termitomyces to clampless taxa with conspicuous hymenial cystidia and mutualistic association with termites. Tephrocybe is a clamp forming sister taxon to the clampless genus Termitomyces and is not associated with termites. Two recently described conidia-producing genera Arthromyces Baroni & Lodge (clampless) [ non “ Arthromyces ” Amano nom. invalid .] and Blastosporella Baroni & Franco-Molano (clamp connections present on hyphae of pileipellis and stipitipellis) are probably part of this “termitomycetoid” clade but their relationships toward the other genera that are part of this clade remain unknown (Baroni et al. , 2006). (2) Two additional taxa previously placed in either Tephrocybe or Lyophyllum , i.e. T. bouderi and T. inolens , like Tephrocybe rancida and Termitomyces species, have nearly free lamellae, and are neither obvious parasites, nitrophiles, nor are they cultivated by insects. Together they form another monophyletic group, possibly with affinities to Tephrocybe and Termitomyces (Fig. 1). Unlike T. rancida , their basidiomata do not originate from a pseudorrhizae deep in the ground. It could reasonably be argued ...
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... al. (2006), with its conserved type. Finally the monophyly of the “termitomycetoid” and “calocyboid” clades (ML-BS = –; PP: 0.95) is weakly supported. The present phylogeny (Fig. 1) suggests, however without significant support, a sister relationship between the Lyophyllaceae sensu Matheny et al. (2006) and the Tricholomataceae. This topology is inferred only by ML analysis of our 6-locus 51 taxa dataset. When using Bayesian inference, Entolomataceae are resolved and supported (PP = 0.95) as sister to Lyophyllaceae with Tricholomataceae occupying a more basal position (not shown). This last topology was also inferred by Matheny et al. (2006) but with C. candicans and C. subditipoda supported as basal to a monophyletic Entolomataceae- Lyophyllaceae clade based on maximum parsimony bootstrap value. Sánchez- García et al. (2014) retreived the Lyophyllaceae basal to Entolomataceae and Tricholomataceae sensu strictus. Their topoplogy suggests, however without support, the monophyly of part of Clitocybe , including C. candicans and C. subditipoda, with Lepista, Collybia and Lyophyllaceae. The present study (Fig. 1) also resolves C. candicans and C. subditipoda at the base of Lyophyllaceae but infer their monophyly with two species that have been previously classified in Lyophyllaceae sensu auct. : Clitocybe connata (Schum.: Fr.) Gillet ( = Lyophyllum connatum (Schum.: Fr.) Singer) and Hypsizygus ulmarius (Bull.: Fr.) Redhead (= Lyophyllum ulmarium [Bull.: Fr.] Kühner). The monophyly of Clitocybe connata and C. candicans has also been inferred in a very recent study (Alvarado et al ., in press) and led the authors to propose genus Leucocybe Vizzini, P. Alvarado, G. Moreno & Consiglio for these two species. Three other recent publications have examined elements of the “Tricholomatoid” clade (Vizzini, Musumeci and Murat, 2010; Vizzini and Ercole, 2012; Yu, Deng and Yao, 2011). These studies inferred a sister relationship between Hypsizygus and Ossicaulis with significant support. The placement of Hypsizygus appears therefore incongruent between previous studies and the phylogeny depicted here (Fig. 1). However, these previous studies only sampled Hypsizygus tessulatus , type species of this genus, while the present study only sampled H. ulmarius . On the other hand, Holec and Kolarík (2013) found H. ulmarius to be monophyletic with maximum support with H. marmoreus . To check for the phylogenetic placement of H. tessulatus , we introduced the sequence data available in GenBank for this taxon in our dataset (AFTOL-ID 1898; RPB 1: DQ917665, nucLSU: DQ917664, ITS: DQ917653) and ran maximum parsimony (MP) analyses. The best MP topology (a single MP tree: length = 12793 steps, CI = 0.228, HI = 0.772, RC = 0.1025; tree not shown) places H. tessulatus sister to Ossicaulis lachnopus with significant support (MP-BS: 98%). Consequently the genus Hypsizygus, as presently delimited, appears to be polyphyletic but the identities of the source materials should be confirmed. The “hemilyophylloid” clade (Fig. 1) includes two species traditionally classified in Lyophyllaceae ( Hypsizygus ulmarius and Clitocybe connata (= Lyophyllum connatum (Schum.: Fr.) Singer), which questions the delimitation of Lyophyllaceae sensu Matheny et al. (2006). These two species have both been included in tribe Lyophylleae (Kühner 1953, Moser 1978, Singer 1986) or family Lyophyllaceae (Bon, 1999) because they exhibit granulation in their basidia, a key character for that tribe or family. However these two species exhibit a granulation of the oligo-type (Clémençon, 1978) compared to the typical Lyophyllaceae, which all exhibit a macro-type granulation except for Ossicaulis in which granulation is absent according to Singer (1947). As Clitocybe candicans and C. subditipoda cluster in the same clade as C. connata , we therefore checked for the presence or absence of granulation in the basidia of these two species and in the basidia of Ossicaulis sp. , O. lignatilis. Staining the basidia of Clitocybe candicans and C. subditopoda with iron-acetocarmine (Clémençon, 1978), granules are absent from the basidia of these two species. Performing the same coloration on basidia of O. lignatilis , a few, very small granules are seen in phase contrast that can easily escape attention when observed in bright field microscopy (Fig. 2). Two possible systematic solutions can be considered: a first solution would be to keep the delimitation of Lyophyllaceae as recovered in Matheny et al . (2006) and Sánchez-García et al. (2014), which would leave the clade including H. ulmarius and part of Clitocybe including C. connata unnamed . This solution would be systematically advantageous if Lyophyllaceae sensu Matheny et al. (2006) only included taxa exhibiting granulation of the macro-type, character largely viewed as the key taxonomic feature for this family. However, Lyophyllaceae as delimited by Matheny et al . (2006) includes Ossicaulis with granulation of the oligo-type (Fig. 2) and Hypsizygus tessulatus (analyses not shown) lacking granulation (Singer, 1947). Also the “hemilyophylloid” clade is clearly separated from the Tricholomataceae and supported to be monophyletic with the Lyophyllaceae (ML-BS: 71%; PP: 1.0). We refrain from proposing a new genus(era) for species of this clade considering our poor sampling of genera Clitocybe and Hypsisygus , the apparent polyphyly of genus Hypsizygus , and the current lack of characters to predict delimitation of the “hemilyophylloid” clade. More extensive taxon and gene sampling for the Lyophyllaceae- Tricholomataceae-Entolomataceae clade are needed to clarify the delimitation of Lyophyllaceae and its relationship with closely related families. Previous studies (Hofstetter et al. 2002; Redhead et al. , 2006) resulted in maintaining the name Lyophyllum with a conserved type, L. semitale , and to place Lyophyllum leucophaeatum in Calocybe as C. gangraenosa (Redhead et al. , 2012). Lyophyllum sensu stricto , once segregated from Lyophyllum sensu lato , forms a monophyletic lineage, the “lyophylloid clade” (Fig. 1), composed only of staining species and more or less characterized by brown or dusky pigments and presence of siderophilous lysosomes of the macro-type (Clémençon, 1978). However Lyophyllum s.s. includes some species from section Tephrophana (Fr.) Singer (now frequently classified in Tephrocybe: i.e. Lyophyllum ambustum, L. anthracophilum and L. atratum; Fig. 1). Tephrocybe , as defined by Donk (1962), was composed of small, more or less hygrophanous, collybioid taxa, several of which have since been transferred back into Lyophyllum . One taxonomic and nomenclatural solution to resolve the conflict between named taxa and results from inferred phylogenies would be to follow Singer (1986) and to lump all Lyophyllaceae species, except the ones that are part of the “hemilyophylloid” clade, under a single generic name. However, a single genus Lyophyllum would not be a reasonable choice based on the following considerations. (1) Several biologically well-characterized taxa are recognized phylogenetically in the complex of siderophilous granule producing agarics (Hofstetter et al., 2002; Moncalvo et al. , 2002), the most well-known, conspicuous genus being Termitomyces (Clémençon, 1984). Termitomyces is the famous agaric genus assiduously cultivated by termites, about which considerable literature has been written (Aanen et al. , 2002; Aanen and Eggleton, 2005; Heim, 1977; Johnson et al. 1981; Nobre and Aanen, 2010; Nobre et al. , 2010). The type for the generic name Podabrella , T. microcarpus (Berk and Broome) R. Heim, is nested by molecular phylogenetic analysis within the greater Termitomyces lineage (Frøslev et al. , 2003), and therefore we follow Heim (1977) and Pegler (1977) who do not recognize Podabrella as a distinct genus (Table 1). The closest ally to the Termitomyces clade is Tephrocybe rancida (Fr.) Donk (Fig. 1), type species of Tephrocybe . This species, which occurs in soil in coniferous forests, is characterized by a long pseudorrhiza and virtually free lamellae (as in Termitomyces ), but unlike Termitomyces has clamp connections. We restrict Termitomyces to clampless taxa with conspicuous hymenial cystidia and mutualistic association with termites. Tephrocybe is a clamp forming sister taxon to the clampless genus Termitomyces and is not associated with termites. Two recently described conidia-producing genera Arthromyces Baroni & Lodge (clampless) [ non “ Arthromyces ” Amano nom. invalid .] and Blastosporella Baroni & Franco-Molano (clamp connections present on hyphae of pileipellis and stipitipellis) are probably part of this “termitomycetoid” clade but their relationships toward the other genera that are part of this clade remain unknown (Baroni et al. , 2006). (2) Two additional taxa previously placed in either Tephrocybe or Lyophyllum , i.e. T. bouderi and T. inolens , like Tephrocybe rancida and Termitomyces species, have nearly free lamellae, and are neither obvious parasites, nitrophiles, nor are they cultivated by insects. Together they form another monophyletic group, possibly with affinities to Tephrocybe and Termitomyces (Fig. 1). Unlike T. rancida , their basidiomata do not originate from a pseudorrhizae deep in the ground. It could reasonably be argued that T. rancida and the T. boudieri-inolens lineage represent together the ancestral group from which Termitomyces originated. Therefore both T. boudieri (Kühner and Romagn.) Derbsch and T. inolens (Fr.) M.M. Moser ought to be retained in Tephrocybe , thus possibly making the genus paraphyletic. However, given an obvious morphological feature (radicating pseudorrhiza) that might be linked to a biological function for T. rancida (unknown hypogeous food source?) shared by T. rancida and Termitomyces , but absent in their sister taxa ( Myochromella gen. nov.) we opt to recognize a separate monotypic small genus, Mycochromella (Fig. 1). This leaves ...
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A wild edible mushroom survey was carried out during latter part of the 2013 rainy season, from 13 August to 16 September in forests near six villages in Phoukout District, Xieng Khouang Province, Lao PDR. Two thirds of the village households (ranging from 10% to 98%) were engaged in collecting wild mushrooms. During the relative short survey perio...
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... The genus Termitomyces R. Heim belongs to the family Lyophyllaceae Jülich (Moncalvo et al. 2000). The mushroom genera in the family have diverse ecologies, encompassing saprotrophic, parasitic to symbiotic lifestyles (Hofstetter et al. 2014). The generic name 'Termitomyces' introduced by Roger Heim refers to the mutualistic symbiotic association of the fungus and termite species (Heim 1942). ...
ei M.N. (2024) The genus Termitomyces: outline, phylogeny, and divergence times estimation with description of a new edible species from Arabian Peninsula.-Sydowia 76: 185-198. The termite-cultivated mushroom genus Termitomyces belongs to the family Lyophyllaceae (Agaricales, Basidiomycota). In this study, we present a phylogeny of Termitomyces based on internal transcribed spacer (ITS1-5.8S-ITS2 = ITS), large subunit of nuclear ribosomal DNA (28S) and translation elongation factor 1 alpha (TEF-1α). The ancestral age of Termitomyces is estimated around 23 million years. Molecular clock analyses also indicate the presence of two clades, each clade with approximate age of 18 million years. We think that these clades can be treated for the infrageneric classification of Termitomyces. We also describe a new species in the genus, Termitomyces dhofarensis from Dhofar region, located in southern Oman. The new species is edible. Local people have been collecting it for generations for food and even trade it in the local markets. The species is described here based on a combination of morphological features of basidiomata and multigene phylogenetic analyses. The divergence time is estimated to be around 1.2 million years for Termitomyces dhofarensis. This is the first report of the genus from Arabian Peninsula .
... Sequences of a total of 78 specimens representing 25 species (Table1). Calocybe persicolor and C. carnea were selected as the outgroup (Hofstetter et al. 2014). Sequences were aligned using MAFFT version 7 (Katoh et al. 2013) first and then minimally adjusted by hand in BioEdit v.7.0.4 (Hall 2007). ...
Lyophyllum lixivium sp. nov., collected from southwestern China, is described, illustrated, and discussed based on morphological characteristics and molecular data (ITS and nrLSU). Lyophyllum lixivium has umbonate to slightly depressed pileus, a cylindrical stipe that slightly enlarges towards the base, fuzz at the base, basidiospores mostly broadly ellipsoid, rarely fusiform lacrymoid or amygdaliform with an acute apex in side view, abundant clamp connections at the pileus surface and hymenophoral trama. Morphologically and phylogenetically similar species were compared with L. lixivium in this study.
... It currently contains about thirty genera, including ectomycorrhizal and nonectomycorrhizal groups (Sánchez-García et al. 2014, Sánchez-García 2016. In addition to the Tricholomataceae (Sánchez-García et al. 2014), it currently encompasses another 10 families: Asproinocybaceae (Bau & Mou 2021), Biannulariaceae (Vizzini et al. 2020a), Callistosporiaceae (Vizzini et al. 2020a), Clitocybaceae (Alvarado et al. 2015, Vizzini et al. 2020b, Entolomataceae (Kluting et al. 2014), Fayodiaceae (Moncalvo et al. 2002), Lyophyllaceae (Hofstetter et al. 2014, Bellanger et al. 2015, Macrocystidiaceae (Dentinger et al. 2016), Omphalinaceae (Vizzini et al. 2020b), and Pseudoclitocybaceae (Alvarado et al. 2018a). ...
... Still, many white-spored clitocyboid and tricholomatoidlooking genera cannot be easily classified within any of these families, and even their position inside suborder Tricholomatineae cannot be confirmed with phylogenetic analyses because of the incomplete data available from some of them (mostly ribosomal DNA sequences). For example, the classification of Asproinocybe, Aspropaxillus, Dendrocollybia, Giacomia, Hertzogia, Hygrophorocybe, Infundibulicybe, Lepistella, Leucocortinarius, Notholepista, Omphaliaster, Omphalina, Paralepista, Paralepistopsis, Pseudoclitopilus, Pseudoomphalina, Resupinatus, Rimbachia, Ripartites, Trichocybe or Tricholosporum is not fully clear , b, 2020, Hofstetter et al. 2014, Sánchez-García et al. 2014, Vizzini 2014a, Alvarado et al. 2018a, b, He et al. 2019, Raj et al. 2019, Varga et al. 2019, Kalichman et al. 2020, Olariaga et al. 2020, He & Yang 2022, Wiest 2022. The classification of these incertae sedis lineages requires the reconstruction of the phylogeny of the entire order Agaricales. ...
... Bayesian analysis of Dataset 1, order Agaricales (Figs 1-3), significantly supported the following hypotheses: 1) family Typhulaceae has a basal position to the remaining suborders analyzed; 2) seven main clades with a significant monophyletic origin were found, matching suborders Agaricineae, Pleurotineae (including Schizophyllineae), Pluteineae, Hygrophorineae, Marasmiineae, Phyllotopsidineae (including Aphroditeola and Sarcomyxineae) and Tricholomatineae; 3) suborder Pleurotineae also encompasses the families Fistulinaceae and Schizophyllaceae, and so it could be considered a synonym of Schizophyllineae; 4) suborder Pluteineae includes Amanitaceae and Leucocortinarius (PP 0.99), as well as the families Pluteaceae, Limnoperdaceae, a strongly supported clade (1.00 PP, 98 % BP) consisting of Melanoleuca and Giacomia, and another including Volvariella; 5) suborder Tricholomatineae has at least twelve families: Macrocystidiaceae (type Macrocystidia, probably related to Pseudoclitopilus); Omphalinaceae (including Infundibulicybe and Omphalina), Pseudoclitocybaceae (including Aspropaxillus), Fayodiaceae (here including Fayodia and Gamundia, but probably also Caulorhiza, Conchomyces and Myxomphalia according to Moncalvo et al. 2002); Biannulariaceae, Callistosporiaceae, Tricholomataceae, Clitocybaceae, Lyophyllaceae sensu lato, Entolomataceae, as well as the unclassified lineages of Neohygrocybe/Pseudoomphalina, Paralepistopsis, Hertzogia and the clade formed by Notholepista, Ripartites, and Paralepista; 6) family Clitocybaceae includes the genera Clitocybe sensu stricto, Lepista, Singerocybe, Collybia sensu lato , and the lineage of C. ditopa; 7) family Lyophyllaceae sensu lato is integrated by Lyophyllaceae sensu stricto as well as the socalled hemilyophylloid lineages (Binder et al. 2010, Hofstetter et al. 2014, including the genera Asproinocybe/Tricholosporum (family Asproinocybaceae), Atractosporocybe, Clitolyophyllum, Leucocybe, Omphaliaster, Rhizocybe, Trichocybe, and several species whose generic status needs to be reviewed; 8) family Mycenaceae is part of the suborder Marasmiineae, where it is sister to the significant clade formed by Xeromphalina and Heimiomyces; 9) the previous concepts of the genera Mycena and Hemimycena are polyphyletic; 10) genus Hygrophorocybe is nested inside suborder Hygrophorineae. ...
The phylogenetic position of several clitocyboid/pleurotoid/tricholomatoid genera previously considered incertae sedis is here resolved using an updated 6-gene dataset of Agaricales including newly sequenced lineages and more complete data from those already analyzed before. Results allowed to infer new phylogenetic relationships, and propose taxonomic novelties to accommodate them, including up to ten new families and a new suborder. Giacomia (for which a new species from China is here described) forms a monophyletic clade with Melanoleuca (Melanoleucaceae) nested inside suborder Pluteineae, together with the families Pluteaceae, Amanitaceae (including Leucocortinarius), Limnoperdaceae and Volvariellaceae. The recently described family Asproinocybaceae is shown to be a later synonym of Lyophyllaceae (which includes also Omphaliaster and Trichocybe) within suborder Tricholomatineae. The families Biannulariaceae, Callistosporiaceae, Clitocybaceae, Fayodiaceae, Macrocystidiaceae (which includes Pseudoclitopilus), Entolomataceae, Pseudoclitocybaceae (which includes Aspropaxillus), Omphalinaceae (Infundibulicybe and Omphalina) and the new families Paralepistaceae and Pseudoomphalinaceae belong also to Tricholomatineae. The delimitation of the suborder Pleurotineae (= Schizophyllineae) is discussed and revised, accepting five distinct families within it, viz. Pleurotaceae, Cyphellopsidaceae, Fistulinaceae, Resupinataceae and Schizophyllaceae. The recently proposed suborder Phyllotopsidineae (= Sarcomyxineae) is found to encompass the families Aphroditeolaceae, Pterulaceae, Phyllotopsidaceae, Radulomycetaceae, Sarcomyxaceae (which includes Tectella), and Stephanosporaceae, all of them unrelated to Pleurotaceae (suborder Pleurotineae) or Typhulaceae (suborder Typhulineae). The new family Xeromphalinaceae, encompassing the genera Xeromphalina and Heimiomyces, is proposed within Marasmiineae. The suborder Hygrophorineae is here reorganized into the families Hygrophoraceae, Cantharellulaceae, Cuphophyllaceae, Hygrocybaceae and Lichenomphaliaceae, to homogenize the taxonomic rank of the main clades inside all suborders of Agaricales. Finally, the genus Hygrophorocybe is shown to represent a distinct clade inside Cuphophyllaceae, and the new combination H. carolinensis is proposed.
... The genus Termitomyces R. Heim belongs to the family Lyophyllaceae Jülich (Moncalvo et al. 2000). The mushroom genera in the family have diverse ecologies, encompassing saprotrophic, parasitic to symbiotic lifestyles (Hofstetter et al. 2014). The generic name 'Termitomyces' introduced by Roger Heim refers to the mutualistic symbiotic association of the fungus and termite species (Heim 1942 Over time, the fungus-growing termites have expanded their distribution range from central Africa to sub-Saharan Africa (Bignell et al. 2010). ...
The termite cultivated mushroom genus Termitomyces belongs to the family Lyophyllaceae (Agaricales, Basidiomycota). In this study, we presented a phylogeny of Termitomyces based on internal transcribed spacer (ITS1-5.8S-ITS2 = ITS), large subunit of nuclear ribosomal DNA (28S) and translation elongation factor 1 alpha (TEF-1α). The ancestral age of Termitomyces was estimated around 23 million years. Molecular clock analyses also indicate the presence of two clades, each clade with approximate stem age of 18 million years. We think that these clades can be treated for the infrageneric classification of Termitomyces . Furthermore, we also described a new species in the genus, Termitomyces dhofarensis from Dhofar region, located in southern Oman. The new species is edible. Local people collect it for generations for food and even trade it in the local markets. The species is described here based on a combination of morphological features of basidiomata and multigene phylogenetic analyses. The divergence time is estimated to be around 1.2 million years for Termitomyces dhofarensis . This is the first report of the genus from Arabian Peninsula.
... Five newly generated and 64 retrieved sequences from GenBank were used as the ingroup. Three sequences of Calocybe gambosa, C. carnea and C. persicolor retrieved from GenBank were used as the outgroup [30]. The ITS was 575 characters in length, of which 284 characters were constant, 291 were variable but parsimony-uninformative and 231 were parsimony-informative. ...
Two new species, Lyophyllum yiqunyang and L. heimogu, that belong to the section Difformia of the genus Lyophyllum, are described based on collections from Tibetan areas, China. The two species are delicious edible low-temperature mushrooms and are widely collected and eaten by local people. Lyophyllum yiqunyang sp. nov. is saprotrophic and has medium-sized basidiomata, olive-grey pileus, cheilocystidia, absent pleurocystidia, globose to subglobose basidiospores (6.12–6.31 × 6.02–6.23 μm) and clamp connections at the pileus context, hymenophoral trama and stipe. Lyophyllum heimogu sp. nov. is saprotrophic and has a dark grey to olive pileus, medium-sized basidiomata and globose to subglobose basidiospores (5.31–5.63 × 5.22–5.41 μm). In the phylogenetic analyses, our two new species formed distinct clades that are well supported by posterior probabilities and bootstrap proportions. Detailed descriptions, colour photos, illustrations and a phylogenetic tree to show the positions of the two new species are presented.
... Singer retrieved from GenBank were used as outgroup (see Table 1). The outgroup taxa were selected based on the phylogeny in Hofstetter et al. (2014). The sequences were aligned with MAFFT version 7 (Katoh and Standley 2013) and checked in Bioedit version 7.0.5 (Hall 2007 Phylogenies and node support were first inferred by Maximum Likelihood (ML) from the three single-gene alignments separately, using RAxML-HPC2 version 8.2.12 (Stamatakis 2014) with 1,000 rapid bootstraps, as implemented on the Cipres portal (Miller et al. 2010). ...
Citation: Tang S-M, Vadthanarat S, He J, Raghoonundon B, Yu F-M, Karunarathna SC, Li S-H, Raspé O (2023) Morphological and molecular analyses reveal two new species of Termitomyces (Agaricales, Lyophyllaceae) and morphological variability of T. intermedius. MycoKeys 95: 61-82. https://doi.org/10.3897/mycokeys.95.97156 Abstract Two new species, Termitomyces tigrinus and T. yunnanensis are described based on specimens collected from southwestern China. Termitomyces yunnanensis is morphologically characterized by a conspicuously venose pileus surface that is grey, olive grey, light grey to greenish grey at center, light grey towards margin, and a cylindrical white stipe. Termitomyces tigrinus is morphologically characterized by a densely tomentose to tomentose-squamulose pileus showing alternating greyish white and dark grey zones, and a stipe that is bulbous at the base. The two new species are supported by phylogenetic analyses of combined nuclear rDNA internal transcribed spacer ITS1-5.8S-ITS2 rDNA (ITS), the mitochondrial rDNA small subunit (mrSSU) and the nuclear rDNA large subunit (nrLSU). The morphological variability of T. intermedius, including five specimens newly collected from Yunnan Province, China, is also discussed. The collections showed variability in colour of the stipe surface and in the shape of cheilocystidia when compared to the original description. Full descriptions of the two new species and of T. intermedius, as well as a taxonomic key to the 14 Termitomyces species reported from China are provided. A peer-reviewed open-access journal Launched to accelerate biodiversity research Song-Ming Tang et al. / MycoKeys 95: 61-82 (2023) 62
... While the importance of fungi as mutualists, decomposers, and pathogens is undisputed, researchers are just beginning to unravel the processes that shape their global species richness and distribution. Previous studies established the monophyly of Lyophyllaceae Jülich and positioned the family within the Tricholomatoid clade, and then Alvarado et al. revealed that Lyophyllaceae may be a putative wider concept or the existence of multiple lineages that are basal to it [1][2][3]. Several new genera established in the past decade are expected to reorganize the system, including Australocybe T.J. Baroni [1,4,5]. ...
... Previous studies established the monophyly of Lyophyllaceae Jülich and positioned the family within the Tricholomatoid clade, and then Alvarado et al. revealed that Lyophyllaceae may be a putative wider concept or the existence of multiple lineages that are basal to it [1][2][3]. Several new genera established in the past decade are expected to reorganize the system, including Australocybe T.J. Baroni [1,4,5]. In addition, some species of Lyophyllum, Calocybe Kühner, Hypsizygus Singer, and Termitomyces R. Heim have edible value [6], and a few species have medicinal and significant economic importance [7][8][9][10]. ...
... Hofstetter et al. found that ancestral states of Lyophyllaceae s. str. and s.l. were unequivocally reconstructed as saprotrophic, while parasitism, ectomycorrhiza, and insect association appear to be derived states in the evolution of Lyophyllaceae [1]. The new species described in this study occupy independent lineages in Lyophyllaceae. ...
The lyophylloid agarics are a group of ecologically highly diversified macrofungi, some of which are very popular edible mushrooms. However, we know little about lyophylloid species diversity in China. In this study, we described four new species from China: Lyophyllum atrofuscum, L. subalpinarum, L. subdecastes, and Ossicaulis sichuanensis. We conducted molecular phylogenetic analyses of Lyophyllaceae based on the nuclear ribosomal RNA gene (nLSU) and the internal transcribed spacer regions (ITS). Phylogenetic analyses by the maximum likelihood method and Bayesian inference showed that the four new species are unique monophyletic species. A key to the species of Lyophyllum from China and a key to Ossicaulis worldwide were given.
... The genus Tephrocybe as introduced by Donk (1962), as a segregate from the genus Lyophyllum P. Karst., contains a number of small collybioid mushrooms, often with a hygrophanous pileus and grey colours (Donk 1962). Various molecular studies have shown the genus to be paraphyletic, which is consistent with the lack of diagnostic features (Hofstetter et al. 2002;Hofstetter et al. 2014;Bellanger et al. 2015). The most recent study which included several members of Tephrocybe found eight distinct evolutionary origins within the Lyophyllaceae (Bellanger et al. 2015). ...
... Tephrocybe species have been transferred back to Lyophyllum, or have been placed in new genera such as Myochromella, Sagaranella and Sphagnurus (Hofstetter et al. 2014). A previous study showed that the type of the genus, Tephrocybe rancida, is part of a termitomycetoid clade which, next to the termite-associated genus Termitomyces, also contains the insect-faecal associated genera Arthromyces and Blastosporella (van de Peppel et al. 2021). ...
... These six criteria are (1) monophyly, (2) sufficient phylogenetic coverage, (3) sufficient branch support, (4) discussion of alternative options, (5) no single marker phylogeny (6) include all supporting evidence and background information. To those criteria we add one additional consideration, viz., the desirability that the approach taken for this clade within the Lyophyllaceae is consistent with earlier treatments of that family (Bellanger et al. 2015;Hofstetter et al. 2014). ...
Four new genera encompassing six new species are described in the Lyophyllaceae ( Agaricales ): the monotypic Australocybe from Australia; the monotypic Phaeotephrocybe from Belize; the monotypic Nigrocarnea from Laos containing the novel conidia-producing species N. radicata and Praearthromyces containing two Asian taxa, the conidia-producing P . corneri known from Malaysia and Singapore and P . griseus from Thailand, which lacks conidial production. In addition, we describe Arthromyces glabriceps , which is the first species in the genus Arthromyces that lacks conidial production on the basidiome. Alternative options for a classification of these lyophylloid taxa are discussed and based on morphological, ecological and biogeographical considerations rejected.
... However, the classification opinion proposed by Singer has not been fully supported by molecular data. In recent years, many phylogenetic investigations on Lyophyllaceae indicated that Calocybe belongs to a monophyletic group [5][6][7][8][9][10][11][12][13][14]. However, the infrageneric classification system of Calocybe remains unclear. ...
... ML and BI resulted in almost identical tree topologies, and the BI tree was selected for display ( Figure 1). The results suggested that the species of Calocybe formed a distinct group supported by a strong value (1.00 PP and 100% BS), corroborated by former research [5][6][7][8][9][10][11][12][13][14]. A total of five clades can be viewed within species of Calocybe, which is in line with Li et al. [11]. ...
... Phylogenetically, species of Calocybe appears to form a monophyletic group, which is consistent with previous studies [5][6][7][8][9][10][11][12][13][14]. Additionally, the three new species determined in this study occupy an independent position in the phylogenetic tree. ...
Three species, Calocybe lilacea, C. longisterigma, and C. subochraceus, were newly discovered in Inner Mongolia, Hunan, and Liaoning provinces of China. Calocybe lilacea is mainly characterized by its small-sized basidiocarp, brownish-orange pileus, lilac gray to dull violet stipe, and noncellular epicutis. The main characteristics of C. longisterigma are its light brown to brownish-orange pileus, long sterigmata, and noncellular epicutis. The main morphological features of C. subochraceus are its small-sized basidiocarp, pale-orange pileus, adnexed lamellae, tortuous stipe, and noncellular epicutis. These morphological features confirmed that the three species of Calocybe all belonged to Sect. Carneoviolaceae. Phylogenetic analysis based on a combined dataset (ITS–nrLSU–RPB2) determined that the three species belong to the genus Calocybe and form a distinct lineage. The morphological differences between the three new species and other related species of Calocybe are also discussed.
... Lyophyllaceae Jülich was suggested by recent molecular phylogenetic analyses to be a putative wider concept that includes several lineages that seem more distantly related, and the generic concepts should be reconsidered [1][2][3][4][5]. Several new genera established in the past ten years were expected to reorganize the system, viz. ...
... Several new genera established in the past ten years were expected to reorganize the system, viz. [2,[6][7][8][9][10]; however, a few of them show a poor species diversity worldwide. Clitolyophyllum was discovered in 2016, a Turkish species fruiting on the dead bark of Picea orientalis. ...
... [13,14]. Several recent molecular phylogenetic analyses have confirmed Calocybe as an independent genus with the merging of Rugosomyces [2,3,[15][16][17]. At least 57 species can be assigned to Calocybe according to Xu et al. [17]; however, the molecular phylogenetic relationships among members of the Calocybe were inconsistent with the morphological classification made by Singer [12]. ...
Four new species, viz. Calocybe coacta, C. fulvipes, C. vinacea and Clitolyophyllum umbilicatum, are described in northern China. Comparisons are made of macro- and micromorphological features among the new species and closely related species within the genus. The new species feature unique morphological characteristics that separate them from the previously described species. Calocybe coacta is characterized by medium- to large-sized basidiocarps, greyish cream, felty pileus and non-cellular epicutis. The key characteristics of C. fulvipes are rose-brown to greyish-brown pileus, stone-brown stipe and non-cellular epicutis. The unique morphological characteristics of C. vinacea that distinguish it from its closely related species are pastel red to dull-red pileus and stipe surface with densely white pruina. The main characteristics of Clitolyophyllum umbilicatum are deeply depressed dark orange to light-brown pileus, central stipe and subglobose-ellipsoid spores. Phylogenetic analyses based on the ITS and 28S regions indicated that the four new species are distinct and monophyletic. Full descriptions, color images, illustrations and a phylogenetic tree that show the placement of the four new species are provided. A key to the Calocybe species reported from China is also given.
