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Morphological characters of Asterinales and Asterotexiales in face view. (A-J) Development. Initiation of ascomata in Asterinales and Asterotexiales usually involves intercalary septation of a superficial generator hypha but is sometimes (D) uninterpretable, (E, F) from spores, or (G) from stomata. (H-J) The highly septate generator hypha usually persists above the scutellum throughout development (white arrowhead). (A-D, H-J) The generator hypha gives rise to the scutellum through (i) the formation of new, lateral hyphal tips that branch with isotomous dichotomies, occasionally (D, I, inset) with one tip that overlaps the other (o). (H, J) The appressed scutellum margin shows lightly pigmented tips and is usually crenulate at maturity (black arrowhead). (K) Hyphae of the appressed marginal of scutella Prillieuxina baccharidincola instead anastomose with surrounding mycelium (white arrowhead). (I, K, L-O) Dehiscence. (I, K, L-M) Asterinales and Asterotexiales ascomata usually dehisce with irregular slits. (K) Mature scutellum with irregular radial slits, black arrowheads. (L-M) Irregular slits follow the longitudinal walls of scutellum cells (white arrowheads). (N-O) Ostiolate openings are less common. (N) Ostiole in young scutellum. (O) Asexual or spermatial stage, with evenly pigmented cells lining ostiole. (P-S) Lateral appressoria showing its characteristic melanized ring at lower focal plane (insets, representing area in the dashed gray boxes). Scale bars: (A-D, H-I) 10 µm; (E-G) 20 µm; (J-K) 50 µm; (L-S) 5 µm. (A, I, J, P, Q) Asterina melastomatis VIC 52822, 1, 4, 6, 2. and 2 focal planes (f.p.), respectively; (B) Asterina chrysophylli VIC 42823, 4 f.p.; (C, O) Asterotexiaceae sp. CBS 143813, 2 f.p. each; (D) Hemigrapha atlantica BR 14014, 1 f.p.; (E, F) Asterotexis cucurbitacearum VIC 42814, 1 f.p. each; (G, L) Rhagadolobiopsis thelypteridis EG156; (H, R) Batistinula gallesiae VIC 42514, 10 and 2 focal planes (f.p.) respectively; (K, N), Prillieuxina baccharidincola VIC 42817; (M) Asterotexiaceae sp. UBC F33036, 11 f.p.; (S) Asterina sp. (VUL. 341b), 2 f.p.

Morphological characters of Asterinales and Asterotexiales in face view. (A-J) Development. Initiation of ascomata in Asterinales and Asterotexiales usually involves intercalary septation of a superficial generator hypha but is sometimes (D) uninterpretable, (E, F) from spores, or (G) from stomata. (H-J) The highly septate generator hypha usually persists above the scutellum throughout development (white arrowhead). (A-D, H-J) The generator hypha gives rise to the scutellum through (i) the formation of new, lateral hyphal tips that branch with isotomous dichotomies, occasionally (D, I, inset) with one tip that overlaps the other (o). (H, J) The appressed scutellum margin shows lightly pigmented tips and is usually crenulate at maturity (black arrowhead). (K) Hyphae of the appressed marginal of scutella Prillieuxina baccharidincola instead anastomose with surrounding mycelium (white arrowhead). (I, K, L-O) Dehiscence. (I, K, L-M) Asterinales and Asterotexiales ascomata usually dehisce with irregular slits. (K) Mature scutellum with irregular radial slits, black arrowheads. (L-M) Irregular slits follow the longitudinal walls of scutellum cells (white arrowheads). (N-O) Ostiolate openings are less common. (N) Ostiole in young scutellum. (O) Asexual or spermatial stage, with evenly pigmented cells lining ostiole. (P-S) Lateral appressoria showing its characteristic melanized ring at lower focal plane (insets, representing area in the dashed gray boxes). Scale bars: (A-D, H-I) 10 µm; (E-G) 20 µm; (J-K) 50 µm; (L-S) 5 µm. (A, I, J, P, Q) Asterina melastomatis VIC 52822, 1, 4, 6, 2. and 2 focal planes (f.p.), respectively; (B) Asterina chrysophylli VIC 42823, 4 f.p.; (C, O) Asterotexiaceae sp. CBS 143813, 2 f.p. each; (D) Hemigrapha atlantica BR 14014, 1 f.p.; (E, F) Asterotexis cucurbitacearum VIC 42814, 1 f.p. each; (G, L) Rhagadolobiopsis thelypteridis EG156; (H, R) Batistinula gallesiae VIC 42514, 10 and 2 focal planes (f.p.) respectively; (K, N), Prillieuxina baccharidincola VIC 42817; (M) Asterotexiaceae sp. UBC F33036, 11 f.p.; (S) Asterina sp. (VUL. 341b), 2 f.p.

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Premise Fossils show that fly‐speck fungi have been reproducing with small, black thyriothecia on leaf surfaces for ~250 million years. We analyzed morphological characters of extant thyriothecial fungi to develop a phylogenetic framework for interpreting fossil taxa. Methods We placed 59 extant fly‐speck fungi in a phylogeny of 320 Ascomycota usi...

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... the most recent common ancestor of Asterotexiales and Aulographaceae ( Figs. 1 and 2; Appendix S5e). A convergent transition from ostioles to irregular slits appeared along a branch nested in Capnodiales that leads to Asterinales. Developmentally, the irregular slits result from cracking between the adjacent hyphae that make up the scutellum (Fig. 2L, M). The precise location of the opening remains unclear until dehiscence. Slit-forming genera sometimes produce ostioles in their asexual or spermatial states (Fig. 2O). In Aulographaceae, and possibly also in species of Lembosia, the slit appears early in development as a line of lightly pigmented cells (Fig. 3), but the crack follows ...
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... along a branch nested in Capnodiales that leads to Asterinales. Developmentally, the irregular slits result from cracking between the adjacent hyphae that make up the scutellum (Fig. 2L, M). The precise location of the opening remains unclear until dehiscence. Slit-forming genera sometimes produce ostioles in their asexual or spermatial states (Fig. 2O). In Aulographaceae, and possibly also in species of Lembosia, the slit appears early in development as a line of lightly pigmented cells (Fig. 3), but the crack follows longitudinal walls and propagates beyond the less-pigmented area. Regular slits that follow a pre-determined pattern without further propagation are uncommon among ...
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... with surrounding mycelium app = lateral appressoria; no, absent; yes, present ini = initiation of sporocarp, position and number of generator hyphae: int, single, intercalary; ter, single, terminal; mult, multiple, intercalary -= equivocal ancestral state reconstruction additional septa delimit the two arms of the "Y" as separate cells (Figs. 2A, 2B, 2H, and 5B). In some cases, one of the two branches overlaps the other, growing downward and disappearing below its neighbors (Figs. 2D, 2I, and 5C). Septa are sometimes aligned around part of the circumference of the scutellum, resulting in partial concentric rings of septa (cf. fig. 3.8 "d" of Hofmann, 2009). Pigmentation in older parts ...
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... generator hyphae: int, single, intercalary; ter, single, terminal; mult, multiple, intercalary -= equivocal ancestral state reconstruction additional septa delimit the two arms of the "Y" as separate cells (Figs. 2A, 2B, 2H, and 5B). In some cases, one of the two branches overlaps the other, growing downward and disappearing below its neighbors (Figs. 2D, 2I, and 5C). Septa are sometimes aligned around part of the circumference of the scutellum, resulting in partial concentric rings of septa (cf. fig. 3.8 "d" of Hofmann, 2009). Pigmentation in older parts of the scutellum contrasts with more translucent tips at the margin of scutellum hyphae (Fig. 2H, J, and K). Scutella with ...
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... downward and disappearing below its neighbors (Figs. 2D, 2I, and 5C). Septa are sometimes aligned around part of the circumference of the scutellum, resulting in partial concentric rings of septa (cf. fig. 3.8 "d" of Hofmann, 2009). Pigmentation in older parts of the scutellum contrasts with more translucent tips at the margin of scutellum hyphae (Fig. 2H, J, and K). Scutella with pseudomonopodial branching are restricted to Aulographaceae. The common ancestor of Aulographaceae is reconstructed with pseudomonopodial branching with 99% proportional likelihood and 28% posterior probability (Table 3; Appendices S3, S4, and S5h). In this growth form, the tips of dominant hyphae grow at the ...
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... of sporocarps lack a direct homolog to the scutellum margin because their outer walls curve up from the substrate. In Asterotexiales, Asterinales, Aulographaceae, in species of Stomiopeltis, and in Muyocopronales, the scutellum margins often look crenulate or finely scalloped because the dome-shaped hyphal tips bulge out around the circumference (Figs. 2H, 2J, 3C, and 3D; Appendix S5i). In some Stomiopeltis-like species (Fig. 3A), tips of hyphae are free, which we interpret as a variant of a crenulate margin (Table 2). In other species, for example in Prillieuxina baccharidincola (Fig. 2K), the scutellum hyphae anastomose with other superficial hyphae on the leaf surface, so that the margin of ...
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... look crenulate or finely scalloped because the dome-shaped hyphal tips bulge out around the circumference (Figs. 2H, 2J, 3C, and 3D; Appendix S5i). In some Stomiopeltis-like species (Fig. 3A), tips of hyphae are free, which we interpret as a variant of a crenulate margin (Table 2). In other species, for example in Prillieuxina baccharidincola (Fig. 2K), the scutellum hyphae anastomose with other superficial hyphae on the leaf surface, so that the margin of the scutellum is poorly defined and continuous with the surface mycelium. In Microthyriales and Zeloasperisporiales, scutellum margins are often entire, the scutellum hyphae ending in truncate tips aligned around the circumference ...
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... Fringe hyphae appear to be more tightly appressed to the host surface than central portions of scutella. Lateral appressoria-Appressoria are widely distributed across thyriothecium-forming Dothideomycetes and vary in their morphology and position. Conspicuous, pigmented, lateral appressoria are only produced by Asterinales and Asterotexiales ( Fig. 2P-S; Appendices S1d and S5j). The appressoria are usually unicellular ( Fig. 2P and R), but in some species they may be two-celled ( Fig. 2Q and S; cf. figs. 3.8, 3.14, 3.21, 3.24, 3.31, 3.56 of Hofmann, 2009). Lateral appressoria may be swollen, lobed or minimally differentiated, and they form as unicellular lateral outgrowths from ...
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... central portions of scutella. Lateral appressoria-Appressoria are widely distributed across thyriothecium-forming Dothideomycetes and vary in their morphology and position. Conspicuous, pigmented, lateral appressoria are only produced by Asterinales and Asterotexiales ( Fig. 2P-S; Appendices S1d and S5j). The appressoria are usually unicellular ( Fig. 2P and R), but in some species they may be two-celled ( Fig. 2Q and S; cf. figs. 3.8, 3.14, 3.21, 3.24, 3.31, 3.56 of Hofmann, 2009). Lateral appressoria may be swollen, lobed or minimally differentiated, and they form as unicellular lateral outgrowths from surface hyphae. We coded thyriothecial taxa of Asterinales and Asterotexiales that lack ...
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... published illustrations (Appendix S1d). Otherwise, data on development is scarce, and ancestral state reconstructions are equivocal where data are missing (Appendices S1d, S4 and S5k). In Asterinales and Asterotexiales a single "generator hypha" (Hofmann, 2009) initiates the thyriothecium by forming multiple, closely spaced intercalary septa ( Figs. 2A-2C, 2H-2J, and 5C). Each of the delimited cells then gives rise to transverse, adjacent, dichotomizing hyphal tips that together form the scutellum. The generator hypha remains above the dorsal surface of the mature scutellum and is visible even in mature thyriothecia (Figs. 2I, 2J, and 5C). Ascomata in one species of Asterotexiales, ...
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... thyriothecium by forming multiple, closely spaced intercalary septa ( Figs. 2A-2C, 2H-2J, and 5C). Each of the delimited cells then gives rise to transverse, adjacent, dichotomizing hyphal tips that together form the scutellum. The generator hypha remains above the dorsal surface of the mature scutellum and is visible even in mature thyriothecia (Figs. 2I, 2J, and 5C). Ascomata in one species of Asterotexiales, Rhagadolobiopsis thelypteridis, are initiated directly over host stomata, as shown by Guatimosim et al. (2014) and in Fig. 2G here; in Asterotexis cucurbitacearum, they are initiated directly from ascospores ( Fig. 2E and F) (shown previously by Guerrero et al., 2011Guerrero et al., ...
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... tips that together form the scutellum. The generator hypha remains above the dorsal surface of the mature scutellum and is visible even in mature thyriothecia (Figs. 2I, 2J, and 5C). Ascomata in one species of Asterotexiales, Rhagadolobiopsis thelypteridis, are initiated directly over host stomata, as shown by Guatimosim et al. (2014) and in Fig. 2G here; in Asterotexis cucurbitacearum, they are initiated directly from ascospores ( Fig. 2E and F) (shown previously by Guerrero et al., 2011Guerrero et al., , ...
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... of the mature scutellum and is visible even in mature thyriothecia (Figs. 2I, 2J, and 5C). Ascomata in one species of Asterotexiales, Rhagadolobiopsis thelypteridis, are initiated directly over host stomata, as shown by Guatimosim et al. (2014) and in Fig. 2G here; in Asterotexis cucurbitacearum, they are initiated directly from ascospores ( Fig. 2E and F) (shown previously by Guerrero et al., 2011Guerrero et al., , ...

Citations

... h Brown and verruculose ascospore. Scale bars: a = 200 µm, b-c = 50 µm, d-e-f = 20 µm, d = 10 µm, g-h = 10 µm families, namely: Asterinaceae and Parmulariaceae (Guatimosim et al. 2015;Giraldo et al. 2017;Phookamsak et al. 2019;Johnston and Park 2019;Hongsanan et al. 2020a;Le Renard et al. 2020;Firmino and Pereira 2021). ...
Article
The description of a new Mediterranean species, Coltricia insularis, is provided, on the basis of material collected in Corsica, Sardinia, Cyprus and Spain
... h Brown and verruculose ascospore. Scale bars: a = 200 µm, b-c = 50 µm, d-e-f = 20 µm, d = 10 µm, g-h = 10 µm families, namely: Asterinaceae and Parmulariaceae (Guatimosim et al. 2015;Giraldo et al. 2017;Phookamsak et al. 2019;Johnston and Park 2019;Hongsanan et al. 2020a;Le Renard et al. 2020;Firmino and Pereira 2021). ...
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This article is the 14th in the Fungal Diversity Notes series, wherein we report 98 taxa distributed in two phyla, seven classes, 26 orders and 50 families which are described and illustrated. Taxa in this study were collected from Australia, Brazil, Burkina Faso, Chile, China, Cyprus, Egypt, France, French Guiana, India, Indonesia, Italy, Laos, Mexico, Russia, Sri Lanka, Thailand, and Vietnam. There are 59 new taxa, 39 new hosts and new geographical distributions with one new combination. The 59 new species comprise Angustimassarina kunmingense, Asterina lopi, Asterina brigadeirensis, Bartalinia bidenticola, Bartalinia caryotae, Buellia pruinocalcarea, Coltricia insularis, Colletotrichum flexuosum, Colletotrichum thasutense, Coniochaeta caraganae, Coniothyrium yuccicola, Dematipyriforma aquatic, Dematipyriforma globispora, Dematipyriforma nilotica, Distoseptispora bambusicola, Fulvifomes jawadhuvensis, Fulvifomes malaiyanurensis, Fulvifomes thiruvannamalaiensis, Fusarium purpurea, Gerronema atrovirens, Gerronema flavum, Gerronema keralense, Gerronema kuruvense, Grammothele taiwanensis, Hongkongmyces changchunensis, Hypoxylon inaequale, Kirschsteiniothelia acutisporum, Kirschsteiniothelia crustaceum, Kirschsteiniothelia extensum, Kirschsteiniothelia septemseptatum, Kirschsteiniothelia spatiosum, Lecanora immersocalcarea, Lepiota subthailandica, Lindgomyces guizhouensis, Marthe asmius pallidoaurantiacus, Marasmius tangerinus, Neovaginatispora mangiferae, Pararamichloridium aquisubtropicum, Pestalotiopsis piraubensis, Phacidium chinaum, Phaeoisaria goiasensis, Phaeoseptum thailandicum, Pleurothecium aquisubtropicum, Pseudocercospora vernoniae, Pyrenophora verruculosa, Rhachomyces cruralis, Rhachomyces hyperommae, Rhachomyces magrinii, Rhachomyces platyprosophi, Rhizomarasmius cunninghamietorum, Skeletocutis cangshanensis, Skeletocutis subchrysella, Sporisorium anadelphiae-leptocomae, Tetraploa dashaoensis, Tomentella exiguelata, Tomentella fuscoaraneosa, Tricholomopsis lechatii, Vaginatispora flavispora and Wetmoreana blastidiocalcarea. The new combination is Torula sundara. The 39 new records on hosts and geographical distribution comprise Apiospora guiyangensis, Aplosporella artocarpi, Ascochyta medicaginicola, Astrocystis bambusicola, Athelia rolfsii, Bambusicola bambusae, Bipolaris luttrellii, Botryosphaeria dothidea, Chlorophyllum squamulosum, Colletotrichum aeschynomenes, Colletotrichum pandanicola, Coprinopsis cinerea, Corylicola italica, Curvularia alcornii, Curvularia senegalensis, Diaporthe foeniculina, Diaporthe longicolla, Diaporthe phaseolorum, Diatrypella quercina, Fusarium brachygibbosum, Helicoma aquaticum, Lepiota metulispora, Lepiota pongduadensis, Lepiota subvenenata, Melanconiella meridionalis, Monotosporella erecta, Nodulosphaeria digitalis, Palmiascoma gregariascomum, Periconia byssoides, Periconia cortaderiae, Pleopunctum ellipsoideum, Psilocybe keralensis, Scedosporium apiospermum, Scedosporium dehoogii, Scedosporium marina, Spegazzinia deightonii, Torula fici, Wiesneriomyces laurinus and Xylaria venosula. All these taxa are supported by morphological and multigene phylogenetic analyses. This article allows the researchers to publish fungal collections which are important for future studies. An updated, accurate and timely report of fungus-host and fungus-geography is important. We also provide an updated list of fungal taxa published in the previous fungal diversity notes. In this list, erroneous taxa and synonyms are marked and corrected accordingly.
... Thus, it is difficult to establish species diversity in the order and whether the genera comprise numerous new taxa or more names than needed . Most taxa of Asterinales are biotrophic that cannot be cultured; thus, DNA extraction of these species must be from fresh fruiting structures (Renard et al. 2020). Therefore, the majority of past studies did not carry out DNA-based analyses. ...
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Asterinales is an important epifoliar order which generally lacks of DNA-based sequence data. There are many genera in Asterinales lacking molecular data and the exact taxonomic placement of those genera is undetermined. In this study, we introduce Brunneofissuraceae fam. nov. and Cirsosia mangiferae sp. nov. and Asterina neomangiferae nom. nov. based on morpho-molecular evidences. All fungal specimens were collected during September (2020) from Chiang Mai, Thailand. The phylogenetic analysis based on 28s (LSU) and 5.8s (ITS) sequence data confirmed the placements of Asterolibertia, Brunneofissura, and Cirsosia in Asterinales. We provide the first molecular data for Asterolibertia (current name is Asterina) and Cirsosia. Comparative morphologies and phylogenetic analyses are provided for each taxon using illustrations and well-supported phylogenetic analyses.
... Some species of rust fungi were cultivated in axenic culture, but only in a very specific medium containing several nutrients (Scott and Maclean, 1969;Wolf, 1974;Cummins and Hiratsuka, 2003). Groups of fungi once considered biotrophic, such as Microthyriaceae and Chaetothyriaceae, have been recently isolated in a systematic way (Chomnunti et al., 2012;Wu et al., 2011;Le Renard et al., 2020). Powdery mildew, mildew, and meliolaceous fungi (Ascomycota, Meliolales) remain biotrophic and without successful cultivation (Göker et al., 2007;Rodriguez and Piepenbring, 2007;Beakes et al., 2012;Pinho et al., 2012aPinho et al., , 2012bPinho et al., , 2013Braun and Cook, 2012;Soares and Dianese, 2013). ...
... They are biotrophs, being parasites on living leaves, stems and fruits, but, do not cause severe damage to the plants, although they penetrate host cells using the superficial appressoria, and can absorb nutrient through the haustoria (von Arx and Müller, 1975;Hofmann, 2010;Hosagoudar, 2012;Hongsanan et al., 2016). Currently, Asterinales stricto sensu is composed of the Asterinaceae and Parmulariaceae families (Giraldo et al., 2017;Phookamsak et al., 2019;Johnston and Park, 2019;Le Renard et al., 2020). Since the first descriptions of Asterinales by Léveillé in 1845 these fungi are treated as being biotrophic (Bezerra, 2004;Hofmann and Piepenbring, 2008;Hofmann et al., 2010;Hofmann and Piepenbring, 2011;Hosagoudar, 2012;Hongsanan et al., 2014;Guatimosim et al., 2015). ...
Article
Although asterinaceous fungi have been studied for many years, all previous attempts to isolate, cultivate, and propagate these fungi in vitro have failed. This paper provides the first reports of in vitro isolation of representative strains of species belonging to four fungi from different genera belonging to Asterinales. To confirm if the sequences of DNA obtained from the mycelia are the same obtained in the direct extraction, a phylogenetic analysis of nuc LSU rDNA was performed. This paper reports for the first time the success of in vitro culturing of asterinaceous fungi using the ascospores ejection technique, opening perspectives of studies of genetics, physiology, among other aspects of the biology for this very understudied group of fungi.
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Yunnan, located in southwestern China, is known for its high fungal diversity, and many of which are endemic to the region. As part of our ongoing studies on fungi in Yunnan, we introduce two new genera in Phaeothecoidiellaceae (Mycosphaerellales), to accommodate one Repetophragma-like and another Stomiopeltis-like taxa. Pseudorepetophragma gen. nov. is introduced herein as a monotypic genus to accommodate P. zygopetali comb. nov.(≡ Repetophragma zygopetali), whereas Pseudostomiopeltis gen. nov. is introduced to accommodate Ps. xishuangbannaensis gen. et sp. nov. and Ps. phyllanthi comb. nov.(≡ Stomiopeltis phyllanthi), based on a new collection from Yunnan. In addition, Stomiopeltis sinensis is transferred to Exopassalora as E. sinensis comb. nov. due to its phylogenetic affinity and grouped with E. zambiae, the generic type of Exopassalora. This study provides new insights into the biodiversity of fungal species in this region and adds to our understanding of their ecological roles, as well as the resolution to ambiguous taxa in Phaeothecoidiellaceae.
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Epifoliar fungi are one of the significant fungal groups typically living on the surface of leaves. They are usually recorded as saprobes, obligate parasites and commensals and are widely distributed in tropical and subtropical regions. Numerous genera within this group remain inadequately understood, primarily attributed to limited taxonomic knowledge and insufficient molecular data. Furthermore, the taxonomic delineation of epifoliar fungi remained uncertain, with scattered and literature-based data often intermixed with other follicolous fungi. Herein, a comprehensive taxonomic monograph of 124 genera in (32) Asterinales, (18) Capnodiales, (15) Chaetothyriales, (8) Meliolales, (8) Micropeltidales, (10) Microthyriales, (32) Parmulariales and (1) Zeloasperisporiales was provided re-describing with illustrations and line drawings. Notes on ecological and economic importance of the families are also provided. Representatives type herbarium materials of Campoa pulcherrima, Cycloschizon brachylaenae, Ferrarisia philippina, Hysterostomella guaranitica, Palawaniella orbiculata and Pseudolembosia orbicularis of Parmulariaceae were re-examined and provided updated illustrations with descriptions. A backbone phylogenetic tree and divergence estimation analysis for epifoliar fungi based on LSU and 5.8s ITS sequence data are provided.
Chapter
Mit ca. 50.400 Arten in ungefähr 2300 Gattungen sind die Basidiomyceten (wissenschaftlich Basidiomycota) nach den Ascomycota die zweitgrößte Abteilung der Fungi (◘ Abb. 2.1). Zu dieser Gruppe gehören die meisten Großpilze, darunter viele Speise- und Giftpilze. Das Schlüsselmerkmal der Arten der Basidiomycota ist die Basidie, eine Zelle, die als kleiner Ständer (lat. basidium = kleine Basis mit Füßen) meist jeweils vier Basidiosporen trägt.
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Premise of research. Mesozoic to Cenozoic fungi forming scutella, shield-like coverings of sporulating tissue on leaf surfaces, offer insights into the diversification of filamentous ascomycetes. We found a new fungal sporocarp type consisting of radiate scutella growing from the stomata of leaves of a conifer in a survey of dispersed plant cuticles from the Lower Cretaceous of Virginia. Here, we interpret the new morphotype and compare it with other fossil groups and with extant taxa. Methodology. We isolated fragments of conifer cuticles colonized by the fungus from macerated clays of the Potomac Group, lower Zone 1 (Aptian, 125–113 Ma), and studied them using light microscopy. We analyzed the anatomy of scutella of living taxa and compared the fossils with extant taxa producing either rhizothyria, asexual sporocarps that release spores from below the outer edge of the scutellum, or thyriothecia, sporocarps that release spores through dorsal cracks or an ostiole. Pivotal results. Stomatothyrium placocentrum gen. et sp. nov. has a radial arrangement of scutellum hyphae, characteristic of extant and fossil thyriothecial species in Dothideomycetes (Ascomycota). The scutella develop cracks on the dorsal surface that may showthe dehiscence mechanism. Unusually for dehiscence in Dothideomycetes, the cracks follow the scutellum circumference rather than its radius. Each scutellum arises from a host plant stoma, appearing first as a hyphal columella that broadens into a flat scutellum disk on the leaf surface. The morphotype is common, and we found more than 63 specimens at different developmental stages on cuticle fragments. Conclusions. Its unique morphology suggests that S. placocentrum gen. et sp. nov. represents a now extinct group of leaf-dwelling Dothideomycetes that formed thyriothecia. This discovery contributes to our understanding of the range of character combinations of early epiphyllous Ascomycota. © 2021 The University of Chicago. All rights reserved. Published by The University of Chicago Press.
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Premise: Fossils can reveal long-vanished characters that inform inferences about the timing and patterns of diversification of living fungi. Through analyzing well-preserved fossil scutella, shield-like covers of fungal sporocarps, we describe a new taxon of early Dothideomycetes with a combination of characters unknown among extant taxa. Methods: Macerated clays from the Potomac Group, lower Zone 1, from the Lower Cretaceous (Aptian, 125-113 Ma) of Virginia USA yielded one gymnospermous leaf cuticle colonized by 21 sporocarps of a single fungal morphotype. We inferred a tree from nuclear ribosomal DNA of extant species, and coded morphological characters to evaluate alternative, equally parsimonious placements of the fossil in a molecular constraint tree of extant species. Results: Bleximothyrium ostiolatum gen. et sp. nov. has an ostiolate scutellum of radiate, dichotomizing hyphae. Unlike otherwise similar extant and fossil taxa, B. ostiolatum has tangled hyphae at its scutellum margin. Scutella of B. ostiolatum are connected to superficial mycelium, to intercalary and lateral appressoria, and to extensive subcuticular "mycélium en palmettes". The gymnospermous host has characters consistent with identity as a non-papillate ginkgophyte or cycad. Conclusions: Bleximothyrium ostiolatum is the oldest known fossil fly-speck fungus that occurs on plant cuticles and has the radiate, ostiolate scutellum known only from Dothideomycetes. Its combination of characters, its scutellum margin, and mycélium en palmettes are unknown in other extant and fossil species, and Bleximothyrium ostiolatum likely represents a new group of fly-speck fungi that may now be extinct.