Ascomycota

Thelocarpon epibolum growing on Baeomyces rufus, from here.

Belongs within: Opisthokonta.
Contains: Taphrinales, Saccharomycetales, Parmulariaceae, Parodiopsidaceae, Schizothyriaceae, Pezizales, Orbiliaceae, Geoglossaceae, Sordariomyceta, Lichinales, Eurotiomycetes, Lecanoromycetes, Xylonales, Dothideomyceta.

The Ascomycota is a diverse group of often multicellular fungi in which spores are produced within saccate asci.

Characters (from Adl et al. 2019): Sexual reproduction within asci (saccate structures); meiosis usually followed by mitosis to produce from one to over 1,000 ascospores, but usually eight; ascospore walls form inside ascus; mating types heterothallic, homothallic (selfing) or both; may reproduce sexually (teleomorph) or asexually (anamorph) only, or both sexually and asexually (holomorph); asci cylindrical, fusiform, clavate or globose, persistent or evanescent, with or without a fruiting structure (ascoma, -ata); asci developing directly from ascogenous hyphae, from a crozier or from a single cell; asexual reproduction by conidiospores (mitospores) formed by fragmentation of vegetative hyphae (thallic), blastically from single cells, hyphae or conidiophores; vegetative body of single cells or tubular, septate filaments (hyphae); septa with simple pores, except for those associated with ascogenous hyphae and asci; cell walls lamellate with a thin electron-dense outer layer and a relatively thick electrontransparent inner layer, consisting of varying proportions of chitin and glucans.

<==Ascomycota [Ascodiomycetes, Ascomycetes, Ascomycotina, Deuteromycotina, Hemiascomycotina, Hyphomycetes]
    |--Taphrinomycota [Archiascomycetes, Protomycetes, Taphrinomycotina] SS09
    |    |--Neolecta Spegazzini 1881 AS12 (see below for synonymy)
    |    |    |--*N. flavovirescens LS03
    |    |    |--N. irregularis [=Geoglossum irregulare] LS03
    |    |    `--N. vitellina [=Geoglossum vitellinum] LS03
    |    `--+--+--Taphrinales SS09
    |       |  `--Saitoella complicata SS09
    |       `--+--Archaeorhizomyces Rosling & James in Rosling, Cox et al. 2011 (see below for synonymy) RC11
    |          |    `--*A. finlayi Rosling & James in Rosling, Cox et al. 2011 RC11
    |          `--+--Pneumocystis Delanoë & Delanoë 1912 RC11, AS12 (see below for synonymy)
    |             |    |--P. carinii Frenkel 1999 KFS03
    |             |    `--P. jirovecii Frenkel 1999 KFS03
    |             `--Schizosaccharomyces Lindner 1893 RC11, AS12 (see below for synonymy)
    |                  |--S. japonicus EB03N
    |                  |    |--S. j. var. japonicus YK99
    |                  |    `--S. j. var. versatilis YK99
    |                  |--S. octosporus SL02
    |                  `--S. pombe EB03N
    `--Saccharomyceta SS09
         |--Saccharomycetales SS09
         `--Pezizomycotina (see below for synonymy) SS09
              |  i. s.: Sesquicillium SL02
              |         Coniocybaceae EB03O
              |           |--Coniocybe EB03O
              |           |--Chaenotheca EB03O
              |           `--Sclerophora EB03O
              |         Malcaria flagellata (Hansford) Hansford 1946 [=Paranectria flagellata Hansford 1941] DP72
              |         Argynnaceae EB03O
              |           |--Argynna EB03O
              |           `--Lepidopterella EB03O
              |         Arthopyreniaceae EB03O
              |           |--Athrisidium EB03O
              |           `--Mycoicrothelia EB03O
              |         Aulographaceae EB03O
              |           |--Aulographum EB03O
              |           `--Polyclypeolina EB03O
              |         Cookellaceae EB03O
              |           |--Pycnoderma EB03O
              |           `--Uleomyces EB03O
              |         Corynesporasca [Corynesporascaceae] EB03O
              |           `--C. cassiicola BSA97
              |         Englerulaceae EB03O
              |           |--Englerula macarangae EB03O, DP72
              |           |--Parenglerula EB03O
              |           |--Rhizotexis EB03O
              |           |--Rhytidenglerula homalanthi EB03O, DP72 [=Englerulella homalanthi DP72]
              |           |--Schiffnerula solani EB03O, DP72
              |           `--Thrauste EB03O
              |         Eremomycetaceae EB03O
              |           |--Eremomyces EB03O
              |           `--Rhexothecium EB03O
              |         Euantennariaceae EB03O
              |           |--Euantennaria EB03O
              |           |--Rasutoria EB03O
              |           |--Strigopodia EB03O
              |           `--Trichopeltheca EB03O
              |         Hypsostromataceae EB03O
              |           |--Hypsostroma EB03O
              |           `--Manglicola EB03O
              |         Leptopeltidaceae EB03O
              |           |--Dothiopeltis EB03O
              |           |--Leptopeltis EB03O
              |           |--Nannfeldtia EB03O
              |           |--Phacidina EB03O
              |           |--Ronnigeria EB03O
              |           `--Staibia EB03O
              |         Lichenotheliaceae EB03O
              |           |--Lichenostigma EB03O
              |           `--Lichenothelia EB03O
              |         Meliolina [Meliolinaceae] ST99
              |           |--M. cladotricha [incl. M. arborescens, M. octospora] DP72
              |           `--M. sydowiana ST99
              |         Mesnieraceae EB03O
              |           |--Bondiella EB03O
              |           |--Helochora EB03O
              |           |--Mesniera EB03O
              |           `--Stegasphaeria EB03O
              |         Microtheliopsis EB03O (see below for synonymy)
              |           `--*M. uleana (see below for synonymy) E99
              |         Moriola [Moriolaceae] EB03O
              |         Mycoporum [Mycoporaceae] EB03O
              |         Naetrocymbaceae EB03O
              |           |--Jarxia EB03O
              |           |--Leptorhaphis EB03O
              |           |--Naetrocymbe EB03O
              |           `--Tomasellia EB03O
              |         Parmulariaceae EB03O
              |         Parodiella [Parodiellaceae] EB03O
              |         Parodiopsidaceae EB03O
              |         Polystomellaceae EB03O
              |           |--Dothidella EB03O
              |           |--Munkiella EB03O
              |           `--Parastigmatea EB03O
              |         Protoscypha [Protoscyphaceae] EB03O
              |         Pyrenotrichaceae EB03O
              |           |--Cyanoporina EB03O
              |           `--Pyrenothrix EB03O
              |         Schizothyriaceae EB03O
              |         Vizellaceae EB03O
              |           |--Blasdalea EB03O
              |           `--Vizella EB03O
              |         Hormiscium myrmecophilum Thaxter 1914 ES12
              |         Xanthopyreniaceae [Collemopsidiales, Collemopsidiomycetes] AB19
              |           |--Collemopsidium EB03O
              |           |--Didymellopsis EB03O
              |           |--Pyrenocollema EB03O
              |           `--Zwackhiomyces EB03O
              |--Pezizales SS09
              `--+--Orbiliaceae SS09
                 `--Leotiomyceta SS09
                      |--Geoglossaceae SS09
                      `--+--Sordariomyceta SS09
                         `--+--+--Lichinales SS09
                            |  `--+--+--Sarcosagium campestre LK04
                            |     |  `--Biatoridium monasteriense LK04
                            |     `--Thelocarpaceae [Thelocarpeae] EB03O
                            |          |--Melanophloea EB03O
                            |          |--Sarcosagium EB03O
                            |          `--Thelocarpon Nylander 1853 LK04, RS99 (see below for synonymy)
                            |               |--*T. albidum Nylander 1853 RS99
                            |               |--T. citrum (Wallroth) Rossman in Rossman, Samuels et al. 1999 (see below for synonymy) RS99
                            |               |--T. epibolum A96
                            |               |--T. laureri LK04
                            |               |--T. lichenicola (Fuckel) Poelt & Hafellner 1975 (see below for synonymy) RS99
                            |               `--‘*Cyanocephalium’ murorum Zukal 1893 RS99
                            `--+--+--Eurotiomycetes JK06
                               |  `--Lecanoromycetes SS09
                               `--+--+--Xylonales SS09
                                  |  `--Dothideomyceta SS09
                                  `--Candelariaceae [Candelariales] SS09
                                       |--Candelina EB03O
                                       |--Clarouxia EB03O
                                       |--Placomaronea EB03O
                                       `--Candelariella EB03O
                                            |--C. antennaria SS09
                                            |--C. reflexa SS09
                                            |--C. terrigena SS09
                                            `--C. vitellina C74

Ascomycota incertae sedis:
  Lahmia [Lahmiaceae, Lahmiales] EB03O
  Medeolaria [Medeolariaceae, Medeolariales] EB03O
  Triblidiaceae [Triblidiales] EB03O
    |--Huangshania EB03O
    `--Pseudographis EB03O
  Amorphotheca [Amorphothecaceae] EB03O
  Aphanopsis [Aphanopsidaceae] EB03O
  Diporotheca Gordon & Shaw 1960 [Diporothecaceae EB03O]
  Eoterfeziaceae EB03O
    |--Acanthogymnomyces EB03O
    `--Eoterfezia EB03O
  Epigloea [Epigloeaceae] EB03O
  Mastodiaceae EB03O
    |--Mastodia EB03O
    `--Turgidosculum EB03O
  Microcalicium [Microcaliciaceae] EB03O
  Phyllobatheliaceae EB03O
    |--Phyllocratera EB03O
    |--Opercularia firma E99
    `--Phyllobathelium [incl. Septoriomyces] E99
         `--P. leguminosae (Cavalc. & Silva) Lücking & Sérus 1998 [=*Septoriomyces leguminosae] E99
  Daruvedia [Pleurotremataceae] EB03O
  Protothelenellaceae EB03O
    |--Mycowinteria EB03O
    `--Prothelenella EB03O
  Saccardiaceae EB03O
    |--Angatia EB03O
    |--Ascolectus EB03O
    |--Cyanodiscus EB03O
    |--Epibelonium EB03O
    |--Johansonia EB03O
    |--Phillipsiella EB03O
    |--Rivilata EB03O
    |--Saccardia EB03O
    `--Vonarxella EB03O
  Seuratiaceae EB03O
    |--Seuratia EB03O
    `--Seuratiopsis EB03O
  Thrombium [Thrombiaceae] EB03O
  Calycidium [Calycidiaceae] EB03O
  Allophoron EB03O
  Antimanoa EB03O
  Argentinomyces EB03O
  Arthopyreniomyces EB03O
  Ascocorticiellum EB03O
  Ascomauritania EB03O
  Ascosorus EB03O
  Ascosubramania EB03O
  Ascoxyta EB03O
  Atractobolus EB03O
  Baculospora EB03O
  Batistospora EB03O
  Berggrenia EB03O
  Biflua EB03O
  Bresadolina EB03O
  Carnia EB03O
  Clathroporinopsis EB03O
  Coryneliella EB03O
  Coscinocladium EB03O
  Crinigera EB03O
  Cyanopyrenia EB03O
  Cystodium EB03O
  Diaboliumbilicus EB03O
  Diehliomyces EB03O
  Dipyrgis EB03O
  Discocera EB03O
  Dryinosphaera EB03O
  Eiona EB03O
  Elaeomyces EB03O
  Endocolium EB03O
  Enduria EB03O
  Erispora Patouillard 1922 (n. d.) EB03O, RS99
    `--*E. parasitica Patouillard 1922 (n. d.) RS99
  Farriolla EB03O
  Feracia Rolland 1905 (n. d.) EB03O, RS99
    `--*F. balearica Rolland 1905 (n. d.) RS99
  Frigidospora EB03O
  Gaeumanniella EB03O
  Gonidiomyces EB03O
  Gyrophthorus EB03O
  Haematomyxa EB03O
  Haplopyrenulomyces EB03O
  Hapsidascus EB03O
  Harmandiana EB03O
  Heterocyphelium EB03O
  Heuflera EB03O
  Hyalodermella EB03O
  Hyalopyrenula EB03O
  Hypnotheca EB03O
  Leucoconiella EB03O
  Leucoconis EB03O
  Lichenopeziza EB03O
  Lithopythium EB03O
  Lohwagiella EB03O
  Ludwigomyces EB03O
  Lyromma EB03O [incl. Anconomyces E99, Lyrommotheca (nom. illeg.) E99]
    |--*L. nectandrae E99
    |--L. dolicobelum E99
    `--L. palmae E99
  Marisolaris EB03O
  Micromastia EB03O
  Molgosphaera EB03O
  Mycotodea EB03O
  Myriococcum EB03O
  Nemacola EB03O
  Normandina EB03O
  Ochrosphaera EB03O
  Phacidiostromella EB03O
  Phaeodothiopsis EB03O
  Phellostroma EB03O
  Phelonitis EB03O
  Phialisphaera EB03O
  Phragmitensis EB03O
  Phthora EB03O
  Porinula EB03O
  Potamomyces EB03O
  Protocalicium EB03O
  Pseudohepatica EB03O
  Pseudoperitheca EB03O
  Psilosphaeria EB03O
  Pteromycula EB03O
  Pycnodermellina EB03O
  Roeslerina EB03O
  Rostafinskia EB03O
  Sachsia EB03O
  Schistophoron EB03O
  Stellifraga EB03O
  Stigmatisphaera EB03O
  Stigmea EB03O
  Syphosphaera EB03O
  Telioclipeum EB03O
  Thallisphaera EB03O
  Trichoplacia EB03O
  Trichosphaera EB03O
  Tylophorella EB03O
  Tylophoron EB03O
  Wadeana EB03O
  Wolkia EB03O
  Xenomyxa EB03O
  Xylobotryum EB03O
  Xylogone EB03O
  *Aciesia xylopiae (n. d.) E99
  Arthrobotryomyces amazonensis (n. d.) E99
  Asbolisiomyces ingae (n. d.) E99
  Astrabomyces amazonensis (n. d.) E99
  Podoxyphiomyces manaosensis (n. d.) E99
  *Pyriomyces protii E99
  Fellhanera E99
  *Pyripnomyces maranhensis E99
  Phyllophiale [incl. Stephosia] E99
    |--P. alba Santesson 1952 LV98
    `--P. fusca LV98
  Puiggarina costaricensis RS99
  Zeus olympius RS99
  Sphaceloma ampelinum [incl. Gloeosporium ampelophagum, Fusarium cesatii, Chrysogluten cesatii] RS99
  Porphyrosoma Patouillard 1928 (n. d.) RS99
    `--*P. episphaeria Patouillard 1928 (n. d.) [=Hypocreopsis episphaeria] RS99
  Lecythium Zukal 1893 non Hertwig & Lesser 1874 (ICZN) [=Lecithium (l. c.)] RS99
    `--*L. aerugineum Zukal 1893 RS99
  Eleutheromyces subulatus (see below for synonymy) RS99
  Kasachstania africana C-S18
  Ramulina yemensis B15
  Clypeoseptoria aparothospermi MS98
  Cryptosporium lunasporum MS98
  Helicodendron giganteum T93

Archaeorhizomyces Rosling & James in Rosling, Cox et al. 2011 [Archaeorhizomycetales, Archaeorhizomycetes] RC11

Eleutheromyces subulatus [=Sphaeronaema subulata, Sphaeronaemella subulata; incl. Sphaeronaema oxysporum, Sphaeronaemella oxyspora] RS99

Microtheliopsis EB03O [incl. Byrsomyces E99, Chaetomodorus E99, Scutomyces E99; Microtheliopsidaceae]

*Microtheliopsis uleana [incl. *Chaetomonodorus brosimi, *Scutomyces concentricus, *Byrsomyces olivaceus] E99

Neolecta Spegazzini 1881 AS12 [incl. Ascocorynium LS03, Spragueola LS03; Neolectaceae, Neolectales, Neolectomycetes SS09]

Pezizomycotina [Ascohymeniales, Calycomycetidae, Discomycetes, Euascomycetae, Euascomycetes, Euascomycotina, Hymenoascomycetes, Loculoascomycetae, Loculoascomycetes, Loculoascomycetidae, Loculomycetes, Meliolineae, Pezomycetidae] SS09

Pneumocystis Delanoë & Delanoë 1912 RC11, AS12 [Pneumocystidaceae, Pneumocystidales, Pneumocystidomycetes]

Schizosaccharomyces Lindner 1893 RC11, AS12 [Schizosaccharomycetaceae, Schizosaccharomycetales, Schizosaccharomycetes]

Thelocarpon Nylander 1853 LK04, RS99 [incl. Ahlesia Fuckel 1870 RS99, Cyanocephalium Zukal 1893 RS99, Metanectria Saccardo 1878 RS99]

Thelocarpon citrum (Wallroth) Rossman in Rossman, Samuels et al. 1999 [=Sphaeria citrum Wallroth 1833, *Metanectria citrum (Wallroth) Saccardo 1878, Nectria citrum (Wallroth) Montagne 1858; incl. Thelocarpon vicinellum Nylander 1885] RS99

Thelocarpon lichenicola (Fuckel) Poelt & Hafellner 1975 [=Peziza lichenicola Fuckel 1864, *Ahlesia lichenicola (Fuckel) Fuckel 1870] RS99

*Type species of generic name indicated

REFERENCES

[AB19] Adl, S. M., D. Bass, C. E. Lane, J. Lukeš, C. L. Schoch, A. Smirnov, S. Agatha, C. Berney, M. W. Brown, F. Burki, P. Cárdenas, I. Čepička, L. Chistyakova, J. del Campo, M. Dunthorn, B. Edvardsen, Y. Eglit, L. Guillou, V. Hampl, A. A. Heiss, M. Hoppenrath, T. Y. James, A. Karnkowska, S. Karpov, E. Kim, M. Kolisko, A. Kudryavtsev, D. J. G. Lahr, E. Lara, L. Le Gall, D. H. Lynn, D. G. Mann, R. Massana, E. A. D. Mitchell, C. Morrow, J. S. Park, J. W. Pawlowski, M. J. Powell, D. J. Richter, S. Rueckert, L. Shadwick, S. Shimano, F. W. Spiegel, G. Torruella, N. Youssef, V. Zlatogursky & Q. Zhang. 2019. Revisions to the classification, nomenclature, and diversity of eukaryotes. Journal of Eukaryotic Microbiology 66: 4–119.

[AS12] Adl, S. M., A. G. B. Simpson, C. E. Lane, J. Lukeš, D. Bass, S. S. Bowser, M. W. Brown, F. Burki, M. Dunthorn, V. Hampl, A. Heiss, M. Hoppenrath, E. Lara, E. Le Gall, D. H. Lynn, H. McManus, E. A. D. Mitchell, S. E. Mozley-Stanridge, L. W. Parfrey, J. Pawlowski, S. Rueckert, L. Shadwick, C. L. Schoch, A. Smirnov & F. W. Spiegel. 2012. The revised classification of eukaryotes. Journal of Eukaryotic Microbiology 59 (5): 429–493.

[A96] Alstrup, V. 1996. Nectria tatrensis sp. nov. and other lichenicolous fungi found on the bryo-lichenological excursion in Slovakia 1993. Biologia 51: 13–14.

[B15] Benjamin, M. S. 1915. A note on the occurrence of urease in legume nodules and other plant parts. Journal and Proceedings of the Royal Society of New South Wales 49 (1): 78–80.

[BSA97] Breton, F., C. Sanier & J. d’Auzac. 1997. Scopoletin production and degradation in relation to resistance of Hevea brasiliensis to Corynespora cassiicola. Journal of Plant Physiology 151: 595–602.

[C-S18] Cavalier-Smith, T. 2018. Kingdom Chromista and its eight phyla: a new synthesis emphasising periplastid protein targeting, cytoskeletal and periplastid evolution, and ancient divergences. Protoplasma 255: 297–357.

[C74] Coineau, Y. 1974. Éléments pour une monographie morphologique, écologique et biologique des Caeculidae (Acariens). Mémoires du Muséum National d'Histoire Naturelle, nouvelle série, Série A, Zoologie 81: 1–299, 24 pls.

[DP72] Deighton, F. C., & K. A. Pirozynski. 1972. Microfungi. V. More hyperparasitic hyphomycetes. Mycological Papers 128: 1–110.

[E99] Eriksson, O. E. (ed.) 1999. Notes on ascomycete systematics. Nos 2440–2755. Myconet 2: 1–41.

[EB03N] Eriksson, O. E., H. O. Barah, R. S. Currah, K. Hansen, C. P. Kurtzman, T. Laessøe & G. Rambold (eds.) 2003. Notes on ascomycete systematics. Nos 3580–3623. Myconet 9: 91–103.

[EB03O] Eriksson, O. E., H. O. Barah, R. S. Currah, K. Hansen, C. P. Kurtzman, G. Rambold & T. Laessøe (eds.) 2003. Outline of Ascomycota—2003. Myconet 9: 1–89.

[ES12] Espadaler, X., & S. Santamaria. 2012. Ecto- and endoparasitic fungi on ants from the Holarctic region. Psyche 2012: 168478.

[KFS03] Keely, S. P., J. M. Fischer & J. R. Stringer. 2003. Evolution and speciation of Pneumocystis. Journal of Eukaryotic Microbiology 50: 624–626.

[LS03] Landvik, S., T. K. Schumacher, O. E. Eriksson & S. T. Moss. 2003. Morphology and ultrastructure of Neolecta species. Mycological Research 107 (9): 1021–1031.

[LV98] Lücking, R., & A. Vězda. 1998. Taxonomic status in foliicolous species of the genus Porina (lichenized Ascomycotina: Trichotheliaceae)—II. The Porina epiphylla group. Willdenowia 28: 181–226.

[LK04] Lutzoni, F., F. Kauff, C. J. Cox, D. McLaughlin, G. Celio, B. Dentinger, M. Padamsee, D. Hibbett, T. Y. James, E. Baloch, M. Grube, V. Reeb, V. Hofstetter, C. Schoch, A. E. Arnold, J. Miadlikowska, J. Spatafora, D. Johnson, S. Hambleton, M. Crockett, R. Shoemaker, G.-H. Sung, R. Lücking, T. Lumbsch, K. O'Donnell, M. Binder, P. Diederich, D. Ertz, C. Gueidan, K. Hansen, R. C. Harris, K. Hosaka, Y.-W. Lim, B. Matheny, H. Nishida, D. Pfister, J. Rogers, A. Rossman, I. Schmitt, H. Sipman, J. Stone, J. Sugiyama, R. Yahr & R. Vilgalys. 2004. Assembling the fungal tree of life: progress, classification, and evolution of subcellular traits. American Journal of Botany 91 (10): 1446–1480.

[MS98] Margulis, L., & K. V. Schwartz. 1998. Five Kingdoms: An Illustrated Guide to the Phyla of Life on Earth 3^rd ed. W. H. Freeman and Company: New York.

[RC11] Rosling, A., F. Cox, K. Cruz-Martinez, K. Ihrmark, G.-A. Grelet, B. D. Lindahl, A. Menkis & T. Y. James. 2011. Archaeorhizomycetes: unearthing an ancient class of ubiquitous soil fungi. Science 333: 876–879.

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