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 3rd 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.

[RS99] Rossman, A. Y., G. J. Samuels, C. T. Rogerson & R. Lowen. 1999. Genera of Bionectriaceae, Hypocreaceae and Nectriaceae (Hypocreales, Ascomycetes). Studies in Mycology 42: 1–248.

[ST99] Saenz, G. S., & J. W. Taylor. 1999. Phylogenetic relationships of Meliola and Meliolina inferred from nuclear small subunit rRNA sequences. Mycological Research 103 (8): 1049–1056.

[SS09] Schoch, C. L., G.-H. Sung, F. López-Giráldez, J. P. Townsend, J. Miadlikowska, V. Hofstetter, B. Robbertse, P. B. Matheny, F. Kauff, Z. Wang, C. Gueidan, R. M. Andrie, K. Trippe, L. M. Ciufetti, A. Wynns, E. Fraker, B. P. Hodkinson, G. Bonito, J. Z. Groenewald, M. Arzanlou, G. S. de Hoog, P. W. Crous, D. Hewitt, D. H. Pfister, K. Peterson, M. Gryzenhout, M. J. Wingfield, A. Aptroot, S.-O. Suh, M. Blackwell, D. M. Hillis, G. W. Griffith, L. A. Castlebury, A. Y. Rossman, H. T. Lumbsch, R. Lücking, B. Büdel, A. Rauhut, P. Diederich, D. Ertz, D. M. Geiser, K. Hosaka, P. Inderbitzin, J. Kohlmeyer, B. Volkmann-Kohlmeyer, L. Mostert, K. O'Donnell, H. Sipman, J. D. Rogers, R. A. Shoemaker, J. Sugiyama, R. C. Summerbell, W. Untereiner, P. R. Johnston, S. Stenroos, A. Zuccaro, P. S. Dyer, P. D. Crittenden, M. S. Cole, K. Hansen, J. M. Trappe, R. Yahr, F. Lutzoni & J. W. Spatafora. 2009. The Ascomycota tree of life: a phylum-wide phylogeny clarifies the origin and evolution of fundamental reproductive and ecological traits. Systematic Biology 58 (2): 224–239.

[SL02] Schweigkofler, W., K. Lopandic, O. Molnár & H. Prillinger. 2002. Analysis of phylogenetic relationships among Ascomycota with yeast phases using ribosomal DNA sequences and cell wall sugars. Organisms Diversity & Evolution 2: 1–17.

[T93] Taylor, T. N. 1993. Fungi. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 9–13. Chapman & Hall: London.

[YK99] Yamada, Y., H. Kawasaki, Y. Nagatsuka, K. Mikata & T. Seki. 1999. The phylogeny of the cactophilic yeasts based on the 18S ribosomal RNA gene sequences: the proposals of Phaffomyces antillensis and Starmera caribaea, new combinations. Bioscience, Biotechnology, and Biochemistry 63: 827–832.

Trichocomaceae

Aspergillus fumigatus, from here.


Belongs within: Eurotiomycetes.
Contains: Penicillium.

The Trichocomaceae are a group of saprobic fungi that are often able to tolerate extreme environmental conditions.

Trichocomaceae [Aspergillaceae]
    |--+--Penicillium SS09
    |  `--Eupenicillium SS09
    |       |--E. javanicum GSSC02
    |       `--E. limosum SS09
    |--+--Byssochlamys nivea LK04
    |  |--Hamigera avellanea SS09
    |  `--Monascaceae EB03
    |       |--Xeromyces bisporus EB03, PHK96
    |       `--Monascus purpureus SS09
    `--Aspergillus SS09
         |  i. s.: A. amstelodami HD91
         |         A. awamori GL98
         |         A. flavipes PG-V03
         |         A. flavus SL02
         |         A. glaucus PHK96
         |         A. niger PHK96
         |         A. ochraceus ZU10
         |         A. oryzae PHK96
         |         A. parasiticus MN03
         |         A. penicilloides HD91
         |         A. soyae PHK96
         |         A. sydowii GSSH02
         |         A. terreus MN03
         |--A. fumigatus SS09
         `--+--A. protuberus SS09
            `--Emericella nidulans SS09 [=Aspergillus nidulans]

Trichocomaceae incertae sedis:
  Eurotium EB03
    |--E. repens [incl. Aspergillus reptans] C91
    `--E. rubrum LS01
  Chaetosartorya EB03
  Chaetotheca EB03
  Cristaspora EB03
  Dactylomyces EB03
  Dendrosphaera EB03
  Dichlaena EB03
  Dichotomomyces EB03
  Edyuillia EB03
  Erythrogymnotheca EB03
  Fennellia EB03
  Hemicarpenteles EB03
  Neosartorya EB03
  Penicilliopsis EB03
  Petromyces EB03
  Sagenoma EB03
  Trichocoma EB03
  Warcupiella EB03
  Chromocleista EB03
    |--C. cinnabarina PA-W02
    `--C. malachitea JC03
  Talaromyces EB03
    |--T. bacillisporus SL02
    `--T. flavus GSSC03
         |--T. f. var. flavus ST02
         `--T. f. var. macrospora ST02
  Thermoascus EB03
    |--T. aurantiacus BY03
    `--T. crustaceus ST99
  Cryptocolax T93
    |--C. clarnensis Scott 1956 T93
    `--C. parvular T93

*Type species of generic name indicated

REFERENCES

[BY03] Baumgartner, M., A. Yapi, R. Gröbner-Ferreira & K. O. Stetter. 2003. Cultivation and properties of Echinamoeba thermarum n. sp., an extremely thermophilic amoeba thriving in hot springs. Extremophiles 7: 267–274.

[C91] Colloff, M. J. 1991. Practical and theoretical aspects of the ecology of house dust mites (Acari: Pyroglyphidae) in relation to the study of mite-mediated allergy. Review of Medical and Veterinary Entomology 79 (11–12): 611–630.

[EB03] 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.

[GSSC02] Gibas, C. F. C., L. Sigler, R. C. Summerbell & R. S. Currah. 2002. Phylogeny of the genus Arachnomyces and its anamorphs and the establishment of Arachnomycetales, a new eurotiomycete order in the Ascomycota. Studies in Mycology 47: 131–139.

[GSSH02] Gibas, C. F. C., L. Sigler, R. C. Summerbell, S. L. R. Hofstader & A. K. Gupta. 2002. Arachnomyces kanei (anamorph Onychocola kanei) sp. nov., from human nails. Medical Mycology 40: 573–580.

[GL98] Giesbert, S., H.-B. Lepping, K. B. Tenberge & P. Tudzynski. 1998. The xylanolytic system of Claviceps purpurea: cytological evidence for secretion of xylanases in infected rye tissue and molecular characterization of two xylanase genes. Phytopathology 88: 1020–1030.

[HD91] Hart, B. J., & A. E. Douglas. 1991. The relationship between house-dust mites and fungi. In: Schuster, R., & P. W. Murphy (eds) The Acari: Reproduction, development and life-history strategies pp. 319–324. Chapman & Hall: London.

[JC03] Jacobs, A., M. P. A. Coetzee, B. D. Wingfield, K. Jacobs & M. J. Wingfield. 2003. Phylogenetic relationships among Phialocephala species and other ascomycetes. Mycologia 95 (4): 637–645.

[LS01] Lumbsch, H. T., I. Schmitt, H. Döring & M. Wedin. 2001. Molecular systematics supports the recognition of an additional order of Ascomycota: the Agyriales. Mycological Research 105 (1): 16–23.

[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.

[MN03] Moffitt, M. C., & B. A. Neilan. 2003. Evolutionary affiliations within the superfamily of ketosynthases reflect complex pathway associations. Journal of Molecular Evolution 56: 446–457.

[PA-W02] Pang, K.-L., M. A. Abdel-Wahab, S. Sivichai, H. M. El-Sharouney & E. B. G. Jones. 2002. Jahnulales (Dothideomycetes, Ascomycota): a new order of lignicolous freshwater ascomycetes. Mycological Research 106 (9): 1031–1042.

[PG-V03] Petrosyan, P., M. García-Varela, A. Luz-Madrigal, C. Huitrón & M. E. Flores. 2003. Streptomyces mexicanus sp. nov., a xylanolytic micro-organism isolated from soil. International Journal of Systematic and Evolutionary Microbiology 53: 269–273.

[PHK96] Prescott, L. M., J. P. Harley & D. A. Klein. 1996. Microbiology 3rd ed. Wm. C. Brown Publishers: Dubuque (Iowa).

[ST99] Saenz, G. S., & J. W. Taylor. 1999. Phylogenetic relationships of Meliola and Meliolina inferred from nuclear small subunit rRNA sequences. Mycological Research 103 (8): 1049–1056.

[SS09] Schoch, C. L., G.-H. Sung, F. López-Giráldez, J. P. Townsend, J. Miadlikowska, V. Hofstetter, B. Robbertse, P. B. Matheny, F. Kauff, Z. Wang, C. Gueidan, R. M. Andrie, K. Trippe, L. M. Ciufetti, A. Wynns, E. Fraker, B. P. Hodkinson, G. Bonito, J. Z. Groenewald, M. Arzanlou, G. S. de Hoog, P. W. Crous, D. Hewitt, D. H. Pfister, K. Peterson, M. Gryzenhout, M. J. Wingfield, A. Aptroot, S.-O. Suh, M. Blackwell, D. M. Hillis, G. W. Griffith, L. A. Castlebury, A. Y. Rossman, H. T. Lumbsch, R. Lücking, B. Büdel, A. Rauhut, P. Diederich, D. Ertz, D. M. Geiser, K. Hosaka, P. Inderbitzin, J. Kohlmeyer, B. Volkmann-Kohlmeyer, L. Mostert, K. O'Donnell, H. Sipman, J. D. Rogers, R. A. Shoemaker, J. Sugiyama, R. C. Summerbell, W. Untereiner, P. R. Johnston, S. Stenroos, A. Zuccaro, P. S. Dyer, P. D. Crittenden, M. S. Cole, K. Hansen, J. M. Trappe, R. Yahr, F. Lutzoni & J. W. Spatafora. 2009. The Ascomycota tree of life: a phylum-wide phylogeny clarifies the origin and evolution of fundamental reproductive and ecological traits. Systematic Biology 58 (2): 224–239.

[SL02] Schweigkofler, W., K. Lopandic, O. Molnár & H. Prillinger. 2002. Analysis of phylogenetic relationships among Ascomycota with yeast phases using ribosomal DNA sequences and cell wall sugars. Organisms Diversity & Evolution 2: 1–17.

[T93] Taylor, T. N. 1993. Fungi. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 9–13. Chapman & Hall: London.

[ZU10] Zvereva, L. V., & L. N. Usheva. 2010. Opportunistic filamentous fungi isolated from pathologically changed internal organs of the mussel Mytilus trossulus (Bivalvia) from Peter the Great Bay, Sea of Japan. In: China-Russia Bilateral Symposium: Proceedings of the China-Russia Bilateral Symposium of "Comparison on Marine Biodiversity in the Northwest Pacific Ocean", 10–11 October 2010, Qingdao (China) pp. 227–231. Institute of Oceanology, Chinese Academy of Sciences; A. V. Zhirmunsky Institute of Marine Biology, Far East Branch of the Russian Academy of Sciences.

Hyalomma

Specimen of Hyalomma marginatum, photographed by Adam Cuerden.


Belongs within: Ixodidae.

Hyalomma is a genus of large, highly active ticks found in Eurasia and Africa (Keirans 2009).

Characters (from Keirans 2009): Scutum inornate; eyes large, bulging upward from distinct, deep sockets. Palpi much longer than basis capituli, palpal segment II at least twice as long as broad. Males with ventral plates, with 7–11 festoons, partially coalesced.

<==Hyalomma Koch 1844 H98
    |--H. aegyptium (Linnaeus 1758) H98 (see below for synonymy)
    |--H. anatolicum KK91
    |--H. asiaticum Sch. & Schl. 1929 P91
    |--H. dentatum Canestrini & Fanzago 1877 CF77
    |--H. detritum KK91
    |--H. dromedarii Koch 1844 [=H. aegyptium dromedarii] A56
    |--H. excavatum Koch 1844 A56
    |--H. lusitanicum BS96
    |--H. marginatum K09
    |    |--H. m. marginatum R91
    |    `--H. m. rufipes (Koch 1844) R91, ADG91
    |--H. plumbeum KK91
    |--H. rufipes A56
    |    |--H. r. rufipes A56
    |    `--H. r. glabrum Delpy 1949 A56
    |--H. schulzei Olenev 1931 A56
    |--H. scupense KK91
    `--H. truncatum (Koch 1844) ADG91

Hyalomma aegyptium (Linnaeus 1758) H98 [=Acarus aegyptius H98, Amblyomma aegyptium H98; incl. Hyalomma syriacum Koch 1844 A56]

*Type species of generic name indicated

REFERENCES

[ADG91] Amoo, A. O. J., O. O. Dipeolu & L. N. Gichuru. 1991. Tick ecology: on-host population dynamics of ticks in the Coast Province of Kenya. In: Dusbábek, F., & V. Bukva (eds) Modern Acarology: Proceedings of the VIII International Congress of Acarology, held in České Budĕjovice, Czechoslovakia, 6–11 August 1990 vol. 2 pp. 461–468. SPB Academic Publishing: The Hague.

[A56] Anastos, G. 1956. The ticks (Acarina: Ixodoidea) of the J. Klapperich Afghanistan Expedition, 1952 and 1953. Journal of the Washington Academy of Sciences 46 (1): 18–19.

[BS96] Burks, C. S., R. L. Stewart Jr, G. R. Needham & R. E. Lee Jr. 1996. Cold hardiness in the ixodid ticks (Ixodidae). In: Mitchell, R., D. J. Horn, G. R. Needham & W. C. Welbourn (eds) Acarology IX vol. 1. Proceedings pp. 85–87. Ohio Biological Survey: Columbus (Ohio).

[CF77] Canestrini, G., & F. Fanzago. 1877. Intorno agli Acari Italiani. Atti del Reale Istituto Veneto di Scienze, Lettere ed Arti, Serie 5, 4: 69–208, pls 2–7.

[H98] Halliday, R. B. 1998. Mites of Australia: A checklist and bibliography. CSIRO Publishing: Collingwood.

[KK91] Kamarinchev, B., T. Kovacheva, T. Christova, G. Georgieva & V. Zlatanova. 1991. Studies on mosquitoes and ticks as carriers of alpha-, flavi- and bunyaviruses. In: Dusbábek, F., & V. Bukva (eds) Modern Acarology: Proceedings of the VIII International Congress of Acarology, held in České Budĕjovice, Czechoslovakia, 6–11 August 1990 vol. 2 pp. 89–92. SPB Academic Publishing: The Hague.

[K09] Keirans, J. E. 2009. Order Ixodida. In: Krantz, G. W., & D. E. Walter (eds) A Manual of Acarology 3rd ed. pp. 111–123. Texas Tech University Press.

[P91] Podboronov, V. M. 1991. Antibacterial protective mechanisms of ixodoid ticks. In: Dusbábek, F., & V. Bukva (eds) Modern Acarology: Proceedings of the VIII International Congress of Acarology, held in České Budĕjovice, Czechoslovakia, 6–11 August 1990 vol. 2 pp. 375–380. SPB Academic Publishing: The Hague.

[R91] Rechav, Y. 1991. The ecology of the vectors of Crimean-Congo haemorrhagic fever in southern Africa. In: Dusbábek, F., & V. Bukva (eds) Modern Acarology: Proceedings of the VIII International Congress of Acarology, held in České Budĕjovice, Czechoslovakia, 6–11 August 1990 vol. 2 pp. 93–99. SPB Academic Publishing: The Hague.

Ptyctima

Lateral view of dorsal plates (A) and ventral plates (B) of Microtritia minima, from Balogh & Balogh (1992, The Oribatid Mites Genera of the World vol. 2).


Belongs within: Acariformes.
Contains: Austrotritia, Mesotritia, Oribotritia, Indotritia, Euphthiracarus, Acrotritia, Phthiracaridae.

The Ptyctima are a group of oribatid mites with a ptychoid body plan: that is, the legs can be fully withdrawn and the aspis rotated ventrad to cover the legs against the notogaster and ventral plate. Two major lineages are recognised in the Ptyctima, the Phthiracaridae and Euphthiracaroidea.

The Euphthiracaroidea are a group of ptychoid mites in which the body is strongly compressed laterally. Retraction of the legs is facilitated by relaxation of the notogaster to increase its volume (Norton 2001). In the family Synichotritiidae, the genital setae are in an antaxial position, in groups or transverse rows, while the Oribotritiidae and Euphthiracaridae both have the genital setae in a paraxial position, in a longitudinal row. The latter two families are distinguished in that the Euphthiracaridae have the ventral (anal and genital) plates fused into a single plate pair, while the Oribotritiidae have them at least partially separated. In the Bornean Temburongia patoi, the sides of the notogaster are produced ventrolaterally into two lobes that contact each other medially, in contrast to the simple notogastral margins leaving the V-shaped ventral region exposed in other genera (Balogh & Balogh 1992).

<==Ptyctima [Euptyctima] PL17
    |--Euphthiracaroidea DW10
    |    |--Synichotritiidae S04
    |    |    |--Apotritia Norton & Lions 1992 S04
    |    |    |    `--*A. walkeri Norton & Lions 1992 S04
    |    |    |--Temburongia Mahunka 1990 [Temburongiidae] S04
    |    |    |    `--*T. patoi Mahunka 1990 S04
    |    |    |--Synichotritia Walker 1965 S04
    |    |    |    |--*S. caroli Walker 1965 S04
    |    |    |    |--S. longipila Niedbała 2002 S04
    |    |    |    |--S. parvipilosa Niedbała 2002 S04
    |    |    |    `--S. spinulosa Walker 1965 S04
    |    |    `--Sabahtritia Mahunka 1987 S04
    |    |         |--*S. hauseri Mahunka 1987 S04
    |    |         |--S. foveolata (Hu, Wang & Aoki 1991) [=Synichotritia foveolata] S04
    |    |         |--S. furcata (Hu & Wang 1992) [=Synichotritia furcata] S04
    |    |         |--S. lienhardi Mahunka 1995 S04
    |    |         |--S. mirabilis Mahunka 1991 S04
    |    |         |--S. sarawak Mahunka 1996 S04
    |    |         `--S. tianmuensis (Hu, Wang & Aoki 1991) [=Synichotritia tianmuensis] S04
    |    |--Oribotritiidae S04
    |    |    |--Austrotritia S04
    |    |    |--Mesotritia S04
    |    |    |--Oribotritia S04
    |    |    |--Indotritia S04
    |    |    |--Paratritia Moritz 1966 S04
    |    |    |    `--*P. baloghi Moritz 1966 S04
    |    |    |--Terratritia Ramsay & Sheals 1969 S04
    |    |    |    |--*T. askewi Ramsay & Sheals 1969 S04
    |    |    |    `--T. seconda Mahunka 1991 S04
    |    |    |--Sabacarus Ramsay & Sheals 1969 S04
    |    |    |    |--*S. corneri Ramsay & Sheals 1969 [incl. S. ranokaoensis Hammer 1970] S04
    |    |    |    `--S. japonicus Shimano & Aoki 1997 S04
    |    |    |--Maerkelotritia Hammer 1967 S04
    |    |    |    |--M. kishidai (Aoki 1958) S04 (see below for synonymy)
    |    |    |    |--M. crytopa (Banks 1904) S04 (see below for synonymy)
    |    |    |    `--M. krivolutskyi Märkel 1968 S04
    |    |    |         |--M. k. krivolutskyi S04
    |    |    |         `--M. k. mongolica Mahunka 1981 S04
    |    |    `--Protoribotritia Jacot 1938 S04
    |    |         |--*P. canadaris Jacot 1938 S04
    |    |         |--P. aberrans (Märkel & Meyer 1959) [=Oribotritia aberrans] S04
    |    |         |--P. africana Mahunka 1985 S04
    |    |         |--P. ensifer Aoki 1969 [=P. aberrans ensifer] S04
    |    |         `--P. oligotricha Märkel 1963 S04
    |    `--Euphthiracaridae [Euphthiracarini] S04
    |         |--Euphthiracarus S04
    |         |--Acrotritia S04
    |         |--Bukitritia Mahunka 1990 S04
    |         |    `--*B. timah Mahunka 1990 S04
    |         |--Sumatrotritia Mahunka 1989 S04
    |         |    |--*S. inusitata Mahunka 1989 S04
    |         |    |--S. elegans Mahunka 1991 S04
    |         |    `--S. murphyi Mahunka 1999 S04
    |         `--Microtritia Märkel 1964 S04
    |              |--*M. minima (Berlese 1904) (see below for synonymy) S04
    |              |--M. contraria Niedbała 1993 S04
    |              |--M. fissurata Märkel 1968 S04
    |              |--M. fusa Niedbała 2000 S04
    |              |--M. glabrata Niedbała 1993 S04
    |              |--M. hauseri Mahunka 1993 [incl. M. altissima Mahunka 1994] S04
    |              |--M. hawaiiensis Niedbała 1994 S04
    |              |--M. incisa Märkel 1964 S04
    |              |--M. neonominata Subías 2004 [=M. glabrata Starý 1993 non Niedbała 1993] S04
    |              |--M. schusteri Märkel 1964 S04
    |              |--M. similis N01
    |              |--M. simplex (Jacot 1930) S04 (see below for synonymy)
    |              |--M. striatissima Mahunka 1999 S04
    |              |--M. tropica Märkel 1964 S04
    |              |    |--M. t. tropica [incl. M. ethiopica Mahunka 1988] S04
    |              |    `--M. t. dusan Ramsay & Sheals 1969 S04
    |              `--M. tumida Niedbała 1988 S04
    `--+--Phthiracaridae DW10
       `--Nehypochthonius Norton & Metz 1980 PL17, S04 [Nehypochthoniidae, Nehypochthonoidea]
            |--*N. porosus Norton & Metz 1980 S04
            `--N. yanoi Aoki 2002 S04

Maerkelotritia crytopa (Banks 1904) S04 [=Phthiracarus crytopus S04; incl. Oribotritia gibbera Walker 1965 S04, Maerkelotritia gibbera A80, Phthiracarus maximus Ewing 1913 S04]

Maerkelotritia kishidai (Aoki 1958) S04 [=Oribotritia kishidai S04; incl. *M. alaskensis Hammer 1967 S04, O. sellnicki Walker 1965 S04, M. sellnicki A80]

*Microtritia minima (Berlese 1904) [=Phthiracarus minimus; incl. Tritia (Pseudotritia) minuta Willmann 1919, Microtritia silofila Subías 1975] S04

Microtritia simplex (Jacot 1930) S04 [=Pseudotritia simplex S04; incl. Rhysotritia paeneminima Walker 1965 S04, Microtritia paeneminima NP91]

*Type species of generic name indicated

REFERENCES

[A80] Aoki, J. 1980. A revision of the oribatid mites of Japan. I. The families Phthiracaridae and Oribotritiidae. Bulletin of the Institute of Environmental Science and Technology, Yokohama National University 6 (2): 1–89.

[DW10] Dabert, M., W. Witalinski, A. Kazmierski, Z. Olszanowski & J. Dabert. 2010. Molecular phylogeny of acariform mites (Acari, Arachnida): strong conflict between phylogenetic signal and long-branch attraction artifacts. Molecular Phylogenetics and Evolution 56 (1): 222–241.

[N01] Niedbała, W. 2001. Study of the diversity of Ptyctima (Acari: Oribatida) and quest for centres of its origin: the fauna of the Oriental and Australian regions. In: Halliday, R. B., D. E. Walter, H. C. Proctor, R. A. Norton & M. J. Colloff (eds) Acarology: Proceedings of the 10th International Congress pp. 143–148. CSIRO Publishing: Melbourne.

[NP91] Norton, R. A., & S. C. Palmer. 1991. The distribution, mechanisms and evolutionary significance of parthenogenesis in oribatid mites. In: Schuster, R., & P. W. Murphy (eds) The Acari: Reproduction, development and life-history strategies pp. 107–136. Chapman & Hall: London.

[PL17] Pachl, P., A. C. Lindl, A. Krause, S. Scheu, I. Schaefer & M. Maraun. 2017. The tropics as an ancient cradle of oribatid mite diversity. Acarologia 57 (2): 309–322.

[S04] Subías, L. S. 2004. Listado sistemático, sinonímico y biogeográfico de los ácaros oribátidos (Acariformes, Oribatida) del mundo (1758–2002). Graellsia 60 (número extraordinario): 3–305.

Steganacarus

Steganacarus magnus, from Micropics.


Belongs within: Phthiracaridae.

Steganacarus is a genus of phthiracarid mites found in the Holarctic, Afrotropical and Neotropical regions in which seta d on tibia IV is long and removed from the solenidion.

<==Steganacarus Ewing 1917 S04
    |--S. (Steganacarus) [incl. Trachyhoplophora Berlese 1923] S04
    |    |--S. (S.) magnus (Nicolet 1855) S04 (see below for synonymy)
    |    |--S. (S.) applicatus (Sellnick 1920) (see below for synonymy) S04
    |    |--S. (S.) balearicus Pérez-Íñigo 1969 S04
    |    |--S. (S.) bicarinatus Jeleva 1970 S04
    |    |--S. (S.) caelestis Niedbała 1984 S04
    |    |--S. (S.) carlosi Niedbała 1984 S04
    |    |--S. (S.) carusoi Bernini & Avanzati 1989 [incl. S. applicatus crassisetosus Willmann 1939 (n. d.)] S04
    |    |--S. (S.) coniunctus Niedbała 1983 [incl. S. danae Mahunka & Miko 1989] S04
    |    |--S. (S.) donatoi Avanzati, Baratti & Bernini 1994 S04
    |    |--S. (S.) doumandji Niedbała 1986 S04
    |    |--S. (S.) flagellatissimus Mahunka 1979 S04
    |    |--S. (S.) grandjeani Mahunka 1977 [incl. S. sacculiferus Parry 1978] S04
    |    |--S. (S.) guanarteme Pérez-Íñigo & Peña 1996 S04
    |    |--S. (S.) herculeanus Willmann 1953 [incl. S. longisetosus Moraza 1985] S04
    |    |--S. (S.) hirsutus Pérez-Íñigo 1974 S04
    |    |    |--S. h. hirsutus [incl. S. perezinigoi Balogh & Mahunka 1979] S04
    |    |    `--S. h. azorensis Pérez-Íñigo 1992 S04
    |    |--S. (S.) inaestimabilis Niedbała 1984 S04
    |    |--S. (S.) incognitus Niedbała 1984 S04
    |    |--S. (S.) incomptus Niedbała 1983 S04
    |    |--S. (S.) insulanus Pérez-Íñigo & Pérez-Íñigo 1996 S04
    |    |--S. (S.) inurbanus Niedbała 1983 S04
    |    |--S. (S.) manganellii Avanzati, Baratti & Bernini 1994 S04
    |    |--S. (S.) maximus (Haupt 1882) (n. d.) [=Hoplophora maxima] S04
    |    |--S. (S.) michaeli Bernini & Avanzati 1987 S04
    |    |--S. (S.) nemorosus Gordeeva 1991 S04
    |    |--S. (S.) patruelis Niedbała 1983 [incl. S. lazitanicus Mahunka & Miko 1989] S04
    |    |--S. (S.) personatus Niedbała 1983 S04
    |    |--S. (S.) similis Willmann 1939 S04
    |    |--S. (S.) simonettae Bernini & Avanzati 1987 S04
    |    |--S. (S.) sol Balogh 1958 S04 [incl. Hoplophthiracarus peracutus Mahunka 1983 S04, S. peracutus N86]
    |    |--S. (S.) spinosus (Sellnick 1920) (see below for synonymy) S04
    |    |--S. (S.) tenerifensis Pérez-Íñigo 1972 S04
    |    |--S. (S.) travei Lions 1968 S04
    |    |--S. (S.) vernaculus Niedbała 1982 S04
    |    |--S. (S.) vestitus Niedbała 1983 [incl. S. complicatus Mahunka 1993] S04
    |    |--S. (S.) wallworki Mahunka 1984 S04
    |    `--S. (S.) werneri Mahunka 1993 S04
    |--S. (Rafacarus Niedbała 1982) (see below for synonymy) S04
    |    |--S. (*R.) rafalskii (Niedbała 1982) S04 (see below for synonymy)
    |    |--S. (R.) angulatus (Balogh & Mahunka 1992) [=Neosteganacarus angulatus] S04
    |    |--S. (R.) niedbalai (Pérez-Íñigo & Baggio 1996) [=Protophthiracarus niedbalai] S04
    |    |--S. (R.) relictus (Balogh & Mahunka 1992) [=Neosteganacarus relictus] S04
    |    `--S. (R.) valeriae (Balogh & Mahunka 1992) [=Nortonacarus valeriae] S04
    `--S. (Tropacarus Ewing 1917) [incl. Calhoplophora Berlese 1923] S04
         |--S. (*T.) carinatus (Koch 1841) S04 (see below for synonymy)
         |--S. (T.) balcanicus Berninni & Avanzati 1989 S04
         |--S. (T.) boulfekhari Niedbała 1986 S04
         |--S. (T.) brevipilus (Berlese 1923) S04 (see below for synonymy)
         |--S. (T.) callainii Bernini, Bernini & Avanzati 1989 S04
         |--S. (T.) desmeti Niedbała 1988 [incl. S. (T.) altitudinis Bernini, Bernini & Avanzati 1989] S04
         |--S. (T.) excavatus (Niedbała 1981) [=Tropacarus excavatus] S04
         |--S. (T.) lasithiensis Mahunka 1979 S04
         |--S. (T.) maghrebinus Bernini, Bernini & Avanzati 1989 S04
         |--S. (T.) malatestai Avanzati, Baratti & Bernini 1994 S04
         `--S. (T.) pseudocarinatus Bernini & Avanzati 1989 S04

Steganacarus (Rafacarus Niedbała 1982) [incl. Neosteganacarus Balogh & Mahunka 1992, Nortonacarus Balogh & Mahunka 1992] S04

Steganacarus (*Rafacarus) rafalskii (Niedbała 1982) S04 [=Rafacarus rafalskii S04; incl. Neosteganacarus cataracta Balogh & Mahunka 1992 S04]

Steganacarus (Steganacarus) applicatus (Sellnick 1920) [=Phthiracarus applicatus; incl. P. berlesei Oudemans 1915 non Michael 1898, Hoplophora quadriseriata Haupt 1882] S04

Steganacarus (Steganacarus) magnus (Nicolet 1855) S04 [=Hoplophorella magna S04, Hoploderma magnum S22, Hoplophora magna BA87; incl. Hoplophora anomala Berlese 1883 S04, Hoploderma anomala CH98, *Steganacarus anomalus S04, Phthiracarus magnus anomaloides Oudemans 1915 S04, S. karamani Tarman 1962 S04, P. magna f. oblonga Oudemans 1915 N86, P. magnus rotundus Oudemans 1915 non Ewing 1908 S04, P. magnus transitans Oudemans 1915 S04]

Steganacarus (Steganacarus) spinosus (Sellnick 1920) [=Phthiracarus spinosus; incl. S. barborae Mahunka & Miko 1989, S. punctulatus Sergienko 1985] S04

Steganacarus (Tropacarus) brevipilus (Berlese 1923) S04 [=Phthiracrus (Trachyhoplophora) brevipilus S04, Tropacarus curtipilus (l. c.) N86; incl. Hoploderma brevipilus perfecta Sellnick 1931 S04]

Steganacarus (*Tropacarus) carinatus (Koch 1841) S04 [=Hoplophora carinata S04, Calhoplophora carinata N86, Hoploderma carinatum BA89, Hoplophora carinatum BA89; incl. Hoplophora carinata var. pulcherrima Berlese 1887 N86, S04, Steganacarus (Tropacarus) pulcherrimus BA91]

*Type species of generic name indicated

REFERENCES

[BA87] Bernini, F., & A. M. Avanzati. 1987. Notulae oribatologicae XLI. Some new Steganacarus species belonging to the magnus group (Acarida, Oribatida). International Journal of Acarology 13 (3): 165–176.

[BA89] Bernini, F., & A. M. Avanzati. 1989. Notulae oribatologicae XLVI. Steganacarus (Tropacarus) carinatus (C. L. Koch, 1841) and two new Mediterranean Tropacarus species (Acarida, Oribatida). Acarologia 30 (2): 143–161.

[BA91] Bernini, F., & A. M. Avanzati. 1991. A new approach to the systematics of the genus Steganacarus (Oribatida). In: Schuster, R., & P. W. Murphy (eds) The Acari: Reproduction, development and life-history strategies pp. 335–342. Chapman & Hall: London.

[CH98] Colloff, M. J., & R. B. Halliday. 1998. Oribatid Mites: A catalogue of Australian genera and species. CSIRO Publishing: Collingwood.

[N86] Niedbała, W. 1986. Catalogue des Phthiracaroidea (Acari), clef pour la détermination des espèces et descriptions d'espèces nouvelles. Annales Zoologici 40 (4): 309–370.

[N91] Niedbała, W. 1991. Preliminary hypotheses on the biogeography of the Phthiracaroidea (Acari, Oribatida). In: Dusbábek, F., & V. Bukva (eds) Modern Acarology: Proceedings of the VIII International Congress of Acarology, held in České Budĕjovice, Czechoslovakia, 6–11 August 1990 vol. 2 pp. 219–231. SPB Academic Publishing: The Hague.

[S22] Schweizer, J. 1922. Beitrag zur Kenntnis der terrestrischen Milbenfauna der Schweiz. Verhandl. Naturf. Ges. Basel 33: 23–112, 4 pls.

[S04] Subías, L. S. 2004. Listado sistemático, sinonímico y biogeográfico de los ácaros oribátidos (Acariformes, Oribatida) del mundo (1758–2002). Graellsia 60 (número extraordinario): 3–305.

Glycyphagidae

Glycyphagus sp., copyright Daktaridudu.


Belongs within: Astigmatina.

The Glycyphagidae are a diverse group of mites mostly associated with the nests of small mammals, mostly characterised by the presence of a series of external ridges on the venter of the subcapitulum (OConnor 2009).

Characters (from OConnor 2009): Ventral subcapitulum with prominent pattern of external transverse and oblique ridges (ridges reduced in heavily sclerotised Fusacarus); palpi reflexed inwards; empodial claws small or absent; condylophores very thin or absent; female usually with epigynal apodeme and often with external copulatory tube, oviporus longitudinal with genital valves fused to body anteriorly, free posteriorly; male without paranal suckers or modified setae on tarsus IV. Deutonymph with attachment organ with setae p1–p2 in the form of ribbed claspers, or if absent, then vestiges of their alveoli present as small sclerotised rings (Baloghella), or attachment organ reduced and tibiae III–IV with large ventral, comblike setae; coxal setae 3a and 4a absent; posterior coxal apodemes with distinct edges.

<==Glycyphagidae [Glycyphagina] O09
    |--Labidophorus P90 [Labidophorinae O09]
    |--Marsupialichus [Marsupialichinae] O09
    |    `--M. johnstoni O09
    |--Baloghella [Melesodectinae] O09
    |    `--B. melis [incl. Melesodectes auricularis] O09
    |--Fusacarinae [Fusacaridae] O09
    |    |--Fusacarus O09
    |    `--Xenoryctes krameri O09, GHG91
    |--Metalabidophorinae O09
    |    |--Metalabidophorus O09
    |    |--Paralabidophorus O09
    |    `--Neolabidophorus yucatanensis Pence & Genoways 1974 O09, QV01
    |--Lophuromyopus [Lophuromyopodinae] O09
    |    `--L. schoutedeni O09
    `--Glycyphagus Hering 1838 H98 [incl. Lepidoglyphus Zakhvatkin 1936 H98; Glycyphaginae O09]
         |--G. destructor (Schrank 1781) W91 (see below for synonymy)
         |--G. domesticus (de Geer 1778) [=Acarus domesticus, Glyciphagus (l. c.) domesticus] H98
         |--G. hypudaei O09
         |--G. latipes Berlese 1910 V25
         |--G. ornatus (Kramer 1881) W91
         `--G. spinipes P94

Glycyphagidae incertae sedis:
  Dermacarus O09
    |--D. hypudaei GHG91
    |--D. ornatus Fain 1967 QV01
    `--D. tamiasciuri O09
  Prolepidoglyphus M-PV-G01
  Gohieria Oudemans 1939 [incl. Ferminia Oudemans 1928 (preoc.)] H98
    `--G. fusca (Oudemans 1902) [=Glycyphagus fuscus, Ferminia fuscus] H98
  Zibethacarus P90
  Ctenoglyphus Berlese 1884 [=Cthenoglyphus] H98
    `--C. plumiger (Koch 1835) [=Acarus plumiger, Cthenoglyphus plumiger, Glycyphagus plumiger] H98
  Orycteroxenus soricis O09
  Dipodomyopus P90
  Mediolabidophorus P90
  Microlabidopus P90
  Scalopacarus P90
  Neoxenoryctes P90
  Rodentopus sciuri O09, WK09
  Grammolichus malukuensis O09
  Pycnoglycyphagus O09
  Lophioglyphus liciosus [incl. Apodemopus apodemi] O09
  Diamesoglyphus intermedius O09
  Cryptomyopus zachvatkini O09

Glycyphagus destructor (Schrank 1781) W91 [=Acarus destructor H98, Lepidoglyphus destructor W91; incl. A. cadaverum Schrank 1781 H98, G. cadaverum H98]

*Type species of generic name indicated

REFERENCES

[GHG91] Gállego, M., E. Hidalgo & J. Ginés. 1991. Mites of the house mouse, Mus musculus L., in the north-eastern part of the Iberian Peninsula in Spain. In: Schuster, R., & P. W. Murphy (eds) The Acari: Reproduction, development and life-history strategies pp. 453–454. Chapman & Hall: London.

[H98] Halliday, R. B. 1998. Mites of Australia: A checklist and bibliography. CSIRO Publishing: Collingwood.

[M-PV-G01 Montiel-Parra, G., G. A. Villegas-Guzman, M. Vargas & O. J. Polaco. 2001. Mites associated with nests of Neotoma albigula Hartley, 1894 (Rodentia: Muridae) in Durango, México. In: Halliday, R. B., D. E. Walter, H. C. Proctor, R. A. Norton & M. J. Colloff (eds) Acarology: Proceedings of the 10th International Congress pp. 586–593. CSIRO Publishing: Melbourne.

[O09] OConnor, B. M. 2009. Cohort Astigmatina. In: Krantz, G. W., & D. E. Walter (eds) A Manual of Acarology 3rd ed. pp. 565–657. Texas Tech University Press.

[P90] Philips, J. R. 1990. Acarina: Astigmata (Acaridida). In: Dindal, D. L. (ed.) Soil Biology Guide pp. 757–778. John Wiley & Sones: New York.

[P94] Pugh, P. J. A. 1994. Non-indigenous Acari of Antarctica and the sub-Antarctic islands. Zoological Journal of the Linnean Society 110: 207–217.

[QV01] Quintero, M. M. T., M. Vargas, B. S. Hernández, P García & N. J. Otero. 2001. Ectoparasitic mites on Heteromys gaumeri in the south of Yucatan, Mexico. In: Halliday, R. B., D. E. Walter, H. C. Proctor, R. A. Norton & M. J. Colloff (eds) Acarology: Proceedings of the 10th International Congress pp. 583–585. CSIRO Publishing: Melbourne.

[V25] Vitzthum, H. 1925. Fauna sumatrensis. (Beitrag no. 5). Acarinae. Supplementa Entomologica 11: 1–79.

[WK09] Walter, D. E., & G. W. Krantz. 2009. Oviposition and life stages. In: Krantz, G. W., & D. E. Walter (eds) A Manual of Acarology 3rd ed. pp. 57–63. Texas Tech University Press.

[W91] Walzl, M. G. 1991. Comparison of the sclerotized structures of Acaridae and Glycyphagidae used for copulation. In: Dusbábek, F., & V. Bukva (eds) Modern Acarology: Proceedings of the VIII International Congress of Acarology, held in České Budĕjovice, Czechoslovakia, 6–11 August 1990 vol. 2 pp. 283–286. SPB Academic Publishing: The Hague.