Mesomycetozoea

Mesomycetozoea
	Sphaeroforma arctica
Scientific classification
Domain:	Eukaryota
(unranked):	Amorphea
(unranked):	Obazoa
(unranked):	Opisthokonta
(unranked):	incertae sedis
Class:	Mesomycetozoea; Mendoza et al. 2002
Orders
	Dermocystida; Ichthyophonida;
Synonyms[1]
	Ichthyosporea Cavalier-Smith 1998

The Mesomycetozoea (or DRIP clade, or Ichthyosporea) are a small group of Opisthokonta in Eukaryota (formerly protists), mostly parasites of fish and other animals.

Significance

They are not particularly distinctive morphologically, appearing in host tissues as enlarged spheres or ovals containing spores, and most were originally classified in various groups as fungi, protozoa, or colorless algae. However, they form a coherent group on molecular trees, closely related to both animals and fungi and so of interest to biologists studying their origins. In a 2008 study they emerge robustly as the sibling-group of the clade Filozoa, which includes the animals.[2][3]

Huldtgren et al., following x-ray tomography of microfossils of the Ediacaran Doushantuo Formation, has interpreted them as mesomycetozoan spore capsules.[4]

Terminology

The name DRIP is an acronym for the first protozoa identified as members of the group,[5] Cavalier-Smith later treated them as the class Ichthyosporea, since they were all parasites of fish.

order Dermocystida
- "D": Dermocystidium. One species, Rhinosporidium seeberi, infects birds and mammals, including humans.
- "R": the "rosette agent", now known as Sphaerothecum destruens
order Ichthyophonida
- "I": Ichthyophonus
- "P": Psorospermium

Since other new members have been added (e.g. the former fungal orders Eccrinales and Amoebidiales), Mendoza et al. suggested changing the name to Mesomycetozoea, which refers to their evolutionary position. On Eukaryota tree, in Opisthokont clade, Mesomycetozoea is in the middle ("Meso-") of the fungi ("-myceto-") and the animals ("-zoea").[6] Note the name Mesomycetozoa (without a third e) is also used to refer to this group, but Mendoza et al. use it as an alternate name for basal Opisthokonts.[7]

Phylogeny

Eukaryota tree. Note "Ichthyosporea" at bottom left, in Opisthokont clade. "Metazoa" are animals, and Choanoflagellates are closely aligned. Fungi is at other end of Opisthokont clade, with Cristidiscoidea closely aligned. Ichthyosporea is in the middle ("Meso-") of the fungi ("-myceto-") and the animals ("-zoea").

Ichthyosporea[8][9]

Dermocystida

Rhinosporidiaceae

Sphaerothecum

Dermocystidium

Valentines

Rhinosporidium

Amphibiocystidium

	Dermotheca

	Dermosporidium

Ichthyophonida

Creolimacidae

Anurofeca

	Creolimax

	Sphaeroforma

Psorospermidae

Psorospermis

Piridae

Abeoforma

	Caullerya

	Pirum

Trichomycina

Ichthyophonidae

Eccrinida

Parataeniellaceae

Parataeniella

Eccrinaceae

Eccrinidus

Enterobryus

Leidyomyces

	Enteromyces

	Arundinula

Enteropogon

Taeniella

Taeniellopsis

	Astreptonema

	Palavascia

Taxonomy

Class Ichthyosporea Cavalier-Smith 1998[10][11]
- Order Dermocystida Cavalier-Smith 1998
  - Family Rhinosporidiaceae Mendoza et al. 2001
    - Genus Amphibiocystidium Pascolini et al. 2003
    - Genus Chromosphaera Grau-Bové et al. 2017
    - Genus Dermocystidium Pérez 1908 [Amphibiothecum Feldman, Wimsatt & Green, 2005; Dermocystis Pérez 1907 non]
    - Genus Dermosporidium Carini 1940
    - Genus Dermotheca
    - Genus Rhinosporidium Minchin & Fantham 1905
    - Genus Sphaerothecum Arkush et al. 2003 (Rosette agent)
    - Genus Valentines Borteiro et al. 2018
- Order Ichthyophonida Cavalier-Smith 1998 [Ichthyophonae Mendoza et al. 2001; Amoebidiales; Eccrinales Léger & Dubosq 1929]
  - Suborder Sphaeroformina Cavalier-Smith 2012
    - Family Creolimacidae Cavalier-Smith 2012
      - Genus Anurofeca Baker, Beebee & Ragan 1999
      - Genus Creolimax Marschall et al. 2008
      - Genus Sphaeroforma Jostensen et al. 2002 [Pseudoperkinsus Figueras et al. 2000]
    - Family Psorospermidae Cavalier-Smith 2012
      - Genus Psorospermis Cavalier-Smith 2012 [Psorospermium Hilgendorf 1883 non Eimer 1870]
    - Family Piridae Cavalier-Smith 2012
      - Genus Caullerya Chatton 1907
      - Genus Abeoforma Marshall & Berbee 2011
      - Genus Pirum Marshall & Berbee 2011
  - Suborder Trichomycina Cavalier-Smith 2012
    - Genus †Paleocadus Poinar 2016
    - Family Amoebidiidae Lichtenstein 1917 ex Kirk et al. 2001
      - Genus Amoebidium Cienkowski 1861
    - Family Ichthyophonidae Cavalier-Smith 2012
      - Genus Ichthyophonus Plehn & Mulsow 1911
    - Family Paramoebidiidae Reynolds et al. 2017
      - Genus Paramoebidium Léger & Duboscq 1929
    - Family Parataeniellaceae Manier & Lichtward 1968
      - Genus Lajassiella Tuzet & Manier 1951 ex Manier 1968
      - Genus Nodocrinella Scheer 1977
      - Genus Parataeniella Poiss. 1929
    - Family Eccrinaceae Leger & Duboscq 1929 [Palavasciaceae Manier & Lichtward 1968]
      - Genus Alacrinella Manier & Ormières ex Manier 1968
      - Genus Arundinula Léger & Duboscq 1906 [Arundinella Léger & Duboscq 1905 non Raddi 1823]
      - Genus Astreptonema Hauptfleisch 1895 [Eccrinella Léger & Duboscq 1933]
      - Genus Eccrinidus Manier 1970
      - Genus Eccrinoides Léger & Duboscq 1929 [Eccrinopsis Léger & Duboscq 1916]
      - Genus Enterobryus [Andohaheloa Manier 1955; Capillus Granata 1908; Cestodella Tuzet, Manier & Jolivet 1957; Daloala Tuzet, Manier & Vog.-Zuber 1952; Eccrina Leidy 1852; Lactella Maessen 1955; Paratrichella Manier 1947; Pistillaria Jeekel et al. 1959 non Fries 1821; Recticoma Scheer 1935; Trichella Léger & Duboscq 1929; Trichellopsis Maessen 1955]
      - Genus Enteromyces Lichtwardt 1961
      - Genus Enteropogon Hibbits 1979 non Nees 1836
      - Genus Leidyomyces Lichtwardt et al. 1999
      - Genus Palavascia Tuzet & Manier 1947 ex Lichtwardt 1964
      - Genus Paramacrinella Manier & Grizel 1971
      - Genus Passalomyces Lichtwardt et al. 1999
      - Genus Ramacrinella Manier & Ormiéres 1962 ex Manier 1968
      - Genus Taeniella Léger & Duboscq 1911
      - Genus Taeniellopsis Poisson 1927

References

Cavalier-Smith, T. 1998. Neomonada and the origin of animals and fungi. In: Coombs GH, Vickerman K, Sleigh MA, Warren A (ed.) Evolutionary relationships among protozoa. Kluwer, London, pp. 375-407,
Shalchian-Tabrizi K., Minge M.A., Espelund M.; et al. (7 May 2008). Aramayo, Rodolfo (ed.). "Multigene Phylogeny of Choanozoa and the Origin of Animals". PLOS ONE. 3 (5): e2098. Bibcode:2008PLoSO...3.2098S. doi:10.1371/journal.pone.0002098. PMC 2346548. PMID 18461162.{{cite journal}}: CS1 maint: multiple names: authors list (link)
Dawkins, Richard; Wong, Yan (2016). The Ancestor's Tale. ISBN 978-0544859937.
Douglas Fox, "How life got complicated", Discover Magazine, December 2012.
Ragan MA, Goggin CL, Cawthorn RJ, et al. (October 1996). "A novel clade of protistan parasites near the animal-fungal divergence". Proc. Natl. Acad. Sci. U.S.A. 93 (21): 11907–12. Bibcode:1996PNAS...9311907R. doi:10.1073/pnas.93.21.11907. PMC 38157. PMID 8876236.
Herr RA, Ajello L, Taylor JW, Arseculeratne SN, Mendoza L (September 1999). "Phylogenetic Analysis of Rhinosporidium seeberi's 18S Small-Subunit Ribosomal DNA Groups This Pathogen among Members of the Protoctistan Mesomycetozoa Clade". J. Clin. Microbiol. 37 (9): 2750–4. doi:10.1128/JCM.37.9.2750-2754.1999. PMC 85368. PMID 10449446.
Mendoza L, Taylor JW, Ajello L (October 2002). "The class mesomycetozoea: a heterogeneous group of microorganisms at the animal-fungal boundary". Annu. Rev. Microbiol. 56: 315–44. doi:10.1146/annurev.micro.56.012302.160950. PMID 12142489.
Borteiro, Claudio; Baldo, Diego; Maronna, Maximiliano Manuel; Ubilla (2018). "Amphibian parasites of the Order Dermocystida (Ichthyosporea): current knowledge, taxonomic review and new records from Brazil". Zootaxa. 4461 (4): 499–518. doi:10.11646/zootaxa.4461.4.3. PMID 30314064.
Reynolds, Nicole K.; Smith, Matthew E.; Tretter, Eric D.; Gause, Justin; Heeney, Dustin; Cafaro, Matías J.; Smith, James F.; Novak, Stephen J.; Bourland, William A.; White, Merlin M. (2017). "Resolving relationships at the animal-fungal divergence: A molecular phylogenetic study of the protist trichomycetes (Ichthyosporea, Eccrinida)". Molecular Phylogenetics and Evolution. 109: 447–464. doi:10.1016/j.ympev.2017.02.007. PMID 28219758.
Cavalier-Smith (May 2012). "Early evolution of eukaryote feeding modes, cell structural diversity, and classification of the protozoan phyla Loukozoa, Sulcozoa, and Choanozoa". European Journal of Protistology. 49 (2): 115–178. doi:10.1016/j.ejop.2012.06.001. PMID 23085100.
Crous PW, Gams W, Stalpers JA, Cannon PF, Kirk PM, David JC, Triebel D (November 2004). "An online database of names and descriptions as an alternative to registration". Mycological Research. 108 (11): 1236–1238. doi:10.1017/S0953756204221554.

"Eccrinida". Species Fungorum. 2016.

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[1] Cavalier-Smith, T. 1998. Neomonada and the origin of animals and fungi. In: Coombs GH, Vickerman K, Sleigh MA, Warren A (ed.) Evolutionary relationships among protozoa. Kluwer, London, pp. 375-407,

[tabrizi-2] Shalchian-Tabrizi K., Minge M.A., Espelund M.; et al. (7 May 2008). Aramayo, Rodolfo (ed.). "Multigene Phylogeny of Choanozoa and the Origin of Animals". PLOS ONE. 3 (5): e2098. Bibcode:2008PLoSO...3.2098S. doi:10.1371/journal.pone.0002098. PMC 2346548. PMID 18461162.{{cite journal}}: CS1 maint: multiple names: authors list (link)

[Ancestor-3] Dawkins, Richard; Wong, Yan (2016). The Ancestor's Tale. ISBN 978-0544859937.

[4] Douglas Fox, "How life got complicated", Discover Magazine, December 2012.

[pmid8876236-5] Ragan MA, Goggin CL, Cawthorn RJ, et al. (October 1996). "A novel clade of protistan parasites near the animal-fungal divergence". Proc. Natl. Acad. Sci. U.S.A. 93 (21): 11907–12. Bibcode:1996PNAS...9311907R. doi:10.1073/pnas.93.21.11907. PMC 38157. PMID 8876236.

[pmid10449446-6] Herr RA, Ajello L, Taylor JW, Arseculeratne SN, Mendoza L (September 1999). "Phylogenetic Analysis of Rhinosporidium seeberi's 18S Small-Subunit Ribosomal DNA Groups This Pathogen among Members of the Protoctistan Mesomycetozoa Clade". J. Clin. Microbiol. 37 (9): 2750–4. doi:10.1128/JCM.37.9.2750-2754.1999. PMC 85368. PMID 10449446.

[pmid12142489-7] Mendoza L, Taylor JW, Ajello L (October 2002). "The class mesomycetozoea: a heterogeneous group of microorganisms at the animal-fungal boundary". Annu. Rev. Microbiol. 56: 315–44. doi:10.1146/annurev.micro.56.012302.160950. PMID 12142489.

[8] Borteiro, Claudio; Baldo, Diego; Maronna, Maximiliano Manuel; Ubilla (2018). "Amphibian parasites of the Order Dermocystida (Ichthyosporea): current knowledge, taxonomic review and new records from Brazil". Zootaxa. 4461 (4): 499–518. doi:10.11646/zootaxa.4461.4.3. PMID 30314064.

[9] Reynolds, Nicole K.; Smith, Matthew E.; Tretter, Eric D.; Gause, Justin; Heeney, Dustin; Cafaro, Matías J.; Smith, James F.; Novak, Stephen J.; Bourland, William A.; White, Merlin M. (2017). "Resolving relationships at the animal-fungal divergence: A molecular phylogenetic study of the protist trichomycetes (Ichthyosporea, Eccrinida)". Molecular Phylogenetics and Evolution. 109: 447–464. doi:10.1016/j.ympev.2017.02.007. PMID 28219758.

[10] Cavalier-Smith (May 2012). "Early evolution of eukaryote feeding modes, cell structural diversity, and classification of the protozoan phyla Loukozoa, Sulcozoa, and Choanozoa". European Journal of Protistology. 49 (2): 115–178. doi:10.1016/j.ejop.2012.06.001. PMID 23085100.

[11] Crous PW, Gams W, Stalpers JA, Cannon PF, Kirk PM, David JC, Triebel D (November 2004). "An online database of names and descriptions as an alternative to registration". Mycological Research. 108 (11): 1236–1238. doi:10.1017/S0953756204221554.