Metamonad: Difference between revisions

Metamonad
	Giardia lamblia, a parasitic diplomonad
Scientific classification
Domain:	Eukaryota
(unranked):	Excavata
Phylum:	Metamonada; Grassé 1952 emend. Cavalier-Smith 2003
Classes & orders
	Anaeramoebidae; Parabasalia Trichonymphea; Trichomonadea; ; Anaeromonada Anaeromonadea; ; Fornicata Carpediemonadida; Retortamonadida; Diplomonadida; ;
Synonyms
	?Archezoa; Centrosomea Chatton Villeneuve 1937; Metamonadina Grassé 1952; Polymastigota Butschli 1884; Tetramastigota Hulsmann & Hausmann 1994;

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Revision as of 21:23, 11 February 2024

The metamonads are a large group of flagellate amitochondriate microscopic eukaryotes. Their composition is not entirely settled, but they include the retortamonads, diplomonads, and possibly the parabasalids and oxymonads as well. These four groups are all anaerobic (many being aerotolerant anaerobes), occurring mostly as symbiotes or parasites of animals, as is the case with Giardia lamblia which causes diarrhea in mammals.^[2]

Characteristics

A number of parabasalids and oxymonads are found in termite guts, and play an important role in breaking down the cellulose found in wood. Some other metamonads are parasites.

These flagellates are unusual in lacking mitochondria. Originally they were considered among the most primitive eukaryotes, diverging from the others before mitochondria appeared. However, they are now known to have lost mitochondria secondarily, and retain both organelles and nuclear genes derived from them. Mitochondrial relics include hydrogenosomes, which produce hydrogen, and small structures called mitosomes.

It now appears the Metamonada are, together with Malawimonas, sister clades of the Podiata.^[3]

All of these groups are united by having flagella or basal bodies in characteristic groups of four, which are often associated with the nucleus, forming a structure called a karyomastigont. In addition, the genera Carpediemonas and Trimastix are now known to be close relatives of the retortamonad-diplomonad line and the oxymonads, respectively. Both are free-living and amitochondriate.

Classification

The metamonads were thought to make up part of the Excavata, a paraphyletic eukaryotic supergroup including flagellates with feeding grooves and their close relatives. Their relationships are uncertain,^[4] and they do not always appear together on molecular trees. It is possible that the metamonads as defined here do not form a monophyletic subgroup.

The following higher level treatment from 2013 is based on works of Cavalier-Smith^[5] with amendments within Fornicata from Yubukia, Simpson & Leander.^[6]

Metamonada were once again proposed to be basal eukaryotes in 2018.^[7]

Phylum Metamonada (Grassé 1952) Cavalier-Smith 1987 emend. Cavalier-Smith 2003
- Family Anaeramoebidae Táborský, Pánek & Čepička 2017
- Subphylum Anaeromonada Cavalier-Smith 1997 emend. 2003
  - Class Anaeromonadea Cavalier-Smith 1997 emend. 1999
    - Family Paratrimastigidae Zhang et al. 2015^[8]
    - Order Trimastigida Cavalier-Smith 2003
      - Family Trimastigidae Saville Kent 1880
    - Order Oxymonadida Grassé 1952 emend. Cavalier-Smith 2003
      - Family Polymastigidae Bütschli 1884
      - Family Saccinobaculidae Brugerolle & Lee 2002 ex Cavalier-Smith 2013
      - Family Pyrsonymphidae Grassé 1892
      - Family Oxymonadidae Kirby 1928
- Subphylum Trichozoa Cavalier-Smith 1996 emend. Cavalier-Smith 2003 stat. n. 2013
  - Superclass Fornicata Simpson 2003 stat. n. Cavalier-Smith 2013
    - Family Kipferliidae Cavalier-Smith 2013
    - Class Carpediemonadea Cavalier-Smith 2013 s.s.
      - Order Carpediemonadida Cavalier-Smith 2003 emend. 2013 s.s.
        Family Carpediemonadidae Cavalier-Smith 2003
    - Class Eopharyngea Cavalier-Smith 1993 stat. n. Cavalier-Smith 2003
      - Order Dysnectida Cavalier-Smith 2013
        Family Dysnectidae Cavalier-Smith 2013
      - Order Retortamonadida Grassé 1952 emend. Cavalier-Smith 2013
        Family Caviomonadidae Cavalier-Smith 2013
        
        Family Chilomastigidae Cavalier-Smith 2013
        
        Family Retortamonadidae Wenrich 1932
      - Order Diplomonadida Wenyon 1926 emend. Brugerolle et al. 1975
        Family Giardiidae Kulda & Nohy´nkova´ 1978
        
        Family Octomitidae Cavalier-Smith 1996
        
        Family Spironucleidae Cavalier-Smith 1996
        
        Family Hexamitidae Kent 1880 emend. Brugerolle et al. 1975
  - Superclass Parabasalia Honigberg 1973 stat. n. Cavalier-Smith 2003
    - Class Trichonymphea Cavalier-Smith 2003
      - Order Lophomonadida Light 1927
        Family Lophomonadidae Saville Kent 1880
      - Order Trichonymphida Poche 1913
        Family †Burmanymphidae Poinar 2009
        
        Family Retractinymphidae Radek & Brune 2023
        
        Family Spirotrichosomidae Hollande & Caruette-Valentin 1971
        
        Family Staurojoeninidae Grassé 1917
        
        Family Trichonymphidae Saville Kent 1880
        
        Family Hoplonymphidae Light 1926
        
        Family Teratonymphidae Koidzumi 1921 [Eucomonymphidae]
    - Class Trichomonadea Kirby 1947 stat. n. Cavalier-Smith 2003
      - Order Pimpavicka Céza & Čepička 2022
        Family Pimpavickidae Céza & Čepička 2022
      - Order Trichomonadida Kirby 1947
        Suborder Trichomonadina (Kirby 1947) Cavalier-Smith 2013
        Family Trichomonadidae Chalmers & Pekkoloa 1918 sensu Hampl et al. 2006
        
        Suborder Honigbergiellina Čepička et al. 2010
        Family Honigbergiellidae Čepička, Hampl & Kulda 2010
        
        Family Hexamastigidae Čepička, Hampl & Kulda 2010
        
        Family Tricercomitidae Čepička, Hampl & Kulda 2010
      - Order Hypotrichomonadida Čepička et al. 2010
        Family Hypotrichomonadidae (Honigberg 1963) Čepička, Hampl & Kulda 2010
      - Order Spirotrichonymphida Grassé 1952
        Family Spirotrichonymphidae Grassé 1917
      - Order Tritrichomonadida Čepička et al. 2010
        Family Dientamoebidae Grassé 1953
        
        Family Monocercomonadidae Kirby 1944
        
        Family Simplicimonadidae Čepička et al. 2010
        
        Family Tritrichomonadidae Honigberg 1963
      - Order Cristamonadida Brugerolle & Patterson 2001 emend. Cavalier-Smith 2013
        Family Calonymphidae Grassé 1911
        
        Family Devescovinidae Doflein 1911

References

^ Stairs, Courtney W.; Táborský, Petr; Salomaki, Eric D.; Kolisko, Martin; Pánek, Tomáš; Eme, Laura; Hradilová, Miluše; Vlček, Čestmír; Jerlström-Hultqvist, Jon; Roger, Andrew J.; Čepička, Ivan (2021-12-20). "Anaeramoebae are a divergent lineage of eukaryotes that shed light on the transition from anaerobic mitochondria to hydrogenosomes". Current Biology. 31 (24): 5605–5612.e5. doi:10.1016/j.cub.2021.10.010. ISSN 0960-9822. PMID 34710348. S2CID 240054026.
^ ^a ^b Al Jewari, Caesar; Baldauf, Sandra L. (2023-04-28). "An excavate root for the eukaryote tree of life". Science Advances. 9 (17): eade4973. doi:10.1126/sciadv.ade4973. ISSN 2375-2548. PMC 10146883. PMID 37115919.
^ Cavalier-Smith, Thomas; Chao, Ema E.; Lewis, Rhodri (2016-06-01). "187-gene phylogeny of protozoan phylum Amoebozoa reveals a new class (Cutosea) of deep-branching, ultrastructurally unique, enveloped marine Lobosa and clarifies amoeba evolution". Molecular Phylogenetics and Evolution. 99: 275–296. doi:10.1016/j.ympev.2016.03.023. PMID 27001604.
^ Cavalier-Smith T (November 2003). "The excavate protozoan phyla Metamonada Grassé emend. (Anaeromonadea, Parabasalia, Carpediemonas, Eopharyngia) and Loukozoa emend. (Jakobea, Malawimonas): their evolutionary affinities and new higher taxa". Int. J. Syst. Evol. Microbiol. 53 (Pt 6): 1741–58. doi:10.1099/ijs.0.02548-0. PMID 14657102.
^ Cavalier-Smith T (2013). "Early evolution of eukaryote feeding modes, cell structural diversity, and classification of the protozoan phyla Loukozoa, Sulcozoa, and Choanozoa". Eur. J. Protistol. 49 (2): 115–178. doi:10.1016/j.ejop.2012.06.001. PMID 23085100.
^ Yubukia; Simpson; Leander (2013). "Comprehensive Ultrastructure of Kipferlia bialata Provides Evidence for Character Evolution within the Fornicata (Excavata)". Protist. 164 (3): 423–439. doi:10.1016/j.protis.2013.02.002. PMID 23517666.
^ Krishnan, Arunkumar; Burroughs, A. Max; Iyer, Lakshminarayan; Aravind, L. (2018-07-04). "The unexpected provenance of components in eukaryotic nucleotide-excision-repair and kinetoplast DNA-dynamics from bacterial mobile elements". bioRxiv: 361121. doi:10.1101/361121.
^ Zhang, Qianqian; Táborský, Petr; Silberman, Jeffrey D.; Pánek, Tomáš; Čepička, Ivan; Simpson, Alastair G.B. (2015). "Marine Isolates of Trimastix marina Form a Plesiomorphic Deep-branching Lineage within Preaxostyla, Separate from Other Known Trimastigids (Paratrimastix n. gen.)". Protist. 166 (4): 468–491. doi:10.1016/j.protis.2015.07.003.

External links

Tree of Life: Fornicata

[1] Stairs, Courtney W.; Táborský, Petr; Salomaki, Eric D.; Kolisko, Martin; Pánek, Tomáš; Eme, Laura; Hradilová, Miluše; Vlček, Čestmír; Jerlström-Hultqvist, Jon; Roger, Andrew J.; Čepička, Ivan (2021-12-20). "Anaeramoebae are a divergent lineage of eukaryotes that shed light on the transition from anaerobic mitochondria to hydrogenosomes". Current Biology. 31 (24): 5605–5612.e5. doi:10.1016/j.cub.2021.10.010. ISSN 0960-9822. PMID 34710348. S2CID 240054026.

[Al_Jewari_Baldauf_2023-2] Al Jewari, Caesar; Baldauf, Sandra L. (2023-04-28). "An excavate root for the eukaryote tree of life". Science Advances. 9 (17): eade4973. doi:10.1126/sciadv.ade4973. ISSN 2375-2548. PMC 10146883. PMID 37115919.

[3] Cavalier-Smith, Thomas; Chao, Ema E.; Lewis, Rhodri (2016-06-01). "187-gene phylogeny of protozoan phylum Amoebozoa reveals a new class (Cutosea) of deep-branching, ultrastructurally unique, enveloped marine Lobosa and clarifies amoeba evolution". Molecular Phylogenetics and Evolution. 99: 275–296. doi:10.1016/j.ympev.2016.03.023. PMID 27001604.

[pmid14657102-4] Cavalier-Smith T (November 2003). "The excavate protozoan phyla Metamonada Grassé emend. (Anaeromonadea, Parabasalia, Carpediemonas, Eopharyngia) and Loukozoa emend. (Jakobea, Malawimonas): their evolutionary affinities and new higher taxa". Int. J. Syst. Evol. Microbiol. 53 (Pt 6): 1741–58. doi:10.1099/ijs.0.02548-0. PMID 14657102.

[5] Cavalier-Smith T (2013). "Early evolution of eukaryote feeding modes, cell structural diversity, and classification of the protozoan phyla Loukozoa, Sulcozoa, and Choanozoa". Eur. J. Protistol. 49 (2): 115–178. doi:10.1016/j.ejop.2012.06.001. PMID 23085100.

[6] Yubukia; Simpson; Leander (2013). "Comprehensive Ultrastructure of Kipferlia bialata Provides Evidence for Character Evolution within the Fornicata (Excavata)". Protist. 164 (3): 423–439. doi:10.1016/j.protis.2013.02.002. PMID 23517666.

[7] Krishnan, Arunkumar; Burroughs, A. Max; Iyer, Lakshminarayan; Aravind, L. (2018-07-04). "The unexpected provenance of components in eukaryotic nucleotide-excision-repair and kinetoplast DNA-dynamics from bacterial mobile elements". bioRxiv: 361121. doi:10.1101/361121.

[8] Zhang, Qianqian; Táborský, Petr; Silberman, Jeffrey D.; Pánek, Tomáš; Čepička, Ivan; Simpson, Alastair G.B. (2015). "Marine Isolates of Trimastix marina Form a Plesiomorphic Deep-branching Lineage within Preaxostyla, Separate from Other Known Trimastigids (Paratrimastix n. gen.)". Protist. 166 (4): 468–491. doi:10.1016/j.protis.2015.07.003.

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[2]

[3]

[4]

[5]

[6]

[7]

[8]

@@ Line 8: / Line 8: @@
 | authority = [[Grassé]] 1952 ''[[emend.]]'' [[Cavalier-Smith]] 2003
 | display_parents = 2
-| synonyms = [[Archezoa]]?
-| synonyms_ref = <ref name="Al Jewari Baldauf 2023"/>
 | subdivision_ranks = Classes & orders
+| subdivision =
-| subdivision = * [[Anaeramoeba]]<ref>{{Cite journal |last1=Stairs |first1=Courtney W. |last2=Táborský |first2=Petr |last3=Salomaki |first3=Eric D. |last4=Kolisko |first4=Martin |last5=Pánek |first5=Tomáš |last6=Eme |first6=Laura |last7=Hradilová |first7=Miluše |last8=Vlček |first8=Čestmír |last9=Jerlström-Hultqvist |first9=Jon |last10=Roger |first10=Andrew J. |last11=Čepička |first11=Ivan |date=2021-12-20 |title=Anaeramoebae are a divergent lineage of eukaryotes that shed light on the transition from anaerobic mitochondria to hydrogenosomes |journal=Current Biology |language=English |volume=31 |issue=24 |pages=5605–5612.e5 |doi=10.1016/j.cub.2021.10.010 |issn=0960-9822 |pmid=34710348|s2cid=240054026 |doi-access=free }}</ref>
+* [[Anaeramoebidae]]<ref>{{Cite journal |last1=Stairs |first1=Courtney W. |last2=Táborský |first2=Petr |last3=Salomaki |first3=Eric D. |last4=Kolisko |first4=Martin |last5=Pánek |first5=Tomáš |last6=Eme |first6=Laura |last7=Hradilová |first7=Miluše |last8=Vlček |first8=Čestmír |last9=Jerlström-Hultqvist |first9=Jon |last10=Roger |first10=Andrew J. |last11=Čepička |first11=Ivan |date=2021-12-20 |title=Anaeramoebae are a divergent lineage of eukaryotes that shed light on the transition from anaerobic mitochondria to hydrogenosomes |journal=Current Biology |language=English |volume=31 |issue=24 |pages=5605–5612.e5 |doi=10.1016/j.cub.2021.10.010 |issn=0960-9822 |pmid=34710348|s2cid=240054026 |doi-access=free }}</ref>
 * [[Parabasalid|Parabasalia]]
 ** [[Trichonymphea]]
@@ Line 19: / Line 18: @@
 * [[Fornicata]]
 ** [[Carpediemonas|Carpediemonadida]]
-** [[Chilomastix|Chilomastigida]]
 ** [[Retortamonad]]ida
 ** [[Diplomonad]]ida
+| synonyms_ref = <ref name="Al Jewari Baldauf 2023"/>
+| synonyms =
+* ?[[Archezoa]]
+* Centrosomea <small>Chatton Villeneuve 1937</small>
+* Metamonadina <small>Grassé 1952</small>
+* Polymastigota <small>Butschli 1884</small>
+* Tetramastigota <small>Hulsmann & Hausmann 1994</small>
 }}
@@ Line 43: / Line 48: @@
 * Phylum '''Metamonada''' <small>(Grassé 1952) Cavalier-Smith 1987 emend. Cavalier-Smith 2003</small>
+** Family [[Anaeramoebidae]] <small>Táborský, Pánek & Čepička 2017</small>
 ** Subphylum [[Anaeromonada]] <small>Cavalier-Smith 1997 emend. 2003</small>
 *** Class [[Anaeromonadea]] <small>Cavalier-Smith 1997 emend. 1999</small>
+**** Family [[Paratrimastigidae]] <small>Zhang et al. 2015</small><ref>{{Cite journal |last1=Zhang |first1=Qianqian |last2=Táborský |first2=Petr |last3=Silberman |first3=Jeffrey D. |last4=Pánek |first4=Tomáš |last5=Čepička |first5=Ivan |last6=Simpson|first6=Alastair G.B. |year=2015 |title=Marine Isolates of Trimastix marina Form a Plesiomorphic Deep-branching Lineage within Preaxostyla, Separate from Other Known Trimastigids (Paratrimastix n. gen.) |url=https://www.sciencedirect.com/science/article/abs/pii/S1434461015000462 |journal=Protist |volume=166 |issue=4 |pages=468-491 |doi=10.1016/j.protis.2015.07.003 |doi-access=}}</ref>
-**** Family [[Paratrimastigidae]] <small>Zhang et al. 2015</small>
 **** Order [[Trimastix|Trimastigida]] <small>Cavalier-Smith 2003</small>
 ***** Family [[Trimastix|Trimastigidae]] <small>Saville Kent 1880</small>
@@ Line 52: / Line 58: @@
 ***** Family [[Saccinobaculidae]] <small>Brugerolle & Lee 2002 ex Cavalier-Smith 2013</small>
 ***** Family [[Pyrsonymphidae]] <small>Grassé 1892</small>
-***** Family [[Streblomastigidae]] <small>Kofoid & Swezy 1919</small>
 ***** Family [[Oxymonadidae]] <small>Kirby 1928</small>
 ** Subphylum [[Trichozoa]] <small>Cavalier-Smith 1996 emend. Cavalier-Smith 2003 stat. n. 2013</small>
 *** Superclass [[Fornicata]] <small>Simpson 2003 stat. n. Cavalier-Smith 2013</small>
-**** Genus ''[[Ergobibamus]]''
-**** Genus ''[[Hicanonectes]]''
 **** Family [[Kipferliidae]] <small>Cavalier-Smith 2013</small>
-**** Order [[Chilomastigida]] <small>Cavalier-Smith 2013</small>
-***** Family [[Chilomastigidae]] <small>Cavalier-Smith 2013</small>
-**** Order [[Dysnectida]] <small>Cavalier-Smith 2013</small>
-***** Family [[Dysnectidae]] <small>Cavalier-Smith 2013</small>
 **** Class [[Carpediemonadea]] <small>Cavalier-Smith 2013 s.s.</small>
 ***** Order [[Carpediemonadida]] <small>Cavalier-Smith 2003 emend. 2013 s.s.</small>
 ****** Family [[Carpediemonadidae]] <small>Cavalier-Smith 2003</small>
 **** Class [[Eopharyngea]] <small>Cavalier-Smith 1993 stat. n. Cavalier-Smith 2003</small>
+***** Order [[Dysnectida]] <small>Cavalier-Smith 2013</small>
+****** Family [[Dysnectidae]] <small>Cavalier-Smith 2013</small>
 ***** Order [[Retortamonadida]] <small>Grassé 1952 emend. Cavalier-Smith 2013</small>
+****** Family [[Caviomonadidae]] <small>Cavalier-Smith 2013</small>
+****** Family [[Chilomastix|Chilomastigidae]] <small>Cavalier-Smith 2013</small>
 ****** Family [[Retortamonadidae]] <small>Wenrich 1932</small>
 ***** Order [[Diplomonadida]] <small>Wenyon 1926 emend. Brugerolle et al. 1975</small>
@@ Line 75: / Line 78: @@
 ****** Family [[Hexamitidae]] <small>Kent 1880 emend. Brugerolle et al. 1975</small>
 *** Superclass [[Parabasalia]] <small>Honigberg 1973 stat. n. Cavalier-Smith 2003</small>
-**** Class [[Chilomitea]]
-***** Order [[Chilomitida]]
-****** Family [[Chilomitidae]]
 **** Class [[Trichonymphea]] <small>Cavalier-Smith 2003</small>
 ***** Order [[Lophomonadida]] <small>Light 1927</small>
 ****** Family [[Lophomonadidae]] <small>Saville Kent 1880</small>
 ***** Order [[Trichonymphida]] <small>Poche 1913</small>
+****** Family †[[Burmanymphidae]] <small>Poinar 2009</small>
+****** Family [[Retractinymphidae]] <small>Radek & Brune 2023</small>
 ****** Family [[Spirotrichosomidae]] <small>Hollande & Caruette-Valentin 1971</small>
 ****** Family [[Staurojoeninidae]] <small>Grassé 1917</small>
 ****** Family [[Trichonymphidae]] <small>Saville Kent 1880</small>
 ****** Family [[Hoplonymphidae]] <small>Light 1926</small>
-****** Family [[Teranymphidae]] <small>Koidzumi 1921</small> [Eucomonymphidae]
+****** Family [[Teratonymphidae]] <small>Koidzumi 1921</small> [Eucomonymphidae]
 **** Class [[Trichomonadea]] <small>Kirby 1947 stat. n. Cavalier-Smith 2003</small>
-***** Order [[Trichocovinida]]
+***** Order [[Pimpavicka]] <small>Céza & Čepička 2022</small>
-****** Family [[Trichocovinidae]]
+****** Family [[Pimpavickidae]] <small>Céza & Čepička 2022</small>
 ***** Order [[Trichomonadida]] <small>Kirby 1947</small>
-****** Suborder [[Hypotrichomonadina]] <small>Čepička et al. 2010 stat. n.</small>
-******* Family [[Hypotrichomonadidae]] <small>(Honigberg 1963) Čepička, Hampl & Kulda 2010 </small>
 ****** Suborder [[Trichomonadina]] <small>(Kirby 1947) Cavalier-Smith 2013</small>
 ******* Family [[Trichomonadidae]] <small>Chalmers & Pekkoloa 1918 sensu Hampl et al. 2006</small>
@@ Line 99: / Line 99: @@
 ******* Family [[Hexamastigidae]] <small>Čepička, Hampl & Kulda 2010</small>
 ******* Family [[Tricercomitidae]] <small>Čepička, Hampl & Kulda 2010</small>
+***** Order [[Hypotrichomonadida]] <small>Čepička et al. 2010</small>
+****** Family [[Hypotrichomonadidae]] <small>(Honigberg 1963) Čepička, Hampl & Kulda 2010 </small>
+***** Order [[Spirotrichonymphida]] <small>Grassé 1952</small>
+****** Family [[Spirotrichonymphidae]] <small>Grassé 1917</small>
 ***** Order [[Tritrichomonadida]] <small>Čepička et al. 2010</small>
 ****** Family [[Dientamoebidae]] <small>Grassé 1953</small>
@@ Line 104: / Line 108: @@
 ****** Family [[Simplicimonadidae]] <small>Čepička et al. 2010</small>
 ****** Family [[Tritrichomonadidae]] <small>Honigberg 1963</small>
-***** Subclass [[Cristomonadia]] <small>Cavalier-Smith, 2013</small>
+***** Order [[Cristamonadida]] <small>Brugerolle & Patterson 2001 emend. Cavalier-Smith 2013</small>
-****** Order [[Spirotrichonymphida]] <small>Grassé 1952</small>
+****** Family [[Calonymphidae]] <small>Grassé 1911</small>
-******* Family [[Spirotrichonymphidae]] <small>Grassé 1917</small>
+****** Family [[Devescovinidae]] <small>Doflein 1911</small>
-****** Order [[Cristamonadida]] <small>Brugerolle & Patterson 2001 emend. Cavalier-Smith 2013</small>
-******* Family [[Calonymphidae]] <small>Grassé 1911</small>
-******* Family [[Devescovinidae]] <small>Doflein 1911</small> [Joeniidae; Rhizonymphidae; Kofoidiidae; Deltotrichonymphidae]
 == References ==