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Excavata

February 15, 2024

For an explanation of very similar terms, see Eukaryota.

Excavata is an extensive and diverse but paraphyletic group of unicellular Eukaryota.[1][2] The group was first suggested by Simpson and Patterson in 1999[3][4] and the name latinized and assigned a rank by Thomas Cavalier-Smith in 2002. It contains a variety of free-living and symbiotic protists, and includes some important parasites of humans such as Giardia and Trichomonas.[5] Excavates were formerly considered to be included in the now obsolete Protista kingdom.[6] They were distinguished from other lineages based on electron-microscopic information about how the cells are arranged (they have a distinctive ultrastructural identity).[4] They are considered to be a basal flagellate lineage.[7]

Excavata
Temporal range: Neoproterozoic–present
Giardia lamblia, a parasitic diplomonad
Scientific classification
(obsolete as paraphyletic)
Domain: Eukaryota
(unranked): Excavata
(Cavalier-Smith), 2002
Phyla and classes

see text

Three types of excavate cells. Top: Jakobida, 1-nucleus, 2-anterior flagellum, 3-ventral/posterior flagellum, 4-ventral feeding groove. Middle: Euglenozoa, 1-nucleus, 2-flagellar pocket/reservoir, 3-dorsal/anterior flagellum, 4-ventral/posterior flagellum, 5-cytostome/feeding apparatus. Bottom: Metamonada, 1-anterior flagella, 2-parabasal body, 3-undulating membrane, 4-posterior flagellum, 5-nucleus, 6-axostyle.

On the basis of phylogenomic analyses, the group was shown to contain three widely separated eukaryote groups, the discobids, metamonads, and malawimonads.[8][9][10][11] A current view of the composition of the excavates is given below, indicating that the group is paraphyletic. Except for some Euglenozoa, all are non-photosynthetic.

Characteristics edit

Most excavates are unicellular, heterotrophic flagellates. Only some Euglenozoa are photosynthetic. In some (particularly anaerobic intestinal parasites), the mitochondria have been greatly reduced.[5] Some excavates lack "classical" mitochondria, and are called "amitochondriate", although most retain a mitochondrial organelle in greatly modified form (e.g. a hydrogenosome or mitosome). Among those with mitochondria, the mitochondrial cristae may be tubular, discoidal, or in some cases, laminar. Most excavates have two, four, or more flagella.[4] Many have a conspicuous ventral feeding groove with a characteristic ultrastructure, supported by microtubules—the "excavated" appearance of this groove giving the organisms their name.[3][6] However, various groups that lack these traits are considered to be derived excavates based on genetic evidence (primarily phylogenetic trees of molecular sequences).[6]

The Acrasidae slime molds are the only excavates to exhibit limited multicellularity. Like other cellular slime molds, they live most of their life as single cells, but will sometimes assemble into larger clusters.

Proposed group edit

See also: Eukaryote § Phylogeny, and wikispecies:Excavata

Excavate relationships were always uncertain, suggesting that they are not a monophyletic group.[12] Phylogenetic analyses often do not place malawimonads on the same branch as the other Excavata.[13]

Excavates were thought to include multiple groups:

Kingdom/Superphylum Included taxa Representative genera (examples) Description
Discoba or JEH or Eozoa Tsukubea Tsukubamonas
Euglenozoa Euglena, Trypanosoma Many important parasites, one large group with plastids (chloroplasts)
Heterolobosea (Percolozoa) Naegleria, Acrasis Most alternate between flagellate and amoeboid forms
Jakobea Jakoba, Reclinomonas Free-living, sometimes loricate flagellates, with very gene-rich mitochondrial genomes
Metamonada or POD Preaxostyla Oxymonads, Trimastix Amitochondriate flagellates, either free-living (Trimastix, Paratrimastix) or living in the hindguts of insects
Fornicata Giardia, Carpediemonas Amitochondriate, mostly symbiotes and parasites of animals.
Parabasalia Trichomonas Amitochondriate flagellates, generally intestinal commensals of insects. Some human pathogens.
Anaeramoeba Anaeramoeba ignava Anaerobic protists with hydrogenosomes instead of mitochondria.
Neolouka Malawimonadida Malawimonas

Discoba or JEH clade edit

Euglenozoa and Heterolobosea (Percolozoa) or Eozoa (as named by Cavalier-Smith[14]) appear to be particularly close relatives, and are united by the presence of discoid cristae within the mitochondria (Superphylum Discicristata). A close relationship has been shown between Discicristata and Jakobida,[15] the latter having tubular cristae like most other protists, and hence were united under the taxon name Discoba, which was proposed for this apparently monophyletic group.[1]

Metamonads edit

Metamonads are unusual in not having classical mitochondria—instead they have hydrogenosomes, mitosomes or uncharacterised organelles. The oxymonad Monocercomonoides is reported to have completely lost homologous organelles. There are competing explanations.[16][17]

Malawimonads edit

The malawimonads have been proposed to be members of Excavata owing to their typical excavate morphology, and phylogenetic affinity to other excavate groups in some molecular phylogenies. However, their position among eukaryotes remains elusive.[2]

Ancyromonads edit

Ancyromonads are small free-living cells with a narrow longitudinal groove down one side of the cell. The ancyromonad groove is not used for "suspension feeding", unlike in "typical excavates" (e.g. malawimonads, jakobids, Trimastix, Carpediemonas, Kiperferlia, etc). Ancyromonads instead capture prokaryotes attached to surfaces. The phylogenetic placement of ancyromonads is poorly understood (in 2020), however some phylogenetic analyses place them as close relatives of malawimonads.[9]

Evolution edit

Origin of the Eukaryotes edit

Further information: Eukaryogenesis

The conventional explanation for the origin of the Eukaryotes is that a heimdallarchaeian or another Archaea acquired an alphaproteobacterium as an endosymbiont, and that this became the mitochondrion, the organelle providing oxidative respiration to the eukaryotic cell.[18]

Caesar al Jewari and Sandra Baldauf argue instead that the Eukaryotes possibly started with an endosymbiosis event of a Deltaproteobacterium or Aammaproteobacterium, accounting for the otherwise unexplained presence of anaerobic bacterial enzymes in Metamonada. The sister of the Preaxostyla within Metamonada represents the rest of the Eukaryotes which acquired an Alphaproteobacterium. In their scenario, the hydrogenosome and mitosome, both conventionally considered "mitochondrion-derived organelles", would predate the mitochondrion, and instead be derived from the earlier symbiotic bacterium.[17]

Phylogeny edit

In 2023, using molecular phylogenetic analysis of 186 taxa, Al Jewari and Baldauf proposed a phylogenetic tree with the metamonad Parabasalia as basal Eukaryotes. Discoba and the rest of the Eukaryota appear to have emerged as sister taxon to the Preaxostyla, incorporating a single alphaproteobacterium as mitochondria by endosymbiosis. Thus the Fornicata are more closely related to e.g. animals than to Parabasalia. The rest of the Eukaryotes emerged within the Excavata as sister of the Discoba; as they are within the same clade but are not cladistically considered part of the Excavata yet, the Excavata are in this analysis highly paraphyletic.[17]

The Anaeramoeba are associated with Parabasalia, but could turn out to be more basal as the root of a tree is often difficult to pinpoint.[19]

See also edit

Metakaryota

Gallery edit

References edit

  1. ^ a b Hampl, Vladimir; Hug, Laura; Leigh, Jessica W.; et al. (2009). "Phylogenomic analyses support the monophyly of Excavata and resolve relationships among eukaryotic "supergroups"". PNAS. 106 (10): 3859–3864. Bibcode:2009PNAS..106.3859H. doi:10.1073/pnas.0807880106. PMC 2656170. PMID 19237557.
  2. ^ a b Simpson, Alastair G. B.; Inagaki, Yuji; Roger, Andrew J. (2006). "Comprehensive multigene phylogenies of excavate protists reveal the evolutionary positions of "primitive" eukaryotes". Molecular Biology and Evolution. 23 (3): 615–625. doi:10.1093/molbev/msj068. PMID 16308337.
  3. ^ a b Simpson, Alastair G.B.; Patterson, David J. (December 1999). "The ultrastructure of Carpediemonas membranifera (Eukaryota) with reference to the 'excavate hypothesis'". European Journal of Protistology. 35 (4): 353–370. doi:10.1016/S0932-4739(99)80044-3.
  4. ^ a b c Simpson, Alastair G. B. (November 2003). "Cytoskeletal organization, phylogenetic affinities and systematics in the contentious taxon Excavata (Eukaryota)". International Journal of Systematic and Evolutionary Microbiology. 53 (6): 1759–1777. doi:10.1099/ijs.0.02578-0. PMID 14657103.
  5. ^ a b Dawkins, Richard; Wong, Yan (2016). The Ancestor's Tale. Houghton Mifflin Harcourt. ISBN 978-0544859937.
  6. ^ a b c Cavalier-Smith, Thomas (2002). "The phagotrophic origin of eukaryotes and phylogenetic classification of Protozoa". International Journal of Systematic and Evolutionary Microbiology. 52 (2): 297–354. doi:10.1099/00207713-52-2-297. PMID 11931142.
  7. ^ Dawson, Scott C.; Paredez, Alexander R. (2013). "Alternative cytoskeletal landscapes: cytoskeletal novelty and evolution in basal excavate protists". Current Opinion in Cell Biology. 25 (1): 134–141. doi:10.1016/j.ceb.2012.11.005. PMC 4927265. PMID 23312067.
  8. ^ Burki, Fabien; Roger, Andrew J.; Brown, Matthew W.; et al. (January 2020). "The New Tree of Eukaryotes". Trends in Ecology & Evolution. 35 (1): 43–55. doi:10.1016/j.tree.2019.08.008. PMID 31606140. S2CID 204545629.
  9. ^ a b Brown, Matthew W.; Heiss, Aaron A.; Kamikawa, Ryoma; et al. (2018-01-19). "Phylogenomics Places Orphan Protistan Lineages in a Novel Eukaryotic Super-Group". Genome Biology and Evolution. 10 (2): 427–433. doi:10.1093/gbe/evy014. PMC 5793813. PMID 29360967.
  10. ^ Heiss, Aaron A.; Kolisko, Martin; Ekelund, Fleming; et al. (4 April 2018). "Combined morphological and phylogenomic re-examination of malawimonads, a critical taxon for inferring the evolutionary history of eukaryotes". Royal Society Open Science. 5 (4): 171707. Bibcode:2018RSOS....571707H. doi:10.1098/rsos.171707. PMC 5936906. PMID 29765641.
  11. ^ Keeling, Patrick J.; Burki, Fabien (19 August 2019). "Progress towards the Tree of Eukaryotes". Current Biology. 29 (16): R808–R817. doi:10.1016/j.cub.2019.07.031. PMID 31430481.
  12. ^ Laura Wegener Parfrey; Erika Barbero; Elyse Lasser; Micah Dunthorn; Debashish Bhattacharya; David J Patterson; Laura A Katz (December 2006). "Evaluating support for the current classification of eukaryotic diversity". PLOS Genetics. 2 (12): e220. doi:10.1371/JOURNAL.PGEN.0020220. ISSN 1553-7390. PMC 1713255. PMID 17194223. Wikidata Q21090155.
  13. ^ Tice, Alexander K.; Žihala, David; Pánek, Tomáš; et al. (2021). "PhyloFisher: A phylogenomic package for resolving eukaryotic relationships". PLOS Biology. 19 (8): e3001365. doi:10.1371/journal.pbio.3001365. PMC 8345874. PMID 34358228.
  14. ^ Cavalier-Smith, Thomas (23 December 2009). "Kingdoms Protozoa and Chromista and the eozoan root of the eukaryotic tree". Biology Letters. The Royal Society. 6 (3): 342–345. doi:10.1098/rsbl.2009.0948. ISSN 1744-9561. PMC 2880060. PMID 20031978.
  15. ^ Rodríguez-Ezpeleta, Naiara; Brinkmann, Henner; Burger, Gertraud; et al. (2007). "Toward Resolving the Eukaryotic Tree: The Phylogenetic Positions of Jakobids and Cercozoans". Current Biology. 17 (16): 1420–1425. doi:10.1016/j.cub.2007.07.036. PMID 17689961.
  16. ^ Bui, Elisabeth T.; Bradley, Peter J.; Johnson, Patricia J. (3 September 1996). "A common evolutionary origin for mitochondria and hydrogenosomes". Proceedings of the National Academy of Sciences. 93 (18): 9651–9656. Bibcode:1996PNAS...93.9651B. doi:10.1073/pnas.93.18.9651. ISSN 0027-8424. PMC 38483. PMID 8790385.
  17. ^ a b c Al Jewari, Caesar; Baldauf, Sandra L. (2023-04-28). "An excavate root for the eukaryote tree of life". Science Advances. 9 (17): eade4973. Bibcode:2023SciA....9E4973A. doi:10.1126/sciadv.ade4973. ISSN 2375-2548. PMC 10146883. PMID 37115919.
  18. ^ a b Eme, Laura; Tamarit, Daniel; Caceres, Eva F.; et al. (2023-03-09). "Inference and reconstruction of the heimdallarchaeial ancestry of eukaryotes". Nature. 618 (7967): 992–999. doi:10.1101/2023.03.07.531504. PMC 10307638. PMID 37316666.
  19. ^ Stairs, Courtney W.; Táborský, Petr; Salomaki, Eric D.; et al. (December 2021). "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.

External links edit

  • Open Tree of Life
  • Taxonomicon
  • Tree of Life Eukaryotes
  • Tree of Life: Jakobida
  • Tree of Life: Fornicata

February 15, 2024
excavata, explanation, very, similar, terms, eukaryota, extensive, diverse, paraphyletic, group, unicellular, eukaryota, group, first, suggested, simpson, patterson, 1999, name, latinized, assigned, rank, thomas, cavalier, smith, 2002, contains, variety, free,. For an explanation of very similar terms see Eukaryota Excavata is an extensive and diverse but paraphyletic group of unicellular Eukaryota 1 2 The group was first suggested by Simpson and Patterson in 1999 3 4 and the name latinized and assigned a rank by Thomas Cavalier Smith in 2002 It contains a variety of free living and symbiotic protists and includes some important parasites of humans such as Giardia and Trichomonas 5 Excavates were formerly considered to be included in the now obsolete Protista kingdom 6 They were distinguished from other lineages based on electron microscopic information about how the cells are arranged they have a distinctive ultrastructural identity 4 They are considered to be a basal flagellate lineage 7 ExcavataTemporal range Neoproterozoic present Pha Proterozoic Archean Had Giardia lamblia a parasitic diplomonadScientific classification obsolete as paraphyletic Domain Eukaryota unranked Excavata Cavalier Smith 2002Phyla and classessee textThree types of excavate cells Top Jakobida 1 nucleus 2 anterior flagellum 3 ventral posterior flagellum 4 ventral feeding groove Middle Euglenozoa 1 nucleus 2 flagellar pocket reservoir 3 dorsal anterior flagellum 4 ventral posterior flagellum 5 cytostome feeding apparatus Bottom Metamonada 1 anterior flagella 2 parabasal body 3 undulating membrane 4 posterior flagellum 5 nucleus 6 axostyle On the basis of phylogenomic analyses the group was shown to contain three widely separated eukaryote groups the discobids metamonads and malawimonads 8 9 10 11 A current view of the composition of the excavates is given below indicating that the group is paraphyletic Except for some Euglenozoa all are non photosynthetic Contents 1 Characteristics 2 Proposed group 2 1 Discoba or JEH clade 2 2 Metamonads 2 3 Malawimonads 2 4 Ancyromonads 3 Evolution 3 1 Origin of the Eukaryotes 3 2 Phylogeny 4 See also 5 Gallery 6 References 7 External linksCharacteristics editMost excavates are unicellular heterotrophic flagellates Only some Euglenozoa are photosynthetic In some particularly anaerobic intestinal parasites the mitochondria have been greatly reduced 5 Some excavates lack classical mitochondria and are called amitochondriate although most retain a mitochondrial organelle in greatly modified form e g a hydrogenosome or mitosome Among those with mitochondria the mitochondrial cristae may be tubular discoidal or in some cases laminar Most excavates have two four or more flagella 4 Many have a conspicuous ventral feeding groove with a characteristic ultrastructure supported by microtubules the excavated appearance of this groove giving the organisms their name 3 6 However various groups that lack these traits are considered to be derived excavates based on genetic evidence primarily phylogenetic trees of molecular sequences 6 The Acrasidae slime molds are the only excavates to exhibit limited multicellularity Like other cellular slime molds they live most of their life as single cells but will sometimes assemble into larger clusters Proposed group editSee also Eukaryote Phylogeny and wikispecies Excavata Excavate relationships were always uncertain suggesting that they are not a monophyletic group 12 Phylogenetic analyses often do not place malawimonads on the same branch as the other Excavata 13 Excavates were thought to include multiple groups Kingdom Superphylum Included taxa Representative genera examples DescriptionDiscoba or JEH or Eozoa Tsukubea TsukubamonasEuglenozoa Euglena Trypanosoma Many important parasites one large group with plastids chloroplasts Heterolobosea Percolozoa Naegleria Acrasis Most alternate between flagellate and amoeboid formsJakobea Jakoba Reclinomonas Free living sometimes loricate flagellates with very gene rich mitochondrial genomesMetamonada or POD Preaxostyla Oxymonads Trimastix Amitochondriate flagellates either free living Trimastix Paratrimastix or living in the hindguts of insectsFornicata Giardia Carpediemonas Amitochondriate mostly symbiotes and parasites of animals Parabasalia Trichomonas Amitochondriate flagellates generally intestinal commensals of insects Some human pathogens Anaeramoeba Anaeramoeba ignava Anaerobic protists with hydrogenosomes instead of mitochondria Neolouka Malawimonadida MalawimonasDiscoba or JEH clade edit Euglenozoa and Heterolobosea Percolozoa or Eozoa as named by Cavalier Smith 14 appear to be particularly close relatives and are united by the presence of discoid cristae within the mitochondria Superphylum Discicristata A close relationship has been shown between Discicristata and Jakobida 15 the latter having tubular cristae like most other protists and hence were united under the taxon name Discoba which was proposed for this apparently monophyletic group 1 Metamonads edit Metamonads are unusual in not having classical mitochondria instead they have hydrogenosomes mitosomes or uncharacterised organelles The oxymonad Monocercomonoides is reported to have completely lost homologous organelles There are competing explanations 16 17 Malawimonads edit The malawimonads have been proposed to be members of Excavata owing to their typical excavate morphology and phylogenetic affinity to other excavate groups in some molecular phylogenies However their position among eukaryotes remains elusive 2 Ancyromonads edit Ancyromonads are small free living cells with a narrow longitudinal groove down one side of the cell The ancyromonad groove is not used for suspension feeding unlike in typical excavates e g malawimonads jakobids Trimastix Carpediemonas Kiperferlia etc Ancyromonads instead capture prokaryotes attached to surfaces The phylogenetic placement of ancyromonads is poorly understood in 2020 however some phylogenetic analyses place them as close relatives of malawimonads 9 Evolution editOrigin of the Eukaryotes edit Further information Eukaryogenesis The conventional explanation for the origin of the Eukaryotes is that a heimdallarchaeian or another Archaea acquired an alphaproteobacterium as an endosymbiont and that this became the mitochondrion the organelle providing oxidative respiration to the eukaryotic cell 18 Caesar al Jewari and Sandra Baldauf argue instead that the Eukaryotes possibly started with an endosymbiosis event of a Deltaproteobacterium or Aammaproteobacterium accounting for the otherwise unexplained presence of anaerobic bacterial enzymes in Metamonada The sister of the Preaxostyla within Metamonada represents the rest of the Eukaryotes which acquired an Alphaproteobacterium In their scenario the hydrogenosome and mitosome both conventionally considered mitochondrion derived organelles would predate the mitochondrion and instead be derived from the earlier symbiotic bacterium 17 Phylogeny edit In 2023 using molecular phylogenetic analysis of 186 taxa Al Jewari and Baldauf proposed a phylogenetic tree with the metamonad Parabasalia as basal Eukaryotes Discoba and the rest of the Eukaryota appear to have emerged as sister taxon to the Preaxostyla incorporating a single alphaproteobacterium as mitochondria by endosymbiosis Thus the Fornicata are more closely related to e g animals than to Parabasalia The rest of the Eukaryotes emerged within the Excavata as sister of the Discoba as they are within the same clade but are not cladistically considered part of the Excavata yet the Excavata are in this analysis highly paraphyletic 17 Hodarchaeales 18 Eukaryota ParabasaliaFornicataPreaxostylaDiscoba JakobidaHeteroloboseaEuglenozoa and alliesNeokaryotes Amorphea inc animals fungi SARArchaeplastida inc plants Excavata The Anaeramoeba are associated with Parabasalia but could turn out to be more basal as the root of a tree is often difficult to pinpoint 19 See also editMetakaryotaGallery edit nbsp Euglena Euglenozoa Euglenoida nbsp Trypanosoma brucei Euglenozoa Kinetoplastida nbsp Bodo sp Euglenozoa Kinetoplastida nbsp Percolomonas sp Percolozoa nbsp Stephanopogon sp Percolozoa nbsp Stages of Naegleria sp Percolozoa Heterolobosea nbsp Acrasis rosea Percolozoa Heterolobosea nbsp Jakobids Jakobida nbsp Trichomonas vaginalis Metamonada Parabasalia nbsp Retortamonas sp left Metamonada Fornicata Retortamonadida nbsp Giardia sp Metamonada Fornicata Diplomonadida References edit a b Hampl Vladimir Hug Laura Leigh Jessica W et al 2009 Phylogenomic analyses support the monophyly of Excavata and resolve relationships among eukaryotic supergroups PNAS 106 10 3859 3864 Bibcode 2009PNAS 106 3859H doi 10 1073 pnas 0807880106 PMC 2656170 PMID 19237557 a b Simpson Alastair G B Inagaki Yuji Roger Andrew J 2006 Comprehensive multigene phylogenies of excavate protists reveal the evolutionary positions of primitive eukaryotes Molecular Biology and Evolution 23 3 615 625 doi 10 1093 molbev msj068 PMID 16308337 a b Simpson Alastair G B Patterson David J December 1999 The ultrastructure of Carpediemonas membranifera Eukaryota with reference to the excavate hypothesis European Journal of Protistology 35 4 353 370 doi 10 1016 S0932 4739 99 80044 3 a b c Simpson Alastair G B November 2003 Cytoskeletal organization phylogenetic affinities and systematics in the contentious taxon Excavata Eukaryota International Journal of Systematic and Evolutionary Microbiology 53 6 1759 1777 doi 10 1099 ijs 0 02578 0 PMID 14657103 a b Dawkins Richard Wong Yan 2016 The Ancestor s Tale Houghton Mifflin Harcourt ISBN 978 0544859937 a b c Cavalier Smith Thomas 2002 The phagotrophic origin of eukaryotes and phylogenetic classification of Protozoa International Journal of Systematic and Evolutionary Microbiology 52 2 297 354 doi 10 1099 00207713 52 2 297 PMID 11931142 Dawson Scott C Paredez Alexander R 2013 Alternative cytoskeletal landscapes cytoskeletal novelty and evolution in basal excavate protists Current Opinion in Cell Biology 25 1 134 141 doi 10 1016 j ceb 2012 11 005 PMC 4927265 PMID 23312067 Burki Fabien Roger Andrew J Brown Matthew W et al January 2020 The New Tree of Eukaryotes Trends in Ecology amp Evolution 35 1 43 55 doi 10 1016 j tree 2019 08 008 PMID 31606140 S2CID 204545629 a b Brown Matthew W Heiss Aaron A Kamikawa Ryoma et al 2018 01 19 Phylogenomics Places Orphan Protistan Lineages in a Novel Eukaryotic Super Group Genome Biology and Evolution 10 2 427 433 doi 10 1093 gbe evy014 PMC 5793813 PMID 29360967 Heiss Aaron A Kolisko Martin Ekelund Fleming et al 4 April 2018 Combined morphological and phylogenomic re examination of malawimonads a critical taxon for inferring the evolutionary history of eukaryotes Royal Society Open Science 5 4 171707 Bibcode 2018RSOS 571707H doi 10 1098 rsos 171707 PMC 5936906 PMID 29765641 Keeling Patrick J Burki Fabien 19 August 2019 Progress towards the Tree of Eukaryotes Current Biology 29 16 R808 R817 doi 10 1016 j cub 2019 07 031 PMID 31430481 Laura Wegener Parfrey Erika Barbero Elyse Lasser Micah Dunthorn Debashish Bhattacharya David J Patterson Laura A Katz December 2006 Evaluating support for the current classification of eukaryotic diversity PLOS Genetics 2 12 e220 doi 10 1371 JOURNAL PGEN 0020220 ISSN 1553 7390 PMC 1713255 PMID 17194223 Wikidata Q21090155 Tice Alexander K Zihala David Panek Tomas et al 2021 PhyloFisher A phylogenomic package for resolving eukaryotic relationships PLOS Biology 19 8 e3001365 doi 10 1371 journal pbio 3001365 PMC 8345874 PMID 34358228 Cavalier Smith Thomas 23 December 2009 Kingdoms Protozoa and Chromista and the eozoan root of the eukaryotic tree Biology Letters The Royal Society 6 3 342 345 doi 10 1098 rsbl 2009 0948 ISSN 1744 9561 PMC 2880060 PMID 20031978 Rodriguez Ezpeleta Naiara Brinkmann Henner Burger Gertraud et al 2007 Toward Resolving the Eukaryotic Tree The Phylogenetic Positions of Jakobids and Cercozoans Current Biology 17 16 1420 1425 doi 10 1016 j cub 2007 07 036 PMID 17689961 Bui Elisabeth T Bradley Peter J Johnson Patricia J 3 September 1996 A common evolutionary origin for mitochondria and hydrogenosomes Proceedings of the National Academy of Sciences 93 18 9651 9656 Bibcode 1996PNAS 93 9651B doi 10 1073 pnas 93 18 9651 ISSN 0027 8424 PMC 38483 PMID 8790385 a b c Al Jewari Caesar Baldauf Sandra L 2023 04 28 An excavate root for the eukaryote tree of life Science Advances 9 17 eade4973 Bibcode 2023SciA 9E4973A doi 10 1126 sciadv ade4973 ISSN 2375 2548 PMC 10146883 PMID 37115919 a b Eme Laura Tamarit Daniel Caceres Eva F et al 2023 03 09 Inference and reconstruction of the heimdallarchaeial ancestry of eukaryotes Nature 618 7967 992 999 doi 10 1101 2023 03 07 531504 PMC 10307638 PMID 37316666 Stairs Courtney W Taborsky Petr Salomaki Eric D et al December 2021 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 External links editOpen Tree of Life Taxonomicon Tree of Life Eukaryotes Tree of Life Jakobida Tree of Life Fornicata Retrieved from https en wikipedia org w index php title Excavata amp oldid 1194661279, wikipedia, wiki, book, books, library,

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