Excavata

Excavata is a former supergroup of unicellular organisms belonging to the domain Eukaryota.[1][2][3] It was first suggested by Simpson and Patterson in 1999[4][5] and introduced by Thomas Cavalier-Smith in 2002 as a formal taxon. It contains a variety of free-living and symbiotic forms, and includes some important parasites of humans, such as Giardia and Trichomonas.[6] Excavates were formerly considered to be included in the now obsolete Protista kingdom.[7] They were classified based on their flagellar structures,[5] and were considered to be the most basal flagellate lineage.[8]

Excavata
Temporal range:
Giardia lamblia, a parasitic diplomonad
Scientific classification e
(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.

Phylogenomic analyses indicate that the Excavata are polyphyletic, not forming a single natural group.[9] Excavata has therefore been split into three groups, not all closely related: Discobids, Metamonads and Malawimonads.[10][11][12] Except for Euglenozoa, they are all non-photosynthetic.

Characteristics

Most excavates are unicellular, heterotrophic flagellates. Only the Euglenozoa are photosynthetic. In some (particularly anaerobic intestinal parasites), the mitochondria have been greatly reduced.[6] 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.[5] 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.[4][7] However, various groups that lack these traits may be considered excavates based on genetic evidence (primarily phylogenetic trees of molecular sequences).[7]

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

Excavate relationships were always uncertain, suggesting that they are not a monophyletic group.[13] The malawimonads often do not branch with the other Excavata.[14]

Excavates were thought to include multiple groups:

Kingdom/SuperphylumIncluded taxaRepresentative genera (examples)Description
Discoba or JEH or EozoaTsukubeaTsukubamonas
Euglenozoa Euglena, Trypanosoma Many important parasites, one large group with plastids (chloroplasts)
Heterolobosea (Percolozoa)Naegleria, AcrasisMost alternate between flagellate and amoeboid forms
JakobeaJakoba, ReclinomonasFree-living, sometimes loricate flagellates, with very gene-rich mitochondrial genomes
Metamonada or PODPreaxostylaOxymonads, TrimastixAmitochondriate flagellates, either free-living (Trimastix, Paratrimastix) or living in the hindguts of insects
FornicataGiardia, CarpediemonasAmitochondriate, mostly symbiotes and parasites of animals.
ParabasaliaTrichomonasAmitochondriate flagellates, generally intestinal commensals of insects. Some human pathogens.
Neolouka MalawimonadidaMalawimonas

Discoba or JEH clade

Euglenozoa and Heterolobosea (Percolozoa) or Eozoa (Cavalier-Smith) 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.[2]

Metamonads

Metamonads are unusual in having lost classical mitochondria—instead they have hydrogenosomes, mitosomes or uncharacterised organelles. The oxymonad Monocercomonoides is reported to have completely lost homologous organelles.

Malawimonads

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.[3]

Ancyromonads

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.[16]

Cladograms

This proposed cladogram for the positioning of the Excavata, with the Eukaryote root in the excavates, mainly follows Cavalier-Smith.[17][18][19][20][21][22][23][24][25]

Eukaryota

Tsukubea

Discicristata

Euglenozoa

Percolozoa

Orthokaryotes

Jakobea

Neokaryotes
Scotokaryotes/

Metamonada

Malawimonas

Podiata/

Planomonadida

CRuMs

Amorphea/

Amoebozoa

Obazoa

Breviata

Apusomonadida

Opisthokonta/Choanozoa

Unikont
Sarcomastigota
Opimoda/Neozoa
Corticata/

Archaeplastida

Chromista

Hacrobia

SAR

Bikont/Diaphoretickes
Excavate taxa

This perspective treats the Excavata as polyphyletic, and some, including Cavalier Smith, argue that it should be abandoned.[26] In an alternative view, the Discoba have been suggested as sister to the rest of the Diphoda, once again breaking up the Excavata.[18][27]

Eukaryota
Opimoda
Podiata

CRuMs

Amorphea

Amoebozoa

Obazoa

Breviata

Apusomonadida

Opisthokonta

Malawimonas

Metamonada

Diphoda
Discoba

Jakobea

Tsukubea

Discicristata

Heterolobosa

Euglenozoa

Archaeplastida

Cryptomonadida

Haptophyta

SAR

Excavate taxa

References

  1. Hampl, V.; Hug, L.; Leigh, J. W.; Dacks, J. B.; Lang, B. F.; Simpson, A. G. B.; Roger, A. J. (2009). "Phylogenomic analyses support the monophyly of Excavata and resolve relationships among eukaryotic "supergroups"". Proceedings of the National Academy of Sciences. 106 (10): 3859–3864. Bibcode:2009PNAS..106.3859H. doi:10.1073/pnas.0807880106. PMC 2656170. PMID 19237557.
  2. Hampl V, Hug L, Leigh JW, et al. (2009). "Phylogenomic analyses support the monophyly of Excavata and resolve relationships among eukaryotic "supergroups"". Proc. Natl. Acad. Sci. U.S.A. 106 (10): 3859–64. Bibcode:2009PNAS..106.3859H. doi:10.1073/pnas.0807880106. PMC 2656170. PMID 19237557.
  3. Simpson, Ag; Inagaki, Y; Roger, Aj (2006). "Comprehensive multigene phylogenies of excavate protists reveal the evolutionary positions of "primitive" eukaryotes". Molecular Biology and Evolution. 23 (3): 615–25. doi:10.1093/molbev/msj068. PMID 16308337.
  4. Simpson, Alastair G.B.; Patterson, David J. (Dec 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.
  5. Simpson, Alastair G.B. (2003-11-01). "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.
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