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Excavata
Temporal range: Neoproterozoic–present
Giardia lamblia.jpg
Giardia lamblia, a parasitic diplomonad
Scientific classification e
Domain: Eukaryota
(unranked): Excavata
(Cavalier-Smith), 2002
Phyla
  • Discoba
  • Loukozoa
    • Metamonada
    • Malawimonas
    • Ancyromonadida
Excavata cell schemes
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.

Excavata is a major supergroup of unicellular organisms belonging to the domain Eukaryota. It was first suggested by Simpson and Patterson in 1999 and introduced by Thomas Cavalier-Smith in 2002 as a formal taxon. It contains a variety of free-living and symbiotic forms, and also includes some important parasites of humans, including Giardia and Trichomonas. Excavates were formerly considered to be included in the now obsolete Protista kingdom. They are classified based on their flagellar structures, and they are considered to be the most basal Flagellate lineage. Phylogenomic analyses split the members of the Excavates into three different and not all closely related groups: Discobids, Metamonads and Malawimonads. 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. 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. 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. However, various groups that lack these traits may be considered excavates based on genetic evidence (primarily phylogenetic trees of molecular sequences).

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.

Classification

See also: Eukaryote#Phylogeny and wikispecies:Excavata

Excavates are classified into six major subdivisions at the phylum/class level. These are shown in the table below. An additional group, Malawimonadida (e.g. Malawimonas), may also be included amongst excavates, though phylogenetic evidence is equivocal.

Kingdom/Superphylum Phylum/Class Representative genera (examples) Description
Discoba or JEH or Eozoa Tsukubea Tsukubamonas
Euglenozoa EuglenaTrypanosoma 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.
Neolouka Malawimonadida Malawimonas

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, 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.

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.

Monophyly

Excavate relationships are still uncertain; it is possible that they are not a monophyletic group. The monophyly of the excavates is far from clear, although there seem to be several clades within the excavates that are monophyletic.

Certain excavates are often considered among the most primitive eukaryotes, based partly on their placement in many evolutionary trees. This could encourage proposals that excavates are a paraphyletic grade that includes the ancestors of other living eukaryotes. However, the placement of certain excavates as 'early branches' may be an analysis artifact caused by long branch attraction, as has been seen with some other groups, for example, microsporidia.

Malawimonads

The malawimonads are generally considered 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.

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. Consequently, it is possible that ancyromonads are relevant for understand the evolution of 'true' excavates.

Cladogram

Here is a proposed cladogram for the positioning of the Excavata, with the Eukaryote root in the excavates, mainly following Cavalier-Smith.

Eukaryota

Tsukubea



Discicristata

Euglenozoa



Percolozoa



Orthokaryotes

Jakobea


Neokaryotes
Scotokaryotes/

Metamonada




Malawimonas


Sulcozoa/Podiata/

Planomonadida




CRuMs


Amorphea/

Amoebozoa


Obazoa

Breviata




Apusomonadida



Opisthokonta/Choanozoa









Corticata/

Archaeplastida


Chromista

Hacrobia



SAR








In this view, excavata is highly paraphyletic, and is proposed to be abandoned. In alternative view, the Discoba are sister to the rest of the Diphoda.

Eukaryota
Opimoda

Metamonada




Malawimonas


Podiata

CRuMs


Amorphea

Amoebozoa


Obazoa

Breviata




Apusomonadida



Opisthokonta








Diphoda
Discoba

Jakobea




Tsukubea


Discicristata

Heterolobosa



Euglenozoa








Archaeplastida



Cryptomonadida





Haptophyta



SAR







Gallery

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