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{{good article}}
{{Short description|Clade containing animals and some protists}}
{{Automatic taxobox 
| name=Holozoans
| display_parents=3
| taxon=Holozoa
| authority=Lang et al., 2002<ref name="Lang et al. 2002"/>
| fossil_range={{long fossil range|1100|0|Early [[Tonian]] - [[Holocene|Present]]}}
| image=<imagemap>
File:Holozoan_diversity.png|300px
rect 1 1 400 300 [[Pluriformea]]
rect 400 1 800 300 [[Dermocystida]]
rect 800 1 1200 300 [[Filasterea]]

rect 1 300 400 600 [[Ichthyophonida]]
rect 400 300 800 600 [[Choanoflagellata]]
rect 800 300 1200 600 [[Choanoflagellata]]

rect 1 600 400 900 [[Porifera]]
rect 400 600 800 900 [[Ctenophora]] 
rect 800 600 1200 900 [[Cnidaria]]

rect 1 900 400 1200 [[Deuterostomia]]
rect 400 900 800 1200 [[Spiralia]]
rect 800 900 1200 1200 [[Ecdysozoa]]
</imagemap>
| image_caption = Holozoan diversity. Top half displays representatives of holozoan protists. Bottom half displays representatives of metazoans.
| subdivision_ranks = Clades<ref name="Adl 2019"/>
| subdivision =
* [[Ichthyosporea]] (=Mesomycetozoea)
* ''[[Tunicaraptor]]''<ref name="ChoanoEvo"/>
* [[Pluriformea]]
** ''[[Corallochytrium]]''
** ''[[Syssomonas]]''
* [[Filozoa]]
** [[Filasterea]]
** [[Choanozoa]]
*** [[Choanoflagellata]]
*** [[Metazoa]] (=animals)
----
{{small|'''Incertae sedis'''}}
*{{extinct}}''[[Bicellum brasieri]]''<ref name="Strother 2021"/>
|synonyms=
*Choanofila {{au|Cavalier-Smith, 2009}}<ref name="CavSmith09">{{cite journal|first1=Thomas|last1=Cavalier-Smith|date=2009|title=Megaphylogeny, Cell Body Plans, Adaptive Zones: Causes and Timing of Eukaryote Basal Radiations|journal=Journal of Eukaryotic Microbiology|volume=56|pages=26–33|doi=10.1111/j.1550-7408.2008.00373.x|doi-access=free}}</ref> (plus animals)
}}
'''Holozoa''' ({{etymology|grc|ὅλος (holos)|whole||ζῷον (zoion)|animal}}) is a [[clade]] of organisms that includes [[animal]]s and their closest [[unicellular|single-celled]] relatives, but excludes [[fungi]] and all other organisms. Together they amount to more than 1.5 million species of purely [[heterotrophic]] organisms, including around 300 [[unicellular organism|unicellular]] species. It consists of various subgroups, namely [[Metazoa]] (or animals) and the [[protist]]s [[Choanoflagellata]], [[Filasterea]], [[Pluriformea]] and [[Ichthyosporea]]. Along with fungi and some other groups, Holozoa is part of the [[Opisthokonta]], a [[supergroup (biology)|supergroup]] of [[eukaryotes]]. '''Choanofila''' was previously used as the name for a group similar in composition to Holozoa, but its usage is discouraged now because it excludes animals and is therefore [[paraphyletic]].

The holozoan protists play a crucial role in understanding the evolutionary steps leading to the emergence of [[multicellular]] animals from single-celled ancestors. Recent [[genomic]] studies have shed light on the evolutionary relationships between the various holozoan [[Lineage (evolution)|lineages]], revealing insights into the origins of [[multicellular organism|multicellularity]]. Some [[fossils]] of possible metazoans have been reinterpreted as holozoan protists.

== Characteristics ==
=== Composition ===
Holozoa is a [[clade]] that includes [[animals]] and their closest relatives, as well as their [[common ancestor]], but excludes [[fungi]]. It is defined on a [[Phylogenetic nomenclature#Phylogenetic definitions of clade names|branch-based]] approach as the clade encompassing all relatives of ''[[Homo sapiens]]'' (an animal), but not ''[[Neurospora crassa]]'' (a [[fungus]]).<ref name="Adl 2019"/> Holozoa, besides animals, primarily comprises unicellular [[protist]] lineages of varied [[Morphology (biology)|morphologies]] such as [[choanoflagellates]], [[filasterean]]s, [[ichthyosporea]]ns, and the distinct genera ''[[Corallochytrium]]'', ''[[Syssomonas]]'', and ''[[Tunicaraptor]]''.<ref name="Hehenberger-2017"/><ref name="ChoanoEvo"/>

* [[Choanoflagellata]], with around 250 species,<ref name="Sebé-Pedrós et al. 2017"/> are the closest living relatives of animals. They are free-living [[unicellular]] or [[Colony (biology)|colonial]] [[flagellate]]s that feed on [[bacteria]] using a characteristic "collar" of [[microvilli]]. The collar of choanoflagellates closely resembles sponge [[choanocyte|collar cells]],<ref name="Steenkamp"/> leading to theories since the 19th century about their relatedness to [[sponges]].<ref name="Handbook of the Protists 1"/> The mysterious ''[[Proterospongia]]'' is an example of a colonial choanoflagellate that was thought to be related to the origin of sponges.<ref name="Brunet & King 2022"/> The affinities of the other single-celled holozoans only began to be recognized in the 1990s.<ref>{{cite journal| doi=10.1073/pnas.93.21.11907| last1=Ragan | first1=Mark A.| last2=Goggin | first2=C. Louise| last3=Cawthorn | first3=Richard J.| last4=Cerenius | first4=Lage| last5=Jamieson | first5=Angela V.C.| last6=Plourde | first6=Susan M.| last7=Rand | first7=Thomas G.| last8=Söoderhäll | first8=Kenneth| last9=Gutell | first9=Robin R.| date=15 October 1996| title=A novel clade of protistan parasites near the animal-fungal divergence| journal=PNAS| volume=93 | pages=11907–11912| pmid=8876236| issue=21| pmc=38157 | bibcode=1996PNAS...9311907R | doi-access=free }}</ref>

* [[Ichthyosporea]], also known as Mesomycetozoea and comprising around 40 species, largely consist of [[parasites]] or [[commensal]]s. They interact with a diverse range of animals, from [[humans]] and [[fish]] to [[marine life|marine]] [[invertebrate]]s. Most reproduce through [[multinucleated]] [[colony (biology)|colonies]] and disperse as flagellates or [[amoeba]]e.<ref name="Sebé-Pedrós et al. 2017"/>

* [[Filasterea]] is a group of 6 amoeboid species belonging to the genera ''[[Ministeria]]'', ''[[Pigoraptor]]'',<ref name="Hehenberger-2017"/> ''[[Capsaspora]]'', and ''[[Txikispora]]'',<ref name="Txikispora"/> united by the structure of their thread-like [[pseudopod]]s.<ref name="pmid18461162"/>

* [[Pluriformea]] is a provisional name for the clade composed by the two species ''[[Corallochytrium limacisporium]]'' and ''[[Syssomonas multiformis]]''. These organisms have varied shapes, including cellular aggregations, amoebae, flagellates, and amoeboflagellates.<ref name="Hehenberger-2017"/> 

* ''[[Tunicaraptor unikontum]]'' is the newest discovered clade, whose position within Holozoa has yet to be resolved. It is a flagellate with a specialized "mouth" structure absent in other holozoans.<ref name="ChoanoEvo"/>

* [[Metazoa]], known as animals, are multicellular organisms that sum more than 1.5 million living species.<ref>{{cite journal|doi=10.11646/zootaxa.3703.1.3|first1=Zhi-Qiang|last1=Zhang|date=2013|journal=Zootaxa|title=Animal biodiversity: an update of classification and diversity in 2013+|volume=3703|issue=1|pages=5–11|doi-access=free}}</ref> They are characterized by a [[blastula]] phase during their [[embryonic development]] and, except for the amorphous [[sponges]], the formation of [[germ layer]]s and differentiated [[tissue (biology)|tissues]].<ref name="Adl 2019"/>

===Genetics===
The first [[DNA sequencing|sequenced]] unicellular holozoan [[genome]] was that of ''[[Monosiga brevicollis]]'', a [[choanoflagellate]]. It measures around 41.6 [[base pair|mega–base-pairs]] (Mbp) and contains around 9200 coding [[gene]]s, making it comparable in size to the genome of filamentous [[fungi]]. Animal genomes are usually larger (e.g. [[human genome]], 2900 Mbp; [[Drosophila melanogaster|fruit fly]], 180 Mbp), with some exceptions.<ref name="King et al 2008">{{cite journal|vauthors=King N, Westbrook M, Young S, etal.|title=The genome of the choanoflagellate ''Monosiga brevicollis'' and the origin of metazoans|journal=Nature|volume=451|pages=783–788|date=2008|doi=10.1038/nature06617|doi-access=free|hdl=2027.42/62649|hdl-access=free}}</ref>

== Evolution ==
=== Phylogeny ===
Holozoa, along with a clade that contains [[fungi]] and their [[protist]] relatives ([[Holomycota]]), are part of the larger [[supergroup (biology)|supergroup]] of eukaryotes known as [[Opisthokonta]]. Holozoa [[divergence time estimation|diverged]] from their opisthokont ancestor around 1070 million years ago (Mya).<ref name="Lawal 2020">{{cite journal|vauthors=Lawal HM, Schilde C, Kin K, etal.|title=Cold climate adaptation is a plausible cause for evolution of multicellular sporulation in ''Dictyostelia''|journal=Scientific Reports|volume=10|pages=8797|date=2020|doi=10.1038/s41598-020-65709-3|doi-access=free|pmc=7260361}}</ref> The choanoflagellates, animals and filastereans group together as the clade [[Filozoa]]. Within Filozoa, the choanoflagellates and animals group together as the clade [[Choanozoa]].<ref name="pmid18461162"/> Based on [[phylogenetic]] and [[phylogenomic]] analyses, the [[cladogram]] of Holozoa is shown below:<ref name="Parfrey-2011"/><ref name="Torruella-2015"/><ref name="Hehenberger-2017"/><ref name="ChoanoEvo"/>

{{Clade|style=font-size:90%;|label1=[[Opisthokonta]]|sublabel1=1250 Mya|1={{clade
  |label1=[[Holomycota]]|sublabel1=1110 Mya|1={{clade
    |1=[[Cristidiscoidea]] [[File:Nuclearia_sp_Nikko_(cropped).jpg|50px]]
    |2=[[Fungi]] [[File:Linear_arrangement_of_ascospores_in_the_asci_of_the_fungus_Sordaria_macrospora_Cropped.jpg|60px]]
    }}
  |label2='''Holozoa'''|sublabel2=1070 Mya|2={{clade
    |label1=[[Ichthyosporea]]|bar1=blue|1={{clade
      |1=[[Dermocystida]] [[File:Chromosphaera_perkinsii.jpg|50px]]
      |2=[[Ichthyophonida]] [[File:Sphaeroforma_arctica.jpg|50 px]]
      }}
    |label2=[[Pluriformea]]|bar2=blue|state2=dashed|2={{clade
        |2=''[[Syssomonas]]'' [[File:Syssomonas_2X_2020.webp|60px]]
        |1=''[[Corallochytrium]]'' [[File:Corallochytrium limacisporum.png|50px]]
        }}
      |3=''[[Tunicaraptor]]''|state3=dashed
      |label4=[[Filozoa]]|4={{clade
        |1=[[Filasterea]] [[File:Ministeria vibrans.jpeg|60 px]] 
        |label2=[[Choanozoa]] |sublabel2=960 Mya |2={{clade 
          |1=[[Choanoflagellata]]  [[File:Desmarella moniliformis.jpg|50px]]
          |2={{clade|sublabel1=''multicellularity''&nbsp;|1=[[Metazoa]] [[File:Comb_jelly.jpg|50px]]}}
          }}
        }}
      }}
    }}
|grouplabel1={{clade labels|label1=[[Teretosporea]]|top1=40%}}
}}

Uncertainty remains around the relationship of the two most [[basal (phylogenetics)|basal]] groups, [[Ichthyosporea]] and [[Pluriformea]].<ref name="Adl 2019"/> They may be [[sister group|sister]] to each other, forming the putative clade [[Teretosporea]].<ref name="Grau-Bové 2017">{{cite journal|first1=Xavier|last1=Grau-Bové|first2=Guifré|last2=Torruella|first3=Stuart|last3=Donachie|first4=Hiroshi|last4=Suga|first5=Guy |last5=Leonard|first6=Thomas A|last6=Richards|first7=Iñaki|last7=Ruiz-Trillo|date=2017|title=Dynamics of genomic innovation in the unicellular ancestry of animals|journal=eLife|volume=6|pages=e26036|doi=10.7554/eLife.26036|doi-access=free|pmc=5560861}}</ref> Alternatively, Ichthyosporea may be the earliest-branching of the two, while Pluriformea is sister to the [[Filozoa]] clade comprising filastereans, choanoflagellates and animals. This second outcome is more [[Robust statistics|strongly supported]] after the discovery of ''[[Syssomonas]]''.<ref name="ChoanoEvo"/><ref name="Hehenberger-2017"/>

The position of ''[[Tunicaraptor]]'', the newest holozoan member, is still unresolved. Three different phylogenetic positions of ''Tunicaraptor'' have been obtained from analyses: as the sister group to [[Filasterea]], as sister to [[Filozoa]], or as the most basal group of all Holozoa.<ref name="ChoanoEvo"/><ref name="Ros-Rocher 2021"/>

[[Environmental DNA]] surveys of oceans have revealed new diverse lineages of Holozoa. Most of them nest within known groups, mainly [[Ichthyosporea]] and [[Choanoflagellata]]. However, one environmental clade does not nest within any known group and is a potential new holozoan lineage. It has been tentatively named MASHOL (for 'marine small Holozoa').<ref>{{cite journal|first1=Alicia S|last1=Arroyo|first2=Romain|last2=Lannes|first3=Eric|last3=Bapteste|first4=Iñaki |last4=Ruiz-Trillo|title=Gene Similarity Networks Unveil a Potential Novel Unicellular Group Closely Related to Animals from the ''Tara'' Oceans Expedition|journal=Genome Biology and Evolution|volume=12|issue=9|date=September 2020|pages=1664–1678|doi=10.1093/gbe/evaa117|doi-access=free|pmc=7533066}}</ref>
=== Unicellular ancestry of animals ===
[[File:RG347357480 Fig1.png|thumb|upright=2|Unicellular holozoans do not go through [[animal embryonic development]], but they display developmental processes using similar molecules. An [[Myofilament|actomyosin]] network controls the cellularization of both an [[ichthyosporea]]n [[coenocyte]] (A) and a [[Drosophila melanogaster|fruit fly]] [[blastoderm]] (B). Similarly, actomyosin contraction allows both the shaping of [[choanoflagellate]] [[Colony (biology)|colonies]] (C) and the [[gastrulation]] of animal [[embryo]]s.]]
The quest to elucidate the [[Animal#Evolutionary_origin|evolutionary origins of animals]] from a unicellular ancestor requires an examination of the transition to [[multicellularity]]. In the absence of a [[fossil record]] documenting this evolution, insights into the unicellular ancestor of animals are obtained from the [[DNA sequencing|analysis]] of shared [[genes]] and [[genetic pathway]]s between animals and their closest living unicellular relatives. The genetic content of these single-celled holozoans has revealed a significant discovery: many genetic characteristics previously thought as unique to animals can also be found in these unicellular relatives. This suggests that the origin of multicellular animals did not happen solely because of the appearance of new genes (i.e. innovation), but because of pre-existing genes that were adapted or utilized in new ways (i.e. co-option).<ref name="Sebé-Pedrós et al. 2017"/><ref name="Hehenberger-2017"/> For example:

* [[Adhesion protein]]s are necessary in allowing cells to stick to each other and to the [[extracellular matrix]], forming layers and tissues in animals. Some unicellular holozoans, like [[choanoflagellates]] and [[filasterea]]ns, possess genes that encode proteins involved in [[cell adhesion#cell-cell junctions|cell-cell adhesion]] and [[cell adhesion#cell-matrix junctions|cell-matrix adhesion]] (e.g. [[cadherin]] and [[integrin]], respectively). Other genes, however, seem to be exclusively found in animals (e.g. [[β-catenin]]).<ref name="Sebé-Pedrós et al. 2017"/>

* ECM-related proteins, involved in the formation of the [[extracellular matrix]], are present in other holozoans (e.g. [[laminin]]s, [[collagen]]s and [[fibronectin]]s).<ref name="Suga et al. 2013"/>

*[[Signal transduction]] proteins are another requirement for metazoan multicellularity. Some animal cytoplasmic [[tyrosine kinase]]s (such as [[focal adhesion kinase]]) and the [[Hippo signaling pathway]] are present in unicellular holozoans. Other [[signaling pathways]] highly conserved in animals (e.g. [[Hedgehog signaling pathway|Hedgehog]], [[Wnt signaling pathway|WNT]], [[Transforming growth factor beta|TGFβ]], [[JAK-STAT]] and [[Notch signaling pathway|Notch]]) are absent in other holozoans, but similar signaling receptors [[convergent evolution|evolved independently]] in [[choanoflagellate]]s, [[filasterean]]s and [[ichthyosporea]]ns (e.g. [[receptor tyrosine kinase]]s).<ref name="Sebé-Pedrós et al. 2017"/>
*A considerable portion of animal [[transcription factor]]s (TF) is already present in unicellular holozoans, including some TF classes previously thought to be animal-specific (e.g. [[p53]] and [[T-box]]).<ref name="Sebé-Pedrós et al. 2017"/>

Additionally, many biological processes seen in animals are already present in their unicellular relatives, such as [[sexual reproduction]] and [[gametogenesis]] in the [[choanoflagellate]] ''[[Salpingoeca rosetta]]'' and several types of multi[[cellular differentiation]].<ref name="Sebé-Pedrós et al. 2017"/>

=== Fossil record ===
[[File:Naked stereoblasts of billion-years-old Bicellum Brasieri.jpg|thumb|A fossilized sample of ''[[Bicellum brasieri]]'', a billion-year-old potential holozoan.]]
A [[billion]]-year-old [[freshwater]] microscopic [[fossil]] named ''[[Bicellum brasieri]]'' is possibly the earliest known holozoan. It shows two differentiated [[cell type]]s or [[biological life cycle|life cycle stages]]. It consists of a spherical ball of tightly packed cells (stereoblasts) enclosed in a single layer of elongated [[cell (biology)|cell]]s. There are also two populations of stereoblasts with mixed shapes, which have been interpreted as [[cellular migration]] to the periphery, a movement that could be explained by differential [[cell adhesion#cell-cell junctions|cell-cell adhesion]]. These occurrences are consistent with extant unicellular holozoans, which are known to form multicellular stages in complex life cycles.<ref name="Strother 2021"/>

Proposed [[Ediacaran]] fossil "[[embryos]]" of early [[metazoa]]ns, discovered in the [[Doushantuo Formation]], have been reinterpreted as non-animal [[protist]]s within Holozoa. According to some authors, although they present possible [[cleavage (embryo)|embryonic cleavage]], they lack metazoan [[synapomorphy|synapomorphies]] such as tissue differentiation and nearby juveniles or adults. Instead, its development is comparable to the [[germination]] stage of non-animal holozoans. They possibly represent an [[evolutionary grade]] in which palintomic cleavage (i.e. rapid [[cell division]]s without [[cytoplasm]]ic growth in between, a characteristic of animal [[cleavage (embryo)|embryonic cleavage]])<ref>{{cite journal|title=Cell differentiation and germ–soma separation in Ediacaran animal embryo-like fossils|vauthors=Chen L, Xiao S, Pang K, Zhou C, Yuan X|pages=238–241|date=September 2014|journal=Nature|volume=516|issue=7530 |doi=10.1038/nature13766|pmid=25252979 |bibcode=2014Natur.516..238C |s2cid=4448316 }}</ref> was the method of [[biological dispersal|dispersal]] and [[reproduction|propagation]].<ref name="Huldtgren et al. 2011"/>

== Taxonomy ==
===History===
Prior to 2002, a relationship between [[Choanoflagellata]], [[Ichthyosporea]] and the [[animal]]-[[fungi]] divergence was considered on the basis of [[morphology (biology)|morphology]] and [[ultrastructure]]. Early phylogenetic analyses gave contradicting results, because the amount of available DNA sequences was insufficient to yield unambiguous results. The taxonomic uncertainty was such that, for example, some Ichthyosporea were traditionally treated as [[trichomycete]] fungi.<ref name="Lang et al. 2002"/>

Holozoa was first recognized as a clade in 2002 through a [[phylogenomic]] analysis by Franz Bernd Lang, Charles J. O'Kelly and other collaborators, as part of a [[scientific paper|paper]] published in the journal ''[[Current Biology]]''. The study used complete [[Mitochondrial genome|mitochondrial genomes]] of a choanoflagellate (''[[Monosiga brevicollis]]'') and an ichthyosporean (''[[Amoebidium parasiticum]]'') to firmly resolve the position of Ichthyosporea as the sister group to Choanoflagellata+Metazoa. This clade was named Holozoa ({{etymology|grc|ὅλος (holos)|whole||ζῷον (zoion)|animal}}), meaning 'whole animal', referencing the wider animal ancestry that it contains.<ref name="Lang et al. 2002"/>

Holozoa has since been supported as a robust clade by every posterior analysis,<ref name="Ros-Rocher 2021"/> even after the discovery of more taxa nested within it (namely [[Filasterea]] since 2008,<ref name="pmid18461162"/> and the [[pluriformea]]n species ''[[Corallochytrium]]'' and ''[[Syssomonas]]'' since 2014<ref>{{cite thesis|vauthors=Torruella G, de Mendoza A, Grau-Bové X, Donachie S, Pérez-Cordón G, Sitjà-Bobadilla A, Paley R, Manohar CS, Nichols K, Eme L, del Campo J|chapter=Phylotranscriptomics reveals ancient and convergent features in ''Corallochytrium'' and ''Ministeria'' (Holozoa, Opisthokonta)|title=Phylogeny and evolutionary perspective of Opisthokonta protists|date=2014|volume=75|pages=1–9|publisher=Universitat de Barcelona|url=https://www.tdx.cat/bitstream/handle/10803/286509/GTiC_PhD_THESIS.pdf?sequence=2.txt#page=80|degree=PhD}}</ref> and 2017<ref name="Hehenberger-2017"/> respectively). As of 2019, the clade is accepted by the International Society of Protistologists, which revises the classification of eukaryotes.<ref name="Adl 2019" />

===Classification===
{{further|Taxonomy of Protista}}
In classifications that use traditional [[taxonomic rank]]s (e.g. kingdom, phylum, class), all holozoan protists are classified as subphylum '''Choanofila''' (phylum [[Choanozoa]],{{efn|name=Choanozoa}} kingdom [[Protozoa]]) while the animals are classified as a separate kingdom [[Metazoa]] or Animalia.<ref name="CavSmith 2021">{{cite journal|vauthors=Cavalier-Smith T|title=Ciliary transition zone evolution and the root of the eukaryote tree: implications for opisthokont origin and classification of kingdoms Protozoa, Plantae, and Fungi|journal=Protoplasma|date=May 2022|volume=259|issue=3|pages=487–593|doi=10.1007/s00709-021-01665-7|pmid=34940909|pmc=9010356}}</ref> This classification excludes animals, even though they descend from the same common ancestor as choanofilan protists, making it a [[paraphyletic]] group rather than a true clade. Modern [[cladistic]] approaches to [[eukaryotic]] classification prioritise [[monophyletic]] groupings over traditional ranks, which are increasingly perceived as redundant and superfluous. Because Holozoa is a clade, its use is preferred over the paraphyletic taxon Choanofila.<ref name="Adl 2019" />

*'''Holozoa''' {{au|Lang et al. 2002}}
**[[Incertae sedis]]: {{extinct}}''[[Bicellum brasieri]]'' {{au|Strother & Wellman 2021}}<ref name="Strother 2021"/> 
**''[[Tunicaraptor]]'' {{au|Tikhonenkov, Mikhailov, Hehenberger, Karpov, Prokina, Esaulov, Belyakova, Mazei, Mylnikov, Aleoshin & Keeling 2020}}<ref name="ChoanoEvo"/>
**[[Ichthyosporea]] {{au|Cavalier-Smith 1998}} [Mesomycetozoea {{au|Mendoza et al. 2002}}]
***[[Dermocystida]] {{au|Cavalier-Smith 1998}}
***[[Ichthyophonida]] {{au|Cavalier-Smith 1998}}
**[[Pluriformea]] {{au|Hehenberger et al. 2017}}
***''[[Corallochytrium]]'' {{au|Raghu-Kumar 1987}}
***''[[Syssomonas]]'' {{au|Tikhonenkov, Hehenberger, Mylnikov & Keeling 2017}}
**[[Filozoa]] {{au|Shalchian-Tabrizi et al. 2008}}
***[[Filasterea]] {{au|Shalchian-Tabrizi et al. 2008}}
****''[[Capsaspora]]'' {{au|Hertel, Bayne & Loker, 2002}}
****''[[Ministeria]]'' {{au|Patterson et al. 1993}}
****''[[Pigoraptor]]'' {{au|Tikhonenkov et al. 2017}}
****''[[Txikispora]]'' {{au|Urrutia, Feist & Bass 2022}}<ref name="Txikispora"/>
***[[Choanozoa]] {{au|Brunet & King 2017}} [Choanozoa {{au|Cavalier-Smith et al. 1991}} (P)]{{efn|name=Choanozoa|The term "[[Choanozoa]]" has been used since 1991 by [[Cavalier-Smith]] as a paraphyletic phylum of opisthokont protists,<ref name="Early Evo Euk"/> and the terms "[[Apoikozoa]]" and "choanimal" were proposed as names for the clade [[Metazoa]]+[[Choanoflagellata]]. However, these terms have not been formally described or adopted, and were rejected in favor of a renamed [[Choanozoa]] to fit the clade Metazoa+Choanoflagellata.<ref name="Adl 2019"/>}}
****[[Choanoflagellata]] {{au|Kent 1880–1882}} [Choanoflagellatea {{au|Cavalier-Smith 1997 emend. Cavalier-Smith 1998}}]
*****[[Craspedida]] {{au|Cavalier-Smith 1997, emend. Nitsche et al. 2011}}
*****[[Acanthoecida]] {{au|Cavalier-Smith 1997, emend. Nitsche et al. 2011}}
****[[Metazoa]] {{au|Haeckel 1874, emend. Adl et al. 2005}} [Animalia {{au|Linnaeus 1758}}]
*****[[Porifera]] {{au|Grant 1836}}
*****[[Placozoa]] {{au|Grell 1971}}
*****[[Ctenophora]] {{au|Eschscholtz 1829}}
*****[[Cnidaria]] {{au|Hatschek 1888}}
*****[[Bilateria]] {{au|Hatschek 1888}}

==Notes==
{{notelist}}

==References==
{{Reflist|refs=

<ref name="Ros-Rocher 2021">{{cite journal|vauthors=Ros-Rocher N, Pérez-Posada A, Michelle LM, Ruiz-Trillo I|date=February 2021|title=The origin of animals: an ancestral reconstruction of the unicellular-to-multicellular transition|doi=10.1098/rsob.200359|journal=Open Biol|pmid=33622103|volume=11|issue=2|doi-access=free|pages=200359|hdl=10261/251922|hdl-access=free}}</ref>

<ref name="pmid18461162">{{cite journal | last1=Shalchian-Tabrizi | first1=Kamran | last2=Minge | first2=Marianne A.| last3=Espelund | first3=Mari| last4=Orr | first4=Russell| last5=Ruden | first5=Torgeir| last6=Jakobsen | first6=Kjetill S.| last7=Cavalier-Smith | first7=Thomas |author7-link=Thomas Cavalier-Smith | title=Multigene phylogeny of choanozoa and the origin of animals |journal=PLOS ONE |volume=3 | editor1-first=Rodolfo  |issue=5 |pages=e2098 |date=7 May 2008 | editor1-last=Aramayo |pmid=18461162 |pmc=2346548 |doi=10.1371/journal.pone.0002098 | last8=Aramayo | first8=Rodolfo| bibcode=2008PLoSO...3.2098S| doi-access=free }}</ref>

<ref name="Steenkamp">{{cite journal | last1=Steenkamp | first1=Emma T.| last2=Wright | first2=Jane| last3=Baldauf | first3=Sandra L.| title=The Protistan Origins of Animals and Fungi| journal=Molecular Biology and Evolution| volume=23 | issue=1 | pages=93–106| doi=10.1093/molbev/msj011 | pmid=16151185| date=January 2006| doi-access=free}}</ref>

<ref name="Early Evo Euk">{{cite journal|title=Early evolution of eukaryote feeding modes, cell structural diversity, and classification of the protozoan phyla Loukozoa, Sulcozoa, and Choanozoa|vauthors=Cavalier-Smith T|journal=European Journal of Protistology|volume=49|issue=2|date=May 2013|pages=115–178|doi=10.1016/j.ejop.2012.06.001|pmid=23085100 }}</ref>

<ref name="Adl 2019">{{cite journal|vauthors=Adl SM, Bass D, Lane CE, Lukeš J, Schoch CL, Smirnov A, Agatha S, Berney C, Brown MW, Burki F, Cárdenas P, Čepička I, Chistyakova L, del Campo J, Dunthorn M, Edvardsen B, Eglit Y, Guillou L, Hampl V, Heiss AA, Hoppenrath M, James TY, Karnkowska A, Karpov S, Kim E, Kolisko M, Kudryavtsev A, ((Lahr DJG)), Lara E, Le Gall L, Lynn DH, Mann DG, Massana R, ((Mitchell EAD)), Morrow C, Park JS, Pawlowski JW, Powell MJ, Richter DJ, Rueckert S, Shadwick L, Shimano S, Spiegel FW, Torruella G, Youssef N, Zlatogursky V, Zhang Q|year=2019|title=Revisions to the Classification, Nomenclature, and Diversity of Eukaryotes|journal=Journal of Eukaryotic Microbiology|volume=66|issue=1 |pages=4–119|doi=10.1111/jeu.12691|pmid=30257078 |pmc=6492006 }}</ref>

<ref name="ChoanoEvo">{{cite journal|volume=30|issue=22|date=2020|pages=4500–4509|vauthors=Tikhonenkov DV, Mikhailov KV, Hehenberger E, Mylnikov AP, Aleoshin VV, Keeling PJ, etal.|doi=10.1016/j.cub.2020.08.061|journal=Current Biology|title=New Lineage of Microbial Predators Adds Complexity to Reconstructing the Evolutionary Origin of Animals|pmid=32976804 |doi-access=free}}</ref>

<ref name="Strother 2021">{{Cite journal |last1=Strother |first1=Paul K. |last2=Brasier |first2=Martin D. |last3=Wacey |first3=David |last4=Timpe |first4=Leslie |last5=Saunders |first5=Martin |last6=Wellman |first6=Charles H. |date=April 2021 |title=A possible billion-year-old holozoan with differentiated multicellularity |journal=Current Biology |volume=31 |issue=12 |pages=2658–2665.e2 |doi=10.1016/j.cub.2021.03.051|pmid=33852871 |doi-access=free }}</ref>

<ref name="Parfrey-2011">{{cite journal|author-link1=Laura Wegener Parfrey|last1=Parfrey|first1=Laura Wegener|last2=Lahr|first2=Daniel J. G.|last3=Knoll|first3=Andrew H.|last4=Katz|first4=Laura A.|author-link4=Laura A. Katz|title=Estimating the timing of early eukaryotic diversification with multigene molecular clocks|journal=Proceedings of the National Academy of Sciences of the United States of America|date=August 16, 2011|volume=108|issue=33|pages=13624&ndash;13629|doi=10.1073/pnas.1110633108|pmid=21810989|pmc=3158185|bibcode=2011PNAS..10813624P |doi-access=free}}</ref>

<ref name="Torruella-2015">{{Cite journal |title=Phylogenomics Reveals Convergent Evolution of Lifestyles in Close Relatives of Animals and Fungi|journal=Current Biology|date=21 September 2015 |issn=0960-9822 |pages=2404–2410 |volume=25 |issue=18 |doi=10.1016/j.cub.2015.07.053 |language=English|first1=Guifré|last1=Torruella|first2=Alex|last2=de Mendoza |first3=Xavier |last3=Grau-Bové |first4=Meritxell|last4=Antó|first5=Mark A. |last5=Chaplin |first6=Javier |last6=del Campo |first7=Laura|last7=Eme |first8=Gregorio |last8=Pérez-Cordón |first9=Christopher M.|last9=Whipps|pmid=26365255|doi-access=free}}</ref>

<ref name="Hehenberger-2017">{{Cite journal |last1=Hehenberger |first1=Elisabeth |last2=Tikhonenkov |first2=Denis V. |last3=Kolisko|first3=Martin|last4=Campo |first4=Javier del|last5=Esaulov|first5=Anton S.|last6=Mylnikov|first6=Alexander P. |last7=Keeling |first7=Patrick J. |title=Novel Predators Reshape Holozoan Phylogeny and Reveal the Presence of a Two-Component Signaling System in the Ancestor of Animals|journal=Current Biology |volume=27 |issue=13 |pages=2043–2050.e6 |doi=10.1016/j.cub.2017.06.006 |pmid=28648822 |year=2017 |doi-access=free}}</ref>

<ref name="Lang et al. 2002">{{cite journal|vauthors=Lang BF, O'Kelly C, Nerad T, Gray MW, Burger G|title=The Closest Unicellular Relatives of Animals|journal=Current Biology|volume=12|issue=20|date=2002|pages=1773–1778|doi=10.1016/S0960-9822(02)01187-9|pmid=12401173 |doi-access=free}}</ref>

<ref name="Brunet & King 2022">{{cite book|vauthors=Brunet T, King N|chapter=The Single-Celled Ancestors of Animals: A History of Hypotheses|title=The Evolution of Multicellularity|veditors=Herron MD, Conlin PL, Ratcliff WC|pages=251–278|doi=10.1201/9780429351907-17|series=Evolutionary Cell Biology|publisher=CRC Press|date=2022|isbn=9780429351907}}</ref>

<ref name="Handbook of the Protists 1">{{Cite book|url=https://www.springer.com/gp/book/9783319281476|title=Handbook of the Protists|date=2017|publisher=Springer International Publishing|isbn=978-3-319-28147-6|veditors=Archibald JM, ((Simpson AGB)), Slamovits CH|edition=2|pages=1–22|language=en|chapter=Chapter 1. Protist Diversity and Eukaryote Phylogeny|vauthors=((Simpson AGB)), Slamovits CH, Archibald JM|volume=1}}</ref>

<ref name="Sebé-Pedrós et al. 2017">{{cite journal|vauthors=Sebé-Pedrós A, Degnan B, Ruiz-Trillo I|title=The origin of Metazoa: a unicellular perspective|journal=Nature Reviews Genetics|volume=18|pages=498–512|date=2017|issue=8 |doi=10.1038/nrg.2017.21|pmid=28479598 |s2cid=30709486 }}</ref>

<ref name="Suga et al. 2013">{{cite journal|vauthors=Suga H, Chen Z, de Mendoza A, Sebé-Pedrós A, Brown MW, Kramer E, Carr M, Kerner P, Vervoot M, Sánchez-Pons N, Torruella G, Derelle R, Manning G, Lang BF, Russ C, Haas BJ, Roger AJ, Nusbaum C, Ruiz-Trillo I|title=The ''Capsaspora'' genome reveals a complex unicellular prehistory of animals|journal=Nature Communications|volume=4|number=2325|date=2013|page=2325 |doi=10.1038/ncomms3325|pmid=23942320 |pmc=3753549 |bibcode=2013NatCo...4.2325S |doi-access=free}}</ref>

<ref name="Txikispora">{{cite journal|vauthors=Urrutia A, Mitsi K, Foster R, Ross S, Carr M, Ward GM, etal.|date=2022|title=''Txikispora philomaios'' n. sp., n. g., a micro-eukaryotic pathogen of amphipods, reveals parasitism and hidden diversity in Class Filasterea|journal=Journal of Eukaryotic Microbiology|volume=69|issue=2 |pages=e12875|doi=10.1111/jeu.12875|pmid=34726818 |s2cid=240422937 |url=https://pure.hud.ac.uk/en/publications/1dd4e1c5-f638-48cf-ab99-95c9c90636a1 }}</ref>

<ref name="Huldtgren et al. 2011">{{cite journal|vauthors=Huldtgren T, Cunningham JA, Yin C, Stampanoni M, Marone F, ((Donoghue PCJ)), Bengtson S|date=2011|title=Fossilized Nuclei and Germination Structures Identify Ediacaran "Animal Embryos" as Encysting Protists|journal=Science|volume=334|issue=6063|pages=1696–1699|doi=10.1126/science.1209537|pmid=22194575 |bibcode=2011Sci...334Q1696H |s2cid=39813961 }}</ref>

}}

{{Eukaryota classification}}
{{Opisthokont protists}}
{{Taxonbar|from=Q1205110}}

[[Category:Holozoa| ]]
[[Category:Opisthokont unranked clades]]
[[Category:Ediacaran first appearances]]
[[Category:Taxa described in 2002]]

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