View source for Eumycetozoa

{{Short description|Taxonomic group of slime molds}}
{{About|a specific [[clade]] of slime molds|the similar group including some unrelated slime molds|Mycetozoa}}
{{Automatic taxobox|taxon=Eumycetozoa|authority=Zopf 1884 ex L.S.Olive 1975 sensu Kang et al. 2017{{efn|[[Friedrich Wilhelm Zopf|Zopf]] was the first to introduce the name Eumycetozoa, but no [[taxonomic rank|rank]] was given.<ref name="Lado_2017"/> [[species:Lindsay Shepherd Olive|Olive]] reintroduced it in 1975 as a class-level rank.<ref name="Mycetozoans"/> Finally, in 2017 Kang and his coauthors renewed the definition of Eumycetozoa into its modern usage.<ref name="BetweenPod"/>}}|synonyms=
*Macromycetozoa {{au|Fiore-Donno et al. 2010}}
*Mycetozoa {{au|de Bary, 1859 ex Rostafinski, 1873}}<ref name="Cutosea"/> (Mycetozoa ''[[sensu stricto]]'')<ref name="Leontyev_2017"/>
|subdivision_ranks = Clades<ref name="Adl 2019"/>
|subdivision=
* [[Dictyostelia]]
* [[Myxogastria]]
* [[Protosporangiida]]
| image = Fuligo septica - Gelbe Lohblüte - Hexenbutter - 02.jpg
| image_caption = ''[[Fuligo septica]]'', a [[myxogastria|myxogastrid]]
}}
'''Eumycetozoa''' ({{etymology|grc|εὖ (eû)|true||μύκης (múkēs)|fungus||ζῷον (zôion)|animal}}), or '''true slime molds''',<ref name="Wijayawardene et al. 2020"/> is a diverse group of [[protist]]s that behave as [[slime mold]]s and develop [[fruiting bodies]], either as [[sorocarp]]s or as [[sporocarp (fungi)|sporocarps]]. It is a [[monophyletic]] group or [[clade]] within the phylum [[Amoebozoa]] that contains the [[Myxogastria|myxogastrids]], [[dictyostelid]]s and [[Protosporangiida|protosporangiids]].

== Characteristics ==
Eumycetozoa is a [[clade]] that includes three groups of amoebozoan protists: [[Myxogastria]], [[Dictyostelia]] and [[Protosporangiida]]—also known as Myxomycetes, Dictyosteliomycetes and Ceratiomyxomycetes, respectively.<ref name="Myxomycetes"/> It is defined on a [[Phylogenetic nomenclature#Phylogenetic definitions of clade names|node-based]] approach as the least inclusive clade containing the species ''[[Dictyostelium discoideum]]'' (a dictyostelid), ''[[Physarum polycephalum]]'' (a myxogastrid) and ''[[Ceratiomyxa fruticulosa]]'' (a protosporangiid).<ref name="BetweenPod"/><ref name="Adl 2019"/>

All known members of Eumycetozoa generate [[fruiting bodies]], either as [[sorocarp]]s (in dictyostelids) or as [[sporocarp (fungi)|sporocarps]] (in myxogastrids and protosporangiids). Within their [[biological life cycle|life cycle]], they may appear as a single [[haploid]] [[amoeboid]] cells (in dictyostelids), or as [[flagellate]]d amoebae with two [[cilia]] that give rise to obligate amoebae with no cilia, from which the sporocarps develop (in myxogastrids and protosporangiids).<ref name="Adl 2019"/>

The flagellated amoebae of myxogastrids and protosporangiids and non-flagellated amoebae of dictyostelids have a flat cell shape. They form wide [[pseudopodia]] with acutely pointed subpseudopodia (i.e. smaller pseudopodia that grow beneath). Unlike other amoebae, the pseudopodia lack a prominent streaming of granular [[cytoplasm]].<ref name="Adl 2019"/>

In eumycetozoans where [[sexual reproduction]] is well studied, the [[zygote]] cannibalizes on haploid amoebae.<ref name="Adl 2019"/>

== Evolution ==
Eumycetozoa is a [[robust statistics|well supported]] [[clade]] within Amoebozoa. In independent [[phylogenetic analyses]], it has been consistently recovered as the [[sister group]] to [[Archamoebae]]. The Eumycetozoa+Archamoebae clade is, in turn, the sister group to [[Variosea]]. Within Eumycetozoa, [[Dictyostelia]] has a [[basal (phylogeny)|basal]] position while [[Myxogastria]] and [[Protosporangiida]] form a clade. Together, these three groups are part of the larger clade [[Conosa]].<ref name="BetweenPod"/> The following cladogram is based on a 2022 analysis:<ref name="Divosa"/>
{{clade|style=font-size:90%;|label1=[[Amoebozoa]]|1={{clade|1=[[Tubulinea]] [[File:Amoeba_proteus_with_many_pseudopodia.jpg|50px]]|2={{clade|1=[[Discosea]] [[File:Parasite140120-fig3_Acanthamoeba_keratitis_Figure_3B.png|50px]]|label2=[[Evosea]]|2={{clade|1=[[Cutosea]]|label2=[[Conosa]]|2={{clade|2=[[Variosea]] [[File:Protostelium mycophaga fruiting body.jpg|35px]]|1={{clade|1=[[Archamoebae]] [[File:Pelomyxa_palustris.jpg|50px]]|label2='''Eumycetozoa'''|2={{clade|1=[[Dictyostelia]] [[File:Dictyostelium_Aggregation.JPG|50px]]|2={{clade|1=[[Myxogastria]] [[File:Physarum album 252229345.jpg|50px]]|2=[[Protosporangiida]] [[File:2014-04-12 Ceratiomyxa fruticulosa (Müll.) Mac 414773.jpg|50px]]}}}}}}}}}}}}}}}}

== Taxonomy ==
The name '''Eumycetozoa''' was first used by German [[mycologist]] [[Friedrich Wilhelm Zopf]] in 1884, although no formal [[taxonomic rank]] was given.<ref name="Lado_2017"/> In 1975, mycologist [[species:Lindsay Shepherd Olive|Lindsay Shepherd Olive]] reintroduced the name Eumycetozoa as a [[class (biology)|class]] containing the three groups of fruiting amoebae traditionally included in this taxon: [[Myxogastria]], [[Dictyostelia]] and [[Protostelia]].<ref name="Mycetozoans"/> Olive hypothesized that all fruiting amoebae were grouped by this monophyletic taxon, and that the Myxogastria and Dictyostelia were also monophyletic taxa that evolved from a paraphyletic [[evolutionary grade|grade]] of Protostelia. This definition of Eumycetozoa, which included protostelids, was maintained in the 2005 [[cladistic]] classification of eukaryotes, where the name was synonymized with [[Mycetozoa]].<ref name="Adl_2005"/>

* [[Amoebozoa]] {{au|Lühe, 1913, emend. Cavalier-Smith, 1998}}
** '''Eumycetozoa''' {{au|Zopf, 1884, emend. L.S.Olive, 1975}} [=Mycetozoa {{au|de Bary, 1873}}]
*** [[Protostelia]] {{au|L.S. Olive, 1975}} ([[paraphyletic|P]])
*** [[Myxogastria]] {{au|Macbride, 1899}} [=Myxomycetes {{au|Link, 1833, emend. Haeckel, 1866}}]
*** [[Dictyostelia]] {{au|Lister, 1909, emend. L.S.Olive, 1970}}
*** [[Incertae sedis]] Eumycetozoa: ''[[Copromyxa]]'', ''[[Copromyxella]]'', ''[[Fonticula]]''

However, studies in the 2000s decade disproved this hypothesis. Both [[morphology (biology)|morphological]] and [[phylogeny|molecular]] studies showed that Eumycetozoa includes a number of non-fruiting amoeboid groups. More importantly, the [[Protostelia]] were discovered to be [[polyphyletic]]. The protosteloid type of fruiting body formation, initially considered the [[synapomorphy|ancestral feature]] shared between all Eumycetozoa, has [[homoplasy|evolved independently]] at least in eight lineages within [[Amoebozoa]] (e.g. soliformoviids, cavosteliids, schizoplasmodiids, protosporangiids).<ref name="Shadwick_2009"/><ref name="Leontyev_2017"/> This discovery lead to the conclusion that the entirety of Amoebozoa became a [[synonym (taxonomy)|synonym]] of Eumycetozoa, and was treated as such in the 2012 cladistic classification of eukaryotes. The term Amoebozoa was conserved as a familiar well-established name of popular usage, despite the term Eumycetozoa having priority as the older name.<ref name="Adl_2012"/>

To preserve this widely used name, biologist Seungho Kang and his coauthors redefined Eumycetozoa in 2017 to include only one group of protosteloid amoebae, the [[Protosporangiida]] (also known as Ceratiomyxomycetes), which are a monophyletic taxon.<ref name="BetweenPod"/><ref name="Wijayawardene et al. 2020">{{cite journal |last1=Wijayawardene |first1=Nalin |last2=Hyde |first2=Kevin |first3=LKT |last3=Al-Ani |first4=S |last4=Dolatabadi |last5=Stadler |first5=Marc |last6=Haelewaters |first6=Danny |last7=Tsurykau |first7=Andrei |last8=Mesic |first8=Armin |last9=Navathe |first9=Sudhir |last10=Papp |first10=Viktor |last11=Oliveira Fiuza |first11=Patrícia |last12=Vázquez |first12=Víctor |last13=Gautam |first13=Ajay |last14=Becerra |first14=Alejandra G. |last15=Ekanayaka |first15=Anusha |last16=K. C. |first16=Rajeshkumar |last17=Bezerra |first17=Jadson |last18=Matočec |first18=Neven |last19=Maharachchikumbura |first19=Sajeewa |last20=Suetrong |first20=Satinee |year=2020 |title=Outline of Fungi and fungus-like taxa |journal=Mycosphere |volume=11 |pages=1060–1456 |doi=10.5943/mycosphere/11/1/8 |doi-access=free|hdl=10481/61998 |hdl-access=free }}</ref> This usage corresponds to the 1975 hypothesis from Olive that postulates a [[clade]] of exclusively fruiting protists that includes myxogastrids, dictyostelids, and some protosteloid amoebae (in this case, the protosporangiids).<ref name="BetweenPod"/> As of 2019, this renewed definition is accepted by the scientific community and appears in the modern cladistic classification of eukaryotes, revised by the International Society of Protistologists.<ref name="Adl 2019"/> The name '''Macromycetozoa''' was suggested earlier,<ref name="Fiore-Donno_2010"/> but Eumycetozoa was chosen for being the oldest term.<ref name="BetweenPod"/>

* [[Amoebozoa]] {{au|Lühe, 1913, emend. [[Cavalier-Smith]], 1998}}
** [[Evosea]] {{au|Kang et al. 2017}}
*** '''Eumycetozoa''' {{au|Zopf 1884 ex L.S. Olive, sensu Kang et al. 2017}} [=Macromycetozoa {{au|Fiore-Donno et al. 2010}}]
**** [[Dictyostelia]] {{au|Lister 1909, sensu Sheikh et al. 2018}}
**** [[Myxogastria]] {{au|Macbride 1899}} [=Myxomycetes {{au|Link 1833, sensu [[Haeckel]] 1866}}]
**** [[Protosporangiida]] {{au|Shadwick & Spiegel in Adl et al. 2012}}

The name [[Mycetozoa]] was maintained in traditional classifications by some authors like [[Thomas Cavalier-Smith]], who also used a renewed definition to include only protosporangiids. However this scheme did not acquire wide usage.<ref name="Cutosea"/>

== Notes ==
{{notelist}}

== References ==
{{Reflist|refs=

<ref name="BetweenPod">{{cite journal|first1=Seungho|last1=Kang|first2=Alexander K |last2=Tice|first3=Frederick W |last3=Spiegel|first4=Jeffrey D |last4=Silberman|first5=Tomáš |last5=Pánek|first6=Ivan |last6=Čepička|first7=Martin |last7=Kostka|first8=Anush |last8=Kosakyan|first9=Daniel M C |last9=Alcântara|first10=Andrew J |last10=Roger|first11=Lora L |last11=Shadwick|first12=Alexey |last12=Smirnov|first13=Alexander |last13=Kudryavtsev|first14=Daniel J G |last14=Lahr|first15=Matthew W |last15=Brown|title=Between a Pod and a Hard Test: The Deep Evolution of Amoebae|journal=Molecular Biology and Evolution|volume=34|issue=9|date=September 2017|pages=2258–2270|pmid=28505375|pmc=5850466|doi=10.1093/molbev/msx162}}</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="Myxomycetes">{{cite journal|title=Towards a phylogenetic classification of the Myxomycetes|first1=Dmitry V|last1=Leontyev|first2=Martin|last2=Schnittler|first3=Steven L|last3=Stephenson|first4=Yuri K|last4=Novozhilov|first5=Oleg N|last5=Shchepin|doi=10.11646/phytotaxa.399.3.5|volume=399|issue=3|date=March 2019|journal=Phytotaxa|page=209 |s2cid=108783142 |doi-access=free }}</ref>

<ref name="Divosa">{{cite journal|vauthors=Tekle YI, Wang F, Wood FC, Anderson OR, Smirnov A|title=New insights on the evolutionary relationships between the major lineages of Amoebozoa|journal=Sci Rep|volume=12|number=11173|date=2022|page=11173 |doi=10.1038/s41598-022-15372-7|pmid=35778543 |pmc=9249873 |bibcode=2022NatSR..1211173T |s2cid=247231712 }}</ref>

<ref name="Shadwick_2009">{{cite journal|title=''Eumycetozoa'' = ''Amoebozoa''?: SSUrDNA Phylogeny of Protosteloid Slime Molds and Its Significance for the Amoebozoan Supergroup|first1=Lora L.|last1=Shadwick|first2=Frederick W.|last2=Spiegel|first3=John D. L.|last3=Shadwick|first4=Matthew W.|last4=Brown|first5=Jeffrey D.|last5=Silberman|date=2009|journal=PLOS ONE|volume=4|issue=8|pages=e6754|doi=10.1371/journal.pone.0006754|doi-access=free}}</ref>

<ref name="Lado_2017">{{cite book|chapter=Taxonomy and Systematics: Current Knowledge and Approaches on the Taxonomic Treatment of Myxomycetes|first1=Carlos|last1=Lado|first2=Uno|last2=Eliasson|doi=10.1016/B978-0-12-805089-7.00007-X|title=Myxomycetes: Biology, Systematics, Biogeography, and Ecology|editor-first1=Steven L.|editor-last1=Stephenson|editor-first2=Carlos|editor-last2=Rojas|publisher=Academic Press, Elsevier|isbn=978-0-12-805089-7|date=2017}}</ref>

<ref name="Leontyev_2017">{{cite book|chapter=The Phylogeny of Myxomycetes|first1=Dmitry V.|last1=Leontyev|first2=Martin|last2=Schnittler|doi=10.1016/B978-0-12-805089-7.00003-2|title=Myxomycetes: Biology, Systematics, Biogeography, and Ecology|editor-first1=Steven L.|editor-last1=Stephenson|editor-first2=Carlos|editor-last2=Rojas|publisher=Academic Press, Elsevier|isbn=978-0-12-805089-7|date=2017}}</ref>

<ref name="Mycetozoans">{{cite book|vauthors=Olive LS|date=1975|title=The Mycetozoans|publisher=Academic Press, Elsevier|isbn=978-0125262507}}</ref>

<ref name="Adl_2005">{{cite journal  |vauthors=Adl SM, Simpson AG, Farmer MA, etal |title=The new higher level classification of eukaryotes with emphasis on the taxonomy of protists |journal=The Journal of Eukaryotic Microbiology |volume=52 |issue=5 |pages=399–451 |year=2005 |pmid=16248873 |doi=10.1111/j.1550-7408.2005.00053.x|doi-access=free }}</ref>

<ref name="Adl_2012">{{cite journal | vauthors = Adl SM, Simpson AG, Lane CE, Lukeš J, Bass D, Bowser SS, Brown MW, Burki F, Dunthorn M, Hampl V, Heiss A, Hoppenrath M, Lara E, Le Gall L, Lynn DH, McManus H, Mitchell EA, Mozley-Stanridge SE, Parfrey LW, Pawlowski J, Rueckert S, Shadwick L, Shadwick L, Schoch CL, Smirnov A, Spiegel FW | title = The revised classification of eukaryotes | journal = The Journal of Eukaryotic Microbiology | volume = 59 | issue = 5 | pages = 429–93 | date = September 2012 | pmid = 23020233 | pmc = 3483872 | doi = 10.1111/j.1550-7408.2012.00644.x }}</ref>

<ref name="Fiore-Donno_2010">{{cite journal|first1=Anna Maria|last1=Fiore-Donno|first2=Sergey I.|last2=Nikolaev|first3=Michaela|last3=Nelson|first4=Jan|last4=Pawlowski|first5=Thomas|last5=Cavalier-Smith|first6=Sandra L|last6=Baldauf|date=January 2010|title=Deep phylogeny and evolution of slime moulds (Mycetozoa)|journal=Protist|volume=161|issue=1|pages=55–70|doi=10.1016/j.protis.2009.05.002}}</ref>

<ref name="Cutosea">{{cite journal | vauthors = Cavalier-Smith T, Chao EE, Lewis R | title = 187-gene phylogeny of protozoan phylum Amoebozoa reveals a new class (Cutosea) of deep-branching, ultrastructurally unique, enveloped marine Lobosa and clarifies amoeba evolution | journal = Molecular Phylogenetics and Evolution | volume = 99 | pages = 275–296 | date = June 2016 | pmid = 27001604 | doi = 10.1016/j.ympev.2016.03.023 | doi-access = free }}</ref>

}}
== External links ==
* [https://eukmap.unieuk.net/taxonomy/110 Eumycetozoa] at UniEuk Taxonomy App.
{{Eukaryota}}
{{taxonbar|from=Q25448563}}
[[Category:Eumycetozoa| ]]
[[Category:Protista]]
[[Category:Taxa described in 1884]]