Myxozoa

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Myxozoa
Triactinomyxon stage of Myxobolus cerebralis
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Cnidaria
Subphylum: Myxozoa
Grassé
, 1970
Classes

Myxozoa (

parasites. It contains the smallest animals ever known to have lived. Over 2,180 species have been described and some estimates have suggested at least 30,000 undiscovered species.[3] Many have a two-host lifecycle, involving a fish and an annelid worm or a bryozoan. The average size of a myxosporean spore usually ranges from 10 μm to 20 μm,[4]
whereas that of a malacosporean (a subclade of the Myxozoa) spore can be up to 2 mm. Myxozoans can live in both freshwater and marine habitats.

Myxozoans are highly

aerobic respiration. The genomes of some myxozoans are now among the smallest genomes of any known animal species.[5][6]

Life cycle and pathology

Myxozoans are endoparasitic animals exhibiting complex life cycles that, in most of the documented cases, involve an

ectoproct
.

Myxozoans Life Cycle

Only about 100 life cycles have been resolved and it is suspected that there may be some exclusively terrestrial.

sporoplasms to penetrate into the epithelium. Subsequently, the parasite undergoes reproduction and development in the gut tissue, and finally produces usually eight actinosporean spore stages (actinospores) within a pansporocyst. After mature actinospores are released from their hosts they float in the water column.[13] Upon contact with skin or gills of fish, sporoplasms penetrate through the epithelium, followed by development of the myxosporean stage. Myxosporean trophozoites are characterized by cell-in-cell state, where the secondary (daughter) cells develop in the mother (primary) cells. The presporogonic stages multiply, migrate via nervous or circulatory systems, and develop into sporogonic stages. At the final site of infection, they produce mature spores within mono- or di-sporic pseudoplasmodia, or poly-sporic plasmodia.[14]

Relationships between myxosporeans and their hosts are often highly evolved and do not usually result in severe diseases of the natural host. Infection in

parasites can be severe, especially where prevalence
rates are high; they may also have a severe impact on wild fish stocks.

The diseases caused by myxosporeas in cultured fish with the most significant economic impact worldwide are

common carp
.

Anatomy

Myxozoans are very small animals, typically 10–300 

μm in length.[15]

Like other cnidarians they possess cnidocysts, which were referred to as "polar capsules" before the discovery that myxozoans are cnidarians. These cnidocysts fire tubules as in other cnidarians; some inject substances into the host. However, the tubules lack hooks or barbs, and in some species are more elastic than in other cnidarians.

Myxozoans have secondarily lost

binary fission is rare, and cells divide instead via endogeny.[15]

In 2020, the myxozoan

aerobic respiration; it was the first animal to be positively identified as such. Its actual metabolism is currently unknown.[17]

Phylogenetics

Myxozoans were originally considered to be

metazoa. Detailed classification within the metazoa was however long hindered by conflicting rDNA evidence: although 18S rDNA suggested an affinity with Cnidaria,[20] other rDNA sampled,[21][22] and the HOX genes of two species,[23] were more similar to those of the Bilateria
.

The discovery that

bryozoans up to 2 mm in length, is a myxozoan[21] initially appeared to strengthen the case for a bilaterian origin, as the body plan is superficially similar. Nevertheless, closer examination reveals that Buddenbrockia's longitudinal symmetry is not twofold, but fourfold, casting doubt on this hypothesis
.

Further testing resolved the genetic conundrum by sourcing the first three previously identified discrepant HOX genes (Myx1-3) to the

nematocysts had been drawn for a long time, but were generally assumed to be the result of convergent evolution
.

Taxonomists now recognize the outdated subgroup Actinosporea as a life-cycle phase of Myxosporea.[25]

Molecular clocks suggest that myxozoans and their closest relatives, the

polypodiozoa, shared their last common ancestor with medusazoans about 600 million years ago, during the Ediacaran period.[3]

Taxonomy

Myxozoan taxonomy has undergone great and important changes in its levels of generic, family and suborder classification. Fiala et al. (2015) proposed a new classification based on spores.[26]

Phylum: Cnidaria
Subphylum: Myxozoa[26]
Class:
Malacosporea
Class: Myxosporea (sequel)
Order: Malacovalvulida Order: Bivalvulida (sequel)
   Family: Saccosporidae    Family:
Myxobilatidae
     Genus:
Buddenbrockia, Tetracapsuloides
     Genus: Myxobilatus, Acauda, Hoferellus
Class: Myxosporea    Family:
Chloromyxidae
Order: Bivalvulida      Genus: Chloromyxum, Caudomyxum, Agarella
  Suborder: Variisporina    Family:
Coccomyxidae
   Family:
Sphaeromyxidae
     Genus: Coccomyxa, Auerbachia, Globospora
     Genus: Sphaeromyxa    Family: Alatosporidae
   Family: Myxidiidae      Genus: Alatospora, Pseudalatospora, Renispora
     Genus: Myxidium, Zschokkella, Enteromyxum, Sigmomyxa, Soricimyxum, Cystodiscus    Family: Parvicapsulidae
   Family:
Ortholineidae
     Genus: Parvicapsula, Neoparvicapsula, Gadimyxa
     Genus: Ortholinea, Neomyxobolus, Cardimyxobolus, Triangula, Kentmoseria   Suborder:
Platysporina
   Family:
Sinuolineidae
   Family: Myxobolidae
     Genus: Sinuolinea, Myxodavisia, Myxoproteus, Bipteria, Paramyxoproteus, Neobipteria, Schulmania, Noblea, Latyspora      Genus: Myxobolus, Spirosuturia, Unicauda, Dicauda, Phlogospora, Laterocaudata, Henneguya, Hennegoides, Tetrauronema, Thelohanellus, Neothelohanellus, Neohenneguya, Trigonosporus
   Family:
Fabesporidae
Order: Multivalvulida
     Genus: Fabespora    Family: Trilosporidae
   Family: Ceratomyxidae      Genus: Trilospora, Unicapsula
     Genus: Ceratomyxa, Meglitschia, Ellipsomyxa, Ceratonova    Family:
Kudoidae
   Family:
Sphaerosporidae
     Genus: Kudoa
     Genus:
Sphaerospora, Wardia, Palliatus
   Family: Spinavaculidae
     Genus: Octospina
Incertae sedis in Multivalvulida: Trilosporoides
 
  Phylum:  
  Cnidaria  
  Subphylum:  
  Myxozoa  
[26]
  Class:  
  Malacosporea  
Order:
   Malacovalvulida  
Family:
   Saccosporidae  

Genus: Buddenbrockia, Tetracapsuloides

  Class:  
  Myxosporea  
Order:
   Bivalvulida  
Suborder:
   Variisporina  
Family:
   Sphaeromyxidae  

Genus: Sphaeromyxa

Family:
   Myxidiidae  

Genus:
Myxidium, Zschokkella, Enteromyxum, Sigmomyxa, Soricimyxum, Cystodiscus

Family:
   Ortholineidae  

Genus:
Ortholinea, Neomyxobolus, Cardimyxobolus, Triangula, Kentmoseria

Family:
   Sinuolineidae  

Genus:
Sinuolinea, Myxodavisia, Myxoproteus, Bipteria, Paramyxoproteus, Neobipteria, Schulmania, Noblea, Latyspora

Family:
   Fabesporidae  

Genus: Fabespora

Family:
   Ceratomyxidae  

Genus:
Ceratomyxa, Meglitschia, Ellipsomyxa, Ceratonova

Family:
   Sphaerosporidae  

Genus:
Sphaerospora, Wardia, Palliatus

Family:
   Myxobilatidae  

Genus:
Myxobilatus, Acauda, Hoferellus

Family:
   Chloromyxidae  

Genus:
Chloromyxum, Caudomyxum, Agarella

Family:
   Coccomyxidae  

Genus:
Coccomyxa, Auerbachia, Globospora

Family:
   Alatosporidae  

Genus:
Alatospora, Pseudalatospora, Renispora

Family:
   Parvicapsulidae  

Genus:
Parvicapsula, Neoparvicapsula, Gadimyxa

Suborder:
   Platysporina  
Family:
   Myxobolidae  

Genus:
Myxobolus, Spirosuturia, Unicauda, Dicauda, Phlogospora, Laterocaudata, Henneguya, Hennegoides, Tetrauronema, Thelohanellus, Neothelohanellus, Neohenneguya, Trigonosporus

Order:
   Multivalvulida  
Family:
   Trilosporidae  

Genus: Trilospora, Unicapsula

Family:
   Kudoidae  

Genus: Kudoa

Family:
   Spinavaculidae  

Genus: Octospina

  incertae sedis:       Trilosporoides  

     other Cnidarians     

    . . .   
 

See also

References

  1. Perseus Project
  2. Perseus Project
  3. ^
    S2CID 52141614
    .
  4. ^ Fiala, Ivan (10 July 2008). "Myxozoa". tolweb.org (under construction). The Tree of Life Web Project.
  5. PMID 26627241
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  13. .
  14. .
  15. ^ .
  16. .
  17. .
  18. ^ Štolc, A. (1899). "Actinomyxidies, nouveau groupe de Mesozoaires parent des Myxosporidies". Bull. Int. l'Acad. Sci. Bohème. 12: 1–12.
  19. ^ Edwin Lanfranco, 2007, A phylogenetic classification of organisms other than animals.
  20. ^ Smothers, J.F.; et al. (September 1994). "Molecular evidence that the myxozoan protists are metazoans".
    PMID 8085160
    .
  21. ^ a b A.S. Monteiro; et al. (1 June 2002). "Orphan worm finds a home: Buddenbrockia is a Myxozoan". Mol. Biol. Evol. 19 (6): 968–71.
    PMID 12032254
    .
  22. ^ J. Zrzavy & V. Hypsa (April 2003). "Myxozoa, Polypodium, and the origin of the Bilateria: The phylogenetic position of "Endocnidozoa" in light of the rediscovery of Buddenbrockia". Cladistics. 19 (2): 164–169.
    S2CID 221583517
    .
  23. ^ C. L. Anderson, E. U. Canning & B. Okamura (March 1999). "A triploblast origin for Myxozoa?".
    S2CID 4426181
    .
  24. ^ a b E. Jímenez-Guri; et al. (July 2007). "Buddenbrockia is a cnidarian worm".
    S2CID 5170702
    .
  25. .
  26. ^ .

External links