ParaHoxozoa

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ParaHoxozoa
Temporal range: 605.2 –0  Ma
Animalia diversity.jpg
Diversity of parahoxozoans
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Subkingdom: Eumetazoa
Clade: ParaHoxozoa
Ryan et al., 2010
Taxa

ParaHoxozoa (or Parahoxozoa) is a clade of animals that consists of Bilateria, Placozoa, and Cnidaria. [1] The relationship of this clade relative to the two other animal lineages Ctenophora and Porifera is debated. Some phylogenomic studies have presented evidence supporting Ctenophora as the sister to Parahoxozoa and Porifera as the sister group to the rest of animals (e.g. [2] [3] [4] ). Other studies have presented evidence supporting Porifera as the sister to Parahoxozoa and Ctenophora as the sister group to the rest of animals (e.g. [5] [6] [7] ), finding that nervous systems either evolved independently in ctenophores and parahoxozoans, [8] or were secondarily lost in poriferans. [9] If ctenophores are taken to have diverged first, Eumetazoa is sometimes used as a synonym for ParaHoxozoa. [10]

Contents

Phylogeny

The tree below, which is congruent with the vast majority of these phylogenomic studies, conveys this uncertainty with a polytomy.

   Choanozoa   

  Choanoflagellata Cronoflagelado2.svg

   Animalia   

  Ctenophora Mertensia ovum.png

  Porifera Reef3859 - Flickr - NOAA Photo Library.jpg

  Parahoxozoa  

  Placozoa Trichoplax adhaerens photograph.png

   Planulozoa   
     

  Cnidaria Medusae of world-vol03 fig360 Atolla chuni.jpg

  Bilateria Acrodipsas brisbanensis.jpg

  
  
  
  

ParaHoxozoa or Parahoxozoa

Though "ParaHox" genes are usually referred to in CamelCase and the original paper that named the clade used "ParaHoxozoa", the single initial capital format "Parahoxozoa" has also come to be used in the literature, [11] as CamelCase is not standard in zoological nomenclature.[ citation needed ]

Characteristics

Parahoxozoa was defined by the presence of several gene (sub)classes (HNF, CUT, PROS, ZF, CERS, K50, S50-PRD), as well as Hox/ParaHox-ANTP from which the name of this clade originated. It was later proposed [12] [13] and contested [14] that a gene of the same class (ANTP) as the Hox/ParaHox, the NK gene and the Cdx Parahox gene, is also present in Porifera, the sponges. Regardless of whether a ParaHox gene is ever definitively identified, Parahoxozoa, as originally defined, is monophyletic and therefore continues to be used as such. [15]

Planula-acoel, triploblasty, and bilaterian similarities

The original Bilateria are hypothesized to be a bottom dwelling worm with a single body opening. [16] A through-gut may already have developed with the Ctenophora however. [17] The through-gut may have developed from the corners of a single opening with lips fusing. E.g. Acoela resemble the planula larvae of some Cnidaria, which exhibit some bilaterian symmetry. They are vermiform, just as the cnidarian Buddenbrockia is. [18] [19] [20] Placozoans have been noted to resemble planula. [21] Usually, "Planulozoa" refers to a Cnidaria–Bilateria clade to the exclusion of Placozoa, but not necessarily. [11] In the other case, when including all three lineages, it appears synonymous with Parahoxozoa. [22] Triploblasty developed before the Cnidaria–Bilateria radiation as well. [23]

ParaHoxozoa/Parahoxozoa taxonomy

The Parahoxozoa has 3 different types of phyla that includes placozoans, cnidarians, and bilaterians.

Placozoans

These are placozoans (e.g. Polyplacotoma mediterranea, Trichoplax adhaerens, Hoilungia hongkongensis, and Cladtertia collaboinventa) are a phylum of this tiny creature.

Cnidarians

Cnidarians (e.g. Jellyfish, Coral, Sea anemone, Stalked jellyfish, Hydra, Box jellyfish, and Sea fans) have stinging cells called cnidocytes.

Bilaterians

Many species of bilterians (e.g. Eels, Flounders, Common dab, Sole (fish), Pufferfish, Stingray, Mantaray,

Related Research Articles

<span class="mw-page-title-main">Cnidaria</span> Aquatic animal phylum having cnydocytes

Cnidaria, is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, including jellyfish, hydroids, sea anemones, corals and some of the smallest marine parasites. Their distinguishing features are a decentralized nervous system distributed throughout a gelatinous body and the presence of cnidocytes or cnidoblasts, specialized cells with ejectable flagella used mainly for envenomation and capturing prey. Their bodies consist of mesoglea, a non-living, jelly-like substance, sandwiched between two layers of epithelium that are mostly one cell thick. Cnidarians are also some of the only animals that can reproduce both sexually and asexually.

<span class="mw-page-title-main">Placozoa</span> Basal form of free-living invertebrate

Placozoa is a phylum of marine and free-living (non-parasitic) animals. They are simple blob-like animals without any body part or organ, and are merely aggregates of cells. Moving in water by ciliary motion, eating food by engulfment, reproducing by fission or budding, placozoans are described as "the simplest animals on Earth." Structural and molecular analyses have supported them as among the most basal animals, thus, constituting the most primitive metazoan phylum.

<span class="mw-page-title-main">Sponge</span> Animals of the phylum Porifera

Sponges, the members of the phylum Porifera, are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate through them, consisting of jelly-like mesohyl sandwiched between two thin layers of cells.

<span class="mw-page-title-main">Bilateria</span> Animals with embryonic bilateral symmetry

Bilateria is a large clade or infrakingdom of animals called bilaterians, characterized by bilateral symmetry during embryonic development. This means their body plans are laid around a longitudinal axis with a front and a rear end, as well as a left–right–symmetrical belly (ventral) and back (dorsal) surface. Nearly all bilaterians maintain a bilaterally symmetrical body as adults; the most notable exception is the echinoderms, which extend to pentaradial symmetry as adults, but are only bilaterally symmetrical as an embryo. Cephalization is also a characteristic feature among most bilaterians, where the special sense organs and central nerve ganglia become concentrated at the front/rostral end.

<span class="mw-page-title-main">Ctenophora</span> Phylum of gelatinous marine animals

Ctenophora comprise a phylum of marine invertebrates, commonly known as comb jellies, that inhabit sea waters worldwide. They are notable for the groups of cilia they use for swimming, and they are the largest animals to swim with the help of cilia.

<span class="mw-page-title-main">Parazoa</span> Ancestral subkingdom of animals

Parazoa are a taxon with sub-kingdom category that is located at the base of the phylogenetic tree of the animal kingdom in opposition to the sub-kingdom Eumetazoa; they group together the most primitive forms, characterized by not having proper tissues or that, in any case, these tissues are only partially differentiated. They generally group a single phylum, Porifera, which lack muscles, nerves and internal organs, which in many cases resembles a cell colony rather than a multicellular organism itself. All other animals are eumetazoans, which do have differentiated tissues.

<span class="mw-page-title-main">Myxozoa</span> Group of marine parasites

Myxozoa is a subphylum of aquatic cnidarian animals – all obligate 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. 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, whereas that of a malacosporean spore can be up to 2 mm. Myxozoans can live in both freshwater and marine habitats.

<span class="mw-page-title-main">Eumetazoa</span> Basal animal clade as a sister group of the Porifera

Eumetazoa, also known as diploblasts, Epitheliozoa or Histozoa, are a proposed basal animal clade as a sister group of Porifera (sponges). The basal eumetazoan clades are the Ctenophora and the ParaHoxozoa. Placozoa is now also seen as a eumetazoan in the ParaHoxozoa. The competing hypothesis is the Myriazoa clade.

<span class="mw-page-title-main">Nerve net</span> Nervous systems lacking a brain

A nerve net consists of interconnected neurons lacking a brain or any form of cephalization. While organisms with bilateral body symmetry are normally associated with a condensation of neurons or, in more advanced forms, a central nervous system, organisms with radial symmetry are associated with nerve nets, and are found in members of the Ctenophora, Cnidaria, and Echinodermata phyla, all of which are found in marine environments. In the Xenacoelomorpha, a phylum of bilaterally symmetrical animals, members of the subphylum Xenoturbellida also possess a nerve net. Nerve nets can provide animals with the ability to sense objects through the use of the sensory neurons within the nerve net.

<i>Trichoplax</i> Genus of Placozoa

Trichoplax adhaerens is one of the four named species in the phylum Placozoa. The others are Hoilungia hongkongensis, Polyplacotoma mediterranea and Cladtertia collaboinventa. Placozoa is a basal group of multicellular animals, possible relatives of Cnidaria. Trichoplax are very flat organisms commonly less than 4 mm in diameter, lacking any organs or internal structures. They have two cellular layers: the top epitheloid layer is made of ciliated "cover cells" flattened toward the outside of the organism, and the bottom layer is made up of cylinder cells that possess cilia used in locomotion, and gland cells that lack cilia. Between these layers is the fibre syncytium, a liquid-filled cavity strutted open by star-like fibres.

<span class="mw-page-title-main">Radiata</span> Taxonomic rank that has been used to classify radially symmetric animals

Radiata or Radiates is a historical taxonomic rank that was used to classify animals with radially symmetric body plans. The term Radiata is no longer accepted, as it united several different groupings of animals that do not form a monophyletic group under current views of animal phylogeny. The similarities once offered in justification of the taxon, such as radial symmetry, are now taken to be the result of either incorrect evaluations by early researchers or convergent evolution, rather than an indication of a common ancestor. Because of this, the term is used mostly in a historical context.

<span class="mw-page-title-main">Symmetry in biology</span> Geometric symmetry in living beings

Symmetry in biology refers to the symmetry observed in organisms, including plants, animals, fungi, and bacteria. External symmetry can be easily seen by just looking at an organism. For example, the face of a human being has a plane of symmetry down its centre, or a pine cone displays a clear symmetrical spiral pattern. Internal features can also show symmetry, for example the tubes in the human body which are cylindrical and have several planes of symmetry.

<span class="mw-page-title-main">Coelenterata</span> Term encompassing animal phyla Cnidaria and Ctenophora

Coelenterata is a term encompassing the animal phyla Cnidaria and Ctenophora. The name comes from Ancient Greek κοῖλος (koîlos) 'hollow', and ἔντερον (énteron) 'intestine', referring to the hollow body cavity common to these two phyla. They have very simple tissue organization, with only two layers of cells along with a middle undifferentiated layer called mesoglea. , and radial symmetry. Some examples are corals, which are typically colonial, and hydrae, jellyfish, and sea anemones, Aurelia which are solitary.And pennatula, Portugese man of war, gorgonia and physalia . Coelenterata lack a specialized circulatory system relying instead on diffusion across the tissue layers.

The ParaHox gene cluster is an array of homeobox genes from the Gsx, Xlox (Pdx) and Cdx gene families.

<span class="mw-page-title-main">Medusozoa</span> Clade of marine invertebrates

Medusozoa is a clade in the phylum Cnidaria, and is often considered a subphylum. It includes the classes Hydrozoa, Scyphozoa, Staurozoa and Cubozoa, and possibly the parasitic Polypodiozoa. Medusozoans are distinguished by having a medusa stage in their often complex life cycle, a medusa typically being an umbrella-shaped body with stinging tentacles around the edge. With the exception of some Hydrozoa, all are called jellyfish in their free-swimming medusa phase.

<i>Mnemiopsis</i> Genus of comb jellies

Mnemiopsis leidyi, the warty comb jelly or sea walnut, is a species of tentaculate ctenophore. It is native to western Atlantic coastal waters, but has become established as an invasive species in European and western Asian regions. Three species have been named in the genus Mnemiopsis, but they are now believed to be different ecological forms of a single species M. leidyi by most zoologists.

<span class="mw-page-title-main">Animal</span> Kingdom of living things

Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, have myocytes and are able to move, can reproduce sexually, and grow from a hollow sphere of cells, the blastula, during embryonic development. Animals form a single clade.

<span class="mw-page-title-main">Planulozoa</span> Planulozoa

Planulozoa is a clade which includes the Placozoa, Cnidaria and the Bilateria. The designation Planulozoa may be considered a synonym to Parahoxozoa. Within Planulozoa, the Placozoa may be a sister of Cnidaria to the exclusion of Bilateria. The clade excludes basal animals such as the Ctenophora, and Porifera (sponges). Although this clade was sometimes used to specify a clade of Cnidaria and Bilateria to the exclusion of Placozoa, this is no longer favoured due to recent data indicating a sister group relationship between Cnidaria and Placozoa.

<span class="mw-page-title-main">Xenacoelomorpha</span> A deep-branching bilaterian clade of animals with a simple body plan

Xenacoelomorpha is a small phylum of bilaterian invertebrate animals, consisting of two sister groups: xenoturbellids and acoelomorphs. This new phylum was named in February 2011 and suggested based on morphological synapomorphies, which was then confirmed by phylogenomic analyses of molecular data.

<i>Hoilungia</i> Species of placozoa

Hoilungia is a genus that contains one of the simplest animals and belongs to the phylum Placozoa. Described in 2018, it has only one named species, H. hongkongensis, although there are possible other species. The animal superficially resembles another placozoan, Trichoplax adhaerens, but genetically distinct from it as mitochondrial DNA analysis revealed.

References

  1. Ryan, Joseph F.; Pang, Kevin; Mullikin, James C.; Martindale, Mark Q.; Baxevanis, Andreas D. (2010-10-04). "The homeodomain complement of the ctenophore Mnemiopsis leidyi suggests that Ctenophora and Porifera diverged prior to the Parahoxozoa". EvoDevo. 1 (1): 9. doi: 10.1186/2041-9139-1-9 . ISSN   2041-9139. PMC   2959044 . PMID   20920347.
  2. Pick, K. S.; Philippe, H.; Schreiber, F.; Erpenbeck, D.; Jackson, D. J.; Wrede, P.; Wiens, M.; Alie, A.; Morgenstern, B.; Manuel, M.; Worheide, G. (2010). "Improved Phylogenomic Taxon Sampling Noticeably Affects Nonbilaterian Relationships". Molecular Biology and Evolution. 27 (9): 1983–1987. doi: 10.1093/molbev/msq089 . ISSN   0737-4038. PMC   2922619 . PMID   20378579.
  3. Feuda, Roberto; Dohrmann, Martin; Pett, Walker; Philippe, Hervé; Rota-Stabelli, Omar; Lartillot, Nicolas; Wörheide, Gert; Pisani, Davide (2017). "Improved Modeling of Compositional Heterogeneity Supports Sponges as Sister to All Other Animals". Current Biology. 6 (24): 3864–3870.e4. Bibcode:2017CBio...27E3864F. doi: 10.1016/j.cub.2017.11.008 . hdl: 11572/302898 . PMID   29199080.
  4. Simion, Paul; Philippe, Hervé; Baurain, Denis; Jager, Muriel; Richter, Daniel J.; Franco, Arnaud Di; Roure, Béatrice; Satoh, Nori; Quéinnec, Éric (3 April 2017). "A Large and Consistent Phylogenomic Dataset Supports Sponges as the Sister Group to All Other Animals" (PDF). Current Biology (Submitted manuscript). 27 (7): 958–967. Bibcode:2017CBio...27..958S. doi: 10.1016/j.cub.2017.02.031 . ISSN   0960-9822. PMID   28318975. S2CID   4560353.
  5. Whelan, Nathan V.; Kocot, Kevin M.; Moroz, Leonid L.; Halanych, Kenneth M. (2015). "Error, signal, and the placement of Ctenophora sister to all other animals". Proceedings of the National Academy of Sciences. 112 (18): 5773–5778. Bibcode:2015PNAS..112.5773W. doi: 10.1073/pnas.1503453112 . ISSN   0027-8424. PMC   4426464 . PMID   25902535.
  6. Borowiec, Marek L.; Lee, Ernest K.; Chiu, Joanna C.; Plachetzki, David C. (2015). "Extracting phylogenetic signal and accounting for bias in whole-genome data sets supports the Ctenophora as sister to remaining Metazoa". BMC Genomics. 16 (1): 987. doi: 10.1186/s12864-015-2146-4 . ISSN   1471-2164. PMC   4657218 . PMID   26596625.
  7. Whelan, Nathan V.; Kocot, Kevin M.; Moroz, Tatiana P.; Mukherjee, Krishanu; Williams, Peter; Paulay, Gustav; Moroz, Leonid L.; Halanych, Kenneth M. (2017). "Ctenophore relationships and their placement as the sister group to all other animals". Nature Ecology & Evolution. 1 (11): 1737–1746. Bibcode:2017NatEE...1.1737W. doi:10.1038/s41559-017-0331-3. ISSN   2397-334X. PMC   5664179 . PMID   28993654.
  8. Moroz, Leonid L.; Kocot, Kevin M.; Citarella, Mathew R.; Dosung, Sohn; Norekian, Tigran P.; Povolotskaya, Inna S.; Grigorenko, Anastasia P.; Dailey, Christopher; Berezikov, Eugene; Buckley, Katherine M.; Ptitsyn, Andrey; Reshetov, Denis; Mukherjee, Krishanu; Moroz, Tatiana P.; Bobkova, Yelena; Yu, Fahong; Kapitonov, Vladimir V.; Jurka, Jerzy; Bobkov, Yuri V.; Swore, Joshua J.; Girardo, David O.; Fodor, Alexander; Gusev, Fedor; Sanford, Rachel; Bruders, Rebecca; Kittler, Ellen; Mills, Claudia E.; Rast, Jonathan P.; Derelle, Romain; Solovyev, Victor V.; Kondrashov, Fyodor A.; Swalla, Billie J.; Sweedler, Jonathan V.; Rogaev, Evgeny I.; Halanych, Kenneth M.; Kohn, Andrea B. (2014). "The ctenophore genome and the evolutionary origins of neural systems". Nature. 510 (7503): 109–114. Bibcode:2014Natur.510..109M. doi: 10.1038/nature13400 . ISSN   0028-0836. PMC   4337882 . PMID   24847885.
  9. Ryan, J. F.; Pang, K.; Schnitzler, C. E.; Nguyen, A.-D.; Moreland, R. T.; Simmons, D. K.; Koch, B. J.; Francis, W. R.; Havlak, P.; Smith, S. A.; Putnam, N. H.; Haddock, S. H. D.; Dunn, C. W.; Wolfsberg, T. G.; Mullikin, J. C.; Martindale, M. Q.; Baxevanis, A. D. (2013). "The Genome of the Ctenophore Mnemiopsis leidyi and Its Implications for Cell Type Evolution". Science. 342 (6164): 1242592. doi:10.1126/science.1242592. ISSN   0036-8075. PMC   3920664 . PMID   24337300.
  10. Erives, Albert; Fritzsch, Bernd (2020). "A Screen for Gene Paralogies Delineating Evolutionary Branching Order of Early Metazoa". G3: Genes, Genomes, Genetics. 10 (2): 811–826. doi:10.1534/g3.119.400951. ISSN   2160-1836. PMC   7003098 . PMID   31879283.
  11. 1 2 Giribet, Gonzalo; Edgecombe, Gregory D. (2020-03-03). The Invertebrate Tree of Life. Princeton University Press. p. 46. doi:10.1515/9780691197067-007. ISBN   978-0-691-19706-7.
  12. Fortunato, Sofia A. V.; Adamski, Marcin; Ramos, Olivia Mendivil; Leininger, Sven; Liu, Jing; Ferrier, David E. K.; Adamska, Maja (2014-10-30). "Calcisponges have a ParaHox gene and dynamic expression of dispersed NK homeobox genes". Nature. 514 (7524): 620–623. Bibcode:2014Natur.514..620F. doi:10.1038/nature13881. hdl: 10023/6597 . ISSN   0028-0836. PMID   25355364. S2CID   205241127.
  13. Larroux, Claire; Fahey, Bryony; Degnan, Sandie M.; Adamski, Marcin; Rokhsar, Daniel S.; Degnan, Bernard M. (1996). "The NK Homeobox Gene Cluster Predates the Origin of Hox Genes". Current Biology. 17 (8): 706–710. doi: 10.1016/j.cub.2007.03.008 . PMID   17379523. S2CID   18994676.
  14. Pastrana, Claudia C.; DeBiasse, Melissa B.; Ryan, Joseph F. (2019). "Sponges lack ParaHox genes". Genome Biology and Evolution. 11 (4): 1250–1257. doi:10.1093/gbe/evz052. PMC   6486804 . PMID   30859199.
  15. Giribet, Gonzalo (1 October 2016). "Genomics and the animal tree of life: conflicts and future prospects". Zoologica Scripta. 45: 14–21. doi: 10.1111/zsc.12215 . ISSN   1463-6409.
  16. Cannon, Johanna Taylor; Vellutini, Bruno Cossermelli; Smith, Julian; Ronquist, Fredrik; Jondelius, Ulf; Hejnol, Andreas (2016). "Xenacoelomorpha is the sister group to Nephrozoa". Nature (Submitted manuscript). 530 (7588): 89–93. Bibcode:2016Natur.530...89C. doi:10.1038/nature16520. PMID   26842059. S2CID   205247296.
  17. Browne, William E.; Amemiya, Chris T.; Swalla, Billie J.; Warren, Kaitlyn J.; Vandepas, Lauren E.; Presnell, Jason S. (2016-10-24). "The Presence of a Functionally Tripartite Through-Gut in Ctenophora Has Implications for Metazoan Character Trait Evolution". Current Biology. 26 (20): 2814–2820. Bibcode:2016CBio...26.2814P. doi: 10.1016/j.cub.2016.08.019 . ISSN   0960-9822. PMID   27568594.
  18. Jiménez-Guri, Eva; Philippe, Hervé; Okamura, Beth; Holland, Peter W. H. (2007-07-06). "Buddenbrockia Is a Cnidarian Worm". Science. 317 (5834): 116–118. Bibcode:2007Sci...317..116J. doi:10.1126/science.1142024. ISSN   0036-8075. PMID   17615357. S2CID   5170702.
  19. Baguñà, Jaume; Martinez, Pere; Paps, Jordi; Riutort, Marta (2008-04-27). "Back in time: a new systematic proposal for the Bilateria". Philosophical Transactions of the Royal Society B: Biological Sciences. 363 (1496): 1481–1491. doi:10.1098/rstb.2007.2238. ISSN   0962-8436. PMC   2615819 . PMID   18192186.
  20. Genikhovich, Grigory; Technau, Ulrich (2017-10-01). "On the evolution of bilaterality". Development. 144 (19): 3392–3404. doi: 10.1242/dev.141507 . ISSN   0950-1991. PMID   28974637.
  21. Syed, Tareq; Schierwater, Bernd (2002). "The evolution of the placozoa: A new morphological model". Senckenbergiana Lethaea. 82 (1): 315–324. doi:10.1007/bf03043791. ISSN   0037-2110. S2CID   16870420.
  22. Wallberg, Andreas; Thollesson, Mikael; Farris, James S.; Jondelius, Ulf (2004). "The phylogenetic position of the comb jellies (Ctenophora) and the importance of taxonomic sampling". Cladistics. 20 (6): 558–578. doi: 10.1111/j.1096-0031.2004.00041.x . ISSN   0748-3007. PMID   34892961. S2CID   86185156.
  23. Boero, F.; Schierwater, B.; Piraino, S. (2007-06-01). "Cnidarian milestones in metazoan evolution". Integrative and Comparative Biology. 47 (5): 693–700. doi:10.1093/icb/icm041. ISSN   1540-7063. PMID   21669750.