Sample Page

Ecdysozoa (/ˌɛkdɪsˈzə/) is a group of protostome animals,[11] including Arthropoda (insects, chelicerates (including arachnids), crustaceans, and myriapods), Nematoda, and several smaller phyla. The grouping of these animal phyla into a single clade was first proposed by Eernisse et al. (1992) based on a phylogenetic analysis of 141 morphological characters of ultrastructural and embryological phenotypes.[12] This clade, that is, a group consisting of a common ancestor and all its descendants, was formally named by Aguinaldo et al. in 1997, based mainly on phylogenetic trees constructed using 18S ribosomal RNA genes.[13]

A large study in 2008 by Dunn et al. strongly supported the monophyly of Ecdysozoa.[14]

The group Ecdysozoa is supported by many morphological characters, including growth by ecdysis, with moulting of the cuticle – without mitosis in the epidermis – under control of the prohormone ecdysone, and internal fertilization.[15]

The group was initially contested by a significant minority of biologists. Some argued for groupings based on more traditional taxonomic techniques,[16] while others contested the interpretation of the molecular data.[17][18]

Etymology

The name Ecdysozoa is scientific Greek, derived from ἔκδυσις (ékdusis) “shedding” + ζῷον (zôion) “animal”.

Characteristics

The most notable characteristic shared by ecdysozoans is a three-layered cuticle (four in Tardigrada[19]) composed of organic material, which is periodically molted as the animal grows. This process of molting is called ecdysis, and gives the group its name. The ecdysozoans lack locomotory cilia and produce mostly amoeboid sperm, and their embryos do not undergo spiral cleavage as in most other protostomes. Ancestrally, the group exhibited sclerotized teeth within the foregut, and a ring of spines around the mouth opening, though these features have been secondarily lost in certain groups.[20][21] An unpaired ventral nerve cord, present in Priapulida and Nematoida, appear to be the ancestral condition, making the paired ventral nerve cord found in Panarthropoda, Kinorhyncha and Loricifera a derived trait.[22] A respiratory and circulatory system is only present in onychophorans and arthropods (often absent in smaller arthropods like mites); in the rest of the groups, both systems are missing. Ecdysozoans rely exclusively on the arginine phosphate/arginine kinase (AP/AK) system, a high-energy phosphate system used to regenerate ATP from ADP. In contrast, vertebrates use only the creatine phosphate/creatine kinase (CP/CK) system, while some other invertebrates may employ both systems.[23][24]

Phylogeny

The Ecdysozoa include the following phyla: Arthropoda, Onychophora, Tardigrada, Kinorhyncha, Priapulida, Loricifera, Nematoda, and Nematomorpha. A few extinct taxa have been classified as stem group ecdysozoans, such as Uncus dzaugisi and Acosmia. Other groups such as the gastrotrichs, have been considered possible members but lack the main characters of the group, and are now placed elsewhere. The Arthropoda, Onychophora, and Tardigrada have been grouped together as the Panarthropoda because they are distinguished by segmented body plans.[25] Dunn et al. in 2008 suggested that the tardigrada could be grouped along with the nematodes, leaving Onychophora as the sister group to the arthropods.[14] The non-panarthropod members of Ecdysozoa have been grouped as Cycloneuralia but they are more usually considered paraphyletic in representing the primitive condition from which the Panarthropoda evolved.[26]

A modern consensus phylogenetic tree for Ecdysozoa is shown below, mainly based on the one supported by Giribet & Edgecombe (2017), which summarizes relationships found in recent ecdysozoan phylogenies:[27][28][29][30]: 21 [2]

Older alternative groupings

Articulata hypothesis

The grouping proposed by Aguinaldo et al. is almost universally accepted, replacing an older hypothesis that Panarthropoda should be classified with Annelida in a group called the Articulata, and that Ecdysozoa are polyphyletic. Nielsen has suggested that a possible solution is to regard Ecdysozoa as a sister-group of Annelida,[31] though later considered them unrelated.[32] Inclusion of the roundworms within the Ecdysozoa was initially contested[17][33][34] but since 2003, a broad consensus has formed supporting the Ecdysozoa[35] and in 2011 the Darwin–Wallace Medal was awarded to James Lake for the discovery of the New Animal Phylogeny consisting of the Ecdysozoa, the Lophotrochozoa, and the Deuterostomia.[36]

Coelomata hypothesis

Before Aguinaldo’s Ecdysozoa proposal, one of the prevailing theories for the evolution of the bilateral animals was based on the morphology of their body cavities. There were three types, or grades of organization: the Acoelomata (no coelom), the Pseudocoelomata (partial coelom), and the Eucoelomata (true coelom). Adoutte and coworkers were among the first to strongly support the Ecdysozoa.[37] With the introduction of molecular phylogenetics, the coelomate hypothesis was abandoned, although some molecular, phylogenetic support for the Coelomata continued until as late as 2005.[38]

References

  1. ^ Hughes, Ian V.; Evans, Scott D.; Droser, Mary L. (November 2024). “An Ediacaran bilaterian with an ecdysozoan affinity from South Australia”. Current Biology. 34 (24): 5782–5788.e1. Bibcode:2024CBio…34.5782H. doi:10.1016/j.cub.2024.10.030. PMID 39561775.
  2. ^ a b c Howard, Richard J.; Giacomelli, Mattia; Lozano-Fernandez, Jesus; Edgecombe, Gregory D.; Fleming, James F.; Kristensen, Reinhardt M.; et al. (10 March 2022). “The Ediacaran origin of Ecdysozoa: Integrating fossil and phylogenomic data”. Journal of the Geological Society. 179 (4). Bibcode:2022JGSoc.179..107H. doi:10.1144/jgs2021-107. hdl:2445/186596. ISSN 0016-7649. S2CID 246494357.
  3. ^ Shao, T.Q.; Qin, J.C.; Shao, Y.; Liu, Y.H.; Waloszek, D.; Maas, A.; Duan, B.C.; Wang, Q.; Xu, Y.; Zhang, H.Q. (October 2020). “New macrobenthic cycloneuralians from the Fortunian (lowermost Cambrian) of South China”. Precambrian Research. 349 105413. Bibcode:2020PreR..34905413S. doi:10.1016/j.precamres.2019.105413.
  4. ^ Bekkouche, Nicolas; Gąsiorowski, Ludwik (31 December 2022). “Careful amendment of morphological data sets improves phylogenetic frameworks: re-evaluating placement of the fossil Amiskwia sagittiformis” (PDF). Journal of Systematic Palaeontology. 20 (1): 1–14. doi:10.1080/14772019.2022.2109217.
  5. ^ a b Liu, Yunhuan; Qin, Jiachen; Wang, Qi; Maas, Andreas; Duan, Baichuan; Zhang, Yanan; Zhang, Hu; Shao, Tiequan; Zhang, Huaqiao (May 2019). “New armoured scalidophorans (Ecdysozoa, Cycloneuralia) from the Cambrian Fortunian Zhangjiagou Lagerstätte, South China”. Papers in Palaeontology. 5 (2): 241–260. Bibcode:2019PPal….5..241L. doi:10.1002/spp2.1239.
  6. ^ Zhang, Hua-Qiao (September 2022). “The evolutionary relationships of the earliest known cycloneuralians and a new record from the Cambrian Fortunian of South China”. Palaeoworld. 31 (3): 389–401. doi:10.1016/j.palwor.2021.09.003.
  7. ^ Shao, T.Q.; Wang, Q.; Liu, Y.H.; Qin, J.C.; Zhang, Y.N.; Liu, M.J.; Shao, Y.; Zhao, J.Y.; Zhang, H.Q. (October 2020). “A new scalidophoran animal from the Cambrian Fortunian Stage of South China and its implications for the origin and early evolution of Kinorhyncha”. Precambrian Research. 349 105616. Bibcode:2020PreR..34905616S. doi:10.1016/j.precamres.2020.105616.
  8. ^ Howard, Richard J.; Edgecombe, Gregory D.; Shi, Xiaomei; Hou, Xianguang; Ma, Xiaoya (23 November 2020). “Ancestral morphology of Ecdysozoa constrained by an early Cambrian stem group ecdysozoan”. BMC Evolutionary Biology. 20 (1): 156. Bibcode:2020BMCEE..20..156H. doi:10.1186/s12862-020-01720-6. ISSN 1471-2148. PMC 7684930. PMID 33228518.
  9. ^ Wang, Deng; Han, Jian; Guo, Junfeng; Qiang, Yaqin (March 2025). “Origin and evolution of bodyplans of ecdysozoans during the Cambrian explosion”. Chinese Journal of Nature. 47 (2): 125–133. doi:10.3969/j.issn.0253-9608.2025.02.004.
  10. ^ Liu, Yunhuan; Carlisle, Emily; Zhang, Huaqiao; Yang, Ben; Steiner, Michael; Shao, Tiequan; et al. (17 August 2022). “Saccorhytus is an early ecdysozoan, and not the earliest deuterostome”. Nature. 609 (7927): 541–546. Bibcode:2022Natur.609..541L. doi:10.1038/s41586-022-05107-z. hdl:1983/454e7bec-4cd4-4121-933e-abeab69e96c1. ISSN 1476-4687. PMID 35978194. S2CID 251646316.
  11. ^ Telford, Maximilian J.; Bourlat, Sarah J.; Economou, Andrew; Papillon, Daniel; Rota-Stabelli, Omar (2008). “The evolution of the Ecdysozoa”. Philosophical Transactions of the Royal Society B: Biological Sciences. 363 (1496): 1529–1537. doi:10.1098/rstb.2007.2243. ISSN 0962-8436. PMC 2614232. PMID 18192181.
  12. ^ Eernisse, D.J.; Albert, J.S.; Anderson, F.E. (1992). “Annelida and Arthropoda are not sister taxa: A phylogenetic analysis of spiralian metazoan morphology”. Systematic Biology. 41 (3): 305–330. doi:10.1093/sysbio/41.3.305.
  13. ^ Aguinaldo, A.M.A.; Turbeville, J.M.; Linford, L.S.; Rivera, M.C.; Garey, J.R.; Raff, R.A.; Lake, J.A. (29 May 1997). “Evidence for a clade of nematodes, arthropods, and other moulting animals”. Nature. 387 (6632): 489–493. Bibcode:1997Natur.387R.489A. doi:10.1038/387489a0. PMID 9168109. S2CID 4334033.
  14. ^ a b Dunn, C.W.; Hejnol, A.; Matus, D.Q.; Pang, K.; Browne, W.E.; Smith, S.A.; et al. (10 April 2008). “Broad phylogenomic sampling improves resolution of the animal tree of life”. Nature. 452 (7188): 745–749. Bibcode:2008Natur.452..745D. doi:10.1038/nature06614. PMID 18322464. S2CID 4397099.
  15. ^ Ax, P. (1985). “The position of the Gnathostomulida and Platyhelminthes in the phylogenetic system of the Bilateria”. In Conway Morris, S.; George, J.D.; Gibson, R.; Platt, H.M. (eds.). The Origins and Relationships of Lower Invertebrates. Systematics Association Special Volume. Vol. 28. New York, NY: Clarendon / Oxford University Press. pp. 168–180. ISBN 0-19-857181-X. OCLC 59186778 – via Internet Archive (archive.org). proceedings of an international symposium held in London, September 1983 ISBN 9780198571810
  16. ^ Nielsen, Claus (1995). Animal Evolution: Interrelationships of the living phyla. Oxford University Press. ISBN 978-0-19-850682-9.
  17. ^ a b Blair, J.E.; Ikeo, Kazuho; Gojobori, Takashi; Hedges, S. Blair (8 April 2002). “The evolutionary position of nematodes”. BMC Evolutionary Biology. 2: 7. doi:10.1186/1471-2148-2-7. PMC 102755. PMID 11985779.
  18. ^ Wägele, J.W.; Erikson, T.; Lockhart, P.; Misof, B. (December 1999). “The Ecdysozoa: Artifact or monophylum?”. Journal of Zoological Systematics and Evolutionary Research. 37 (4): 211–223. doi:10.1111/j.1439-0469.1999.tb00985.x.
  19. ^ Barnes, Robert D. (1982). Invertebrate Zoology. Philadelphia, PA: Holt-Saunders International. pp. 877–880. ISBN 0-03-056747-5.
  20. ^ Smith, Martin R.; Caron, Jean-Bernard (2 July 2015). “Hallucigenia’s head and the pharyngeal armature of early ecdysozoans” (PDF). Nature. 523 (7558): 75–78. Bibcode:2015Natur.523…75S. doi:10.1038/nature14573. PMID 26106857. S2CID 205244325.
  21. ^ Vinther, Jakob; Porras, Luis; Young, Fletcher J.; Budd, Graham E.; Edgecombe, Gregory D. (2016). “The mouth apparatus of the Cambrian gilled lobopodian Pambdelurion whittingtoni. Palaeontology. 59 (6): 841–849. Bibcode:2016Palgy..59..841V. doi:10.1111/pala.12256. hdl:1983/16da11f1-5231-4d6c-9968-69ddc5633a8a.
  22. ^ Wang, Deng; Vannier, Jean; Martín-Durán, José M.; Herranz, María; Yu, Chiyang (2025). “Preservation and early evolution of scalidophoran ventral nerve cord”. Science Advances. 11 (2) eadr0896. Bibcode:2025SciA…11R.896W. doi:10.1126/sciadv.adr0896. PMC 11721716. PMID 39792685.
  23. ^ Lukić-Bilela, Lada; Perović-Ottstadt, Sanja; Walenta, Stefan; Natalio, Filipe; Pleše, Bruna; Link, Thorben; Müller, Werner E. G. (2011). “ATP distribution and localization of mitochondria in Suberites domuncula (Olivi 1792) tissue”. Journal of Experimental Biology. 214 (10): 1748–1753. Bibcode:2011JExpB.214.1748L. doi:10.1242/jeb.053439. PMID 21525322.
  24. ^ Deligio, J.T.; Ellington, W.R. (2006). “The capacity for the de novo biosynthesis of creatine is present in the tunicate Ciona intestinalis and is likely widespread in other protochordate and invertebrate groups”. Comparative Biochemistry and Physiology Part D: Genomics and Proteomics. 1 (2): 167–178. doi:10.1016/j.cbd.2005.08.006. PMID 20483248.
  25. ^ “Panarthropoda”. Paleos (palaeos.com). Invertebrates. Archived from the original on 2007-02-07. Retrieved 17 February 2007.
  26. ^ Webster, Bonnie L.; Copley, Richard R.; Jenner, Ronald A.; Mackenzie-Dodds, Jacqueline A.; Bourlat, Sarah J.; Rota-Stabelli, Omar; et al. (November 2006). “Mitogenomics and phylogenomics reveal priapulid worms as extant models of the ancestral Ecdysozoan”. Evolution & Development. 8 (6): 502–510. doi:10.1111/j.1525-142X.2006.00123.x. PMID 17073934. S2CID 22823313.
  27. ^ Giribet, Gonzalo; Edgecombe, Gregory D. (September 2017). “Current Understanding of Ecdysozoa and its Internal Phylogenetic Relationships”. Integrative and Comparative Biology. 57 (3): 455–466. doi:10.1093/icb/icx072. ISSN 1540-7063.
  28. ^ Giribet, Gonzalo (2025-11-18). “Evolución de ecdisozoos y una hoja de ruta para resolver su filogenia”. eVOLUCIÓN: Boletín Electrónico de la SESBE (in Spanish). 20 (II). doi:10.65120/evo.25. ISSN 1989-046X.
  29. ^ Giribet, Gonzalo (October 2016). “Genomics and the animal tree of life: conflicts and future prospects”. Zoologica Scripta. 45 (S1): 14–21. doi:10.1111/zsc.12215. ISSN 0300-3256.
  30. ^ Giribet, Gonzalo; Edgecombe, Gregory D. (2020-03-03). The Invertebrate Tree of Life. Princeton University Press. doi:10.2307/j.ctvscxrhm. ISBN 978-0-691-19706-7.
  31. ^ Nielsen, C. (September 2003). “Proposing a solution to the Articulata–Ecdysozoa controversy”. Zoologica Scripta. 32 (5): 475–482. doi:10.1046/j.1463-6409.2003.00122.x. S2CID 1416582.
  32. ^ Nielsen, Claus (2012). Animal Evolution: Interrelationships of the living phyla (3rd ed.). Oxford University Press. ISBN 978-0-19-960603-0.
  33. ^ Zrzavý, J. (12 January 2002). “Ecdysozoa versus Articulata: Clades, artifacts, prejudices”. Journal of Zoological Systematics and Evolutionary Research. 39 (3): 159–163. doi:10.1046/j.1439-0469.2001.00168.x. — in support of clade Ecdysozoa
  34. ^ Wägele, J.W.; Misof, B. (September 2001). “On quality of evidence in phylogeny reconstruction: A reply to Zrzavý’s defence of the ‘Ecdysozoa’ hypothesis”. Journal of Zoological Systematics and Evolutionary Research. 39 (3): 165–176. doi:10.1046/j.1439-0469.2001.00177.x.
  35. ^ Telford, Maximilian J.; Littlewood, D. Timothy J. (27 April 2008). “The evolution of the animals: Introduction to a Linnean tercentenary celebration”. Philosophical Transactions of the Royal Society B: Biological Sciences. 363 (1496): 1421–1424. doi:10.1098/rstb.2007.2231. PMC 2394567. PMID 18192193.
  36. ^ “The Darwin-Wallace Medal” (Press release). The Linnean Society of London. Retrieved 28 February 2018.
  37. ^ Adoutte, A.; Balavoine, G.; Lartillot, N.; Lespinet, O.; Prud’homme, B.; de Rosa, R. (25 April 2000). “The new animal phylogeny: Reliability and implications”. Special feature. Proceedings of the National Academy of Sciences of the USA. 97 (9): 4453–4456. Bibcode:2000PNAS…97.4453A. doi:10.1073/pnas.97.9.4453. PMC 34321. PMID 10781043.
  38. ^ Philip, G.K.; Creevey, C.J.; McInerney, J.O. (9 February 2005). “The Opisthokonta and the Ecdysozoa may not be clades: Stronger support for the grouping of plant and animal than for animal and fungi, and stronger support for the Coelomata than Ecdysozoa”. Molecular Biology and Evolution. 22 (5): 1175–1184. doi:10.1093/molbev/msi102. PMID 15703245.

Further reading

  • Nielsen, C. (2002). Animal Evolution: Interrelationships of the Living Phyla (2nd ed.). Oxford University Press. ISBN 0-19-850682-1.