The Asterozoa are a subphylum in the phylum Echinodermata, within the Eleutherozoa. Characteristics include a star-shaped body and radially divergent axes of symmetry. The subphylum includes the classes Asteroidea (the starfish or sea stars), Ophiuroidea (the brittle stars and basket stars), Somasteroidea (early asterozoans from which the other classes most likely evolved), and Stenuroidea (early asterozoans with unclear relationships to extant classes).[2] A fifth class, Concentricycloidea, was proposed for the unusual genus Xyloplax (sea daisies),[4] but was later demoted to the status of infraclass as the sister of Neoasteroidea within the asteroidean sublcass Ambuloasteroidea.[5]
Asterozoa | |
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A brittle star on a starfish | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Echinodermata |
Subphylum: | Asterozoa von Zittel, 1895[1] |
Classes[2] | |
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Synonyms | |
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Taxonomy
editAsterozoa was originally proposed in the late 1800s, but was not used in F.A. Bather's two-subphylum echinoderm taxonomy in 1900. However, it was adopted as part of a four-subphylum taxonomy by the Treatise on Invertebrate Paleontology in 1966.[6]
Asterozoa is generally thought to be a monophyletic clade; its sister group within Eleutherozoa is Echinozoa.[7] The external affinities of Asterozoa are unclear. It has been proposed to derive from either the Edrioasteroidea or the Crinoidea, but no form of either proposal has gained wide acceptance due to the lack of any transitional fossils.[8]
Distinguishing the classes
editSomasteroids and stenuroids are distinguished from each other and from the extant asteroids and ophiuroids by the arrangements of a specific type of ossicle known as virgals: Somasteroids possess a varying number of virgals per series extending laterally from the ambulacral ossicles, with at least some series exceeding three virgals. Stenuroids possess series of exactly two (or rarely three or four) virgals for each ambulacral ossicle, with other ossicular specializations. Asteroid and ophiuroid virgal series are each reduced to a single ossicle in addition to their more obvious morphological differences.[9]
Somasteroidea
editMost authors consider Somasteroidea to be the basal stock from which the other three classes evolved.,[10][11] but an argument in favor of monophyly and a position closer to stenuroids and ophiuroids than to asteroids has also been made.[12]
Somasteroids are "more or less petaloid," with arm shape reflecting virgal series lengths.[13] Somasteroids have been described as being more rigid in shape than derived asterozoans, although this apparent structure could be exaggerated by tissue changes at the time of death.[14] S
Stenuroidea
editStenuroids were initially seen as early ophiuroids[15] before being promoted to class level, but their evaluation is challenging due to wide variations in morphologies. Some stenuroids appear closer to asteroids, others to ophiuroids, and others do not closely resemble either extant class.[16] A recent examination of Stenuroidea found it to be monophyletic, but allowed that (as with other asterozoan classes), paraphyly or polyphyly could not be entirely ruled out.[17] Other authors have considered that Stenuroidea is likely paraphyletic.[2]
Asteroidea
editAsteroids have a permanently vaulted ambulacral furrow down the center of the underside of each arm. They use their tube feet for locomotion and (in many species) to pry open shells and access food.[18] Their arms touch at the base, and there is no clear border between the arms and the central disc.[19]
Ophiuroidea
editOphiuroid arms have evolved joints within their arms allowing lateral, snake-like movements for locomotion, while the tube feet are significantly reduced.[18] The central disc is clearly marked off from the arms.[19]
References
edit- ^ K. A. von Zittel. 1895. Grundzuge der Palaeontologie 1-971
- ^ a b c Nanglu et al. 2023, p. 331
- ^ Spencer & Wright 1966, p. U39
- ^ Rowe, Baker & Clark 1988
- ^ Mah 2006
- ^ Spencer & Wright 1966
- ^ Escriva et al. 2015
- ^ Blake 2013, p. 357
- ^ Blake 2024, p. 13
- ^ Mooi & David 2000
- ^ Blake & Guensburg 2015
- ^ Dean Shackleton 2005
- ^ Blake & Hotchkiss 2022, p. 29
- ^ Blake 2024, p. 2
- ^ Blake 2024, p. 18
- ^ Blake 2013, p. 361
- ^ Blake 2024, p. 1
- ^ a b Blake 2013, p. 359
- ^ a b Giribet & Edgecombe 2020, p. 120
Works cited
edit- Blake, Daniel B. (2013). "Early asterozoan (Echinodermata) diversification: a paleontologic quandry". Journal of Paleontology. 87 (3): 353–372. doi:10.1666/12-042.1.
- Blake, Daniel B. (2024). "A review of the class Stenuroidea (Echinodermata: Asterozoa)". Bulletins of American Paleontology. 409. doi:10.32857/bap.2024.409.01.
- Blake, Daniel B.; Guensburg, Thomas E. (2015). "The class Somasteroidea (Echinodermata, Asterozoa): morphology and occurrence". Journal of Paleontology. 89 (3): 465–486. doi:10.1017/jpa.2015.22.
- Blake, Daniel B.; Hotchkiss, Frederick H.C. (2022). "Origin of the subphylum Asterozoa and redescription of a Moroccan Ordovician somasteroid". Geobios. 72–73: 22–36. doi:10.1016/j.geobios.2022.07.002.
- Dean Shackleton, Juliette (2005). "Skeletal homologies, phylogeny and classification of the earliest asterozoan echinoderms". Journal of Systematic Palaeontology. 3 (1): 29–114. doi:10.1017/S1477201905001525.
- Escriva, Hector; Reich, Adrian; Dunn, Casey; Akasaka, Koji; Wessel, Gary (2015). "Phylogenomic Analyses of Echinodermata Support the Sister Groups of Asterozoa and Echinozoa". PLOS ONE. 10 (3): e0119627. Bibcode:2015PLoSO..1019627R. doi:10.1371/journal.pone.0119627. PMC 4368666. PMID 25794146.
- Giribet, Gonzalo; Edgecombe, Gregory D. (2020). "Echinodermata". The Invertebrate Tree of Life. Princeton University Press. pp. 115–138.
- Mah, Christopher L. (June 2006). "A new species of Xyloplax (Echinodermata: Asteroidea: Concentricycloidea) from the northeast Pacific: comparative morphology and a reassessment of phylogeny". Invertebrate Biology. 125 (2): 83–176. doi:10.1111/j.1744-7410.2006.00048.x.
- Mooi, Rich; David, Bruno (June 2000). "What a new model of skeletal homologies tells us about asteroid evolution". American Zoologist. 40 (3): 326–339. doi:10.1668/0003-1569(2000)040[0326:WANMOS]2.0.CO;2.
- Nanglu, Karma; Cole, Selina R.; Wright, David F.; Souto, Camilla (2023). "Worms and gills, plates and spines: the evolutionary origins and incredible disparity of deuterostomes revealed by fossils, genes, and development". Biological Reviews. 98: 316–351. doi:10.1111/brv.12908.
- Rowe, F. W. E.; Baker, A. N.; Clark, H. E. S. (23 May 1988). "The Morphology, Development and Taxonomic Status of Xyloplax Baker, Rowe and Clark (1986) (Echinodermata: Concentricycloidea), with the Description of a New Species". Proceedings of the Royal Society of London. Series B, Biological Sciences. 233 (1237): 431–459. doi:10.1098/rspb.1988.0032.
- Shackleton, Juliette Dean (2005). "Skeletal homologies, phylogeny and classification of the earliest asterozoan echinoderms". Journal of Systematic Palaeontology. 3 (1): 29–114. doi:10.1017/S1477201905001525.
- Spencer, W. K.; Wright, C. W. (1966). "Asterozoans". In Moore; Raymond C. (eds.). Treatise on Invertebrate Paleontology, Part U: Echinodermata 3. Vol. 1. University of Kansas Press. pp. U5 – U107. Retrieved 28 October 2024.