Echinoderm
Echinoderms are among the most architecturally strange animals on the planet. A starfish has no brain, no left or right side, and can digest its prey outside its own body. Sea cucumbers eject their internal organs at predators, then grow them back. Some species can regenerate an entire animal from a single severed limb. The phylum Echinodermata contains about 7,600 living species, and another roughly 13,000 that exist only in the fossil record. They are the largest marine-only phylum on Earth. Yet for all their diversity, every adult echinoderm is built around the same strange architectural principle: a body arranged in five parts around a central axis. What is this body plan, where did it come from, and why has it persisted from the Cambrian to the present day?
Every echinoderm larva is bilaterally symmetrical, possessing a left side and a right side just like a fish or a human. Then metamorphosis arrives. During that transformation, the left side of the body begins to grow at the expense of the right side, which is eventually absorbed entirely. What remains is a creature arranged in five parts around a central axis, a form called pentaradial symmetry. This shift is among the more dramatic developmental events in the animal kingdom. After metamorphosis, genetic studies of starfish show that the genes directing head development are expressed along the arms' central ambulacra, with trunk genes limited to interior tissue rather than the body surface. By this logic, a starfish body can be understood as consisting almost entirely of a head. The five-armed pattern has exceptions: most starfish in the genus Leptasterias have six arms, and Labidiaster annulatus can possess up to fifty arms. The sea lily Comaster schlegelii reaches two hundred. But five remains the foundation.
Echinoderm skeletons are embedded in the dermis rather than outside the body, composed of calcite-based plates called ossicles. These ossicles are not solid. They have a sponge-like porous structure called stereom, which keeps them light. Ossicles may fuse into the rigid test of a sea urchin, or they may articulate into the flexible joints of a starfish arm. In sea urchins, some ossicles are gathered into a specialized chewing structure called Aristotle's lantern. Sea cucumbers incorporate a structural ring of lime. The epidermis contains pigment cells that create the vivid colours common across the phylum, including deep red, stripes of black and white, and intense purple. Those cells can be light-sensitive: the sea urchin Centrostephanus longispinus changes colour in just fifty minutes when exposed to light. Perhaps the most unusual tissue in the phylum is what researchers call catch connective tissue, a collagen-based material that can shift between flaccid and rigid states under nervous control, without the involvement of muscles. This allows a starfish to move flexibly across the seabed, then lock itself rigid while prying open a bivalve, or resist being pulled from a crevice.
No other animal phylum uses a fluid-filled canal network as its primary system for movement, feeding, gas exchange, and sensory reception, yet that is precisely what echinoderms do. The water vascular system typically opens to the exterior through a sieve-like plate called a madreporite on the animal's upper surface. From there, a stone canal connects to a ring canal encircling the mouth, which branches into radial canals extending into each arm or around the body. Short lateral branches off those radial canals end in chambers called ampullae. Fluid can be pushed from the ampulla into the foot, extending it outward; withdrawing the fluid contracts the foot. Thousands of these tube feet, extended and contracted in waves, propel the animal across the seabed. The arrangement varies considerably between groups. In brittle stars, the madreporite sits on the oral surface and the tube feet lack suckers. In sea cucumbers, the whole system is reduced, with the madreporite opening into the body cavity rather than the exterior. Crinoids use their tube feet not for locomotion but in a back-and-forth wafting motion, sweeping food particles toward the central mouth.
Sea cucumbers can discharge parts of their internal organs when threatened, then regenerate them over a period of several months. During that interval, they live off stored nutrients and absorb dissolved organic matter directly from the surrounding water. Sea urchins constantly replace spines lost to damage. In most starfish species, regenerating a complete animal from a single severed arm requires at least a portion of the central disc, but a few species can produce a complete individual from an arm alone. When doing so asexually, some individuals produce a characteristic form: a large original arm with several tiny new ones growing from the stub, a shape that has earned them the common name "comets." Holothuria parvula reproduces asexually by transverse fission, splitting into two halves slightly ahead of the midpoint; each half then regenerates its missing organs. Even echinoderm larvae can clone themselves. Larvae of some sand dollars initiate cloning when they detect dissolved fish mucus in the water, a chemical signal indicating predators nearby. The cloned larvae are smaller and escape planktivorous fish more effectively, suggesting the behaviour functions as an anti-predator adaptation. The regeneration process itself involves two mechanisms: epimorphosis, in which stem cells form a blastema and produce new tissue, and morphallaxis, in which existing tissue is reorganised and remodelled to replace what was lost.
Echinoderms sequester about 0.1 gigatonnes of carbon dioxide per year as calcium carbonate, giving them a measurable role in the global carbon cycle. Their population swings can reorder entire ecosystems. In 1983, mass mortality of the tropical sea urchin Diadema antillarum in the Caribbean converted a coral-dominated reef into an alga-dominated one. On the Great Barrier Reef, unexplained increases in the crown-of-thorns starfish Acanthaster planci have driven significant coral mortality. In 2019, harvests of echinoderms reached 129,052 tonnes globally. Sea cucumbers accounted for 59,262 tonnes and sea urchins for 66,341 tonnes, consumed mainly as food and in traditional Chinese medicine. Popular species include the pineapple roller Thelenota ananas and the red sea cucumber Holothuria edulis, collectively known in parts of Asia as bêche de mer or trepang. Both male and female gonads of sea urchins are eaten in Japan and France, described as soft and melting, like a mixture of seafood and fruit. In research, sea urchins are widely used as model organisms in developmental biology and ecotoxicology; Strongylocentrotus purpuratus and Arbacia punctulata appear frequently in embryological studies. The arm regeneration capacity of brittle stars is currently being studied for what it might reveal about neurodegenerative diseases in humans.
The oldest potential echinoderm fossil is Arkarua, a disc-like organism from the late Ediacaran of Australia, dating to around 555 million years ago. Its fossils display radial ridges and a five-pointed central depression, but they lack the stereom structure that would confirm it as a true echinoderm. Whether Arkarua belongs in the phylum remains contested. The next candidates are the vetulocystids, from the early to mid Cambrian, roughly 541-501 million years ago. Yanjiahella, from approximately 539-529 million years ago, is notable for possessing a plated theca, though without confirmed stereom. The first universally accepted echinoderms appear in the Lower Cambrian. The phylum name Echinodermata was originated by Jacob Theodor Klein in 1734, initially applied only to echinoids. Jean Guillaume Bruguière expanded it to the full phylum level, first informally in 1789 and then in formal Latin in 1791. A 2024 review of echinoderm phylogeny identified two competing schools of thought: one that considers pentaradiality a primitive trait shared by all early members, and another that treats it as a derived feature that evolved later. Supporters of the second position point to recently discovered fossils including Ctenoimbricata and Helicocystis as evidence that non-radial forms predate the five-pointed body plan. The debate centres on whether the early dominance of pentaradial forms in the fossil record reflects true evolutionary history or simply the incompleteness of what has been preserved.
Common questions
What is an echinoderm and what animals belong to the phylum Echinodermata?
Echinoderms are marine animals of the phylum Echinodermata, which includes starfish, brittle stars, sea urchins, sand dollars, sea cucumbers, and sea lilies. The phylum contains about 7,600 living species and roughly 13,000 known extinct species, making it the largest marine-only phylum on Earth.
Why do echinoderms have five-pointed radial symmetry?
Adult echinoderms develop pentaradial symmetry during metamorphosis: the left side of the larval body grows at the expense of the right, which is absorbed, and the remaining tissue reorganises around a central axis in five parts. Larvae are bilaterally symmetrical before this process begins.
How do echinoderms move without muscles or a brain?
Echinoderms move using a water vascular system, a network of fluid-filled canals connected to external tube feet. Fluid pushed into a tube foot extends it; withdrawing the fluid contracts it. Thousands of tube feet operating in coordinated waves propel the animal across the seabed. There is no central brain; nerves radiate from rings around the mouth into each arm.
Can echinoderms really regenerate lost body parts?
Most echinoderms have strong regenerative abilities. Sea cucumbers can expel and then regrow internal organs over several months. Sea urchins continuously replace lost spines. A few starfish species can regenerate a complete individual from a single severed arm, though most require at least part of the central disc.
What is the ecological role of echinoderms in marine ecosystems?
Echinoderms sequester about 0.1 gigatonnes of carbon dioxide per year as calcium carbonate. Sea urchins are primary herbivores on coral reefs; in 1983 the mass mortality of Diadema antillarum in the Caribbean shifted reefs from coral-dominated to alga-dominated systems. Sea cucumbers process large volumes of seabed sediment as deposit feeders.
How are echinoderms used by humans as food and in research?
In 2019, global echinoderm harvests totalled 129,052 tonnes, primarily sea urchins (66,341 tonnes) and sea cucumbers (59,262 tonnes), consumed as food and in traditional Chinese medicine. In research, sea urchins such as Strongylocentrotus purpuratus and Arbacia punctulata serve as model organisms in developmental biology. Brittle star arm regeneration is studied for its potential relevance to human neurodegenerative diseases.
All sources
163 references cited across the entry
- 1harvnbSmith, Zamora, Álvaro (2013) p. 2Smith, Zamora, Álvaro — 2013
- 2harvnbHall (2022)Hall — 2022
- 3dictionaryechinoderm
- 4harvnbStokes (2021)Stokes — 2021
- 5harvnbITIS p. EchinodermataITIS
- 6harvnbUbaghs (1967) p. S5Ubaghs — 1967
- 7harvnbBather (1900)Bather — 1900
- 8eolEchinoderms
- 9harvnbU. of Michigan Museum of Zoology
- 10harvnbUthicke, Schaffelke, Byrne (2009)Uthicke, Schaffelke, Byrne — 2009
- 11harvnbArnone, Byrne, Martinez (2015)Arnone, Byrne, Martinez — 2015
- 12harvnbRuppert, Fox, Barnes (2004) p. 873Ruppert, Fox, Barnes — 2004
- 13harvnbByrne, O'Hara, CSIRO (2017) p. 322–324Byrne, O'Hara, CSIRO — 2017
- 14harvnbHolló, Novák (2012)Holló, Novák — 2012
- 15harvnbMessing (2004)Messing — 2004
- 16harvnbFormery, Peluso, Kohnle (2023)Formery, Peluso, Kohnle — 2023
- 17harvnbLacali (2023)Lacali — 2023
- 18harvnbBehrens, Bäuerlein (2007) p. 393Behrens, Bäuerlein — 2007
- 19harvnbBrusca, Moore, Shuster (2016) p. 979–980Brusca, Moore, Shuster — 2016
- 20harvnbDavies (1925) p. 240–241Davies — 1925
- 21harvnbPerillo, Oulhen, Foster (2020)Perillo, Oulhen, Foster — 2020
- 22harvnbWeber, Dambach (1974)Weber, Dambach — 1974
- 23harvnbMotokawa (1984)Motokawa — 1984
- 24harvnbBrusca, Moore, Shuster (2016) p. 980Brusca, Moore, Shuster — 2016
- 25harvnbDorit, Walker, Barnes (1991) p. 780–791Dorit, Walker, Barnes — 1991
- 26harvnbBrusca, Moore, Shuster (2016) p. 980–982Brusca, Moore, Shuster — 2016
- 27harvnbDorit, Walker, Barnes (1991) p. 784–785Dorit, Walker, Barnes — 1991
- 28harvnbBrusca, Moore, Shuster (2016) p. 982Brusca, Moore, Shuster — 2016
- 29harvnbDorit, Walker, Barnes (1991) p. 790–793Dorit, Walker, Barnes — 1991
- 30harvnbJangoux (1982) p. 244Jangoux — 1982
- 31harvnbRigby, Iken, Shirayama (2007) p. 44Rigby, Iken, Shirayama — 2007
- 32harvnbRuppert, Fox, Barnes (2004) p. 885Ruppert, Fox, Barnes — 2004
- 33harvnbRuppert, Fox, Barnes (2004) p. 891Ruppert, Fox, Barnes — 2004
- 34harvnbRuppert, Fox, Barnes (2004) p. 902–904Ruppert, Fox, Barnes — 2004
- 35harvnbRuppert, Fox, Barnes (2004) p. 912Ruppert, Fox, Barnes — 2004
- 36harvnbRuppert, Fox, Barnes (2004) p. 920Ruppert, Fox, Barnes — 2004
- 37harvnbMoore, Overhill (2006) p. 245Moore, Overhill — 2006
- 38harvnbHickman, Roberts, Larson (2003) p. 271Hickman, Roberts, Larson — 2003
- 39harvnbMacrobenthos of the North Sea p. EchinodermataMacrobenthos of the North Sea
- 40harvnbNielsen (2012) p. 78Nielsen — 2012
- 41harvnbRamirez-Gomez (2010)Ramirez-Gomez — 2010
- 42harvnbSmith (2010)Smith — 2010
- 43harvnbSmith, Ghosh, Buckley (2010)Smith, Ghosh, Buckley — 2010
- 44harvnbRuppert, Fox, Barnes (2004) p. 872–929Ruppert, Fox, Barnes — 2004
- 45harvnbJames, Siikavuopio, Johansson (2018)James, Siikavuopio, Johansson — 2018
- 46harvnbEdmondson (1935)Edmondson — 1935
- 47harvnbMcAlary (1993)McAlary — 1993
- 48harvnbHotchkiss (2000) p. last paragraph in review above AnalysisHotchkiss — 2000
- 49harvnbFisher (1925)Fisher — 1925
- 50harvnbDobson, Stancyk, Clements (1991)Dobson, Stancyk, Clements — 1991
- 51harvnbCandia-Carnevali, Thorndyke, Matranga (2009)Candia-Carnevali, Thorndyke, Matranga — 2009
- 52harvnbMashanov, Dolmatov, Heinzeller (2005)Mashanov, Dolmatov, Heinzeller — 2005
- 53harvnbYoung, Eckelbarger (1994) p. 179–194Young, Eckelbarger — 1994
- 54harvnbRuppert, Fox, Barnes (2004) p. 887–888Ruppert, Fox, Barnes — 2004
- 55harvnbRuppert, Fox, Barnes (2004) p. 895Ruppert, Fox, Barnes — 2004
- 56harvnbRuppert, Fox, Barnes (2004) p. 888Ruppert, Fox, Barnes — 2004
- 57harvnbRuppert, Fox, Barnes (2004) p. 908Ruppert, Fox, Barnes — 2004
- 58harvnbRuppert, Fox, Barnes (2004) p. 916Ruppert, Fox, Barnes — 2004
- 59harvnbRuppert, Fox, Barnes (2004) p. 922Ruppert, Fox, Barnes — 2004
- 60harvnbYamaguchi, Lucas (1984)Yamaguchi, Lucas — 1984
- 61harvnbMcGovern (2002)McGovern — 2002
- 62harvnbMonks (1904)Monks — 1904
- 63harvnbKille (1942)Kille — 1942
- 64harvnbEaves, Palmer (2003)Eaves, Palmer — 2003
- 65harvnbJaeckle (1994)Jaeckle — 1994
- 66harvnbVaughn (2009)Vaughn — 2009
- 67harvnbMcDonald, Vaughn (2010)McDonald, Vaughn — 2010
- 68harvnbVaughn, Strathmann (2008)Vaughn, Strathmann — 2008
- 69harvnbVaughn (2010)Vaughn — 2010
- 70harvnbDorit, Walker, Barnes (1991) p. 778Dorit, Walker, Barnes — 1991
- 71harvnbBrusca, Moore, Shuster (2016) p. 997–998Brusca, Moore, Shuster — 2016
- 72harvnbvan Egmond (2000)van Egmond — 2000
- 73harvnbBrusca, Moore, Shuster (2016) p. 968Brusca, Moore, Shuster — 2016
- 74harvnbWang, Hagdorn, Wang (2006)Wang, Hagdorn, Wang — 2006
- 75harvnbSmith (1937)Smith — 1937
- 76harvnbAstley (2012)Astley — 2012
- 77harvnbBrusca, Moore, Shuster (2016) p. 982–983Brusca, Moore, Shuster — 2016
- 78harvnbSea Stars of the Pacific Northwest p. Sand Star – ''Luidia foliolata''Sea Stars of the Pacific Northwest
- 79harvnbRuppert, Fox, Barnes (2004) p. 899–900Ruppert, Fox, Barnes — 2004
- 80harvnbRuppert, Fox, Barnes (2004) p. 911–912Ruppert, Fox, Barnes — 2004
- 81harvnbMessing (2006)Messing — 2006
- 82harvnbRuppert, Fox, Barnes (2004) p. 893Ruppert, Fox, Barnes — 2004
- 83harvnbRuppert, Fox, Barnes (2004) p. 914Ruppert, Fox, Barnes — 2004
- 84harvnbRuppert, Fox, Barnes (2004) p. 884–885Ruppert, Fox, Barnes — 2004
- 85harvnbBarnes (1982) p. 997–1007Barnes — 1982
- 86harvnbCarefoot, 2011a
- 87harvnbCarefoot, 2011b
- 88harvnbCarefoot, 2011c
- 89harvnbMiller (1998)Miller — 1998
- 90harvnbDorit, Walker, Barnes (1991) p. 779Dorit, Walker, Barnes — 1991
- 91harvnbDorit, Walker, Barnes (1991) p. 789–790Dorit, Walker, Barnes — 1991
- 92harvnbMladenov, Igdoura, Asotra (1989)Mladenov, Igdoura, Asotra — 1989
- 93harvnbHerrera-Escalante, López-Pérez, Leyte-Morales (2005)Herrera-Escalante, López-Pérez, Leyte-Morales — 2005
- 94harvnbLebrato, Iglesias-Rodríguez, Feely (2010)Lebrato, Iglesias-Rodríguez, Feely — 2010
- 95harvnbOsborne (2000) p. 464Osborne — 2000
- 96harvnbLawrence (1975)Lawrence — 1975
- 97harvnbBirkeland, Lucas (1990)Birkeland, Lucas — 1990
- 98harvnbWray (1999)Wray — 1999
- 99harvnbUbaghs (1978)Ubaghs — 1978
- 100harvnbSmith (1984)Smith — 1984
- 101harvnbRahman, Zamora (2024) p. 308–309Rahman, Zamora — 2024
- 102harvnbNanglu, Cole, Wright (2023)Nanglu, Cole, Wright — 2023
- 103harvnbDorit, Walker, Barnes (1991) p. 777–779Dorit, Walker, Barnes — 1991
- 104harvnbFox (2007)Fox — 2007
- 105harvnbEdgecombe, Giribet, Dunn (2011)Edgecombe, Giribet, Dunn — 2011
- 106harvnbShu, Conway Morris, Han (2004) p. 426–427Shu, Conway Morris, Han — 2004
- 107harvnbTopper, Guo, Clausen (2019) p. 5–7Topper, Guo, Clausen — 2019
- 108harvnbConway Morris, Halgedahl, Selden (2015) p. 4–6Conway Morris, Halgedahl, Selden — 2015
- 109harvnbZamora, Wright, Mooi (2020) p. 2Zamora, Wright, Mooi — 2020
- 110harvnbMussini, Smith, Vinther (2024) p. supplemental data figure S4Mussini, Smith, Vinther — 2024
- 111harvnbLi, Dunn, Murdock (2023) p. 159Li, Dunn, Murdock — 2023
- 112harvnbTelford, Lowe, Cameron (2014)Telford, Lowe, Cameron — 2014
- 113harvnbEscriva, Reich, Dunn (2015)Escriva, Reich, Dunn — 2015
- 114harvnbSmith (1984) p. 433–434Smith — 1984
- 115harvnbMoore (1966) p. U2–U3Moore — 1966
- 116harvnbSprinkle (1973) p. 3Sprinkle — 1973
- 117harvnbSprinkle (1973)Sprinkle — 1973
- 118harvnbSmith (1984) p. 436Smith — 1984
- 119harvnbMoore (1966)Moore — 1966
- 120harvnbUbaghs (1967)Ubaghs — 1967
- 121harvnbSprinkle, 1980a
- 122harvnbSprinkle, 1980b
- 123harvnbSmith (1984) p. 436–439, 453–454Smith — 1984
- 124harvnbPBDB Somasteroidea
- 125harvnbPBDB Stenuroidea
- 126harvnbPBDB Coronoidea
- 127harvnbRowe, Baker, Clark (1988)Rowe, Baker, Clark — 1988
- 128harvnbRahman (2009)Rahman — 2009
- 129harvnbRozhnov (2022) p. 1310Rozhnov — 2022
- 130harvnbGee (1996) p. 223Gee — 1996
- 131harvnbLefebvre, Guensburg, Martin (2019)Lefebvre, Guensburg, Martin — 2019
- 132harvnbRahman, Zamora (2024) p. 308Rahman, Zamora — 2024
- 133harvnbDavid, Mooi (1999) p. 100David, Mooi — 1999
- 134harvnbDavid, Lefebvre, Mooi (2000) p. 549David, Lefebvre, Mooi — 2000
- 135harvnbRahman, Zamora (2024) p. 300Rahman, Zamora — 2024
- 136harvnbDavid, Lefebvre, Mooi (2000) p. 550David, Lefebvre, Mooi — 2000
- 137harvnbRahman, Zamora (2024) p. 310Rahman, Zamora — 2024
- 138harvnbRahman, Zamora (2024) p. 311Rahman, Zamora — 2024
- 139harvnbRahman, Zamora (2024) p. 298Rahman, Zamora — 2024
- 140harvnbRahman, Zamora (2024) p. 296Rahman, Zamora — 2024
- 141harvnbWaggoner (1995)Waggoner — 1995
- 142harvnbRahman, Zamora (2024) p. 299–303Rahman, Zamora — 2024
- 143harvnbSmith (2005)Smith — 2005
- 144harvnbZamora, Wright, Mooi (2020)Zamora, Wright, Mooi — 2020
- 145harvnbTopper, Guo, Clausen (2020)Topper, Guo, Clausen — 2020
- 146harvnbDorit, Walker, Barnes (1991) p. 792–793Dorit, Walker, Barnes — 1991
- 147harvnbDornbos, Bottjer (2000)Dornbos, Bottjer — 2000
- 148harvnbUCMP Berkeley p. Echinodermata: MorphologyUCMP Berkeley
- 149harvnbFAO p. B-76FAO
- 150harvnbPangestuti, Arifin (2018)Pangestuti, Arifin — 2018
- 151harvnbNYTimes (2009)NYTimes — 2009
- 152harvnbLawrence (2001)Lawrence — 2001
- 153harvnbDavidson (2014) p. 730Davidson — 2014
- 154harvnbSartori, Scuderi, Sansone (2015)Sartori, Scuderi, Sansone — 2015
- 155harvnbExploratorium
- 156harvnbSartori, Gaion (2015)Sartori, Gaion — 2015
- 157harvnbGaion, Scuderi, Pellegrini (2013)Gaion, Scuderi, Pellegrini — 2013
- 158harvnbHart (2002)Hart — 2002
- 159harvnbLongo, Anderson (1969)Longo, Anderson — 1969
- 160harvnbArshinoff, Cary, Karimi (2022)Arshinoff, Cary, Karimi — 2022
- 161harvnbTelmer, Karimi, Chess (2024)Telmer, Karimi, Chess — 2024
- 162harvnbWild Singapore p. Sea StarsWild Singapore
- 163harvnbBarkhouse, Niles, Davidson (2007)Barkhouse, Niles, Davidson — 2007