View source for Ostracod

{{Short description|Class of crustaceans}}
{{Automatic taxobox
| fossil_range = {{fossil range|Ordovician|Recent}}<ref>{{cite journal |last1=Williams |first1=Mark |last2=Siveter |first2=David J. |last3=Salas |first3=María José |last4=Vannier |first4=Jean |last5=Popov |first5=Leonid E. |last6=Ghobadi Pour |first6=Mansoureh |title=The earliest ostracods: the geological evidence |journal=Senckenbergiana Lethaea |date=June 2008 |volume=88 |issue=1 |pages=11–21 |doi=10.1007/BF03043974 }}</ref>
| image = Ostracod.JPG
| image_alt = 
| image_caption = 
| taxon = Ostracoda
| authority = [[Pierre André Latreille|Latreille]], 1802
| synonyms = 
| synonyms_ref = 
| subdivision_ranks = Subclasses and orders
| subdivision = 
*[[Myodocopa]] <small> [[Georg Ossian Sars|Sars]], 1866</small>
**[[Myodocopida]] <small>Sars, 1866</small>
**[[Halocyprida]] <small>[[James Dwight Dana|Dana]], 1853</small>
*[[Podocopa]] <small> Sars, 1866</small>
**[[Palaeocopida]] <small>Henningsmoen, 1953</small>
**[[Platycopida]] <small>Sars, 1866</small>
**[[Podocopida]] <small>Sars, 1866</small>
}}

'''Ostracods''', or ostracodes, are a [[Class (biology)|class]] of the [[crustacean|Crustacea]] (class '''Ostracoda'''), sometimes known as '''seed shrimp'''. Some 33,000 species (only 13,000 of which are [[extant taxon|extant]]) have been identified,<ref name=":0">'''Brandão, S.N.; Antonietto, L.S; Nery, D.G.; Santos, S.G.; Karanovic, I. (2023). World Ostracoda Database. Accessed at https://www.marinespecies.org/ostracoda  on 2023-09-12. {{doi|10.14284/364}}'''</ref> grouped into 7 valid orders.<ref name=":0" /> They are small crustaceans, typically around {{convert|1|mm|in|abbr=on|sigfig=1}} in size, but varying from {{convert|0.2|to|32|mm|in|abbr=on|sigfig=2}}, the latter in the case of the marine ''[[Gigantocypris]].'' The largest known freshwater species is ''[[Megalocypris princeps]]'', which reach 8&nbsp;mm in length.<ref>[https://books.google.com/books?id=5Q1bDwAAQBAJ&dq=South+Africa+pride+largest+living+Megalocypris+princeps&pg=PT837 Freshwater Life: A field guide to the plants and animals of southern Africa]</ref><ref>[https://books.google.com/books?id=ulvmCQAAQBAJ&dq=Ostracods+adult+interstitial+forms+0.2+mm&pg=PA427 The Triassic-Jurassic Terrestrial Transition: 37]</ref> In most cases, their bodies are flattened from side to side and protected by a [[bivalve]]-like valve or "shell" made of [[chitin]], and often calcium carbonate. The family [[Entocytheridae]] and many planktonic forms do not have calcium carbonate.<ref>{{Cite thesis |last=Jöst |first=Anna B. |title=Ecological Evaluation of Ostracode (Crustacea) Occurrence in the City of Munich & Phylogenetic Relationships between Ostracode Species and Populations from Bavaria |date=January 2012 |degree=master |publisher=[[Ludwig Maximilian University of Munich]] |url=https://www.researchgate.net/publication/331485946}}</ref><ref>{{cite book |doi=10.1093/oso/9780195365764.003.0010 |chapter=Ostracoda |title=Evolution and Phylogeny of Pancrustacea |date=2022 |last1=Schram |first1=Frederick R. |last2=Koenemann |first2=Stefan |pages=119–140 |isbn=978-0-19-536576-4 }}</ref> The hinge of the two valves is in the upper (dorsal) region of the body. Ostracods are grouped together based on shell and soft part morphology, and molecular studies have not unequivocally supported the group's [[monophyly]].<ref name="oakley2013">{{cite journal |last1=Oakley |first1=Todd H. |last2=Wolfe |first2=Joanna M. |last3=Lindgren |first3=Annie R. |last4=Zaharoff |first4=Alexander K. |title=Phylotranscriptomics to Bring the Understudied into the Fold: Monophyletic Ostracoda, Fossil Placement, and Pancrustacean Phylogeny |journal=Molecular Biology and Evolution |date=January 2013 |volume=30 |issue=1 |pages=215–233 |doi=10.1093/molbev/mss216 |pmid=22977117 }}</ref> They have a wide range of diets, and the class includes carnivores, herbivores, scavengers and filter feeders, but most ostracods are [[Detritivores|deposit feeders]].

==Etymology==
''Ostracod'' comes from the [[Greek (language)|Greek]] ''óstrakon'' meaning shell or tile.

==Fossils==
[[File:HerrmanninaSilurianEstonia.jpg|thumb|The large ostracod ''Herrmannina'' from the [[Silurian]] (Ludlow) Soeginina Beds (Paadla Formation) on eastern [[Saaremaa]] Island, [[Estonia]]]]
Ostracods are "by far the most common arthropods in the fossil record"<ref>{{cite journal |author=[[David J. Siveter]] |author2=[[Derek E. G. Briggs]] |author3=Derek J. Siveter |author4=Mark D. Sutton |year=2010 |title=An exceptionally preserved myodocopid ostracod from the Silurian of Herefordshire, UK |journal=[[Proceedings of the Royal Society B]] |volume=277 |issue=1687 |pages=1539–1544 |doi=10.1098/rspb.2009.2122 |pmid=20106847 |pmc=2871837}}</ref> with fossils being found from the early [[Ordovician]] to the present. An outline [[microfauna]]l zonal scheme based on both [[Foraminifera]] and Ostracoda was compiled by M. B. Hart.<ref>{{cite journal |author=Malcolm B. Hart |year=1972 |title=A correlation of the macrofaunal and microfaunal zonations of the Gault Clay in southeast England |journal=[[Geological Journal]] |issue=Special Issue 5 |pages=267–288 |editor1=R. Casey |editor2=P. F. Rawson}}</ref> Freshwater ostracods have even been found in [[Baltic amber]] of [[Eocene]] age, having presumably been washed onto trees during floods.<ref>{{cite journal <!-- |book-title=Evolution and Diversity of Ostracoda. Theme 2 of the 14th International Symposium on Ostracoda (ISO 2001), Shizuoka, Japan --> |editor1=Noriyuki Ikeya |editor2=Akira Tsukagoshi |editor3=David J. Horne |title=Preface: The phylogeny, fossil record and ecological diversity of ostracod crustaceans |journal=[[Hydrobiologia]] |volume=538 |issue=1–3 |year=2005 |doi=10.1007/s10750-004-4914-z |pages=vii–xiii |author1=Noriyuki Ikeya, Akira Tsukagoshi  |author2=David J. Horne |s2cid=43836792 |name-list-style=amp }}</ref>

Ostracods have been particularly useful for the [[biostratigraphy|biozonation]] of marine strata on a local or regional scale, and they are invaluable indicators of paleoenvironments because of their widespread occurrence, small size, easily preservable, generally moulted, calcified bivalve carapaces; the valves are a commonly found [[microfossil]].

A find in [[Queensland, Australia]], in 2013, announced in May 2014, at the [[Bicentennary of Australia|Bicentennary]] Site in the [[Australian Fossil Mammal Sites (Riversleigh)|Riversleigh]] [[World Heritage]] area, revealed both male and female specimens with very well preserved soft tissue. This set the [[Guinness World Record]] for the oldest penis.<ref>[http://www.guinnessworldrecords.com/world-records/oldest-penis Oldest penis]:<br/>''The oldest fossilised penis discovered to date dates back around 100 million years. It belongs to a crustacean called an ostracod, discovered in Brazil and measuring just 1mm across.''</ref> Males had observable sperm that is the oldest yet seen and, when analysed, showed internal structures and has been assessed as being the largest sperm (per body size) of any animal recorded. It was assessed that the fossilisation was achieved within several days, due to phosphorus in the bat droppings of the cave where the ostracods were living.<ref>[http://www.abc.net.au/science/articles/2014/05/14/4003454.htm World's oldest sperm 'preserved in bat poo'], Anna Salleh, [[ABC Online]] Science, 14 May 2014, accessed 15 May 2014</ref>

==Description==
[[File:CypridinaMediterranea.png|thumb|left|upright|Anatomy of ''[[Cypridina mediterranea]]'']]
The body of an ostracod is encased by a carapace originating from the head region, and consists of two valves superficially resembling the shell of a clam. A distinction is made between the valve (hard parts) and the body with its appendages (soft parts). Studies of the embryonic development in [[Myodocopida]] show that the bivalved carapace develops from two independent buds of the carapace valves. As the two halves grow, they meet in the middle. In ''[[Manawa (crustacean)|Manawa]]'', an ostracod in the order [[Palaeocopida]], the carapace originates as a single element and during growth folds at the midline.<ref>{{cite journal |last1=Betts |first1=Marissa J. |last2=Brock |first2=Glenn A. |last3=Paterson |first3=John R. |title=Butterflies of the Cambrian benthos? Shield position in bradoriid arthropods |journal=Lethaia |date=October 2016 |volume=49 |issue=4 |pages=478–491 |doi=10.1111/let.12160 |bibcode=2016Letha..49..478B }}</ref><ref>{{cite journal |last1=Wakayama |first1=N. |title=Embryonic development clarifies polyphyly in ostracod crustaceans |journal=Journal of Zoology |date=December 2007 |volume=273 |issue=4 |pages=406–413 |doi=10.1111/j.1469-7998.2007.00344.x }}</ref>

===Body parts===
[[File:Ostracod_swimming_motions_20200520.gif|thumb|right|upright|Ostracod swimming motions (in real time)]]
The body consists of a head and [[thorax]], separated by a slight constriction. Unlike many other crustaceans, the body is not clearly divided into [[segmentation (biology)|segments]]. Most species have completely or partly lost their trunk segmentation, and there are no boundaries between the thorax and [[Abdomen#Other animals|abdomen]], and it has therefore been impossible to tell if the first pair of limbs after the maxillae belongs to the head or the thorax. With a few exceptions, like [[Platycopida|platycopids]] which have an 11-segmented trunk, the abdomen in ostracods has no visible segments.<ref>{{cite journal |last1=Ikuta |first1=Kyosuke |title=Expression of two ''engrailed'' genes in the embryo of ''Vargula hilgendorfii'' (Müller, 1890) (Ostracoda: Myodocopida) |journal=Journal of Crustacean Biology |date=18 January 2018 |volume=38 |issue=1 |pages=23–26 |doi=10.1093/jcbiol/rux099 }}</ref>

The head is the largest part of the body, and bears four pairs of appendages. Two pairs of well-developed [[antenna (biology)|antennae]] are used to swim through the water. In addition, there are a pair of [[mandible (arthropod)|mandibles]] and a pair of [[Maxilla (arthropod mouthpart)|maxillae]]. The thorax has three primary pairs of appendages. The first of these has different functions in different groups. It can be used for feeding ([[Cypridoidea]]) or for walking ([[Cytheroidea]]), and in some species it has evolved into a male clasping organ. The second pair is mainly used for locomotion, and the third is used for walking or cleaning, but can also be reduced or absent. Both the second and third pair are absent in suborder [[Cladocopina]].<ref>{{cite journal |last1=Kornicker |first1=Louis S. |last2=Sohn |first2=I. G. |title=Phylogeny, ontogeny & morphology of living and fossil Thaumatocypridacea (Myodocopa: Ostracoda) |journal=Smithsonian Contributions to Zoology |date=1976 |issue=219 |pages=1–124 |doi=10.5479/si.00810282.219 }}</ref> In the [[Myodocopina]] the third pair is a multisegmented cleaning organ that resembles a worm. Their external genitals seem to originate from the fusion of three to five appendages. The two "rami", or projections, from the tip of the tail point downward and slightly forward from the rear of the shell.<ref name=IZ>{{cite book |author=Robert D. Barnes |year=1982 |title= Invertebrate Zoology |publisher=[[Holt-Saunders International]] |location=[[Philadelphia]] |pages=680–683 |isbn=978-0-03-056747-6}}</ref><ref>{{cite book |doi=10.1016/B978-012370626-3.00184-8 |chapter=Ostracoda |title=Encyclopedia of Inland Waters |date=2009 |last1=Martens |first1=K. |last2=Horne |first2=D.J. |pages=405–414 |isbn=978-0-12-370626-3 }}</ref><ref>{{cite book|title=Recent Freshwater Ostracods of the World: Crustacea, Ostracoda, Podocopida|last=Karanovic|first=Ivana|publisher=Springer|year=2012|isbn=978-3-642-21809-5|doi=10.1007/978-3-642-21810-1|lccn=2011944255|pages=5–47|s2cid=40120445 }} [https://books.google.com/books?id=orzLHglZlEUC&dq=%22This%20appendage%20is%20absent%20in%20the%20order%20Platycopida%20(Podocopa)%20and%20Cladocopina%20(Myodocopa).%22&pg=PA40 Limited preview via Google Books]</ref>{{rp|40}}

All ostracods have a pair of "ventilatory appendages" that beat rhythmically, which create a water current between the body and the inner surface of the carapace. [[Podocopa]], the largest subclass, have no gills, [[heart]] or circulatory system, so the gas exchange take place all over the surface. The other subclass of ostracods, the Myodocopa, do have a heart, and the family [[Cylindroleberididae]] also have 6–8 lamellar gills. Certain other larger members of Myodocopa, even if they don't have gills, have a circulatory system where hemolymph sinuses absorbs oxygen through special areas on the inner wall of the carapace.<ref>{{Cite journal |last1=Williams |first1=Mark |last2=Vannier |first2=Jean |last3=Corbari |first3=Laure |last4=Massabuau |first4=Jean-Charles |date=2011 |title=Oxygen as a driver of early arthropod micro-benthos evolution |journal=PLOS ONE |volume=6 |issue=12 |pages=e28183 |doi=10.1371/journal.pone.0028183 |doi-access=free |pmc=3229522 |pmid=22164241|bibcode=2011PLoSO...628183W }}</ref><ref>{{cite journal|title=The early life history of tissue oxygenation in crustaceans: the strategy of the myodocopid ostracod ''Cylindroleberis mariae''|last1=Corbari|first1=Laure|last2=Carbonel|first2=Pierre|last3=Massabuau|first3=Jean-Charles|journal=Journal of Experimental Biology|year=2005|volume=208|issue=4|pages=661–670|doi=10.1242/jeb.01427|pmid=15695758 |s2cid=30226212 |doi-access=|bibcode=2005JExpB.208..661C }}</ref> In addition, the respiratory protein [[hemocyanin]] has been found in the two orders Myodocopida and Platycopida.<ref>{{Cite journal |last1=Marxen |first1=Julia C. |last2=Pick |first2=Christian |last3=Oakley |first3=Todd H. |last4=Burmester |first4=Thorsten |date=August 2014 |title=Occurrence of hemocyanin in ostracod crustaceans |url=https://pubmed.ncbi.nlm.nih.gov/25135304 |journal=Journal of Molecular Evolution |volume=79 |issue=1–2 |pages=3–11 |doi=10.1007/s00239-014-9636-x |pmid=25135304|bibcode=2014JMolE..79....3M }}</ref> Nitrogenous waste is excreted through glands on the maxillae, antennae, or both.<ref name=IZ/>

The primary sense of ostracods is likely touch, as they have several sensitive hairs on their bodies and appendages. Compound eyes are only found in [[Myodocopina]] within the Myodocopa.<ref>{{cite journal|last1=Oakley|first1=Todd H.|last2=Cunningham|first2=Clifford W.|title=Molecular phylogenetic evidence for the independent evolutionary origin of an arthropod compound eye|journal=Proceedings of the National Academy of Sciences USA|volume=99|issue=3|pages=1426–1430|year=2002|pmid=11818548|pmc=122207|doi=10.1073/pnas.032483599|bibcode=2002PNAS...99.1426O |doi-access=free}}</ref> The order [[Halocyprida]] in the same subclass is eyeless.<ref>[https://www.zobodat.at/pdf/BioEco_11_0123-0143.pdf The darkness syndrome in subsurface-shallow and deep-sea dwelling Ostracoda (Crustacea)]</ref> Podocopid ostracods have just a [[naupliar eye]] consisting of two lateral ocelli and a single ventral ocellus, but the ventral one is absent in some species.<ref name=IZ/><ref>{{cite journal|title=Functional morphology and light-gathering ability of podocopid ostracod eyes and the palaeontological implications|last=Tanaka|first=Gengo|journal=Zoological Journal of the Linnean Society|volume=147|issue=1|year=2006|pages=97–108|doi=10.1111/j.1096-3642.2006.00216.x|doi-access=free}}</ref><ref>{{cite journal|title=A gigantic marine ostracod (Crustacea: Myodocopa) trapped in mid-Cretaceous Burmese amber|last1=Xing|first1=Lida|last2=Sames|first2=Benjamin|last3=McKellar|first3=Ryan C.|last4=Xi|first4=Dangpeng|last5=Bai|first5=Ming|last6=Wan|first6=Xiaoqiao|journal=Scientific Reports|year=2018|volume=8|issue=1 |page=1365 |id=1365|doi=10.1038/s41598-018-19877-y|pmid=29358761 |pmc=5778021 |bibcode=2018NatSR...8.1365X |doi-access=free}}</ref> Platycopida was assumed to be completely eyeless, but two species, ''[[Keijcyoidea infralittoralis]]'' and ''[[Cytherella sordida]]'', have been found to both possess a nauplius eye too.<ref>{{cite journal|doi=10.1144/jm.29.1.81|doi-access=free|title=Talking about a re-evolution: blind alleys in ostracod phylogeny|last=Horne|first=David J.|year=2010|journal=Journal of Micropalaeontology|volume=29|issue=1|pages=81-85}}</ref>

==Palaeoclimatic reconstruction==
[[File:PermianOstracod.jpg|thumb|Articulated ostracod valves in cross-section from the [[Permian]] of central Texas; typical [[thin section]] view of an ostracod fossil]]
A new method is in development called mutual ostracod temperature range (MOTR), similar to the [[mutual climatic range]] (MCR) used for beetles, which can be used to infer palaeotemperatures.<ref name="Horne">{{cite journal |author=D. J. Horne |url=http://www.cosis.net/abstracts/EGU2007/00093/EGU2007-J-00093.pdf?PHPSESSID=e |title=A mutual temperature range method for European Quaternary non-marine Ostracoda |journal=Geophysical Research Abstracts |volume=9 |pages=00093 |year=2007 |format=[[Portable Document Format|PDF]]}}</ref> The ratio of oxygen-18 to oxygen-16 (δ18O) and the ratio of magnesium to calcium (Mg/Ca) in the calcite of ostracod valves can be used to infer information about past hydrological regimes, global ice volume and water temperatures.

==Distribution==
Ecologically, marine ostracods can be part of the [[zooplankton]] or (most commonly) are part of the [[benthos]], living on or inside the upper layer of the sea floor. Ostracods has been found as deep as 9,307 m (genus Krithe in family [[Krithidae]]).<ref>{{cite journal |last1=Brandão |first1=Simone N. |last2=Hoppema |first2=Mario |last3=Kamenev |first3=Gennady M. |last4=Karanovic |first4=Ivana |last5=Riehl |first5=Torben |last6=Tanaka |first6=Hayato |last7=Vital |first7=Helenice |last8=Yoo |first8=Hyunsu |last9=Brandt |first9=Angelika |title=Review of Ostracoda (Crustacea) living below the Carbonate Compensation Depth and the deepest record of a calcified ostracod |journal=Progress in Oceanography |date=November 2019 |volume=178 |pages=102144 |doi=10.1016/j.pocean.2019.102144 |bibcode=2019PrOce.17802144B }}</ref> Subclass Myodocopa and the two podocop orders Palaeocopida and Platycopida are restricted to marine environments (except for Platycopida which have a few brackish species),<ref>[https://ns-zooplankton.linnaeus.naturalis.nl/linnaeus_ng/app/views/species/taxon.php?id=131459&epi=210 Zooplankton and Micronekton of the North Sea 2.0 – Ordo Platycopida]</ref><ref>{{cite journal |last1=Horne |first1=David J. |title=Key Events in the Ecological Radiation of the Ostracoda |journal=The Paleontological Society Papers |date=November 2003 |volume=9 |pages=181–202 |doi=10.1017/S1089332600002205 }}</ref><ref>{{Cite web |title=News from mid-Cretaceous 'Burmese Amber' |url=https://fgga.univie.ac.at/news/news-views/detailansicht/news/news-from-mid-cretaceous-burmese-amber-1/ |website=fgga.univie.ac.at |language=de}}</ref> but we find non-marine forms in the four superfamilies Terrestricytheroidea, Cypridoidea, Darwinuloidea, and Cytheroidea in the order [[Podocopida]]. Terrestricytheroidea is semi-terrestrial and usually found in brackish and marine-influenced environments such as salt marshes, but not in freshwater.<ref>[https://books.google.com/books?id=LB-OAwAAQBAJ&dq=terrestricytheroidea+terrestrial+salt+marshes&pg=PA759 Thorp and Covich's Freshwater Invertebrates: Ecology and General Biology]</ref> The other three superfamilies also live in freshwater (Darwinuloidea is exclusively non-marine).<ref>[http://evolbiol.ru/docs/docs/large_files/lost_sex.pdf Lost Sex]</ref><ref>[https://edis.ifas.ufl.edu/publication/IN1260 Seed Shrimp, Mussel Shrimp (Freshwater Ostracods) scientific name: (Crustacea: Ostracoda: Podocopa)]</ref><ref>{{cite journal |last1=Jeffery |first1=Nicholas W |last2=Ellis |first2=Emily A |last3=Oakley |first3=Todd H |last4=Gregory |first4=T Ryan |title=The Genome Sizes of Ostracod Crustaceans Correlate with Body Size and Evolutionary History, but not Environment |journal=Journal of Heredity |date=September 2017 |volume=108 |issue=6 |pages=701–706 |doi=10.1093/jhered/esx055 |pmid=28595313 }}</ref> Of these three, only Cypridoidea have freshwater species able to swim.<ref>[https://books.google.com/books?id=PgFQCwAAQBAJ&dq=Nonswimmers+darwinulids+cytherids&pg=PA633 Thorp and Covich's Freshwater Invertebrates, Volume 5: Keys to Neotropical and Antarctic Fauna]</ref> Representatives living in terrestrial habitats are also found in all three freshwater groups,<ref>{{cite journal | pmc=3428786 | year=2012 | last1=Karanovic | first1=I. | last2=Eberhard | first2=S. | last3=Perina | first3=G. | title=''Austromesocypris bluffensis'' sp. n. (Crustacea, Ostracoda, Cypridoidea, Scottiinae) from subterranean aquatic habitats in Tasmania, with a key to world species of the subfamily | journal=ZooKeys | issue=215 | pages=1–31 | doi=10.3897/zookeys.215.2987 | pmid=22936868 | doi-access=free | bibcode=2012ZooK..215....1K }}</ref> such as genus ''[[Mesocypris]]'' which is known from humid forest soils of [[South Africa]], [[Australia]] and [[New Zealand]].<ref>{{cite journal |url=https://nzetc.victoria.ac.nz/tm/scholarly/tei-Bio11Tuat02-t1-body-d1.html |journal=[[Tuatara (journal)|Tuatara]] |volume=11 |issue=2|year=1963 |title=The Terrestrial Plankton |author=J. D. Stout |pages=57–65}}</ref>

As of 2008, around 2000 species and 200 genera of non-marine ostracods are found.<ref name=Martens08>{{cite journal |journal=Hydrobiologia |volume=595 |issue=1|year=2008|title=Global diversity of ostracods (Ostracoda, Crustacea) in freshwater |author1=K. Martens |author2=I. Schon |author3=C. Meisch |author4=D. J. Horne |s2cid=207150861 |pages=185–193 |doi=10.1007/s10750-007-9245-4|bibcode=2008HyBio.595..185M }}</ref> However, a large portion of diversity is still undescribed, indicated by undocumented diversity hotspots of temporary habitats in Africa and Australia.<ref name=Martens12>{{cite journal |url=http://www.europeanjournaloftaxonomy.eu/index.php/ejt/article/view/98 |journal=[[European Journal of Taxonomy (journal)|European Journal of Taxonomy]] |issue=8 |year=2012 |title=Nine new species of Bennelongia De Deckker & McKenzie, 1981 (Crustacea, Ostracoda) from Western Australia, with the description of a new subfamily |author=K. Martens, S. A. Halse & I. Schon |pages=1–56|doi=10.5852/ejt.2012.8 }}</ref> Non-marine species have been found to live in sulfidic cave ecosystems such as the [[Movile Cave]], deep groundwaters, hypersaline waters, acidic waters with pH as low as 3.4, [[phytotelma]]ta in plants like [[Bromeliaceae|bromeliads]], and in temperatures varying from almost freezing to more than 50&nbsp;°C in hot springs.<ref>{{cite journal | doi=10.1038/s41598-023-32573-w | title=A new extremophile ostracod crustacean from the Movile Cave sulfidic chemoautotrophic ecosystem in Romania | year=2023 | last1=Iepure | first1=Sanda | last2=Wysocka | first2=Anna | last3=Sarbu | first3=Serban M. | last4=Kijowska | first4=Michalina | last5=Namiotko | first5=Tadeusz | journal=Scientific Reports | volume=13 | issue=1 | page=6112 | pmid=37059813 | pmc=10104858 | bibcode=2023NatSR..13.6112I }}</ref><ref>{{cite journal | doi=10.1371/journal.pone.0248863 | doi-access=free | title=Copepods and ostracods associated with bromeliads in the Yucatán Peninsula, Mexico | year=2021 | last1=Mercado-Salas | first1=Nancy F. | last2=Khodami | first2=Sahar | last3=Martínez Arbizu | first3=Pedro | journal=PLOS ONE | volume=16 | issue=3 | pages=e0248863 | pmid=33735283 | pmc=7971893 | bibcode=2021PLoSO..1648863M }}</ref> Of the known specific and generic diversity of non-marine ostracods, half (1000 species, 100 genera) belongs to one family (of 13 families), [[Cyprididae]].<ref name=Martens12/> Many Cyprididae occur in temporary water bodies and have drought-resistant eggs, mixed/[[parthenogenetic]] reproduction, and the ability to swim.  These biological attributes preadapt them to form successful radiations in these habitats.<ref>{{cite book |last1=Horne |first1=D. J. |last2=Martens |first2=Koen |editor-last1=Brendonck |editor-first1=L. |editor-last2=De Meester |editor-first2=L. |editor-last3=Hairston |editor-first3=N. |title=Evolutionary and ecological aspects of crustacean diapause |publisher=Advances in Limnology |year=1998 |volume=52 |pages=549–561 |chapter=An assessment of the importance of resting eggs for the evolutionary success of non-marine Ostracoda (Crustacea) |chapter-url=http://www.schweizerbart.de/publications/detail/isbn/9783510470549/Evolutionary_and_ecological_aspects_of_crustacean_diapause |isbn=9783510470549}}</ref>

==Ecology==

===Lifecycle===
Male ostracods have two [[penis]]es, corresponding to two genital openings ([[gonopore]]s) on the female. The individual sperm are often large, and are coiled up within the [[testis]] prior to mating; in some cases, the uncoiled sperm can be up to six times the length of the male ostracod itself. Mating typically occurs during swarming, with large numbers of females swimming to join the males. Some species are partially or wholly [[Parthenogenesis|parthenogenetic]].<ref name=IZ/> Superfamily Darwinuloidea was assumed to have reproduced asexually  for the last 200 million years, but rare males have since been discovered in one of the species.<ref>{{cite journal |last1=Smith |first1=Robin J |last2=Kamiya |first2=Takahiro |last3=Horne |first3=David J |title=Living males of the 'ancient asexual' Darwinulidae (Ostracoda: Crustacea) |journal=Proceedings of the Royal Society B: Biological Sciences |date=22 June 2006 |volume=273 |issue=1593 |pages=1569–1578 |doi=10.1098/rspb.2005.3452 |pmid=16777754 |pmc=1560310 }}</ref>
[[File:Karpvähiline.jpg|thumb|Ostracod]]
In the subclass Myodocopa, all members of the order Myodocopida have brood care, releasing their offspring as first instars, allowing a pelagic lifestyle. In the order Halocyprida the eggs are released directly into the sea, except for a single genus with brood care. In the subclass Podocopa, brood care is only found in Darwinulocopina and some Cytherocopina in the order Podocopida. In the remaining Podocopa it is common to glue the eggs to a firm surface, like vegetation or the substratum. These eggs are often resting eggs, and remain dormant during desiccation and extreme temperatures, only hatching when exposed to more favorable conditions.<ref>[https://books.google.com/books?id=XYOPpkBuuTIC&dq=%22avoid+brooding+and+simply+glue+the+eggs+to+a+firm+surface+and+then+abandon+them%22&pg=PA141 Field Guide to Freshwater Invertebrates of North America]</ref><ref>[https://books.google.com/books?id=orzLHglZlEUC&dq=%22In+the+subclass+Myodocopa%2C+all+representatives+of+Myodocopida+have+brood+care%22&pg=PA59 Recent Freshwater Ostracods of the World: Crustacea, Ostracoda, Podocopida]</ref> Species adapted to [[vernal pool]]s can reach sexual maturity in just 30 days after hatching.<ref>[https://www.theguardian.com/environment/2019/mar/26/specieswatch-ancient-crustaceans-still-going-strong-after-450m-years Specieswatch: ancient crustaceans still going strong after 450m years]</ref> There is no larval stage or metamorphosis ([[direct development]]). Instead they hatch from the egg as juveniles with the bivalved carapace and at least three functional limbs. As the juvenile grows through a series of molts they acquire more limbs and develop further the already existing ones.<ref>[https://marinescience.ucdavis.edu/sites/g/files/dgvnsk9221/files/inline-files/BMR_Ostracoda.pdf The Light and Smith Manual : Intertidal Invertebrates from Central California to Oregon – Page 419]</ref> They reach sexual maturity in the final instar and then never molts again. The number of instars they go through before adulthood varies. In Podocopa it is eight or nine (but family Entocytheridae and suborder Bairdiocopina has only seven),<ref>[https://books.google.com/books?id=orzLHglZlEUC&dq=Entocytheridae+Bairdiocopina+seven+instars&pg=PA62 Recent Freshwater Ostracods of the World: Crustacea, Ostracoda, Podocopida]</ref> the Halocyprida goes through six or seven, and Myodocopida only four to six. They are able to produce several offspring many times as adults ([[Semelparity and iteroparity#Iteroparity|iteroparity]]).<ref>[https://media.australian.museum/media/Uploads/Journals/17619/378_complete.pdf Strategies of crustacean growth - Australian Museum]</ref><ref>[https://books.google.com/books?id=1FQPEAAAQBAJ&dq=%22Ostracods+become+mature+only+in+the+final+instar%22&pg=PT131 Crustacean Issues 3: Factors in Adult Growth]</ref><ref>[https://books.google.com/books?id=Z2tEEAAAQBAJ&dq=Podocopa+eight+nine+stages+Myodocopa+Halocyprida+six+seven+Myodocopida+four&pg=PA130 Evolution and Phylogeny of Pancrustacea: A Story of Scientific Method]</ref>

===Predators===
A variety of fauna prey upon ostracods in both aquatic and terrestrial environments. An example of predation in the marine environment is the action of certain Cytherocopina in the [[Cuspidariidae|cuspidariid clams]] in detecting ostracods with [[cilia]] protruding from inhalant structures, thence drawing the ostracod prey in by a violent suction action.<ref>{{cite book |author1=John D. Gage  |author2=Paul A. Tyler  |name-list-style=amp |title=Deep-Sea Biology: A Natural History of Organisms at the Deep-Sea Floor |publisher=[[University of Southampton]] |isbn=978-0-521-33665-9|date=1992-09-28  }}</ref> Predation from higher animals also occurs; for example, amphibians such as the [[rough-skinned newt]] prey upon certain ostracods.<ref>{{cite web |author=C. Michael Hogan |year=2008 |title=''Rough-skinned Newt ("Taricha granulosa")'' |website=[[Globaltwitcher]], ed. N. Stromberg |url=http://www.globaltwitcher.com/artspec_information.asp?thingid=43182 |url-status=dead |archive-url=https://web.archive.org/web/20090527153302/http://www.globaltwitcher.com/artspec_information.asp?thingid=43182 |archive-date=2009-05-27 }}</ref> Whale sharks also seem to eat them as part of their filter feeding process.<ref>{{cite journal |last1=Rohner |first1=Ca |last2=Couturier |first2=Lie |last3=Richardson |first3=Aj |last4=Pierce |first4=Sj |last5=Prebble |first5=Cem |last6=Gibbons |first6=Mj |last7=Nichols |first7=Pd |title=Diet of whale sharks Rhincodon typus inferred from stomach content and signature fatty acid analyses |journal=Marine Ecology Progress Series |date=20 November 2013 |volume=493 |pages=219–235 |doi=10.3354/meps10500 |bibcode=2013MEPS..493..219R }}</ref>



===Bioluminescence===
Some ostracods, such as ''[[Vargula hilgendorfii]]'', have a light organ in which they produce luminescent chemicals.<ref>{{Cite book |title=Bioluminescence: Chemical Principles and Methods |chapter=The ostracod ''Cypridina'' (''Vargula'') and other luminous crustaceans |author=Osamu Shimomura |chapter-url=https://books.google.com/books?id=DNrTfH5PcWoC&pg=PA49 |isbn=978-981-256-801-4 |publisher=[[World Scientific]] |year=2006 |pages=47–89}}</ref> These ostracods are called "blue sand" or "blue tears" and glow blue in the dark. Their [[bioluminescent]] properties made them valuable to the Japanese during [[World War II]], when the Japanese army collected large amounts from the ocean to use as a convenient light for reading maps and other papers at night. The light from these ostracods, called ''umihotaru'' in Japanese, was sufficient to read by but not bright enough to give away troops' position to enemies.<ref name="jabr">{{cite web|last1=Jabr|first1=Ferris|title=The Secret History of Bioluminescence|url=https://www.hakaimagazine.com/article-long/secret-history-bioluminescence|website=Hakai Magazine|access-date=6 July 2016}}</ref> Bioluminescence has evolved twice in ostracods; once in [[Cypridinidae]], and once in [[Halocyprididae]].<ref>{{cite journal |last1=Cohen |first1=Anne C. |last2=Oakley |first2=Todd H. |title=Collecting and processing marine ostracods |journal=Journal of Crustacean Biology |date=May 2017 |volume=37 |issue=3 |pages=347–352 |doi=10.1093/jcbiol/rux027 }}</ref> In bioluminescent Halocyprididae a green light is produced within carapace glands, and in Cypridinidae a blue light is produced and extruded from the upper lip.<ref>{{cite journal |last1=Cohen |first1=Anne C. |last2=Morin |first2=James G. |title=Sexual Morphology, Reproduction and the Evolution of Bioluminescence in Ostracoda |journal=The Paleontological Society Papers |date=November 2003 |volume=9 |pages=37–70 |doi=10.1017/S108933260000214X }}</ref><ref>{{cite journal |last1=Morin |first1=James G. |last2=Cohen |first2=Anne C. |title=Two New Bioluminescent Ostracode Genera, Enewton And Photeros (Myodocopida: Cypridinidae), with Three New Species from Jamaica |journal=Journal of Crustacean Biology |date=2010 |volume=30 |issue=1 |pages=1–55 |doi=10.1651/08-3075.1 |bibcode=2010JCBio..30....1M }}</ref> Most species use the light as predation defense, but the male of at least 75 known species of the Cypridinidae, restricted to the Caribbean, use pulses of light to attract females. Some species are the opposite where the females use pulses of light to attract males. This is seen in one example such as the [[Lampyris noctiluca|glow worm]]. This bioluminescent [[courtship display]] has only evolved once in ostracods, in a cypridinid group named Luxorina that originated at least 151 million years ago.<ref>{{cite journal |last1=Ellis |first1=Emily A |last2=Goodheart |first2=Jessica A |last3=Hensley |first3=Nicholai M |last4=González |first4=Vanessa L |last5=Reda |first5=Nicholas J |last6=Rivers |first6=Trevor J |last7=Morin |first7=James G |last8=Torres |first8=Elizabeth |last9=Gerrish |first9=Gretchen A |last10=Oakley |first10=Todd H |title=Sexual Signals Persist over Deep Time: Ancient Co-option of Bioluminescence for Courtship Displays in Cypridinid Ostracods |journal=Systematic Biology |date=16 June 2023 |volume=72 |issue=2 |pages=264–274 |doi=10.1093/sysbio/syac057 |pmid=35984328 |pmc=10448971 }}</ref><ref>{{cite journal |last1=Rivers |first1=Trevor J. |last2=Morin |first2=James G. |title=Female ostracods respond to and intercept artificial conspecific male luminescent courtship displays |journal=Behavioral Ecology |date=2013 |volume=24 |issue=4 |pages=877–887 |doi=10.1093/beheco/art022 }}</ref> Ostracods with bioluminescent courtship show higher rates of [[speciation]] than those who simply use light as protection against predators.<ref>[https://nautil.us/bioluminescence-is-natures-love-light-243320/ Bioluminescence Is Nature's Love Light]</ref> The male will continue to swim after releasing its small ball of bioluminescent mucus, but the female is able to read the display to pinpoint the male's location.<ref>[https://www.nytimes.com/2023/11/28/science/sea-fireflies-glowing.html Watch ‘Sea Fireflies’ Make Underwater Fireworks as They Seek Mates]</ref> In one species hundreds of thousands of males synchronize their light display, and when one male creates a pattern of light, the new pattern will spread out as the neighboring males repeat it.<ref>[https://www.cbc.ca/radio/asithappens/sea-firefly-mating-dance-1.7046972 Mating dance of sea fireflies is 'the coolest fireworks show that you've ever seen']</ref>

==Classification==

Early work indicated that Ostracoda may not be [[monophyly|monophyletic]],<ref>{{cite book |author1=Richard A. Fortey  |author2=Richard H. Thomas  |name-list-style=amp |title=Arthropod Relationships |year=1998 |isbn=978-0-412-75420-3 |publisher=[[Chapman & Hall]]}}</ref> and early [[molecular phylogeny]] was ambiguous on this front.<ref>{{cite journal |author1=S. Yamaguchi  |author2=K. Endo  |s2cid=83831572 |name-list-style=amp |title=Molecular phylogeny of Ostracoda (Crustacea) inferred from 18S ribosomal DNA sequences: implication for its origin and diversification |journal=[[Marine Biology (journal)|Marine Biology]] |volume=143 |issue=1 |pages=23–38 |year=2003 |doi=10.1007/s00227-003-1062-3|bibcode=2003MarBi.143...23Y }}</ref> Recent combined analyses of molecular and morphological data suggested monophyly in analyses with broadest taxon sampling, but this monophyly had no or very little support (Fig. 1 - bootstrap 0, 17 and 46, often values above 95 are considered sufficient for the taxon support).<ref name="oakley2013"/>

Class Ostracoda is divided into following living clades:<ref>[https://www.biwahaku.jp/smith/about.html Ostracods - what are they? - Lake Biwa Museum]</ref>
*Subclass Myodocopa
**Order Myodocopida
***Suborder [[Myodocopina]]
****Superfamily [[Cypridinoidea]] (1 family)
****Superfamily [[Cylindroleberidoidea]] (1 family)
****Superfamily [[Sarsielloidea]] (3 families)
**Order Halocyprida
***Suborder [[Halocypridina]]
****Superfamily [[Thaumatocypridoidea]] (1 family)
****Superfamily [[Halocypridoidea]] (1 family)
***Suborder [[Cladocopina]]
****Superfamily [[Cladocopoidea]] (1 family)
*Subclass Podocopa
**Order Palaeocopida
***Suborder [[Kirkbyocopina]]
****Superfamily [[Puncioidea]] (1 family)
**Order Platycopida
***Suborder [[Platycopina]]
****Superfamily [[Cytherelloidea]] (1 family)
**Order Podocopida
***Suborder [[Cytherocopina]]
****Superfamily [[Cytheroidea]] (27 families)
****Superfamily [[Terrestricytheroidea]] (1 family)
***Suborder [[Cypridocopina]]
****Superfamily [[Macrocypridoidea]] (1 family)
****Superfamily [[Pontocypridoidea]] (1 family)
****Superfamily [[Cypridoidea]] (4 families)
***Suborder [[Darwinulocopina]]
****Superfamily [[Darwinuloidea]] (1 family)
***Suborder [[Bairdiocopina]]
****Superfamily [[Bairdioidea]] (3 families)
***Suborder [[Sigilliocopina]]
****Superfamily [[Sigillioidea]] (1 family)

==See also==
{{Portal|Crustaceans}}
*[[Mari Mari Group]], fossil formation in the [[Amazonas (Brazilian state)|state of Amazonas]] of northwestern Brazil

==References==
{{Reflist|32em}}

==External links==
{{Wikispecies|Ostracoda}}
* [http://ostracoda-on.tripod.com Kempf Database Ostracoda] 
* [https://web.archive.org/web/20081212060559/http://www.tafi.org.au/zooplankton/imagekey/ostracoda/ Ostracoda fact sheet],  Guide to the Marine Zooplankton of South-eastern Australia
* [http://home.comcast.net/~fireflea2/OstracodeKeyindex.html Key to the two subclasses]
* [http://userpage.fu-berlin.de/%7Epalaeont/irgo/irgohome.html International Research Group on Ostracoda]
* [https://web.archive.org/web/20090917190408/http://www.tafi.org.au/zooplankton/imagekey/ostracoda/index.html Ostracoda Fact Sheet]
* [http://news.bbc.co.uk/2/hi/science/nature/8107515.stm Huge sperm of ancient crustaceans]
* [http://www.marinespecies.org/ostracoda/index.php World Ostracoda Database]

{{Arthropods}}
{{Taxonbar|from=Q276412}}
{{Authority control}}

[[Category:Ostracods| ]]
[[Category:Late Ordovician first appearances]]
[[Category:Extant Ordovician first appearances]]