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{{short description|Order of crustaceans}}
{{Short description|Order of crustaceans}}
{{Automatic taxobox
{{Automatic taxobox
| fossil_range = {{fossil range|Ordovician|Recent}}
| fossil_range = {{fossil range|Devonian|Recent|earliest=500}}
| image = Artemia salina 4.jpg
| image = Artemia salina 4.jpg
| image_caption = ''[[Artemia salina]]''
| image_caption = ''[[Artemia salina]]''
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| taxon = Anostraca
| taxon = Anostraca
| authority = [[Georg Ossian Sars|G. O. Sars]], 1867
| authority = [[Georg Ossian Sars|G. O. Sars]], 1867
| subdivision_ranks = [[Family (biology)|Families]]&nbsp;<ref name="M&D">{{cite book |url=http://atiniui.nhm.org/pdfs/3839/3839.pdf |title=An Updated Classification of the Recent Crustacea |author1=Joel W. Martin |author2=George E. Davis |year=2001 |pages=1–132 |publisher=[[Natural History Museum of Los Angeles County]]}}</ref><ref name="Weekers">{{cite journal |author1=Peter H. H. Weekers |author2=Gopal Murugan |author3=Jacques R. Vanfleteren |author4=Denton Belk |author5=Henri J. Dumont |year=2002 |title=Phylogenetic analysis of anostracans (Branchiopoda: Anostraca) inferred from nuclear 18S ribosomal DNA (18S rDNA) sequences |journal=[[Molecular Phylogenetics and Evolution]] |volume=25 |issue=3 |pages=535–544 |pmid=12450757 |doi=10.1016/S1055-7903(02)00289-0 |url=http://arthroinfo.org/pdfs/3663/3663.pdf }}</ref>
| subdivision_ranks = [[Family (biology)|Families]]&nbsp;<ref name="M&D">{{cite book |url=http://atiniui.nhm.org/pdfs/3839/3839.pdf |title=An Updated Classification of the Recent Crustacea |author1=Joel W. Martin |author2=George E. Davis |year=2001 |pages=1–132 |publisher=[[Natural History Museum of Los Angeles County]] |access-date=2011-03-03 |archive-date=2013-05-12 |archive-url=https://web.archive.org/web/20130512091254/http://atiniui.nhm.org/pdfs/3839/3839.pdf |url-status=dead }}</ref><ref name="Weekers">{{cite journal |author1=Peter H. H. Weekers |author2=Gopal Murugan |author3=Jacques R. Vanfleteren |author4=Denton Belk |author5=Henri J. Dumont |year=2002 |title=Phylogenetic analysis of anostracans (Branchiopoda: Anostraca) inferred from nuclear 18S ribosomal DNA (18S rDNA) sequences |journal=[[Molecular Phylogenetics and Evolution]] |volume=25 |issue=3 |pages=535–544 |pmid=12450757 |doi=10.1016/S1055-7903(02)00289-0 |url=http://arthroinfo.org/pdfs/3663/3663.pdf }}</ref>
| subdivision =
| subdivision = *[[Brine shrimp|Artemiidae]] <small>Grochowski, 1896</small>
*[[Brine shrimp|Artemiidae]] <small>Grochowski, 1896</small>
* [[Branchinecta|Branchinectidae]] <small>Daday, 1910</small>
*[[Branchinecta|Branchinectidae]] <small>Daday, 1910</small>
* [[Branchipodidae]] <small>Simon, 1886</small>
*[[Branchipodidae]] <small>Simon, 1886</small>
* [[Chirocephalidae]] <small>Daday, 1910</small>
*[[Chirocephalidae]] <small>Daday, 1910</small>
* [[Parartemia|Parartemiidae]] <small>Daday, 1910</small>
*[[Parartemia|Parartemiidae]] <small>Daday, 1910</small>
* [[Streptocephalus|Streptocephalidae]] <small>Daday, 1910</small>
*[[Streptocephalus|Streptocephalidae]] <small>Daday, 1910</small>
* [[Tanymastigidae]] <small>Weekers ''et al.'', 2002</small>
*[[Tanymastigidae]] <small>Weekers ''et al.'', 2002</small>
* [[Thamnocephalidae]] <small>Simon, 1886</small>
*[[Thamnocephalidae]] <small>Simon, 1886</small>
}}
}}


'''Anostraca''' is one of the four [[order (biology)|orders]] of [[crustacean]]s in the class [[Branchiopoda]]; its members are also known as '''fairy shrimp'''. They live in [[vernal pool]]s and [[hypersaline lake]]s across the world, and they have even been found in [[desert]]s, ice-covered mountain lakes and [[Antarctica|Antarctic]] ice. They are usually {{convert|6|-|25|mm|abbr=on}} long (exceptionally up to {{convert|170|mm|abbr=on|disp=or}}). Most species have 20 body segments, bearing 11 pairs of leaf-like ''phyllopodia'' (swimming legs), and the body lacks a [[carapace]]. They swim "upside-down" and [[filter feeder|feed by filtering]] organic particles from the water or by scraping [[algae]] from surfaces. They are an important food for many birds and fish, and some are cultured and harvested for use as fish food. There are 300 species spread across 8 [[family (biology)|families]].
'''Anostraca''' is one of the four [[order (biology)|orders]] of [[crustacean]]s in the class [[Branchiopoda]]; its members are referred to as '''fairy shrimp'''. They live in [[vernal pool]]s<ref>{{Cite web |title=Vernal Pool Fairy Shrimp (Branchinecta lynchi) {{!}} U.S. Fish & Wildlife Service |url=https://www.fws.gov/species/vernal-pool-fairy-shrimp-branchinecta-lynchi |access-date=2024-11-06 |website=www.fws.gov |language=en}}</ref> and [[hypersaline lake]]s across the world, and they have even been found in [[desert]]s, ice-covered mountain lakes, and [[Antarctica|Antarctic]] ice.<ref name=":10">{{Cite web |title=Cape Ann Vernal Pond Team - Fairy Shrimp |url=https://www.capeannvernalpondteam.org/fairy-shrimp |access-date=2024-11-06 |website=www.capeannvernalpondteam.org}}</ref> They are usually {{convert|6|-|25|mm|abbr=on}} long (exceptionally up to {{convert|170|mm|abbr=on|disp=or}}). Most species have 20 body segments, bearing 11 pairs of leaf-like ''phyllopodia'' (swimming legs), and the body lacks a [[carapace]]. They swim "upside-down" and [[filter feeder|feed by filtering]] organic particles from the water or by scraping [[algae]] from surfaces,<ref name=":10" /> with the exception of ''[[Branchinecta gigas]],'' or "giant fairy shrimp", which is itself a predator of other species of anostracans. They are an important food for many birds and fish, and some are cultured and harvested for use as fish food. There are 300 species spread across 8 [[family (biology)|families]].<ref name=":10" />


==Description==
==Description==
The body of a fairy shrimp is elongated and divided into segments.<ref name="Belk"/> The whole animal is typically {{convert|6|-|25|mm}} long, but one species, ''[[Branchinecta gigas]]'' does not reach [[sexual maturity]] until it reaches {{convert|50|mm|abbr=on}} long, and can grow to {{convert|170|mm|abbr=on}} long.<ref name="Belk"/> The [[exoskeleton]] is thin and flexible,<ref name="Belk"/> and lacks any sign of a [[carapace]].<ref name="Australia">{{cite book |author=William David Williams |year=1980 |title=Australian Freshwater Life: the Invertebrates of Australian Inland Waters |edition=2nd |publisher=[[Palgrave Macmillan Australia]] |isbn=978-0-333-29894-7 |chapter=Arachnids and Crustaceans |pages=118–184 |chapter-url=https://books.google.com/books?id=2qqrV5WaJ4sC&pg=PA126}}</ref> The body can be divided into three distinct parts ([[Tagma (biology)|tagmata]]) – head, thorax and abdomen.<ref name="Australia"/>
The body of a fairy shrimp is elongated and divided into segments.<ref name="Belk"/> The whole animal is typically {{convert|6|-|25|mm}} long, but one species, ''[[Branchinecta gigas]]'' does not reach [[sexual maturity]] until it reaches {{convert|50|mm|abbr=on}} long, and can grow to {{convert|170|mm|abbr=on}} long.<ref name="Belk"/> The [[exoskeleton]] is thin and flexible,<ref name="Belk"/> and lacks any sign of a [[carapace]].<ref name="Australia">{{cite book |author=William David Williams |year=1980 |title=Australian Freshwater Life: the Invertebrates of Australian Inland Waters |edition=2nd |publisher=Palgrave Macmillan Australia |isbn=978-0-333-29894-7 |chapter=Arachnids and Crustaceans |pages=118–184 |chapter-url=https://books.google.com/books?id=2qqrV5WaJ4sC&pg=PA126}}</ref> The body can be divided into three distinct parts ([[Tagma (biology)|tagmata]]) – head, thorax and abdomen.<ref name="Australia"/>


===Head===
===Head===
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===Thorax and abdomen===
===Thorax and abdomen===
[[File:EubranchipusGrubiiMale+Female.jpg|thumb|upright|Male (top) and female (bottom) ''[[Eubranchipus grubii]]'' ([[Chirocephalidae]]): the female is holding eggs on her genital appendages.]]
[[File:EubranchipusGrubiiMale+Female.jpg|thumb|upright|Male (top) and female (bottom) ''[[Eubranchipus grubii]]'' ([[Chirocephalidae]]): the female is holding eggs on her genital appendages.]]
The thorax of most anostracans has 13 segments (19 in ''[[Polyartemiella]]'' and 21 in ''[[Polyartemia]]'').<ref name="Khanna">{{cite book |author=D. R. Khanna |year=2004 |title=Biology of Arthropoda |publisher=[[Discovery Publishing House]] |isbn=978-81-7141-897-8 |chapter=Segmentation in arthropods |pages=316–394 |chapter-url=https://books.google.com/books?id=Hd4OEDo4gbwC&pg=PA347}}</ref> All but the last two are very similar, with a pair of biramous phyllopods (flattened, leaf-like [[appendage]]s).<ref name="Australia"/> The last two segments are fused together,<ref name="Belk"/> and their appendages are specialised for reproduction.<ref name="Khanna"/> Most anostracans have separate sexes ([[gonochorism]]), but a few reproduce by [[parthenogenesis]].<ref name="Bell">{{Cite book |author=Graham Bell |year=1982 |title=The Masterpiece of Nature: the Evolution and Genetics of Sexuality |publisher=[[Cambridge University Press]] |isbn=978-0-85664-753-6 |chapter=Arthropoda: Crustacea Branchiopoda |pages=239–248 |chapter-url=https://books.google.com/books?id=q5g9AAAAIAAJ&pg=PA239}}</ref> The abdomen comprises 6 segments without appendages, and a [[telson]],<ref name="Khanna"/> which bears two flattened caudal rami or "cercopods".<ref name="Belk"/>
The thorax of most anostracans has 13 segments (19 in ''[[Polyartemiella]]'' and 21 in ''[[Polyartemia]]'').<ref name="Khanna">{{cite book |author=D. R. Khanna |year=2004 |title=Biology of Arthropoda |publisher=Discovery Publishing House |isbn=978-81-7141-897-8 |chapter=Segmentation in arthropods |pages=316–394 |chapter-url=https://books.google.com/books?id=Hd4OEDo4gbwC&pg=PA347}}</ref> All but the last two are very similar, with a pair of biramous phyllopods (flattened, leaf-like [[appendage]]s).<ref name="Australia"/> The last two segments are fused together,<ref name="Belk"/> and their appendages are specialised for reproduction.<ref name="Khanna"/> Most anostracans have separate sexes ([[gonochorism]]), but a few reproduce by [[parthenogenesis]].<ref name="Bell">{{Cite book |author=Graham Bell |year=1982 |title=The Masterpiece of Nature: the Evolution and Genetics of Sexuality |publisher=[[Cambridge University Press]] |isbn=978-0-85664-753-6 |chapter=Arthropoda: Crustacea Branchiopoda |pages=239–248 |chapter-url=https://books.google.com/books?id=q5g9AAAAIAAJ&pg=PA239}}</ref> The abdomen comprises 6 segments without appendages, and a [[telson]],<ref name="Khanna"/> which bears two flattened caudal rami or "cercopods".<ref name="Belk"/>


===Internal anatomy===
===Internal anatomy===
The head contains two digestive glands and the small lobate [[stomach]] that they empty into. This is connected to a long [[intestine]], which terminates in a short [[rectum]], with the [[anus]] located on the [[telson]].<ref name="Pennak"/> The [[haemocoel]] of anostracans is pumped by a long, tubular [[heart]], which runs through most of the animal's length.<ref name="Pennak"/> A series of slits allow haemocoel into the heart, which is then pumped out of the anterior opening by [[peristalsis]].<ref name="Pennak"/> The nervous system consists of two nerve cords which run the length of the body, with two [[ganglion|ganglia]] and two transverse [[commissure]]s in most of the body segments.<ref name="Pennak"/>
The head contains two digestive glands and the small lobate [[stomach]] into which they empty. This is connected to a long [[intestine]], which terminates in a short [[rectum]], with the [[anus]] located on the [[telson]].<ref name="Pennak"/> The [[haemocoel]] of anostracans is pumped by a long, tubular [[heart]], which runs through most of the animal's length.<ref name="Pennak"/> A series of slits allow haemocoel into the heart, which is then pumped out of the anterior opening by [[peristalsis]].<ref name="Pennak"/> The nervous system consists of two nerve cords which run the length of the body, with two [[ganglion|ganglia]] and two transverse [[commissure]]s in most of the body segments.<ref name="Pennak"/>


[[Gas exchange]] is thought to take place through the entire body surface, but especially that of the phyllopodia and their associated gills, which may also be responsible for [[osmoregulation|osmotic regulation]].<ref name="Pennak"/> Two coiled [[gland]]s at the bases of the maxillae are used to [[excretion|excrete]] [[nitrogenous waste]], typically in the form of [[urea]].<ref name="Pennak"/> Most of the animal's nitrogenous waste is, however, in the form of [[ammonia]], which probably diffuses into the environment through the phyllopodia and gills.<ref name="Pennak"/>
[[Gas exchange]] is thought to take place through the entire body surface, but especially that of the phyllopodia and their associated gills, which may also be responsible for [[osmoregulation|osmotic regulation]].<ref name="Pennak"/> Two coiled [[gland]]s at the bases of the maxillae are used to [[excretion|excrete]] [[nitrogenous waste]], typically in the form of [[urea]].<ref name="Pennak"/> Most of the animal's nitrogenous waste is, however, in the form of [[ammonia]], which probably diffuses into the environment through the phyllopodia and gills.<ref name="Pennak"/>


==Ecology and behaviour==
==Ecology and behaviour==
Anostracans inhabit inland waters ranging from [[hypersaline lake]]s to lakes that are almost devoid of dissolved substances;<ref name="Belk">{{cite book |editor1=Sol Felty Light |editor2=James T. Carlton |year=2007 |title=The Light and Smith Manual: Intertidal Invertebrates from Central California to Oregon |edition=4th |publisher=[[University of California Press]] |isbn=978-0-520-23939-5|chapter=Branchiopoda |author=Denton Belk |pages=414–417 |chapter-url=https://books.google.com/books?id=64jgZ1CfmB8C&pg=PA416}}</ref> they are "the most archetypal crustaceans" in ephemeral waters.<ref name="Dumont">{{cite book |year=2009 |editor=Henri J. Dumont |title=The Nile: Origins, Environments, Limnology and Human Use |series=Volume 89 of Monographiae Biologicae |publisher=[[Springer Science+Business Media|Springer]] |isbn=978-1-4020-9725-6 |chapter=The crustacean zooplankton (Copepoda, Branchiopoda), atyid Decapoda, and Syncarida of the Nile Basin |author=Henri J. Dumont |pages=521–546 |chapter-url=https://books.google.com/books?id=iF_U1NoknHoC&pg=PA536}}</ref> The relatively large size of fairy shrimp, together with their slow means of locomotion, makes them an easy target for [[predator]]y [[fish]] and [[water bird|waterfowl]].<ref name="Dumont"/> This has led to their distribution being restricted to environments with fewer predators, such as [[vernal pool]]s, [[salt lake]]s and lakes at high altitudes or latitudes.<ref name="Dumont"/> The southernmost recorded fairy shrimp is ''[[Branchinecta gaini]]'' from the [[Antarctic Peninsula]],<ref>{{cite journal |author=T.C. Hawes |date=October 2009 |title=Origins and dispersal of the Antarctic fairy shrimp |journal=[[Antarctic Science]] |volume=21 |issue=5 |pages=477–482 |doi=10.1017/S095410200900203X|bibcode=2009AntSc..21..477H }}</ref> while the altitude record is held by ''[[Branchinecta brushi|B. brushi]]'', which lives at {{convert|5930|m}} in the [[Chile]]an [[Andes]].<ref name="Hegna">{{cite journal|author1=Thomas A. Hegna |author2=Eric A. Lazo-Wasem |date=1 July 2010 |title=''Branchinecta brushi'' n. sp. (Branchiopoda: Anostraca: Branchinectidae) from a volcanic crater in northern Chile (Antofagasta Province): a new altitude record for crustaceans |journal=[[Journal of Crustacean Biology]] |volume=30 |issue=3 |pages=445–464 |doi=10.1651/09-3236.1 |s2cid=85961054 |url=https://yale.academia.edu/documents/0068/6445/Hegna___Lazo-Wasem.pdf }}{{dead link|date=July 2017 |bot=InternetArchiveBot |fix-attempted=yes }}</ref> Other genera, such as ''[[Streptocephalus]]'', occur in [[desert]]s throughout the world.<ref>{{cite book |author=David Ward |year=2009 |title=The Biology of Deserts |publisher=[[Oxford University Press]] |isbn=978-0-19-921147-0 |chapter=Biodiversity and biogeography of deserts |pages=192–216 |chapter-url=https://books.google.com/books?id=3ixJ5GwT5R4C&pg=PA204}}</ref>
Anostracans inhabit inland waters ranging from [[hypersaline lake]]s to lakes that are almost devoid of dissolved substances;<ref name="Belk">{{cite book |editor1=Sol Felty Light |editor2=James T. Carlton |year=2007 |title=The Light and Smith Manual: Intertidal Invertebrates from Central California to Oregon |edition=4th |publisher=[[University of California Press]] |isbn=978-0-520-23939-5|chapter=Branchiopoda |author=Denton Belk |pages=414–417 |chapter-url=https://books.google.com/books?id=64jgZ1CfmB8C&pg=PA416}}</ref> they are "the most archetypal crustaceans" in ephemeral waters.<ref name="Dumont">{{cite book |year=2009 |editor=Henri J. Dumont |title=The Nile: Origins, Environments, Limnology and Human Use |series=Volume 89 of Monographiae Biologicae |publisher=[[Springer Science+Business Media|Springer]] |isbn=978-1-4020-9725-6 |chapter=The crustacean zooplankton (Copepoda, Branchiopoda), atyid Decapoda, and Syncarida of the Nile Basin |author=Henri J. Dumont |pages=521–546 |chapter-url=https://books.google.com/books?id=iF_U1NoknHoC&pg=PA536}}</ref> The relatively large size of fairy shrimp, together with their slow means of locomotion, makes them an easy target for [[predator]]y [[fish]] and [[water bird|waterfowl]].<ref name="Dumont"/> This has led to their distribution being restricted to environments with fewer predators, such as [[vernal pool]]s, [[salt lake]]s and lakes at high altitudes or latitudes.<ref name="Dumont"/> The southernmost recorded fairy shrimp is ''[[Branchinecta gaini]]'' from the [[Antarctic Peninsula]],<ref>{{cite journal |author=T.C. Hawes |date=October 2009 |title=Origins and dispersal of the Antarctic fairy shrimp |journal=[[Antarctic Science]] |volume=21 |issue=5 |pages=477–482 |doi=10.1017/S095410200900203X|bibcode=2009AntSc..21..477H |s2cid=84382536 }}</ref> while the altitude record is held by ''[[Branchinecta brushi|B. brushi]]'', which lives at {{convert|5930|m}} in the [[Chile]]an [[Andes]].<ref name="Hegna">{{cite journal|author1=Thomas A. Hegna |author2=Eric A. Lazo-Wasem |date=1 July 2010 |title=''Branchinecta brushi'' n. sp. (Branchiopoda: Anostraca: Branchinectidae) from a volcanic crater in northern Chile (Antofagasta Province): a new altitude record for crustaceans |journal=[[Journal of Crustacean Biology]] |volume=30 |issue=3 |pages=445–464 |doi=10.1651/09-3236.1 |s2cid=85961054 |doi-access=free }}</ref> Other genera, such as ''[[Streptocephalus]]'', occur in [[desert]]s throughout the world.<ref>{{cite book |author=David Ward |year=2009 |title=The Biology of Deserts |publisher=[[Oxford University Press]] |isbn=978-0-19-921147-0 |chapter=Biodiversity and biogeography of deserts |pages=192–216 |chapter-url=https://books.google.com/books?id=3ixJ5GwT5R4C&pg=PA204}}</ref>


Anostracans swim gracefully by movements of their phyllopodia (thoracic appendages) in a [[metachronal rhythm]].<ref name="Pennak"/> When swimming, the animal's ventral side is normally uppermost (often described as swimming "upside-down").<ref name="Belk"/> They [[filter feeder|filter food]] indiscriminately from the water as they swim, but also scrape algae and other organic materials from solid surfaces, for which they turn to have their ventral side against the food surface.<ref name="Belk"/>
Anostracans swim gracefully by movements of their phyllopodia (thoracic appendages) in a [[metachronal rhythm]].<ref name="Pennak"/> When swimming, the animal's ventral side is normally uppermost (often described as swimming "upside-down").<ref name="Belk"/> They [[filter feeder|filter food]] indiscriminately from the water as they swim, but also scrape algae and other organic materials from solid surfaces, for which they turn to have their ventral side against the food surface.<ref name="Belk"/>


Another important aspect of the fairy shrimp’s life cycle is their universal ability to enter diapause,<ref name=":0">{{Cite journal|last1=Hairston|first1=Nelson G.|last2=Cáceres|first2=Carla E.|date=1996-03-01|title=Distribution of crustacean diapause: micro- and macroevolutionary pattern and process|journal=Hydrobiologia|language=en|volume=320|issue=1–3|pages=27–44|doi=10.1007/bf00016802|s2cid=44914111|issn=0018-8158}}</ref><ref name=":1">{{Cite journal|last=Fryer|first=Geoffrey|date=1996-03-01|title=Diapause, a potent force in the evolution of freshwater crustaceans|journal=Hydrobiologia|language=en|volume=320|issue=1–3|pages=1–14|doi=10.1007/bf00016800|s2cid=24128518|issn=0018-8158}}</ref> a state of biological dormancy where growth and metabolism are arrested,<ref name=":2">{{Cite journal|last1=Alekseev|first1=Victor R.|last2=Starobogatov|first2=Yaroslav I.|date=1996-03-01|title=Types of diapause in Crustacea: definitions, distribution, evolution|journal=Hydrobiologia|language=en|volume=320|issue=1–3|pages=15–26|doi=10.1007/bf00016801|s2cid=253609|issn=0018-8158}}</ref> as an egg (or cyst). This trait assists in both species' dispersal and in overcoming adverse environmental conditions.<ref name=":1" /><ref name=":2" /> Once dormant, these cysts can withstand conditions as harsh and diverse as droughts, frosts, hypersalinity, complete desiccation, exposure to UV radiation and the vacuum of space.<ref>{{Cite journal|last1=Czyż|first1=M.|last2=Woliński|first2=P.|last3=Gołdyn|first3=B.|date=2016|title=Cyst morphology of large branchiopod crustaceans (Anostraca, Notostraca, Laevicaudata, Spinicaudata) in western Poland|url=|journal=Biological Letters|volume=53|issue=2|pages=79–88|doi=10.1515/biolet-2017-0006|doi-access=free}}</ref><ref name="Belk" /><ref name=":2" /> It is also the only way for the fairy shrimps to colonize new habitats—facilitated by a variety of conditions including wind, predators, currents<ref name=":3">{{Cite journal|last1=Parnov|first1=V.|last2=Krylov|first2=P.|last3=Riccardi|first3=N.|date=2004|title=Role of diapause in dispersal and invasion success by aquatic invertebrates|url=|journal=Journal of Limnology|volume=63|pages=59–69|via=}}</ref><ref name=":4">{{Cite journal|last=Rogers|first=C.|date=2015|title=A conceptual model for Anostracan biogeography|url=|journal=Journal of Crustacean Biology|volume=35|issue=5|pages=686–699|doi=10.1163/1937240x-00002369|doi-access=free}}</ref><ref>{{Cite journal|last1=Brendonck|first1=Luc|last2=Riddoch|first2=Bruce J.|date=1999-05-01|title=Wind-borne short-range egg dispersal in anostracans (Crustacea: Branchiopoda)|journal=Biological Journal of the Linnean Society|volume=67|issue=1|pages=87–95|doi=10.1111/j.1095-8312.1999.tb01931.x|issn=0024-4066|doi-access=free}}</ref>—as the soft-bodied adults are unable to leave the freshwater system.<ref name=":4" /> Once in diapause, these cysts can remain viable for centuries,<ref name=":3" /> and the mixing of system sediment results in the hatching of different aged cysts in each generation.<ref name=":5">{{Cite journal|last1=Remigio|first1=E. A.|last2=Hebert|first2=P. D. N.|last3=Savage|first3=A.|date=2001-09-01|title=Phylogenetic relationships and remarkable radiation in Parartemia (Crustacea: Anostraca), the endemic brine shrimp of Australia: evidence from mitochondrial DNA sequences|journal=Biological Journal of the Linnean Society|volume=74|issue=1|pages=59–71|doi=10.1006/bijl.2001.0567|issn=0024-4066|doi-access=free}}</ref><ref name=":4" /><ref>{{Cite journal|last1=Kraus|first1=Holger|last2=Eder|first2=Erich|last3=Sten Møller|first3=Ole|last4=Werding|first4=Bernd|date=2004-07-01|title=Cyst Deposition Behaviour and the Functional Morphology of the Brood Pouch in Streptocephalus Torvicornis (Branchiopoda: Anostraca)|journal=Journal of Crustacean Biology|volume=24|issue=3|pages=393–397|doi=10.1651/c-2470|issn=0278-0372|doi-access=free}}</ref> This inbreeding slows the rate of selection by resisting gene flow and minimizing phenotypic variation, in turn promoting the stability of the existing, successful phenotype.<ref name=":4" />
Another important aspect of the fairy shrimp’s life cycle is their universal ability to enter diapause,<ref name=":0">{{Cite journal|last1=Hairston|first1=Nelson G.|last2=Cáceres|first2=Carla E.|date=1996-03-01|title=Distribution of crustacean diapause: micro- and macroevolutionary pattern and process|journal=Hydrobiologia|language=en|volume=320|issue=1–3|pages=27–44|doi=10.1007/bf00016802|s2cid=44914111|issn=0018-8158}}</ref><ref name=":1">{{Cite journal|last=Fryer|first=Geoffrey|date=1996-03-01|title=Diapause, a potent force in the evolution of freshwater crustaceans|journal=Hydrobiologia|language=en|volume=320|issue=1–3|pages=1–14|doi=10.1007/bf00016800|s2cid=24128518|issn=0018-8158|doi-access=free}}</ref> a state of biological dormancy where growth and metabolism are arrested,<ref name=":2">{{Cite journal|last1=Alekseev|first1=Victor R.|last2=Starobogatov|first2=Yaroslav I.|date=1996-03-01|title=Types of diapause in Crustacea: definitions, distribution, evolution|journal=Hydrobiologia|language=en|volume=320|issue=1–3|pages=15–26|doi=10.1007/bf00016801|s2cid=253609|issn=0018-8158}}</ref> as an egg (or cyst). This trait assists in both species' dispersal and in overcoming adverse environmental conditions.<ref name=":1" /><ref name=":2" /> Once dormant, these cysts can withstand conditions as harsh and diverse as droughts, frosts, hypersalinity, complete desiccation, exposure to UV radiation and the vacuum of space.<ref>{{Cite journal|last1=Czyż|first1=M.|last2=Woliński|first2=P.|last3=Gołdyn|first3=B.|date=2016|title=Cyst morphology of large branchiopod crustaceans (Anostraca, Notostraca, Laevicaudata, Spinicaudata) in western Poland|journal=Biological Letters|volume=53|issue=2|pages=79–88|doi=10.1515/biolet-2017-0006|doi-access=free}}</ref><ref name="Belk" /><ref name=":2" /> It is also the only way for the fairy shrimps to colonize new habitats—facilitated by a variety of conditions including wind, predators, currents<ref name=":3">{{Cite journal|last1=Parnov|first1=V.|last2=Krylov|first2=P.|last3=Riccardi|first3=N.|date=2004|title=Role of diapause in dispersal and invasion success by aquatic invertebrates|journal=Journal of Limnology|volume=63|pages=59–69}}</ref><ref name=":4">{{Cite journal|last=Rogers|first=C.|date=2015|title=A conceptual model for Anostracan biogeography|journal=Journal of Crustacean Biology|volume=35|issue=5|pages=686–699|doi=10.1163/1937240x-00002369|doi-access=free}}</ref><ref>{{Cite journal|last1=Brendonck|first1=Luc|last2=Riddoch|first2=Bruce J.|date=1999-05-01|title=Wind-borne short-range egg dispersal in anostracans (Crustacea: Branchiopoda)|journal=Biological Journal of the Linnean Society|volume=67|issue=1|pages=87–95|doi=10.1111/j.1095-8312.1999.tb01931.x|issn=0024-4066|doi-access=free}}</ref>—as the soft-bodied adults are unable to leave the freshwater system.<ref name=":4" /> Once in diapause, these cysts can remain viable for centuries,<ref name=":3" /> and the mixing of system sediment results in the hatching of different aged cysts in each generation.<ref name=":5">{{Cite journal|last1=Remigio|first1=E. A.|last2=Hebert|first2=P. D. N.|last3=Savage|first3=A.|date=2001-09-01|title=Phylogenetic relationships and remarkable radiation in Parartemia (Crustacea: Anostraca), the endemic brine shrimp of Australia: evidence from mitochondrial DNA sequences|journal=Biological Journal of the Linnean Society|volume=74|issue=1|pages=59–71|doi=10.1006/bijl.2001.0567|issn=0024-4066|doi-access=free|bibcode=2001BJLS...74...59R }}</ref><ref name=":4" /><ref>{{Cite journal|last1=Kraus|first1=Holger|last2=Eder|first2=Erich|last3=Sten Møller|first3=Ole|last4=Werding|first4=Bernd|date=2004-07-01|title=Cyst Deposition Behaviour and the Functional Morphology of the Brood Pouch in Streptocephalus Torvicornis (Branchiopoda: Anostraca)|journal=Journal of Crustacean Biology|volume=24|issue=3|pages=393–397|doi=10.1651/c-2470|issn=0278-0372|doi-access=free}}</ref> This inbreeding slows the rate of selection by resisting gene flow and minimizing phenotypic variation, in turn promoting the stability of the existing, successful phenotype.<ref name=":4" />


Anostracans are an important food source for many birds and fish. For example, they provide much of the food for female [[Northern pintail|pintails]] and [[mallard]]s in the [[Prairie Pothole Region]] of the [[Great Plains]] in [[North America]], especially in years when temporary wetlands are abundant.<ref name="Batt">{{cite book |editor=Bruce D. J. Batt |year=1992 |title=Ecology and Management of Breeding Waterfowl |chapter=Foraging ecology and nutrition |author1=Gary L. Krapu |author2=Kenneth J. Reinecke |pages=1–29 |publisher=[[University of Minnesota Press]] |isbn=978-0-8166-2001-2 |chapter-url=https://books.google.com/books?id=MroyihsZQoIC&pg=PA9}}</ref> Similarly, ''[[Artemia]]'' forms an important part of the diet of [[flamingo]]s wherever it can be found.<ref>{{cite book |editor1=Hans-Jörg Barth |editor2=Benno Böer |year=2002 |title=Sabkha Ecosystems, Volume 2 |series=Tasks for Vegetation science |publisher=[[Springer Science+Business Media|Springer]] |isbn=978-1-4020-0504-6 |chapter=Saline wetland reserve management: a case study from the United Arab Emirates |author1=Simon Aspinall |author2=Peter Hellyer |pages=335–340 |chapter-url=https://books.google.com/books?id=O7fIy_Rwgc8C&pg=PA338}}</ref>
Anostracans are an important food source for many birds and fish. For example, they provide much of the food for female [[Northern pintail|pintails]] and [[mallard]]s in the [[Prairie Pothole Region]] of the [[Great Plains]] in [[North America]], especially in years when temporary wetlands are abundant.<ref name="Batt">{{cite book |editor=Bruce D. J. Batt |year=1992 |title=Ecology and Management of Breeding Waterfowl |chapter=Foraging ecology and nutrition |author1=Gary L. Krapu |author2=Kenneth J. Reinecke |pages=1–29 |publisher=[[University of Minnesota Press]] |isbn=978-0-8166-2001-2 |chapter-url=https://books.google.com/books?id=MroyihsZQoIC&pg=PA9}}</ref> Similarly, ''[[Artemia]]'' forms an important part of the diet of [[flamingo]]s wherever it can be found.<ref>{{cite book |editor1=Hans-Jörg Barth |editor2=Benno Böer |year=2002 |title=Sabkha Ecosystems, Volume 2 |series=Tasks for Vegetation science |publisher=[[Springer Science+Business Media|Springer]] |isbn=978-1-4020-0504-6 |chapter=Saline wetland reserve management: a case study from the United Arab Emirates |author1=Simon Aspinall |author2=Peter Hellyer |pages=335–340 |chapter-url=https://books.google.com/books?id=O7fIy_Rwgc8C&pg=PA338}}</ref>


==Uses==
==Uses==
[[File:Artemia breeding ponds.jpg|thumb|Salt evaporation ponds at [[Redwood City, California|Redwood City]] on [[San Francisco Bay]]: the orange colour is produced by the presence of ''[[brine shrimp|Artemia]]''.]]
[[File:Artemia breeding ponds.jpg|thumb|[[San Francisco Bay Salt Ponds]]: the orange colour is produced by the presence of ''[[brine shrimp|Artemia]]''.]]
[[Brine shrimp]] are used as food for [[fish]] and other organisms in [[aquarium|aquaria]] and [[aquaculture]].<ref name="Jørgensen">{{cite book |editor=Sven Erik Jørgensen |title=Ecosystem Ecology |year=2009 |publisher=[[Academic Press]] |isbn=978-0-444-53466-8 |chapter=Saline and soda lakes |author=J. M. Melack |pages=380–384 |chapter-url=https://books.google.com/books?id=a-LjEvwWmOEC&pg=PA384}}</ref> Their drought-resistant eggs are collected from lakeshores and are stored and transported dry. They hatch readily when submerged in salt water. This is a multimillion-dollar industry, centred on the [[Great Salt Lake]] in [[Utah]] and [[San Francisco Bay]] in [[California]];<ref>{{cite book |author=Hugh F. Clifford |year=1991 |title=Aquatic Invertebrates of Alberta: an Illustrated Guide |publisher=[[University of Alberta]] |isbn=978-0-88864-234-9 |chapter=Anostraca |pages=140–143 |chapter-url=https://books.google.com/books?id=8UQ4jHev6voC&pg=PA140}}</ref> adults are collected from [[Mono Lake]] and transported frozen.<ref name="Jørgensen"/>
[[Brine shrimp]] are used as food for [[fish]] and other organisms in [[aquarium|aquaria]] and [[aquaculture]].<ref name="Jørgensen">{{cite book |editor=Sven Erik Jørgensen |title=Ecosystem Ecology |year=2009 |publisher=[[Academic Press]] |isbn=978-0-444-53466-8 |chapter=Saline and soda lakes |author=J. M. Melack |pages=380–384 |chapter-url=https://books.google.com/books?id=a-LjEvwWmOEC&pg=PA384}}</ref> Their drought-resistant eggs are collected from lakeshores and are stored and transported dry. They hatch readily when submerged in salt water. This is a multimillion-dollar industry, centred on the [[Great Salt Lake]] in [[Utah]] and [[San Francisco Bay]] in [[California]];<ref>{{cite book |author=Hugh F. Clifford |year=1991 |title=Aquatic Invertebrates of Alberta: an Illustrated Guide |publisher=[[University of Alberta]] |isbn=978-0-88864-234-9 |chapter=Anostraca |pages=140–143 |chapter-url=https://books.google.com/books?id=8UQ4jHev6voC&pg=PA140}}</ref> adults are collected from [[Mono Lake]] and transported frozen.<ref name="Jørgensen"/>


==Fossil record and evolution==
==Fossil record and evolution==
Fairy shrimp are believed to have diverged from the main line of Branchiopoda during the Ordovician period,<ref name=":6">{{Cite journal|last=Olesen|first=Jørgen|date=2007-04-01|title=Monophyly and Phylogeny of Branchiopoda, with Focus on Morphology and Homologies of Branchiopod Phyllopodous Limbs|journal=Journal of Crustacean Biology|volume=27|issue=2|pages=165–183|doi=10.1651/s-2727.1|issn=0278-0372|doi-access=free}}</ref><ref name=":7">{{Cite journal|last1=Daniels|first1=Savel R.|last2=Hamer|first2=Michelle|last3=Rogers|first3=Christopher|date=2004-07-01|title=Molecular evidence suggests an ancient radiation for the fairy shrimp genus Streptocephalus (Branchiopoda: Anostraca)|journal=Biological Journal of the Linnean Society|volume=82|issue=3|pages=313–327|doi=10.1111/j.1095-8312.2004.00359.x|issn=0024-4066|doi-access=free}}</ref> around the same time it is thought they colonized freshwater and estuarine ecosystems.<ref name=":2" /> This transition is believed to have been the result of selection pressure to escape predation in the Early Paleozoic seas.<ref name="Diversity" /><ref name="Belk" /><ref name=":1" />
Fairy shrimp are believed to have diverged from the main line of Branchiopoda during the Ordovician period,<ref name=":6">{{Cite journal|last=Olesen|first=Jørgen|date=2007-04-01|title=Monophyly and Phylogeny of Branchiopoda, with Focus on Morphology and Homologies of Branchiopod Phyllopodous Limbs|journal=Journal of Crustacean Biology|volume=27|issue=2|pages=165–183|doi=10.1651/s-2727.1|issn=0278-0372|doi-access=free}}</ref><ref name=":7">{{Cite journal|last1=Daniels|first1=Savel R.|last2=Hamer|first2=Michelle|last3=Rogers|first3=Christopher|date=2004-07-01|title=Molecular evidence suggests an ancient radiation for the fairy shrimp genus Streptocephalus (Branchiopoda: Anostraca)|journal=Biological Journal of the Linnean Society|volume=82|issue=3|pages=313–327|doi=10.1111/j.1095-8312.2004.00359.x|issn=0024-4066|doi-access=free}}</ref> around the same time it is thought they colonized freshwater and estuarine ecosystems.<ref name=":2" /> This transition is believed to have resulted from selection pressure to escape predation in the Early Paleozoic seas.<ref name="Diversity" /><ref name="Belk" /><ref name=":1" /> ''[[Lepidocaris]]'' from the early [[Devonian]] [[Rhynie chert]] is likely a close relative of Anostraca. The oldest known modern-looking ansotracan is ''[[Haltinnaias]]'' from the late Devonian ([[Famennian]]) Strud locality of Belgium, around 365 million years old.<ref>{{Cite journal|last1=Gueriau|first1=Pierre|last2=Rabet|first2=Nicolas|last3=Clément|first3=Gaël|last4=Lagebro|first4=Linda|last5=Vannier|first5=Jean|last6=Briggs|first6=Derek E.G. |last7=Charbonnier|first7=Sylvain|last8=Olive|first8=Sébastien|last9=Béthoux|first9=Olivier|date=February 2016|title=A 365-Million-Year-Old Freshwater Community Reveals Morphological and Ecological Stasis in Branchiopod Crustaceans|journal=Current Biology|language=en|volume=26|issue=3|pages=383–390|doi=10.1016/j.cub.2015.12.039|pmid=26776738 |doi-access=free|bibcode=2016CBio...26..383G }}</ref><ref>{{Cite journal|last1=Luo|first1=Cihang|last2=Jarzembowski|first2=Edmund A.|last3=Fang|first3=Yan|last4=Wang|first4=Bo|last5=Xiao|first5=Chuantao|date=February 2020|title=First Anostraca (Crustacea: Branchiopoda) from the Middle Jurassic of Daohugou, China|url=https://linkinghub.elsevier.com/retrieve/pii/S0016787819301300|journal=Proceedings of the Geologists' Association|language=en|volume=131|issue=1|pages=67–72|doi=10.1016/j.pgeola.2019.11.004|bibcode=2020PrGA..131...67L |s2cid=214226902 }}</ref>


Some studies point to fossils resembling fairy shrimp in the Upper Cambrian,<ref name=":8">{{Cite journal|last1=Richter|first1=Stefan|last2=Olesen|first2=Jørgen|last3=Wheeler|first3=Ward C.|date=2007-08-01|title=Phylogeny of Branchiopoda (Crustacea) based on a combined analysis of morphological data and six molecular loci|journal=Cladistics|language=en|volume=23|issue=4|pages=301–336|doi=10.1111/j.1096-0031.2007.00148.x|s2cid=55997584|issn=1096-0031}}</ref><ref>{{Cite journal|last=Brendonck|first=L.|date=1996|title=Diapause, quiescence, hatching requirements: what we can learn from large freshwater branchiopods (Crustacea Branciopoda: Anostraca, Notostraca, Conchostraca)|doi=10.1007/bf00016809|journal=Hydrobiologia|volume=320|issue=1–3|pages=85–97|s2cid=26327230}}</ref> specifically the oldest known branchiopod fossil, ''Rebachiella kinnekullensis'', from Orsten marine deposits.<ref>{{cite journal|author1=Joel W. Martin|author2=Michael S. Laverack|date=December 1992|title=On the distribution of the crustacean dorsal organ|url=http://crustacea.nhm.org/people/martin/publications/pdf/59.pdf|url-status=dead|journal=[[Acta Zoologica: Morphology and Evolution|Acta Zoologica]]|volume=73|issue=5|pages=357–368|doi=10.1111/j.1463-6395.1992.tb01108.x|archiveurl=https://web.archive.org/web/20110719085139/http://crustacea.nhm.org/people/martin/publications/pdf/59.pdf|archivedate=2011-07-19}}</ref> Despite its seeming resemblance to modern fairy shrimp, this fossil is still considered by most to be an outlying member of the ancestral marine Branchiopoda rather than a true fairy shrimp.<ref name=":6" />
Some studies point to fossils resembling fairy shrimp in the Upper Cambrian,<ref name=":8">{{Cite journal|last1=Richter|first1=Stefan|last2=Olesen|first2=Jørgen|last3=Wheeler|first3=Ward C.|date=2007-08-01|title=Phylogeny of Branchiopoda (Crustacea) based on a combined analysis of morphological data and six molecular loci|journal=Cladistics|language=en|volume=23|issue=4|pages=301–336|doi=10.1111/j.1096-0031.2007.00148.x|pmid=34905837 |s2cid=55997584|issn=1096-0031|doi-access=free}}</ref><ref>{{Cite journal|last=Brendonck|first=L.|date=1996|title=Diapause, quiescence, hatching requirements: what we can learn from large freshwater branchiopods (Crustacea Branciopoda: Anostraca, Notostraca, Conchostraca)|doi=10.1007/bf00016809|journal=Hydrobiologia|volume=320|issue=1–3|pages=85–97|s2cid=26327230}}</ref> specifically the oldest known branchiopod fossil, ''[[Rehbachiella|Rehbachiella kinnekullensis]]'', from Orsten marine deposits.<ref>{{cite journal|author1=Joel W. Martin|author2=Michael S. Laverack|date=December 1992|title=On the distribution of the crustacean dorsal organ|url=http://crustacea.nhm.org/people/martin/publications/pdf/59.pdf|url-status=dead|journal=[[Acta Zoologica: Morphology and Evolution|Acta Zoologica]]|volume=73|issue=5|pages=357–368|doi=10.1111/j.1463-6395.1992.tb01108.x|archive-url=https://web.archive.org/web/20110719085139/http://crustacea.nhm.org/people/martin/publications/pdf/59.pdf|archive-date=2011-07-19}}</ref> Despite its seeming resemblance to modern fairy shrimp, this fossil is still considered by most to be an outlying member of the ancestral marine Branchiopoda rather than an actual fairy shrimp.<ref name=":6" />


The monophyly of this order is well supported,<ref name="Diversity" /><ref>{{Cite book|title=Arthropod Relationships|last1=Fortey|first1=R.|last2=Thomas|first2=H.|publisher=Springer Science & Business Media|year=2012|isbn=|location=Berlin|pages=104–105}}</ref><ref>{{Cite book|title=Perspectives in Animal Phylogeny and Evolution|url=https://archive.org/details/perspectivesanim00mine|url-access=limited|last=Minelli|first=A.|publisher=Oxford University Press|year=2009|isbn=|location=Oxford|pages=[https://archive.org/details/perspectivesanim00mine/page/n57 98]–100}}</ref><ref name=":6" /><ref name=":8" /><ref>{{Cite journal|last=Weekers|first=P|title=Phylogenetic analysis of anostracans (Branchiopoda: Anostraca) inferred from nuclear 18S ribosomal DNA (18S rDNA) sequences|journal=Molecular Phylogenetics and Evolution|volume=25|issue=3|pages=535–544|doi=10.1016/s1055-7903(02)00289-0|year=2002|pmid=12450757}}</ref><ref name=":9">{{Cite journal|last1=Regier|first1=Jerome C.|last2=Shultz|first2=Jeffrey W.|last3=Zwick|first3=Andreas|last4=Hussey|first4=April|last5=Ball|first5=Bernard|last6=Wetzer|first6=Regina|last7=Martin|first7=Joel W.|last8=Cunningham|first8=Clifford W.|date=2010-02-10|title=Arthropod relationships revealed by phylogenomic analysis of nuclear protein-coding sequences|journal=Nature|language=en|volume=463|issue=7284|pages=1079–1083|doi=10.1038/nature08742|pmid=20147900|bibcode=2010Natur.463.1079R|s2cid=4427443|issn=1476-4687}}</ref> and the scientific community has reached consensus that Anostraca was the first group to branch off from the Branchiopoda.<ref name=":0" /><ref name=":8" /><ref name=":1" /><ref name=":9" /><ref name=":7" />
The monophyly of this order is well supported,<ref name="Diversity" /><ref>{{Cite book|title=Arthropod Relationships|last1=Fortey|first1=R.|last2=Thomas|first2=H.|publisher=Springer Science & Business Media|year=2012|location=Berlin|pages=104–105}}</ref><ref>{{Cite book|title=Perspectives in Animal Phylogeny and Evolution|url=https://archive.org/details/perspectivesanim00mine|url-access=limited|last=Minelli|first=A.|publisher=Oxford University Press|year=2009|location=Oxford|pages=[https://archive.org/details/perspectivesanim00mine/page/n57 98]–100}}</ref><ref name=":6" /><ref name=":8" /><ref>{{Cite journal|last=Weekers|first=P|title=Phylogenetic analysis of anostracans (Branchiopoda: Anostraca) inferred from nuclear 18S ribosomal DNA (18S rDNA) sequences|journal=Molecular Phylogenetics and Evolution|volume=25|issue=3|pages=535–544|doi=10.1016/s1055-7903(02)00289-0|year=2002|pmid=12450757}}</ref><ref name=":9">{{Cite journal|last1=Regier|first1=Jerome C.|last2=Shultz|first2=Jeffrey W.|last3=Zwick|first3=Andreas|last4=Hussey|first4=April|last5=Ball|first5=Bernard|last6=Wetzer|first6=Regina|last7=Martin|first7=Joel W.|last8=Cunningham|first8=Clifford W.|date=2010-02-10|title=Arthropod relationships revealed by phylogenomic analysis of nuclear protein-coding sequences|journal=Nature|language=en|volume=463|issue=7284|pages=1079–1083|doi=10.1038/nature08742|pmid=20147900|bibcode=2010Natur.463.1079R|s2cid=4427443|issn=1476-4687}}</ref> and the scientific community has reached consensus that Anostraca was the first group to branch off from the Branchiopoda.<ref name=":0" /><ref name=":8" /><ref name=":1" /><ref name=":9" /><ref name=":7" />


The radiation hypothesis championing rapid spread and colonization during the Gondwana fragmentation closely echoes the current distribution of the order.<ref name=":7" /><ref name=":4" /> Presently, Anostraca are found on all seven continents.<ref name="Diversity" /> Most of the extant genera have restricted geographical distributions. Only three genera are widespread across the remnants of the former supercontinent [[Pangaea]]: ''[[Artemia]]'', ''[[Branchinella]]'' and ''[[Branchinecta]]'', while the remaining genera are found only throughout former Laurasia.<ref name="Williams">{{cite book|chapter-url=https://books.google.com/books?id=gKgOAAAAQAAJ&pg=PA25|title=The Ecology of Temporary Waters|author=D. Dudley Williams|publisher=[[Taylor & Francis]]|year=1987|isbn=978-0-7099-5211-4|pages=21–67|chapter=The Biota}}</ref> This suggests that much of the potential habitat in this supercontinent, now occupied by Anostraca, to have been unoccupied by ecologically similar species, or to have been inhabited by species with less adaptive ability.<ref name=":4" /> Studies have found Anostraca capable of rapid colonization<ref>{{Cite journal|last1=Kappas|first1=Ilias|last2=Mura|first2=Graziella|last3=Synefiaridou|first3=Dimitra|last4=Marrone|first4=Federico|last5=Alfonso|first5=Giuseppe|last6=Alonso|first6=Miguel|last7=Abatzopoulos|first7=Theodore J.|date=2017-10-01|title=Molecular and morphological data suggest weak phylogeographic structure in the fairy shrimp Streptocephalus torvicornis (Branchiopoda, Anostraca)|journal=Hydrobiologia|language=en|volume=801|issue=1|pages=21–32|doi=10.1007/s10750-017-3203-6|s2cid=25271723|issn=0018-8158}}</ref> and speciation.<ref name=":5" />
The radiation hypothesis championing rapid spread and colonization during the Gondwana fragmentation closely echoes the current distribution of the order.<ref name=":7" /><ref name=":4" /> Presently, Anostraca are found on all seven continents.<ref name="Diversity" /> Most extant genera have restricted geographical distributions. Only three genera are widespread across the remnants of the former supercontinent [[Pangaea]]: ''[[Artemia]]'', ''[[Branchinella]]'' and ''[[Branchinecta]]'', while the remaining genera are found only throughout former Laurasia.<ref name="Williams">{{cite book|chapter-url=https://books.google.com/books?id=gKgOAAAAQAAJ&pg=PA25|title=The Ecology of Temporary Waters|author=D. Dudley Williams|publisher=[[Taylor & Francis]]|year=1987|isbn=978-0-7099-5211-4|pages=21–67|chapter=The Biota}}</ref> This suggests that much of the potential habitat in this supercontinent, now occupied by Anostraca, was previously unoccupied by ecologically similar species, or inhabited by species with less adaptive ability.<ref name=":4" /> Studies have found Anostraca capable of rapid colonization<ref>{{Cite journal|last1=Kappas|first1=Ilias|last2=Mura|first2=Graziella|last3=Synefiaridou|first3=Dimitra|last4=Marrone|first4=Federico|last5=Alfonso|first5=Giuseppe|last6=Alonso|first6=Miguel|last7=Abatzopoulos|first7=Theodore J.|date=2017-10-01|title=Molecular and morphological data suggest weak phylogeographic structure in the fairy shrimp Streptocephalus torvicornis (Branchiopoda, Anostraca)|journal=Hydrobiologia|language=en|volume=801|issue=1|pages=21–32|doi=10.1007/s10750-017-3203-6|s2cid=25271723|issn=0018-8158|url=https://research.rug.nl/en/publications/11d3e994-3461-4015-9a3a-6d1f80d5c321 }}</ref> and speciation.<ref name=":5" />


==Diversity==
==Diversity==
[[File:Branchinecta paludosa (plate).jpg|thumb|upright|Anatomical drawings of ''[[Branchinecta paludosa]]'' ([[Branchinectidae]]) from [[Georg Ossian Sars]]' ''Fauna Norvegiae'' (1896)]]
[[File:Branchinecta paludosa (plate).jpg|thumb|upright|Anatomical drawings of ''[[Branchinecta paludosa]]'' ([[Branchinectidae]]) from [[Georg Ossian Sars]]' ''Fauna Norvegiae'' (1896)]]
Anostraca is the most diverse of the four orders of Branchiopoda. It comprises around 300 species, grouped into 26 genera in eight families:<ref name="Diversity">{{cite book |author1=Luc Brendonck |author2=D. Christopher Rogers |author3=Jorgen Olesen |author4=Stephen Weeks |author5=Walter R. Hoch |year=2008 |chapter=Global diversity of large branchiopods (Crustacea: Branchiopoda) in freshwater |id=Reprinted from [[Hydrobiologia]], Volume 595 |pages=167–176 |doi=10.1007/s10750-007-9119-9 |title=Freshwater Animal Diversity Assessment |editor1=Estelle V. Balian |editor2=Christian Lévêque |editor3=Hendrik Segers |editor4=Koen Martens |series=Developments in Hydrobiology 198 |isbn=978-1-4020-8258-0 |s2cid=46608816 |chapter-url=https://books.google.com/books?id=Dw4H6DBHnAgC&pg=PA167}}</ref>
Anostraca is the most diverse of the four orders of Branchiopoda. It comprises around 313 species, grouped into 26 genera in eight families:<ref name="Diversity">{{cite book |author1=Luc Brendonck |author2=D. Christopher Rogers |author3=Jorgen Olesen |author4=Stephen Weeks |author5=Walter R. Hoch |year=2008 |chapter=Global diversity of large branchiopods (Crustacea: Branchiopoda) in freshwater |id=Reprinted from [[Hydrobiologia]], Volume 595 |pages=167–176 |doi=10.1007/s10750-007-9119-9 |title=Freshwater Animal Diversity Assessment |editor1=Estelle V. Balian |editor2=Christian Lévêque |editor3=Hendrik Segers |editor4=Koen Martens |series=Developments in Hydrobiology 198 |isbn=978-1-4020-8258-0 |s2cid=46608816 |chapter-url=https://books.google.com/books?id=Dw4H6DBHnAgC&pg=PA167}}</ref>
*[[Brine shrimp|Artemiidae]] – 1 genus, 9 species
* [[Brine shrimp|Artemiidae]] – 1 genus, 8 species
*[[Branchinecta|Branchinectidae]] – 1 genus, 45 species
* [[Branchinecta|Branchinectidae]] – 1 genus, 45 species
*[[Branchipodidae]] – 5 genera, 35 species
* [[Branchipodidae]] – 5 genera, 35 species
*[[Chirocephalidae]] – 9 genera, 81 species
* [[Chirocephalidae]] – 9 genera, 81 species
*[[Parartemia|Parartemiidae]] – 1 genus, 13 species
* [[Parartemia|Parartemiidae]] – 1 genus, 18 species
*[[Streptocephalus|Streptocephalidae]] – 1 genus, 56 species
* [[Streptocephalus|Streptocephalidae]] – 1 genus, 56 species
*[[Tanymastigidae]] – 2 genera, 8 species
* [[Tanymastigidae]] – 2 genera, 8 species
*[[Thamnocephalidae]] – 6 genera, 62 species
* [[Thamnocephalidae]] – 6 genera, 62 species


{{clear}}
{{clear}}

==Fairy Shrimp in Popular Culture==
Fairy Shrimp have been a student favorite to be the mascot of [[University of California, Merced|UC Merced]]. There had been several efforts to make this animal the official mascot of the campus, but in 2001 the bobcat was chosen instead.<ref>{{Cite web |last=Writer |first=Steve Rubenstein, Chronicle Staff |date=2000-05-25 |title=Bobcat Is Top Dog at UC Merced / Girl wins free tuition at future campus for suggesting mascot name |url=https://www.sfgate.com/education/article/Bobcat-Is-Top-Dog-at-UC-Merced-Girl-wins-free-3304310.php |access-date=2024-11-06 |website=SFGATE |language=en}}</ref><ref>{{Cite web |title=UC Merced Unveils Big Rufus, a Monument to Resilience, Diversity and Hope {{!}} Newsroom |url=https://news.ucmerced.edu/news/2024/uc-merced-unveils-big-rufus-monument-resilience-diversity-and-hope |access-date=2024-11-06 |website=news.ucmerced.edu |language=en}}</ref> Fairy shrimp had also been the focus of a challenge to the location of where the campus would be built because of their nearby vernal pool habitat.<ref>{{Cite web |last=Derby |first=John |date=2023-07-25 |title=‘University Community Development’ dream is alive and well |url=https://mercedcountytimes.com/university-community-development-dream-is-alive-and-well/ |access-date=2024-11-06 |website=Merced County Times |language=en-US}}</ref><ref>{{Cite web |last=Schevitz |first=Tanya |date=2007-02-20 |title=MERCED / UC expansion plans again run up against protected fairy shrimp |url=https://www.sfgate.com/green/article/MERCED-UC-expansion-plans-again-run-up-against-2647615.php |access-date=2024-11-06 |website=SFGATE |language=en}}</ref>


==References==
==References==
{{Reflist|30em}}
{{Reflist}}


==External links==
==External links==
{{Portal|Crustaceans|Arthropods}}
{{Portal|Crustaceans|Arthropods}}
*{{Wikispecies-inline|Anostraca}}
* {{Wikispecies-inline|Anostraca}}
*{{commons category-inline|Anostraca}}
* {{Commons category-inline|Anostraca}}


{{Branchiopoda}}
{{Branchiopoda}}
{{Taxonbar|from=Q853383}}
{{Taxonbar|from=Q853383}}
{{Authority control}}


[[Category:Anostraca| ]]
[[Category:Anostraca| ]]

Latest revision as of 20:06, 3 December 2024

Anostraca
Temporal range: Devonian–Recent
Artemia salina
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Branchiopoda
Subclass: Sarsostraca
Tasch, 1969
Order: Anostraca
G. O. Sars, 1867
Families [1][2]

Anostraca is one of the four orders of crustaceans in the class Branchiopoda; its members are referred to as fairy shrimp. They live in vernal pools[3] and hypersaline lakes across the world, and they have even been found in deserts, ice-covered mountain lakes, and Antarctic ice.[4] They are usually 6–25 mm (0.24–0.98 in) long (exceptionally up to 170 mm or 6.7 in). Most species have 20 body segments, bearing 11 pairs of leaf-like phyllopodia (swimming legs), and the body lacks a carapace. They swim "upside-down" and feed by filtering organic particles from the water or by scraping algae from surfaces,[4] with the exception of Branchinecta gigas, or "giant fairy shrimp", which is itself a predator of other species of anostracans. They are an important food for many birds and fish, and some are cultured and harvested for use as fish food. There are 300 species spread across 8 families.[4]

Description

[edit]

The body of a fairy shrimp is elongated and divided into segments.[5] The whole animal is typically 6–25 millimetres (0.24–0.98 in) long, but one species, Branchinecta gigas does not reach sexual maturity until it reaches 50 mm (2.0 in) long, and can grow to 170 mm (6.7 in) long.[5] The exoskeleton is thin and flexible,[5] and lacks any sign of a carapace.[6] The body can be divided into three distinct parts (tagmata) – head, thorax and abdomen.[6]

[edit]
Drawing of the head of Chirocephalus diaphanus (Chirocephalidae), showing the first antenna (A1), second antenna (A2) and frontal appendage (Ap)

The head is morphologically distinct from the thorax. It bears two compound eyes on prominent stalks, and two pairs of antennae.[7] The first pair of antennae are small, usually unsegmented, and uniramous. The second pair are long and cylindrical in females, but in males they are enlarged and specialised for holding the female during mating.[7] In some groups, males have an additional frontal appendage.[7]

Thorax and abdomen

[edit]
Male (top) and female (bottom) Eubranchipus grubii (Chirocephalidae): the female is holding eggs on her genital appendages.

The thorax of most anostracans has 13 segments (19 in Polyartemiella and 21 in Polyartemia).[8] All but the last two are very similar, with a pair of biramous phyllopods (flattened, leaf-like appendages).[6] The last two segments are fused together,[5] and their appendages are specialised for reproduction.[8] Most anostracans have separate sexes (gonochorism), but a few reproduce by parthenogenesis.[9] The abdomen comprises 6 segments without appendages, and a telson,[8] which bears two flattened caudal rami or "cercopods".[5]

Internal anatomy

[edit]

The head contains two digestive glands and the small lobate stomach into which they empty. This is connected to a long intestine, which terminates in a short rectum, with the anus located on the telson.[7] The haemocoel of anostracans is pumped by a long, tubular heart, which runs through most of the animal's length.[7] A series of slits allow haemocoel into the heart, which is then pumped out of the anterior opening by peristalsis.[7] The nervous system consists of two nerve cords which run the length of the body, with two ganglia and two transverse commissures in most of the body segments.[7]

Gas exchange is thought to take place through the entire body surface, but especially that of the phyllopodia and their associated gills, which may also be responsible for osmotic regulation.[7] Two coiled glands at the bases of the maxillae are used to excrete nitrogenous waste, typically in the form of urea.[7] Most of the animal's nitrogenous waste is, however, in the form of ammonia, which probably diffuses into the environment through the phyllopodia and gills.[7]

Ecology and behaviour

[edit]

Anostracans inhabit inland waters ranging from hypersaline lakes to lakes that are almost devoid of dissolved substances;[5] they are "the most archetypal crustaceans" in ephemeral waters.[10] The relatively large size of fairy shrimp, together with their slow means of locomotion, makes them an easy target for predatory fish and waterfowl.[10] This has led to their distribution being restricted to environments with fewer predators, such as vernal pools, salt lakes and lakes at high altitudes or latitudes.[10] The southernmost recorded fairy shrimp is Branchinecta gaini from the Antarctic Peninsula,[11] while the altitude record is held by B. brushi, which lives at 5,930 metres (19,460 ft) in the Chilean Andes.[12] Other genera, such as Streptocephalus, occur in deserts throughout the world.[13]

Anostracans swim gracefully by movements of their phyllopodia (thoracic appendages) in a metachronal rhythm.[7] When swimming, the animal's ventral side is normally uppermost (often described as swimming "upside-down").[5] They filter food indiscriminately from the water as they swim, but also scrape algae and other organic materials from solid surfaces, for which they turn to have their ventral side against the food surface.[5]

Another important aspect of the fairy shrimp’s life cycle is their universal ability to enter diapause,[14][15] a state of biological dormancy where growth and metabolism are arrested,[16] as an egg (or cyst). This trait assists in both species' dispersal and in overcoming adverse environmental conditions.[15][16] Once dormant, these cysts can withstand conditions as harsh and diverse as droughts, frosts, hypersalinity, complete desiccation, exposure to UV radiation and the vacuum of space.[17][5][16] It is also the only way for the fairy shrimps to colonize new habitats—facilitated by a variety of conditions including wind, predators, currents[18][19][20]—as the soft-bodied adults are unable to leave the freshwater system.[19] Once in diapause, these cysts can remain viable for centuries,[18] and the mixing of system sediment results in the hatching of different aged cysts in each generation.[21][19][22] This inbreeding slows the rate of selection by resisting gene flow and minimizing phenotypic variation, in turn promoting the stability of the existing, successful phenotype.[19]

Anostracans are an important food source for many birds and fish. For example, they provide much of the food for female pintails and mallards in the Prairie Pothole Region of the Great Plains in North America, especially in years when temporary wetlands are abundant.[23] Similarly, Artemia forms an important part of the diet of flamingos wherever it can be found.[24]

Uses

[edit]
San Francisco Bay Salt Ponds: the orange colour is produced by the presence of Artemia.

Brine shrimp are used as food for fish and other organisms in aquaria and aquaculture.[25] Their drought-resistant eggs are collected from lakeshores and are stored and transported dry. They hatch readily when submerged in salt water. This is a multimillion-dollar industry, centred on the Great Salt Lake in Utah and San Francisco Bay in California;[26] adults are collected from Mono Lake and transported frozen.[25]

Fossil record and evolution

[edit]

Fairy shrimp are believed to have diverged from the main line of Branchiopoda during the Ordovician period,[27][28] around the same time it is thought they colonized freshwater and estuarine ecosystems.[16] This transition is believed to have resulted from selection pressure to escape predation in the Early Paleozoic seas.[29][5][15] Lepidocaris from the early Devonian Rhynie chert is likely a close relative of Anostraca. The oldest known modern-looking ansotracan is Haltinnaias from the late Devonian (Famennian) Strud locality of Belgium, around 365 million years old.[30][31]

Some studies point to fossils resembling fairy shrimp in the Upper Cambrian,[32][33] specifically the oldest known branchiopod fossil, Rehbachiella kinnekullensis, from Orsten marine deposits.[34] Despite its seeming resemblance to modern fairy shrimp, this fossil is still considered by most to be an outlying member of the ancestral marine Branchiopoda rather than an actual fairy shrimp.[27]

The monophyly of this order is well supported,[29][35][36][27][32][37][38] and the scientific community has reached consensus that Anostraca was the first group to branch off from the Branchiopoda.[14][32][15][38][28]

The radiation hypothesis championing rapid spread and colonization during the Gondwana fragmentation closely echoes the current distribution of the order.[28][19] Presently, Anostraca are found on all seven continents.[29] Most extant genera have restricted geographical distributions. Only three genera are widespread across the remnants of the former supercontinent Pangaea: Artemia, Branchinella and Branchinecta, while the remaining genera are found only throughout former Laurasia.[39] This suggests that much of the potential habitat in this supercontinent, now occupied by Anostraca, was previously unoccupied by ecologically similar species, or inhabited by species with less adaptive ability.[19] Studies have found Anostraca capable of rapid colonization[40] and speciation.[21]

Diversity

[edit]
Anatomical drawings of Branchinecta paludosa (Branchinectidae) from Georg Ossian Sars' Fauna Norvegiae (1896)

Anostraca is the most diverse of the four orders of Branchiopoda. It comprises around 313 species, grouped into 26 genera in eight families:[29]

[edit]

Fairy Shrimp have been a student favorite to be the mascot of UC Merced. There had been several efforts to make this animal the official mascot of the campus, but in 2001 the bobcat was chosen instead.[41][42] Fairy shrimp had also been the focus of a challenge to the location of where the campus would be built because of their nearby vernal pool habitat.[43][44]

References

[edit]
  1. ^ Joel W. Martin; George E. Davis (2001). An Updated Classification of the Recent Crustacea (PDF). Natural History Museum of Los Angeles County. pp. 1–132. Archived from the original (PDF) on 2013-05-12. Retrieved 2011-03-03.
  2. ^ Peter H. H. Weekers; Gopal Murugan; Jacques R. Vanfleteren; Denton Belk; Henri J. Dumont (2002). "Phylogenetic analysis of anostracans (Branchiopoda: Anostraca) inferred from nuclear 18S ribosomal DNA (18S rDNA) sequences" (PDF). Molecular Phylogenetics and Evolution. 25 (3): 535–544. doi:10.1016/S1055-7903(02)00289-0. PMID 12450757.
  3. ^ "Vernal Pool Fairy Shrimp (Branchinecta lynchi) | U.S. Fish & Wildlife Service". www.fws.gov. Retrieved 2024-11-06.
  4. ^ a b c "Cape Ann Vernal Pond Team - Fairy Shrimp". www.capeannvernalpondteam.org. Retrieved 2024-11-06.
  5. ^ a b c d e f g h i j Denton Belk (2007). "Branchiopoda". In Sol Felty Light; James T. Carlton (eds.). The Light and Smith Manual: Intertidal Invertebrates from Central California to Oregon (4th ed.). University of California Press. pp. 414–417. ISBN 978-0-520-23939-5.
  6. ^ a b c William David Williams (1980). "Arachnids and Crustaceans". Australian Freshwater Life: the Invertebrates of Australian Inland Waters (2nd ed.). Palgrave Macmillan Australia. pp. 118–184. ISBN 978-0-333-29894-7.
  7. ^ a b c d e f g h i j k Douglas Grant Smith (2001). "Phyllopodous Branchiopoda (fairy, tadpole, and clam shrimps)". In Douglas Grant Smith (ed.). Pennak's Freshwater Invertebrates of the United States: Porifera to Crustacea (4th ed.). John Wiley and Sons. pp. 427–452. ISBN 978-0-471-35837-4.
  8. ^ a b c D. R. Khanna (2004). "Segmentation in arthropods". Biology of Arthropoda. Discovery Publishing House. pp. 316–394. ISBN 978-81-7141-897-8.
  9. ^ Graham Bell (1982). "Arthropoda: Crustacea Branchiopoda". The Masterpiece of Nature: the Evolution and Genetics of Sexuality. Cambridge University Press. pp. 239–248. ISBN 978-0-85664-753-6.
  10. ^ a b c Henri J. Dumont (2009). "The crustacean zooplankton (Copepoda, Branchiopoda), atyid Decapoda, and Syncarida of the Nile Basin". In Henri J. Dumont (ed.). The Nile: Origins, Environments, Limnology and Human Use. Volume 89 of Monographiae Biologicae. Springer. pp. 521–546. ISBN 978-1-4020-9725-6.
  11. ^ T.C. Hawes (October 2009). "Origins and dispersal of the Antarctic fairy shrimp". Antarctic Science. 21 (5): 477–482. Bibcode:2009AntSc..21..477H. doi:10.1017/S095410200900203X. S2CID 84382536.
  12. ^ Thomas A. Hegna; Eric A. Lazo-Wasem (1 July 2010). "Branchinecta brushi n. sp. (Branchiopoda: Anostraca: Branchinectidae) from a volcanic crater in northern Chile (Antofagasta Province): a new altitude record for crustaceans". Journal of Crustacean Biology. 30 (3): 445–464. doi:10.1651/09-3236.1. S2CID 85961054.
  13. ^ David Ward (2009). "Biodiversity and biogeography of deserts". The Biology of Deserts. Oxford University Press. pp. 192–216. ISBN 978-0-19-921147-0.
  14. ^ a b Hairston, Nelson G.; Cáceres, Carla E. (1996-03-01). "Distribution of crustacean diapause: micro- and macroevolutionary pattern and process". Hydrobiologia. 320 (1–3): 27–44. doi:10.1007/bf00016802. ISSN 0018-8158. S2CID 44914111.
  15. ^ a b c d Fryer, Geoffrey (1996-03-01). "Diapause, a potent force in the evolution of freshwater crustaceans". Hydrobiologia. 320 (1–3): 1–14. doi:10.1007/bf00016800. ISSN 0018-8158. S2CID 24128518.
  16. ^ a b c d Alekseev, Victor R.; Starobogatov, Yaroslav I. (1996-03-01). "Types of diapause in Crustacea: definitions, distribution, evolution". Hydrobiologia. 320 (1–3): 15–26. doi:10.1007/bf00016801. ISSN 0018-8158. S2CID 253609.
  17. ^ Czyż, M.; Woliński, P.; Gołdyn, B. (2016). "Cyst morphology of large branchiopod crustaceans (Anostraca, Notostraca, Laevicaudata, Spinicaudata) in western Poland". Biological Letters. 53 (2): 79–88. doi:10.1515/biolet-2017-0006.
  18. ^ a b Parnov, V.; Krylov, P.; Riccardi, N. (2004). "Role of diapause in dispersal and invasion success by aquatic invertebrates". Journal of Limnology. 63: 59–69.
  19. ^ a b c d e f Rogers, C. (2015). "A conceptual model for Anostracan biogeography". Journal of Crustacean Biology. 35 (5): 686–699. doi:10.1163/1937240x-00002369.
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