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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}}</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}}</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>
Line 53: Line 53:
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 been the result of selection pressure to escape predation in the Early Paleozoic seas.<ref name="Diversity" /><ref name="Belk" /><ref name=":1" />


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|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|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 a true 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 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" />

Revision as of 15:29, 5 December 2020

Anostraca
Temporal range: Ordovician–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 also known as fairy shrimp. They live in vernal pools and hypersaline lakes across the world, and they have even been found in deserts, ice-covered mountain lakes and Antarctic ice. 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. 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.

Description

The body of a fairy shrimp is elongated and divided into segments.[3] 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.[3] The exoskeleton is thin and flexible,[3] and lacks any sign of a carapace.[4] The body can be divided into three distinct parts (tagmata) – head, thorax and abdomen.[4]

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.[5] 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.[5] In some groups, males have an additional frontal appendage.[5]

Thorax and abdomen

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).[6] All but the last two are very similar, with a pair of biramous phyllopods (flattened, leaf-like appendages).[4] The last two segments are fused together,[3] and their appendages are specialised for reproduction.[6] Most anostracans have separate sexes (gonochorism), but a few reproduce by parthenogenesis.[7] The abdomen comprises 6 segments without appendages, and a telson,[6] which bears two flattened caudal rami or "cercopods".[3]

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.[5] The haemocoel of anostracans is pumped by a long, tubular heart, which runs through most of the animal's length.[5] A series of slits allow haemocoel into the heart, which is then pumped out of the anterior opening by peristalsis.[5] 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.[5]

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.[5] Two coiled glands at the bases of the maxillae are used to excrete nitrogenous waste, typically in the form of urea.[5] 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.[5]

Ecology and behaviour

Anostracans inhabit inland waters ranging from hypersaline lakes to lakes that are almost devoid of dissolved substances;[3] they are "the most archetypal crustaceans" in ephemeral waters.[8] The relatively large size of fairy shrimp, together with their slow means of locomotion, makes them an easy target for predatory fish and waterfowl.[8] 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.[8] The southernmost recorded fairy shrimp is Branchinecta gaini from the Antarctic Peninsula,[9] while the altitude record is held by B. brushi, which lives at 5,930 metres (19,460 ft) in the Chilean Andes.[10] Other genera, such as Streptocephalus, occur in deserts throughout the world.[11]

Anostracans swim gracefully by movements of their phyllopodia (thoracic appendages) in a metachronal rhythm.[5] When swimming, the animal's ventral side is normally uppermost (often described as swimming "upside-down").[3] 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.[3]

Another important aspect of the fairy shrimp’s life cycle is their universal ability to enter diapause,[12][13] a state of biological dormancy where growth and metabolism are arrested,[14] as an egg (or cyst). This trait assists in both species' dispersal and in overcoming adverse environmental conditions.[13][14] 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.[15][3][14] It is also the only way for the fairy shrimps to colonize new habitats—facilitated by a variety of conditions including wind, predators, currents[16][17][18]—as the soft-bodied adults are unable to leave the freshwater system.[17] Once in diapause, these cysts can remain viable for centuries,[16] and the mixing of system sediment results in the hatching of different aged cysts in each generation.[19][17][20] 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.[17]

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.[21] Similarly, Artemia forms an important part of the diet of flamingos wherever it can be found.[22]

Uses

Salt evaporation ponds at Redwood City on San Francisco Bay: 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.[23] 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;[24] adults are collected from Mono Lake and transported frozen.[23]

Fossil record and evolution

Fairy shrimp are believed to have diverged from the main line of Branchiopoda during the Ordovician period,[25][26] around the same time it is thought they colonized freshwater and estuarine ecosystems.[14] This transition is believed to have been the result of selection pressure to escape predation in the Early Paleozoic seas.[27][3][13]

Some studies point to fossils resembling fairy shrimp in the Upper Cambrian,[28][29] specifically the oldest known branchiopod fossil, Rebachiella kinnekullensis, from Orsten marine deposits.[30] 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.[25]

The monophyly of this order is well supported,[27][31][32][25][28][33][34] and the scientific community has reached consensus that Anostraca was the first group to branch off from the Branchiopoda.[12][28][13][34][26]

The radiation hypothesis championing rapid spread and colonization during the Gondwana fragmentation closely echoes the current distribution of the order.[26][17] Presently, Anostraca are found on all seven continents.[27] 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.[35] 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.[17] Studies have found Anostraca capable of rapid colonization[36] and speciation.[19]

Diversity

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:[27]

References

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  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.
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  25. ^ a b c Olesen, Jørgen (2007-04-01). "Monophyly and Phylogeny of Branchiopoda, with Focus on Morphology and Homologies of Branchiopod Phyllopodous Limbs". Journal of Crustacean Biology. 27 (2): 165–183. doi:10.1651/s-2727.1. ISSN 0278-0372.
  26. ^ a b c Daniels, Savel R.; Hamer, Michelle; Rogers, Christopher (2004-07-01). "Molecular evidence suggests an ancient radiation for the fairy shrimp genus Streptocephalus (Branchiopoda: Anostraca)". Biological Journal of the Linnean Society. 82 (3): 313–327. doi:10.1111/j.1095-8312.2004.00359.x. ISSN 0024-4066.
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  35. ^ D. Dudley Williams (1987). "The Biota". The Ecology of Temporary Waters. Taylor & Francis. pp. 21–67. ISBN 978-0-7099-5211-4.
  36. ^ Kappas, Ilias; Mura, Graziella; Synefiaridou, Dimitra; Marrone, Federico; Alfonso, Giuseppe; Alonso, Miguel; Abatzopoulos, Theodore J. (2017-10-01). "Molecular and morphological data suggest weak phylogeographic structure in the fairy shrimp Streptocephalus torvicornis (Branchiopoda, Anostraca)". Hydrobiologia. 801 (1): 21–32. doi:10.1007/s10750-017-3203-6. ISSN 0018-8158. S2CID 25271723.
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