Remipedia: Difference between revisions

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'''Remipedia''' is a [[class (biology)|class]] of blind [[crustacean]]s, closely related to [[Hexapoda|hexapods]], found in coastal [[aquifer]]s which contain saline groundwater, with populations identified in almost every ocean basin so far explored, including in [[Australia]], the [[Caribbean Sea]], and the [[Atlantic Ocean]]. The first described remipede was the fossil ''[[Tesnusocaris goldichi]]'' ([[Early Pennsylvanian|Lower Pennsylvanian]]). Since 1979, at least seventeen living species have been identified in [[subtropical]] regions around the world.<ref name="KSHI">{{cite journal |author1=Stefan Koenemann |author2=Frederick R. Schram |author3=Mario Hönemann |author4=Thomas M. Iliffe  |name-list-style=amp |year=2007 |title=Phylogenetic analysis of Remipedia (Crustacea) |journal=[[Organisms Diversity & Evolution]] |volume=7 |issue=1 |pages=33–51 |doi=10.1016/j.ode.2006.07.001|bibcode=2007ODivE...7...33K }}</ref>
'''Remipedia''' is a [[class (biology)|class]] of blind [[crustacean]]-like animals, closely related to [[Hexapoda|hexapods]]. They are found in coastal [[aquifer]]s which contain saline groundwater, with populations identified in almost every ocean basin so far explored, including in [[Australia]], the [[Caribbean Sea]], and the [[Atlantic Ocean]]. The first described remipede was the fossil ''[[Tesnusocaris goldichi]]'' ([[Early Pennsylvanian|Lower Pennsylvanian]]). Since 1979, at least seventeen living species have been identified in [[subtropical]] regions around the world.<ref name="KSHI">{{cite journal |author1=Stefan Koenemann |author2=Frederick R. Schram |author3=Mario Hönemann |author4=Thomas M. Iliffe  |name-list-style=amp |year=2007 |title=Phylogenetic analysis of Remipedia (Crustacea) |journal=[[Organisms Diversity & Evolution]] |volume=7 |issue=1 |pages=33–51 |doi=10.1016/j.ode.2006.07.001|bibcode=2007ODivE...7...33K }}</ref>


==Description==
==Description==
Remipedes are {{convert|1|-|4|cm|in|1}} long and comprise a head and an elongate trunk of up to thirty-two similar [[body segment]]s.<ref>{{cite web |url=http://cameronmccormick.blogspot.com/2008/11/remipedia.html |title=Remipedia  |work=The Lord Geekington |author=Cameron McCormick |date=November 10, 2008}}</ref> Pigmentation and eyes are absent.<ref name=Yager2013>{{cite web| author=Yager, J. | title=Lasionectes entrichoma Yager & Schram, 1986 | url=http://www.tamug.edu/cavebiology/fauna/remipedes/L_entrichoma.html | date=18 September 2013 | publisher=tamug.edu | access-date=9 February 2018 }}</ref> Biramous swimming [[appendage]]s are laterally present on each segment. The animals swim on their backs and are generally slow-moving.<ref name=":0">{{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=February 2010|title=Arthropod relationships revealed by phylogenomic analysis of nuclear protein-coding sequences|journal=Nature|volume=463|issue=7284|pages=1079–1083|doi=10.1038/nature08742|pmid=20147900|issn=0028-0836|bibcode=2010Natur.463.1079R|s2cid=4427443}}</ref> They are the only known venomous crustaceans, and have fangs connected to secretory glands, which inject a combination of [[digestive enzyme]]s and [[venom]] into their prey,<ref>{{cite journal |title=First venomous crustacean discovered |journal=[[Nature News]] |last=Kaplan |first=Matt |date=22 October 2013 |access-date=10 May 2015 |url=http://www.nature.com/news/first-venomous-crustacean-discovered-1.13985 |doi=10.1038/nature.2013.13985|s2cid=87091184 |url-access=subscription }}</ref> but they also feed through [[Filter feeder|filter feeding]]. Being [[hermaphrodite]]s, the female pore is located on the seventh trunk segment and the male pore on the fourteenth.<ref>{{Cite journal |last1=Hinderstein |first1=Lara M. |last2=Iliffe |first2=Thomas M. |last3=Schram |first3=Frederick R. |last4=Bloechl |first4=Armin |last5=Koenemann |first5=Stefan |date=2007 |title=Behavior of Remipedia in the Laboratory, with Supporting Field Observations |url=https://academic.oup.com/jcb/article-lookup/doi/10.1651/S-2809A.1 |journal=Journal of Crustacean Biology |language=en |volume=27 |issue=4 |pages=534–542 |doi=10.1651/S-2809A.1|bibcode=2007JCBio..27..534H |url-access=subscription }}</ref>
Remipedes are {{convert|1|-|4|cm|in|1}} long and comprise a head and an elongate trunk of up to thirty-two similar [[body segment]]s.<ref>{{cite web |url=https://cameronmccormick.blogspot.com/2008/11/remipedia.html |title=Remipedia  |work=The Lord Geekington |author=Cameron McCormick |date=November 10, 2008}}</ref> Pigmentation and eyes are absent.<ref name=Yager2013>{{cite web | author=Yager, J. | title=Lasionectes entrichoma Yager & Schram, 1986 | url=http://www.tamug.edu/cavebiology/fauna/remipedes/L_entrichoma.html | date=18 September 2013 | publisher=tamug.edu | access-date=9 February 2018 | archive-date=24 November 2019 | archive-url=https://web.archive.org/web/20191124023131/http://www.tamug.edu/cavebiology/fauna/remipedes/L_entrichoma.html }}</ref> Biramous swimming [[appendage]]s are laterally present on each segment. The animals swim on their backs and are generally slow-moving.<ref name=":0">{{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=February 2010|title=Arthropod relationships revealed by phylogenomic analysis of nuclear protein-coding sequences|journal=Nature|volume=463|issue=7284|pages=1079–1083|doi=10.1038/nature08742|pmid=20147900|issn=0028-0836|bibcode=2010Natur.463.1079R|s2cid=4427443}}</ref> They are the only known venomous crustaceans, and have fangs connected to secretory glands, which inject a combination of [[digestive enzyme]]s and [[venom]] into their prey,<ref>{{cite journal |title=First venomous crustacean discovered |journal=[[Nature News]] |last=Kaplan |first=Matt |date=22 October 2013 |access-date=10 May 2015 |url=http://www.nature.com/news/first-venomous-crustacean-discovered-1.13985 |doi=10.1038/nature.2013.13985|s2cid=87091184 |url-access=subscription }}</ref> but they also feed through [[Filter feeder|filter feeding]]. Being [[hermaphrodite]]s, the female pore is located on the seventh trunk segment and the male pore on the fourteenth.<ref>{{Cite journal |last1=Hinderstein |first1=Lara M. |last2=Iliffe |first2=Thomas M. |last3=Schram |first3=Frederick R. |last4=Bloechl |first4=Armin |last5=Koenemann |first5=Stefan |date=2007 |title=Behavior of Remipedia in the Laboratory, with Supporting Field Observations |url=https://academic.oup.com/jcb/article-lookup/doi/10.1651/S-2809A.1 |journal=Journal of Crustacean Biology |language=en |volume=27 |issue=4 |pages=534–542 |doi=10.1651/S-2809A.1|bibcode=2007JCBio..27..534H |url-access=subscription }}</ref>


Remipedia have a generally primitive body plan compared to other extant crustaceans, and are the only extant [[pancrustacea]]ns to lack significant postcephalic [[Tagma (biology)|tagmosis]].<ref name=":0" /> External respiratory structures like gills are absent.<ref>{{cite book | url=https://books.google.com/books?id=0aLWAQAAQBAJ&dq=remipedia+gills&pg=PA149 | title=Treatise on Zoology - Anatomy, Taxonomy, Biology. The Crustacea, Volume 4 part A | isbn=978-90-474-4045-1 | last1=Klein | first1=Carel von Vaupel | last2=Charmantier-Daures | first2=Mireille | date=24 October 2013 | publisher=BRILL }}</ref> Previously regarded as 'primitive', Remipedia have since been shown to have enhanced olfactory nerve centers (a common feature for species that live in dark environments).<ref>{{cite journal |author1=Martin Fanenbruck |author2=Steffen Harzsch  |author3=Johann Wolfgang Wägele  |name-list-style=amp |year=2004 |title=The brain of the Remipedia (Crustacea) and an alternative hypothesis on their phylogenetic relationships |journal=[[Proceedings of the National Academy of Sciences]] |doi=10.1073/pnas.0306212101 |volume=101 |issue=11 |pages=3868–3873 |pmid=15004272 |pmc=374336|doi-access=free }}</ref>
Remipedia have a generally primitive body plan compared to other extant crustaceans, and are the only extant [[pancrustacea]]ns to lack significant postcephalic [[Tagma (biology)|tagmosis]].<ref name=":0" /> External respiratory structures like gills are absent.<ref>{{cite book | url=https://books.google.com/books?id=0aLWAQAAQBAJ&dq=remipedia+gills&pg=PA149 | title=Treatise on Zoology - Anatomy, Taxonomy, Biology. The Crustacea, Volume 4 part A | isbn=978-90-474-4045-1 | last1=Klein | first1=Carel von Vaupel | last2=Charmantier-Daures | first2=Mireille | date=24 October 2013 | publisher=BRILL }}</ref> Previously regarded as 'primitive', Remipedia have since been shown to have enhanced olfactory nerve centers (a common feature for species that live in dark environments).<ref>{{cite journal |author1=Martin Fanenbruck |author2=Steffen Harzsch  |author3=Johann Wolfgang Wägele  |name-list-style=amp |year=2004 |title=The brain of the Remipedia (Crustacea) and an alternative hypothesis on their phylogenetic relationships |journal=[[Proceedings of the National Academy of Sciences]] |doi=10.1073/pnas.0306212101 |volume=101 |issue=11 |pages=3868–3873 |pmid=15004272 |pmc=374336|doi-access=free }}</ref>
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The larvae are free-living and appear to be lecithotrophic (non-feeding). Mouths, guts, and anuses appear in the juvenile stage. Because of the energy and nutrients required for swimming, molting, and to grow in size and length, it has been speculated that the larvae may have other sources of growth than its yolk; possibly symbiotic bacteria.<ref>{{Cite book |last1=Klein |first1=Carel von Vaupel |url=https://books.google.com/books?id=0aLWAQAAQBAJ&dq=remipedia+lecithotrophic+yolk+carbohydrates+lipid&pg=PA155 |title=Treatise on Zoology - Anatomy, Taxonomy, Biology. The Crustacea, Volume 4 part A |last2=Charmantier-Daures |first2=Mireille |date=2013-10-24 |publisher=BRILL |isbn=978-90-474-4045-1 |language=en}}</ref><ref>{{cite book | url=https://books.google.com/books?id=Z2tEEAAAQBAJ&dq=remipedia+anoxic+zone+halocline+bacterial+symbionts&pg=PA741 | title=Evolution and Phylogeny of Pancrustacea: A Story of Scientific Method | isbn=978-0-19-971092-8 | last1=Schram | first1=Frederick R. | last2=Koenemann | first2=Stefan | date=16 October 2021 | publisher=Oxford University Press }}</ref>
The larvae are free-living and appear to be lecithotrophic (non-feeding). Mouths, guts, and anuses appear in the juvenile stage. Because of the energy and nutrients required for swimming, molting, and to grow in size and length, it has been speculated that the larvae may have other sources of growth than its yolk; possibly symbiotic bacteria.<ref>{{Cite book |last1=Klein |first1=Carel von Vaupel |url=https://books.google.com/books?id=0aLWAQAAQBAJ&dq=remipedia+lecithotrophic+yolk+carbohydrates+lipid&pg=PA155 |title=Treatise on Zoology - Anatomy, Taxonomy, Biology. The Crustacea, Volume 4 part A |last2=Charmantier-Daures |first2=Mireille |date=2013-10-24 |publisher=BRILL |isbn=978-90-474-4045-1 |language=en}}</ref><ref>{{cite book | url=https://books.google.com/books?id=Z2tEEAAAQBAJ&dq=remipedia+anoxic+zone+halocline+bacterial+symbionts&pg=PA741 | title=Evolution and Phylogeny of Pancrustacea: A Story of Scientific Method | isbn=978-0-19-971092-8 | last1=Schram | first1=Frederick R. | last2=Koenemann | first2=Stefan | date=16 October 2021 | publisher=Oxford University Press }}</ref>


With the exception of ''Speleonectes kakuki'', which inhabits a fully marine, sub-seafloor cave in the Bahamas, all known species of remipedians have been found exclusively in [[Anchialine system|anchialine cave systems]].<ref>{{Cite journal |last1=Neiber |first1=Marco T. |last2=Hartke |first2=Tamara R. |last3=Stemme |first3=Torben |last4=Bergmann |first4=Alexandra |last5=Rust |first5=Jes |last6=Iliffe |first6=Thomas M. |last7=Koenemann |first7=Stefan |date=2011 |title=Global Biodiversity and Phylogenetic Evaluation of Remipedia (Crustacea) |journal=PLOS ONE |volume=6 |issue=5 |pages=e19627 |doi=10.1371/journal.pone.0019627 |doi-access=free |pmc=3098257 |pmid=21625553|bibcode=2011PLoSO...619627N }}</ref>
With the exception of ''Speleonectes kakuki'', which inhabits a fully marine, sub-seafloor cave in the Bahamas, all known species of remipedians have been found exclusively in [[Anchialine system|anchialine cave systems]].<ref>{{Cite journal |last1=Neiber |first1=Marco T. |last2=Hartke |first2=Tamara R. |last3=Stemme |first3=Torben |last4=Bergmann |first4=Alexandra |last5=Rust |first5=Jes |last6=Iliffe |first6=Thomas M. |last7=Koenemann |first7=Stefan |date=2011 |title=Global Biodiversity and Phylogenetic Evaluation of Remipedia (Crustacea) |journal=PLOS ONE |volume=6 |issue=5 |article-number=e19627 |doi=10.1371/journal.pone.0019627 |doi-access=free |pmc=3098257 |pmid=21625553|bibcode=2011PLoSO...619627N }}</ref>


== History of classification ==
== History of classification ==
The first species in this group to be described was ''[[Speleonectes lucayensis]]'', discovered by [[Jill Yager]] while [[cave diving]] in [[Lucayan Caverns]] on the [[Grand Bahama Island]] in 1979 and described in a paper in the ''[[Journal of Crustacean Biology]]'' in 1981. The novel nature of this species was recognized and the class Remipedia was erected in the same paper.<ref name="Yager">{{cite journal|author=Jill Yager|date=August 1981|title=Remipedia, a new class of Crustacea from a marine cave in the Bahamas|journal=[[Journal of Crustacean Biology]]|volume=1|issue=3|pages=328–333|doi=10.2307/1547965|jstor=1547965|bibcode=1981JCBio...1..328Y }}</ref><ref>{{cite web|url=http://ww2.odu.edu/vhosts/sci/biology/directory/Holsinger/jrh/SUBTERR.HTM |title=What are subterranean amphipods? |author=John R. Holsinger |work=Systematics of amphipod crustaceans in the families Crangonyctidae and Hadziidae |access-date=October 25, 2013 |publisher=[[Old Dominion University]] |url-status=dead |archive-url=https://web.archive.org/web/20131029194258/http://ww2.odu.edu/vhosts/sci/biology/directory/Holsinger/jrh/SUBTERR.HTM |archive-date=October 29, 2013 }}</ref><ref>{{cite web |url=http://invertebrates.si.edu/Yager.html |title=Jill Yager, Research Associate |publisher=[[Smithsonian Institution]] |work=Invertebrate Zoology Staff |access-date=October 25, 2013}}</ref> The name "Remipedia" is from the [[Latin]] ''{{lang|la|remipedes}}'', meaning "oar-footed".<ref name="Yager"/>
The first species in this group to be described was ''[[Speleonectes lucayensis]]'', discovered by [[Jill Yager]] while [[cave diving]] in [[Lucayan Caverns]] on the [[Grand Bahama Island]] in 1979 and described in a paper in the ''[[Journal of Crustacean Biology]]'' in 1981. The novel nature of this species was recognized and the class Remipedia was erected in the same paper.<ref name="Yager">{{cite journal|author=Jill Yager|date=August 1981|title=Remipedia, a new class of Crustacea from a marine cave in the Bahamas|journal=[[Journal of Crustacean Biology]]|volume=1|issue=3|pages=328–333|doi=10.2307/1547965|jstor=1547965|bibcode=1981JCBio...1..328Y }}</ref><ref>{{cite web|url=http://ww2.odu.edu/vhosts/sci/biology/directory/Holsinger/jrh/SUBTERR.HTM |title=What are subterranean amphipods? |author=John R. Holsinger |work=Systematics of amphipod crustaceans in the families Crangonyctidae and Hadziidae |access-date=October 25, 2013 |publisher=[[Old Dominion University]] |archive-url=https://web.archive.org/web/20131029194258/http://ww2.odu.edu/vhosts/sci/biology/directory/Holsinger/jrh/SUBTERR.HTM |archive-date=October 29, 2013 }}</ref><ref>{{cite web |url=http://invertebrates.si.edu/Yager.html |title=Jill Yager, Research Associate |publisher=[[Smithsonian Institution]] |work=Invertebrate Zoology Staff |access-date=October 25, 2013}}</ref> The name "Remipedia" is from the [[Latin]] ''{{lang|la|remipedes}}'', meaning "oar-footed".<ref name="Yager"/>


Historical phylogeny based on morphology and physiology has placed Remipedia under [[Mandibulata]], in the subphylum [[Crustacean|Crustacea]], and distinct from [[Hexapoda]].
Historical phylogeny based on morphology and physiology has placed Remipedia under [[Mandibulata]], in the subphylum [[Crustacean|Crustacea]], and distinct from [[Hexapoda]].
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*** Genus ''[[Kaloketos]]'' <small>Koenemann, Iliffe & Yager 2004</small>
*** Genus ''[[Kaloketos]]'' <small>Koenemann, Iliffe & Yager 2004</small>
**** ''[[Kaloketos pilosus]]'' <small>Koenemann, Iliffe & Yager 2004</small>
**** ''[[Kaloketos pilosus]]'' <small>Koenemann, Iliffe & Yager 2004</small>
*** Genus ''[[Angirasu]]'' <small>Hoenemann et al. 2013</small>
*** Genus ''[[Angirasu (crustacean)|Angirasu]]'' <small>Hoenemann et al. 2013</small>
**** ''[[Angirasu benjamini]]'' <small>(Yager 1987) Hoenemann et al. 2013</small> [''Speleonectes benjamini'' <small>Yager 1987</small>]
**** ''[[Angirasu benjamini]]'' <small>(Yager 1987) Hoenemann et al. 2013</small> [''Speleonectes benjamini'' <small>Yager 1987</small>]
**** ''[[Angirasu parabenjamini]]'' <small>(Koenemann, Iliffe & van der Ham 2003) Hoenemann et al. 2013</small> [''Speleonectes parabenjamini'' <small>Koenemann, Iliffe & van der Ham 2003</small>]
**** ''[[Angirasu parabenjamini]]'' <small>(Koenemann, Iliffe & van der Ham 2003) Hoenemann et al. 2013</small> [''Speleonectes parabenjamini'' <small>Koenemann, Iliffe & van der Ham 2003</small>]

Latest revision as of 12:17, 17 October 2025

Template:Short description Template:Automatic taxobox

Remipedia is a class of blind crustacean-like animals, closely related to hexapods. They are found in coastal aquifers which contain saline groundwater, with populations identified in almost every ocean basin so far explored, including in Australia, the Caribbean Sea, and the Atlantic Ocean. The first described remipede was the fossil Tesnusocaris goldichi (Lower Pennsylvanian). Since 1979, at least seventeen living species have been identified in subtropical regions around the world.[1]

Description

Remipedes are Script error: No such module "convert". long and comprise a head and an elongate trunk of up to thirty-two similar body segments.[2] Pigmentation and eyes are absent.[3] Biramous swimming appendages are laterally present on each segment. The animals swim on their backs and are generally slow-moving.[4] They are the only known venomous crustaceans, and have fangs connected to secretory glands, which inject a combination of digestive enzymes and venom into their prey,[5] but they also feed through filter feeding. Being hermaphrodites, the female pore is located on the seventh trunk segment and the male pore on the fourteenth.[6]

Remipedia have a generally primitive body plan compared to other extant crustaceans, and are the only extant pancrustaceans to lack significant postcephalic tagmosis.[4] External respiratory structures like gills are absent.[7] Previously regarded as 'primitive', Remipedia have since been shown to have enhanced olfactory nerve centers (a common feature for species that live in dark environments).[8]

The larvae are free-living and appear to be lecithotrophic (non-feeding). Mouths, guts, and anuses appear in the juvenile stage. Because of the energy and nutrients required for swimming, molting, and to grow in size and length, it has been speculated that the larvae may have other sources of growth than its yolk; possibly symbiotic bacteria.[9][10]

With the exception of Speleonectes kakuki, which inhabits a fully marine, sub-seafloor cave in the Bahamas, all known species of remipedians have been found exclusively in anchialine cave systems.[11]

History of classification

The first species in this group to be described was Speleonectes lucayensis, discovered by Jill Yager while cave diving in Lucayan Caverns on the Grand Bahama Island in 1979 and described in a paper in the Journal of Crustacean Biology in 1981. The novel nature of this species was recognized and the class Remipedia was erected in the same paper.[12][13][14] The name "Remipedia" is from the Latin Script error: No such module "Lang"., meaning "oar-footed".[12]

Historical phylogeny based on morphology and physiology has placed Remipedia under Mandibulata, in the subphylum Crustacea, and distinct from Hexapoda.

New research in evolution and development reveals similarities between larvae and postembryonic development of remipedes and Malacostraca, singling Remipedia as a potential crustacean sister group of Hexapoda. Similarities in brain anatomy further support this affinity, and hexapod-type hemocyanins have been discovered in remipedes.[15]

Recent molecular studies have grouped Remipedia with Cephalocarida, Branchiopoda, and Hexapoda in a clade named Allotriocarida.[16][17] Remipedia was found as the sister group to Hexapoda both in phylogenomic[18][17] and combined morphological and transcriptome studies.[16] In other studies Remipedia and Cephalocarida are grouped together form the clade Xenocarida, which in turn was sister to Hexapoda in a clade named Anartiopoda[19] or Miracrustacea ('surprising crustaceans').[4]

The relationship of Remipedia and other crustacean classes and insects is shown in the following phylogenetic tree, which shows Allotriocarida, along with Oligostraca and Multicrustacea, as the three main divisions of subphylum Pancrustacea, embracing the traditional crustaceans and the hexapods (including insects).[17]

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Classification

Thirty extant species are recognized as of early 2022, divided among eight families and twelve genera.[20][21] All are placed in the order Nectiopoda. The second order, Enantiopoda, comprises the fossil species Tesnusocaris goldichi and Cryptocaris hootchi.[1]

Geographic distribution of extant Remipedia

References

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External links

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