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Temporal range: Lower Pennsylvanian–Recent
Speleonectes tanumekes unlabeled.png
Speleonectes tanumekes
Scientific classification e
Kingdom: Animalia
Phylum: Arthropoda
Subphylum: Crustacea
Class: Remipedia
J. Yager 1981
Orders & families
  • †Enantiopoda
    • †Tesnusocarididae
  • Nectiopoda

Remipedia is a class of blind crustaceans 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]


Remipedes are 1–4 centimetres (0.4–1.6 in) long and comprise a head and an elongate trunk of up to thirty-two similar body segments.[2] They lack eyes and pigmentation.[3] Biramous swimming appendages are laterally present on each segment, and the animals swim on their backs. They 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]

Remipedia have a generally primitive body plan compared to other extant crustaceans, and are the only extant pancrustaceans to lack significant postcephalic tagmosis.[4] 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).[6]

History of classification[edit]

The class Remipedia was erected in 1981 by Jill Yager, in describing Speleonectes lucayensis from the Bahamas.[7] The name "Remipedia" is from the Latin remipedes, meaning "oar-footed".[7]

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.[8]

Recent analysis based on nuclear protein-coding genes implies that remipedes (along with Cephalocarida) are the sister group of arthropods most closely related to insects. Remipedia and Cephalocarida are grouped together form the clade Xenocarida.[9] The extant lineages Xenocarida, Vericrustacia, Oligostraca, and Hexapoda together form the proposed class Pancrustacea. The clade names Anartiopoda[10] or Miracrustacea ('surprising crustaceans') has been proposed for the monophyletic clade containing xenocarids and hexapods.[4]














Phylogenetic position of Remipedia within Pancrustacea[11]

Thirty extant species are recognized as of early 2022, divided among eight families and twelve genera.[12][13] 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[edit]


  1. ^ a b Stefan Koenemann; Frederick R. Schram; Mario Hönemann & Thomas M. Iliffe (2007). "Phylogenetic analysis of Remipedia (Crustacea)". Organisms Diversity & Evolution. 7 (1): 33–51. doi:10.1016/j.ode.2006.07.001.
  2. ^ Cameron McCormick (November 10, 2008). "Remipedia". The Lord Geekington.
  3. ^ Yager, J. (18 September 2013). "Lasionectes entrichoma Yager & Schram, 1986". Retrieved 9 February 2018.
  4. ^ a b c Regier, Jerome C.; Shultz, Jeffrey W.; Zwick, Andreas; Hussey, April; Ball, Bernard; Wetzer, Regina; Martin, Joel W.; Cunningham, Clifford W. (February 2010). "Arthropod relationships revealed by phylogenomic analysis of nuclear protein-coding sequences". Nature. 463 (7284): 1079–1083. Bibcode:2010Natur.463.1079R. doi:10.1038/nature08742. ISSN 0028-0836. PMID 20147900. S2CID 4427443.
  5. ^ Kaplan, Matt (22 October 2013). "First venomous crustacean discovered". Nature News. doi:10.1038/nature.2013.13985. S2CID 87091184. Retrieved 10 May 2015.
  6. ^ Martin Fanenbruck; Steffen Harzsch & Johann Wolfgang Wägele (2004). "The brain of the Remipedia (Crustacea) and an alternative hypothesis on their phylogenetic relationships". Proceedings of the National Academy of Sciences. 101 (11): 3868–3873. doi:10.1073/pnas.0306212101. PMC 374336. PMID 15004272.
  7. ^ a b Jill Yager (August 1981). "Remipedia, a new class of Crustacea from a marine cave in the Bahamas". Journal of Crustacean Biology. 1 (3): 328–333. doi:10.2307/1547965. JSTOR 1547965.
  8. ^ Giribet, Gonzalo; Edgecombe, Gregory D. (2012-01-07). "Reevaluating the Arthropod Tree of Life". Annual Review of Entomology. 57 (1): 167–186. doi:10.1146/annurev-ento-120710-100659. ISSN 0066-4170. PMID 21910637. S2CID 207597767.
  9. ^ Bjoern M. von Reumont; Ronald A. Jenner; Matthew A. Wills; Emiliano Dell'Ampio; Günther Pass; Ingo Ebersberger; Benjamin Meyer; Stefan Koenemann; Thomas M. Iliffe; Alexandros Stamatakis; Oliver Niehuis; Karen Meusemann & Bernhard Misof (March 2012). "Pancrustacean phylogeny in the light of new phylogenomic data: support for Remipedia as the possible sister group of Hexapoda". Molecular Biology and Evolution. 29 (3): 1031–1045. doi:10.1093/molbev/msr270. PMID 22049065.
  10. ^ Engel, Michael (2015). "Insect evolution". Current Biology. 25 (19): R868–R872. doi:10.1016/j.cub.2015.07.059. PMID 26439349. S2CID 14406214.
  11. ^ Lozano-Fernandez, Jesus; Giacomelli, Mattia; Fleming, James F; Chen, Albert; Vinther, Jakob; Thomsen, Philip Francis; Glenner, Henrik; Palero, Ferran; Legg, David A; Iliffe, Thomas M; Pisani, Davide; Olesen, Jørgen (2019). "Pancrustacean Evolution Illuminated by Taxon-Rich Genomic-Scale Data Sets with an Expanded Remipede Sampling". Genome Biology and Evolution. 11 (8): 2055–2070. doi:10.1093/gbe/evz097. ISSN 1759-6653. PMC 6684935. PMID 31270537.
  12. ^ Koenemann, S.; Hoenemann, M.; Stemme T. (2022). "World Remipedia Database". Vlaams Instituut voor de Zee. Retrieved 7 February 2022.
  13. ^ World Remipedia Database. "Remipedia". World Register of Marine Species. Retrieved 7 February 2022.
  14. ^ Dennis Hazerli; Stefan Koenemann & Thomas M. Iliffe (2010). "Cryptocorynetes elmorei, a new species of Remipedia (Crustacea) from an anchialine cave on Eleuthera, Bahamas". Marine Biodiversity. 40 (2): 71–78. doi:10.1007/s12526-009-0033-4. S2CID 8082592.
  15. ^ Tamara R. Hartke; Stefan Koenemann & Jill Yager (2011). "Speleonectes williamsi, a new species of Remipedia (Crustacea) from the Bahamas" (PDF excerpt). Zootaxa. 3115: 21–28. doi:10.11646/zootaxa.3115.1.2.
  16. ^ Yager J (2013). "Speleonectes cokei, new species of Remipedia (Crustacea: Speleonectidae) from a submerged ocean cave near Caye Chapel, Belize". Zootaxa. 3710 (4): 354–362. doi:10.11646/zootaxa.3710.4.4. PMID 26106696. S2CID 10850210.
  17. ^ Marco T. Neiber; Finja C. Hansen; Thomas M. Iliffe; Brett C. Gonzalez & Stefan Koenemann (2012). "Molecular taxonomy of Speleonectes fuchscockburni, a new pseudocryptic species of Remipedia (Crustacea) from an anchialine cave system on the Yucatán Peninsula, Quintana Roo, Mexico" (PDF excerpt). Zootaxa. 3190: 31–46. doi:10.11646/zootaxa.3190.1.2.
  18. ^ Lorentzen, Dörte; Koenemann, Stefan; Iliffe, Thomas M. (2007). "Speleonectes emersoni, A New Species of Remipedia (Crustacea) from the Dominican Republic". Zootaxa. 1543: 61–68. doi:10.11646/zootaxa.1543.1.3.

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