Remipedia

Life-forms

Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Ecdysozoa
Cladus: Panarthropoda
Phylum: Arthropoda
Cladus: Pancrustacea
Cladus: Allotriocarida
Classis: Remipedia
Ordines (2): Nectiopoda – †Enantiopoda

Name

Remipedia Yager, 1981
References
Primary references

Yager, J. 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 Paywall. JSTOR Hybrid open access journal. Reference page.

Additional references

Emerson, M.J. & Schram, F.R. 1991. Remipedia. Part 2. Paleontology. Proceedings of the San Diego Society of Natural History 7: 1–52. BHL. Reference page.
Koenemann, S.; Schram, F.R.; Hönemann, M.; Iliffe, T.M. 2006: Phylogenetic analysis of Remipedia (Crustacea). Organisms diversity & evolution, 7(1): 33–51. DOI: 10.1016/j.ode.2006.07.001 PDF
Schram, F.R., Yager, J. & Emerson, M.J. 1986. Remipedia. Part 1. Systematics. Memoirs of the San Diego Society of Natural History 15: 1–60. BHL. Reference page.

Links

Remipedia in the World Remipedia Database

Vernacular names
Ελληνικά: Ρεμίποδα
English: remipedes
中文: 桨足纲

Remipedia is a class of blind crustaceans, closely related to hexapods, 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 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] 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] 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).[7]

Based on studies of the free-living larvae, they appear to be lecithotrophic (non-feeding). Mouths, guts, and anuses appear in the juvenile stage. Because of the energy and nutrients required for swimming, molt several times 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.[8][9]

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.[10]
History of classification

The class Remipedia was erected in 1981 by Jill Yager, in describing Speleonectes lucayensis from the Bahamas.[11] The name "Remipedia" is from the Latin remipedes, meaning "oar-footed".[11]

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.[12]

Recent molecular studies have grouped Remipedia with Cephalocarida, Branchiopoda, and Hexapoda in a clade named Allotriocarida.[13][14] Remipedia was found as the sister group to Hexapoda both in phylogenomic[15] [14] and combined morphological and transcriptome studies.[13] 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[16] 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).[14]

Pancrustacea	Oligostraca Multicrustacea Thecostraca Copepoda Malacostraca Allotriocarida Cephalocarida Branchiopoda Remipedia Hexapoda Protura Diplura Collembola Insecta

Classification

Thirty extant species are recognized as of early 2022, divided among eight families and twelve genera.[17][18] All are placed in the order Nectiopoda. The second order, Enantiopoda, comprises the fossil species Tesnusocaris goldichi and Cryptocaris hootchi.[1]

†Order Enantiopoda Birshtein 1960
†Family Tesnusocarididae Brooks 1955 [Cryptocarididae Sieg 1980]
Genus †Tesnusocaris Brooks 1955
†Tesnusocaris goldichi Brooks 1955
Genus †Cryptocaris Schram 1974
†Cryptocaris hootchi Schram 1974
Order Nectiopoda Schram 1986
Family Micropacteridae Koenemann, Iliffe & van der Ham 2007
Genus Micropacter Koenemann, Iliffe & van der Ham 2007
Micropacter yagerae Koenemann, Iliffe & van der Ham 2007
Family Godzilliidae Schram, Yager & Emerson 1986
Genus Godzilliognomus Yager 1989
Godzilliognomus frondosus Yager, 1989
Godzillognomus schrami Iliffe, Otten & Koenemann 2010
Genus Godzillius Schram et al., 1986
Godzillius fuchsi Gonzalez, Singpiel & Schlagner 2013
Godzillius robustus Schram, Yager & Emerson 1986
Family Kumongidae Hoenemann et al. 2013
Genus Kumonga Hoenemann et al. 2013
Kumonga exleyi (Yager & Humphreys 1996) Hoenemann et al. 2013 [Lasionectes exleyi Yager & Humphreys 1996]
Family Cryptocorynetidae Hoenemann et al. 2013
Genus Kaloketos Koenemann, Iliffe & Yager 2004
Kaloketos pilosus Koenemann, Iliffe & Yager 2004
Genus Angirasu Hoenemann et al. 2013
Angirasu benjamini (Yager 1987) Hoenemann et al. 2013 [Speleonectes benjamini Yager 1987]
Angirasu parabenjamini (Koenemann, Iliffe & van der Ham 2003) Hoenemann et al. 2013 [Speleonectes parabenjamini Koenemann, Iliffe & van der Ham 2003]
Genus Cryptocorynetes Yager 1987
Cryptocorynetes elmorei Hazerli, Koenemann & Iliffe 2009 [19]
Cryptocorynetes haptodiscus Yager 1987
Cryptocorynetes longulus Wollermann, Koenemann & Iliffe 2007
Family Morlockiidae García-Valdecasas 1984
Genus Morlockia García-Valdecasas 1984
Morlockia williamsi (Hartke, Koenemann & Yager 2011) [Speleonectes williamsi Hartke, Koenemann & Yager 2011][20]
Morlockia emersoni (Lorentzen, Koenemann & Iliffe 2007) [Speleonectes emersoni Lorentzen, Koenemann & Iliffe 2007]
Morlockia atlantida (Koenemann et al. 2009) Hoenemann et al. 2012 [Speleonectes atlantidus Koenemann et al. 2009]
Morlockia ondinae García-Valdecasas 1984 [Speleonectes ondinae (Garcia-Valdecasas 1984)]
Family Speleonectidae Yager 1981
Genus Lasionectes Yager & Schram, 1986
Lasionectes entrichoma Yager & Schram, 1986
Genus Speleonectes Yager 1981
Speleonectes epilimnius Yager & Carpenter, 1999
Speleonectes gironensis Yager, 1994
Speleonectes kakuki Daenekas et al., 2009
Speleonectes lucayensis Yager, 1981
Speleonectes minnsi Koenemann, Iliffe & van der Ham, 2003
Speleonectes tanumekes Koenemann, Iliffe & van der Ham, 2003
Family Xibalbanidae Olesen et al. 2017
Genus Xibalbanus Hoenemann et al. 2013
Xibalbanus cokei (Yager, 2013) Olesen et al. 2017 [Speleonectes cokei Yager, 2013][21]
Xibalbanus cozumelensis Olesen, Meland, Glenner, van Hengstum & Iliffe, 2017
Xibalbanus fuchscockburni (Neiber et al. 2012) Hoenemann et al. 2013 [Speleonectes fuchscockburni Neiber et al. 2012][22]
Xibalbanus tulumensis (Yager 1987) Hoenemann et al. 2013 [Speleonectes tulumensis Yager 1987]
Family Pleomothridae Hoenemann et al. 2013
Genus Pleomothra Yager 1989
Pleomothra apletocheles Yager 1989
Pleomothra fragilis Koenemann, Ziegler & Iliffe 2008

Geographic distribution of extant Remipedia

Bahamas – Andros, Sweetings Cay, Grand Bahama, Great Exuma, Great Guana Cay (Exuma Cays), Cat Island, Abaco Islands, San Salvador Island
Turks and Caicos Islands – North Caicos, Providenciales
Australia – North West Cape (Western Australia)
Cuba – Matanzas Province
Spain – Lanzarote (Canary Islands)
Mexico – Quintana Roo
Belize - Caye_Chapel
Dominican Republic – Distrito Nacional Cueva Taína, Santo Domingo Este.[23]

References

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.
Cameron McCormick (November 10, 2008). "Remipedia". The Lord Geekington.
Yager, J. (18 September 2013). "Lasionectes entrichoma Yager & Schram, 1986". tamug.edu. Retrieved 9 February 2018.
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.
Kaplan, Matt (22 October 2013). "First venomous crustacean discovered". Nature News. doi:10.1038/nature.2013.13985. S2CID 87091184. Retrieved 10 May 2015.
Behavior of Remipedia in the Laboratory, with Supporting Field Observations
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.
Treatise on Zoology - Anatomy, Taxonomy, Biology. The Crustacea, Volume 4 part A
Evolution and Phylogeny of Pancrustacea: A Story of Scientific Method
Global Biodiversity and Phylogenetic Evaluation of Remipedia (Crustacea)
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.
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.
Oakley, Todd H.; Wolfe, Joanna M.; Lindgren, Annie R.; Zaharoff, Alexander K. (2013). "Phylotranscriptomics to Bring the Understudied into the Fold: Monophyletic Ostracoda, Fossil Placement, and Pancrustacean Phylogeny". Molecular Biology and Evolution. 30: 215–233. doi:10.1093/molbev/mss216. PMID 22977117.
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. PMC 6684935. PMID 31270537.
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.
Engel, Michael (2015). "Insect evolution". Current Biology. 25 (19): R868–R872. doi:10.1016/j.cub.2015.07.059. PMID 26439349. S2CID 14406214.
Koenemann, S.; Hoenemann, M.; Stemme T. (2022). "World Remipedia Database". Vlaams Instituut voor de Zee. Retrieved 7 February 2022.
World Remipedia Database. "Remipedia". World Register of Marine Species. Retrieved 7 February 2022.
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.
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.
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.
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.
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.