Remipedes Temporal range: | |
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Speleonectes tanumekes | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Superclass: | Allotriocarida |
Class: | Remipedia J. Yager 1981 |
Orders & families | |
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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]
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] 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]
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, 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]
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 remipedes, 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]
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]
Branchiopoda is a class of crustaceans. It comprises fairy shrimp, clam shrimp, Diplostraca, Notostraca, the Devonian Lepidocaris and possibly the Cambrian Rehbachiella. They are mostly small, freshwater animals that feed on plankton and detritus.
The Stenopodidea or boxer shrimps are a small group of decapod crustaceans. Often confused with Caridea shrimp or Dendrobranchiata prawns, they are neither, belonging to their own group.
Tesnusocaris goldichi is an extinct species of remipedian crustacean that lived in the Pennsylvanian period, one of the two representatives of the extinct remipedian order Enantiopoda. Its fossil is from the Lower Pennsylvanian Tesnus formation, Texas. The other known enantiopod remipedian is Cryptocaris hootchi of the Mazon Creek fauna.
Atelocerata is a proposed clade of arthropods that includes Hexapoda and Myriapoda, but excludes Crustacea and Chelicerata. The name is currently used interchangeably with Tracheata. or Uniramia sensu stricto. It is an extensive division of arthropods comprising all those that breathe by tracheae, as distinguished from Crustacea, which breathe by means of gills.
Pancrustacea is the clade that comprises all crustaceans, and all hexapods. This grouping is contrary to the Atelocerata hypothesis, in which Hexapoda and Myriapoda are sister taxa, and Crustacea are only more distantly related. As of 2010, the Pancrustacea taxon was considered well accepted, with most studies recovering Hexapoda within Crustacea. The clade has also been called Tetraconata, referring to having four cone cells in the ommatidia. The term "Tetraconata" is preferred by some scientists in order to avoid confusion with the use of "pan-" to indicate a clade that includes a crown group and all of its stem group representatives.
Speleonectidae is a family of remipedes in the order Nectiopoda. There are at least two genera and about seven described species in Speleonectidae.
Godzilliidae is a family of remipedes in the order Nectiopoda. There are at least two genera and four described species in Godzilliidae.
The hairy stone crab is a crab-like anomuran crustacean that lives in the littoral zone of southern Australia from Bunbury, Western Australia, to the Bass Strait. It is the only species in the family Lomisidae. It is 1.5–2.5 cm (0.6–1.0 in) wide, slow-moving, and covered in brown hair which camouflages it against the rocks upon which it lives.
The subphylum Mandibulata constitutes one of the major subdivisions of the phylum Arthropoda, alongside Chelicerata. Mandibulates include the crustaceans, myriapods, and all true insects. The name "Mandibulata" refers to the mandibles, a modified pair of limbs used in food processing, the presence of which are characteristic of most members of the group.
Morlockia atlantida is a species of eyeless crustacean in the order Nectiopoda. It was discovered in August 2009 in the Túnel de la Atlántida, the world's longest submarine lava tube on Lanzarote in the Canary Islands off the west coast of North Africa. Originally named Speleonectes atlantida, it was transferred to genus Morlockia in 2012. Like other remipedes, the species is equipped with venomous fangs.
Xenocarida is a proposed clade inside the subphylum Crustacea that comprises two classes that were discovered in the 20th century: Remipedia and Cephalocarida. Both groups are marine hermaphrodites. The clade was recovered as the sister groups to Hexapoda.
Crustaceans may pass through a number of larval and immature stages between hatching from their eggs and reaching their adult form. Each of the stages is separated by a moult, in which the hard exoskeleton is shed to allow the animal to grow. The larvae of crustaceans often bear little resemblance to the adult, and there are still cases where it is not known what larvae will grow into what adults. This is especially true of crustaceans which live as benthic adults, more-so than where the larvae are planktonic, and thereby easily caught.
Godzilliognomus schrami is a species of Remipedia discovered in 2007, representing one of five extant species of the family Godzilliidae. It generally reaches about 7 millimetres (0.28 in) in length and inhabits a single anchialine cave on Eleuthera Island. The specific epithet commemorates Frederick Schram.
The subphylum Hexapoda or hexapods comprises the largest clade of arthropods and includes most of the extant arthropod species. It includes the crown group class Insecta, as well as the much smaller clade Entognatha, which includes three classes of wingless arthropods that were once considered insects: Collembola (springtails), Protura (coneheads) and Diplura. The insects and springtails are very abundant and are some of the most important pollinators, basal consumers, scavengers/detritivores and micropredators in terrestrial environments.
Crustaceans are invertebrate animals that constitute one group of arthropods that are a part of the subphylum Crustacea, a large, diverse group of mainly aquatic arthropods including decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, opossum shrimps, amphipods and mantis shrimp. The crustacean group can be treated as a subphylum under the clade Mandibulata. It is now well accepted that the hexapods emerged deep in the Crustacean group, with the completed pan-group referred to as Pancrustacea. The three classes Cephalocarida, Branchiopoda and Remipedia are more closely related to the hexapods than they are to any of the other crustaceans.
Xibalbanus tulumensis is a venomous, hermaphroditic crustacean found in anchialine caves on the Yucatán Peninsula in the Caribbean Sea. This blind remipede liquefies the body contents of other crustaceans with a venom similar to that of rattlesnakes, and which includes digestive enzymes and a paralysing toxin.
Pleomothra is a genus of crustacean found in the Caribbean. First described in 1989, the genus has 2 identified species as of 2008. It is the sole member of family Pleomothridae, but was previously placed in family Godzilliidae.
The clade Multicrustacea constitutes the largest superclass of crustaceans, containing approximately four-fifths of all described non-hexapod crustacean species, including crabs, lobsters, crayfish, shrimp, krill, prawns, woodlice, barnacles, copepods, amphipods, mantis shrimp and others. The largest branch of multicrustacea is the class Malacostraca.
Morlockia is a genus of eyeless crustaceans in the family Morlockiidae. The genus contains 4 described species.
Allotriocarida is a clade of Pancrustacea, containing Hexapoda. It also contains three non-hexapod classes: Remipedia, Cephalocarida, and Branchiopoda. Newer studies also relocate copepoda, which traditionally has belonged to the multicrustaceans, to the Allotriocarida.