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Loricifera (from
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
, '' lorica'', corselet (armour) + ''ferre'', to bear) is a
phylum In biology, a phylum (; plural: phyla) is a level of classification or taxonomic rank below kingdom and above class. Traditionally, in botany the term division has been used instead of phylum, although the International Code of Nomenclature f ...
of very small to microscopic marine
cycloneuralia Cycloneuralia is a clade of ecdysozoan animals including the Scalidophora (Kinorhynchans, Loriciferans, Priapulids) and the Nematoida ( nematodes, Nematomorphs). It may be paraphyletic, or may be a sister group to Panarthropoda. Or perhaps P ...
n sediment-dwelling animals that had been determined to be 37 described species, in 9 genera, but in 2021 has increased to 43 species. Aside from these described species, there are approximately 100 more that have been collected and not yet described. Their sizes range from 100 μm to ca. 1 mm. They are characterised by a protective outer case called a lorica and their habitat is in the spaces between marine gravel to which they attach themselves. The phylum was discovered in 1983 by R.M. Kristensen, near
Roscoff Roscoff (; br, Rosko) is a commune in the Finistère département of Brittany in northwestern France. Roscoff is renowned for its picturesque architecture, labelled (small town of character) since 2009. Roscoff is also a traditional departure ...
,
France France (), officially the French Republic ( ), is a country primarily located in Western Europe. It also comprises of Overseas France, overseas regions and territories in the Americas and the Atlantic Ocean, Atlantic, Pacific Ocean, Pac ...
. They are among the most recently discovered groups of animals. They attach themselves quite firmly to the substrate, and hence remained undiscovered for so long. The first specimen was collected in the 1970s, and later described in 1983. They are found at all depths, in different sediment types, and in all latitudes.


Morphology

The animals have a head, mouth, and digestive system, as well as the lorica. The head (which contains the mouth and the brain), a trunk region surrounded by six plates that make up the 'lorica' or corselet and – in between these two – the neck region. Loricifera have a well developed brain and each scalid is individually connected to the brain by nerves. The armor-like lorica consists of a protective external shell or case of encircling plicae. There is no circulatory system and no endocrine system. Many of the larvae are
acoelomate The coelom (or celom) is the main body cavity in most animals and is positioned inside the body to surround and contain the digestive tract and other organs. In some animals, it is lined with mesothelium. In other animals, such as molluscs, it r ...
, with some adults being
pseudocoelomate The coelom (or celom) is the main body cavity in most animals and is positioned inside the body to surround and contain the digestive tract and other organs. In some animals, it is lined with mesothelium. In other animals, such as molluscs, it ...
, and some remaining acoelomate. Development is generally direct, though there are so-called Higgins larvae, which differ from adults in several respects. As adults, the animals are
gonochoric In biology, gonochorism is a sexual system where there are only two sexes and each individual organism is either male or female. The term gonochorism is usually applied in animal species, the vast majority of which are gonochoric. Gonochorism c ...
. Very complex and plastic life cycles of pliciloricids include also paedogenetic stages with different forms of
parthenogenetic Parthenogenesis (; from the Greek grc, παρθένος, translit=parthénos, lit=virgin, label=none + grc, γένεσις, translit=génesis, lit=creation, label=none) is a natural form of asexual reproduction in which growth and development ...
reproduction. Fossils have been dated to the late Cambrian.


Taxonomic affinity

Morphological studies have traditionally placed the phylum in the
vinctiplicata The Vinctiplicata is a clade of Scalidophora uniting the Loricifera and the Priapulida, and representing the sister group to the Kinorhyncha. Its monophyly is supported on morphological grounds, although recent molecular studies indicate that the ...
with the
Priapulida Priapulida (priapulid worms, from Gr. πριάπος, ''priāpos'' 'Priapus' + Lat. ''-ul-'', diminutive), sometimes referred to as penis worms, is a phylum of unsegmented marine worms. The name of the phylum relates to the Greek god of fertility ...
; this plus the
Kinorhyncha Kinorhyncha ( grc, κινέω, kīnéō, I move, ' "snout") is a phylum of small marine invertebrates that are widespread in mud or sand at all depths as part of the meiobenthos. They are also called mud dragons. Modern species are or less, ...
constitutes the taxon
Scalidophora Scalidophora is a group of marine pseudocoelomate protostomes that was proposed on morphological grounds to unite three phyla: the Kinorhyncha, the Priapulida and the Loricifera. The three phyla have four characters in common — chitinous cuti ...
. The three phyla share four characters in common – chitinous cuticle, rings of scalids on the introvert, flosculi, and two rings of introvert retracts. However, despite a 2015 study showing the phylum's closest relatives being the
Panarthropoda Panarthropoda is a proposed animal clade containing the extant phyla Arthropoda, Tardigrada (water bears) and Onychophora (velvet worms). Panarthropods also include extinct marine legged worms known as lobopodians ("Lobopodia"), a paraphyletic g ...
, a 2022 study again showed that it belonged to the Scalidophora and told that further, more comprehensive genetic tests will be required to find its actual position in
Ecdysozoa Ecdysozoa () is a group of protostome animals, including Arthropoda (insects, chelicerata, crustaceans, and myriapods), Nematoda, and several smaller phyla. They were first defined by Aguinaldo ''et al.'' in 1997, based mainly on phylogenetic tr ...
.Howard, R. J., Giacomelli, M., Lozano-Fernandez, J., Edgecombe, G. D., Fleming, J. F., Kristensen, R. M., Ma, X., Olesen, J., Sørensen, M. V., Thomsen, P. F., Wills, M. A., Donoghue, P. C., & Pisani, D. (2022). The Ediacaran origin of ecdysozoa: Integrating fossil and Phylogenomic Data. ''Journal of the Geological Society'', 179 (4). https://doi.org/10.1144/jgs2021-107


Evolutionary history

The loriciferans are believed to be miniaturized descendants of a larger organism, perhaps resembling the
Cambrian The Cambrian Period ( ; sometimes symbolized C with bar, Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million ...
fossil ''
Sirilorica ''Sirilorica'' is a genus of stem-group scalidophoran from the Sirius Passet. Morphology ''Sirilorica'' was a worm-like animal ranging from three to eight centimetres in length. Ecology ''Sirilorica'' probably burrowed on the sediment surfa ...
''. However, the fossil record of the microscopic non-mineralized group is (perhaps unsurprisingly) scarce, so it is difficult to trace out the evolutionary history of the phylum in any detail. The 2017 discovery of Cambrian Period '' Eolorica deadwoodensis'' may shed some light on the group's history.


In anoxic environments

Three species of Loricifera have been found in the oxygen-free sediments at the bottom of the
L'Atalante basin L'Atalante basin is a hypersaline brine lake at the bottom of the Mediterranean Sea about west of the island of Crete. It is named for the French ''L'Atalante'', one of the oceanographic research vessels involved in its discovery in 1993. L'Atala ...
in
Mediterranean Sea The Mediterranean Sea is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean Basin and almost completely enclosed by land: on the north by Western and Southern Europe and Anatolia, on the south by North Africa, and on the ea ...
, more than 3,000 meters down, the first multicellular organisms known to spend their entire lives in an anoxic environment. Initially, it was thought that they are able to do this because their
mitochondria A mitochondrion (; ) is an organelle found in the Cell (biology), cells of most Eukaryotes, such as animals, plants and Fungus, fungi. Mitochondria have a double lipid bilayer, membrane structure and use aerobic respiration to generate adenosi ...
act like
hydrogenosome A hydrogenosome is a membrane-enclosed organelle found in some anaerobic ciliates, flagellates, and fungi. Hydrogenosomes are highly variable organelles that have presumably evolved from protomitochondria to produce molecular hydrogen and ATP in ...
s, allowing them to respire anaerobically. However, by 2021, questions arose as to whether or not they have mitochondria. The newly reported animals complete their life cycle in the total absence of light and oxygen, and they are less than a millimetre in size. They were collected from a deep basin at the bottom of the Mediterranean Sea, where they inhabit a nearly salt-saturated
brine Brine is a high-concentration solution of salt (NaCl) in water (H2O). In diverse contexts, ''brine'' may refer to the salt solutions ranging from about 3.5% (a typical concentration of seawater, on the lower end of that of solutions used for br ...
that, because of its density (> 1.2 g/cm³), does not mix with the waters above. As a consequence, this environment is completely anoxic and, due to the activity of
sulfate The sulfate or sulphate ion is a polyatomic anion with the empirical formula . Salts, acid derivatives, and peroxides of sulfate are widely used in industry. Sulfates occur widely in everyday life. Sulfates are salts of sulfuric acid and many ar ...
reducers, contains sulphide at a concentration of 2.9 mM. Despite such harsh conditions, this anoxic and sulphidic environment is teeming with microbial life, both chemosynthetic prokaryotes that are
primary producer Primary or primaries may refer to: Arts, entertainment, and media Music Groups and labels * Primary (band), from Australia * Primary (musician), hip hop musician and record producer from South Korea * Primary Music, Israeli record label Works ...
s, and a broad diversity of eukaryotic
heterotroph A heterotroph (; ) is an organism that cannot produce its own food, instead taking nutrition from other sources of organic carbon, mainly plant or animal matter. In the food chain, heterotrophs are primary, secondary and tertiary consumers, but ...
s at the next
trophic level The trophic level of an organism is the position it occupies in a food web. A food chain is a succession of organisms that eat other organisms and may, in turn, be eaten themselves. The trophic level of an organism is the number of steps it i ...
.


Taxa


References


Further reading

* * * * * * * {{Authority control Animal phyla Extant Cambrian first appearances