Phoronis Architecta
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''Phoronis'' is one of the two
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
of the horseshoe worm family (
Phoronidae Phoronids (scientific name Phoronida, sometimes called horseshoe worms) are a small phylum of marine animals that filter-feed with a lophophore (a "crown" of tentacles), and build upright tubes of chitin to support and protect their soft bodie ...
), in the
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 ...
Phoronida. The body has two sections, each with its own
coelom 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 ...
. There is a specialist feeding structure, the
lophophore The lophophore () is a characteristic feeding organ possessed by four major groups of animals: the Brachiopoda, Bryozoa, Hyolitha, and Phoronida, which collectively constitute the protostome group Lophophorata.filter feeder Filter feeders are a sub-group of suspension feeding animals that feed by straining suspended matter and food particles from water, typically by passing the water over a specialized filtering structure. Some animals that use this method of feedin ...
s. They live on hard substrates or soft sediments in marine environments throughout the world. They have different modes of reproduction which help with their success. The scientific name of the larval form is ''
Actinotrocha ''Phoronis psammophila'' is a species of marine horseshoe worm in the phylum Phoronida. It lives in a tube projecting from the sea floor in shallow seas around the world. Description ''Phoronis psammophila'' constructs and lives in a rigid, chit ...
''.


Etymology

The generic name refers to
Phoronis ''Phoronis'' is one of the two genera of the horseshoe worm family (Phoronidae), in the phylum Phoronida. The body has two sections, each with its own coelom. There is a specialist feeding structure, the lophophore, which is an extension of the ...
(better known as Io), a Greek mythological character sometimes conflated with
Isis Isis (; ''Ēse''; ; Meroitic: ''Wos'' 'a''or ''Wusa''; Phoenician: 𐤀𐤎, romanized: ʾs) was a major goddess in ancient Egyptian religion whose worship spread throughout the Greco-Roman world. Isis was first mentioned in the Old Kingd ...
. Thomas Strethill Wright, of Edinburgh, did not give a specific reason for choosing the name.


Species

* ''
Phoronis australis ''Phoronis australis'' is a species of marine horseshoe worm in the phylum Phoronida. It is found in shallow warm-temperate and tropical waters in the eastern Atlantic Ocean and the Indo-Pacific region and was first detected in the Mediterranean S ...
'' Haswell 1883 * ''
Phoronis embryolabi ''Phoronis embryolabi'' is a species of marine horseshoe worm in the phylum Phoronida. It is found in Vostok Bay, where it lives together with '' Nihonotrypaea japonica'', an Axiidea Axiidea is an infraorder of decapod crustaceans. They are ...
'' Temereva & Chichvarkhin, 2017 * ''
Phoronis emigi ''Phoronis emigi'' is a species of marine horseshoe worm in the phylum Phoronida. It is found off the coast of Japan and is morphologically similar to ''Phoronis psammophila ''Phoronis psammophila'' is a species of marine horseshoe worm in the ...
'' Hirose et al. 2014 * ''
Phoronis hippocrepia ''Phoronis hippocrepia'' is a species of marine horseshoe worm in the phylum Phoronid Phoronids (scientific name Phoronida, sometimes called horseshoe worms) are a small phylum of marine animals that filter-feed with a lophophore (a "crown" o ...
'' Wright 1856 * ''
Phoronis ijimai ''Phoronis ijimai'' is a species of marine horseshoe worm in the phylum Phoronid Phoronids (scientific name Phoronida, sometimes called horseshoe worms) are a small phylum of marine animals that filter-feed with a lophophore (a "crown" of ten ...
'' Oka 1897 * ''
Phoronis muelleri ''Phoronis muelleri'' is a species of marine horseshoe worm in the phylum Phoronid Phoronids (scientific name Phoronida, sometimes called horseshoe worms) are a small phylum of marine animals that filter-feed with a lophophore (a "crown" of t ...
'' Selys-Lonchamps 1903 * ''
Phoronis ovalis ''Phoronis ovalis'' is a species of marine horseshoe worm in the phylum Phoronida. It is found in shallow waters in the northeastern Atlantic Ocean, the southeastern Atlantic Ocean, Argentina, and other scattered locations worldwide. These worms s ...
'' Wright 1856 * ''
Phoronis pallida ''Phoronis pallida'' is a species of marine horseshoe worm in the phylum Phoronid Phoronids (scientific name Phoronida, sometimes called horseshoe worms) are a small phylum of marine animals that filter-feed with a lophophore (a "crown" of te ...
'' (Schneider 1862) Silén 1952 * ''
Phoronis psammophila ''Phoronis psammophila'' is a species of marine horseshoe worm in the phylum Phoronida. It lives in a tube projecting from the sea floor in shallow seas around the world. Description ''Phoronis psammophila'' constructs and lives in a rigid, chit ...
'' Cori 1889World Register of Marine Species
/ref> * ''
Phoronis savinkini ''Phoronis savinkini'' is a species of marine horseshoe worm in the phylum Phoronida. It is found in the Gulf of Tonkin. The holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the s ...
'' Temereva & Neklyudov, 2018 ''Phoronis architecta'' is no longer accepted and is considered to be a synonym of ''
Phoronis psammophila ''Phoronis psammophila'' is a species of marine horseshoe worm in the phylum Phoronida. It lives in a tube projecting from the sea floor in shallow seas around the world. Description ''Phoronis psammophila'' constructs and lives in a rigid, chit ...
''. Similarly ''Phoronis vancouverensis'' is considered to be a synonym of ''
Phoronis ijimai ''Phoronis ijimai'' is a species of marine horseshoe worm in the phylum Phoronid Phoronids (scientific name Phoronida, sometimes called horseshoe worms) are a small phylum of marine animals that filter-feed with a lophophore (a "crown" of ten ...
''. A 2018 phylogenetic analysis indicates that ''Phoronis'' is
paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
, while ''
Phoronopsis ''Phoronopsis'' is a genus of horseshoe worm in the family Phoronidae, in the phylum Phoronida. The members of the genus live in tubes at the bottom of the sea. Characteristics Like other phoronids, members of this genus are benthic filter feed ...
'' is a monophyletic genus.Temereva, E. N.; Neklyudov, B. V. (2018). A New Phoronid Species, ''Phoronis savinkini'' sp. n., from the South China Sea and an Analysis of the Taxonomic Diversity of Phoronida. ''Biology Bulletin''. 45(7): 617-639.


Distribution

Horseshoe worms are not very sensitive to environmental conditions and are therefore considered to be both eurythermic and euryhaline animals. While they are not very sensitive, there are some factors that can impact their distribution such as current strength (as they are filter feeders), the result of asexual reproduction, predation in the area, and local fauna which they may have to compete with. However, horseshoe worms have a relatively global distribution as they can be found in the Atlantic, Pacific, and Indian oceans.


Structure


Tube

The horseshoe worms live in tubes that are fully imbedded in the soil and are formed during digging or boring. The tubes are produced by the worm's epidermal glands. These tubes have an inner organic cylinder and an outer layer. The inner layer is composed of three parts: the inner film, middle film, and outer film. These thin films are composed of fibers from the worms that form nets that make up these tubes.


Nervous and digestive systems

These worms have a nervous system that consists of a dorsal ganglion, a tentacle nerve ring, an inner nerve ring, intertentacular groups of perikarya, and tentacle nerves. These tentacles nerves have three groups: abfrontal, frontal, and laterofrontal. They have a U-shaped gut, meaning their anus is close in distance to their mouths and their digestive tract consists of three parts: the mouth, the stomach, and the anus.


Lophophore

The lophophore structure has evolved from a simple, oval shape to the horseshoe shape from which the worms get their name, even into a complex spiral structure. These worms can regenerate their lophophores when injured or even voluntarily drop their lophophores as ''Phoronis ovalis'' does when it is laying eggs. The lophophore is used for both respiration and feeding. The tentacles of the lophophores have cilia on them which move water towards their mouth to filter out small particulates. Through moving water over the tentacles of the lophophore, they act as an organ for gas exchange and allow oxygen into the blood as it passes through.


Development


Embryos

Generally, Phoronis species have benthic adults with lecithotrophic larvae. These species incubate their embryos in various ways. Some species release their embryos into the water column and they develop there. In other species, embryos are incubated among the tentacles of the adult worms. Recently, it was discovered that some species even exhibit viviparity, where the embryos are incubated in the coelom of their mother's tube and the "hatch" to produce feeding actinotroch larvae. There is a correlation between egg size and development type: allowing embryos into the water column to develop takes less energy and can therefore produce more eggs. Incubating in the tube's coelom takes more energy so less eggs are produced.


Larvae

The lecithotrophic larvae, given the name actinotroch, are pelagic and float in the water column. This allows for wider dispersal of species. The larvae have also developed different morphologies over time such as differing pigmentation and arrangement of blood masses.


Adult

When they are full grown, these worms typically have thin but long bodies that can grow up to 50 cm. The adults are typically colonial and can grow around each other in order to support each other.


Reproduction

Phoronis can be either gonochoristic or hermaphroditic. The worms create sperm and eggs through the processes of spermiogenesis and oogenesis. Spermiogenesis leads to the creation of a spermatophore, which is inserted into the metacoelom to combine with the eggs in two ways. The small fertilized eggs, numbering up to 500, are then released through spawning into the water column where the larvae develop over a period of 3 weeks before settling to change into the adult stage. Others lay fewer, larger eggs, and after fertilization, do not release them until they have hatched. These larvae are only in the water column for a period of 4 days before settling to find their home.


Parasite

Some phoronid species have microsporidia-like spores that were discovered in 2017. This was the first recorded instance of phoronids being hosts to parasites. These microsporidia were named ''Microsporidium phoronidi.''


References

{{Taxonbar, from=Q2341489 Phoronids