Horseshoe worm
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Phoronids (scientific name Phoronida, sometimes called horseshoe worms) are a small
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 marine animals that filter-feed with a lophophore (a "crown" of tentacles), and build upright tubes of
chitin Chitin ( C8 H13 O5 N)n ( ) is a long-chain polymer of ''N''-acetylglucosamine, an amide derivative of glucose. Chitin is probably the second most abundant polysaccharide in nature (behind only cellulose); an estimated 1 billion tons of chit ...
to support and protect their soft bodies. They live in most of the oceans and seas, including the Arctic Ocean but excluding the Antarctic Ocean, and between the intertidal zone and about 400 meters down. Most adult phoronids are 2 cm long and about 1.5 mm wide, although the largest are 50 cm long. The name of the group comes from its type genus: ''
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 ...
''.


Overview

The bottom end of the body is an ampulla (a flask-like swelling), which anchors the animal in the tube and enables it to retract its body very quickly when threatened. When the lophophore is extended at the top of the body,
cilia The cilium, plural cilia (), is a membrane-bound organelle found on most types of eukaryotic cell, and certain microorganisms known as ciliates. Cilia are absent in bacteria and archaea. The cilium has the shape of a slender threadlike projecti ...
(little hairs) on the sides of the tentacles draw food particles to the mouth, which is inside and slightly to one side of the base of the lophophore. Unwanted material can be excluded by closing a lid above the mouth or be rejected by the tentacles, whose cilia can switch into reverse. The food then moves down to the stomach, which is in the ampulla. Solid wastes are moved up the intestine and out through the
anus The anus (Latin, 'ring' or 'circle') is an opening at the opposite end of an animal's digestive tract from the mouth. Its function is to control the expulsion of feces, the residual semi-solid waste that remains after food digestion, which, d ...
, which is outside and slightly below the lophophore. A blood vessel leads up the middle of the body from the stomach to a circular vessel at the base of the lophophore, and from there a single blind vessel runs up each tentacle. A pair of blood vessels near the body wall lead downward from the lophophore ring to the stomach and also to blind branches throughout the body. There is no heart, but the major vessels can contract in waves to move the blood. Phoronids do not ventilate their trunks with oxygenated water, but rely on respiration through the lophophore. The blood contains hemoglobin, which is unusual in such small animals and seems to be an adaptation to anoxic and hypoxic environments. The blood of ''Phoronis architecta'' carries twice as much oxygen as a human of the same weight. Two metanephridia filter the body fluid, returning any useful products and dumping the remaining soluble wastes through a pair of pores beside the anus. One species builds colonies by budding or by splitting into top and bottom sections, and all phoronids
reproduce sexually Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete ( haploid reproductive cells, such as a sperm or egg cell) with a single set of chromosomes combines with another gamete to produce a zygote tha ...
from spring to autumn. The eggs of most species form free-swimming actinotroch larvae, which feed on plankton. An actinotroch settles to the seabed after about 20 days and then undergoes a radical change in 30 minutes: the larval tentacles are replaced by the adult lophophore; the anus moves from the bottom to just outside the lophophore; and this changes the gut from upright to a U-bend, with the stomach at the bottom of the body. One species forms a "slug-like" larva, and the larvae of a few species are not known. Phoronids live for about one year. Some species live separately, in vertical tubes embedded in soft sediment, while others form tangled masses buried in or encrusting rocks and shells. In some habitats populations of phoronids reach tens of thousand of individuals per square meter. The actinotroch larvae are familiar among plankton, and sometimes account for a significant proportion of the zooplankton biomass. Predators include fish,
gastropod The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. T ...
s (snails), and
nematode The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-Parasitism, parasitic nematodes also known as eelworms. They are a diverse animal phylum inhab ...
s (tiny roundworms). One phoronid species is unpalatable to many epibenthic predators. Various parasites infest phoronids' body cavities, digestive tract and tentacles. It is unknown whether phoronids have any significance for humans. The International Union for Conservation of Nature (IUCN) has not listed any phoronid species as endangered. As of 2010 there are no indisputable body fossils of phoronids. There is good evidence that phoronids created trace fossils found in the
Silurian The Silurian ( ) is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the shortest period of the Paleozo ...
,
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
, Permian, Jurassic and Cretaceous periods, and possibly in the Ordovician and Triassic. Phoronids, brachiopods and bryozoans (ectoprocts) have collectively been called
lophophorate The Lophophorata are a Lophotrochozoan clade consisting of the Brachiozoa and the Bryozoa. They have a lophophore. Molecular phylogenetic analyses suggest that lophophorates are protostomes Protostomia () is the clade of animals once though ...
s, because all use lophophores to feed. From about the 1940s to the 1990s, family trees based on embryological and morphological features placed lophophorates among or as a
sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...
to the
deuterostome Deuterostomia (; in Greek) are animals typically characterized by their anus forming before their mouth during embryonic development. The group's sister clade is Protostomia, animals whose digestive tract development is more varied. Some exampl ...
s, a super-phylum which includes chordates and echinoderms. While a minority adhere to this view, most researchers now regard phoronids as members of the protostome super-phylum
Lophotrochozoa Lophotrochozoa (, "crest/wheel animals") is a clade of protostome animals within the Spiralia. The taxon was established as a monophyletic group based on molecular evidence. The clade includes animals like annelids, molluscs, bryozoans, brachi ...
. Although analysts using molecular phylogeny are confident that members of Lophotrochozoa are more closely related to each other than of non-members, the relationships between members are mostly unclear. Some analyses regard phoronids and brachiopods as sister-groups, while others place phoronids as a sub-group within brachiopoda.


Comparison of similar phyla


Description


Body structure

Most adult phoronids are 2 to 20 cm long and about 1.5 mm wide, although the largest are 50 cm long. Their skins have no
cuticle A cuticle (), or cuticula, is any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticle" are non- homologous, differing in their origin, structu ...
but secrete rigid tubes of
chitin Chitin ( C8 H13 O5 N)n ( ) is a long-chain polymer of ''N''-acetylglucosamine, an amide derivative of glucose. Chitin is probably the second most abundant polysaccharide in nature (behind only cellulose); an estimated 1 billion tons of chit ...
, similar to the material used in arthropods' exoskeletons, and sometimes reinforced with sediment particles and other debris. Most species' tubes are erect, but those of ''Phoronis vancouverensis'' are horizontal and tangled. Phoronids can move within their tubes but never leave them. The bottom end of the body is an ampulla (a flask-like swelling in a tube-like structure), which anchors the animal in the tube and enables it to retract its body when threatened, reducing the body to 20 percent of its maximum length. Longitudinal muscles retract the body very quickly, while circular muscles slowly extend the body by compressing the internal fluid. For feeding and respiration each phoronid has at the top end a lophophore, a "crown" of tentacles with which the animal filter-feeds. In small species the "crown" is a simple circle, in medium-size species it is bent into the shape of a
horseshoe A horseshoe is a fabricated product designed to protect a horse hoof from wear. Shoes are attached on the palmar surface (ground side) of the hooves, usually nailed through the insensitive hoof wall that is anatomically akin to the human toen ...
with tentacles on the outer and inner sides, and in the largest species the ends of the horseshoe wind into complex spirals. These more elaborate shapes increase the area available for feeding and respiration. The tentacles are hollow, held upright by fluid pressure, and can be moved individually by muscles. The mouth is inside the base of the crown of tentacles but to one side. The gut runs from the mouth to one side of the stomach, in the bottom of the ampulla. The intestine runs from the stomach, up the other side of the body, and exits at the anus, outside and a little below the crown of tentacles. The gut and intestine are both supported by two mesenteries (partitions that run the length of the body) connected to the body wall, and another mesentery connects the gut to the intestine. The body is divided into
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 ...
s, compartments lined with
mesothelium The mesothelium is a membrane composed of simple squamous epithelial cells of mesodermal origin, which forms the lining of several body cavities: the pleura (pleural cavity around the lungs), peritoneum (abdominopelvic cavity including the mesente ...
. The main body cavity, under the crown of tentacles, is called the metacoelom, and the tentacles and their base share the mesocoelom. Above the mouth is the epistome, a hollow lid which can close the mouth. The cavity in the epistome is sometimes called the protocoelom, although other authors disagree that it is a coelom and Ruppert, Fox and Barnes think it is built by a different process. The tube comprises a three-layered organic inner cylinder, and an agglutinated external layer.


Feeding, circulation and excretion

When the lophophore is extended,
cilia The cilium, plural cilia (), is a membrane-bound organelle found on most types of eukaryotic cell, and certain microorganisms known as ciliates. Cilia are absent in bacteria and archaea. The cilium has the shape of a slender threadlike projecti ...
(little hairs) on the sides of the tentacles draw water down between the tentacles and out at the base of the lophophore. Shorter cilia on the inner sides of the tentacles flick food particles into a groove in a circle under and just inside the tentacles, and cilia in the groove push the particles into the mouth. Phoronids direct their lophophores into the water current, and quickly reorient to maximize the food-catching area when currents change. Their diet includes
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...
,
diatom A diatom (Neo-Latin ''diatoma''), "a cutting through, a severance", from el, διάτομος, diátomos, "cut in half, divided equally" from el, διατέμνω, diatémno, "to cut in twain". is any member of a large group comprising sev ...
s,
flagellate A flagellate is a cell or organism with one or more whip-like appendages called flagella. The word ''flagellate'' also describes a particular construction (or level of organization) characteristic of many prokaryotes and eukaryotes and their ...
s,
peridinian Peridiniaceae is a family of dinoflagellates belonging to the order Peridiniales. Genera Peridiniaceae contains the following genera: * '' Alterbia'' * '' Alterbidinium'' * '' Amphidiadema'' * '' Andalusiela'' * '' Apectodinium'' * '' Arch ...
s, small invertebrate larvae, and detritus. Unwanted material can be excluded by closing the epistome (lid above the mouth) or be rejected by the tentacles, whose cilia can switch into reverse. The gut uses cilia and muscles to move food towards the stomach and secretes enzymes that digest some of the food, but the stomach digests the majority of the food. Phoronids also absorb amino acids (the building blocks of proteins) through their skins, mainly in summer. Solid wastes are moved up the intestine and out through the
anus The anus (Latin, 'ring' or 'circle') is an opening at the opposite end of an animal's digestive tract from the mouth. Its function is to control the expulsion of feces, the residual semi-solid waste that remains after food digestion, which, d ...
, which is outside and slightly below the lophophore. A blood vessel starts from the peritoneum (the membrane that loosely encloses the stomach), with blind capillaries supplying the stomach. The blood vessel leads up the middle of the body to a circular vessel at the base of the lophophore, and from there a single blind vessel runs up each tentacle. A pair of blood vessels near the body wall lead downward from the lophophore ring, and in most species these are combined into one a little below the lophophore ring. The downward vessel(s) leads back to the peritoneum, and also to blind branches throughout the body. There is no heart, but muscles in the major vessels contract in waves to move the blood. Unlike many animals that live in tubes, phoronids do not ventilate their trunks with oxygenated water, but rely on respiration by the lophophore, which extends above hypoxic sediments. The blood has hemocytes containing hemoglobin, which unusual in such small animals and seems to be an adaptation to anoxic and hypoxic environments. The blood of ''Phoronis architecta'' carries as much oxygen per cm3 as that of most vertebrates; the blood's volume in cm3 per gm of body weight is twice that of a human. Podocytes on the walls of the blood vessels perform first-stage filtration of soluble wastes into the main coelom's fluid. Two metanephridia, each with a funnel-like intake, filter the fluid a second time, returning any useful products to the coelom and dumping the remaining wastes through a pair of nephridiopores beside the anus.


Nervous system and movement

There is a nervous center between the mouth and anus, and a nerve ring at the base of the lophophore. The ring supplies nerves to the tentacles and, just under the skin, to the body-wall muscles. ''
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 ...
'' has two nerve trunks under the skin, whereas other species have one. The trunk(s) have giant axons (nerves that transmit signals very fast) which co-ordinate the retraction of the body when danger threatens. Except for retracting the body into the tube, phoronids have limited and slow movement: partial emerging from the tube; bending the body when extended; and the lophophore's flicking of food into the mouth.


Reproduction and lifecycle

Only ''
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 ...
'' naturally builds colonies by budding or by splitting into top and bottom sections which then grow into full bodies. In experiments, other species have split successfully, but only when both parts have enough
gonad A gonad, sex gland, or reproductive gland is a mixed gland that produces the gametes and sex hormones of an organism. Female reproductive cells are egg cells, and male reproductive cells are sperm. The male gonad, the testicle, produces sper ...
al (reproductive) tissue. All phoronids breed sexually from spring to autumn. Some species are hermaphroditic (have both male and female reproductive organs) but cross-fertilize (fertilize the eggs of other members), while others are
dioecious Dioecy (; ; adj. dioecious , ) is a characteristic of a species, meaning that it has distinct individual organisms (unisexual) that produce male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproductio ...
(have separate sexes). The gametes (
sperm Sperm is the male reproductive cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm with a tail known as a flagellum, whi ...
s and ova) are produced in the swollen gonads, around the stomach. The gametes swim through the metacoelom to the metanephridia. Sperm exit by the nephridiopores and some are captured by the lophophores of individuals of the same species. Species that lay small fertilized eggs release them into the water as plankton, while species with larger eggs brood them either in the body's tube or stuck in the center of the lophophore by adhesive. The brooded eggs are released to feed on plankton when they develop into larvae. Development of the eggs is a mixture of
deuterostome Deuterostomia (; in Greek) are animals typically characterized by their anus forming before their mouth during embryonic development. The group's sister clade is Protostomia, animals whose digestive tract development is more varied. Some exampl ...
and protostome characteristics. Early divisions of the egg are holoblastic (the cells divide completely) and radial (they gradually form a stack of circles). The process is regulative (the fate of each cell depends on interaction with other cells, not on a rigid program in each cell), and experiments that divided early embryos produced complete larvae.
Mesoderm The mesoderm is the middle layer of the three germ layers that develops during gastrulation in the very early development of the embryo of most animals. The outer layer is the ectoderm, and the inner layer is the endoderm.Langman's Medical E ...
is formed from
mesenchyme Mesenchyme () is a type of loosely organized animal embryonic connective tissue of undifferentiated cells that give rise to most tissues, such as skin, blood or bone. The interactions between mesenchyme and epithelium help to form nearly every o ...
originating from the archenteron. The coelom is formed by
schizocoely Schizocoely (adjective forms: schizocoelous or schizocoelic) is a process by which some animal embryos develop. The schizocoely mechanism occurs when secondary body cavities (coeloms) are formed by splitting a solid mass of mesodermal embryonic ti ...
, and the
blastopore Gastrulation is the stage in the early embryonic development of most animals, during which the blastula (a single-layered hollow sphere of cells), or in mammals the blastocyst is reorganized into a multilayered structure known as the gastrula. Be ...
(a dent in the embryo) becomes the mouth. The slug-like larva of ''Phoronis ovalis'' swims for about 4 days, creeps on the seabed for 3 to 4 days, then bores into a carbonate floor. Nothing is known about three species. The remaining species develop free-swimming actinotroch larvae, which feed on plankton. The actinotroch is an upright cylinder with the anus at the bottom and fringed with cilia. At the top is a lobe or hood, under which are: a ganglion, connected to a patch of cilia outside the apex of the hood; a pair of protonephridia (smaller and simpler than the metanephridia in the adult); the mouth; and feeding tentacles that encircle the mouth. After swimming for about 20 days, the actinotroch settles on the seabed and undergoes a catastrophic metamorphosis (radical change) in 30 minutes: the hood and larval tentacles are absorbed and the adult lophophore is created around the mouth, and both now point upward; the gut develops a U-bend so that the anus is just under and outside the lophophore. Finally the adult phoronid builds a tube. Phoronids live for about one year.


Ecology

Phoronids live in all the oceans and seas including the Arctic and excepting the Antarctic Ocean, and appear between the intertidal zone and about 400 meters down. Some occur separately, in vertical tubes embedded in soft sediment such as sand, mud, or fine gravel. Others form tangled masses of many individuals buried in or encrusting rocks and shells. In some habitats populations of phoronids reach tens of thousand of individuals per square meter. The actinotroch larvae are familiar among plankton, and sometimes account for a significant proportion of the zooplankton biomass. ''
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 ...
'' bores into the wall of the tube of a cerianthid anemone, ''Ceriantheomorphe brasiliensis'', and uses this as a foundation for building its own tube. One cerianthid can house up to 100 phoronids. In this unequal relationship, the anemone experiences no significant benefits nor harm, while the phoronid benefits from: a foundation for its tube; food (both animals are filter-feeders); and protection, as the cerianthid withdraws into its tube when danger threatens, and this alerts the phoronid to retract into its own tube. Although predators of phoronids are not well known, they include fish,
gastropod The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. T ...
s (snails), and
nematode The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-Parasitism, parasitic nematodes also known as eelworms. They are a diverse animal phylum inhab ...
s (tiny roundworms). ''Phoronopsis viridis'', which reaches densities of 26,500 per square meter on tidal flats in California (USA), is unpalatable to many epibenthic predators, including fish and crabs. The unpalatability is strongest in the top section, including the lophophore, which is exposed to predators when phoronids feed. When the lophophores were removed in an experiment, the phoronids were more palatable, but this effect reduced over 12 days as the lophophores regenerated. These broadly effective defenses, which appear unusual among invertebrates inhabiting soft sediment, may be important in allowing ''Phoronopsis viridis'' to reach high densities. Some parasites infest phoronids: progenetic metacercariae and
cysts A cyst is a closed sac, having a distinct envelope and division compared with the nearby tissue. Hence, it is a cluster of cells that have grouped together to form a sac (like the manner in which water molecules group together to form a bubble) ...
of trematodes in phoronids'
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 ...
ic cavities; unidentified
gregarine The gregarines are a group of Apicomplexan alveolates, classified as the Gregarinasina or Gregarinia. The large (roughly half a millimeter) parasites inhabit the intestines of many invertebrates. They are not found in any vertebrates. However, gr ...
s in phoronids' digestive tract; and an ancistrocomid ciliate parasite, '' Heterocineta'', in the tentacles. It is unknown whether phoronids have any significance for humans. The International Union for Conservation of Nature (IUCN) has not listed any phoronid species as endangered.


Evolutionary history


Fossil record

As of 2016 there are no indisputable body fossils of phoronids. Researching the Lower
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 ...
Chengjiang fossils, in 1997 Chen and Zhou interpreted '' Iotuba chengjiangensis'' as a phoronid since it had tentacles and a U-shaped gut, – cited by Emig (Mar 2010) and Taylor, Vinn and Wilson(2010) and in 2004 Chen interpreted ''
Eophoronis ''Iotuba chengjiangensis'' (sometimes misspelt ''Lotuba'') is a Cambrian fossil with a U-shaped gut and tentacles, known from the Chengjiang biota The Maotianshan Shales are a series of Early Cambrian deposits in the Chiungchussu Formation, ...
'' as a phoronid. However, in 2006 Conway Morris regarded ''Iotuba'' and ''Eophoronis'' as synonyms for the same genus, which in his opinion looked like the priapulid '' Louisella''. In 2009 Balthasar and Butterfield found in western Canada two specimens from about 505 million years ago of a new fossil, ''Lingulosacculus nuda'', which had two shells like those of brachiopods but not mineralized. In the authors' opinion, the U-shaped gut extended beyond the hinge line and outside the smaller shell. This would have precluded the attachment of muscles to close and open the shells, and the 50% of the animal's length beyond the hinge line would have needed longitudinal muscles and also a
cuticle A cuticle (), or cuticula, is any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticle" are non- homologous, differing in their origin, structu ...
for protection. Hence they suggest that ''Lingulosacculus'' may have been a member of a phoronid stem group within the linguliform brachiopods. Another alternative is that ''
Eccentrotheca ''Eccentrotheca'' is a genus of "tommotiid" known from Cambrian deposits. Its sclerites form rings that are stacked to produce a widening-upwards conical scleritome. Individual plates have been homologized with the valves of brachiopods, and a r ...
'' lies somewhere in the phoronid stem lineage. There is good evidence that species of ''Phoronis'' created the trace fossils of the ichnogenus ''Talpina'', which have been found in the
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
, Jurassic and Cretaceous periods. The Talpina animal bored into calcareous
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...
, corals, echinoid
tests Test(s), testing, or TEST may refer to: * Test (assessment), an educational assessment intended to measure the respondents' knowledge or other abilities Arts and entertainment * ''Test'' (2013 film), an American film * ''Test'' (2014 film), ...
(shells), mollusc shells and the rostra of belemnites.
Hederellid Hederellids are extinct colonial animals with calcitic tubular branching exoskeletons. They range from the Silurian to the Permian and were most common in the Devonian period. They are more properly known as "hederelloids" because they were origin ...
s or Hederelloids are fossilized tubes, usually curved and between 0.1 and 1.8 mm wide, found from the
Silurian The Silurian ( ) is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the shortest period of the Paleozo ...
to the Permian, and possibly in the Ordovician and Triassic. Their branching colonies may have been made by phoronids.


Family tree

Phoronids, brachiopods and bryozoans (ectoprocts) are collectively called
lophophorate The Lophophorata are a Lophotrochozoan clade consisting of the Brachiozoa and the Bryozoa. They have a lophophore. Molecular phylogenetic analyses suggest that lophophorates are protostomes Protostomia () is the clade of animals once though ...
s, because all feed using lophophores. From about the 1940s to the 1990s, family trees based on embryological and morphological features placed lophophorates among or as a
sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...
to the
deuterostome Deuterostomia (; in Greek) are animals typically characterized by their anus forming before their mouth during embryonic development. The group's sister clade is Protostomia, animals whose digestive tract development is more varied. Some exampl ...
s, a super-phylum that includes chordates and echinoderms. In the early development of their embryos, deuterostomes form the
anus The anus (Latin, 'ring' or 'circle') is an opening at the opposite end of an animal's digestive tract from the mouth. Its function is to control the expulsion of feces, the residual semi-solid waste that remains after food digestion, which, d ...
before the mouth, while protostomes form the mouth first. Nielsen (2002) views the phoronids and brachiopods as affiliated with the deuterostome
pterobranchs Pterobranchia is a class of small worm-shaped animals. They belong to the Hemichordata, and live in secreted tubes on the ocean floor. Pterobranchia feed by filtering plankton out of the water with the help of cilia attached to tentacles. The ...
, which also filter-feed by tentacles, because the current-driving cells of the lophophores of all three have one cilium per cell, while lophophores of bryozoans, which he regards as protostomes, have multiple cilia per cell. Helmkampf, Bruchhaus and Hausdorf (2008) summarise several authors' embryological and morphological analyses which doubt or disagree that phoronids and brachiopods are deuterostomes: * While deuterostomes have three
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 ...
ic cavities, lophophorates such as phoronids and brachiopods have only two. * Pterobranchs may be a sub-group of
enteropneust The acorn worms or Enteropneusta are a hemichordate class of invertebrates consisting of one order of the same name. The closest non-hemichordate relatives of the Enteropneusta are the echinoderms. There are 111 known species of acorn worm in the ...
s ("acorn worms"). This suggests that the ancestral deuterostome looks more like a mobile worm-like enteropneust than a sessile colonial pterobranch. The fact that lophophorates and pterobranchs both use tentacles for feeding is probably not a synapomorphy of lophophorates and deuterostomes, but evolved independently as convergent adaptations to a sessile lifestyle. * The mesoderm does not form by enterocoely in phoronids and bryozoans, but does in deuterostomes, while there are disagreements about whether brachiopods form the mesoderm by enterocoely.
Relationships of Phoronida to other
Bilateria The Bilateria or bilaterians are animals with bilateral symmetry as an embryo, i.e. having a left and a right side that are mirror images of each other. This also means they have a head and a tail (anterior-posterior axis) as well as a belly and ...
:
From 1988 onwards analyses based on molecular phylogeny, which compares
biochemical Biochemistry or biological chemistry is the study of chemical processes within and relating to living organisms. A sub-discipline of both chemistry and biology, biochemistry may be divided into three fields: structural biology, enzymology an ...
features such as similarities in DNA, have placed phoronids and brachiopods among the
Lophotrochozoa Lophotrochozoa (, "crest/wheel animals") is a clade of protostome animals within the Spiralia. The taxon was established as a monophyletic group based on molecular evidence. The clade includes animals like annelids, molluscs, bryozoans, brachi ...
, a protostome super-phylum that includes
mollusc Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is esti ...
s,
annelid The annelids (Annelida , from Latin ', "little ring"), also known as the segmented worms, are a large phylum, with over 22,000 extant species including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecol ...
s and
flatworm The flatworms, flat worms, Platyhelminthes, or platyhelminths (from the Greek πλατύ, ''platy'', meaning "flat" and ἕλμινς (root: ἑλμινθ-), ''helminth-'', meaning "worm") are a phylum of relatively simple bilaterian, unsegment ...
s but excludes the other main protostome super-phylum
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 ...
, whose members include arthropods. Cohen wrote, "This inference, if true, undermines virtually all morphology–based reconstructions of phylogeny made during the past century or more." While analyses by molecular phylogeny are confident that members of Lophotrochozoa are more closely related to each other than of non-members, the relationships between members are mostly unclear. The
Lophotrochozoa Lophotrochozoa (, "crest/wheel animals") is a clade of protostome animals within the Spiralia. The taxon was established as a monophyletic group based on molecular evidence. The clade includes animals like annelids, molluscs, bryozoans, brachi ...
are generally divided into: Lophophorata (animals that have lophophores), including Phoronida and Brachiopoda; Trochozoa (animals many of which have trochophore larvae), including molluscs,
annelid The annelids (Annelida , from Latin ', "little ring"), also known as the segmented worms, are a large phylum, with over 22,000 extant species including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecol ...
s, echiurans, sipunculans and nemerteans; and some other phyla (such as Platyhelminthes, Gastrotricha, Gnathostomulida, Micrognathozoa, and Rotifera). Molecular phylogeny indicates that Phoronida are closely related to Brachiopoda, but Bryozoa (Ectoprocta) are not closely related to this group, despite using a similar lophophore for feeding and respiration. This implies that the traditional definition "Lophophorata" is not
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
. Recently the term "Lophophorata" has been applied only to the Phoronida and Brachiopoda, and Halanych thinks this change will cause confusion. Some analyses regard Phoronida and Brachiopoda as sister-groups, while others place Phoronida as a sub-group within Brachiopoda, implying that Brachiopoda 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 ...
. Cohen and Weydman's analysis (2005) concludes that phoronids are a sub-group of inarticulate brachiopods (those in which the hinge between the two valves have no teeth and sockets) and sister-group of the other inarticulate sub-groups. The authors also suggest that the ancestors of
mollusc Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is esti ...
s and the brachiopod+phoronid clade diverged between 900 Ma and 560 Ma, most probably about 685 Ma.


Taxonomy

The phylum has 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 ...
, with no
class Class or The Class may refer to: Common uses not otherwise categorized * Class (biology), a taxonomic rank * Class (knowledge representation), a collection of individuals or objects * Class (philosophy), an analytical concept used differentl ...
or
order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...
names. Zoologists have given the larvae, usually called an actinotroch, a separate genus name from the adults. In 1999 Temereva and Malakhov described ''
Phoronis svetlanae ''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 ...
''. In 2000 Temereva described a new species, ''
Phoronopsis malakhovi ''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 f ...
'', while Emig regards it as a synonym for ''
Phoronopsis harmeri ''Phoronopsis harmeri'' is a species of marine horseshoe worm in the phylum Phoronida. It was first described by H.L.M. Pixell in 1912, and was found off of Vancouver Island. Ecology This species has been found around the world in coastal hab ...
''. Santagata thinks ''
Phoronis architecta ''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 ...
'' is a different species from both ''
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 ...
'' and ''
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 ...
'', and that "
he phoronids' He or HE may refer to: Language * He (pronoun), an English pronoun * He (kana), the romanization of the Japanese kana へ * He (letter), the fifth letter of many Semitic alphabets * He (Cyrillic), a letter of the Cyrillic script called ''He'' ...
species diversity is currently underestimated". In 2009 Temereva described what may be larvae of ''
Phoronopsis albomaculata ''Phoronopsis albomaculata'' 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 "c ...
'' and ''
Phoronopsis californica ''Phoronopsis californica'' is a species of marine horseshoe worm in the phylum Phoronida. It was first described as a new species by William Hilton in 1930 when he found it at Balboa Bay in Newport Beach, California.Hilton, W. A. (1930)A new ' ...
''. She wrote that, while there are 12 undisputed adult phoronid species, 25 morphological types of larvae have been identified.


Notes


References


External links


PHORONIDAPhoronida World database
ttps://web.archive.org/web/20080930064242/http://www.tafi.org.au/ Tasmanian Aquaculture & Fisheries Institutebr>Bioerosion website at The College of Wooster
{{Good article * Taxa named by Berthold Hatschek