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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 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.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 The Southern Ocean, also known as the Antarctic Ocean, comprises the southernmost waters of the World Ocean, generally taken to be south of 60° S latitude and encircling Antarctica. With a size of , it is regarded as the second-small ...
, and between the
intertidal The intertidal zone, also known as the foreshore, is the area above water level at low tide and underwater at high tide (in other words, the area within the tidal range). This area can include several types of Marine habitat, habitats with var ...
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 In biological taxonomy, the type genus is the genus which defines a biological family and the root of the family name. Zoological nomenclature According to the International Code of Zoological Nomenclature, "The name-bearing type of a nominal f ...
: ''
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 ...
''.


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 Respiration may refer to: Biology * Cellular respiration, the process in which nutrients are converted into useful energy in a cell ** Anaerobic respiration, cellular respiration without oxygen ** Maintenance respiration, the amount of cellul ...
through the lophophore. The blood contains
hemoglobin Hemoglobin (haemoglobin BrE) (from the Greek word αἷμα, ''haîma'' 'blood' + Latin ''globus'' 'ball, sphere' + ''-in'') (), abbreviated Hb or Hgb, is the iron-containing oxygen-transport metalloprotein present in red blood cells (erythrocyte ...
, 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 Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements as wel ...
as a human of the same weight. Two
metanephridia The nephridium (plural ''nephridia'') is an invertebrate organ, found in pairs and performing a function similar to the vertebrate kidneys (which originated from the chordate nephridia). Nephridia remove metabolic wastes from an animal's body. Ne ...
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 Budding or blastogenesis is a type of asexual reproduction in which a new organism develops from an outgrowth or bud due to cell division at one particular site. For example, the small bulb-like projection coming out from the yeast cell is know ...
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 Sediment is a naturally occurring material that is broken down by processes of weathering and erosion, and is subsequently transported by the action of wind, water, or ice or by the force of gravity acting on the particles. For example, sand an ...
, 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 Plankton are the diverse collection of organisms found in Hydrosphere, water (or atmosphere, air) that are unable to propel themselves against a Ocean current, current (or wind). The individual organisms constituting plankton are called plankt ...
, 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 Benthos (), also known as benthon, is the community of organisms that live on, in, or near the bottom of a sea, river, lake, or stream, also known as the benthic zone.parasite Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has ...
s 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 fossil A trace fossil, also known as an ichnofossil (; from el, ἴχνος ''ikhnos'' "trace, track"), is a fossil record of biological activity but not the preserved remains of the plant or animal itself. Trace fossils contrast with body fossils, ...
s 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 The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleoz ...
,
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
and
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...
periods, and possibly in the
Ordovician The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start ...
and
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
. Phoronids,
brachiopod Brachiopods (), phylum Brachiopoda, are a phylum of trochozoan animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear end, w ...
s and
bryozoa Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic invertebrate animals, nearly all living in sedentary colonies. Typically about long, they have a special feeding structure called a l ...
ns (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 A family tree, also called a genealogy or a pedigree chart, is a chart representing family relationships in a conventional tree structure. More detailed family trees, used in medicine and social work, are known as genograms. Representations of ...
based on
embryological Embryology (from Greek ἔμβρυον, ''embryon'', "the unborn, embryo"; and -λογία, ''-logia'') is the branch of animal biology that studies the prenatal development of gametes (sex cells), fertilization, and development of embryos and ...
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 deuterostomes, a super-phylum which includes
chordate A chordate () is an animal of the phylum Chordata (). All chordates possess, at some point during their larval or adult stages, five synapomorphies, or primary physical characteristics, that distinguish them from all the other taxa. These fiv ...
s and
echinoderm An echinoderm () is any member of the phylum Echinodermata (). The adults are recognisable by their (usually five-point) radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the sea ...
s. While a minority adhere to this view, most researchers now regard phoronids as members of the
protostome Protostomia () is the clade of animals once thought to be characterized by the formation of the organism's mouth before its anus during embryogenesis, embryonic development. This nature has since been discovered to be extremely variable among Pro ...
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 Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
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 440px Secretion is the movement of material from one point to another, such as a secreted chemical substance from a cell or gland. In contrast, excretion is the removal of certain substances or waste products from a cell or organism. The classical ...
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
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
s'
exoskeleton An exoskeleton (from Greek ''éxō'' "outer" and ''skeletós'' "skeleton") is an external skeleton that supports and protects an animal's body, in contrast to an internal skeleton (endoskeleton) in for example, a human. In usage, some of the ...
s, and sometimes reinforced with
sediment Sediment is a naturally occurring material that is broken down by processes of weathering and erosion, and is subsequently transported by the action of wind, water, or ice or by the force of gravity acting on the particles. For example, sand an ...
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 Respiration may refer to: Biology * Cellular respiration, the process in which nutrients are converted into useful energy in a cell ** Anaerobic respiration, cellular respiration without oxygen ** Maintenance respiration, the amount of cellul ...
each phoronid has at the top end a
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-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 Respiration may refer to: Biology * Cellular respiration, the process in which nutrients are converted into useful energy in a cell ** Anaerobic respiration, cellular respiration without oxygen ** Maintenance respiration, the amount of cellul ...
. 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 The stomach is a muscular, hollow organ in the gastrointestinal tract of humans and many other animals, including several invertebrates. The stomach has a dilated structure and functions as a vital organ in the digestive system. The stomach i ...
, 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 In zoology, a mesentery is a membrane inside the body cavity of an animal. The term identifies different structures in different phyla: in vertebrates it is a double fold of the peritoneum enclosing the intestines; in other organisms it forms com ...
(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, peridinians, 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
enzyme Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as products. A ...
s that digest some of the food, but the stomach digests the majority of the food. Phoronids also absorb
amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha am ...
s (the building blocks of
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...
s) 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 peritoneum is the serous membrane forming the lining of the abdominal cavity or coelom in amniotes and some invertebrates, such as annelids. It covers most of the intra-abdominal (or coelomic) organs, and is composed of a layer of mesoth ...
(the
membrane A membrane is a selective barrier; it allows some things to pass through but stops others. Such things may be molecules, ions, or other small particles. Membranes can be generally classified into synthetic membranes and biological membranes. B ...
that loosely encloses the stomach), with blind
capillaries A capillary is a small blood vessel from 5 to 10 micrometres (μm) in diameter. Capillaries are composed of only the tunica intima, consisting of a thin wall of simple squamous endothelial cells. They are the smallest blood vessels in the body: ...
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 Respiration may refer to: Biology * Cellular respiration, the process in which nutrients are converted into useful energy in a cell ** Anaerobic respiration, cellular respiration without oxygen ** Maintenance respiration, the amount of cellul ...
by the lophophore, which extends above
hypoxic Hypoxia means a lower than normal level of oxygen, and may refer to: Reduced or insufficient oxygen * Hypoxia (environmental), abnormally low oxygen content of the specific environment * Hypoxia (medical), abnormally low level of oxygen in the t ...
sediments. The blood has hemocytes containing
hemoglobin Hemoglobin (haemoglobin BrE) (from the Greek word αἷμα, ''haîma'' 'blood' + Latin ''globus'' 'ball, sphere' + ''-in'') (), abbreviated Hb or Hgb, is the iron-containing oxygen-transport metalloprotein present in red blood cells (erythrocyte ...
, 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
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
s; the blood's volume in cm3 per gm of body weight is twice that of a human.
Podocyte Podocytes are cells in Bowman's capsule in the kidneys that wrap around capillaries of the glomerulus. Podocytes make up the epithelial lining of Bowman's capsule, the third layer through which filtration of blood takes place. Bowman's capsule ...
s on the walls of the blood vessels perform first-stage filtration of soluble wastes into the main coelom's fluid. Two
metanephridia The nephridium (plural ''nephridia'') is an invertebrate organ, found in pairs and performing a function similar to the vertebrate kidneys (which originated from the chordate nephridia). Nephridia remove metabolic wastes from an animal's body. Ne ...
, 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
nephridiopore A nephridiopore is part of the nephridium, an excretory organ found in many organisms, such as flatworms and annelids. Polychaetes typically release their gametes into the water column using nephridiopores. Nephridia are homologous to nephrons o ...
s 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'' 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'' naturally builds colonies by
budding Budding or blastogenesis is a type of asexual reproduction in which a new organism develops from an outgrowth or bud due to cell division at one particular site. For example, the small bulb-like projection coming out from the yeast cell is know ...
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 In reproductive biology, a hermaphrodite () is an organism that has both kinds of reproductive organs and can produce both gametes associated with male and female sexes. Many taxonomic groups of animals (mostly invertebrates) do not have separ ...
(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
gamete A gamete (; , ultimately ) is a haploid cell that fuses with another haploid cell during fertilization in organisms that reproduce sexually. Gametes are an organism's reproductive cells, also referred to as sex cells. In species that produce t ...
s (
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 , abbreviated as OVA and sometimes as OAV (original animation video), are Japanese animated films and series made specially for release in home video formats without prior showings on television or in theaters, though the first part of an OVA s ...
) 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 and
protostome Protostomia () is the clade of animals once thought to be characterized by the formation of the organism's mouth before its anus during embryogenesis, embryonic development. This nature has since been discovered to be extremely variable among Pro ...
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 primary gut that forms during gastrulation in a developing embryo is known as the archenteron, the gastrocoel or the primitive digestive tube. It develops into the endoderm and mesoderm of an animal. Formation in sea urchins As primary mesen ...
. 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 ...
, and the blastopore (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 The nephridium (plural ''nephridia'') is an invertebrate organ, found in pairs and performing a function similar to the vertebrate kidneys (which originated from the chordate nephridia). Nephridia remove metabolic wastes from an animal's body. Neph ...
(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 The Arctic ( or ) is a polar regions of Earth, polar region located at the northernmost part of Earth. The Arctic consists of the Arctic Ocean, adjacent seas, and parts of Canada (Yukon, Northwest Territories, Nunavut), Danish Realm (Greenla ...
and excepting the
Antarctic Ocean The Southern Ocean, also known as the Antarctic Ocean, comprises the southernmost waters of the World Ocean, generally taken to be south of 60° S latitude and encircling Antarctica. With a size of , it is regarded as the second-small ...
, and appear between the
intertidal The intertidal zone, also known as the foreshore, is the area above water level at low tide and underwater at high tide (in other words, the area within the tidal range). This area can include several types of Marine habitat, habitats with var ...
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 Plankton are the diverse collection of organisms found in Hydrosphere, water (or atmosphere, air) that are unable to propel themselves against a Ocean current, current (or wind). The individual organisms constituting plankton are called plankt ...
, and sometimes account for a significant proportion of the zooplankton biomass. '' Phoronis australis'' bores into the wall of the tube of a cerianthid
anemone ''Anemone'' () is a genus of flowering plants in the buttercup family Ranunculaceae. Plants of the genus are commonly called windflowers. They are native to the temperate and subtropical regions of all continents except Australia, New Zealand an ...
, ''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 California is a U.S. state, state in the Western United States, located along the West Coast of the United States, Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the List of states and territori ...
(USA), is unpalatable to many
epibenthic Benthos (), also known as benthon, is the community of organisms that live on, in, or near the bottom of a sea, river, lake, or stream, also known as the benthic zone.parasite Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has ...
s infest phoronids: progenetic
metacercariae Trematodes are parasitic flatworms of the class ''Trematoda'', specifically parasitic flukes with two suckers: one ventral and the other oral. Trematodes are covered by a tegument, that protects the organism from the environment by providing secr ...
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
trematode Trematoda is a class of flatworms known as flukes. They are obligate internal parasites with a complex life cycle requiring at least two hosts. The intermediate host, in which asexual reproduction occurs, is usually a snail. The definitive host ...
s 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 The ciliates are a group of alveolates characterized by the presence of hair-like organelles called cilia, which are identical in structure to flagellum, eukaryotic flagella, but are in general shorter and present in much larger numbers, with a ...
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 ''Iotuba chengjiangensis'' (sometimes misspelt ''Lotuba'') is a Cambrian fossil with a U-shaped gut and tentacles, known from the Chengjiang biota. Originally interpreted as a phoronid marine animal, its affinity is currently disputed. It, and ...
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 A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bon ...
'' 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 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 ...
''
Louisella ''Louisella'' is a genus of worm known from the Middle Cambrian Burgess Shale. It was originally described by Charles Walcott in 1911 as a holothurian echinoderm, and represents a senior synonym of ''Miskoia'', which was originally described as ...
''. 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
brachiopod Brachiopods (), phylum Brachiopoda, are a phylum of trochozoan animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear end, w ...
s 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 In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...
within the
linguliform Linguliformea is a subphylum of inarticulate brachiopods. These were the earliest of brachiopods, ranging from the Cambrian into the Holocene. They rapidly diversified during the Cambrian into the Ordovician, but most families became extinct by ...
brachiopods. Another alternative is that '' Eccentrotheca'' lies somewhere in the phoronid stem lineage. There is good evidence that species of ''Phoronis'' created the
trace fossil A trace fossil, also known as an ichnofossil (; from el, ἴχνος ''ikhnos'' "trace, track"), is a fossil record of biological activity but not the preserved remains of the plant or animal itself. Trace fossils contrast with body fossils, ...
s of the
ichnogenus An ichnotaxon (plural ichnotaxa) is "a taxon based on the fossilized work of an organism", i.e. the non-human equivalent of an artifact. ''Ichnotaxa'' comes from the Greek ίχνος, ''ichnos'' meaning ''track'' and ταξις, ''taxis'' meaning ...
''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 The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
and
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...
periods. The Talpina animal bored into
calcareous Calcareous () is an adjective meaning "mostly or partly composed of calcium carbonate", in other words, containing lime or being chalky. The term is used in a wide variety of scientific disciplines. In zoology ''Calcareous'' is used as an adje ...
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 ...
,
coral Corals are marine invertebrates within the class Anthozoa of the phylum Cnidaria. They typically form compact colonies of many identical individual polyps. Coral species include the important reef builders that inhabit tropical oceans and sec ...
s,
echinoid Sea urchins () are spine (zoology), spiny, globular echinoderms in the class Echinoidea. About 950 species of sea urchin live on the seabed of every ocean and inhabit every depth zone from the intertidal seashore down to . The spherical, hard s ...
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 The rostra ( it, Rostri, links=no) was a large platform built in the city of Rome that stood during the republican and imperial periods. Speakers would stand on the rostra and face the north side of the comitium towards the senate house and deli ...
of
belemnite Belemnitida (or the belemnite) is an extinct order of squid-like cephalopods that existed from the Late Triassic to Late Cretaceous. Unlike squid, belemnites had an internal skeleton that made up the cone. The parts are, from the arms-most ...
s. Hederellids 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 The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleoz ...
, and possibly in the
Ordovician The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start ...
and
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
. Their branching colonies may have been made by phoronids.


Family tree

Phoronids,
brachiopod Brachiopods (), phylum Brachiopoda, are a phylum of trochozoan animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear end, w ...
s and
bryozoa Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic invertebrate animals, nearly all living in sedentary colonies. Typically about long, they have a special feeding structure called a l ...
ns (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 A family tree, also called a genealogy or a pedigree chart, is a chart representing family relationships in a conventional tree structure. More detailed family trees, used in medicine and social work, are known as genograms. Representations of ...
based on
embryological Embryology (from Greek ἔμβρυον, ''embryon'', "the unborn, embryo"; and -λογία, ''-logia'') is the branch of animal biology that studies the prenatal development of gametes (sex cells), fertilization, and development of embryos and ...
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 deuterostomes, a super-phylum that includes
chordate A chordate () is an animal of the phylum Chordata (). All chordates possess, at some point during their larval or adult stages, five synapomorphies, or primary physical characteristics, that distinguish them from all the other taxa. These fiv ...
s and
echinoderm An echinoderm () is any member of the phylum Echinodermata (). The adults are recognisable by their (usually five-point) radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the sea ...
s. 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
protostome Protostomia () is the clade of animals once thought to be characterized by the formation of the organism's mouth before its anus during embryogenesis, embryonic development. This nature has since been discovered to be extremely variable among Pro ...
s 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 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 ...
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 enteropneusts ("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 In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...
of lophophorates and deuterostomes, but evolved independently as convergent adaptations to a sessile lifestyle. * The mesoderm does not form by
enterocoely Enterocoely (adjective forms: enterocoelic and enterocoelous) describes both the process by which some animal embryos develop and the origin of the cells involved. In enterocoely, a mesoderm (middle layer) is formed in a developing embryo, in whic ...
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 Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
, 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, whose members include
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
s. 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 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, but on morphological grounds they have been assessed ...
(animals that have lophophores), including Phoronida and Brachiopoda;
Trochozoa The Trochozoa are a proposed Lophotrochozoa clade that is a sister clade of Bryozoa. The trochozoans clade include animals in six phyla: the Nemertea, the Annelida, the Cycliophora, the Mollusca, and the two Brachiozoa Brachiozoa is a groupin ...
(animals many of which have
trochophore A trochophore (; also spelled trocophore) is a type of free-swimming planktonic marine larva with several bands of cilia. By moving their cilia rapidly, they make a water eddy, to control their movement, and to bring their food closer, to captur ...
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,
echiura The Echiura, or spoon worms, are a small group of marine animals. Once treated as a separate phylum, they are now considered to belong to Annelida. Annelids typically have their bodies divided into segments, but echiurans have secondarily los ...
ns,
sipuncula The Sipuncula or Sipunculida (common names sipunculid worms or peanut worms) is a class containing about 162 species of unsegmented marine annelid worms. The name ''Sipuncula'' is from the genus name ''Sipunculus'', and comes from the Latin ' ...
ns and
nemertea Nemertea is a phylum of animals also known as ribbon worms or proboscis worms, consisting of 1300 known species. Most ribbon worms are very slim, usually only a few millimeters wide, although a few have relatively short but wide bodies. Many h ...
ns; and some other phyla (such as
Platyhelminthes The flatworms, flat worms, Platyhelminthes, or platyhelminths (from the Greek language, Greek πλατύ, ''platy'', meaning "flat" and ἕλμινς (root: ἑλμινθ-), ''helminth-'', meaning "worm") are a Phylum (biology), phylum of relati ...
,
Gastrotricha The gastrotrichs (phylum Gastrotricha), commonly referred to as hairybellies or hairybacks, are a group of microscopic (0.06-3.0 mm), worm-like, acoelomate animals, and are widely distributed and abundant in freshwater and marine environmen ...
,
Gnathostomulida Gnathostomulids, or jaw worms, are a small phylum of nearly microscopic marine animals. They inhabit sand and mud beneath shallow coastal waters and can survive in relatively anoxic environments. They were first recognised and described in 1956. ...
,
Micrognathozoa ''Limnognathia maerski'' is a microscopic freshwater animal, discovered living in warm springs on Disko Island, Greenland, in 1994. Since then, it was also found in Crozet Islands of Antarctica.). With an average length of 100 micrometers ...
, and
Rotifera The rotifers (, from the Latin , "wheel", and , "bearing"), commonly called wheel animals or wheel animalcules, make up a phylum (Rotifera ) of microscopic and near-microscopic pseudocoelomate animals. They were first described by Rev. John Ha ...
). Molecular phylogeny indicates that Phoronida are closely related to Brachiopoda, but
Bryozoa Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic invertebrate animals, nearly all living in sedentary colonies. Typically about long, they have a special feeding structure called a l ...
(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 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''. 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'' is a different species from both '' Phoronis psammophila'' and '' Phoronis muelleri'', and that " he phoronids'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