Bryozoa
Encyclopedia
The Bryozoa, also known as Ectoprocta or commonly as moss animals, are a phylum
of aquatic
invertebrate
animal
s. Typically about 0.5 millimetre (0.0196850393700787 in) long, they are filter feeder
s that sieve food particles out of the water using a retractable lophophore
, a "crown" of tentacle
s lined with cilia. Most marine
species live in tropical waters, but a few occur in oceanic trench
es, and others are found in polar
waters. One class
lives only in a variety of freshwater
environments, and a few members of a mostly marine class prefer brackish water. Over 4,000 living species are known. One genus
is solitary and the rest colonial
.
The phylum was originally called "Polyzoa", but this term was superseded by "Bryozoa" in 1831. Another group of animals discovered subsequently, whose filtering mechanism looked similar, was also included in "Bryozoa" until 1869, when the two groups were noted to be very different internally. The more recently discovered group were given the name Entoprocta
, while the original "Bryozoa" were called "Ectoprocta". However, "Bryozoa" has remained the more widely used term for the latter group.
Individuals in bryozoan (ectoproct) colonies are called zooids, since they are not fully independent animals. All colonies contain autozooids, which are responsible for feeding and excretion
. Colonies of some classes
have various types of non-feeding specialist zooids, some of which are hatcheries for fertilized eggs, and some classes also have special zooids for defense of the colony. The class Cheilostomata
have the largest number of species, possibly because they have the widest range of specialist zooids. A few species can creep very slowly by using spiny defensive zooids as legs. Autozooids supply nutrients to non-feeding zooids by channels that vary between classes. All zooids, including those of the solitary species, consist of a cystid that provides the body wall and produces the exoskeleton
and a polypide that contains the internal organs and the lophophore or other specialist extensions. Zooids have no special excretory organs, and the polypides of autozooids are scrapped when the polypides become overloaded by waste products; usually the body wall then grows a replacement polypide. In autozooids the gut is U-shaped, with the mouth inside the "crown" of tentacles and the anus outside it. Colonies take a variety of forms, including fans, bushes and sheets. The Cheilostomata produce mineralized exoskeletons and form single-layered sheets that encrust over surfaces.
Zooids of all the freshwater species are simultaneous hermaphrodite
s. Although those of many marine species function first as males and then as females, their colonies always contain a combination of zooids that are in their male and female stages. All species emit sperm
into the water. Some also release ova
into the water, while others capture sperm via their tentacles to fertilize their ova internally. In some species the larva
e have large yolks, go to feed, and quickly settle on a surface. Others produce larvae that have little yolk but swim and feed for a few days before settling. After settling, all larvae undergo a radical metamorphosis
that destroys and rebuilds almost all the internal tissues. Freshwater species also produce statoblasts that lie dormant until conditions are favorable, which enables a colony's lineage to survive even if severe conditions kill the mother colony.
Predators of marine bryozoans include nudibranch
s (sea slugs), fish, sea urchin
s, pycnogonids, crustacean
s, mite
s and starfish. Freshwater bryozoans are preyed on by snails, insects, and fish. In Thailand
, many populations of one freshwater species have been wiped out by an introduced species
of snail. A fast-growing invasive bryozoan off the northeast and northwest coasts of the USA has reduced kelp
forests so much that it has affected local fish and invertebrate populations. Bryozoans have spread diseases to fish farms and fishermen. Chemicals extracted from a marine bryozoan species have been investigated for treatment of cancer
and Alzheimer's Disease
, but analyses have not been encouraging.
Mineralized skeletons of bryozoans first appear in rocks from latest Cambrian
period, about , making it the last major phylum to appear in the fossil record. This has led researchers to suspect that bryozoans had arisen earlier but were initially unmineralized, and may have differed significantly from fossilized and modern forms. Early fossils are mainly of erect forms, but encrusting forms gradually became dominant. It is uncertain whether the phylum is monophyletic. Bryozoans' evolutionary relationships to other phyla are also unclear, partly because scientists' view of the family tree of animals is mainly influenced by better-known phyla. Both morphological
and molecular phylogeny
analyses disagree over bryozoans' relationships with entoprocts, about whether bryozoans should be grouped with brachiopod
s and phoronid
s in Lophophorata, and whether bryozoans should be considered protostome
s or deuterostome
s.
Bryozoans, phoronids and brachiopods strain food out of the water by means of a lophophore
, a "crown" of hollow tentacles. Bryozoans form colonies consisting of clone
s called zooids that are typically about 0.5 millimetre (0.0196850393700787 in) long. Phoronids resemble bryozoan zooids but are 2 to 20 cm (0.78740157480315 to 7.9 in) long and, although they often grow in clumps, do not form colonies consisting of clones. Brachiopods, generally thought to be closely related to bryozoans and phoronids, are distinguished by having shells rather like those of bivalves. All three of these phyla
have a coelom
, an internal cavity lined by mesothelium
.
Some encrusting bryozoan colonies with mineralized exoskeleton
s look very like small corals. However, bryozoan colonies are founded by an ancestrula, which is round rather than shaped like a normal zooid of that species. On the other hand the founding polyp of a coral has a shape like that of its daughter polyps, and coral zooids have no coelom or lophophore.
Entoprocts
, another phylum of filter-feeders, look rather like bryozoans but their lophophore-like feeding structure has solid tentacles, their anus
lies inside rather than outside the base of the "crown" and they have no coelom
.
, Monobryozoon. Individual members of a bryozoan colony are about 0.5 millimetre (0.0196850393700787 in) long and are known as zooids, since they are not fully independent animals. All colonies contain feeding zooids, known as autozooids, and those of some groups also contain non-feeding specialist heterozooids; colony members are genetically identical and co-operate, rather like the organs of larger animals. What type of zooid grows where in a colony is determined by chemical signals from the colony as a whole or sometimes in response to the scent of predators or rival colonies.
The bodies of all types have two main parts. The cystid consists of the body wall and whatever type of exoskeleton
is secreted
by the epidermis
. The exoskeleton may be organic (chitin
, polysaccharide
or protein
) or made of the mineral calcium carbonate
. The body wall consists of the epidermis, basal lamina
(a mat of non-cellular material), connective tissue
, muscles, and the mesothelium
which lines the coelom
(main body cavity) – except that in one class
, the mesothelium is split into two separate layers, the inner one forming a membranous sac that floats freely and contains the coelom, and the outer one attached to the body wall and enclosing the membranous sac in a pseudocoelom. The other main part of the bryozoan body, known as the polypide and situated almost entirely within the cystid, contains the nervous system, digestive system, some specialized muscles and the feeding apparatus or other specialized organs that take the place of the feeding apparatus.
, which captures food particles from the water. In all colonies a large percentage of zooids are autozooids, and some consist entirely of autozooids, some of which also engage in reproduction.
The basic shape of the "crown" is a full circle. In the class
Phylactolaemata
the crown appears U-shaped, but this impression is created by a deep dent in the rim of the crown, which has no gap in the fringe of tentacles. The sides of the tentacles bear fine hairs called cilia, whose beating drives a water current from the tips of the tentacles to their bases, where it exits. Food particles that collide with the tentacles are trapped by mucus
, and further cilia on the inner surfaces of the tentacles convey the particles towards the mouth, which lies in the center of the base of the "crown". The method used by ectoprocts is known as "upstream collecting", as food particles are captured before they pass through the field of cilia that creates the feeding current. This method is also used by phoronid
s, brachiopod
s and pterobranchs.
The lophophore and mouth are mounted on a flexible tube, called the "invert" because it can be turned inside-out and withdrawn into the polypide, rather like the finger of a rubber glove; in this position the lophophore lies inside the invert and is folded like the spokes of an umbrella. The invert is withdrawn, sometimes within 60 millisecond
s, by a pair of retractor muscles that are anchored at the far end of the cystid. Sensors at the tips of the tentacles may check for signs of danger before the invert and lophophore are fully extended. Extension is driven by an increase in internal fluid pressure, which species with flexible exoskeletons produce by contracting circular muscles that lie just inside the body wall, while species with a membranous sac use circular muscles to squeeze this. Some species with rigid exoskeletons have a flexible membrane that replaces part of the exoskeleton, and transverse muscles anchored on the far side of the exoskeleton increase the fluid pressure by pulling the membrane inwards. In others there is no gap in the protective skeleton, and the transverse muscles pull on a flexible sac which is connected to the water outside by a small pore; the expansion of the sac increases the pressure inside the body and pushes the invert and lophophore out. In some species the retracted invert and lophophore are protected by an operculum ("lid"), which is closed by muscles and opened by fluid pressure. In one class
, a hollow lobe called the "epistome" overhands the mouth.
The gut is U-shaped, running from the mouth, in the center of the lophophore, down into the animal's interior and then back to the anus
, which is located on the invert, outside and usually below the lophophore. A network of strands of mesothelium
called "funiculi" ("little ropes") connects the mesothelium covering the gut with that lining the body wall. The wall of each strand is made of mesothelium, and surrounds a space filled with fluid, thought to be blood. A colony's zooids are connected, enabling autozooids to share food with each other and with any non-feeding heterozooids. The method of connection varies between the different classes of bryozoans, ranging from quite large gaps in the body walls to small pores through which nutrients are passed by funiculi.
There is a nerve ring round the pharynx (throat) and a ganglion
that serves as a brain to one side of this. Nerves run from the ring and ganglion to the tentacles and to the rest of the body. Bryozoans have no specialized sense organs, but cilia on the tentacles act as sensors. Members of the genus
Bugula
grow towards the sun
, and therefore must be able to detect light. In colonies of some species, signals are transmitted between zooids through nerves that pass through pores in the body walls, and coordinate activities such as feeding and the retraction of lophophores.
The solitary individuals of Monobryozoon are autozooids with pear-shaped bodies. The wider ends have up to 15 short, muscular projections by which the animals anchor themselves to sand or gravel and pull themselves through the sediments.
s of others. In some species the snapping zooids are mounted on a peduncle (stalk), their bird-like appearance responsible for the term – Charles Darwin
described these as like "the head and beak of a vulture in miniature, seated on a neck and capable of movement". Stalked avicularia are placed upside-down on their stalks. The "lower jaws" are modified versions of the opercula that protect the retracted lophophores in autozooids of some species, and are snapped shut "like a mousetrap" by similar muscles, while the beak-shaped upper jaw is the inverted body wall. In other species the avicularia are stationary box-like zooids laid the normal way up, so that the modified operculum snaps down against the body wall. In both types the modified operculum is opened by other muscles that attach to it, or by internal muscles that raise the fluid pressure by pulling on a flexible membrane. The actions of these snapping zooids are controlled by small, highly modified polypides that are located inside the "mouth" and bear tufts of short sensory cilia. These zooids appear in various positions: some take the place of autozooids, some fit into small gaps between autozooids, and small avicularia may occur on the surfaces of other zooids.
In vibracula, regarded by some as a type of avicularia, the operculum is modified to form a long bristle that has a wide range of motion. They may function as defenses against predators and invaders, or as cleaners. In some species that form mobile colonies, vibracula around the edges are used as legs for burrowing and walking.
Spinozooids form defensive spines, and sometimes appear on top of autozooids. Gonozooids act as brood chambers for fertilized eggs. Some species have miniature nanozooids with small single-tentacled polypides, and these may grow on other zooids or within the body walls of autozooids that have degenerated.
.
Some marine species are bush-like or fan-like, supported by "trunks" and "branches" formed by kenozooids, with feeding autozooids growing from these. Colonies of these types are generally unmineralized but may have exoskeleton
s made of chitin
. Others look like small coral
s, producing heavy lime skeletons. Many species form colonies which consist of sheets of autozooids. These sheets may form leaves, tufts or, in the genus
Thalmoporella, structures that resemble an open head of lettuce
.
The most common marine form, however, is encrusting, in which a one-layer sheet of zooids spreads over a hard surface or over seaweed. Some encrusting colonies may grow to over 50 centimetres (1.6 ft) and contain about 2,000,000 zooids. These species generally have exoskeletons reinforced with calcium carbonate
, and the openings through which the lophophores protrude are on the top or outer surface. The moss-like appearance of encrusting colonies is responsible for the phylum's alternative name Bryozoa (Ancient Greek
words βρυος bryos meaning "moss" and ζωον zoon meaning "animal"). Large colonies of encrusting species often have "chimney
s", gaps in the canopy of lophophores, through which they swiftly expel water that has been sieved, and thus avoid re-filtering water that is already exhausted. They are formed by patches of non-feeding heterozooids. New chimneys appear near the edges of expanding colonies, at points where the speed of the outflow is already high, and do not change position if the water flow changes.
Some freshwater species secrete a mass of gelatinous material, up to 1 metres (3.3 ft) in diameter, to which the zooids stick. Other freshwater species have plant-like shapes with "trunks" and "branches", which may stand erect or spread over the surface. A few species can creep at about 2 centimetre (0.78740157480315 in) per day.
Each colony grows by asexual
budding
from a single zooid known as the ancestrula, which is round rather than shaped like a normal zooid. This occurs at the tips of "trunks" or "branches" in forms that have this structure. Encrusting colonies grow round their edges. In species with calcareous
exoskeletons, these do not mineralize until the zooids are fully grown. Colony lifespans range from one to about 12 years, and the short-lived species pass though several generations in one season.
Species that produce defensive zooids do so only when threats have already appeared, and may do so within 48 hours. The theory of "induced defenses" suggests that production of defenses is expensive and that colonies which defend themselves too early or too heavily will have reduced growth rates and lifespans. This "last minute" approach to defense is feasible because the loss of zooids to a single attack is unlikely to be significant. Colonies of some encrusting species also produce special heterozooids to limit the expansion of other encrusting organisms, especially other bryozoans. In some cases this response is more belligerent if the opposition is smaller, which suggests that zooids on the edge of a colony can somehow sense the size of the opponent. Some species consistently prevail against certain others, but most turf war
s are indecisive and the combatants soon turn to growing in uncontested areas. Bryozoans competing for territory do not use the sophisticated techniques employed by sponges or coral
s, possibly because the shortness of bryozoan lifespans makes heavy investment in turf wars unprofitable.
lies outside the "crown" of tentacles (based on the Ancient Greek
prefix ἐκτο meaning "outside" and word πρωκτος meaning "anus"). After the discovery of the Entoprocta
, in which the anus lies within a "crown" of tentacles (based on the Ancient Greek prefix ἐντο meaning "inside" and word πρωκτος meaning "anus"), the name "Bryozoa" was used at phylum
level to include the two classes
Ectoprocta and Entoprocta. However, in 1869 Hinrich Nitsche regarded the two groups as quite distinct for a variety of reasons, and coined the name "Ectoprocta" for Ehrenberg's "Bryozoa". Despite their apparently similar methods of feeding, they differed markedly anatomically; in addition to the different positions of the anus, ectoprocts have hollow tentacles and a coelom
, while entoprocts have solid tentacles and no coelom. Hence the two groups are now widely regarded as separate phyla, and the name "Bryozoa" is now synonymous with "Ectoprocta". This has remained the majority view ever since, although most publications have preferred the name "Bryozoa" rather than "Ectoprocta". Nevertheless some notable scientists have continued to regard the "Ectoprocta" and Entoprocta as close relatives and group them under "Bryozoa".
The ambiguity about the scope of the name "Bryozoa" led to proposals in the 1960s and 1970s that it should be avoided and the unambiguous term "Ectoprocta" should be used. However, the change would have made it harder to find older works about in which the phylum was called "Bryozoa", and the desire to avoid ambiguity, if applied consistently to all classifications, would have necessitated renaming of several other phyla and many lower-level groups. In practice, zoological naming of split or merged groups of animals is complex and not completely consistent. Works since 2000 have used various names to resolve the ambiguity, including: "Bryozoa", "Ectoprocta", "Bryozoa (Ectoprocta)", and "Ectoprocta (Bryozoa)". Some have used more than one approach in the same work.
The common name "moss animals" is based on the Greek βρυόν (moss) and ζῷα (animals), and refers to the mossy appearance of encrusting species.
, which is still used as a convenient way to label groups of organisms, living members of the phylum
Bryozoa are divided into:
Fossils of about 15,000 bryozoan species have been found. The oldest species with a mineralized skeleton occurs in the uppermost Cambrian of Mexico. It is likely that the first bryozoans appeared much earlier and were entirely soft-bodied, and the Cambrian fossils record the appearance of mineralized skeletons in this phylum. By the Arenigian stage of the Early Ordovician
period, about , all the modern orders
of stenolaemates
were present, and the ctenostome
order of gymnolaemates
had appeared by the Middle Ordovician, about . The Early Ordovician fossils may also represent forms that had already become significantly different from the original members of the phylum. Other types of filter feeder
s appeared around the same time, which suggests that some change made the environment more favorable for this lifestyle. Fossils of cheilostomates
, another order of gymnolaemates, first appear in the Mid Jurassic
, about , and these have been the most abundant and diverse bryozoans from the Cretaceous
to the present. Evidence compiled from the last 100 million years show that cheilostomates consistently grew over cyclostomates in territorial struggles, which may help to explain how cheilostomates replaced cyclostomates as the dominant marine bryozoans. Marine fossils from the Paleozoic
era, which ended , are mainly of erect forms, those from the Mesozoic
are fairly equally divided by erect and encrusting forms, and more recent ones are predominantly encrusting. Fossils of the soft, freshwater phylactolaemates
are very rare, appear in and after the Late Permian (which began about ) and consist entirely of their durable statoblasts. There are no known fossils of freshwater members of other classes.
Since all the other phyla that have left fossils are found in Cambrian
rocks, it is surprising that the earliest bryozoan fossil dates from the Ordovician, which immediately followed the Cambrian. This suggests that the first bryozoans appeared much earlier and were entirely soft-bodied, and the Ordovician fossils record the appearance of mineralized skeletons in this phylum. The Early Ordovician fossils may also represent forms that had already become significantly different from the original members of the phylum.
s or deuterostome
s, the two major groups that account for all moderately complex animals.
The traditional view is that the Bryozoa are a monophyletic group, in which the class
Phylactolaemata
is most closely related to Stenolaemata
and Ctenostomata
, the classes that appear earliest in the fossil record. However, in 2005 a molecular phylogeny
study that focused on phylactolaemates concluded that these are more closely related to the phylum Phoronid
a, and especially to the only phoronid species that is colonial, than they are to the other ectoproct classes. That implies that the Entoprocta are not monophyletic, as the Phoronida are a sub-group of ectoprocts but the standard definition of Entoprocta excludes the Phoronida. In 2009 another molecular phylogeny
study, using a combination of genes from mitochondria and the cell nucleus
, concluded that Bryozoa is a monophyletic phylum, in other words includes all the descendants of a common ancestor that is itself a bryozoan. The analysis also concluded that the classes
Phylactolaemata
, Stenolaemata
and Gymnolaemata
are also monophyletic, but could not determine whether Stenolaemata
are more closely related to Phylactolaemata or Gymnolaemata
. The Gymnolaemata are traditionally divided into the soft-bodied Ctenostomata
and mineralized Cheilostomata, but the 2009 analysis considered it more likely that neither of these orders
is monophyletic and that mineralized skeleton
s probably evolved more than once within the early Gymnolaemata.
Bryozoans' relationships with other phyla are uncertain and controversial. Traditional phylogeny, based on anatomy
and on the development of the adult forms from embryo
s, has produced no enduring consensus about the position of ectoprocts. Attempts to reconstruct the family tree of animals have largely ignored ectoprocts and other "minor phyla", which have received little scientific study because they are generally tiny, have relatively simple body plans, and have little impact on human economies – despite the fact that the "minor phyla" include most of the variety in the evolutionary history of animals. In the opinion of Ruth Dewel, Judith Winston and Frank McKinney, "Our standard interpretation of bryozoan morphology
and embryology
is a construct resulting from over 100 years of attempts to synthesize a single framework for all invertebrates," and takes little account of some peculiar features of ectoprocts. In ectoprocts, all of the larva's internal organs are destroyed during the metamorphosis to the adult form and the adult's organs are built from the larva's epidermis and mesoderm
, while in other bilateria
ns some organs including the gut are built from endoderm
. In most bilaterian embryos the blastopore, a dent in the outer wall, deepens to become the larva's gut, but in ectoprocts the blastopore disappears and a new dent becomes the point from which the gut grows. The ectoproct coelom is formed by neither of the processes used by other bilaterians, enterocoely
, in which pouches that form on the wall of the gut become separate cavities, nor schizocoely, in which the tissue between the gut and the body wall splits, forming paired cavities.
Molecular phylogeny, which attempts to work out the evolutionary family tree of organisms by comparing their biochemistry
and especially their gene
s, has done much to clarify the relationships between the better-known invertebrate
phyla. However, the shortage of genetic data about "minor phyla" such as bryozoans and entoprocts
has left their relationships to other groups unclear.
When entoprocts were discovered in the 19th century, they and bryozoans (ectoprocts) were regarded as classes within the phylum Bryozoa, because both groups were sessile
animals that filter-fed
by means of a crown of tentacles that bore cilia. However, from 1869 onwards increasing awareness of differences, including the position of the entoproct anus
inside the feeding structure and the difference in the early pattern of division
of cells in their embryo
s, caused scientists to regard the two groups as separate phyla, and "Bryozoa" became just an alternative name for ectoprocts, in which the anus is outside the feeding organ. A series of molecular phylogeny studies from 1996 to 2006 have also concluded that bryozoans (ectoprocts) and entoprocts are not sister groups. However, two well-known zoologists, Claus Nielsen and Thomas Cavalier-Smith
, maintain on anatomical and developmental grounds that bryozoans and entoprocts are member of the same phylum, Bryozoa. A molecular phylogeny study in 2007 also supported this old idea, while its conclusions about other phyla agreed with those of several other analyses.
By 1891 bryozoans (ectoprocts) were grouped with phoronid
s in a super-phylum called "Tentaculata". In the 1970s comparisons between phoronid larvae and the cyphonautes larva of some gymnolaete bryozoans produced suggestions that the bryozoans, most of which are colonial, evolved from a semi-colonial species of phoronid. Brachiopod
s were also assigned to the "Tentaculata", which were renamed Lophophorata as they all use a lophophore
for filter feeding. Although the majority of scientists accept this, Claus Nielsen thinks these similarities are superficial. The Lophophorata are usually defined as animals with a lophophore, a three-part coelom and a U-shaped gut. In Nielsen's opinion, phoronids' and brachiopods' lophophores are more like those of pterobranchs, which are members of the phylum Hemichordata
. Bryozoan's tentacles bear cells with multiple cilia, while the corresponding cells of phoronids', brachiopods' and pterobranchs' lophophores have one cilium per cell; and bryozoan tentacles have no hemal canal ("blood vessel"), which those of the other three phyla have.
If the grouping of bryozoans with phoronids and brachiopods into Lophophorata is correct, the next issue is whether the Lophophorata are protostome
s, along with most invertebrate phyla, or deuterostome
s, along with chordate
s, hemichordates and echinoderm
s. The traditional view was that lophophorates were a mix of protostome and deuterostome features. Research from the 1970s onwards suggested they were deuterostomes, because of some features that were thought characteristic of deuterostomes: a three-part coelom; radial rather than spiral cleavage in the development of the embryo; and formation of the coelom by enterocoely
. However the coelom of ectoproct larvae shows no sign of division into three sections, and that of adult ectoprocts is different from that of other coelomate phyla as it is built anew from epidermis and mesoderm after metamorphosis has destroyed the larval coelom.
Molecular phylogeny analyses from 1995 onwards, using a variety of biochemical evidence and analytical techniques, placed the lophophorates as protostomes and closely related to annelid
s and molluscs in a super-phylum called Lophotrochozoa
. "Total evidence" analyses, which used both morphological features and a relatively small set of genes, came to various conclusions, mostly favoring a close relationship between lophophorates and Lophotrochozoa. A study in 2008, using a larger set of genes, concluded that the lophophorates were closer to the Lophotrochozoa than to deuterostomes, but also that the lophophorates were not monophyletic. Instead, it concluded that brachiopods and phoronids formed a monophyletic group, but bryozoans (ectoprocts) were closest to entoprocts, supporting the original definition of "Bryozoa".
s that sieve small particles, mainly phytoplankton
(microscopic floating plants), out of the water. The freshwater species Plumatella emarginata feeds on diatom
s, green algae
, cyanobacteria, non-photosynthetic bacteria
, dinoflagellate
s, rotifer
s, protozoa
, small nematode
s, and microscopic crustacean
s. While the currents that bryozoans generate to draw food towards the mouth are well understood, the exact method of capture is still debated. All species also flick larger particles towards the mouth with a tentacle, and a few capture zooplankton
(planktonic animals) by using their tentacles as cages. In addition the tentacles, whose surface area is increased by microvilli (small hairs and pleats), absorb organic compound
s dissolved in the water. Unwanted particles may be flicked away by tentacles or shut out by closing the mouth. A study in 2008 showed that both encrusting and erect colonies fed more quickly and grew faster in gentle than in strong currents.
In some species the first part of the stomach forms a muscular gizzard
lined with chitin
ous teeth that crush armored prey such as diatom
s. Wave-like peristaltic
contractions move the food through the stomach for digestion. The final section of the stomach is lined with cilia (minute hairs) that compress undigested solids, which then pass through the intestine
and out through the anus
.
There are no nephridia ("little kidneys") or other excretory
organs in bryzoa, and it is thought that ammonia
diffuses out through the body wall and lophophore. More complex waste products are not excreted but accumulate in the polypide
, which degenerates after a few weeks. Some of the old polypide is recycled, but much of it remains as a large mass of dying cells containing accumulated wastes, and this is compressed into a "brown body". When the degeneration is complete, the cystid (outer part of the animal) produces a new polypide, and the brown body remains in the coelom
, or in the stomach of the new polypide and is expelled next time the animal defecates.
or blood vessel
s. Instead zooids absorb oxygen
and eliminate carbon dioxide
through the body wall and especially the lophophore. The fluid in the coelom transports gases and nutrients and its circulation is passive, except that some relatively large species use cilia to boost its speed. The different bryozoan groups use various methods to share nutrients and oxygen between zooids: some have quite large gaps in the body walls, allowing the coelomic fluid to circulate freely; in others the funiculi (internal "little ropes") of adjacent zooids connect via small pores in the body wall.
s. Although those of many marine species are protandric, in other words function first as males and then as females, their colonies contain a combination of zooids that are in their male and female stages. In all species the ovaries
develop on the inside of the body wall, and the testes on the funiculus connecting the stomach to the body wall. Eggs and sperm are released into the coelom, and sperm exit into the water through pores in the tips of some of the tentacles, and then are captured by the feeding currents of zooids that are producing eggs. Some species' eggs are fertilized externally after being released through a pore between two tentacles, which in some cases is at the tip of a small projection called the "intertentacular organ" in the base of a pair of tentacles. Others' are fertilized internally, in the intertentacular organ or in the coelom. In ctenostomes the mother provides a brood chamber for the fertilized eggs, and her polypide disintegrates, providing nourishment to the embryo
. Stenolaemates produce specialized zooids to serve as brood chambers, and their eggs divide within this to produce up to 100 identical embryos.
The cleavage
of bryozoan eggs is biradial, in other words the early stages are bilaterally symmetrical. It is unknown how the coleom forms, since the metamorphosis
from larva
to adult destroys all of the larva's internal tissues. In many animals the blastopore
, an opening in the surface of the early embryo, tunnels through to form the gut. However, in bryozoans the blastopore closes, and a new opening develops to create the mouth.
Bryozoan larvae vary in form, but all have a band of cilia round the body which enables them to swim, a tuft of cilia at the top, and an adhesive sac that everts and anchors them when they settle on a surface. Some gymnolaemate species produce cyphonautes larvae which have little yolk but a well-developed mouth and gut, and live as plankton
for a considerable time before settling. These larvae have triangular shells of chitin, with one corner at the top and the base open, forming a hood round the downward-facing mouth. In 2006 it was reported that the cilia of cyphonautes larvae use the same range of techniques as those of adults to capture food. Species that brood their embryos form larvae that are nourished by large yolks, have no gut and do not feed, and such larvae quickly settle on a surface. In all marine species the larvae produce cocoons in which they metamorphose
completely after settling: the larva's epidermis becomes the lining of the coelom
, and the internal tissues are converted to a food reserve that nourishes the developing zooid until it is ready to feed. The larvae of phylactolaemates produce multiple polypides, so that each new colony starts with several zooids. In all species the founder zooids then grow the new colonies by budding
clone
s of themselves. In phylactolaemates, zooids die after producing several clones, so that living zooids are found only round the edges of a colony.
Phylactolaemates also reproduce asexually by a method that enables a colony's lineage to survive the variable and uncertain conditions of freshwater environments. Throughout summer and autumn they produce disc-shaped statoblasts, masses of cells that function as "survival pods" rather like the gemmules of sponges. Statoblasts form on the funiculus connected to the parent's gut, which nourishes them. As they grow, statoblasts develop protective bivalve-like
shells made of chitin
. When they mature, some statoblasts stick to the parent colony, some fall to the bottom ("sessoblasts"), some contain air spaces that enable them to float ("floatoblasts"), and some remain in the parent's cystid to re-build the colony if it dies. Statoblasts can remain dormant for considerable periods, and while dormant can survive harsh conditions such as freezing and desiccation
. They can be transported across long distances by animals, floating vegetation, currents and winds, and even in the guts of larger animals. When conditions improve, the valves of the shell separate and the cells inside develop into a zooid that tries to form a new colony. Plumatella emarginata produces both "sessoblasts", which enable the lineage to control a good territory even if hard times decimate the parent colonies, and "floatoblasts", which spread to new sites. New colonies of Plumatella repens produce mainly "sessoblasts" while mature ones switch to "floatoblasts". A study estimated that one group of colonies in a patch measuring 1 square metres (10.8 sq ft) produced 800,000 statoblasts.
, especially around cold seep
s, and others near the poles
. The great majority are sessile
. Encrusting forms are much the commonest of these in shallow seas, but erect forms become more common as the depth increases. A few marine species can move, and an Antarctic
species forms floating colonies.
The phylactolaemates live in all types of freshwater environment – lakes and ponds, rivers and streams, and estuaries
– and are among the most abundant sessile freshwater animals. Some ctenostomes are exclusively freshwater while others prefer brackish water but can survive in freshwater. Scientists' knowledge of freshwater bryozoan populations in many parts of the world is incomplete, even in some parts of Europe. It was long thought that some freshwater species occurred worldwide, but since 2002 all of these have been split into more localized species.
Marine species are common on coral reef
s, but seldom a significant proportion of the total biomass
. In temperate waters, the skeletons of dead colonies form a significant component of shell gravel
s, and live ones are abundant in these areas. The marine lace-like bryozoan Membranipora membranacea
produces spines in response to predation by several species of nudibranch
s ("sea slugs"). Other predators on marine bryozoans include fish, sea urchin
s, pycnogonids, crustacean
s, mite
s and starfish. In general marine echinoderms and molluscs eat masses of zooids by gouging pieces of colonies, breaking their mineralized "houses", while most arthropod
predators on bryozoans eat individual zooids.
In freshwater, bryozoans are among the most important filter feeder
s, along with sponges and mussel
s. Freshwater bryozoans are attacked by many predators, including snails, insects, and fish.
In Thailand
the introduced species
Pomacea canaliculata
(golden apple snail), which is generally a destructive herbivore
, has wiped out phylactolaemate populations wherever it has appeared. P. canaliculata also preys on a common freshwater gymnolaemate, but with less devastating effect. Indigenous snails do not feed on bryozoans.
Several species of the hydroid
family Zancleidae have symbiotic relationships with bryozoans, some of which are beneficial to the hydroids while others are parasitic. Modifications appear in the shapes of some these hydroids, for example smaller tentacles or encrustation of the roots by bryozoans. The bryozoan Alcyonidium nodosum protects the whelk
Burnupena papyracea
against predation by the powerful and voracious rock lobster
Jasus lalandii
. While whelk shells encrusted by the bryozoans are stronger than those without this reinforcement, chemical defenses produced by the bryozoans are probably the more significant deterrent.
Some phylactolaemate species are parasitized by a group of myxozoa
that have also been found to cause Proliferative Kidney Disease, which is often fatal in salmonid fish, and has severely reduced wild fish populations in Europe and North America.
Membranipora membranacea, whose colonies feed and grow exceptionally fast in a wide range of current speeds, was first noticed in the Gulf of Maine
in 1987 and quickly became the most abundant organism living on kelp
s. This invasion
reduced the kelp population by breaking their fronds, so that its place as the dominant "vegetation" in some areas was taken by another invader, the large alga Codium fragile
tomentosoides. These changes reduced the area of habitat available for local fish and invertebrates. M. membranacea has also invaded the northwest coast of the U.S.A. A few freshwater species have been also found thousands of kilometers from their native ranges. Some may have been transported naturally as statoblasts. Others more probably were spread by humans, for example on imported water plants or as stowaways on ships.
Fishermen in the North Sea
have had to find other work because of a form of eczema
(a skin disease) known as "Dogger Bank
itch", caused by contact with bryozoans that have stuck to nets and lobster pots.
Marine bryozoans are often responsible for biofouling
on ships' hulls, on docks and marinas, and on offshore structures. They are among the first colonizers of new or recently cleaned structures. Freshwater species are occasional nuisances in water pipes, drinking water purification equipment, sewage treatment facilities, and the cooling pipes of power stations.
A group of chemicals called bryostatin
s can be extracted from the marine bryozoan Bugula neritina. In 2001 pharmaceutical company GPC Biotech
licensed Bryostatin 1 from Arizona State University
for commercial development as a treatment for cancer
. GPC Biotech canceled development in 2003, saying that Bryostatin 1 showed little effectiveness and some toxic side-effects. In January 2008 a clinical trial
was submitted to the United States
National Institutes of Health
to measure the safety and effectiveness of Bryostatin 1 in the treatment of Alzheimer's Disease
. However, no participants had been recruited by the end of December 2008, when the study was scheduled for completion. About 1 tonnes (157.5 st) of bryozoans must processed to extract 1 gram (0.035273962105112 oz) of bryostatin. As a result, synthetic equivalents have been developed that are simpler to produce and apparently at least as effective.
Phylum
In biology, a phylum The term was coined by Georges Cuvier from Greek φῦλον phylon, "race, stock," related to φυλή phyle, "tribe, clan." is a taxonomic rank below kingdom and above class. "Phylum" is equivalent to the botanical term division....
of aquatic
Aquatic animal
An aquatic animal is an animal, either vertebrate or invertebrate, which lives in water for most or all of its life. It may breathe air or extract its oxygen from that dissolved in water through specialised organs called gills, or directly through its skin. Natural environments and the animals that...
invertebrate
Invertebrate
An invertebrate is an animal without a backbone. The group includes 97% of all animal species – all animals except those in the chordate subphylum Vertebrata .Invertebrates form a paraphyletic group...
animal
Animal
Animals are a major group of multicellular, eukaryotic organisms of the kingdom Animalia or Metazoa. Their body plan eventually becomes fixed as they develop, although some undergo a process of metamorphosis later on in their life. Most animals are motile, meaning they can move spontaneously and...
s. Typically about 0.5 millimetre (0.0196850393700787 in) long, they are filter feeder
Filter feeder
Filter feeders are animals that feed by straining suspended matter and food particles from water, typically by passing the water over a specialized filtering structure. Some animals that use this method of feeding are clams, krill, sponges, baleen whales, and many fish and some sharks. Some birds,...
s that sieve food particles out of the water using a retractable lophophore
Lophophore
The lophophore is a characteristic feeding organ possessed by four major groups of animals: the Brachiopoda, Bryozoa, Entoprocta, and Phoronida. All lophophores are found in aquatic organisms.-Characteristics:...
, a "crown" of tentacle
Tentacle
A tentacle or bothrium is one of usually two or more elongated flexible organs present in animals, especially invertebrates. The term may also refer to the hairs of the leaves of some insectivorous plants. Usually, tentacles are used for feeding, feeling and grasping. Anatomically, they work like...
s lined with cilia. Most marine
Marine (ocean)
Marine is an umbrella term. As an adjective it is usually applicable to things relating to the sea or ocean, such as marine biology, marine ecology and marine geology...
species live in tropical waters, but a few occur in oceanic trench
Oceanic trench
The oceanic trenches are hemispheric-scale long but narrow topographic depressions of the sea floor. They are also the deepest parts of the ocean floor....
es, and others are found in polar
Geographical pole
A geographical pole is either of the two points—the north pole and the south pole—on the surface of a rotating planet where the axis of rotation meets the surface of the body...
waters. One class
Class (biology)
In biological classification, class is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, order, family, genus, and species, with class fitting between phylum and order...
lives only in a variety of freshwater
Freshwater
Fresh water is naturally occurring water on the Earth's surface in ice sheets, ice caps, glaciers, bogs, ponds, lakes, rivers and streams, and underground as groundwater in aquifers and underground streams. Fresh water is generally characterized by having low concentrations of dissolved salts and...
environments, and a few members of a mostly marine class prefer brackish water. Over 4,000 living species are known. One genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
is solitary and the rest colonial
Colony (biology)
In biology, a colony reference to several individual organisms of the same species living closely together, usually for mutual benefit, such as stronger defense or the ability to attack bigger prey. Some insects live only in colonies...
.
The phylum was originally called "Polyzoa", but this term was superseded by "Bryozoa" in 1831. Another group of animals discovered subsequently, whose filtering mechanism looked similar, was also included in "Bryozoa" until 1869, when the two groups were noted to be very different internally. The more recently discovered group were given the name Entoprocta
Entoprocta
Entoprocta, whose name means "anus inside", is a phylum of mostly sessile aquatic animals, ranging from long. Mature individuals are goblet-shaped, on relatively long stalks. They have a "crown" of solid tentacles whose cilia generate water currents that draw food particles towards the mouth, and...
, while the original "Bryozoa" were called "Ectoprocta". However, "Bryozoa" has remained the more widely used term for the latter group.
Individuals in bryozoan (ectoproct) colonies are called zooids, since they are not fully independent animals. All colonies contain autozooids, which are responsible for feeding and excretion
Excretion
Excretion is the process by which waste products of metabolism and other non-useful materials are eliminated from an organism. This is primarily carried out by the lungs, kidneys and skin. This is in contrast with secretion, where the substance may have specific tasks after leaving the cell...
. Colonies of some classes
Class (biology)
In biological classification, class is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, order, family, genus, and species, with class fitting between phylum and order...
have various types of non-feeding specialist zooids, some of which are hatcheries for fertilized eggs, and some classes also have special zooids for defense of the colony. The class Cheilostomata
Cheilostomata
Cheilostomata, an order of Bryozoa in the class Gymnolaemata, are exclusively marine, colonial invertebrate animals. Cheilostome colonies are composed of calcium carbonate and grow on a variety of surfaces, including rocks, shells, seagrass and kelps. The colony shapes range from simple encrusting...
have the largest number of species, possibly because they have the widest range of specialist zooids. A few species can creep very slowly by using spiny defensive zooids as legs. Autozooids supply nutrients to non-feeding zooids by channels that vary between classes. All zooids, including those of the solitary species, consist of a cystid that provides the body wall and produces the exoskeleton
Exoskeleton
An exoskeleton is the external skeleton that supports and protects an animal's body, in contrast to the internal skeleton of, for example, a human. In popular usage, some of the larger kinds of exoskeletons are known as "shells". Examples of exoskeleton animals include insects such as grasshoppers...
and a polypide that contains the internal organs and the lophophore or other specialist extensions. Zooids have no special excretory organs, and the polypides of autozooids are scrapped when the polypides become overloaded by waste products; usually the body wall then grows a replacement polypide. In autozooids the gut is U-shaped, with the mouth inside the "crown" of tentacles and the anus outside it. Colonies take a variety of forms, including fans, bushes and sheets. The Cheilostomata produce mineralized exoskeletons and form single-layered sheets that encrust over surfaces.
Zooids of all the freshwater species are simultaneous hermaphrodite
Hermaphrodite
In biology, a hermaphrodite is an organism that has reproductive organs normally associated with both male and female sexes.Many taxonomic groups of animals do not have separate sexes. In these groups, hermaphroditism is a normal condition, enabling a form of sexual reproduction in which both...
s. Although those of many marine species function first as males and then as females, their colonies always contain a combination of zooids that are in their male and female stages. All species emit sperm
Sperm
The term sperm is derived from the Greek word sperma and refers to the male reproductive cells. In the types of sexual reproduction known as anisogamy and oogamy, there is a marked difference in the size of the gametes with the smaller one being termed the "male" or sperm cell...
into the water. Some also release ova
Ovum
An ovum is a haploid female reproductive cell or gamete. Both animals and embryophytes have ova. The term ovule is used for the young ovum of an animal, as well as the plant structure that carries the female gametophyte and egg cell and develops into a seed after fertilization...
into the water, while others capture sperm via their tentacles to fertilize their ova internally. In some species the larva
Larva
A larva is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle...
e have large yolks, go to feed, and quickly settle on a surface. Others produce larvae that have little yolk but swim and feed for a few days before settling. After settling, all larvae undergo a radical metamorphosis
Metamorphosis
Metamorphosis is a biological process by which an animal physically develops after birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation...
that destroys and rebuilds almost all the internal tissues. Freshwater species also produce statoblasts that lie dormant until conditions are favorable, which enables a colony's lineage to survive even if severe conditions kill the mother colony.
Predators of marine bryozoans include nudibranch
Nudibranch
A nudibranch is a member of what is now a taxonomic clade, and what was previously a suborder, of soft-bodied, marine gastropod mollusks which shed their shell after their larval stage. They are noted for their often extraordinary colors and striking forms...
s (sea slugs), fish, sea urchin
Sea urchin
Sea urchins or urchins are small, spiny, globular animals which, with their close kin, such as sand dollars, constitute the class Echinoidea of the echinoderm phylum. They inhabit all oceans. Their shell, or "test", is round and spiny, typically from across. Common colors include black and dull...
s, pycnogonids, crustacean
Crustacean
Crustaceans form a very large group of arthropods, usually treated as a subphylum, which includes such familiar animals as crabs, lobsters, crayfish, shrimp, krill and barnacles. The 50,000 described species range in size from Stygotantulus stocki at , to the Japanese spider crab with a leg span...
s, mite
Mite
Mites, along with ticks, are small arthropods belonging to the subclass Acari and the class Arachnida. The scientific discipline devoted to the study of ticks and mites is called acarology.-Diversity and systematics:...
s and starfish. Freshwater bryozoans are preyed on by snails, insects, and fish. In Thailand
Thailand
Thailand , officially the Kingdom of Thailand , formerly known as Siam , is a country located at the centre of the Indochina peninsula and Southeast Asia. It is bordered to the north by Burma and Laos, to the east by Laos and Cambodia, to the south by the Gulf of Thailand and Malaysia, and to the...
, many populations of one freshwater species have been wiped out by an introduced species
Introduced species
An introduced species — or neozoon, alien, exotic, non-indigenous, or non-native species, or simply an introduction, is a species living outside its indigenous or native distributional range, and has arrived in an ecosystem or plant community by human activity, either deliberate or accidental...
of snail. A fast-growing invasive bryozoan off the northeast and northwest coasts of the USA has reduced kelp
Kelp
Kelps are large seaweeds belonging to the brown algae in the order Laminariales. There are about 30 different genera....
forests so much that it has affected local fish and invertebrate populations. Bryozoans have spread diseases to fish farms and fishermen. Chemicals extracted from a marine bryozoan species have been investigated for treatment of cancer
Cancer
Cancer , known medically as a malignant neoplasm, is a large group of different diseases, all involving unregulated cell growth. In cancer, cells divide and grow uncontrollably, forming malignant tumors, and invade nearby parts of the body. The cancer may also spread to more distant parts of the...
and Alzheimer's Disease
Alzheimer's disease
Alzheimer's disease also known in medical literature as Alzheimer disease is the most common form of dementia. There is no cure for the disease, which worsens as it progresses, and eventually leads to death...
, but analyses have not been encouraging.
Mineralized skeletons of bryozoans first appear in rocks from latest Cambrian
Cambrian
The Cambrian is the first geological period of the Paleozoic Era, lasting from Mya ; it is succeeded by the Ordovician. Its subdivisions, and indeed its base, are somewhat in flux. The period was established by Adam Sedgwick, who named it after Cambria, the Latin name for Wales, where Britain's...
period, about , making it the last major phylum to appear in the fossil record. This has led researchers to suspect that bryozoans had arisen earlier but were initially unmineralized, and may have differed significantly from fossilized and modern forms. Early fossils are mainly of erect forms, but encrusting forms gradually became dominant. It is uncertain whether the phylum is monophyletic. Bryozoans' evolutionary relationships to other phyla are also unclear, partly because scientists' view of the family tree of animals is mainly influenced by better-known phyla. Both morphological
Morphology (biology)
In biology, morphology is a branch of bioscience dealing with the study of the form and structure of organisms and their specific structural features....
and molecular phylogeny
Molecular phylogeny
Molecular phylogenetics is the analysis of hereditary molecular differences, mainly in DNA sequences, to gain information on an organism's evolutionary relationships. The result of a molecular phylogenetic analysis is expressed in a phylogenetic tree...
analyses disagree over bryozoans' relationships with entoprocts, about whether bryozoans should be grouped with brachiopod
Brachiopod
Brachiopods are a phylum of marine animals that have hard "valves" on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear end, while the front can be opened for feeding or closed for protection...
s and phoronid
Phoronid
Phoronids are a phylum of marine animals that filter-feed with a lophophore , and build upright tubes of chitin to support and protect their soft bodies. They live in all the oceans and seas including the Arctic Ocean but excluding the Antarctic Ocean, and between the intertidal zone and about...
s in Lophophorata, and whether bryozoans should be considered protostome
Protostome
Protostomia are a clade of animals. Together with the deuterostomes and a few smaller phyla, they make up the Bilateria, mostly comprising animals with bilateral symmetry and three germ layers...
s or deuterostome
Deuterostome
Deuterostomes are a superphylum of animals. They are a subtaxon of the Bilateria branch of the subregnum Eumetazoa, and are opposed to the protostomes...
s.
Distinguishing features
Lophophore
The lophophore is a characteristic feeding organ possessed by four major groups of animals: the Brachiopoda, Bryozoa, Entoprocta, and Phoronida. All lophophores are found in aquatic organisms.-Characteristics:...
, a "crown" of hollow tentacles. Bryozoans form colonies consisting of clone
Cloning
Cloning in biology is the process of producing similar populations of genetically identical individuals that occurs in nature when organisms such as bacteria, insects or plants reproduce asexually. Cloning in biotechnology refers to processes used to create copies of DNA fragments , cells , or...
s called zooids that are typically about 0.5 millimetre (0.0196850393700787 in) long. Phoronids resemble bryozoan zooids but are 2 to 20 cm (0.78740157480315 to 7.9 in) long and, although they often grow in clumps, do not form colonies consisting of clones. Brachiopods, generally thought to be closely related to bryozoans and phoronids, are distinguished by having shells rather like those of bivalves. All three of these phyla
Phylum
In biology, a phylum The term was coined by Georges Cuvier from Greek φῦλον phylon, "race, stock," related to φυλή phyle, "tribe, clan." is a taxonomic rank below kingdom and above class. "Phylum" is equivalent to the botanical term division....
have a coelom
Coelom
The coelom is a fluid-filled cavity formed within the mesoderm. Coeloms developed in triploblasts but were subsequently lost in several lineages. Loss of coelom is correlated with reduction in body size...
, an internal cavity lined by mesothelium
Mesothelium
The mesothelium is a membrane that forms the lining of several body cavities: the pleura , peritoneum and pericardium . Mesothelial tissue also surrounds the male internal reproductive organs and covers the internal reproductive organs of women...
.
Some encrusting bryozoan colonies with mineralized exoskeleton
Exoskeleton
An exoskeleton is the external skeleton that supports and protects an animal's body, in contrast to the internal skeleton of, for example, a human. In popular usage, some of the larger kinds of exoskeletons are known as "shells". Examples of exoskeleton animals include insects such as grasshoppers...
s look very like small corals. However, bryozoan colonies are founded by an ancestrula, which is round rather than shaped like a normal zooid of that species. On the other hand the founding polyp of a coral has a shape like that of its daughter polyps, and coral zooids have no coelom or lophophore.
Entoprocts
Entoprocta
Entoprocta, whose name means "anus inside", is a phylum of mostly sessile aquatic animals, ranging from long. Mature individuals are goblet-shaped, on relatively long stalks. They have a "crown" of solid tentacles whose cilia generate water currents that draw food particles towards the mouth, and...
, another phylum of filter-feeders, look rather like bryozoans but their lophophore-like feeding structure has solid tentacles, their anus
Anus
The anus 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, unwanted semi-solid matter produced during digestion, which, depending on the type of animal, may be one or more of: matter which the animal cannot digest,...
lies inside rather than outside the base of the "crown" and they have no coelom
Coelom
The coelom is a fluid-filled cavity formed within the mesoderm. Coeloms developed in triploblasts but were subsequently lost in several lineages. Loss of coelom is correlated with reduction in body size...
.
| Bryozoa (Ectoprocta) | | Other lophophorates | Other Lophotrochozoa Lophotrochozoa The Lophotrochozoa are a major grouping of protostome animals. The taxon was discovered based on molecular data. Molecular evidence such as a result of studies of the evolution of small-subunit ribosomal RNA supports the monophyly of the phyla listed in the infobox shown at right.-Terminology:The... | | Similar-looking phyla | |||
|---|---|---|---|---|---|---|
| Phoronida | Brachiopoda | Annelida, Mollusca Mollusca The Mollusca , common name molluscs or mollusksSpelled mollusks in the USA, see reasons given in Rosenberg's ; for the spelling mollusc see the reasons given by , is a large phylum of invertebrate animals. There are around 85,000 recognized extant species of molluscs. Mollusca is the largest... | Entoprocta Entoprocta Entoprocta, whose name means "anus inside", is a phylum of mostly sessile aquatic animals, ranging from long. Mature individuals are goblet-shaped, on relatively long stalks. They have a "crown" of solid tentacles whose cilia generate water currents that draw food particles towards the mouth, and... | Coral Coral Corals are marine animals in class Anthozoa of phylum Cnidaria typically living in compact colonies of many identical individual "polyps". The group includes the important reef builders that inhabit tropical oceans and secrete calcium carbonate to form a hard skeleton.A coral "head" is a colony of... s (class in phylum Cnidaria Cnidaria Cnidaria is a phylum containing over 9,000 species of animals found exclusively in aquatic and mostly marine environments. Their distinguishing feature is cnidocytes, specialized cells that they use mainly for capturing prey. Their bodies consist of mesoglea, a non-living jelly-like substance,... ) |
||
| Coelom Coelom The coelom is a fluid-filled cavity formed within the mesoderm. Coeloms developed in triploblasts but were subsequently lost in several lineages. Loss of coelom is correlated with reduction in body size... |
Three-part, if the cavity of the epistome is included | Three-part | One per segment in basic form; merged in some taxa Taxon |thumb|270px|[[African elephants]] form a widely-accepted taxon, the [[genus]] LoxodontaA taxon is a group of organisms, which a taxonomist adjudges to be a unit. Usually a taxon is given a name and a rank, although neither is a requirement... |
none | ||
| Formation of coelom | Uncertain because metamorphosis Metamorphosis Metamorphosis is a biological process by which an animal physically develops after birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation... of larvae into adults makes this impossible to trace |
Enterocoely Enterocoely Enterocoely is a process by which some animal embryos develop. In enterocoely, a mesoderm is formed in a developing embryo, in which the coelom forms from pouches "pinched" off of the digestive tract... |
Schizocoely | not applicable | ||
| Lophophore | With hollow tentacles | none | Similar-looking feeding structure, but with solid tentacles | none | ||
| Feeding current | From tips to bases of tentacles | not applicable | From bases to tips of tentacles | not applicable | ||
| Multiciliated cells in epithelium Epithelium Epithelium is one of the four basic types of animal tissue, along with connective tissue, muscle tissue and nervous tissue. Epithelial tissues line the cavities and surfaces of structures throughout the body, and also form many glands. Functions of epithelial cells include secretion, selective... |
Yes | no | Yes | not applicable | ||
| Position of anus Anus The anus 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, unwanted semi-solid matter produced during digestion, which, depending on the type of animal, may be one or more of: matter which the animal cannot digest,... |
Outside base of lophophore | Varies, none in some species | Rear end, but none in Siboglinidae Siboglinidae Siboglinidae, also known as the beard worms, is a family of polychaete annelid worms whose members made up the former phyla Pogonophora and Vestimentifera. They are composed of about 100 species of vermiform creatures and live in thin tubes buried in sediments at ocean depths from 100 to 10,000 m... |
Inside base of lophophore-like organ | none | |
| Colonial | Colonies of clones in most; one solitary genus Genus In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia... |
Sessile Sessility (zoology) In zoology, sessility is a characteristic of animals which are not able to move about. They are usually permanently attached to a solid substrate of some kind, such as a part of a plant or dead tree trunk, a rock, or the hull of a ship in the case of barnacles. Corals lay down their own... species often form clumps, but with no active co-operation |
Colonies of clones in some species; some solitary species | Colonies of clones | ||
| Shape of founder zooid | Round, unlike normal zooids | not applicable | Same as other zooids | |||
| Mineralized exoskeleton Exoskeleton An exoskeleton is the external skeleton that supports and protects an animal's body, in contrast to the internal skeleton of, for example, a human. In popular usage, some of the larger kinds of exoskeletons are known as "shells". Examples of exoskeleton animals include insects such as grasshoppers... s |
Some taxa Taxon |thumb|270px|[[African elephants]] form a widely-accepted taxon, the [[genus]] LoxodontaA taxon is a group of organisms, which a taxonomist adjudges to be a unit. Usually a taxon is given a name and a rank, although neither is a requirement... |
no | Bivalve-like shells | Some sessile annelids build mineralized tubes; most molluscs have shells, but most modern cephalopod Cephalopod A cephalopod is any member of the molluscan class Cephalopoda . These exclusively marine animals are characterized by bilateral body symmetry, a prominent head, and a set of arms or tentacles modified from the primitive molluscan foot... s have internal shells or none. |
no | Some taxa |
Types of zooid
All bryozoans are colonial except for one genusGenus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
, Monobryozoon. Individual members of a bryozoan colony are about 0.5 millimetre (0.0196850393700787 in) long and are known as zooids, since they are not fully independent animals. All colonies contain feeding zooids, known as autozooids, and those of some groups also contain non-feeding specialist heterozooids; colony members are genetically identical and co-operate, rather like the organs of larger animals. What type of zooid grows where in a colony is determined by chemical signals from the colony as a whole or sometimes in response to the scent of predators or rival colonies.
The bodies of all types have two main parts. The cystid consists of the body wall and whatever type of exoskeleton
Exoskeleton
An exoskeleton is the external skeleton that supports and protects an animal's body, in contrast to the internal skeleton of, for example, a human. In popular usage, some of the larger kinds of exoskeletons are known as "shells". Examples of exoskeleton animals include insects such as grasshoppers...
is secreted
Secretion
Secretion is the process of elaborating, releasing, and oozing chemicals, or a secreted chemical substance from a cell or gland. In contrast to excretion, the substance may have a certain function, rather than being a waste product...
by the epidermis
Epidermis (zoology)
The Epidermis is an epithelium that covers the body of an eumetazoan . Eumetazoa have a cavity lined with a similar epithelium, the gastrodermis, which forms a boundary with the epidermis at the mouth.Sponges have no epithelium, and therefore no epidermis or gastrodermis...
. The exoskeleton may be organic (chitin
Chitin
Chitin n is a long-chain polymer of a N-acetylglucosamine, a derivative of glucose, and is found in many places throughout the natural world...
, polysaccharide
Polysaccharide
Polysaccharides are long carbohydrate molecules, of repeated monomer units joined together by glycosidic bonds. They range in structure from linear to highly branched. Polysaccharides are often quite heterogeneous, containing slight modifications of the repeating unit. Depending on the structure,...
or protein
Protein
Proteins are biochemical compounds consisting of one or more polypeptides typically folded into a globular or fibrous form, facilitating a biological function. A polypeptide is a single linear polymer chain of amino acids bonded together by peptide bonds between the carboxyl and amino groups of...
) or made of the mineral calcium carbonate
Calcium carbonate
Calcium carbonate is a chemical compound with the formula CaCO3. It is a common substance found in rocks in all parts of the world, and is the main component of shells of marine organisms, snails, coal balls, pearls, and eggshells. Calcium carbonate is the active ingredient in agricultural lime,...
. The body wall consists of the epidermis, basal lamina
Basal lamina
The basal lamina is a layer of extracellular matrix secreted by the epithelial cells, on which the epithelium sits. It is often confused with the basement membrane, and sometimes used inconsistently in the literature, see below....
(a mat of non-cellular material), connective tissue
Connective tissue
"Connective tissue" is a fibrous tissue. It is one of the four traditional classes of tissues . Connective Tissue is found throughout the body.In fact the whole framework of the skeleton and the different specialized connective tissues from the crown of the head to the toes determine the form of...
, muscles, and the mesothelium
Mesothelium
The mesothelium is a membrane that forms the lining of several body cavities: the pleura , peritoneum and pericardium . Mesothelial tissue also surrounds the male internal reproductive organs and covers the internal reproductive organs of women...
which lines the coelom
Coelom
The coelom is a fluid-filled cavity formed within the mesoderm. Coeloms developed in triploblasts but were subsequently lost in several lineages. Loss of coelom is correlated with reduction in body size...
(main body cavity) – except that in one class
Class (biology)
In biological classification, class is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, order, family, genus, and species, with class fitting between phylum and order...
, the mesothelium is split into two separate layers, the inner one forming a membranous sac that floats freely and contains the coelom, and the outer one attached to the body wall and enclosing the membranous sac in a pseudocoelom. The other main part of the bryozoan body, known as the polypide and situated almost entirely within the cystid, contains the nervous system, digestive system, some specialized muscles and the feeding apparatus or other specialized organs that take the place of the feeding apparatus.
Feeding zooids
The most common type of zooid is the feeding autozooid, in which the polypide bears a "crown" of hollow tentacles called a lophophoreLophophore
The lophophore is a characteristic feeding organ possessed by four major groups of animals: the Brachiopoda, Bryozoa, Entoprocta, and Phoronida. All lophophores are found in aquatic organisms.-Characteristics:...
, which captures food particles from the water. In all colonies a large percentage of zooids are autozooids, and some consist entirely of autozooids, some of which also engage in reproduction.
The basic shape of the "crown" is a full circle. In the class
Class (biology)
In biological classification, class is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, order, family, genus, and species, with class fitting between phylum and order...
Phylactolaemata
Phylactolaemata
Phylactolaemata is a class of the phylum Bryozoa whose members live only in freshwater environments. Like all bryozoans, they filter feed by means of an extensible "crown" of ciliated tentacles called a lophophore. They live in colonies, each of which consists of clones of the founding member...
the crown appears U-shaped, but this impression is created by a deep dent in the rim of the crown, which has no gap in the fringe of tentacles. The sides of the tentacles bear fine hairs called cilia, whose beating drives a water current from the tips of the tentacles to their bases, where it exits. Food particles that collide with the tentacles are trapped by mucus
Mucus
In vertebrates, mucus is a slippery secretion produced by, and covering, mucous membranes. Mucous fluid is typically produced from mucous cells found in mucous glands. Mucous cells secrete products that are rich in glycoproteins and water. Mucous fluid may also originate from mixed glands, which...
, and further cilia on the inner surfaces of the tentacles convey the particles towards the mouth, which lies in the center of the base of the "crown". The method used by ectoprocts is known as "upstream collecting", as food particles are captured before they pass through the field of cilia that creates the feeding current. This method is also used by phoronid
Phoronid
Phoronids are a phylum of marine animals that filter-feed with a lophophore , and build upright tubes of chitin to support and protect their soft bodies. They live in all the oceans and seas including the Arctic Ocean but excluding the Antarctic Ocean, and between the intertidal zone and about...
s, brachiopod
Brachiopod
Brachiopods are a phylum of marine animals that have hard "valves" on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear end, while the front can be opened for feeding or closed for protection...
s and pterobranchs.
The lophophore and mouth are mounted on a flexible tube, called the "invert" because it can be turned inside-out and withdrawn into the polypide, rather like the finger of a rubber glove; in this position the lophophore lies inside the invert and is folded like the spokes of an umbrella. The invert is withdrawn, sometimes within 60 millisecond
Millisecond
A millisecond is a thousandth of a second.10 milliseconds are called a centisecond....
s, by a pair of retractor muscles that are anchored at the far end of the cystid. Sensors at the tips of the tentacles may check for signs of danger before the invert and lophophore are fully extended. Extension is driven by an increase in internal fluid pressure, which species with flexible exoskeletons produce by contracting circular muscles that lie just inside the body wall, while species with a membranous sac use circular muscles to squeeze this. Some species with rigid exoskeletons have a flexible membrane that replaces part of the exoskeleton, and transverse muscles anchored on the far side of the exoskeleton increase the fluid pressure by pulling the membrane inwards. In others there is no gap in the protective skeleton, and the transverse muscles pull on a flexible sac which is connected to the water outside by a small pore; the expansion of the sac increases the pressure inside the body and pushes the invert and lophophore out. In some species the retracted invert and lophophore are protected by an operculum ("lid"), which is closed by muscles and opened by fluid pressure. In one class
Class (biology)
In biological classification, class is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, order, family, genus, and species, with class fitting between phylum and order...
, a hollow lobe called the "epistome" overhands the mouth.
The gut is U-shaped, running from the mouth, in the center of the lophophore, down into the animal's interior and then back to the anus
Anus
The anus 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, unwanted semi-solid matter produced during digestion, which, depending on the type of animal, may be one or more of: matter which the animal cannot digest,...
, which is located on the invert, outside and usually below the lophophore. A network of strands of mesothelium
Mesothelium
The mesothelium is a membrane that forms the lining of several body cavities: the pleura , peritoneum and pericardium . Mesothelial tissue also surrounds the male internal reproductive organs and covers the internal reproductive organs of women...
called "funiculi" ("little ropes") connects the mesothelium covering the gut with that lining the body wall. The wall of each strand is made of mesothelium, and surrounds a space filled with fluid, thought to be blood. A colony's zooids are connected, enabling autozooids to share food with each other and with any non-feeding heterozooids. The method of connection varies between the different classes of bryozoans, ranging from quite large gaps in the body walls to small pores through which nutrients are passed by funiculi.
There is a nerve ring round the pharynx (throat) and a ganglion
Ganglion
In anatomy, a ganglion is a biological tissue mass, most commonly a mass of nerve cell bodies. Cells found in a ganglion are called ganglion cells, though this term is also sometimes used to refer specifically to retinal ganglion cells....
that serves as a brain to one side of this. Nerves run from the ring and ganglion to the tentacles and to the rest of the body. Bryozoans have no specialized sense organs, but cilia on the tentacles act as sensors. Members of the genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
Bugula
Bugula
Spiral tufted bryozoa or Bugula turrita are a very common colonial marine animal found from Maine to North Carolina.Bugula neritina is of current interest as a source of cytotoxic chemicals, bryostatins, under clinical investigation as anti-cancer agents.Dried Bugula are commonly used as...
grow towards the sun
Phototropism
Phototropism is directional growth in which the direction of growth is determined by the direction of the light source. In other words, it is the growth and response to a light stimulus. Phototropism is most often observed in plants, but can also occur in other organisms such as fungi...
, and therefore must be able to detect light. In colonies of some species, signals are transmitted between zooids through nerves that pass through pores in the body walls, and coordinate activities such as feeding and the retraction of lophophores.
The solitary individuals of Monobryozoon are autozooids with pear-shaped bodies. The wider ends have up to 15 short, muscular projections by which the animals anchor themselves to sand or gravel and pull themselves through the sediments.
Avicularia and vibracula
Some authorities use the term avicularia to refer to any type of zooid in which the lophophore is replaced by an extension that serves some protective function, while others restrict the term to those that defend the colony by snapping at invaders and small predators, killing some and biting the appendageAppendage
In invertebrate biology, an appendage is an external body part, or natural prolongation, that protrudes from an organism's body . It is a general term that covers any of the homologous body parts that may extend from a body segment...
s of others. In some species the snapping zooids are mounted on a peduncle (stalk), their bird-like appearance responsible for the term – Charles Darwin
Charles Darwin
Charles Robert Darwin FRS was an English naturalist. He established that all species of life have descended over time from common ancestry, and proposed the scientific theory that this branching pattern of evolution resulted from a process that he called natural selection.He published his theory...
described these as like "the head and beak of a vulture in miniature, seated on a neck and capable of movement". Stalked avicularia are placed upside-down on their stalks. The "lower jaws" are modified versions of the opercula that protect the retracted lophophores in autozooids of some species, and are snapped shut "like a mousetrap" by similar muscles, while the beak-shaped upper jaw is the inverted body wall. In other species the avicularia are stationary box-like zooids laid the normal way up, so that the modified operculum snaps down against the body wall. In both types the modified operculum is opened by other muscles that attach to it, or by internal muscles that raise the fluid pressure by pulling on a flexible membrane. The actions of these snapping zooids are controlled by small, highly modified polypides that are located inside the "mouth" and bear tufts of short sensory cilia. These zooids appear in various positions: some take the place of autozooids, some fit into small gaps between autozooids, and small avicularia may occur on the surfaces of other zooids.
In vibracula, regarded by some as a type of avicularia, the operculum is modified to form a long bristle that has a wide range of motion. They may function as defenses against predators and invaders, or as cleaners. In some species that form mobile colonies, vibracula around the edges are used as legs for burrowing and walking.
Other types of colonial zooid
Kenozooids (from Greek κενος meaning "empty") consist only of the body wall and funicular strands crossing the interior, and no polypide. In some species they form the stems of branching structures, while in others they act as spacers that enable colonies to grow quickly in a new direction.Spinozooids form defensive spines, and sometimes appear on top of autozooids. Gonozooids act as brood chambers for fertilized eggs. Some species have miniature nanozooids with small single-tentacled polypides, and these may grow on other zooids or within the body walls of autozooids that have degenerated.
Colony forms and composition
Although zooids are microscopic, colonies range in size from 1 centimetre (0.393700787401575 in) to over 1 metres (3.3 ft). However, the majority are under 10 centimetres (3.9 in) across. The shapes of colonies vary widely, depend on the pattern of budding by which they grow, the variety of zooids present and the type and amount of skeletal material they secreteSecretion
Secretion is the process of elaborating, releasing, and oozing chemicals, or a secreted chemical substance from a cell or gland. In contrast to excretion, the substance may have a certain function, rather than being a waste product...
.
Some marine species are bush-like or fan-like, supported by "trunks" and "branches" formed by kenozooids, with feeding autozooids growing from these. Colonies of these types are generally unmineralized but may have exoskeleton
Exoskeleton
An exoskeleton is the external skeleton that supports and protects an animal's body, in contrast to the internal skeleton of, for example, a human. In popular usage, some of the larger kinds of exoskeletons are known as "shells". Examples of exoskeleton animals include insects such as grasshoppers...
s made of chitin
Chitin
Chitin n is a long-chain polymer of a N-acetylglucosamine, a derivative of glucose, and is found in many places throughout the natural world...
. Others look like small coral
Coral
Corals are marine animals in class Anthozoa of phylum Cnidaria typically living in compact colonies of many identical individual "polyps". The group includes the important reef builders that inhabit tropical oceans and secrete calcium carbonate to form a hard skeleton.A coral "head" is a colony of...
s, producing heavy lime skeletons. Many species form colonies which consist of sheets of autozooids. These sheets may form leaves, tufts or, in the genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
Thalmoporella, structures that resemble an open head of lettuce
Lettuce
Lettuce is a temperate annual or biennial plant of the daisy family Asteraceae. It is most often grown as a leaf vegetable. It is eaten either raw, notably in salads, sandwiches, hamburgers, tacos, and many other dishes, or cooked, as in Chinese cuisine in which the stem becomes just as important...
.
The most common marine form, however, is encrusting, in which a one-layer sheet of zooids spreads over a hard surface or over seaweed. Some encrusting colonies may grow to over 50 centimetres (1.6 ft) and contain about 2,000,000 zooids. These species generally have exoskeletons reinforced with calcium carbonate
Calcium carbonate
Calcium carbonate is a chemical compound with the formula CaCO3. It is a common substance found in rocks in all parts of the world, and is the main component of shells of marine organisms, snails, coal balls, pearls, and eggshells. Calcium carbonate is the active ingredient in agricultural lime,...
, and the openings through which the lophophores protrude are on the top or outer surface. The moss-like appearance of encrusting colonies is responsible for the phylum's alternative name Bryozoa (Ancient Greek
Ancient Greek
Ancient Greek is the stage of the Greek language in the periods spanning the times c. 9th–6th centuries BC, , c. 5th–4th centuries BC , and the c. 3rd century BC – 6th century AD of ancient Greece and the ancient world; being predated in the 2nd millennium BC by Mycenaean Greek...
words βρυος bryos meaning "moss" and ζωον zoon meaning "animal"). Large colonies of encrusting species often have "chimney
Chimney
A chimney is a structure for venting hot flue gases or smoke from a boiler, stove, furnace or fireplace to the outside atmosphere. Chimneys are typically vertical, or as near as possible to vertical, to ensure that the gases flow smoothly, drawing air into the combustion in what is known as the...
s", gaps in the canopy of lophophores, through which they swiftly expel water that has been sieved, and thus avoid re-filtering water that is already exhausted. They are formed by patches of non-feeding heterozooids. New chimneys appear near the edges of expanding colonies, at points where the speed of the outflow is already high, and do not change position if the water flow changes.
Some freshwater species secrete a mass of gelatinous material, up to 1 metres (3.3 ft) in diameter, to which the zooids stick. Other freshwater species have plant-like shapes with "trunks" and "branches", which may stand erect or spread over the surface. A few species can creep at about 2 centimetre (0.78740157480315 in) per day.
Each colony grows by asexual
Asexual reproduction
Asexual reproduction is a mode of reproduction by which offspring arise from a single parent, and inherit the genes of that parent only, it is reproduction which does not involve meiosis, ploidy reduction, or fertilization. A more stringent definition is agamogenesis which is reproduction without...
budding
Budding
Budding is a form of asexual reproduction in which a new organism grows on another one. The new organism remains attached as it grows, separating from the parent organism only when it is mature. Since the reproduction is asexual, the newly created organism is a clone and is genetically identical...
from a single zooid known as the ancestrula, which is round rather than shaped like a normal zooid. This occurs at the tips of "trunks" or "branches" in forms that have this structure. Encrusting colonies grow round their edges. In species with calcareous
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:...
exoskeletons, these do not mineralize until the zooids are fully grown. Colony lifespans range from one to about 12 years, and the short-lived species pass though several generations in one season.
Species that produce defensive zooids do so only when threats have already appeared, and may do so within 48 hours. The theory of "induced defenses" suggests that production of defenses is expensive and that colonies which defend themselves too early or too heavily will have reduced growth rates and lifespans. This "last minute" approach to defense is feasible because the loss of zooids to a single attack is unlikely to be significant. Colonies of some encrusting species also produce special heterozooids to limit the expansion of other encrusting organisms, especially other bryozoans. In some cases this response is more belligerent if the opposition is smaller, which suggests that zooids on the edge of a colony can somehow sense the size of the opponent. Some species consistently prevail against certain others, but most turf war
Turf war
According to Wordnet the definition of a turf war is "a bitter struggle for territory or power or control or rights". For example: a turf war erupted between street gangs; the president's resignation was the result of a turf war with the board of directors. In larger companies Turf wars could...
s are indecisive and the combatants soon turn to growing in uncontested areas. Bryozoans competing for territory do not use the sophisticated techniques employed by sponges or coral
Coral
Corals are marine animals in class Anthozoa of phylum Cnidaria typically living in compact colonies of many identical individual "polyps". The group includes the important reef builders that inhabit tropical oceans and secrete calcium carbonate to form a hard skeleton.A coral "head" is a colony of...
s, possibly because the shortness of bryozoan lifespans makes heavy investment in turf wars unprofitable.
Taxonomy and evolution
The phylum was originally called "Polyzoa", but this name was soon replaced by Ehrenberg's term "Bryozoa". The name "Bryozoa" was originally applied only to the animals also known as "Ectoprocta", in which the anusAnus
The anus 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, unwanted semi-solid matter produced during digestion, which, depending on the type of animal, may be one or more of: matter which the animal cannot digest,...
lies outside the "crown" of tentacles (based on the Ancient Greek
Ancient Greek
Ancient Greek is the stage of the Greek language in the periods spanning the times c. 9th–6th centuries BC, , c. 5th–4th centuries BC , and the c. 3rd century BC – 6th century AD of ancient Greece and the ancient world; being predated in the 2nd millennium BC by Mycenaean Greek...
prefix ἐκτο meaning "outside" and word πρωκτος meaning "anus"). After the discovery of the Entoprocta
Entoprocta
Entoprocta, whose name means "anus inside", is a phylum of mostly sessile aquatic animals, ranging from long. Mature individuals are goblet-shaped, on relatively long stalks. They have a "crown" of solid tentacles whose cilia generate water currents that draw food particles towards the mouth, and...
, in which the anus lies within a "crown" of tentacles (based on the Ancient Greek prefix ἐντο meaning "inside" and word πρωκτος meaning "anus"), the name "Bryozoa" was used at phylum
Phylum
In biology, a phylum The term was coined by Georges Cuvier from Greek φῦλον phylon, "race, stock," related to φυλή phyle, "tribe, clan." is a taxonomic rank below kingdom and above class. "Phylum" is equivalent to the botanical term division....
level to include the two classes
Class (biology)
In biological classification, class is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, order, family, genus, and species, with class fitting between phylum and order...
Ectoprocta and Entoprocta. However, in 1869 Hinrich Nitsche regarded the two groups as quite distinct for a variety of reasons, and coined the name "Ectoprocta" for Ehrenberg's "Bryozoa". Despite their apparently similar methods of feeding, they differed markedly anatomically; in addition to the different positions of the anus, ectoprocts have hollow tentacles and a coelom
Coelom
The coelom is a fluid-filled cavity formed within the mesoderm. Coeloms developed in triploblasts but were subsequently lost in several lineages. Loss of coelom is correlated with reduction in body size...
, while entoprocts have solid tentacles and no coelom. Hence the two groups are now widely regarded as separate phyla, and the name "Bryozoa" is now synonymous with "Ectoprocta". This has remained the majority view ever since, although most publications have preferred the name "Bryozoa" rather than "Ectoprocta". Nevertheless some notable scientists have continued to regard the "Ectoprocta" and Entoprocta as close relatives and group them under "Bryozoa".
The ambiguity about the scope of the name "Bryozoa" led to proposals in the 1960s and 1970s that it should be avoided and the unambiguous term "Ectoprocta" should be used. However, the change would have made it harder to find older works about in which the phylum was called "Bryozoa", and the desire to avoid ambiguity, if applied consistently to all classifications, would have necessitated renaming of several other phyla and many lower-level groups. In practice, zoological naming of split or merged groups of animals is complex and not completely consistent. Works since 2000 have used various names to resolve the ambiguity, including: "Bryozoa", "Ectoprocta", "Bryozoa (Ectoprocta)", and "Ectoprocta (Bryozoa)". Some have used more than one approach in the same work.
The common name "moss animals" is based on the Greek βρυόν (moss) and ζῷα (animals), and refers to the mossy appearance of encrusting species.
Classification and diversity
Counts of formally described species range between 4,000 and 4,500. The Gymnolaemata and especially Cheilostomata have the greatest numbers of species, possibly because of their wide range of specialist zooids. Under the Linnaean system of classificationLinnaean taxonomy
Linnaean taxonomy can mean either of two related concepts:# the particular form of biological classification set up by Carl Linnaeus, as set forth in his Systema Naturæ and subsequent works...
, which is still used as a convenient way to label groups of organisms, living members of the phylum
Phylum
In biology, a phylum The term was coined by Georges Cuvier from Greek φῦλον phylon, "race, stock," related to φυλή phyle, "tribe, clan." is a taxonomic rank below kingdom and above class. "Phylum" is equivalent to the botanical term division....
Bryozoa are divided into:
| Class Class (biology) In biological classification, class is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, order, family, genus, and species, with class fitting between phylum and order... | Phylactolaemata Phylactolaemata Phylactolaemata is a class of the phylum Bryozoa whose members live only in freshwater environments. Like all bryozoans, they filter feed by means of an extensible "crown" of ciliated tentacles called a lophophore. They live in colonies, each of which consists of clones of the founding member... | Stenolaemata Stenolaemata Stenolaemata are a class of marine bryozoans with tubular with strongly calcified walls. They are characterized by a lophophore which is protruded by the action of annular muscles. Most forms lack an operculum.... | | Gymnolaemata Gymnolaemata Gymnolaemata is a class of the phylum Bryozoa. Gymnolaemata typically live under seawater and grow on surfaces of rocks, kelps, and even in some cases, on animal species like fish. They are mostly marine bryozoans with cylindrical or flattened zooids. The lophophore is protruded by action of... |
|
|---|---|---|---|---|
| Order Order (biology) In scientific classification used in biology, the order is# a taxonomic rank used in the classification of organisms. Other well-known ranks are life, domain, kingdom, phylum, class, family, genus, and species, with order fitting in between class and family... | Plumatellida | Cyclostomata Cyclostomatida Cyclostomatida, or cyclostomes, are an ancient order of stenolaemate bryozoans which first appeared in the Lower Ordovician. It consists of 7+ suborders, 59+ families, 373+ genera, and 666+ species. The cyclostome bryozoans were dominant in the Mesozoic; since that era, they have decreased... | Ctenostomata Ctenostomata The Ctenostomata are an order of bryozoans in the class Gymnolaemata. The great majority of ctenostome species are marine, although Paludicella inhabits freshwater.... | Cheilostomata Cheilostomata Cheilostomata, an order of Bryozoa in the class Gymnolaemata, are exclusively marine, colonial invertebrate animals. Cheilostome colonies are composed of calcium carbonate and grow on a variety of surfaces, including rocks, shells, seagrass and kelps. The colony shapes range from simple encrusting... |
| Environments | Freshwater Freshwater Fresh water is naturally occurring water on the Earth's surface in ice sheets, ice caps, glaciers, bogs, ponds, lakes, rivers and streams, and underground as groundwater in aquifers and underground streams. Fresh water is generally characterized by having low concentrations of dissolved salts and... |
Marine | Mostly marine | |
| Lip-like epistome overhanging mouth | Yes | none | ||
| Colony shapes | Gelatinous masses or tubular branching structures | Erect or encrusting | Erect, encrusting or free-living | |
| Exoskeleton material | Gelatinous or membranous; unmineralized | Mineralized | Chitin Chitin Chitin n is a long-chain polymer of a N-acetylglucosamine, a derivative of glucose, and is found in many places throughout the natural world... , gelatinous or membranous; unmineralized |
Mineralized |
| Operculum ("lid") | none | none (except in family Eleidae) | None in most species | Yes (except in genus Bugula Bugula Spiral tufted bryozoa or Bugula turrita are a very common colonial marine animal found from Maine to North Carolina.Bugula neritina is of current interest as a source of cytotoxic chemicals, bryostatins, under clinical investigation as anti-cancer agents.Dried Bugula are commonly used as... ) |
| Shape of lophophore | U-shaped appearance (except in genus Genus In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia... Fredericella, whose lophophore is circular) |
Circular | ||
| How lophophore extended | Compressing the whole body wall | Compressing the membranous sac (separate inner layer of epithelium Epithelium Epithelium is one of the four basic types of animal tissue, along with connective tissue, muscle tissue and nervous tissue. Epithelial tissues line the cavities and surfaces of structures throughout the body, and also form many glands. Functions of epithelial cells include secretion, selective... that lines the coelom) |
Compressing the whole body wall | Pulling inwards of a flexible section of body wall, or making an internal sac expand. |
| Types of zooid | Autozooids only | Limited heterozooids, mainly gonozooids | Stolon Stolon In biology, stolons are horizontal connections between organisms. They may be part of the organism, or of its skeleton; typically, animal stolons are external skeletons.-In botany:... s and spines as well as autozooids |
Full range of types |
Fossil record
Ordovician
The Ordovician is a geologic period and system, the second of six of the Paleozoic Era, and covers the time between 488.3±1.7 to 443.7±1.5 million years ago . It follows the Cambrian Period and is followed by the Silurian Period...
period, about , all the modern orders
Order (biology)
In scientific classification used in biology, the order is# a taxonomic rank used in the classification of organisms. Other well-known ranks are life, domain, kingdom, phylum, class, family, genus, and species, with order fitting in between class and family...
of stenolaemates
Stenolaemata
Stenolaemata are a class of marine bryozoans with tubular with strongly calcified walls. They are characterized by a lophophore which is protruded by the action of annular muscles. Most forms lack an operculum....
were present, and the ctenostome
Ctenostomata
The Ctenostomata are an order of bryozoans in the class Gymnolaemata. The great majority of ctenostome species are marine, although Paludicella inhabits freshwater....
order of gymnolaemates
Gymnolaemata
Gymnolaemata is a class of the phylum Bryozoa. Gymnolaemata typically live under seawater and grow on surfaces of rocks, kelps, and even in some cases, on animal species like fish. They are mostly marine bryozoans with cylindrical or flattened zooids. The lophophore is protruded by action of...
had appeared by the Middle Ordovician, about . The Early Ordovician fossils may also represent forms that had already become significantly different from the original members of the phylum. Other types of filter feeder
Filter feeder
Filter feeders are animals that feed by straining suspended matter and food particles from water, typically by passing the water over a specialized filtering structure. Some animals that use this method of feeding are clams, krill, sponges, baleen whales, and many fish and some sharks. Some birds,...
s appeared around the same time, which suggests that some change made the environment more favorable for this lifestyle. Fossils of cheilostomates
Cheilostomata
Cheilostomata, an order of Bryozoa in the class Gymnolaemata, are exclusively marine, colonial invertebrate animals. Cheilostome colonies are composed of calcium carbonate and grow on a variety of surfaces, including rocks, shells, seagrass and kelps. The colony shapes range from simple encrusting...
, another order of gymnolaemates, first appear in the Mid Jurassic
Jurassic
The Jurassic is a geologic period and system that extends from about Mya to Mya, that is, from the end of the Triassic to the beginning of the Cretaceous. The Jurassic constitutes the middle period of the Mesozoic era, also known as the age of reptiles. The start of the period is marked by...
, about , and these have been the most abundant and diverse bryozoans from the Cretaceous
Cretaceous
The Cretaceous , derived from the Latin "creta" , usually abbreviated K for its German translation Kreide , is a geologic period and system from circa to million years ago. In the geologic timescale, the Cretaceous follows the Jurassic period and is followed by the Paleogene period of the...
to the present. Evidence compiled from the last 100 million years show that cheilostomates consistently grew over cyclostomates in territorial struggles, which may help to explain how cheilostomates replaced cyclostomates as the dominant marine bryozoans. Marine fossils from the Paleozoic
Paleozoic
The Paleozoic era is the earliest of three geologic eras of the Phanerozoic eon, spanning from roughly...
era, which ended , are mainly of erect forms, those from the Mesozoic
Mesozoic
The Mesozoic era is an interval of geological time from about 250 million years ago to about 65 million years ago. It is often referred to as the age of reptiles because reptiles, namely dinosaurs, were the dominant terrestrial and marine vertebrates of the time...
are fairly equally divided by erect and encrusting forms, and more recent ones are predominantly encrusting. Fossils of the soft, freshwater phylactolaemates
Phylactolaemata
Phylactolaemata is a class of the phylum Bryozoa whose members live only in freshwater environments. Like all bryozoans, they filter feed by means of an extensible "crown" of ciliated tentacles called a lophophore. They live in colonies, each of which consists of clones of the founding member...
are very rare, appear in and after the Late Permian (which began about ) and consist entirely of their durable statoblasts. There are no known fossils of freshwater members of other classes.
Since all the other phyla that have left fossils are found in Cambrian
Cambrian
The Cambrian is the first geological period of the Paleozoic Era, lasting from Mya ; it is succeeded by the Ordovician. Its subdivisions, and indeed its base, are somewhat in flux. The period was established by Adam Sedgwick, who named it after Cambria, the Latin name for Wales, where Britain's...
rocks, it is surprising that the earliest bryozoan fossil dates from the Ordovician, which immediately followed the Cambrian. This suggests that the first bryozoans appeared much earlier and were entirely soft-bodied, and the Ordovician fossils record the appearance of mineralized skeletons in this phylum. The Early Ordovician fossils may also represent forms that had already become significantly different from the original members of the phylum.
Evolutionary family tree
Scientists are divided about whether the Bryozoa (Ectoprocta) are a monophyletic group (whether they include all and only a single ancestor species and all its descendants), about what are the phylum's closest relatives in the family tree of animals, and even about whether they should be regarded as members of the protostomeProtostome
Protostomia are a clade of animals. Together with the deuterostomes and a few smaller phyla, they make up the Bilateria, mostly comprising animals with bilateral symmetry and three germ layers...
s or deuterostome
Deuterostome
Deuterostomes are a superphylum of animals. They are a subtaxon of the Bilateria branch of the subregnum Eumetazoa, and are opposed to the protostomes...
s, the two major groups that account for all moderately complex animals.
Class (biology)
In biological classification, class is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, order, family, genus, and species, with class fitting between phylum and order...
Phylactolaemata
Phylactolaemata
Phylactolaemata is a class of the phylum Bryozoa whose members live only in freshwater environments. Like all bryozoans, they filter feed by means of an extensible "crown" of ciliated tentacles called a lophophore. They live in colonies, each of which consists of clones of the founding member...
is most closely related to Stenolaemata
Stenolaemata
Stenolaemata are a class of marine bryozoans with tubular with strongly calcified walls. They are characterized by a lophophore which is protruded by the action of annular muscles. Most forms lack an operculum....
and Ctenostomata
Ctenostomata
The Ctenostomata are an order of bryozoans in the class Gymnolaemata. The great majority of ctenostome species are marine, although Paludicella inhabits freshwater....
, the classes that appear earliest in the fossil record. However, in 2005 a molecular phylogeny
Molecular phylogeny
Molecular phylogenetics is the analysis of hereditary molecular differences, mainly in DNA sequences, to gain information on an organism's evolutionary relationships. The result of a molecular phylogenetic analysis is expressed in a phylogenetic tree...
study that focused on phylactolaemates concluded that these are more closely related to the phylum Phoronid
Phoronid
Phoronids are a phylum of marine animals that filter-feed with a lophophore , and build upright tubes of chitin to support and protect their soft bodies. They live in all the oceans and seas including the Arctic Ocean but excluding the Antarctic Ocean, and between the intertidal zone and about...
a, and especially to the only phoronid species that is colonial, than they are to the other ectoproct classes. That implies that the Entoprocta are not monophyletic, as the Phoronida are a sub-group of ectoprocts but the standard definition of Entoprocta excludes the Phoronida. In 2009 another molecular phylogeny
Molecular phylogeny
Molecular phylogenetics is the analysis of hereditary molecular differences, mainly in DNA sequences, to gain information on an organism's evolutionary relationships. The result of a molecular phylogenetic analysis is expressed in a phylogenetic tree...
study, using a combination of genes from mitochondria and the cell nucleus
Cell nucleus
In cell biology, the nucleus is a membrane-enclosed organelle found in eukaryotic cells. It contains most of the cell's genetic material, organized as multiple long linear DNA molecules in complex with a large variety of proteins, such as histones, to form chromosomes. The genes within these...
, concluded that Bryozoa is a monophyletic phylum, in other words includes all the descendants of a common ancestor that is itself a bryozoan. The analysis also concluded that the classes
Class (biology)
In biological classification, class is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, order, family, genus, and species, with class fitting between phylum and order...
Phylactolaemata
Phylactolaemata
Phylactolaemata is a class of the phylum Bryozoa whose members live only in freshwater environments. Like all bryozoans, they filter feed by means of an extensible "crown" of ciliated tentacles called a lophophore. They live in colonies, each of which consists of clones of the founding member...
, Stenolaemata
Stenolaemata
Stenolaemata are a class of marine bryozoans with tubular with strongly calcified walls. They are characterized by a lophophore which is protruded by the action of annular muscles. Most forms lack an operculum....
and Gymnolaemata
Gymnolaemata
Gymnolaemata is a class of the phylum Bryozoa. Gymnolaemata typically live under seawater and grow on surfaces of rocks, kelps, and even in some cases, on animal species like fish. They are mostly marine bryozoans with cylindrical or flattened zooids. The lophophore is protruded by action of...
are also monophyletic, but could not determine whether Stenolaemata
Stenolaemata
Stenolaemata are a class of marine bryozoans with tubular with strongly calcified walls. They are characterized by a lophophore which is protruded by the action of annular muscles. Most forms lack an operculum....
are more closely related to Phylactolaemata or Gymnolaemata
Gymnolaemata
Gymnolaemata is a class of the phylum Bryozoa. Gymnolaemata typically live under seawater and grow on surfaces of rocks, kelps, and even in some cases, on animal species like fish. They are mostly marine bryozoans with cylindrical or flattened zooids. The lophophore is protruded by action of...
. The Gymnolaemata are traditionally divided into the soft-bodied Ctenostomata
Ctenostomata
The Ctenostomata are an order of bryozoans in the class Gymnolaemata. The great majority of ctenostome species are marine, although Paludicella inhabits freshwater....
and mineralized Cheilostomata, but the 2009 analysis considered it more likely that neither of these orders
Order (biology)
In scientific classification used in biology, the order is# a taxonomic rank used in the classification of organisms. Other well-known ranks are life, domain, kingdom, phylum, class, family, genus, and species, with order fitting in between class and family...
is monophyletic and that mineralized skeleton
Skeleton
The skeleton is the body part that forms the supporting structure of an organism. There are two different skeletal types: the exoskeleton, which is the stable outer shell of an organism, and the endoskeleton, which forms the support structure inside the body.In a figurative sense, skeleton can...
s probably evolved more than once within the early Gymnolaemata.
Bryozoans' relationships with other phyla are uncertain and controversial. Traditional phylogeny, based on anatomy
Anatomy
Anatomy is a branch of biology and medicine that is the consideration of the structure of living things. It is a general term that includes human anatomy, animal anatomy , and plant anatomy...
and on the development of the adult forms from embryo
Embryo
An embryo is a multicellular diploid eukaryote in its earliest stage of development, from the time of first cell division until birth, hatching, or germination...
s, has produced no enduring consensus about the position of ectoprocts. Attempts to reconstruct the family tree of animals have largely ignored ectoprocts and other "minor phyla", which have received little scientific study because they are generally tiny, have relatively simple body plans, and have little impact on human economies – despite the fact that the "minor phyla" include most of the variety in the evolutionary history of animals. In the opinion of Ruth Dewel, Judith Winston and Frank McKinney, "Our standard interpretation of bryozoan morphology
Morphology (biology)
In biology, morphology is a branch of bioscience dealing with the study of the form and structure of organisms and their specific structural features....
and embryology
Embryology
Embryology is a science which is about the development of an embryo from the fertilization of the ovum to the fetus stage...
is a construct resulting from over 100 years of attempts to synthesize a single framework for all invertebrates," and takes little account of some peculiar features of ectoprocts. In ectoprocts, all of the larva's internal organs are destroyed during the metamorphosis to the adult form and the adult's organs are built from the larva's epidermis and mesoderm
Mesoderm
In all bilaterian animals, the mesoderm is one of the three primary germ cell layers in the very early embryo. The other two layers are the ectoderm and endoderm , with the mesoderm as the middle layer between them.The mesoderm forms mesenchyme , mesothelium, non-epithelial blood corpuscles and...
, while in other bilateria
Bilateria
The bilateria are all animals having a bilateral symmetry, i.e. they have a front and a back end, as well as an upside and downside. Radially symmetrical animals like jellyfish have a topside and downside, but no front and back...
ns some organs including the gut are built from endoderm
Endoderm
Endoderm is one of the three primary germ cell layers in the very early embryo. The other two layers are the ectoderm and mesoderm , with the endoderm as the intermost layer...
. In most bilaterian embryos the blastopore, a dent in the outer wall, deepens to become the larva's gut, but in ectoprocts the blastopore disappears and a new dent becomes the point from which the gut grows. The ectoproct coelom is formed by neither of the processes used by other bilaterians, enterocoely
Enterocoely
Enterocoely is a process by which some animal embryos develop. In enterocoely, a mesoderm is formed in a developing embryo, in which the coelom forms from pouches "pinched" off of the digestive tract...
, in which pouches that form on the wall of the gut become separate cavities, nor schizocoely, in which the tissue between the gut and the body wall splits, forming paired cavities.
Molecular phylogeny, which attempts to work out the evolutionary family tree of organisms by comparing their biochemistry
Biochemistry
Biochemistry, sometimes called biological chemistry, is the study of chemical processes in living organisms, including, but not limited to, living matter. Biochemistry governs all living organisms and living processes...
and especially their gene
Gene
A gene is a molecular unit of heredity of a living organism. It is a name given to some stretches of DNA and RNA that code for a type of protein or for an RNA chain that has a function in the organism. Living beings depend on genes, as they specify all proteins and functional RNA chains...
s, has done much to clarify the relationships between the better-known invertebrate
Invertebrate
An invertebrate is an animal without a backbone. The group includes 97% of all animal species – all animals except those in the chordate subphylum Vertebrata .Invertebrates form a paraphyletic group...
phyla. However, the shortage of genetic data about "minor phyla" such as bryozoans and entoprocts
Entoprocta
Entoprocta, whose name means "anus inside", is a phylum of mostly sessile aquatic animals, ranging from long. Mature individuals are goblet-shaped, on relatively long stalks. They have a "crown" of solid tentacles whose cilia generate water currents that draw food particles towards the mouth, and...
has left their relationships to other groups unclear.
When entoprocts were discovered in the 19th century, they and bryozoans (ectoprocts) were regarded as classes within the phylum Bryozoa, because both groups were sessile
Sessility (zoology)
In zoology, sessility is a characteristic of animals which are not able to move about. They are usually permanently attached to a solid substrate of some kind, such as a part of a plant or dead tree trunk, a rock, or the hull of a ship in the case of barnacles. Corals lay down their own...
animals that filter-fed
Filter feeder
Filter feeders are animals that feed by straining suspended matter and food particles from water, typically by passing the water over a specialized filtering structure. Some animals that use this method of feeding are clams, krill, sponges, baleen whales, and many fish and some sharks. Some birds,...
by means of a crown of tentacles that bore cilia. However, from 1869 onwards increasing awareness of differences, including the position of the entoproct anus
Anus
The anus 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, unwanted semi-solid matter produced during digestion, which, depending on the type of animal, may be one or more of: matter which the animal cannot digest,...
inside the feeding structure and the difference in the early pattern of division
Cleavage (embryo)
In embryology, cleavage is the division of cells in the early embryo. The zygotes of many species undergo rapid cell cycles with no significant growth, producing a cluster of cells the same size as the original zygote. The different cells derived from cleavage are called blastomeres and form a...
of cells in their embryo
Embryo
An embryo is a multicellular diploid eukaryote in its earliest stage of development, from the time of first cell division until birth, hatching, or germination...
s, caused scientists to regard the two groups as separate phyla, and "Bryozoa" became just an alternative name for ectoprocts, in which the anus is outside the feeding organ. A series of molecular phylogeny studies from 1996 to 2006 have also concluded that bryozoans (ectoprocts) and entoprocts are not sister groups. However, two well-known zoologists, Claus Nielsen and Thomas Cavalier-Smith
Thomas Cavalier-Smith
Professor Thomas Cavalier-Smith , FRS, FRSC, NERC Professorial Fellow, is a Professor of Evolutionary Biology in the Department of Zoology, at the University of Oxford...
, maintain on anatomical and developmental grounds that bryozoans and entoprocts are member of the same phylum, Bryozoa. A molecular phylogeny study in 2007 also supported this old idea, while its conclusions about other phyla agreed with those of several other analyses.
By 1891 bryozoans (ectoprocts) were grouped with phoronid
Phoronid
Phoronids are a phylum of marine animals that filter-feed with a lophophore , and build upright tubes of chitin to support and protect their soft bodies. They live in all the oceans and seas including the Arctic Ocean but excluding the Antarctic Ocean, and between the intertidal zone and about...
s in a super-phylum called "Tentaculata". In the 1970s comparisons between phoronid larvae and the cyphonautes larva of some gymnolaete bryozoans produced suggestions that the bryozoans, most of which are colonial, evolved from a semi-colonial species of phoronid. Brachiopod
Brachiopod
Brachiopods are a phylum of marine animals that have hard "valves" on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear end, while the front can be opened for feeding or closed for protection...
s were also assigned to the "Tentaculata", which were renamed Lophophorata as they all use a lophophore
Lophophore
The lophophore is a characteristic feeding organ possessed by four major groups of animals: the Brachiopoda, Bryozoa, Entoprocta, and Phoronida. All lophophores are found in aquatic organisms.-Characteristics:...
for filter feeding. Although the majority of scientists accept this, Claus Nielsen thinks these similarities are superficial. The Lophophorata are usually defined as animals with a lophophore, a three-part coelom and a U-shaped gut. In Nielsen's opinion, phoronids' and brachiopods' lophophores are more like those of pterobranchs, which are members of the phylum Hemichordata
Hemichordata
Hemichordata is a phylum of marine deuterostome animals, generally considered the sister group of the echinoderms. They date back to the Lower or Middle Cambrian and include two main classes: Enteropneusta , and Pterobranchia. A third class, Planctosphaeroidea, is known only from the larva of a...
. Bryozoan's tentacles bear cells with multiple cilia, while the corresponding cells of phoronids', brachiopods' and pterobranchs' lophophores have one cilium per cell; and bryozoan tentacles have no hemal canal ("blood vessel"), which those of the other three phyla have.
If the grouping of bryozoans with phoronids and brachiopods into Lophophorata is correct, the next issue is whether the Lophophorata are protostome
Protostome
Protostomia are a clade of animals. Together with the deuterostomes and a few smaller phyla, they make up the Bilateria, mostly comprising animals with bilateral symmetry and three germ layers...
s, along with most invertebrate phyla, or deuterostome
Deuterostome
Deuterostomes are a superphylum of animals. They are a subtaxon of the Bilateria branch of the subregnum Eumetazoa, and are opposed to the protostomes...
s, along with chordate
Chordate
Chordates are animals which are either vertebrates or one of several closely related invertebrates. They are united by having, for at least some period of their life cycle, a notochord, a hollow dorsal nerve cord, pharyngeal slits, an endostyle, and a post-anal tail...
s, hemichordates and echinoderm
Echinoderm
Echinoderms are a phylum of marine animals. Echinoderms are found at every ocean depth, from the intertidal zone to the abyssal zone....
s. The traditional view was that lophophorates were a mix of protostome and deuterostome features. Research from the 1970s onwards suggested they were deuterostomes, because of some features that were thought characteristic of deuterostomes: a three-part coelom; radial rather than spiral cleavage in the development of the embryo; and formation of the coelom by enterocoely
Enterocoely
Enterocoely is a process by which some animal embryos develop. In enterocoely, a mesoderm is formed in a developing embryo, in which the coelom forms from pouches "pinched" off of the digestive tract...
. However the coelom of ectoproct larvae shows no sign of division into three sections, and that of adult ectoprocts is different from that of other coelomate phyla as it is built anew from epidermis and mesoderm after metamorphosis has destroyed the larval coelom.
Molecular phylogeny analyses from 1995 onwards, using a variety of biochemical evidence and analytical techniques, placed the lophophorates as protostomes and closely related to annelid
Annelid
The annelids , formally called Annelida , are a large phylum of segmented worms, with over 17,000 modern species including ragworms, earthworms and leeches...
s and molluscs in a super-phylum called Lophotrochozoa
Lophotrochozoa
The Lophotrochozoa are a major grouping of protostome animals. The taxon was discovered based on molecular data. Molecular evidence such as a result of studies of the evolution of small-subunit ribosomal RNA supports the monophyly of the phyla listed in the infobox shown at right.-Terminology:The...
. "Total evidence" analyses, which used both morphological features and a relatively small set of genes, came to various conclusions, mostly favoring a close relationship between lophophorates and Lophotrochozoa. A study in 2008, using a larger set of genes, concluded that the lophophorates were closer to the Lophotrochozoa than to deuterostomes, but also that the lophophorates were not monophyletic. Instead, it concluded that brachiopods and phoronids formed a monophyletic group, but bryozoans (ectoprocts) were closest to entoprocts, supporting the original definition of "Bryozoa".
Feeding and excretion
Most species are filter feederFilter feeder
Filter feeders are animals that feed by straining suspended matter and food particles from water, typically by passing the water over a specialized filtering structure. Some animals that use this method of feeding are clams, krill, sponges, baleen whales, and many fish and some sharks. Some birds,...
s that sieve small particles, mainly phytoplankton
Phytoplankton
Phytoplankton are the autotrophic component of the plankton community. The name comes from the Greek words φυτόν , meaning "plant", and πλαγκτός , meaning "wanderer" or "drifter". Most phytoplankton are too small to be individually seen with the unaided eye...
(microscopic floating plants), out of the water. The freshwater species Plumatella emarginata feeds on diatom
Diatom
Diatoms are a major group of algae, and are one of the most common types of phytoplankton. Most diatoms are unicellular, although they can exist as colonies in the shape of filaments or ribbons , fans , zigzags , or stellate colonies . Diatoms are producers within the food chain...
s, green algae
Green algae
The green algae are the large group of algae from which the embryophytes emerged. As such, they form a paraphyletic group, although the group including both green algae and embryophytes is monophyletic...
, cyanobacteria, non-photosynthetic bacteria
Bacteria
Bacteria are a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria have a wide range of shapes, ranging from spheres to rods and spirals...
, dinoflagellate
Dinoflagellate
The dinoflagellates are a large group of flagellate protists. Most are marine plankton, but they are common in fresh water habitats as well. Their populations are distributed depending on temperature, salinity, or depth...
s, rotifer
Rotifer
The rotifers make up a phylum of microscopic and near-microscopic pseudocoelomate animals. They were first described by Rev. John Harris in 1696, and other forms were described by Anton van Leeuwenhoek in 1703...
s, protozoa
Protozoa
Protozoa are a diverse group of single-cells eukaryotic organisms, many of which are motile. Throughout history, protozoa have been defined as single-cell protists with animal-like behavior, e.g., movement...
, small nematode
Nematode
The nematodes or roundworms are the most diverse phylum of pseudocoelomates, and one of the most diverse of all animals. Nematode species are very difficult to distinguish; over 28,000 have been described, of which over 16,000 are parasitic. It has been estimated that the total number of nematode...
s, and microscopic crustacean
Crustacean
Crustaceans form a very large group of arthropods, usually treated as a subphylum, which includes such familiar animals as crabs, lobsters, crayfish, shrimp, krill and barnacles. The 50,000 described species range in size from Stygotantulus stocki at , to the Japanese spider crab with a leg span...
s. While the currents that bryozoans generate to draw food towards the mouth are well understood, the exact method of capture is still debated. All species also flick larger particles towards the mouth with a tentacle, and a few capture zooplankton
Zooplankton
Zooplankton are heterotrophic plankton. Plankton are organisms drifting in oceans, seas, and bodies of fresh water. The word "zooplankton" is derived from the Greek zoon , meaning "animal", and , meaning "wanderer" or "drifter"...
(planktonic animals) by using their tentacles as cages. In addition the tentacles, whose surface area is increased by microvilli (small hairs and pleats), absorb organic compound
Organic compound
An organic compound is any member of a large class of gaseous, liquid, or solid chemical compounds whose molecules contain carbon. For historical reasons discussed below, a few types of carbon-containing compounds such as carbides, carbonates, simple oxides of carbon, and cyanides, as well as the...
s dissolved in the water. Unwanted particles may be flicked away by tentacles or shut out by closing the mouth. A study in 2008 showed that both encrusting and erect colonies fed more quickly and grew faster in gentle than in strong currents.
In some species the first part of the stomach forms a muscular gizzard
Gizzard
The gizzard, also referred to as the ventriculus, gastric mill, and gigerium, is an organ found in the digestive tract of some animals, including birds, reptiles, earthworms and some fish. This specialized stomach constructed of thick, muscular walls is used for grinding up food; often rocks are...
lined with chitin
Chitin
Chitin n is a long-chain polymer of a N-acetylglucosamine, a derivative of glucose, and is found in many places throughout the natural world...
ous teeth that crush armored prey such as diatom
Diatom
Diatoms are a major group of algae, and are one of the most common types of phytoplankton. Most diatoms are unicellular, although they can exist as colonies in the shape of filaments or ribbons , fans , zigzags , or stellate colonies . Diatoms are producers within the food chain...
s. Wave-like peristaltic
Peristalsis
Peristalsis is a radially symmetrical contraction and relaxation of muscles which propagates in a wave down the muscular tube, in an anterograde fashion. In humans, peristalsis is found in the contraction of smooth muscles to propel contents through the digestive tract. Earthworms use a similar...
contractions move the food through the stomach for digestion. The final section of the stomach is lined with cilia (minute hairs) that compress undigested solids, which then pass through the intestine
Intestine
In human anatomy, the intestine is the segment of the alimentary canal extending from the pyloric sphincter of the stomach to the anus and, in humans and other mammals, consists of two segments, the small intestine and the large intestine...
and out through the anus
Anus
The anus 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, unwanted semi-solid matter produced during digestion, which, depending on the type of animal, may be one or more of: matter which the animal cannot digest,...
.
There are no nephridia ("little kidneys") or other excretory
Excretory system
The excretory system is a passive biological system that removes excess, unnecessary or dangerous materials from an organism, so as to help maintain homeostasis within the organism and prevent damage to the body. It is responsible for the elimination of the waste products of metabolism as well as...
organs in bryzoa, and it is thought that ammonia
Ammonia
Ammonia is a compound of nitrogen and hydrogen with the formula . It is a colourless gas with a characteristic pungent odour. Ammonia contributes significantly to the nutritional needs of terrestrial organisms by serving as a precursor to food and fertilizers. Ammonia, either directly or...
diffuses out through the body wall and lophophore. More complex waste products are not excreted but accumulate in the polypide
Polypide
The polypide in bryozoans encompasses most of the organs and tissues of each individual zooid. This includes the tentacles, tentacle sheath, U-shaped digestive tract, musculature and nerve cells. It is housed in the zooidal skeleton, which in cyclostomes is tubular and in cheilostomes is box-shaped....
, which degenerates after a few weeks. Some of the old polypide is recycled, but much of it remains as a large mass of dying cells containing accumulated wastes, and this is compressed into a "brown body". When the degeneration is complete, the cystid (outer part of the animal) produces a new polypide, and the brown body remains in the coelom
Coelom
The coelom is a fluid-filled cavity formed within the mesoderm. Coeloms developed in triploblasts but were subsequently lost in several lineages. Loss of coelom is correlated with reduction in body size...
, or in the stomach of the new polypide and is expelled next time the animal defecates.
Respiration and circulation
There are no respiratory organs, heartHeart
The heart is a myogenic muscular organ found in all animals with a circulatory system , that is responsible for pumping blood throughout the blood vessels by repeated, rhythmic contractions...
or blood vessel
Blood vessel
The blood vessels are the part of the circulatory system that transports blood throughout the body. There are three major types of blood vessels: the arteries, which carry the blood away from the heart; the capillaries, which enable the actual exchange of water and chemicals between the blood and...
s. Instead zooids absorb oxygen
Oxygen
Oxygen is the element with atomic number 8 and represented by the symbol O. Its name derives from the Greek roots ὀξύς and -γενής , because at the time of naming, it was mistakenly thought that all acids required oxygen in their composition...
and eliminate carbon dioxide
Carbon dioxide
Carbon dioxide is a naturally occurring chemical compound composed of two oxygen atoms covalently bonded to a single carbon atom...
through the body wall and especially the lophophore. The fluid in the coelom transports gases and nutrients and its circulation is passive, except that some relatively large species use cilia to boost its speed. The different bryozoan groups use various methods to share nutrients and oxygen between zooids: some have quite large gaps in the body walls, allowing the coelomic fluid to circulate freely; in others the funiculi (internal "little ropes") of adjacent zooids connect via small pores in the body wall.
Reproduction and life cycles
Zooids of all phylactolaemate species are simultaneous hermaphroditeHermaphrodite
In biology, a hermaphrodite is an organism that has reproductive organs normally associated with both male and female sexes.Many taxonomic groups of animals do not have separate sexes. In these groups, hermaphroditism is a normal condition, enabling a form of sexual reproduction in which both...
s. Although those of many marine species are protandric, in other words function first as males and then as females, their colonies contain a combination of zooids that are in their male and female stages. In all species the ovaries
Ovary
The ovary is an ovum-producing reproductive organ, often found in pairs as part of the vertebrate female reproductive system. Ovaries in anatomically female individuals are analogous to testes in anatomically male individuals, in that they are both gonads and endocrine glands.-Human anatomy:Ovaries...
develop on the inside of the body wall, and the testes on the funiculus connecting the stomach to the body wall. Eggs and sperm are released into the coelom, and sperm exit into the water through pores in the tips of some of the tentacles, and then are captured by the feeding currents of zooids that are producing eggs. Some species' eggs are fertilized externally after being released through a pore between two tentacles, which in some cases is at the tip of a small projection called the "intertentacular organ" in the base of a pair of tentacles. Others' are fertilized internally, in the intertentacular organ or in the coelom. In ctenostomes the mother provides a brood chamber for the fertilized eggs, and her polypide disintegrates, providing nourishment to the embryo
Embryo
An embryo is a multicellular diploid eukaryote in its earliest stage of development, from the time of first cell division until birth, hatching, or germination...
. Stenolaemates produce specialized zooids to serve as brood chambers, and their eggs divide within this to produce up to 100 identical embryos.
The cleavage
Cleavage (embryo)
In embryology, cleavage is the division of cells in the early embryo. The zygotes of many species undergo rapid cell cycles with no significant growth, producing a cluster of cells the same size as the original zygote. The different cells derived from cleavage are called blastomeres and form a...
of bryozoan eggs is biradial, in other words the early stages are bilaterally symmetrical. It is unknown how the coleom forms, since the metamorphosis
Metamorphosis
Metamorphosis is a biological process by which an animal physically develops after birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation...
from larva
Larva
A larva is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle...
to adult destroys all of the larva's internal tissues. In many animals the blastopore
Blastopore
A blastopore is an opening into the archenteron during the embryonic stages of an organism. The distinction between protostomes and deuterostomes is based on the direction in which the mouth develops in relation to the blastopore...
, an opening in the surface of the early embryo, tunnels through to form the gut. However, in bryozoans the blastopore closes, and a new opening develops to create the mouth.
Bryozoan larvae vary in form, but all have a band of cilia round the body which enables them to swim, a tuft of cilia at the top, and an adhesive sac that everts and anchors them when they settle on a surface. Some gymnolaemate species produce cyphonautes larvae which have little yolk but a well-developed mouth and gut, and live as plankton
Plankton
Plankton are any drifting organisms that inhabit the pelagic zone of oceans, seas, or bodies of fresh water. That is, plankton are defined by their ecological niche rather than phylogenetic or taxonomic classification...
for a considerable time before settling. These larvae have triangular shells of chitin, with one corner at the top and the base open, forming a hood round the downward-facing mouth. In 2006 it was reported that the cilia of cyphonautes larvae use the same range of techniques as those of adults to capture food. Species that brood their embryos form larvae that are nourished by large yolks, have no gut and do not feed, and such larvae quickly settle on a surface. In all marine species the larvae produce cocoons in which they metamorphose
Metamorphosis
Metamorphosis is a biological process by which an animal physically develops after birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation...
completely after settling: the larva's epidermis becomes the lining of the coelom
Coelom
The coelom is a fluid-filled cavity formed within the mesoderm. Coeloms developed in triploblasts but were subsequently lost in several lineages. Loss of coelom is correlated with reduction in body size...
, and the internal tissues are converted to a food reserve that nourishes the developing zooid until it is ready to feed. The larvae of phylactolaemates produce multiple polypides, so that each new colony starts with several zooids. In all species the founder zooids then grow the new colonies by budding
Budding
Budding is a form of asexual reproduction in which a new organism grows on another one. The new organism remains attached as it grows, separating from the parent organism only when it is mature. Since the reproduction is asexual, the newly created organism is a clone and is genetically identical...
clone
Cloning
Cloning in biology is the process of producing similar populations of genetically identical individuals that occurs in nature when organisms such as bacteria, insects or plants reproduce asexually. Cloning in biotechnology refers to processes used to create copies of DNA fragments , cells , or...
s of themselves. In phylactolaemates, zooids die after producing several clones, so that living zooids are found only round the edges of a colony.
Phylactolaemates also reproduce asexually by a method that enables a colony's lineage to survive the variable and uncertain conditions of freshwater environments. Throughout summer and autumn they produce disc-shaped statoblasts, masses of cells that function as "survival pods" rather like the gemmules of sponges. Statoblasts form on the funiculus connected to the parent's gut, which nourishes them. As they grow, statoblasts develop protective bivalve-like
Bivalve shell
A bivalve shell is part of the body, the exoskeleton or shell, of a bivalve mollusk. In life, the shell of this class of mollusks is composed of two parts, two valves which are hinged together...
shells made of chitin
Chitin
Chitin n is a long-chain polymer of a N-acetylglucosamine, a derivative of glucose, and is found in many places throughout the natural world...
. When they mature, some statoblasts stick to the parent colony, some fall to the bottom ("sessoblasts"), some contain air spaces that enable them to float ("floatoblasts"), and some remain in the parent's cystid to re-build the colony if it dies. Statoblasts can remain dormant for considerable periods, and while dormant can survive harsh conditions such as freezing and desiccation
Desiccation
Desiccation is the state of extreme dryness, or the process of extreme drying. A desiccant is a hygroscopic substance that induces or sustains such a state in its local vicinity in a moderately sealed container.-Science:...
. They can be transported across long distances by animals, floating vegetation, currents and winds, and even in the guts of larger animals. When conditions improve, the valves of the shell separate and the cells inside develop into a zooid that tries to form a new colony. Plumatella emarginata produces both "sessoblasts", which enable the lineage to control a good territory even if hard times decimate the parent colonies, and "floatoblasts", which spread to new sites. New colonies of Plumatella repens produce mainly "sessoblasts" while mature ones switch to "floatoblasts". A study estimated that one group of colonies in a patch measuring 1 square metres (10.8 sq ft) produced 800,000 statoblasts.
Habitats and distribution
Most marine species live in tropical waters at depths less than 100 metres (328.1 ft). However, a few have been found in deep-sea trenchesOceanic trench
The oceanic trenches are hemispheric-scale long but narrow topographic depressions of the sea floor. They are also the deepest parts of the ocean floor....
, especially around cold seep
Cold seep
A cold seep is an area of the ocean floor where hydrogen sulfide, methane and other hydrocarbon-rich fluid seepage occurs, often in the form of a brine pool...
s, and others near the poles
Geographical pole
A geographical pole is either of the two points—the north pole and the south pole—on the surface of a rotating planet where the axis of rotation meets the surface of the body...
. The great majority are sessile
Sessility (zoology)
In zoology, sessility is a characteristic of animals which are not able to move about. They are usually permanently attached to a solid substrate of some kind, such as a part of a plant or dead tree trunk, a rock, or the hull of a ship in the case of barnacles. Corals lay down their own...
. Encrusting forms are much the commonest of these in shallow seas, but erect forms become more common as the depth increases. A few marine species can move, and an Antarctic
Antarctic
The Antarctic is the region around the Earth's South Pole, opposite the Arctic region around the North Pole. The Antarctic comprises the continent of Antarctica and the ice shelves, waters and island territories in the Southern Ocean situated south of the Antarctic Convergence...
species forms floating colonies.
The phylactolaemates live in all types of freshwater environment – lakes and ponds, rivers and streams, and estuaries
– and are among the most abundant sessile freshwater animals. Some ctenostomes are exclusively freshwater while others prefer brackish water but can survive in freshwater. Scientists' knowledge of freshwater bryozoan populations in many parts of the world is incomplete, even in some parts of Europe. It was long thought that some freshwater species occurred worldwide, but since 2002 all of these have been split into more localized species.
Interactions with non-human organisms
Coral reef
Coral reefs are underwater structures made from calcium carbonate secreted by corals. Coral reefs are colonies of tiny living animals found in marine waters that contain few nutrients. Most coral reefs are built from stony corals, which in turn consist of polyps that cluster in groups. The polyps...
s, but seldom a significant proportion of the total biomass
Biomass
Biomass, as a renewable energy source, is biological material from living, or recently living organisms. As an energy source, biomass can either be used directly, or converted into other energy products such as biofuel....
. In temperate waters, the skeletons of dead colonies form a significant component of shell gravel
Gravel
Gravel is composed of unconsolidated rock fragments that have a general particle size range and include size classes from granule- to boulder-sized fragments. Gravel can be sub-categorized into granule and cobble...
s, and live ones are abundant in these areas. The marine lace-like bryozoan Membranipora membranacea
Membranipora membranacea
Membranipora membranacea is a very widely distributed species of marine bryozoan known from the Atlantic and Pacific Oceans, usually in temperate zone environments. This bryozoan is a colonial organism characterized by a thin, mat-like encrustation, white to gray in color...
produces spines in response to predation by several species of nudibranch
Nudibranch
A nudibranch is a member of what is now a taxonomic clade, and what was previously a suborder, of soft-bodied, marine gastropod mollusks which shed their shell after their larval stage. They are noted for their often extraordinary colors and striking forms...
s ("sea slugs"). Other predators on marine bryozoans include fish, sea urchin
Sea urchin
Sea urchins or urchins are small, spiny, globular animals which, with their close kin, such as sand dollars, constitute the class Echinoidea of the echinoderm phylum. They inhabit all oceans. Their shell, or "test", is round and spiny, typically from across. Common colors include black and dull...
s, pycnogonids, crustacean
Crustacean
Crustaceans form a very large group of arthropods, usually treated as a subphylum, which includes such familiar animals as crabs, lobsters, crayfish, shrimp, krill and barnacles. The 50,000 described species range in size from Stygotantulus stocki at , to the Japanese spider crab with a leg span...
s, mite
Mite
Mites, along with ticks, are small arthropods belonging to the subclass Acari and the class Arachnida. The scientific discipline devoted to the study of ticks and mites is called acarology.-Diversity and systematics:...
s and starfish. In general marine echinoderms and molluscs eat masses of zooids by gouging pieces of colonies, breaking their mineralized "houses", while most arthropod
Arthropod
An arthropod is an invertebrate animal having an exoskeleton , a segmented body, and jointed appendages. Arthropods are members of the phylum Arthropoda , and include the insects, arachnids, crustaceans, and others...
predators on bryozoans eat individual zooids.
In freshwater, bryozoans are among the most important filter feeder
Filter feeder
Filter feeders are animals that feed by straining suspended matter and food particles from water, typically by passing the water over a specialized filtering structure. Some animals that use this method of feeding are clams, krill, sponges, baleen whales, and many fish and some sharks. Some birds,...
s, along with sponges and mussel
Mussel
The common name mussel is used for members of several families of clams or bivalvia mollusca, from saltwater and freshwater habitats. These groups have in common a shell whose outline is elongated and asymmetrical compared with other edible clams, which are often more or less rounded or oval.The...
s. Freshwater bryozoans are attacked by many predators, including snails, insects, and fish.
In Thailand
Thailand
Thailand , officially the Kingdom of Thailand , formerly known as Siam , is a country located at the centre of the Indochina peninsula and Southeast Asia. It is bordered to the north by Burma and Laos, to the east by Laos and Cambodia, to the south by the Gulf of Thailand and Malaysia, and to the...
the introduced species
Introduced species
An introduced species — or neozoon, alien, exotic, non-indigenous, or non-native species, or simply an introduction, is a species living outside its indigenous or native distributional range, and has arrived in an ecosystem or plant community by human activity, either deliberate or accidental...
Pomacea canaliculata
Pomacea canaliculata
Pomacea canaliculata, common name the channeled applesnail, is a species of large freshwater snail with gills and an operculum, an aquatic gastropod mollusk in the family Ampullariidae, the apple snails....
(golden apple snail), which is generally a destructive herbivore
Herbivore
Herbivores are organisms that are anatomically and physiologically adapted to eat plant-based foods. Herbivory is a form of consumption in which an organism principally eats autotrophs such as plants, algae and photosynthesizing bacteria. More generally, organisms that feed on autotrophs in...
, has wiped out phylactolaemate populations wherever it has appeared. P. canaliculata also preys on a common freshwater gymnolaemate, but with less devastating effect. Indigenous snails do not feed on bryozoans.
Several species of the hydroid
Hydroid
-Marine biology:Hydroids are a life stage for most animals of class Hydrozoa, small predators related to jellyfish.-Botany:In mosses, hydroids form the innermost layer of the stem of long, colourless, thin walled cells of small diameter.The cells are dead and lack protoplasm.They function as water...
family Zancleidae have symbiotic relationships with bryozoans, some of which are beneficial to the hydroids while others are parasitic. Modifications appear in the shapes of some these hydroids, for example smaller tentacles or encrustation of the roots by bryozoans. The bryozoan Alcyonidium nodosum protects the whelk
Whelk
Whelk, also spelled welk or even "wilks", is a common name used to mean one or more kinds of sea snail. The species, genera and families referred to using this common name vary a great deal from one geographic area to another...
Burnupena papyracea
Burnupena papyracea
Burnupena papyracea, common name the papery burnupena, is a species of sea snail, a marine gastropod mollusk in the family Buccinidae, the true whelks....
against predation by the powerful and voracious rock lobster
Rock lobster
Jasus edwardsii, the southern rock lobster, red rock lobster, or spiny rock lobster, is a species of spiny lobster found throughout coastal waters of southern Australia and New Zealand including the Chatham Islands. This species is commonly called crayfish or crays in New Zealand and in Māori...
Jasus lalandii
Jasus lalandii
Jasus lalandii is a species of spiny lobster found off the coast of Southern Africa. It is not known whom the specific epithet lalandii commemorates, although it may be the French astronomer Jérôme Lalande.J...
. While whelk shells encrusted by the bryozoans are stronger than those without this reinforcement, chemical defenses produced by the bryozoans are probably the more significant deterrent.
Some phylactolaemate species are parasitized by a group of myxozoa
Myxozoa
The Myxozoa are a group of parasitic animals of aquatic environments. Over 1300 species have been described and many have a two-host lifecycle, involving a fish and an annelid worm or bryozoan. The average size of a Myxosporea spore usually ranges from 10 μm to 20 μm and Malacosporea up...
that have also been found to cause Proliferative Kidney Disease, which is often fatal in salmonid fish, and has severely reduced wild fish populations in Europe and North America.
Membranipora membranacea, whose colonies feed and grow exceptionally fast in a wide range of current speeds, was first noticed in the Gulf of Maine
Gulf of Maine
The Gulf of Maine is a large gulf of the Atlantic Ocean on the east coast of North America.It is delineated by Cape Cod at the eastern tip of Massachusetts in the southwest and Cape Sable at the southern tip of Nova Scotia in the northeast. It includes the entire coastlines of the U.S...
in 1987 and quickly became the most abundant organism living on kelp
Kelp
Kelps are large seaweeds belonging to the brown algae in the order Laminariales. There are about 30 different genera....
s. This invasion
Invasive species
"Invasive species", or invasive exotics, is a nomenclature term and categorization phrase used for flora and fauna, and for specific restoration-preservation processes in native habitats, with several definitions....
reduced the kelp population by breaking their fronds, so that its place as the dominant "vegetation" in some areas was taken by another invader, the large alga Codium fragile
Codium fragile
Codium fragile, also known as Green sea fingers, Dead man's fingers, felty fingers, felt-alga, Green sponge and Green fleece is an invasive species of seaweed in the family Codiaceae....
tomentosoides. These changes reduced the area of habitat available for local fish and invertebrates. M. membranacea has also invaded the northwest coast of the U.S.A. A few freshwater species have been also found thousands of kilometers from their native ranges. Some may have been transported naturally as statoblasts. Others more probably were spread by humans, for example on imported water plants or as stowaways on ships.
Interaction with humans
Fish farms and hatcheries have lost stock to proliferative kidney disease, which is apparently caused by one or more myxozoans that also parasitize bryozoans.Fishermen in the North Sea
North Sea
In the southwest, beyond the Straits of Dover, the North Sea becomes the English Channel connecting to the Atlantic Ocean. In the east, it connects to the Baltic Sea via the Skagerrak and Kattegat, narrow straits that separate Denmark from Norway and Sweden respectively...
have had to find other work because of a form of eczema
Eczema
Eczema is a form of dermatitis, or inflammation of the epidermis . In England, an estimated 5.7 million or about one in every nine people have been diagnosed with the disease by a clinician at some point in their lives.The term eczema is broadly applied to a range of persistent skin conditions...
(a skin disease) known as "Dogger Bank
Dogger Bank
Dogger Bank is a large sandbank in a shallow area of the North Sea about off the east coast of England. It extends over approximately , with its dimensions being about long and up to broad. The water depth ranges from 15 to 36 metres , about shallower than the surrounding sea. It is a...
itch", caused by contact with bryozoans that have stuck to nets and lobster pots.
Marine bryozoans are often responsible for biofouling
Biofouling
Biofouling or biological fouling is the undesirable accumulation of microorganisms, plants, algae, or animals on wetted structures.-Impact:...
on ships' hulls, on docks and marinas, and on offshore structures. They are among the first colonizers of new or recently cleaned structures. Freshwater species are occasional nuisances in water pipes, drinking water purification equipment, sewage treatment facilities, and the cooling pipes of power stations.
A group of chemicals called bryostatin
Bryostatin
Bryostatins are a group of macrolide lactones first isolated in the 1960s by George Pettit from extracts of a species of bryozoan, Bugula neritina. The structure of bryostatin 1 was determined in 1982. Until today 20 different bryostatins have been isolated. Bryostatins are a potent modulators of...
s can be extracted from the marine bryozoan Bugula neritina. In 2001 pharmaceutical company GPC Biotech
GPC Biotech
GPC Biotech is a German biopharmaceutical company. The company's mission statement reads "... to discover, develop and commercialize new anticancer drugs."...
licensed Bryostatin 1 from Arizona State University
Arizona State University
Arizona State University is a public research university located in the Phoenix Metropolitan Area of the State of Arizona...
for commercial development as a treatment for cancer
Cancer
Cancer , known medically as a malignant neoplasm, is a large group of different diseases, all involving unregulated cell growth. In cancer, cells divide and grow uncontrollably, forming malignant tumors, and invade nearby parts of the body. The cancer may also spread to more distant parts of the...
. GPC Biotech canceled development in 2003, saying that Bryostatin 1 showed little effectiveness and some toxic side-effects. In January 2008 a clinical trial
Clinical trial
Clinical trials are a set of procedures in medical research and drug development that are conducted to allow safety and efficacy data to be collected for health interventions...
was submitted to the United States
United States
The United States of America is a federal constitutional republic comprising fifty states and a federal district...
National Institutes of Health
National Institutes of Health
The National Institutes of Health are an agency of the United States Department of Health and Human Services and are the primary agency of the United States government responsible for biomedical and health-related research. Its science and engineering counterpart is the National Science Foundation...
to measure the safety and effectiveness of Bryostatin 1 in the treatment of Alzheimer's Disease
Alzheimer's disease
Alzheimer's disease also known in medical literature as Alzheimer disease is the most common form of dementia. There is no cure for the disease, which worsens as it progresses, and eventually leads to death...
. However, no participants had been recruited by the end of December 2008, when the study was scheduled for completion. About 1 tonnes (157.5 st) of bryozoans must processed to extract 1 gram (0.035273962105112 oz) of bryostatin. As a result, synthetic equivalents have been developed that are simpler to produce and apparently at least as effective.
Further reading
- Hayward, P.G., J.S. Ryland and P.D. Taylor (eds.), 1992. Biology and Palaeobiology of Bryozoans, Olsen and Olsen, Fredensborg, Denmark.
- Robison, R.A. (ed.), 1983. Treatise on Invertebrate PaleontologyTreatise on Invertebrate PaleontologyThe Treatise on Invertebrate Paleontology published by the Geological Society of America and the University of Kansas Press, is a definitive multi-authored work of some 50 volumes, written by more than 300 paleontologists, and covering every phylum, class, order, family, and genus of fossil and...
, Part G, Bryozoa (revised). Geological Society of America and University of Kansas Press. - Woollacott, R.M. and R.L. Zimmer (eds), 1977. The Biology of Bryozoans, Academic Press, New York.
External links
- Index to Bryozoa Bryozoa Home Page, was at RMIT; now bryozoa.net
- Other Bryozoan WWW Resources
- International Bryozoology Association official website
- Bryozoan Introduction
- The Phylum Ectoprocta (Bryozoa)
- Phylum Bryozoa at Wikispecies
- Bryozoans in the Connecticut RiverConnecticut RiverThe Connecticut River is the largest and longest river in New England, and also an American Heritage River. It flows roughly south, starting from the Fourth Connecticut Lake in New Hampshire. After flowing through the remaining Connecticut Lakes and Lake Francis, it defines the border between the...
- Bryozoa Fact Sheet