Bryozoa - Knowledge

1199:, 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 1501:

encrusting, dome-shaped, palmate, foliose, fenestrate, robust branching, delicate branching, articulated and free-living. Most of these sediments come from two distinct groups of colonies: domal, delicate branching, robust branching and palmate; and fenestrate. Fenestrate colonies generate rough particles both as sediment and components of stromatoporoids coral reefs. The delicate colonies however, create both coarse sediment and form the cores of deep-water, subphotic biogenic mounds. Nearly all post- bryozoan sediments are made up of growth forms, with the addition to free-living colonies which include significant numbers of various colonies. "In contrast to the Palaeozoic, post-Palaeozoic bryozoans generated sediment varying more widely with the size of their grains; they grow as they moved from mud, to sand, to gravel."

1535: 3240:. Typically, sessile bryozoans live on hard substrates including rocks, sand or shells. Encrusting forms are much the commonest of these in shallow seas, but erect forms become more common as the depth increases. An example of incrustation on pebbles and cobbles is found in the diverse Pleistocene bryozoans found in northern Japan, where fossils have been found of single stones covered with more than 20 bryozoan species. Sediments with smaller particles, like sand or silt, are usually unsuitable habitat for bryozoans, but tiny colonies have been found encrusting grains of coarse sand. Some bryozoan species specialize in colonizing marine algae, seagrasses, and even mangrove roots; the genus 3258: 3469: 1272:

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

2604:. "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". 2800: 1294:'empty') consist only of the body wall and funicular strands crossing the interior, and no polypide. The functions of these zooids include forming the stems of branching structures, acting as spacers that enable colonies to grow quickly in a new direction, strengthening the colony's branches, and elevating the colony slightly above its substrate for competitive advantages against other organisms. Some kenozooids are hypothesized to be capable of storing nutrients for the colony. Because kenozooids' function is generally structural, they are called "structural polymorphs." 2409: 3254:

evolved similarly shaped colonies capable of movement. When observed in an aquarium, Selenaria maculata colonies were recorded to crawl at a speed of one meter per hour, climb over each other, move toward light, and right themselves when turned upside-down. Later study of this genus showed that neuroelectrical activity in the colonies increased in correlation with movement toward light sources. It is theorized that the capacity for movement arose as a side effect when colonies evolved longer setae for unburying themselves from sediment.

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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

937:, since they are not fully independent animals. All colonies contain feeding zooids, known as autozooids. Those of some groups also contain non-feeding heterozooids, also known as polymorphic zooids, which serve a variety of functions other than feeding; 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. 2111:

from the Ordovician onward. However, unlike corals and other colonial animals found in the fossil record, Bryozoan colonies did not reach large sizes. Fossil bryozoan colonies are typically found highly fragmented and scattered; the preservation of complete zoaria is uncommon in the fossil record, and relatively little study has been devoted to reassembling fragmented zoaria. The largest known fossil colonies are branching trepostome bryozoans from Ordovician rocks in the United States, reaching 66 centimeters in height.

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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.

1945: 2655: 1602:, 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". 3310:

colonies can be formed through sexual reproduction and the generation of free swimming larvae. When colonies grow too large, however, they can split in two. This is the only case where asexual reproduction results in a new colony separate from its predecessor. Most colonies are stationary. Indeed, these colonies tend to be settled on immobile substances such as sediment and coarse substances. There are some colonies of freshwater species such as

2394:, 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. 2683: 1516: 1353: 135: 2219: 3065: 2424: 2088: 2081: 2046: 2011: 1985: 3281:, which is disc-shaped and similarly free-living, inhabit muddy seabeds in the Arctic and can sequester sand grains they have engulfed, potentially using the sand as ballast to turn themselves right-side-up after they have been overturned. Some bryozoan species can form bryoliths, sphere-shaped free-living colonies that grow outward in all directions as they roll about on the seabed. 1452:", 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. 3187:. 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 3277:, consists of floating colonies. The pelagic species is between 5.0 and 23.0 mm (0.20 and 0.91 in) in diameter, has the shape of a hollow sphere and consists of a single layer of autozooids. It is still not known if these colonies are pelagic their whole life or only represents a temporarily and previously undescribed juvenile stage. Colonies of the species 397:", which holds the organs. Zooids have no special excretory organs, and autozooids' polypides are scrapped when they become overloaded with waste products; usually the body wall then grows a replacement polypide. Their gut is U-shaped, with the mouth inside the crown of tentacles and the anus outside it. Zooids of all the freshwater species are simultaneous 3023:, 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 3633:. However, no participants had been recruited by the end of December 2008, when the study was scheduled for completion. More recent work shows it has positive effects on cognition in patients with Alzheimer's disease with few side effects. About 1,000 kilograms (2,200 lb) of bryozoans must be processed to extract 1 gram ( 2498:, 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. 572: in) long. Phoronids resemble bryozoan zooids but are 2 to 20 cm (1 to 8 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 1176:, and further cilia on the inner surfaces of the tentacles move the particles towards the mouth in the center. The method used by ectoprocts is called "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 3301:

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'

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The phylogenetic position of the ectoproct bryozoans remains uncertain, but it remains certain that they belong to the Protostomia and more specifically to the Lophotrochozoa. This implies that the ectoproct larva is a trochophore with the corona being a homologue of the prototroch; this is supported

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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

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Spinozooids are hollow, movable spines, like very slender, small tubes, present on the surface of colonies, which probably are for defense. 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

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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

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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

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has also invaded the northwest coast of the US. 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

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colonies. A larval Bryozoan settles on a hard substance and produces a colony asexually through budding. These colonies can grow thousands of individual zooids in a relatively short period of time. Even though colonies of zooids grow through asexual reproduction, Bryozoans are hermaphrodites and new

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Fossils of about 15,000 bryozoan species have been found. Bryozoans are among the three dominant groups of Paleozoic fossils. Bryozoans with calcitic skeletons were a major source of the carbonate minerals that make up limestones, and their fossils are incredibly common in marine sediments worldwide

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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

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Some freshwater species secrete a mass of gelatinous material, up to 1 m (3 ft 3 in) 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

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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

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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

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They are the only major phylum of exclusively clonal animals, composed of modular units known as zooids. Because they thrive in colonies, colonial growth allows them to develop unrestricted variations in form. Despite this, only a small number of basic growth forms have been found and have commonly

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can move. Lunulitiform cheilostomes are one group of free-living bryozoans with mobile colonies. They form small round colonies un-attached to any substrate; colonies of the genus Selenaria have been observed to "walk" around using setae. Another cheilostome family, the Cupuladriidae, convergently

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Phylactolaemates can 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

3147:, 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 1417:

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 cm (1 ft 8 in) and contain about 2,000,000 zooids. These species generally have exoskeletons reinforced

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and lives on their shells. These zooids are smaller than the others and have four short tentacles and four long tentacles, unlike the autozooids which have 15–16 tentacles. Androzooids are also found in species with mobile colonies that can crawl around. It is possible that androzooids are used to

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Some heterozooids found in extinct trepostome bryozoans, called mesozooids, are thought to have functioned to space the feeding autozooids an appropriate distance apart. In thin sections of trepostome fossils, mesozooids can be seen in between the tubes that held autozooids; they are smaller tubes

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The terms Polyzoa and Bryozoa were introduced in 1830 and 1831, respectively. Soon after it was named, another group of animals was discovered whose filtering mechanism looked similar, so it was included in Bryozoa until 1869, when the two groups were noted to be very different internally. The new

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Occasionally, the impact of an invertebrate invader can be substantial; such has been documented for Membranipora membranacea, a fast-growing invasive bryozoan species which has reduced kelp forests off the northeast and northwest coasts of the USA to an extent that it has affected local fish and

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Until 2008 there were "inadequately known and misunderstood type species belonging to the Cyclostome Bryozoan family Oncousoeciidae." Modern research and experiments have been done using low-vacuum scanning electron microscopy of uncoated type material to critically examine and perhaps revise the

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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 in which the phylum was called "Bryozoa", and the desire to

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Gonozooids act as brood chambers for fertilized eggs. Almost all modern cyclostome bryozoans have them, but they can be hard to locate on a colony because there are so few gonozooids in one colony. The aperture in gonozooids, which is called an ooeciopore, acts as a point for larvae to exit. Some

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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

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The lophophore and mouth are mounted on a flexible tube called the "invert", which 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

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Cupuladriid Bryozoa are capable of both sexual and asexual reproduction. The sexually reproducing colonies (aclonal) are the result of a larval cupuladriid growing into an adult stage whereas the asexual colonies(clonal) are a result of a fragment of a colony of cupuladriids growing into its own

3163:, 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 1597:

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

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All phylactolaemates and stenolaemates, and most gymnolaemates, exhibit placentation, and has therefore lecithotrophic (non-feeding) larvae. Except for Cyclostomata and the small gymnolaemate family Epistomiidae, which are viviparous, all are brooders. Phylactolaemata brood their embryos in an

2455:. 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, 2536:

The majority of scientists accept this, but 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

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Bryozoans have contributed to carbonate sedimentation in marine life since the Ordovician period. Bryozoans take responsibility for many of the colony forms, which have evolved in different taxonomic groups and vary in sediment producing ability. The nine basic bryozoan colony-forms include:

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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

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Planktotrophic (feeding) larvae are only found in class Gymnolaemata: In the cheilostomatan suborder Malacostegina they are found in the two families Membraniporidae and Electridae, and in the three ctenostome families Alcyonidiidae, Farrellidae, and Hislopiidae. In addition there are a few

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to the present. Evidence compiled from the last 100 million years show that cheilostomatids consistently grew over cyclostomatids in territorial struggles, which may help to explain how cheilostomatids replaced cyclostomatids as the dominant marine bryozoans. Marine fossils from the

1382: in) to over 1 m (3 ft 3 in). However, the majority are under 10 cm (4 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 3101:

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.

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living species are known. Originally all of the crown group Bryozoa were colonial, but as an adaptation to a mesopsammal (interstitial spaces in marine sand) life or to deep-sea habitats, secondarily solitary forms have since evolved. Solitary species has been described in four

2305:, 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. 595:

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

3191:. 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. 485:

period, making it the last major phylum to appear in the fossil record. This has led researchers to suspect that bryozoans arose earlier but were initially unmineralized, and may have differed significantly from fossilized and modern forms. In 2021, some research suggested

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resulting in egg-size structures known as bryoliths. Nucleating on an empty gastropod shell, the bryozoan colonies form multilamellar skeletal crusts that produce spherical encrustations and extend the living chamber of the hermit crab through helicospiral tubular growth.

2505:, 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. 988:(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 3302:

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.

1263:) 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 1238:, 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. 1318:

Cheilostome bryozoans also brood their embryos; one of the common methods is through ovicells, capsules attached to autozooids. The autozooids possessing ovicells are normally still able to feed, however, so these are not considered heterozooids.

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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

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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.

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and some polyzoa) or through pseudopores located on the outer dermis of the zooid. 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

2957:. 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 2245:

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

2160:. The Early Ordovician fossils may also represent forms that had already become significantly different from the original members of the phylum. Ctenostomes with phosphatized soft tissue are known from the Devonian. Other types of 8035: 6454:

Jackson, Patrick N. Wyse; Key, Marcus M. Key Jr. (2019). "Epizoan and endoskeletozoan distribution across reassembled ramose stenolaemate bryozoan zoaria from the Upper Ordovician (Katian) of the Cincinnati Arch region, USA".

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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.

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internal brood sac, but Gymnolaemata both external membranous sacs, skeletal chambers (ovicells) and internal brooding sacs exist. The developing embryo relies on egg's yolk, extraembryonic nutrition (matrotrophy) or both.

3092:. 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 1725:

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

2549:, 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. 996:

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.

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gonozooids have very complex shapes with autozooidal tubes passing through chambers within them. All larvae released from a gonozooid are clones created by division of a single egg; this is called monozygotic

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produce mainly "sessoblasts" while mature ones switch to "floatoblasts". A study estimated that one group of colonies in a patch measuring 1 square meter (11 sq ft) produced 800,000 statoblasts.

371:, and supplying nutrients to the colony through diverse channels. Some classes have specialist zooids like hatcheries for fertilized eggs, colonial defence structures, and root-like attachment structures. 8480:

Wender, P.A.; Baryza, JL; Bennett, CE; Bi, FC; Brenner, SE; Clarke, MO; Horan, JC; Kan, C; et al. (20 November 2002). "The Practical Synthesis of a Novel and Highly Potent Analogue of Bryostatin".

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The phylum was originally called "Polyzoa", but this name was eventually replaced by Ehrenberg's term "Bryozoa". The name "Bryozoa" was originally applied only to the animals also known as Ectoprocta (

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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 through several generations in one season.

401:. 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 6161: 1245:

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.

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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

1168:) 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 3197:

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

1157:, 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. 5967:

Yokobori, S.; Iseto, T; Asakawa, S; Sasaki, T; Shimizu, N; Yamagishi, A; Oshima, T; Hirose, E (May 2008). "Complete nucleotide sequences of mitochondrial genomes of two solitary entoprocts,

2122:. It is likely 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. By the 5308: 3465:. 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. 1389:

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

1276:. 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. 8017: 4253:

Lauer, Antje (2016). "Watersipora subtorquata and the Possible Role of Its Associated Microbes: An Attempt to Explain the Extraordinary Invasion Success of This Marine Bryozoan Species".

417:, 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 502:. 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 367:, live throughout the colony and are not fully independent. These individuals can have unique and diverse functions. All colonies have "autozooids", which are responsible for feeding, 1322:"Female" polymorphs are more common than "male" polymorphs, but specialized zooids that produce sperm are also known. These are called androzooids, and some are found in colonies of 2405:

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.

6400: 3911:

Fuchs, J.; Obst, M.; Sundberg, P (July 2009). "The first comprehensive molecular phylogeny of Bryozoa (Ectoprocta) based on combined analyses of nuclear and mitochondrial genes".

2887:

from the similarity between the coronate larvae and the Type 1 pericalymma larvae of some molluscs and sipunculans, where the prototroch zone is expanded to cover the hyposphere.

2592:

Molecular phylogeny analyses from 1995 onwards, using a variety of biochemical evidence and analytical techniques, placed the lophophorates as protostomes and closely related to

1422:, 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 name ( 6780: 1222:

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

360:", while the original Bryozoa were called "Ectoprocta". Disagreements about terminology persisted well into the 20th century, but "Bryozoa" is now the generally accepted term. 6541: 3135:, 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. 3345:. In temperate waters, the skeletons of dead colonies form a significant component of shell gravels, and live ones are abundant in these areas. The marine lace-like bryozoan 8133:

Gray, C.A.; McQuaid, CD; Davies-Coleman, MT (December 2005). "A symbiotic shell-encrusting bryozoan provides subtidal whelks with chemical defence against rock lobsters".

7553: 6640: 1673:, using this modern method of low vacuum scanning, with which it has been inaccurately synonymized with in the past. A new genus has also been recently discovered called 7939: 6223: 3047:. Instead, zooids absorb oxygen and eliminate carbon dioxide through diffusion. Bryozoa accomplish diffusion through the use of either a thin membrane (in the case of 3491: 3474: 1172:, 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 3208:

colony. The different forms of reproduction in cupuladriids are achieved through a variety of methods depending on the morphology and classification of the zooid.

1489:

are indecisive and the combatants soon turn to growing in uncontested areas. Bryozoans competing for territory do not use the sophisticated techniques employed by

2471:

When entoprocts were discovered in the 19th century, they and bryozoans (ectoprocts) were regarded as classes within the phylum Bryozoa, because both groups were

4883: 3193: 2917: 5525: 5093: 2439:

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

5267: 5226: 11185: 6142: 3543:

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

2521:

larva of some gymnolaete bryozoans produced suggestions that the bryozoans, most of which are colonial, evolved from a semi-colonial species of phoronid.

386:

exoskeletons and form single-layered sheets which encrust over surfaces, and some colonies can creep very slowly by using spiny defensive zooids as legs.

3199: 8178:"Bryoliths constructed by bryozoans in symbiotic associations with hermit crabs in a tropical heterozoan carbonate system, Golfe d'Arguin, Mauritania" 4209: 2580:. 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 6333:

Jablonski, D.; Lidgard, S.; Taylor, P.D. (1997). "Comparative Ecology of Bryozoan Radiations: Origin of novelties in cyclostomes and Cheilostomes".

10421: 3441:. Modifications appear in the shapes of some these hydroids, for example smaller tentacles or encrustation of the roots by bryozoans. The bryozoan 1405:, producing heavy lime skeletons. Many species form colonies which consist of sheets of autozooids. These sheets may form leaves, tufts or, in the 3643: oz) of bryostatin, As a result, synthetic equivalents have been developed that are simpler to produce and apparently at least as effective. 11052: 7030: 6565:"Exceptional soft-tissue preservation in boring ctenostome bryozoans and associated "fungal" borings from the Early Devonian of Podolia, Ukraine" 3221:

Most marine species live in tropical waters at depths less than 100 meters (330 ft; 55 fathoms). However, a few have been found in deep-sea

3112:. Stenolaemates produce specialized zooids to serve as brood chambers, and their eggs divide within this to produce up to 100 identical embryos. 3116:

unconfirmed records, like the solitary form Aethozoid where larvae has never been observed, but which is assumed to have planktotrophic larvae.

8667: 4316:

Zhang, Zhiliang; Zhang, Zhifei; Ma, J.; Taylor, P. D.; Strotz, L. C.; Jacquet, S. M.; Skovsted, C. B.; Chen, F.; Han, J.; Brock, G. A. (2021).

425:

that lie dormant until conditions are favorable, which enables a colony's lineage to survive even if severe conditions kill the mother colony.

11078: 8073: 2199:

are fairly equally divided by erect and encrusting forms, and more recent ones are predominantly encrusting. Fossils of the soft, freshwater

8814: 8321: 4079:"Aethozooides uraniae, a new deep-sea genus and species of solitary bryozoan from the Mediterranean Sea, with a revision of the Aethozoidae" 11180: 3151:, 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 1298:

that are divided along their length by diaphragms, making them look like rows of box-like chambers sandwiched between autozooidal tubes.

11175: 6596:

McKinney, F.K. (1994). "One hundred million years of competitive interactions between bryozoan clades: asymmetrical but not escalating".

3123:

of bryozoan eggs is biradial, in other words the early stages are bilaterally symmetrical. It is unknown how the coelom forms, since the

3108:

In ctenostomes the mother provides a brood chamber for the fertilized eggs, and her polypide disintegrates, providing nourishment to the

2552:

If the grouping of bryozoans with phoronids and brachiopods into Lophophorata is correct, the next issue is whether the Lophophorata are

2343:

phylum, in other words includes all the descendants of a common ancestor that is itself a bryozoan. The analysis also concluded that the

6382: 1267:

of others. In some species the snapping zooids are mounted on a peduncle (stalk), their bird-like appearance responsible for the term –

8111: 7898: 5820: 8805: 8275: 7291: 3179:. Statoblasts form on the funiculus connected to the parent's gut, which nourishes them. As they grow, statoblasts develop protective 6802: 6762: 5463:

Eckman, J.E.; Okamura, B (December 1998). "A Model of Particle Capture by Bryozoans in Turbulent Flow: Significance of Colony Form".

1857: 1637:. This method permits data to be obtained that would be difficult to recognize with an optical microscope. The valid type species of 498:

alga. Early fossils are mainly of erect forms, but encrusting forms gradually became dominant. It is uncertain whether the phylum is

8679: 8407: 8240:"Molecular data implicate bryozoans as hosts for PKX (Phylum Myxozoa) and identify a clade of bryozoan parasites within the Myxozoa" 7766: 6523: 3056:

to circulate freely; in others, the funiculi (internal "little ropes") of adjacent zooids connect via small pores in the body wall.

11026: 5290: 5149: 1206:

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

7723: 2164:

appeared around the same time, which suggests that some change made the environment more favorable for this lifestyle. Fossils of

11065: 10127: 7535: 7486: 7169:"The placental analogue and the pattern of sexual reproduction in the cheilostome bryozoan Bicellariella ciliata (Gymnolaemata)" 6929:(Bryozoa: Cheilostomatida) determined with next-generation sequencing: The first representative of the suborder Malacostegina". 6704:

Pohowsky, R.A. (1978). "The boring ctenostomate bryozoa: taxonomy and paleobiology based on cavities in calcareous substrata".

6624: 5548: 8546:

Hall, S.R.; Taylor, P.D.; Davis, SA; Mann, S (2002). "Electron diffraction studies of the calcareous skeletons of bryozoans".

5185: 3426:

also preys on a common freshwater gymnolaemate, but with less devastating effect. Indigenous snails do not feed on bryozoans.

1335:

exchange sperm between colonies when two mobile colonies or bryozoan-encrusted hermit crabs happen to encounter one another.

8618: 8029: 7933: 7921: 7844: 7547: 7504: 7301: 6774: 6634: 6535: 6394: 6317: 6203: 6155: 6081: 5765: 5740: 5519: 5447: 5415: 5302: 5261: 5220: 5087: 4990: 4965: 4877: 4848: 4817: 4726: 4690: 4647: 4593: 4555: 4517: 4270: 3652: 3244:

lives on the leaves of mangrove trees and is called "amphibious" because it can survive regular exposure to air at low tide.

11070: 7797: 6219: 4294: 8213: 7577: 1153:

The most common type of zooid is the feeding autozooid, in which the polypide bears a "crown" of hollow tentacles called a

8377: 5856:

Hausdorf, B.; Helmkampf, M.; Meyer, A.; Witek, A.; Herlyn, H.; Bruchhaus, I.; Hankeln, T.; Struck, T.H.; Lieb, B. (2007).

6679: 3657: 7362:

Strathmann, R.R. (March 2006). "Versatile ciliary behaviour in capture of particles by the bryozoan cyphonautes larva".

6925:

Shen, X.; Tian, M.; Meng, X.; Liu, H.; Cheng, H.; Zhu, C.; Zhao, F. (September 2012). "Complete mitochondrial genome of

5403: 2210:) and consist entirely of their durable statoblasts. There are no known fossils of freshwater members of other classes. 1534: 8841: 5043: 8346:

Wood, T.S.; Marsh, Terrence G (February 1999). "Biofouling of wastewater treatment plants by the freshwater bryozoan,

4865: 3807: 2961:(planktonic animals) by using their tentacles as cages. In addition the tentacles, whose surface area is increased by 409:

into the water, while others capture sperm via their tentacles to fertilize their ova internally. In some species the

8529: 8404:"Safety, Efficacy, Pharmacokinetics, and Pharmacodynamics Study of bryostatin 1 in Patients With Alzheimer's Disease" 5507: 5075: 3969: 1145: 474:

and fishermen. Chemicals extracted from a marine bryozoan species have been investigated for treatment of cancer and

7575:

Kuklinski, P.; Bader, Beate (2007). "Comparison of bryozoan assemblages from two contrasting Arctic shelf regions".

11137: 11005: 8878: 6882:

Nielsen, C.; Worsaae, K. (September 2010). "Structure and occurrence of cyphonautes larvae (Bryozoa, Ectoprocta)".

1510: 11083: 7826: 6835:"Phylogenomic analyses of lophophorates (brachiopods, phoronids and bryozoans) confirm the Lophotrochozoa concept" 4257:. Advances in Environmental Microbiology. Vol. 2. Cham: Springer International Publishing. pp. 239–268. 11018: 8873: 6969:

Callaghan, T.P.; R., Karlson (June 2002). "Summer dormancy as a refuge from mortality in the freshwater bryozoan

5249: 5208: 3756:(Cryptostomata) from the lower Tremadocian (Lower Ordovician) of Liujiachang, south-western Hubei, central China" 494:, could be an example of an early bryozoan, but later research suggested that this taxon may instead represent a 510:

analyses disagree over bryozoans' relationships with entoprocts, about whether bryozoans should be grouped with

8431:"Bryostatin Effects on Cognitive Function and PKCɛ in Alzheimer's Disease Phase IIA and Expanded Access Trials" 6499: 6475: 8299:"Professional photosensitive eczema of fishermen by contact with bryozoans: disabling occupational dermatosis" 4629: 3728: 3531:, whose colonies feed and grow exceptionally fast in a wide range of current speeds, was first noticed in the 8957: 8947: 8930: 3626: 3019:

out through the body wall and lophophore. More complex waste products are not excreted but accumulate in the

2989:

contractions move the food through the stomach for digestion. The final section of the stomach is lined with

2257:

Molecular phylogeny, which attempts to work out the evolutionary family tree of organisms by comparing their

214: 6106:

Taylor, Paul D (October 2008). "Taxonomy of the bryozoan genera Oncousoecia, Microeciella and Eurystrotos".

6260:

Massard, J.A.; Geimer, Gaby (2008). "Global diversity of bryozoans (Bryozoa or Ectoprocta) in freshwater".

4030:"Key novelties in the evolution of the aquatic colonial phylum Bryozoa: evidence from soft body morphology" 3570: 3516: 3437:

have symbiotic relationships with bryozoans, some of which are beneficial to the hydroids while others are

2584:

phyla as it is built anew from epidermis and mesoderm after metamorphosis has destroyed the larval coelom.

2397:

In the opinion of Ruth Dewel, Judith Winston, and Frank McKinney, "Our standard interpretation of bryozoan

2386:

Bryozoans' relationships with other phyla are uncertain and controversial. Traditional phylogeny, based on

1009: 4228: 10974: 10961: 10120: 3247:

There are a variety of "free-living" bryozoans that live un-attached to a substrate. A few forms such as

1497:, possibly because the shortness of bryozoan lifespans makes heavy investment in turf wars unprofitable. 4167: 470:

so much that it has affected local fish and invertebrate populations. Bryozoans have spread diseases to

4133:

Zoological researches and illustrations; or Natural history of nondescript or imperfectly known animals

1649:

by establishing a type species that corresponds to the general usage of the genus. Fellow Oncousoeciid

7259:"Morphology and life cycle of an epiphytic pherusellid ctenostome bryozoan from the Mediterranean Sea" 11170: 11142: 11093: 10264: 8628: 7118:"Three in one: Evolution of viviparity, coenocytic placenta and polyembryony in cyclostome bryozoans" 17: 10979: 3383:

eat masses of zooids by gouging pieces of colonies, breaking their mineralized "houses", while most

1210:, which is located on the invert, outside and usually below the lophophore. A network of strands of 164: 8058: 7116:

Nekliudova, U. A.; Schwaha, T. F.; Kotenko, O. N.; Gruber, D.; Cyran, N.; Ostrovsky, A. N. (2021).

5858:"Spiralian Phylogenomics Supports the Resurrection of Bryozoa Comprising Ectoprocta and Entoprocta" 4189: 4127: 3618: 3527: 3347: 3330: 2301:

study that focused on phylactolaemates concluded that these are more closely related to the phylum

462: 8298: 4149:"Pars zoologica. Animalia evertebrata exclusis insectis [Invertebrata other than Insecta]" 10349: 8962: 7219: 3257: 2517:

in a super-phylum called "Tentaculata". In the 1970s comparisons between phoronid larvae and the

466:, a fast-growing invasive bryozoan off the northeast and northwest coasts of the US, has reduced 7836: 7830: 4754: 4752: 4750: 4748: 4746: 4744: 4742: 4740: 4738: 382:

of bryozoan, possibly because its members have the widest range of specialist zooids. They have

11119: 10901: 10113: 8834: 4153:

Symbolae physicae, seu lcones et descriptiones corporum naturalium novorum aut minus cognitorum

3630: 2486:

From 1869 onwards increasing awareness of differences, including the position of the entoproct

475: 8723:"The earliest bryozoan parasite: Middle Ordovician (Darriwilian) of Osmussaar Island, Estonia" 7496: 7490: 4077:

Schwaha, Thomas; Bernhard, Joan M.; Edgcomb, Virginia P.; Todaro, M. Antonio (1 August 2019).

1730:, which is still used as a convenient way to label groups of organisms, living members of the 1657:, as previously suggested, but shows enough similarities to be considered a junior synonym of 972:. The latter is always absent in freshwater species. The body wall consists of the epidermis, 11114: 10577: 10227: 8925: 6669:"The higher phylogeny of phylactolaemate bryozoans inferred from 18S ribosomal DNA sequences" 6309: 6301: 5816: 5797: 5616:"Secular changes in colony-forms and bryozoan carbonate sediments through geological history" 4735: 4639: 4633: 2502: 2102: 149: 11129: 11106: 8429:

Nelson, T. J.; Sun, M. K.; Lim, C.; Sen, A.; Khan, T.; Chirila, F. V.; Alkon, D. L. (2017).

8099: 7868: 3402:. Freshwater bryozoans are attacked by many predators, including snails, insects, and fish. 1589:), in which the anus lies within a "crown" of tentacles, the name "Bryozoa" was promoted to 421:

that destroys and rebuilds almost all the internal tissues. Freshwater species also produce

363:

Colonies take a variety of forms, including fans, bushes and sheets. Single animals, called

10948: 10242: 8734: 8694: 8239: 8189: 8142: 7974: 7880: 7699: 7586: 7446: 7334: 7231: 7082: 6982: 6487: 6344: 6031: 6018:

Reynolds, K.T. (2000). "Taxonomically Important Features on the Surface of Floatoblasts in

5984: 5131: 4329: 4090: 3920: 3870: 3819: 3767: 3704: 3414: 2398: 1470: 503: 11057: 8710: 8403: 7758: 6806: 4840: 4834: 4809: 4801: 4718: 4710: 4682: 4674: 4585: 4577: 4547: 4539: 4509: 4501: 3068:

Encrusting cyclostome bryozoans (B), the one on the right showing swollen gonozooids; T =

2890:

A study of the mitochondrial DNA sequence suggests that the Bryozoa may be related to the

2408: 1677:

in the family Stomachetosellidae, along with 10 relatively new species of bryozoa such as

892:

Some sessile annelids build mineralized tubes; most molluscs have shells, but most modern

8: 7317:

Schwaha, Thomas; Zeppilli, Daniela; González-Casarrubios, Alberto; Cepeda, Diego (2024).

5116: 3468: 3452: 3443: 3286: 3237: 3016: 2799: 2472: 2463:, in which the tissue between the gut and the body wall splits, forming paired cavities. 2328: 2298: 953: 841: 507: 10105: 8738: 8698: 8193: 8146: 7978: 7884: 7869:"Specificity of cues inducing defensive spines in the bryozoan Membranipora membranacea" 7703: 7612:

Brusca, R; Brusca, G. "Invertebrates (2nd Edition)". Sunderland, MA: Sinauer Associates.

7590: 7450: 7338: 7235: 7086: 6986: 6828: 6826: 6824: 6491: 6348: 6195: 6187: 6035: 5988: 5135: 4333: 4094: 3924: 3874: 3823: 3771: 3708: 3553:. These changes reduced the area of habitat available for local fish and invertebrates. 3269:

Other free-living bryozoans are moved freely by waves, currents, or other phenomena. An

11165: 10572: 10254: 9390: 8942: 8920: 8908: 8883: 8457: 8430: 8267: 8205: 8158: 7998: 7715: 7687: 7464: 7415: 7195: 7168: 7144: 7117: 7098: 7006: 6907: 6859: 6834: 6609: 6360: 6279: 6123: 6068: 5949: 5914: 5713: 5675: 5637: 5576: 5488: 4363: 4350: 4317: 4054: 4029: 3886: 3843: 3785: 3720: 3410: 2269:

phyla. However, the shortage of genetic data about "minor phyla" such as bryozoans and

457: 238: 159: 31: 8706: 8559: 8363: 6528:

Bryozoan studies 2001: proceedings of the Twelfth International Bryozoology Conference

4148: 3861:

Ernst, A. (2007). "A cystoporate bryozoan species from the Zechstein (Late Permian)".

2281:

The traditional view is that the Bryozoa are a monophyletic group, in which the class

1913:

Pulling inwards of a flexible section of body wall, or making an internal sac expand.

11101: 10939: 10364: 9812: 9732: 9082: 8952: 8915: 8903: 8893: 8827: 8809: 8799: 8722: 8659: 8605: 8563: 8525: 8498: 8462: 8313: 8259: 8025: 7990: 7929: 7840: 7543: 7500: 7459: 7434: 7375: 7297: 7200: 7149: 7055: 6998: 6946: 6899: 6864: 6821: 6770: 6630: 6531: 6390: 6313: 6199: 6151: 6077: 6047: 6000: 5879: 5761: 5736: 5641: 5584: 5568: 5515: 5480: 5443: 5411: 5298: 5257: 5216: 5083: 5039: 4986: 4961: 4873: 4844: 4813: 4782: 4722: 4686: 4643: 4589: 4551: 4513: 4367: 4355: 4266: 4131: 4059: 3994: 3975: 3965: 3956: 3936: 3890: 3847: 3835: 3789: 3724: 3430: 3230: 3120: 2491: 2375:

Cheilostomata, but the 2009 analysis considered it more likely that neither of these

2372: 2115: 1727: 1419: 1390: 977: 969: 869: 589: 383: 376: 8209: 8162: 8002: 7719: 7468: 7010: 6911: 6676:

Bryozoan Studies 2004: Proceedings of the 13th International Bryozoology Association

6283: 6127: 5851: 5849: 5847: 5845: 5843: 5841: 5592: 10640: 10519: 10411: 10354: 10294: 10289: 10217: 10202: 8867: 8742: 8702: 8651: 8643: 8595: 8555: 8490: 8452: 8442: 8359: 8271: 8251: 8197: 8150: 7982: 7888: 7707: 7594: 7492:

Planet Earth: Cosmology, Geology, & the Evolution of Life & the Environment

7454: 7419: 7407: 7371: 7342: 7270: 7239: 7190: 7180: 7139: 7129: 7102: 7090: 7045: 6990: 6938: 6891: 6854: 6846: 6736: 6605: 6576: 6495: 6352: 6271: 6115: 6039: 5992: 5941: 5906: 5869: 5812: 5705: 5667: 5627: 5560: 5492: 5472: 5177: 5139: 5031: 5005: 4772: 4345: 4337: 4258: 4106: 4098: 4049: 4041: 4010: 3928: 3878: 3827: 3775: 3712: 3662: 3589: 3540: 3482: 3156: 3009: 2966: 2440: 2127: 491: 280: 9086: 4128:"Memoir V: On Polyzoa, a new animal discovered as an inhabitant of some Zoophites" 3523:

fish, and has severely reduced wild fish populations in Europe and North America.

2459:, in which pouches that form on the wall of the gut become separate cavities, nor 10813: 10718: 10680: 10635: 10567: 10324: 10284: 10269: 10162: 10093: 9999: 9850: 9753: 9610: 9143: 8789: 8746: 8233: 8231: 7789: 7482: 6425:

McKinney; Frank K; Jeremy. "Bryozoan Evolution". Boston: Unwin & Hyman, 1989.

6150:. Department of the Environment and Heritage, Australian Government. p. 34. 5897:

Cuffey, R. J. (1969). "Bryozoa versus Ectoprocta – The Necessity for Precision".

5838: 4761:"The Phylogenetic Position of Entoprocta, Ectoprocta, Phoronida, and Brachiopoda" 4290: 3964:(revised & expanded, 2nd ed.). Tucson, AZ: University of Arizona Press. 3609: 3495:, which is generally growing attached to gravel and hard-substrate, has formed a 2751: 2633: 2376: 2360: 2344: 2294: 2282: 2200: 2138: 1762: 1745: 1740: 1594: 1200: 1165: 1161: 1095: 389:

Each zooid consists of a "cystid", which provides the body wall and produces the

379: 314: 10953: 8794: 8177: 5996: 4262: 3932: 3143:

for a considerable time before settling. These larvae have triangular shells of

2821: 2168:, an order of gymnolaemates with mineralized skeletons, first appear in the Mid 1364: 352:). The latter having a statocyst-like organ with a supposed excretory function. 11013: 11000: 10966: 10873: 10818: 10808: 10650: 10464: 10416: 10329: 10304: 10299: 10279: 10274: 10177: 10167: 9758: 9736: 9554: 9466: 9444: 9407: 8898: 8381: 7598: 7433:

O'Dea, Aaron; Jackson, Jeremy B. C.; Taylor, Paul D.; Rodríguez, Felix (2008).

7347: 7318: 7275: 7258: 7167:

Moosbrugger, M.; Schwaha, T.; Walzl, M. G.; Obst, M.; Ostrovsky, A. N. (2012).

7134: 6668: 4341: 4102: 4014: 3831: 3622: 3548: 3461: 3291: 3222: 2938: 2934: 2615: 2601: 2368: 2290: 2146: 2067: 2032: 1777: 1772: 1609:

The common name "moss animals" is the literal meaning of "Bryozoa", from Greek

1410: 1268: 961: 644: 551: 322: 306: 302: 273: 122: 8255: 8228: 8201: 8154: 7411: 7243: 6994: 6942: 6725:"The Essential Role of "Minor" Phyla in Molecular Studies of Animal Evolution" 6275: 6119: 6043: 5215:. Synopses of the British fauna. Linnean Society of London. pp. 106–107. 3979: 3592:", caused by contact with bryozoans that have stuck to nets and lobster pots. 3167:

clones of themselves. In phylactolaemates, zooids die after producing several

3053: 11159: 10924: 10597: 10582: 10544: 10469: 10405: 10391: 10339: 10334: 10248: 10237: 10232: 10212: 10207: 10182: 9903: 9807: 9306: 7822: 7398: 6262: 5572: 5117:"Downstream collecting in ciliary suspension feeders: the catch-up principle" 4078: 3906: 3904: 3902: 3900: 3532: 3472:

Mauritanian bryolith formed by circumrotatory growth of the bryozoan species

3391: 3180: 3152: 3124: 3089: 3027:, or in the stomach of the new polypide and is expelled next time the animal 2930: 2912: 2908: 2730: 2476: 2227: 2165: 2161: 1782: 1423: 719: 418: 398: 372: 298: 143: 77: 5874: 5857: 5438:

Little, W.; Fowler, H.W.; Coulson, J. & Onions, C.T. (1959). "Bryozoa".

5035: 3629:

to measure the safety and effectiveness of Bryostatin 1 in the treatment of

2993:(minute hairs) that compress undigested solids, which then pass through the 1486: 10793: 10788: 10763: 10738: 10728: 10655: 10524: 10398: 10359: 10344: 10319: 10314: 10259: 10192: 10009: 10004: 9892: 9748: 9634: 9520: 9423: 9402: 9272: 9223: 9179: 9160: 8663: 8609: 8567: 8502: 8466: 8317: 8263: 7994: 7817: 7815: 7204: 7185: 7153: 7059: 7002: 6950: 6903: 6868: 6850: 6741: 6724: 6255: 6253: 6251: 6249: 6247: 6245: 6243: 6241: 6051: 6004: 5883: 5588: 5484: 4786: 4777: 4760: 4359: 4063: 3940: 3839: 3750:

Ma, Junye; Taylor, Paul D.; Xia, Fengsheng; Zhan, Renbin (September 2015).

3500: 3457: 3324: 3131:

to adult destroys all of the larva's internal tissues. In many animals the

3044: 2891: 2703: 2565: 2557: 2542: 2526: 2364: 2356: 2352: 2348: 2340: 2336: 2332: 2316: 2308: 2286: 2266: 2258: 2251: 2242: 2150: 2142: 1885: 1767: 1755: 1750: 1308: 1235: 1137: 989: 973: 919: 815: 638: 527: 519: 499: 495: 348: 318: 310: 276: 199: 8784: 7926:

Cyclostome bryozoans: keys and notes for the identification of the species

6581: 6564: 5756:

Little, W.; Fowler, H.W.; Coulson, J. & Onions, C.T. (1964). "Ento-".

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Schwaha, Thomas F.; Ostrovsky, Andrew N.; Wanninger, Andreas (June 2020).

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lies inside rather than outside the base of the "crown" and they have no

610:, another phylum of filter-feeders, look rather like bryozoans but their 592: 585: 548: 467: 390: 52: 8494: 7812: 6524:"Deconstructing byozoans: origin and consequences of a unique body plan" 6238: 5730: 4980: 3806:

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3298:, and this species is particularly adept at recolonizing scoured areas. 2915:(microscopic floating plants), out of the water. The freshwater species 2254:, the two major groups that account for all moderately complex animals. 2059: 2024: 1998: 1972: 1598:

different positions of the anus, ectoprocts have hollow tentacles and a

1372:

Although zooids are microscopic, colonies range in size from 1 cm (

481:

Mineralized skeletons of bryozoans first appear in rocks from the Early

10881: 10843: 10773: 10753: 10748: 10733: 10703: 10675: 10630: 10620: 10592: 10587: 10529: 10504: 10197: 10172: 10067: 10052: 10047: 10028: 10023: 9951: 9898: 9868: 9860: 9802: 9676: 9668: 9651: 9431: 9343: 9296: 9236: 9198: 9172: 9018: 9003: 8447: 7986: 7893: 7711: 7031:"Living where the flow is right: How flow affects feeding in bryozoans" 6931:

Comparative Biochemistry and Physiology Part D: Genomics and Proteomics

6895: 6756: 6754: 6752: 6364: 5953: 5918: 5717: 5679: 5580: 5144: 3882: 3780: 3751: 3604: 3596: 3566: 3486: 3434: 3364: 3360: 3356: 3352: 3338: 3176: 3132: 3069: 2962: 2954: 2816: 2794: 2778: 2713: 2569: 2553: 2530: 2522: 2402: 2320: 2270: 2247: 2223: 2203:

are very rare, appear in and after the Late Permian (which began about

2180: 2179:, and these have been the most abundant and diverse bryozoans from the 2119: 1949: 1904: 1792: 1570: 1523: 1478: 1181: 1154: 893: 821: 806: 780: 740: 673: 611: 607: 601: 558:, a "crown" of hollow tentacles. Bryozoans form colonies consisting of 555: 523: 511: 482: 456:, many populations of one freshwater species have been wiped out by an 441: 437: 433: 429: 422: 357: 294: 97: 62: 8819: 8176:

Klicpera, André; Taylor, Paul D.; Westphal, Hildegard (30 July 2013).

7094: 6662: 6660: 6658: 6506: 5632: 5615: 4045: 3808:"Protomelission is an early dasyclad alga and not a Cambrian bryozoan" 3422:, has wiped out phylactolaemate populations wherever it has appeared. 2265:, has done much to clarify the relationships between the better-known 452:. Freshwater bryozoans are preyed on by snails, insects, and fish. In 11044: 10987: 10798: 10778: 10670: 10549: 10484: 10474: 10459: 10454: 10435: 10187: 10033: 9980: 9975: 9946: 9935: 9884: 9879: 9831: 9743: 9530: 9525: 9487: 9336: 9328: 9257: 9209: 9130: 9117: 9098: 9093: 9052: 8655: 8647: 8584:"Life in the Colonies: Learning the Alien Ways of Colonial Organisms" 6964: 6962: 6960: 6526:. In Wyse Jacksdon, P.E.; Buttler, C.E.; Spencer Jones, M.E. (eds.). 3581: 3496: 3419: 3384: 3270: 3226: 3073: 3005: 2994: 2746: 2682: 2581: 2448: 2432: 2205: 2190: 2185: 2174: 2155: 2132: 2123: 1953: 1443: 1432: 1383: 1327: 1312: 1289: 1264: 949: 654: 544: 471: 368: 176: 102: 10895: 8778: 6749: 6356: 5945: 5910: 5709: 5671: 5564: 4956:

Doherty, P.J. (2001). "The Lophophorates". In Anderson, D.T. (ed.).

4210:"Heavy Predation on Freshwater Bryozoans by the Golden Apple Snail, 3171:, so that living zooids are found only round the edges of a colony. 2654: 2241:

Scientists are divided about whether the Bryozoa (Ectoprocta) are a

1448:

meaning 'animal'). Large colonies of encrusting species often have "

10992: 10918: 10853: 10833: 10828: 10803: 10783: 10758: 10723: 10713: 10708: 10539: 10514: 10494: 10154: 10057: 10041: 9964: 9956: 9911: 9874: 9855: 9844: 9826: 9820: 9681: 9622: 9598: 9495: 9376: 9265: 9149: 9040: 9031: 8937: 8721:

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Ernst, Andrej (2020). "2- Fossil record and evolution of Bryozoa".

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Some phylactolaemate species are intermediate hosts for a group of

3438: 3406: 3380: 3372: 3140: 3077: 3028: 3020: 2950: 2946: 2677: 2650: 2561: 2514: 2452: 2444: 2380: 2302: 2196: 2169: 1542: 1515: 1352: 1219: 1177: 1051: 682: 668: 664: 515: 453: 449: 394: 116: 92: 87: 72: 67: 57: 11031: 8680:"Palaeoecology and evolution of marine hard substrate communities" 7686:

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6957: 6185: 5691: 5689: 5401: 4919: 4917: 4915: 4913: 4911: 4909: 4907: 4905: 4903: 4901: 4832: 4799: 4708: 4704: 4702: 4672: 4575: 4537: 4499: 2367:. The Gymnolaemata are traditionally divided into the soft-bodied 2218: 1621:('animals'), based on the mossy appearance of encrusting species. 1569:

lies outside the "crown" of tentacles. After the discovery of the

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4293:. Berkeley, CA: University of California Museum of Paleontology. 3512: 3355:(nudibranchs). Other predators on marine bryozoans include fish, 3342: 3306: 3295: 3168: 3164: 3064: 3013: 2974: 2942: 2597: 2593: 2387: 1957: 1527: 1473: 1449: 1017: 965: 923:. Individual members of a bryozoan colony are about 0.5 mm ( 886: 573: 559: 107: 82: 8773: 8018:"Predation on Bryozoans and its Reflection in the Fossil Record" 6521: 3535:

in 1987 and quickly became the most abundant organism living on

10698: 10690: 10559: 10062: 9796: 8996: 8983: 8854: 8851: 8015: 7432: 7319:"The first deep-sea ctenostome bryozoan from the Indian Ocean: 7257:

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produces spines in response to predation by several species of

3184: 3160: 3144: 3109: 3097: 3081: 3048: 3024: 2982: 2978: 2973:

In some species the first part of the stomach forms a muscular

2922: 2495: 2423: 2391: 2383:

probably evolved more than once within the early Gymnolaemata.

1927: 1868: 1839: 1731: 1599: 1590: 1490: 1398: 1231: 1078: 1065: 985: 957: 713: 690: 619: 597: 581: 577: 293: in) long, they have a special feeding structure called a 269: 186: 8059:"Freshwater Bryozoans of Thailand (Ectoprocta and Entoprocta)" 7115: 6674:. In Moyano, H. I.; Cancino, J.M.; Wyse-Jackson, P.N. (eds.). 6376: 6374: 6308:(2 ed.). Cambridge: Cambridge University Press. pp. 5791: 5789: 5787: 5785: 5783: 5781: 5779: 5777: 4571: 4569: 4567: 4318:"Fossil evidence unveils an early Cambrian origin for Bryozoa" 2297:

that appear earliest in the fossil record. However, in 2005 a

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inside the feeding structure and the difference in the early

2480: 1881: 1625:

taxonomy of three genera belonging to this family, including

1616: 1610: 1522:, a trepostome bryozoan from the Whitewater Formation (Upper 1494: 1438: 1427: 1406: 1402: 1273: 1227: 1223: 1173: 1169: 1108: 1034: 914: 878: 836: 776: 702: 678: 410: 402: 364: 331: 7387: 7385: 7073:

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were also assigned to the "Tentaculata", which were renamed

27:

Phylum of colonial aquatic invertebrates called moss animals

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6371: 5855: 5774: 4960:(2nd ed.). Oxford University Press. pp. 363–373. 4564: 4495: 4493: 4491: 4489: 4487: 4485: 4483: 4481: 4479: 4477: 4475: 4473: 4471: 4469: 4467: 4465: 4463: 4461: 4459: 4457: 4455: 4453: 4451: 4449: 4447: 4445: 4443: 4441: 4439: 4437: 4435: 4433: 4431: 4429: 4427: 4425: 4423: 4421: 4419: 4417: 4415: 4413: 4411: 4409: 4407: 4405: 4403: 4401: 4399: 4397: 4155:(in Latin). Berolini Ex officina Academica. pp. 1–126. 3544: 3536: 3368: 3148: 2998: 2487: 2416: 2323:

valve; Cincinnatian of southeastern Indiana, United States.

2319:

of a Late Ordovician ctenostome bryozoan on a strophomenid

2262: 1566: 1207: 1203:, a hollow lobe called the "epistome" overhangs the mouth. 1160:

The basic shape of the "crown" is a full circle. Among the

1058: 800: 751:

Similar-looking feeding structure, but with solid tentacles

615: 445: 414: 406: 246:

Ectoprocta (Nitsche, 1869) (formerly subphylum of Bryozoa)

7790:"Bryozoans: The Fascinating Colonies Of Phylum Ectoprocta" 7652: 7640: 7024: 7022: 7020: 6832: 6500:

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development of the embryo; and formation of the coelom by

844:

species often form clumps, but with no active co-operation

305:

bryozoans live in tropical waters, but a few are found in

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Colonies of clones in some species; some solitary species

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6473: 5323: 5073: 4602: 4374: 4194:. London: British Museum (Natural History). p. 13. 3418:(golden apple snail), which is generally a destructive 3088:

Zooids of all phylactolaemate species are simultaneous

2587: 2355:

are also monophyletic, but could not determine whether

1903:

Compressing the membranous sac(separate inner layer of

1218:

There is a nerve ring round the pharynx (throat) and a

8767: 8237: 8175: 7738: 7628: 7616: 7256: 6332: 5172: 5170: 3390:

In freshwater, bryozoans are among the most important

3290:

had become a dominant species in parts of Antarctica.

2501:

However, two well-known zoologists, Claus Nielsen and

2273:

has left their relationships to other groups unclear.

309:

and polar waters. The bryozoans are classified as the

30:"Polyzoa" redirects here. For the tunicate genus, see 8479: 7961:

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Three-part, if the cavity of the epistome is included

6530:. M.E. Lisse: Swets and Zeitlinger. pp. 93–96. 6186: