(invertebrate zoology) A group of bryozoans, sometimes considered to be a subphylum, having a pseudocoelomate visceral cavity and the anus opening inside the circlet of tentacles.
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(invertebrate zoology) A group of bryozoans, sometimes considered to be a subphylum, having a pseudocoelomate visceral cavity and the anus opening inside the circlet of tentacles.
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| Animal Classification: Entoprocta |
(Entoprocts)
Phylum: Entoprocta (Kamptozoa)
Number of families: 4
Thumbnail description
Colonial or solitary tiny benthic animals with a tentacular crown on top, and a slender stalk that attaches basally to the substratum
Evolution and systematics
There has been only one fossil record of Entoprocta, belonging to the extant genus Barentsia, which was collected from the Upper Jurassic of England. The phylogenetic relationships of Entoprocta to other invertebrate phyla are still obscure, but Entoprocta may have affinities to spiralians (animal groups that show spiral cleavage patterns). The phylum encompasses two orders, four families, sixteen genera, and approximately 170 species. The four families are: Barentsiidae, colonial species with a muscular swelling at the base of the stalk; Pedicellinidae, colonial species without basal muscular swelling, each zooid of a colony interrupted by stolon; Loxokalypodidae, colonial species without basal muscular swelling, component zooids of a colony erect from a common basal plate, not interrupted by stolons; and Loxosomatidae, which encompasses all solitary species.
The phylum name Entoprocta means "inside anus;" the phylum has this name because of its unique plan. The animal's anus opens inside its tentacular crown. Kamptozoa is another scientific name for this phylum; the name means "bending animal," and comes from the very active movement of these animals.
Physical characteristics
The calyx, or main body, of an entoproct contains a U-shaped gut, a ganglion, a pair of gonads, a pair of protonephridia, and has a tentacular crown on top. Both the mouth and anus open inside the tentacular crown. The calyx is supported by a slender stalk that attaches basally to the substratum. In colonial species, zooids of a colony are generally connected by a highly branched stolon that creeps over the substratum. The solitary species have an attaching organ at the base of the stalk. In some species, however, the attaching organ degenerates in the adult stage and the adult animals are cemented onto the substratum.
Distribution
Entoprocts have been reported from tropical, temperate, and polar marine waters, and from shallow seashore to deep seas of more than 1,640 ft (500 m). One colonial species, Urnatella gracilis, occurs worldwide in inland waters.
Habitat
Colonial species live on a wide variety of substrata, including rocks, stones, shell remains, human-made objects, and occasionally on other animals. Most solitary species have been known to live on the bodies of specific host animals, such as polychaetes, bryozoans, sponges, and sipunculans, and on the inner side of the tube of polychaetes.
Behavior
In response to irritation, entoprocts contract their tentacles and bend at the stalk. Some solitary species can glide over the substratum as slugs do. One solitary species (Loxosoma agile) somersaults across the substratum, and another species (Loxosomella bifida) can walk on the substratum similar to the way humans do, using a unique foot with two elongated, leglike extensions. Newly liberated buds of solitary species often swim using ciliary tentacles, contributing to the dispersal of the species.
Feeding ecology and diet
All entoprocts are suspension feeders, feeding on phytoplankton or other organic particles in a water current they create using the cilia along their tentacles.
Reproductive biology
Each zooid of a colonial species is generally dioecious, male or female, but both sexes occur in a single colony. Solitary species are generally protandrous hermaphrodites, namely, animals are males in the early stage but later convert into females. Eggs are fertilized in the ovary and transferred to a brood pouch, a deep depression between the mouth and the anus, where embryos develop to trochophorelike larvae. Asexual reproduction (budding) is vigorous in all entoprocts. Buds occur from the tips of developing stolon, or from a basal disc and stalks in colonial species. In solitary species budding usually occurs at two latero-frontal areas of the calyx.
Conservation status
Entoprocts may be common in worldwide seas. However, their distribution and abundance are still poorly documented, and their responses to human activities have not been monitored. No species is listed by the IUCN.
Significance to humans
Entoprocts have no significance to humans.
Species accounts
Marine colonial entoproctResources
Books:Hyman, L. H. The Invertebrates: Acanthocephala, Aschelminthes, and Entoprocta. Vol. 3, The Pseudocoelomate Bilateria. New York: McGraw-Hill, 1951.
Nielsen, C. "Entoprocts." In Synopses of the British Fauna (New Series) 41, edited by Doris M. Kermack and R. S. K. Barnes. Leiden, The Netherlands: E. J. Brill., 1989.
Periodicals:Wasson, K. "A Review of the Invertebrate Phylum Kamptozoa (Entoprocta) and Synopses of Kamptozoan Diversity in Australia and New Zealand." Transactions of the Royal Society of South Australia 126 (2002): 1–20.
[Article by: Tohru Iseto, PhD]
| Sci-Tech Encyclopedia: Entoprocta |
A phylum of sessile, aquatic, often colonial invertebrates having a looped gut with both mouth and anus situated inside a circlet of tentacles, a pseudocoelomate body cavity, and no hardened skeleton. The phylum is divided into two orders, Loxosomatida (or Solitaria), with about 100 species, and Pedicellinida (or Coloniales), with about 30. No fossils are known.
Entoprocts are mainly marine, and may be encountered as small colonies on seaweeds and bryozoans near low tidemark. The colonies consist of threadlike, creeping stolons and erect zooids arising at intervals from them (see illustration). Noncolonial forms have similar zooids but lack the stolons. Each zooid is goblet-shaped, consisting of a basal stalk supporting a rounded or pyriform viscera-containing dilatation, the calyx.

Morphological features of an entoproct zooid, in vertical cross section.
The entoprocts feed on particles filtered from the surrounding water. Long lateral cilia drive water inward between the tentacles, while shorter cilia on the inner face convey particles trapped in mucus down the tentacles. These frontal tracts join an annular ciliated groove, which dips into the mouth.
Entoprocts reproduce asexually and sexually. Dispersal is by means of sexually produced larvae. Zooids may be unisexual or hermaphroditic. Gonads are paired, with the ducts uniting before opening into the coelom. Fertilization is internal.
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Barentsia discreta
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Barentsiidae (Urnatellidae) |
Entoprocta, whose name means "anus inside", is a phylum of mostly sessile aquatic animals, ranging from 0.1 to 7 millimetres (0.0039 to 0.28 in) 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 both the mouth and anus lie inside the "crown". The superficially similar Bryozoa (Ectoprocta) have the anus outside a "crown" of hollow tentacles. Most families of entoprocts are colonial, and all but 2 of the 150 species are marine. A few solitary species can move slowly.
Some species eject unfertilized ova into the water while others keep their ova in brood chambers until they hatch, and some of these species use placenta-like organs to nourish the developing eggs. After hatching, the larvae swim for a short time and then settle on a surface. There they metamorphose, and the larval gut generally rotates by up to 180°, so that the mouth and anus face upwards. Both colonial and solitary species also reproduce by cloning – solitary species grow clones in the space between the tentacles and then release them when developed, while colonial ones produce new members from the stalks or from corridor-like stolons.
Some species of nudibranchs ("sea slugs") and turbellarian flatworms prey on entoprocts. A few entoproct species have been found living in close association with other animals. It is uncertain whether any are invasive species.
Fossils of entoprocts are very rare and the earliest specimens that have been identified with confidence date from the Late Jurassic. Most studies from 1996 onwards have regarded entoprocts as members of the Trochozoa, which also includes molluscs and annelids. However, a study in 2008 concluded that entoprocts are closely related to bryozoans.
Contents |
"Entoprocta", coined in 1870,[1] means "anus inside".[2] The alternative name "Kamptozoa", meaning "bent" or "curved" animals,[3] was assigned in 1929.[1] Some authors use "Entoprocta",[4][5] while others prefer "Kamptozoa".[2][6]
Most species are colonial, and their members are known as "zooids",[2] since they are not fully-independent animals.[7] Zooids are typically 1 millimetre (0.039 in) long but ranging from 0.1 to 7 millimetres (0.0039 to 0.28 in) long.[2]
Entoprocts are superficially like bryozoans (ectoproct)s, as both groups have a "crown" of tentacles whose cilia generate water currents that draw food particles towards the mouth. However, they have different feeding mechanisms and internal anatomy, and ectoprocts undergo a a metamorphosis from larva to adult that destroys most of the larval tissues, and their colonies also have a founder zooid that is different from its "daughters".[2][8]
| Entoprocta[2] | Bryozoa (Ectoprocta)[8] | |
|---|---|---|
| Tentacles | Solid | Hollow |
| Feeding current | From bases to tips of tentacles | From tips to bases of tentacles |
| Position of anus | Inside "crown" of tentacles | Outside "crown" of tentacles |
| Coelom | none | Three-part |
| Shape of founder zooid in a colony | Same as other zooids | Round, unlike normal zooids[9] |
| Metamorphosis to adult | Retains most larval structures | Destroys most larval structures |
| Excretory organs | Protonephridia | None |
The body of a mature entoproct zooid has a goblet-like structure with a calyx mounted on a relatively long stalk that attaches to a surface. The rim of the calyx bears a "crown" of 8 to 30 solid tentacles, which are extensions of the body wall. The base of the "crown" of tentacles is surrounded by a membrane that partially covers the tentacles when they retract. The mouth and anus lie on opposite sides of the atrium (space enclosed by the "crown" of tentacles), and both can be closed by sphincter muscles. The gut is U-shaped, curving down towards the base of the calyx, where it broadens to form the stomach. This is lined with a membrane consisting of a single layer of cells, each of which has multiple cilia.[2]
The stalks of colonial species arise from shared attachment plates or from a network of stolons, tubes that run across a surface.[2] In solitary species the stalk ends in a muscular sucker, or a flexible foot, or is cemented to a surface.[5] The stalk is muscular and produces a characteristic nodding motion. In some species it is segmented. Some solitary species can move, either by creeping on the muscular foot or by somersaulting.[2]
The body wall consists of the epidermis and an external cuticle,[2] which consists mainly of criss-cross collagen fibers. The epidermis contains only a single layer of cells, each of which bears multiple cilia ("hairs") and microvilli (tiny "pleats") that penetrate through the cuticle.[2] The stolons and stalks of colonial species have thicker cuticles, stiffened with chitin.[5]
There is no coelom (internal fluid-filled cavity lined with peritoneum) and the other internal organs are embedded in connective tissue that lies between the stomach and the base of the "crown" of tentacles. The nervous system runs through the connective tissue and just below the epidermis, and is controlled by a pair of ganglia. Nerves run from these to the calyx, tentacles and stalk, and to sense organs in all these areas.[2]
A band of cells, each with multiple cilia, runs along the sides of the tentacles, connecting each tentacle to its neighbors, except that there is a gap in the band nearest the anus. A separate band of cilia runs along a groove that runs close to the inner side of the base of the "crown", with a narrow extension up the inner surface of each tentacle.[5] The cilia on the sides of the tentacles create a current that flows into the "crown" at the bases of the tentacles and exits above the center of the "crown".[2] These cilia pass food particles to the cilia on the inner surface of the tentacles, and the inner cillia produce a downward current that drives particles into and round the groove, and then to the mouth.[5]
Entoprocts generally use one or both of: ciliary sieving, in which one band of cilia creates the feeding current and another traps food particles (the "seive"); and downstream collecting, in which food articles are trapped as they are about to exit past them. In entoprocts, downstream collecting is done by the same bands of cilia that generate the current; trochozoan larvae also use downstream collecting, but use a separate set of cilia to trap food particles.[10]
In addition glands in the tentacles secrete sticky threads that capture large particles.[2] A non-colonial species reported from around the Antarctic Peninsula in 1993 has cells that superficially resemble the cnidocytes of cnidaria, and fire sticky threads. These unusual cells lie around the mouth, and may provide an additional means of capturing prey.[11]
The stomach and intestine are lined with microvilli, which are thought to absorb nutrients. The anus, which opens inside the "crown", ejects solid wastes into the outgoing current after the tentacles have filtered food out of the water; in some families it is raised on a cone above the level of the groove that conducts food to the mouth.[2][12] Most species have a pair of protonephridia which extract soluble wastes from the internal fluids and eliminate them through pores near the mouth. However the freshwater species Urnatella gracilis has multiple nephridia in the calyx and stalk.[2]
The zooids absorb oxygen and emit carbon dioxide by diffusion,[2] which works well for small animals.[13]
Most species are simultaneous hermaphrodites, but some switch from males to females as they mature, while individuals of some species remain of the same sex all their lives. Individuals have one or two pairs of gonads, placed between the atrium and stomach, and opening into a single gonopore in the atrium.[5] Scientists think the eggs are fertilized in the ovaries. Most species release eggs that hatch into planktonic larvae, but a few brood their eggs in the gonopore. Those that brood small eggs nourish them by a placenta-like organ, while larvae of species with larger eggs live on stored yolk.[2] The development of the fertilized egg into a larva follows a typical spiralian pattern: the cells divide by spiral cleavage, and mesoderm develops from a specific cell labelled "4d" in the early embryo.[15] There is no coelom at any stage.[2]
In some species the larva is a trochophore which is planktonic and feeds on floating food particles by using the two bands of cilia round its "equator" to sweep food into the mouth, which uses more cilia to drive them into the stomach, which uses further cilia to expel undigested remains through the anus.[16] In some species of the genera Loxosomella and Loxosoma, the larva produces one or two buds that separate and form new individuals, while the trochophore disintegrates. However, most produce a larva with sensory tufts at the top and front, a pair of pigment-cup ocelli ("little eyes"), a pair of protonephridia, and a large, cilia-bearing foot at the bottom.[5] After settling, the foot and frontal tuft attach to the surface. Larvae of most species undergo a complex metamorphosis, and the internal organs may rotate by up to 180°, so that the mouth and anus point upwards.[2]
All species can produce clones by budding. Colonial species produce new zooids from the stolon or from the stalks, and can form large colonies in this way.[2] In solitary species, clones form on the floor of the atrium, and are released when their organs are developed.[5]
The phylum consists of about 150 recognized species, grouped into 4 families:[2][4]
| Family | Barentsiidae | Pedicellinidae | Loxokalypodidae | Loxosomatidae |
|---|---|---|---|---|
| Genera | Barentsia, Coriella, Pedicellinopsis, Pseudopedicellina, Urnatella[17] | Chitaspis, Loxosomatoides, Myosoma, Pedicellina[18] | Loxokalypus[19] | Loxocore, Loxomitra, Loxosoma, Loxosomella, Loxosomespilon[20] |
| Colonial[6] | Colonial | Solitary | ||
| Septum between calyx and stalk[6] | Yes | no | ||
| Star-cell organ[6] | Yes | no | ||
| Anus on cone[2] | no | Yes | ||
| Stolons present[6] | Yes | No, colonies grow on shared baseplate | not colonial | |
| Segmented stems[6][2] | Yes | no | ||
All species are sessile.[2] While the great majority are marine, two species live in freshwater, Loxosomatoides sirindhornae reported in 2004 in central Thailand and Urnatella gracilis found in in all the continents except Antarctica.[1] Colonial species are found in all the oceans, living on rocks, shells, algae and underwater buildings.[2] The solitary species, which are marine,[1] live on other animals that feed by producing water currents, such as sponges, ectoprocts and sessile annelids. [5]
Some species of nudibranchs ("sea slugs") and turbellarian flatworms prey on entoprocts.[21]
Small colonies of the freshwater entoproct Urnatella gracilis have been found living on the aquatic larvae of the dobsonfly Corydalus cornutus. The ectoprocts gain a means of dispersal, protection from predators and possibly a source of water that is rich in oxygen and nutrients, as colonies often live next to the gills of the larval flies.[22] In the White Sea, the non-colonial entoproct Loxosomella nordgaardi prefers to live attached to bryozoan (ectoproct) colonies, mainly on the edges of colonies or in the "chimneys", gaps by which large bryozoan colonies expel water from which they have seived food. Observation suggests that both the entoprocts and the bryozoans benefit from the association: each enhances the water flow that the other needs for feeding; and the longer cilia of the entoprocts may help them to capture different food from that caught by the bryozoans, so that the animals do not compete for the same food.[23]
Entoprocts are small and have been little studied by zoologists. Hence it is hard to determine whether a specimen belongs to a species that already occurs in the same area or is an invader, possibly as a result of human activities.[24]
Since entoprocts are small and soft-bodied, fossils have been extremely rare.[26] In 1977 Simon Conway Morris provided the first description of Dinomishcus, a sessile animal with calyx, stalk and holdfast, found in Canada's Burgess Shale, which was formed about . Morris regarded this animal as the earliest known entoproct, since its mouth and anus lay inside a ring of structures above the calyx, but noted that these structures were flat and rather stiff, while the tentacles of modern entoprocts are flexible and have a round cross-section.[25]
In 1992 J.A. Todd and P.D. Taylor concluded that Dinomishcus was not an entoproct, because it did not have the typical rounded, flexible tentacles and the fossils showed no other features that clearly resembled those of entoprocts. In their opinion the earliest fossil entoprocts were specimens they found from Late Jurassic rocks in England. These resemble the modern colonial genus Barentsia in many ways, including: upright zooids linked by a network of stolons encrusting the surface to which the colony is attached; straight stalks joined to the stolons by bulky sockets with transverse bands of wrinkles; overall size and proportions similar to that of modern species of Barentsia.[26]
When entoprocts were discovered in the nineteenth 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 embryos, caused scientists to regard the two groups as separate phyla.[27] "Bryozoa" then became just an alternative name for ectoprocts, in which the anus is outside the feeding organ.[28] However studies by one team in 2007 and 2008 [27][29]
The consensus of studies from 1996 onwards has been that entoprocts are part of the Trochozoa, a protostome "superphylum" whose members are united in having as their most basic larval form the trochophore type. The trochozoa also include molluscs, annelids, flatworms, nemertines and others. However, scientists disagree about which phylum is mostly closely related to enctoprocts within the trochozoans.[30] An analysis in 2008 re-introduced the pre-1869 meaning of the term "Bryozoa", for a group in which entoprocts and ectoprocts are each other's closest relatives.[27]
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| Endoprocta (invertebrate zoology) | |
| Kamptozoa (invertebrate zoology) | |
| Calyssozoa (invertebrate zoology) |
| Apa Struktur dan fungsi tubuh pada entoprocta? |
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