Entoprocta (), or Kamptozoa , commonly called goblet worms, is a phylum of mostly-sessile aquatic animals, ranging from long. Mature individuals are goblet-shaped, on relatively long stalks. They have a "crown" of solid tentacles whose cilia generate water currents that draw food particles towards the mouth, and 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 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.

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. Other studies place them in a clade Tetraneuralia, together with molluscs.

Names

"Entoprocta", coined in 1870, means "anus inside". The alternative name "Kamptozoa", meaning "bent" or "curved" animals, was assigned in 1929. since they are not fully independent animals. Zooids are typically long but range from long.

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! Metamorphosis to adult

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Zooids

thumb|280px|Barentsia laxa

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. 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.

In addition, glands in the tentacles secrete sticky threads that capture large particles.

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. 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.

Reproduction and life cycle

Most species are simultaneous hermaphrodites, but some switch from male to female 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. There is no coelom at any stage. 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.

| Chitaspis, Loxosomatoides, Myosoma, Pedicellina

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| Loxocore, Loxomitra, Loxosoma, Loxosomella,<!-- ZoolSci25:1171. --> Loxosomespilon

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! Colonial

| colspan="3" | Colonial || Solitary

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! Septum between calyx and stalk

In 1992 J.A. Todd and P.D. Taylor concluded that Dinomischus 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.

Another species, Cotyledion tylodes, first described in 1999, was larger than extant entoprocts, reaching 8–56&nbsp;mm in height, and unlike modern species, was "armored" with sclerites, scale-like structures. C. tylodes did have a similar sessile lifestyle to modern entoprocts. The identified fossils of C. tylodes were found in 520-million-year-old rocks from southern China. This places early entoprocts in the period of the Cambrian explosion.

Family tree

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. "Bryozoa" then became just an alternative name for ectoprocts, in which the anus is outside the feeding organ. However, studies by one team in 2007 and 2008 argue for sinking Entoprocta into Bryozoa as a class, and resurrecting Ectoprocta as a name for the currently identified bryozoans.

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 entoprocts within the trochozoans. 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.

Interaction with other organisms

Some species of nudibranchs ("sea slugs"), particularly those of the genus Trapania, as well as turbellarian flatworms, prey on entoprocts.

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. 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 sieved 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.

Entoprocts are small and have been little studied by zoologists. Hence it is difficult 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.

References

Further reading

  • Information on Entoprocta from earthlife.net