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ADW: Entoprocta: INFORMATION
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Marine species are found throughout the world鈥檚 oceans, and <a class="taxon-link rank-species" href="/accounts/Urnatella_gracilis/">Urnatella gracilis</a> is found on all continents except Antarctica. These organisms superficially resemble cnidarian hydroids and bryozoans (Phylum <a class="taxon-link rank-phylum" href="/accounts/Bryozoa/">Bryozoa</a>); the main body consists of a cup-like calyx that bears a ring of ciliated tentacles, which is attached on its dorsal surface to the substrate (or stolon in colonial species) by a long, thin stalk. Whether solitary or colonial, all entoprocts are sessile suspension feeders. Entoprocts exhibit a range of reproductive modes and behaviors, including asexual clonal reproduction, as well as hermaphroditic and dioecious sexual reproduction. <span class="citations"> (<a href="#874637AB-7DE7-11E2-B377-002500F14F28" class="citation">Appeltans, et al., 2012</a>; <a href="#0FCD0A17-7DE7-11E2-A279-002500F14F28" class="citation">Brusca and Brusca, 2003</a>; <a href="#A9D68773-B1A7-11E2-9BFE-002500F14F28" class="citation">Nielsen, 2002</a>; <a href="#3AE5F4E3-7DE7-11E2-8E1C-002500F14F28" class="citation">Ruppert, et al., 2004</a>; <a href="#7E83343A-B1A2-11E2-808D-002500F14F28" class="citation">Schwaha, et al., 2010</a>; <a href="#BFE4DA6B-7DE9-11E2-84B6-002500F14F28" class="citation">Zhang, 2011</a>)</span></p> </section> <section class="hyphenate"> <h3 id="geographic_range">Geographic Range</h3> <p audience="advanced intermediate">The majority of <span class="taxon-name">entoprocts</span> live in coastal, marine environments throughout the world. <a class="taxon-link rank-species" href="/accounts/Urnatella_gracilis/">Urnatella gracilis</a> is found in rivers on every continent except Antarctica, and <span rank="Species" class="taxon-name rank-species">Loxosomatoides sirindhorne</span> has been identified from rivers in Thailand. <span class="citations"> (<a href="#0FCD0A17-7DE7-11E2-A279-002500F14F28" class="citation">Brusca and Brusca, 2003</a>; <a href="#C1003402-B1A4-11E2-ACA6-002500F14F28" class="citation">Ramel, 2012</a>; <a href="#7E83343A-B1A2-11E2-808D-002500F14F28" class="citation">Schwaha, et al., 2010</a>; <a href="#EADBD5EE-7DE8-11E2-9D21-002500F14F28" class="citation">Wood, 2005</a>)</span></p> <ul class="keywords donthyphenate "> <li class="keywords-header">Biogeographic Regions</li> <li> <a class="gloss" href="#20020914235803">nearctic</a> <ul> <li> <a class="gloss" href="#20020904145365">native</a> </li> </ul> </li> <li> <a class="gloss" href="#20020915001330">palearctic</a> <ul> <li> <a class="gloss" href="#20020904145365">native</a> </li> </ul> </li> <li> <a class="gloss" href="#20020915001348">oriental</a> <ul> <li> <a class="gloss" href="#20020904145365">native</a> </li> </ul> </li> <li> <a class="gloss" href="#20020916134510">ethiopian</a> <ul> <li> <a class="gloss" href="#20020904145365">native</a> </li> </ul> </li> <li> <a class="gloss" href="#20020915000403">neotropical</a> <ul> <li> <a class="gloss" href="#20020904145365">native</a> </li> </ul> </li> <li> <a class="gloss" href="#20020916135737">australian</a> <ul> <li> <a class="gloss" href="#20020904145365">native</a> </li> </ul> </li> </ul> <ul class="keywords donthyphenate last"> <li class="keywords-header">Other Geographic Terms</li> <li> <a class="gloss" href="#200304045530">holarctic</a> </li> <li> <a class="gloss" href="#20020904145620">cosmopolitan</a> </li> </ul> </section> <section class="hyphenate"> <h3 id="habitat">Habitat</h3> <p audience="advanced intermediate">Colonial <span class="taxon-link">entoprocts</span> are found living on rocks, shells, algae, and underwater structures, preferring more solid substrates to softer sediments. Solitary species are found living on other animals such as sponges, sessile annelids, and ectoprocts, taking advantage of the feeding currents produced by these animals. They are found from intertidal areas to depths of up to 500 m. While marine species may be found in brackish environments, the two freshwater species are found only in fully freshwater environments. <span class="citations"> (<a href="#0FCD0A17-7DE7-11E2-A279-002500F14F28" class="citation">Brusca and Brusca, 2003</a>; <a href="#A9D68773-B1A7-11E2-9BFE-002500F14F28" class="citation">Nielsen, 2002</a>; <a href="#3AE5F4E3-7DE7-11E2-8E1C-002500F14F28" class="citation">Ruppert, et al., 2004</a>; <a href="#F1A1720C-B1AB-11E2-925A-002500F14F28" class="citation">Weise, 1961</a>; <a href="#EADBD5EE-7DE8-11E2-9D21-002500F14F28" class="citation">Wood, 2005</a>)</span></p> <ul class="keywords donthyphenate "> <li class="keywords-header">Habitat Regions</li> <li> <a class="gloss" href="#20020904145595">temperate</a> </li> <li> <a class="gloss" href="#20020904145598">tropical</a> </li> <li> <a class="gloss" href="#20020904145582">saltwater or marine</a> </li> <li> <a class="gloss" href="#20020904145747">freshwater</a> </li> </ul> <ul class="keywords donthyphenate "> <li class="keywords-header">Aquatic Biomes</li> <li> <a class="gloss" href="#20020904145431">benthic</a> </li> <li> <a class="gloss" href="#20020904145675">reef</a> </li> <li> <span>rivers and streams</span> </li> <li> <a class="gloss" href="#20020904145825">coastal</a> </li> <li> <a class="gloss" href="#20020904145435">brackish water</a> </li> </ul> <ul class="keywords donthyphenate last"> <li class="keywords-header">Other Habitat Features</li> <li> <a class="gloss" href="#20020904145633">intertidal or littoral</a> </li> </ul> </section> <section class="hyphenate"> <h3 id="taxonomic_history">Systematic and Taxonomic History</h3> <p audience="advanced intermediate">Following their discovery, these organisms were alternatively classified as special, polyp-like rotifers, or they were classified along with <a class="taxon-link rank-phylum" href="/accounts/Bryozoa/">Bryozoa</a>, in phylum <a class="taxon-link rank-phylum" href="/accounts/Bryozoa/">Bryozoa</a>. The name <span class="taxon-link rank-unspecified">Entoprocta</span> was first applied to these organisms by the German zoologist Hinrich Nitsche in 1870 to separate them from <a class="taxon-link rank-phylum" href="/accounts/Bryozoa/">Bryozoa</a>, based upon the position of the anus (inside the lophophore ring, rather than outside), embryonic cleavage pattern (spiral vs. radial), and differences in body cavity type (acoelomate vs. coelomate). These differences resulted in Hatschek elevating <span class="taxon-link rank-unspecified">Entoprocta</span> to phylum level in 1888. The name <span class="taxon-name">Kamptozoa</span> would later be applied to this group by Carl Cori in 1929, and is still used interchangeably with <span class="taxon-link rank-unspecified">Entoprocta</span>. <span class="citations"> (<a href="#1B3104B0-4494-11E3-887F-002500F14F28" class="citation">Cori, 1929</a>; <a href="#DC8A1821-44A3-11E3-9070-002500F14F28" class="citation">Ellis, 1755</a>; <a href="#3AB82FD9-449F-11E3-B181-002500F14F28" class="citation">Hatschek, 1888</a>; <a href="#5E180C82-4498-11E3-A3AC-002500F14F28" class="citation">Nitsche, 1870</a>; <a href="#7C80210A-449F-11E3-BCED-002500F14F28" class="citation">Pallas, 1774</a>; <a href="#2627487D-44A4-11E3-930B-002500F14F28" class="citation">Sars, 1835</a>; <a href="#911BCEE1-44A2-11E3-8D1E-002500F14F28" class="citation">van Beneden, 1845</a>)</span></p> <p audience="advanced intermediate">The monophyly of <span class="taxon-link rank-unspecified">Entoprocta</span>, as well as that of its two orders, <span class="taxon-name">Solitaria</span> and <span class="taxon-name">Coloniales</span>, is well supported by morphological characters and a recent molecular phylogenetic analysis. However, the evolutionary relationships of entoprocts to other phyla remain a matter of some debate. Recent molecular phylogenetic studies have suggested that entoprocts and ectoprocts should in fact be considered classes within a single taxon called <a class="taxon-link rank-genus" href="/accounts/Polyzoa/">Polyzoa</a>, in agreement with their original classification. These, and additional molecular studies have suggested that <a class="taxon-link rank-genus" href="/accounts/Polyzoa/">Polyzoa</a> should also include the obscure phylum <a class="taxon-link rank-unspecified" href="/accounts/Cycliophora/">Cycliophora</a>, which appears to share a sister relationship with <span class="taxon-link rank-unspecified">Entoprocta</span>. These studies also suggest that <a class="taxon-link rank-genus" href="/accounts/Polyzoa/">Polyzoa</a> be placed in a larger superphylum, <a class="taxon-link rank-unspecified" href="/accounts/Lophotrochozoa/">Lophotrochozoa</a>, based on the common presence of the trochophore larval state in member phyla (which include annelid worms, mollusks, flatworms, and nemertean worms, among others). There is no consensus, however, on which of the lophotrochozoan phyla represent the closest relative of modern polyzoans. <span class="citations"> (<a href="#0C076851-4492-11E3-89A1-002500F14F28" class="citation">Fuchs, et al., 2010</a>; <a href="#AD67661E-426C-11E3-914A-002500F14F28" class="citation">Funch and Kristensen, 1995</a>; <a href="#EBA00845-426C-11E3-8975-002500F14F28" class="citation">Giribet, et al., 2004</a>; <a href="#50F21C6B-7DE8-11E2-A5A8-002500F14F28" class="citation">Hausdorf, et al., 2007</a>; <a href="#57DC2CE1-44A7-11E3-9054-002500F14F28" class="citation">Hejnol, et al., 2009</a>; <a href="#A57EB914-7DE8-11E2-B772-002500F14F28" class="citation">Helmkampf, et al., 2008</a>; <a href="#26C4AB78-44A9-11E3-BF56-002500F14F28" class="citation">Paps, et al., 2009a</a>; <a href="#AF251028-44A9-11E3-AF4A-002500F14F28" class="citation">Paps, et al., 2009b</a>)</span></p> <ul class="aside block-grid donthyphenate two-up"> <li> <dl> <dt>Synonyms</dt> <dd> <ul> <li><span class="taxon-name">Kamptozoa</span> (Cori, 1929)</li> </ul> </dd> </dl> </li> <li> <dl> <dt>Synapomorphies</dt> <dd> <ul> <li>anus located inside ring of lophophore tentacles</li> </ul> </dd> </dl> </li> </ul> </section> <section class="hyphenate"> <h3 id="physical_description">Physical Description</h3> <p audience="advanced intermediate">These organisms superficially resemble cnidarian hydroids and bryozoans (phylum <a class="taxon-link rank-phylum" href="/accounts/Bryozoa/">Bryozoa</a>), with the main body consisting of a cup-like calyx that bears a ring of 6 to 36 ciliated tentacles, attached on its dorsal surface to the substrate (or stolon in colonial species) by a long, thin stalk. The calyx and stalk are covered by a thin, collagenous cuticle, which does not extend over the tentacles, and is underlaid by a cellular epidermis. Muscle bands beneath the epidermis allow the organism to compress its calyx and stalk, extend its tentacles, and bend its stalk. These organisms are functionally acoelomate, lacking a fluid-filled body cavity. It is uncertain whether this condition results from a secondary filling of an embryonic blastocoel with mesenchyme, or whether these organisms lack a blastocoel throughout development. The viscera are located entirely in the calyx, with the mouth and anus found on the ventral surface (vestibule), surrounded by the tentacular crown. The gut is U-shaped and lined with a layer of ciliated cells, widening into a stomach near the internal base of the calyx, followed by an intestinal segment that leads to the anus. Entoprocts are quite small, with individual zooids ranging from 0.1 to 7 mm in length. <span class="citations"> (<a href="#0FCD0A17-7DE7-11E2-A279-002500F14F28" class="citation">Brusca and Brusca, 2003</a>; <a href="#C1003402-B1A4-11E2-ACA6-002500F14F28" class="citation">Ramel, 2012</a>; <a href="#3AE5F4E3-7DE7-11E2-8E1C-002500F14F28" class="citation">Ruppert, et al., 2004</a>)</span></p> <ul class="keywords donthyphenate "> <li class="keywords-header">Other Physical Features</li> <li> <a class="gloss" href="#20020916125902">heterothermic</a> </li> <li> <a class="gloss" href="#20020904145642">bilateral symmetry</a> </li> </ul> <ul class="keywords donthyphenate last"> <li class="keywords-header">Sexual Dimorphism</li> <li> <span>sexes alike</span> </li> </ul> </section> <section class="hyphenate"> <h3 id="development">Development</h3> <p audience="advanced intermediate">Both colonial and solitary species are capable of asexual clonal reproduction by budding; species may reproduce sexually and be dioecious, simultaneous, or protandrous hermaphrodites. Eggs either hatch into planktonic larvae, or in some species, complete early development in a brood chamber (located in the atrium), attached by secretions of cement glands to the chamber's wall. In some brooding species, nutrition is provided to developing embryos via special placental cells; in others, the egg yolk nourishes the growing larvae. Planktonic larvae may remain free swimming for up to 7 months before settling, while those raised in a brood chamber will settle relatively quickly (within a few days of hatching). <span class="citations"> (<a href="#0FCD0A17-7DE7-11E2-A279-002500F14F28" class="citation">Brusca and Brusca, 2003</a>; <a href="#3AE5F4E3-7DE7-11E2-8E1C-002500F14F28" class="citation">Ruppert, et al., 2004</a>; <a href="#790754CA-B26F-11E2-9028-002500F14F28" class="citation">Shanks, 2001</a>)</span></p> <p audience="advanced intermediate">Entoproct embryonic development follows the holoblastic, spiral cleavage pattern typical of protostome organisms, with the mesoderm forming from the 4d mesentoblast. Development continues to a coeloblastula stage, after which, the embryos of most species proceed into a free swimming, feeding planktonic larval stage strongly resembling the trochophore larva of protostome species. Some species produce lecithotrophic or benthic crawling larvae. Trochophore larvae have equatorial ciliary bands (used for feeding on suspended particles), apical and ventral sensory tufts of cilia, pigment-cup ocelli that serve as light-sensing organs, a complete digestive system, and a pair of protonephridia for waste excretion. After their larval period, most entoprocts settle, attach to the substrate, and undergo metamorphosis. Unequal growth of the body directs the vestibular surface away from the substrate and the mouth, anus, and gut may rotate up to 180掳, to face the vestibular surface; however, no rotation or unequal growth is necessary for some species. In these, an asexual bud forms from the attached larvae to form the zooid, which is already oriented in the correct position. <span class="citations"> (<a href="#0FCD0A17-7DE7-11E2-A279-002500F14F28" class="citation">Brusca and Brusca, 2003</a>; <a href="#A9D68773-B1A7-11E2-9BFE-002500F14F28" class="citation">Nielsen, 2002</a>)</span></p> <p audience="advanced intermediate">Some species in family <a class="taxon-link rank-family" href="/accounts/Loxosomatidae/">Loxosomatidae</a> produce free swimming larvae that may produce adult buds precociously; the adults are held in a body pocket of the larvae until their release through the body wall, usually a few days. After release, the larvae die. In some species, adult buds have been noticed developing from larvae while the larvae were still developing in their parents' ovaries. In at least one species, males and females may be produced via budding. <span class="citations"> (<a href="#790754CA-B26F-11E2-9028-002500F14F28" class="citation">Shanks, 2001</a>)</span></p> <ul class="keywords donthyphenate last"> <li class="keywords-header">Development - Life Cycle</li> <li> <a class="gloss" href="#20020904145364">metamorphosis</a> </li> </ul> </section> <section class="hyphenate"> <h3 id="reproduction">Reproduction</h3> <p audience="advanced intermediate">Gonads, when present, are located just beneath the vestibular surface and empty into the water via the gonopore. Male zooids release sperm into the water, where it can be drawn into the reproductive tract of female zooids. Fertilization occurs in the ovaries or oviducts. <span class="citations"> (<a href="#0FCD0A17-7DE7-11E2-A279-002500F14F28" class="citation">Brusca and Brusca, 2003</a>)</span></p> <ul class="keywords donthyphenate last"> <li class="keywords-header">Mating System</li> <li> <a class="gloss" href="#20020904145483">polygynandrous (promiscuous)</a> </li> </ul> <p audience="advanced intermediate"><span class="taxon-link">Entoprocts</span> exhibit a wide range of reproductive modes and behaviors. Both colonial and solitary species are capable of asexual clonal reproduction by budding. In colonial species, new zooids are formed from the stalk or the stolons joining individuals, while solitary species produce buds from the base of the calyx. Sexual reproduction also occurs and individuals may be dioecious, simultaneous, or protandrous hermaphrodites. Mating seasons depend on location; entoprocts from warmer waters may reproduce year-round, while many of those in more temperate waters show a breeding peak in late summer and early fall (experimental evidence supports increased water temperature as a trigger for sexual maturity). <span class="citations"> (<a href="#0FCD0A17-7DE7-11E2-A279-002500F14F28" class="citation">Brusca and Brusca, 2003</a>; <a href="#03A9324C-B275-11E2-880F-002500F14F28" class="citation">Emschermann, 1985</a>; <a href="#A9D68773-B1A7-11E2-9BFE-002500F14F28" class="citation">Nielsen, 2002</a>; <a href="#7552844A-B271-11E2-915E-002500F14F28" class="citation">Winston, 1982</a>)</span></p> <ul class="keywords donthyphenate last"> <li class="keywords-header">Key Reproductive Features</li> <li> <a class="gloss" href="#20020904145584">seasonal breeding</a> </li> <li> <a class="gloss" href="#20020904145698">year-round breeding</a> </li> <li> <span>gonochoric/gonochoristic/dioecious (sexes separate)</span> </li> <li> <span>simultaneous hermaphrodite</span> </li> <li> <span>sequential hermaphrodite</span> <ul> <li> <a class="gloss" href="#20020904145373">protandrous</a> </li> </ul> </li> <li> <a class="gloss" href="#20020904145786">sexual</a> </li> <li> <a class="gloss" href="#20020904145314">asexual</a> </li> <li> <a class="gloss" href="#20020904145546">fertilization</a> <ul> <li> <a class="gloss" href="#20020904145427">internal</a> </li> </ul> </li> <li> <a class="gloss" href="#20020904145572">oviparous</a> </li> </ul> <p audience="advanced intermediate">Beyond production of gametes and, in some cases, development of larvae in a brooding pouch (sometimes fed by placental cells), <span class="taxon-link">entoprocts</span> do not display parental investment. <span class="citations"> (<a href="#0FCD0A17-7DE7-11E2-A279-002500F14F28" class="citation">Brusca and Brusca, 2003</a>; <a href="#3AE5F4E3-7DE7-11E2-8E1C-002500F14F28" class="citation">Ruppert, et al., 2004</a>; <a href="#790754CA-B26F-11E2-9028-002500F14F28" class="citation">Shanks, 2001</a>)</span></p> <ul class="keywords donthyphenate last"> <li class="keywords-header">Parental Investment</li> <li> <span>no parental involvement</span> </li> <li> <span>pre-hatching/birth</span> <ul> <li> <span>provisioning</span> <ul> <li> <span>female</span> </li> </ul> </li> <li> <span>protecting</span> <ul> <li> <span>female</span> </li> </ul> </li> </ul> </li> </ul> </section> <section class="hyphenate"> <h3 id="lifespan_longevity">Lifespan/Longevity</h3> <p audience="advanced intermediate">Although no definitive studies regarding the lifespan of <span class="taxon-link">entoprocts</span> are currently available, feeding planktonic larvae of some species live for more than 7 months (at which point they settle and attach to the substrate). <span class="citations"> (<a href="#03A9324C-B275-11E2-880F-002500F14F28" class="citation">Emschermann, 1985</a>; <a href="#790754CA-B26F-11E2-9028-002500F14F28" class="citation">Shanks, 2001</a>)</span></p> </section> <section class="hyphenate"> <h3 id="behavior">Behavior</h3> <p audience="advanced intermediate"><span class="taxon-link">Entoprocts</span> may be colonial or solitary. An entoproct may shed (typically under adverse environmental conditions) and regenerate its calyx, sometimes changing sex when it does; most entoprocts with this ability are colonial (only one species of solitary entoproct, <span rank="Species" class="taxon-name rank-species">Loxosomella antarctia</span> is noted to regenerate the entire calyx in this way). It is possible to fold the tentacles and move the calyx using subepidermal musculature of the body and stalk. Once larvae have settled, they are most typically sessile, using a "foot" to attach to a substrate. A few species are reportedly capable of movement even after adulthood, for instance members of genus <span rank="Genus" class="taxon-name rank-genus">Loxasomella</span> inside the tubes of marine annelids. <span class="citations"> (<a href="#0FCD0A17-7DE7-11E2-A279-002500F14F28" class="citation">Brusca and Brusca, 2003</a>; <a href="#409A4661-B27E-11E2-B2AD-002500F14F28" class="citation">Emschermann, 1993</a>; <a href="#449105AB-B278-11E2-B909-002500F14F28" class="citation">Margulis, et al., 1999</a>; <a href="#C1003402-B1A4-11E2-ACA6-002500F14F28" class="citation">Ramel, 2012</a>)</span></p> <ul class="keywords donthyphenate last"> <li class="keywords-header">Key Behaviors</li> <li> <a class="gloss" href="#20020904145414">diurnal</a> </li> <li> <a class="gloss" href="#20020904145503">nocturnal</a> </li> <li> <a class="gloss" href="#20020904145752">crepuscular</a> </li> <li> <a class="gloss" href="#20020904145586">sessile</a> </li> <li> <a class="gloss" href="#20020904145472">motile</a> </li> <li> <a class="gloss" href="#20020904145585">sedentary</a> </li> <li> <a class="gloss" href="#20020904145381">solitary</a> </li> <li> <a class="gloss" href="#20020904145440">colonial</a> </li> </ul> </section> <section class="hyphenate"> <h3 id="communication">Communication and Perception</h3> <p audience="advanced intermediate"><span class="taxon-link">Entoprocts</span> have greatly reduced nervous systems, consisting of a single subenteric ganglion between the vestibular surface and stomach. This enervates the tentacles, calyx, and stalk via several pairs of nerves arising from the central ganglionic mass. Tactile receptors are scattered over the body and concentrated on the tentacles. Some species in family <a class="taxon-link rank-family" href="/accounts/Loxosomatidae/">Loxosomatidae</a> have ciliated papillae, which form lateral sense organs as well. <span class="citations"> (<a href="#0FCD0A17-7DE7-11E2-A279-002500F14F28" class="citation">Brusca and Brusca, 2003</a>; <a href="#3AE5F4E3-7DE7-11E2-8E1C-002500F14F28" class="citation">Ruppert, et al., 2004</a>)</span></p> <ul class="keywords donthyphenate "> <li class="keywords-header">Communication Channels</li> <li> <a class="gloss" href="#20020904145500">tactile</a> </li> </ul> <ul class="keywords donthyphenate last"> <li class="keywords-header">Perception Channels</li> <li> <a class="gloss" href="#20020904145500">tactile</a> </li> </ul> </section> <section class="hyphenate"> <h3 id="food_habits">Food Habits</h3> <p audience="advanced intermediate">All <span class="taxon-link">entoprocts</span> are sessile suspension feeders. The cilia on their tentacles generate water currents that carry small food particles (particularly phytoplankton, as well as diatoms and algae) to a ciliated food groove at the base of the tentacular crown. The tentacles also have glandular cells that secrete a sticky substance to capture larger food particles, which are also moved to the food groove. The cilia in the food groove carry food to the mouth, where a combination of ciliary action and muscle contractions move the food through the digestive system. One species of entoproct, <a class="taxon-link rank-species" href="/accounts/Loxocorone_brochobola/">Loxocorone brochobola</a>, has nematocyst-type organs; their function is unknown but may be related to feeding. <span class="citations"> (<a href="#0FCD0A17-7DE7-11E2-A279-002500F14F28" class="citation">Brusca and Brusca, 2003</a>; <a href="#409A4661-B27E-11E2-B2AD-002500F14F28" class="citation">Emschermann, 1993</a>; <a href="#F1A1720C-B1AB-11E2-925A-002500F14F28" class="citation">Weise, 1961</a>)</span></p> <ul class="keywords donthyphenate last"> <li class="keywords-header">Primary Diet</li> <li> <a class="gloss" href="#20020904145368">planktivore</a> </li> </ul> </section> <section class="hyphenate"> <h3 id="predation">Predation</h3> <p audience="advanced intermediate">As very small invertebrates, <span class="taxon-link">entoprocts</span> are likely prey to many larger marine and freshwater animals. Known predators include small <a class="taxon-link" href="/accounts/Crustacea/">crustaceans</a> and <a class="taxon-link" href="/accounts/Mollusca/">mollusks</a>, <a class="taxon-link" href="/accounts/Platyhelminthes/">flatworms</a>, and <a class="taxon-link" href="/accounts/Gastropoda/">nudibranchs</a>. <span class="citations"> (<a href="#B499BC8F-B336-11E2-BF71-002500F14F28" class="citation">Canning and Carlton, 2000</a>; <a href="#C1003402-B1A4-11E2-ACA6-002500F14F28" class="citation">Ramel, 2012</a>; <a href="#D56D99BA-B338-11E2-AF75-002500F14F28" class="citation">Wood, et al., 2006</a>)</span></p> <ul class="aside block-grid donthyphenate one-up"> <li> <dl> <dt>Known Predators</dt> <dd> <ul> <li><a class="taxon-link rank-species" href="/accounts/Ancula_gibbosa/">Ancula gibbosa</a> (Class <a class="taxon-link rank-class" href="/accounts/Gastropoda/">Gastropoda</a>, Phylum <a class="taxon-link rank-unspecified" href="/accounts/Mollusca/">Mollusca</a>)</li> <li><a class="taxon-link rank-species" href="/accounts/Janolus_hyalinus/">Janolus hyalinus</a> (Class <a class="taxon-link rank-class" href="/accounts/Gastropoda/">Gastropoda</a>, Phylum <a class="taxon-link rank-unspecified" href="/accounts/Mollusca/">Mollusca</a>)</li> <li><a class="taxon-link rank-species" href="/accounts/Onchidoris_sparsa/">Onchidoris sparsa</a> (Class <a class="taxon-link rank-class" href="/accounts/Gastropoda/">Gastropoda</a>, Phylum <a class="taxon-link rank-unspecified" href="/accounts/Mollusca/">Mollusca</a>)</li> <li><a class="taxon-link rank-species" href="/accounts/Pomacea_canaliculata/">Pomacea canaliculata</a> (Class <a class="taxon-link rank-class" href="/accounts/Gastropoda/">Gastropoda</a>, Phylum <a class="taxon-link rank-unspecified" href="/accounts/Mollusca/">Mollusca</a>)</li> <li><a class="taxon-link rank-species" href="/accounts/Trapania_pallida/">Trapania pallida</a> (Class <a class="taxon-link rank-class" href="/accounts/Gastropoda/">Gastropoda</a>, Phylum <a class="taxon-link rank-unspecified" href="/accounts/Mollusca/">Mollusca</a>)</li> <li><a class="taxon-link rank-genus" href="/accounts/Plagiostomum/">Plagiostomum</a> sp. (Order <a class="taxon-link rank-order" href="/accounts/Prolecithophora/">Prolecithophora</a>, Phylum <a class="taxon-link rank-unspecified" href="/accounts/Platyhelminthes/">Platyhelminthes</a>)</li> </ul> </dd> </dl> </li> </ul> </section> <section class="hyphenate"> <h3 id="ecosystem_roles">Ecosystem Roles</h3> <p audience="advanced intermediate">Solitary <span class="taxon-link">entoprocts</span> are most often commensal on invertebrates such as <a class="taxon-link" href="/accounts/Porifera/">sponges</a>, <a class="taxon-link" href="/accounts/Annelida/">annelids</a>, <a class="taxon-link" href="/accounts/Sipuncula/">sipunculans</a>, <a class="taxon-link" href="/accounts/Ascidiacea/">ascidians</a>, and <a class="taxon-link" href="/accounts/Bryozoa/">ectoprocts</a>. There is a high level of host specificity. Entoproct colonies may also be found on mollusk shells. There are no parasitic forms currently recognized and there is no data currently available regarding parasites of these animals. <span class="citations"> (<a href="#2762BD11-B33C-11E2-86B1-002500F14F28" class="citation">Bleidorn, 2008</a>; <a href="#0FCD0A17-7DE7-11E2-A279-002500F14F28" class="citation">Brusca and Brusca, 2003</a>; <a href="#409A4661-B27E-11E2-B2AD-002500F14F28" class="citation">Emschermann, 1993</a>; <a href="#C0A1149C-B33C-11E2-A2C7-002500F14F28" class="citation">Kristensen, 1970</a>; <a href="#F1A1720C-B1AB-11E2-925A-002500F14F28" class="citation">Weise, 1961</a>; <a href="#EADBD5EE-7DE8-11E2-9D21-002500F14F28" class="citation">Wood, 2005</a>)</span></p> <p audience="advanced intermediate">A phoretic relationship has been identified between <a class="taxon-link rank-species" href="/accounts/Urnatella_gracilis/">Urnatella gracilis</a> and larval <span rank="Species" class="taxon-name rank-species">Cordalus cornutus</span> (commonly called hellgrammites, the larval form of Eastern dobsonflies); as a result of this relationship, entoprocts gain a means of dispersal, protection from predators, and nutrition. <span class="citations"> (<a href="#B9D67594-B337-11E2-AC82-002500F14F28" class="citation">Tracy and Hazelwood, 1983</a>)</span></p> <div class="note"> <strong>Species Used as Host</strong> <br /> <ul> <li><a class="taxon-link rank-species" href="/accounts/Clymenella_zonalis/">Clymenella zonalis</a> (Class <a class="taxon-link rank-class" href="/accounts/Polychaeta/">Polychaeta</a>, Phylum <a class="taxon-link rank-unspecified" href="/accounts/Annelida/">Annelida</a>)</li> <li><a class="taxon-link rank-family" href="/accounts/Nephtyidae/">Nephtyidae</a> sp. (Class <a class="taxon-link rank-class" href="/accounts/Polychaeta/">Polychaeta</a>, Phylum <a class="taxon-link rank-unspecified" href="/accounts/Annelida/">Annelida</a>)</li> <li><a class="taxon-link rank-family" href="/accounts/Polynoidae/">Polynoidae</a> sp. (Class <a class="taxon-link rank-class" href="/accounts/Polychaeta/">Polychaeta</a>, Phylum <a class="taxon-link rank-unspecified" href="/accounts/Annelida/">Annelida</a>)</li> <li><a class="taxon-link rank-genus" href="/accounts/Amblema/">Amblema</a> sp. (Class <a class="taxon-link rank-class" href="/accounts/Bivalvia/">Bivalvia</a>, Phylum <a class="taxon-link rank-unspecified" href="/accounts/Mollusca/">Mollusca</a>)</li> <li><a class="taxon-link rank-species" href="/accounts/Dreissena_polymorpha/">Dreissena polymorpha</a> (Class <a class="taxon-link rank-class" href="/accounts/Bivalvia/">Bivalvia</a>, Phylum <a class="taxon-link rank-unspecified" href="/accounts/Mollusca/">Mollusca</a>)</li> <li><span rank="Species" class="taxon-name rank-species">Pleurocera caniculatum</span> (Class <a class="taxon-link rank-class" href="/accounts/Gastropoda/">Gastropoda</a>, Phylum <a class="taxon-link rank-unspecified" href="/accounts/Mollusca/">Mollusca</a>)</li> <li><a class="taxon-link rank-species" href="/accounts/Ophioplinthus_gelida/">Ophioplinthus gelida</a> (Class <a class="taxon-link rank-class" href="/accounts/Ophiuroidea/">Ophiuroidea</a>, Phylum <a class="taxon-link rank-unspecified" href="/accounts/Echinodermata/">Echinodermata</a>)</li> <li><span rank="Species" class="taxon-name rank-species">Phascolion strombi</span> (Class <span class="taxon-name">Sipunculidea</span>, Phylum <a class="taxon-link rank-unspecified" href="/accounts/Sipuncula/">Sipuncula</a>)</li> </ul> </div> <div class="note"> <strong>Mutualist Species</strong> <br /> <ul> <li>eastern dobsonflies (<span rank="Species" class="taxon-name rank-species">Cordalus cornutus</span>)</li> </ul> </div> </section> <section class="hyphenate"> <h3 id="economic_importance_positive">Economic Importance for Humans: Positive</h3> <p audience="advanced intermediate">Beyond the potential for scientific research, there are no known positive effects of <span class="taxon-link">entoproct</span> species on humans. <span class="citations"> (<a href="#0FCD0A17-7DE7-11E2-A279-002500F14F28" class="citation">Brusca and Brusca, 2003</a>)</span></p> <ul class="keywords donthyphenate last"> <li class="keywords-header">Positive Impacts</li> <li> <span>research and education</span> </li> </ul> </section> <section class="hyphenate"> <h3 id="economic_importance_negative">Economic Importance for Humans: Negative</h3> <p audience="advanced intermediate"><span class="taxon-link">Entoprocts</span> may be a cause of water and vessel fouling. <span class="citations"> (<a href="#B499BC8F-B336-11E2-BF71-002500F14F28" class="citation">Canning and Carlton, 2000</a>; <a href="#83522419-B340-11E2-9E48-002500F14F28" class="citation">Soule and Soule, 1968</a>)</span></p> </section> <section class="hyphenate"> <h3 id="conservation_status">Conservation Status</h3> <p audience="advanced intermediate">There is currently no concern regarding <span class="taxon-link">entoproct</span> species becoming threatened or endangered. <span class="citations"> (<a href="#0FCD0A17-7DE7-11E2-A279-002500F14F28" class="citation">Brusca and Brusca, 2003</a>)</span></p> <ul class="aside block-grid donthyphenate one-up"> <li> <dl> <dt>IUCN Red List <a href="http://www.iucnredlist.org/">[Link]</a></dt> <dd>Not Evaluated</dd> </dl> </li> </ul> </section> <section class="hyphenate"> <h3 id="comments">Other Comments</h3> <p audience="advanced intermediate">While rare, <span class="taxon-link">entoproct</span> fossils date back to approximately 520 million years ago, during the Cambrian period. <span class="citations"> (<a href="#15A74A87-7DE9-11E2-8D5D-002500F14F28" class="citation">Zhang, et al., 2013</a>)</span></p> </section> <section class="hyphenate"> <h3 id="contributors">Contributors</h3> <p>Jeremy Wright (author), University of Michigan-Ann Arbor, Leila Siciliano Martina (editor), Animal Diversity Web Staff. </p> </section> <section class="offscreen"> <h3 id="glossary">Glossary</h3> <div id="20020916135737"> <dl> <dt>Australian</dt> <dd> <p>Living in Australia, New Zealand, Tasmania, New Guinea and associated islands.</p> <p align="center"> <img alt="World Map" src="/images/worldmap.2001.jpg" /> </p> </dd> </dl> </div> <div id="20020916134510"> <dl> <dt>Ethiopian</dt> <dd> <p>living in sub-Saharan Africa (south of 30 degrees north) and Madagascar.</p> <p align="center"> <img alt="World Map" src="/images/worldmap.2001.jpg" /> </p> </dd> </dl> </div> <div id="20020914235803"> <dl> <dt>Nearctic</dt> <dd> <p>living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.</p> <p align="center"> <img alt="World Map" src="/images/worldmap.2001.jpg" /> </p> </dd> </dl> </div> <div id="20020915000403"> <dl> <dt>Neotropical</dt> <dd> <p>living in the southern part of the New World. In other words, Central and South America.</p> <p align="center"> <img alt="World Map" src="/images/worldmap.2001.jpg" /> </p> </dd> </dl> </div> <div id="20020915001330"> <dl> <dt>Palearctic</dt> <dd> <p>living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.</p> <p align="center"> <img alt="World Map" src="/images/worldmap.2001.jpg" /> </p> </dd> </dl> </div> <div id="20020904145314"> <dl> <dt>asexual</dt> <dd> <p>reproduction that is not sexual; that is, reproduction that does not include recombining the genotypes of two parents</p> </dd> </dl> </div> <div id="20020904145431"> <dl> <dt>benthic</dt> <dd> <p>Referring to an animal that lives on or near the bottom of a body of water. Also an aquatic biome consisting of the ocean bottom below the pelagic and coastal zones. Bottom habitats in the very deepest oceans (below 9000 m) are sometimes referred to as the abyssal zone. see also oceanic vent.</p> </dd> </dl> </div> <div id="20020904145642"> <dl> <dt>bilateral symmetry</dt> <dd> <p>having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.</p> </dd> </dl> </div> <div id="20020904145435"> <dl> <dt>brackish water</dt> <dd> <p>areas with salty water, usually in coastal marshes and estuaries.</p> </dd> </dl> </div> <div id="20020904145825"> <dl> <dt>coastal</dt> <dd> <p>the nearshore aquatic habitats near a coast, or shoreline.</p> </dd> </dl> </div> <div id="20020904145440"> <dl> <dt>colonial</dt> <dd> <p>used loosely to describe any group of organisms living together or in close proximity to each other - for example nesting shorebirds that live in large colonies. More specifically refers to a group of organisms in which members act as specialized subunits (a continuous, modular society) - as in clonal organisms.</p> </dd> </dl> </div> <div id="20020904145620"> <dl> <dt>cosmopolitan</dt> <dd> <p>having a worldwide distribution. Found on all continents (except maybe Antarctica) and in all biogeographic provinces; or in all the major oceans (Atlantic, Indian, and Pacific.</p> </dd> </dl> </div> <div id="20020904145752"> <dl> <dt>crepuscular</dt> <dd> <p>active at dawn and dusk</p> </dd> </dl> </div> <div id="20020904145414"> <dl> <dt>diurnal</dt> <dd> <ol class="arabic simple"> <li>active during the day, 2. lasting for one day.</li> </ol> </dd> </dl> </div> <div id="20020904145546"> <dl> <dt>fertilization</dt> <dd> <p>union of egg and spermatozoan</p> </dd> </dl> </div> <div id="20020904145747"> <dl> <dt>freshwater</dt> <dd> <p>mainly lives in water that is not salty.</p> </dd> </dl> </div> <div id="20020916125902"> <dl> <dt>heterothermic</dt> <dd> <p>having a body temperature that fluctuates with that of the immediate environment; having no mechanism or a poorly developed mechanism for regulating internal body temperature.</p> </dd> </dl> </div> <div id="200304045530"> <dl> <dt>holarctic</dt> <dd> <p>a distribution that more or less circles the Arctic, so occurring in both the Nearctic and Palearctic biogeographic regions.</p> <p align="center"> <img alt="World Map" src="/images/worldmap.2001.jpg" /> </p> <p>Found in northern North America and northern Europe or Asia.</p> </dd> </dl> </div> <div id="20020904145427"> <dl> <dt>internal fertilization</dt> <dd> <p>fertilization takes place within the female's body</p> </dd> </dl> </div> <div id="20020904145633"> <dl> <dt>intertidal or littoral</dt> <dd> <p>the area of shoreline influenced mainly by the tides, between the highest and lowest reaches of the tide. An aquatic habitat.</p> </dd> </dl> </div> <div id="20020904145364"> <dl> <dt>metamorphosis</dt> <dd> <p>A large change in the shape or structure of an animal that happens as the animal grows. In insects, "incomplete metamorphosis" is when young animals are similar to adults and change gradually into the adult form, and "complete metamorphosis" is when there is a profound change between larval and adult forms. Butterflies have complete metamorphosis, grasshoppers have incomplete metamorphosis.</p> </dd> </dl> </div> <div id="20020904145472"> <dl> <dt>motile</dt> <dd> <p>having the capacity to move from one place to another.</p> </dd> </dl> </div> <div id="20020904145365"> <dl> <dt>native range</dt> <dd> <p>the area in which the animal is naturally found, the region in which it is endemic.</p> </dd> </dl> </div> <div id="20020904145503"> <dl> <dt>nocturnal</dt> <dd> <p>active during the night</p> </dd> </dl> </div> <div id="20020915001348"> <dl> <dt>oriental</dt> <dd> <p>found in the oriental region of the world. In other words, India and southeast Asia.</p> <p align="center"> <img alt="World Map" src="/images/worldmap.2001.jpg" /> </p> </dd> </dl> </div> <div id="20020904145572"> <dl> <dt>oviparous</dt> <dd> <p>reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.</p> </dd> </dl> </div> <div id="20020904145368"> <dl> <dt>planktivore</dt> <dd> <p>an animal that mainly eats plankton</p> </dd> </dl> </div> <div id="20020904145483"> <dl> <dt>polygynandrous</dt> <dd> <p>the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.</p> </dd> </dl> </div> <div id="20020904145373"> <dl> <dt>protandrous</dt> <dd> <p>condition of hermaphroditic animals (and plants) in which the male organs and their products appear before the female organs and their products</p> </dd> </dl> </div> <div id="20020904145675"> <dl> <dt>reef</dt> <dd> <p>structure produced by the calcium carbonate skeletons of coral polyps (Class Anthozoa). Coral reefs are found in warm, shallow oceans with low nutrient availability. They form the basis for rich communities of other invertebrates, plants, fish, and protists. The polyps live only on the reef surface. Because they depend on symbiotic photosynthetic algae, zooxanthellae, they cannot live where light does not penetrate.</p> </dd> </dl> </div> <div id="20020904145582"> <dl> <dt>saltwater or marine</dt> <dd> <p>mainly lives in oceans, seas, or other bodies of salt water.</p> </dd> </dl> </div> <div id="20020904145584"> <dl> <dt>seasonal breeding</dt> <dd> <p>breeding is confined to a particular season</p> </dd> </dl> </div> <div id="20020904145585"> <dl> <dt>sedentary</dt> <dd> <p>remains in the same area</p> </dd> </dl> </div> <div id="20020904145586"> <dl> <dt>sessile</dt> <dd> <p>non-motile; permanently attached at the base.</p> <p>Attached to substratum and moving little or not at all. Synapomorphy of the Anthozoa</p> </dd> </dl> </div> <div id="20020904145786"> <dl> <dt>sexual</dt> <dd> <p>reproduction that includes combining the genetic contribution of two individuals, a male and a female</p> </dd> </dl> </div> <div id="20020904145381"> <dl> <dt>solitary</dt> <dd> <p>lives alone</p> </dd> </dl> </div> <div id="20020904145500"> <dl> <dt>tactile</dt> <dd> <p>uses touch to communicate</p> </dd> </dl> </div> <div id="20020904145595"> <dl> <dt>temperate</dt> <dd> <p>that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).</p> </dd> </dl> </div> <div id="20020904145598"> <dl> <dt>tropical</dt> <dd> <p>the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.</p> </dd> </dl> </div> <div id="20020904145698"> <dl> <dt>year-round breeding</dt> <dd> <p>breeding takes place throughout the year</p> </dd> </dl> </div> </section> <section class="hyphenate"> <h3 id="references">References</h3> <p id="874637AB-7DE7-11E2-B377-002500F14F28">Appeltans, W., P. 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Histoire naturelle du genre Pedicellina. <span style="font-style: italic">Nouveaux M茅moires de l'Acad茅mie Royale des Sciences et des Belles-Lettres de Bruxelles</span>, 19: 1-31. </p> </section> </div> </div> <div class="span2_5 right sidebar"> <div class="well" id="pocket-guide-link"> <a href="https://itunes.apple.com/us/app/adw-pocket-guide/id916179421?mt=8" target="_blank"><i class="icon-download icon-white" style="vertical-align: bottom"></i> ADW Pocket Guides on the iOS App Store! </a> <p>The Animal Diversity Web team is excited to announce ADW Pocket Guides!</p> <p> <a href="/news/5009059054038197651/">Read more...</a> </p> </div> <h2 class="offscreen">Search</h2> <div id="site-search"> <form action="/search" method="GET"> <label for="q" class="offscreen">Enter search text</label> <input value="" id="q" name="q" class="span12" size="15" type="text" placeholder="Search ADW" /> <button class="btn"> <i class="icon-zoom-in"></i> <span class="offscreen">Search</span> </button> <p style="max-height: 35px"> <label for="feature-list" class="offscreen">Search in feature</label> <select size="1" name="feature" class="feature-list input-medium" id="feature-list"> <option value="INFORMATION">Taxon Information</option> <option value="COLLECTIONS">Contributor Galleries</option> <option value="TOPICS">Topics</option> <option value="CLASSIFICATION">Classification</option> </select> </p> </form> <ul class="unstyled"> <li> <a href="https://animaldiversity.ummz.umich.edu/quaardvark/">Explore Data @ Quaardvark</a> </li> <li> <a href="/search_guide/">Search Guide</a> </li> </ul> </div> <h2 class="offscreen">Navigation Links</h2> <dl class="tabbed features"> <dd class="feature-information active"> <a name="feature-information" href="/accounts/Entoprocta/" class="active" id="feature-information"> Information </a> </dd> <dd class="feature-pictures"> <a name="feature-pictures" href="/accounts/Entoprocta/pictures/" id="feature-pictures"> Pictures </a> </dd> <dd class="feature-specimens"> <a name="feature-specimens" href="/accounts/Entoprocta/specimens/" id="feature-specimens"> Specimens </a> </dd> <dd class="feature-classification"> <a name="feature-classification" href="/accounts/Entoprocta/classification/#Entoprocta" id="feature-classification"> Classification </a> </dd> </dl> <div class="classification well"> <h3>Classification</h3> <ul class="unstyled"> <li> <span class="rank">Kingdom</span> <a href="/accounts/Animalia/" class="taxon-name rank-kingdom">Animalia</a> <span class="vernacular-name">animals</span> <div class="features"> <a href="/accounts/Animalia/" class="feature feature-information" rel="tooltip" data-original-title="Animalia: information (1)" data-delay="250" data-total="4749">Animalia: information (1)</a> <a href="/accounts/Animalia/pictures/" class="feature feature-pictures" rel="tooltip" data-original-title="Animalia: pictures (22861)" data-delay="250">Animalia: pictures (22861)</a> <a href="/accounts/Animalia/specimens/" class="feature feature-specimens" rel="tooltip" data-original-title="Animalia: specimens (7109)" data-delay="250">Animalia: specimens (7109)</a> <a href="/accounts/Animalia/sounds/" class="feature feature-sounds" rel="tooltip" data-original-title="Animalia: sounds (722)" data-delay="250">Animalia: sounds (722)</a> <a href="/accounts/Animalia/maps/" class="feature feature-maps" rel="tooltip" data-original-title="Animalia: maps (42)" data-delay="250">Animalia: maps (42)</a> </div> </li> <li class="active"> <span class="rank">Unspecified</span> <a href="/accounts/Entoprocta/" class="taxon-name rank-unspecified">Entoprocta</a> <span class="vernacular-name">hairy back worms</span> <div class="features"> <a href="/accounts/Entoprocta/" class="feature feature-information" rel="tooltip" data-original-title="Entoprocta: information (1)" data-delay="250" data-total="1">Entoprocta: information (1)</a> <a href="/accounts/Entoprocta/pictures/" class="feature feature-pictures" rel="tooltip" data-original-title="Entoprocta: pictures (4)" data-delay="250">Entoprocta: pictures (4)</a> <a href="/accounts/Entoprocta/specimens/" class="feature feature-specimens" rel="tooltip" data-original-title="Entoprocta: specimens (2)" data-delay="250">Entoprocta: specimens (2)</a> <span class="feature-off feature-sounds"></span> <span class="feature-off feature-maps"></span> </div> </li> </ul> </div> </div> </div> </div> </div> <div id="footer" aria-role="contentinfo"> <div class="inner-footer"> <div class="page-citation"> <p class="content">To cite this page: Wright, J. 2014. 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