scyphozoans
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The Scyphozoa are an exclusively marine class of the phylum
Cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter. Their distinguishing feature is cnidocytes, specialized cells that ...
, referred to as the true
jellyfish Jellyfish and sea jellies are the informal common names given to the medusa-phase of certain gelatinous members of the subphylum Medusozoa, a major part of the phylum Cnidaria. Jellyfish are mainly free-swimming marine animals with umbrell ...
(or "true jellies"). The class name Scyphozoa comes from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
word ''
skyphos A ''skyphos'' ( grc, σκύφος; plural ''skyphoi'') is a two-handled deep wine-cup on a low flanged base or none. The handles may be horizontal ear-shaped thumbholds that project from the rim (in both Corinthian and Athenian shapes), or they ma ...
'' (), denoting a kind of drinking cup and alluding to the cup shape of the organism. Scyphozoans have existed from the earliest Cambrian to the present.


Biology

Most species of Scyphozoa have two life-history phases, including the planktonic
medusa In Greek mythology, Medusa (; Ancient Greek: Μέδουσα "guardian, protectress"), also called Gorgo, was one of the three monstrous Gorgons, generally described as winged human females with living venomous snakes in place of hair. Those ...
or polyp form, which is most evident in the warm summer months, and an inconspicuous, but longer-lived, bottom-dwelling polyp, which seasonally gives rise to new medusae. Most of the large, often colorful, and conspicuous jellyfish found in coastal waters throughout the world are Scyphozoa. They typically range from in diameter, but the largest species, ''
Cyanea capillata The lion's mane jellyfish (''Cyanea capillata''), also known as the giant jellyfish, arctic red jellyfish, or the hair jelly, is one of the largest known species of jellyfish. Its range is confined to cold, boreal waters of the Arctic, norther ...
'' can reach across. Scyphomedusae are found throughout the world's oceans, from the surface to great depths; no Scyphozoa occur in freshwater (or on land). As medusae, they eat a variety of crustaceans and fish, which they capture using stinging cells called
nematocysts A cnidocyte (also known as a cnidoblast or nematocyte) is an explosive cell containing one large secretory organelle called a cnidocyst (also known as a cnida () or nematocyst) that can deliver a sting to other organisms. The presence of this c ...
. The nematocysts are located throughout the tentacles that radiate downward from the edge of the umbrella dome, and also cover the four or eight oral arms that hang down from the central mouth. Some species, however, are instead filter feeders, using their tentacles to strain
plankton Plankton are the diverse collection of organisms found in water (or air) that are unable to propel themselves against a current (or wind). The individual organisms constituting plankton are called plankters. In the ocean, they provide a crucia ...
from the water.


Anatomy

Scyphozoans usually display a four-part symmetry and have an internal gelatinous material called
mesoglea Mesoglea refers to the extracellular matrix found in cnidarians like coral or jellyfish that functions as a hydrostatic skeleton. It is related to but distinct from mesohyl, which generally refers to extracellular material found in sponges. Desc ...
, which provides the same structural integrity as a skeleton. The mesoglea includes mobile amoeboid cells originating from the epidermis. Scyphozoans have no durable hard parts, including no head, no skeleton, and no specialized organs for respiration or excretion. Marine jellyfish can consist of as much as 98% water, so are rarely found in fossil form. Unlike the
hydrozoan Hydrozoa (hydrozoans; ) are a taxonomic class of individually very small, predatory animals, some solitary and some colonial, most of which inhabit saline water. The colonies of the colonial species can be large, and in some cases the specialize ...
jellyfish, Hydromedusae, Scyphomedusae lack a vellum, which is a circular membrane beneath the umbrella that helps propel the (usually smaller) Hydromedusae through the water. However, a ring of muscle fibres is present within the mesoglea around the rim of the dome, and the jellyfish swims by alternately contracting and relaxing these muscles. The periodic contracting and relaxing propels the jellyfish through the water, allowing it to escape predation or catch its prey. The mouth opens into a central stomach, from which four interconnected
diverticula In medicine or biology, a diverticulum is an outpouching of a hollow (or a fluid-filled) structure in the body. Depending upon which layers of the structure are involved, diverticula are described as being either true or false. In medicine, t ...
radiate outwards. In many species, this is further elaborated by a system of radial canals, with or without an additional ring canal towards the edge of the dome. Some genera, such as ''
Cassiopea ''Cassiopea'' (upside-down jellyfish) is a genus of true jellyfish and the only members of the family Cassiopeidae. They are found in warmer coastal regions around the world, including shallow mangrove swamps, mudflats, canals, and turtle gra ...
'', even have additional, smaller mouths in the oral arms. The lining of the digestive system includes further stinging nematocysts, along with cells that secrete
digestive enzyme Digestive enzymes are a group of enzymes that break down polymeric macromolecules into their smaller building blocks, in order to facilitate their absorption into the cells of the body. Digestive enzymes are found in the digestive tracts of anima ...
s. The nervous system usually consists of a distributed net of cells, although some species possess more organised nerve rings. In species lacking nerve rings, the nerve cells are instead concentrated into small structures called ''
rhopalia Rhopalia (singular: rhopalium) are small sensory structures of certain Scyphozoan (true jellyfish) and Cubozoan (box jellyfish) species. Description The structures typically occur in multiples of four, are bell shaped and face outward from inva ...
''. There are between four and sixteen of these small lobes arranged around the rim of the umbrella, where they coordinate the muscular action allowing the animal to move. Each rhopalium is typically associated with a pair of sensory pits, a
statocyst The statocyst is a balance sensory receptor present in some aquatic invertebrates, including bivalves, cnidarians, ctenophorans, echinoderms, cephalopods, and crustaceans. A similar structure is also found in '' Xenoturbella''. The statocyst c ...
, and sometimes a pigment-cup
ocellus A simple eye (sometimes called a pigment pit) refers to a form of eye or an optical arrangement composed of a single lens and without an elaborate retina such as occurs in most vertebrates. In this sense "simple eye" is distinct from a multi-l ...
.


Reproduction

Most species appear to be
gonochorists In biology, gonochorism is a sexual system where there are only two sexes and each individual organism is either male or female. The term gonochorism is usually applied in animal species, the vast majority of which are gonochoric. Gonochorism c ...
, with separate male and female individuals. The gonads are located in the stomach lining, and the mature
gamete A gamete (; , ultimately ) is a haploid cell that fuses with another haploid cell during fertilization in organisms that reproduce sexually. Gametes are an organism's reproductive cells, also referred to as sex cells. In species that produce ...
s are expelled through the mouth. After fertilization, some species brood their young in pouches on the oral arms, but they are more commonly planktonic.


Growth and development

The fertilized egg produces a
planula A planula is the free-swimming, flattened, ciliated, bilaterally symmetric larval form of various cnidarian species and also in some species of Ctenophores. Some groups of Nemerteans also produce larvae that are very similar to the planula, which ...
r larva which, in most species, quickly attaches itself to the sea bottom. The larva develops into the hydroid stage of the lifecycle, a tiny sessile polyp called a scyphistoma. The scyphistoma reproduces asexually, producing similar polyps by budding, and then either transforming into a medusa, or budding several medusae off from its upper surface via a process called strobilation. The medusae are initially microscopic and may take years to reach sexual maturity.


Commercial importance

Scyphozoa include the moon jelly ''
Aurelia aurita ''Aurelia aurita'' (also called the common jellyfish, moon jellyfish, moon jelly or saucer jelly) is a species of the genus '' Aurelia''. All species in the genus are very similar, and it is difficult to identify ''Aurelia'' medusae without genet ...
'', in the order Semaeostomeae, and the enormous '' Nemopilema nomurai'', in the order Rhizostomeae, found between Japan and China and which in some years causes major fisheries disruptions. The jellyfish fished commercially for food are Scyphomedusae in the order Rhizostomeae. Most rhizostome jellyfish live in warm water.


Taxonomy

Although the Scyphozoa were formerly considered to include the animals now referred to as the classes
Cubozoa Box jellyfish (class Cubozoa) are cnidarian invertebrates distinguished by their box-like (i.e. cube-shaped) body. Some species of box jellyfish produce potent venom delivered by contact with their tentacles. Stings from some species, including ...
and
Staurozoa Staurozoa is a class of Medusozoa, jellyfishes and hydrozoans. It has one extant order: Stauromedusae (stalked jellyfishes) with a total of 50 known species. A fossil group called Conulariida has been proposed as a second order, although this is ...
, they now include just three extant orders (two of which are in Discomedusae, a subclass of Scyphozoa).Daly, Brugler, Cartwright, Collins, Dawson, Fautin, France, McFadden, Opresko, Rodriguez, Romano & Stake (2007).
The phylum Cnidaria: A review of phylogenetic patterns and diversity 300 years after Linnaeus.
' Zootaxa 1668: 127–182
About 200 extant species are recognized at present, but the true diversity is likely to be at least 400 species. Class Scyphozoa *Subclass Coronamedusae :* Order
Coronatae Crown jellyfishes are the six families of true jellyfish that belong to the order Coronatae.Daly, Brugler, Cartwright, Collins, Dawson, Fautin, France, McFadden, Opresko, Rodriguez, Romano & Stake (2007). The phylum Cnidaria: A review of phylog ...
:::* Family Atollidae :::* Family Atorellidae :::* Family Linuchidae :::* Family Nausithoidae :::* Family Paraphyllinidae :::* Family Periphyllidae * Subclass
Discomedusae Discomedusae is a subclass of jellyfish in the class Scyphozoa. It is the sister taxon of Coronamedusae. Discomedusae contains about 155 named species and there are likely to be many more as yet undescribed. Jellyfish in this subclass are much ...
:* Order Rhizostomeae ::* Suborder Daktyliophorae :::* Family Catostylidae :::* Family Lobonematidae :::* Family Lychnorhizidae :::* Family
Rhizostomatidae Rhizostomatidae is a family of cnidarians in the class Scyphozoa. Genera and species According to the World Register of Marine Species, 11 extant species are in four extant genera within this family: *Genus ''Eupilema'' **'' Eupilema inexpect ...
:::* Family Stomolophidae ::* Suborder Kolpophorae :::* Family Cassiopeidae :::* Family
Cepheidae Cepheidae may refer to: * Cepheidae (jellyfish) Cepheidae is a family of jellyfish Jellyfish and sea jellies are the informal common names given to the medusa-phase of certain gelatinous members of the subphylum Medusozoa, a major part o ...
:::* Family Mastigiidae :::* Family Thysanostomatidae :::* Family Versurigidae :* Order
Semaeostomeae Semaeostomeae (literally "flag mouths") is an order of large jellyfish characterized by four long, frilly oral arms flanking their quadrate mouths. The umbrella is domed with scalloped margins, and the gastrovascular system consists of four unbra ...
:::* Family
Cyaneidae The Cyaneidae are a family of true jellyfish. About 20 species are in this family, including the well-known lion's-mane jellyfish. Species The following species are recognized within the family Cyaneidae. Formerly, this family also included th ...
:::* Family
Drymonematidae ''Drymonema'' is a genus of true jellyfish, placed in its own family, the Drymonematidae. There are three species: '' Drymonema dalmatinum'', '' Drymonema gorgo'', and '' Drymonema larsoni'', which are found in the Gulf of Mexico, Atlantic Ocean ...
Bayha, K. M., and M. N. Dawson (2010). ''New family of allomorphic jellyfishes, Drymonematidae (Scyphozoa, Discomedusae), emphasizes evolution in the functional morphology and trophic ecology of gelatinous zooplankton.''
The Biological Bulletin ''The Biological Bulletin'' is a peer-reviewed scientific journal covering the field of biology. The journal was established in 1897 as the ''Zoological Bulletin'' by Charles Otis Whitman and William Morton Wheeler. In 1899 the title was changed to ...
219(3): 249–267
:::* Family Pelagiidae :::* Family Phacellophoridae :::* Family Ulmaridae


References


External links


The Classification and Distribution of the Class Scyphozoa
* {{Authority control Medusozoa Ediacaran first appearances