cnidarian

/nuy dair"ee euhn/, n.

1. any invertebrate animal, as a hydra, jellyfish, sea anemone, or coral, considered as belonging to the phylum Cnidaria, characterized by the specialized stinging structures in the tentacles surrounding the mouth; a coelenterate.

adj.

2. belonging or pertaining to the Cnidaria.

[CNIDARI(A) + -AN]

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or coelenterate

Any of about 9,000 species of mostly marine aquatic invertebrates, constituting the phylum Cnidaria (or Coelenterata), that are unique in possessing specialized stinging cells (cnidocytes) borne on the tentacles.

Cnidocytes contain fluid-filled capsules (nematocysts) with a harpoonlike coiled thread used for stinging, paralyzing, and capturing prey. Cnidarians have no well-defined separate respiratory, circulatory, or excretory organs; their tissues, composed of two cell layers, surround a cavity known as a coelenteron (gastrovascular cavity), which is the basic internal organ. Tentacles surrounding the mouth are used to capture and ingest food. Cnidarians are carnivorous, feeding mostly on zooplankton but also on small crustaceans, fish eggs, worms, smaller cnidarians, and even small fish. Cnidarians range in size from nearly microscopic to more than 100 ft (30 m) long and more than a ton (910 kg) in weight. There are two basic body forms: the polyp (e.g., coral) and the medusa (e.g., jellyfish). See also hydra; Portuguese man-of-war; sea anemone.

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▪ invertebrate

Introduction

also called  coelenterate 

 any member of the phylum Cnidaria (Coelenterata), a group made up of more than 9,000 living species. Mostly marine animals, the cnidarians include the corals, hydras, jellyfish, Portuguese men-of-war, sea anemones, sea pens, sea whips, and sea fans.

      The phylum Cnidaria is made up of four classes: Hydrozoa (hydrozoans); Scyphozoa (scyphozoans); Anthozoa (anthozoans); and Cubozoa (cubozoans). All cnidarians share several attributes, supporting the theory that they had a single origin. Variety and symmetry of body forms, varied coloration, and the sometimes complex life histories of cnidarians fascinate layperson and scientist alike. Inhabiting all marine and some freshwater environments, these animals are most abundant and diverse in tropical waters. Their calcareous skeletons form the frameworks of the reefs (coral reef) and atolls in most tropical seas, including the Great Barrier Reef that extends more than 2,000 kilometres along the northeastern coast of Australia.

      Only cnidarians manufacture microscopic intracellular stinging capsules, known as nematocysts (nematocyst) or cnidae, which give the phylum its name. The alternative name, coelenterate, refers to their simple organization around a central body cavity (the coelenteron). As first defined, coelenterates included not only the animals now designated cnidarians but also sponges (phylum Porifera) and comb jellies (phylum Ctenophora). In contemporary usage, “coelenterate” generally refers only to cnidarians, but the latter term is used in order to avoid ambiguity.

General features

Size range and diversity of structure

      Cnidarians are radially symmetrical (i.e., similar parts are arranged symmetrically around a central axis). They lack cephalization (concentration of sensory organs in a head), their bodies have two cell layers rather than the three of so-called higher animals, and the saclike coelenteron has one opening (the mouth). They are the most primitive of animals whose cells are organized into distinct tissues, but they lack organs. Cnidarians have two body forms— polyp and medusa—which often occur within the life cycle of a single cnidarian.

      The body of a medusa, commonly called a jellyfish, usually has the shape of a bell or an umbrella, with tentacles hanging downward at the margin. The tubelike manubrium hangs from the centre of the bell, connecting the mouth at the lower end of the manubrium to the coelenteron within the bell. Most medusae are slow-swimming, planktonic animals. In contrast, the mouth and surrounding tentacles of polyps face upward, and the cylindrical body is generally attached by its opposite end to a firm substratum. The mouth is at the end of a manubrium in many hydrozoan polyps. Anthozoan polyps have an internal pharynx, or stomodaeum, connecting the mouth to the coelenteron.

      Most species of cubozoans, hydrozoans, and scyphozoans pass through the medusoid and polypoid body forms, with medusae giving rise sexually to larvae that metamorphose into polyps, while polyps produce medusae asexually. Thus, the polyp is essentially a juvenile form, while the medusa is the adult form. In contrast, anthozoans are polypoid cnidarians and do not have a medusa stage. Commonly polyps, and in some species medusae too, can produce more of their own kind asexually.

      One body form may be more conspicuous than the other. For example, scyphozoans are commonly known as true jellyfishes, for the medusa form is larger and better known than the polyp form. In hydrozoans, the polyp phase is more conspicuous than the medusa phase in groups such as hydroids (hydroid) and hydrocorals. Hydromedusae are smaller and more delicate than scyphomedusae or cubomedusae; they may be completely absent from the life cycle of some hydrozoan species. Some other species produce medusae, but the medusae never separate themselves from the polyps. Cubozoans have medusae commonly known as box jellyfish, from their shape. Some of these are responsible for human fatalities, mostly in tropical Australia and Southeast Asia, and include the so-called sea wasps. The polyp is tiny and inconspicuous.

      Many cnidarian polyps are individually no more than a millimetre or so across. Polyps of most hydroids, hydrocorals, and soft and hard corals (coral), however, proliferate asexually into colonies, which can attain much greater size and longevity than their component polyps. Certain tropical sea anemones (sea anemone) (class Anthozoa) may be a metre in diameter, and some temperate ones are nearly that tall. Anthozoans are long-lived, both individually and as colonies; some sea anemones are centuries old. All medusae and sea anemones occur only as solitary individuals. Scyphomedusae can weigh more than a ton, whereas hydromedusae are, at most, a few centimetres across. Tentacles of medusae, however, may be numerous and extensible, which allows the animals to influence a considerably greater range than their body size might suggest. Large populations of hydroids can build up on docks, boats, and rocks. Similarly, some medusae attain remarkable densities—up to thousands per litre of water—but only for relatively brief periods.

Distribution and abundance

      Many of the world's benthic (bottom-dwelling) ecosystems are dominated by anthozoans. Although soft and hard corals coexist in virtually all tropical areas appropriate for either, coral reefs of the tropical Indo-Pacific are built mainly by members of the anthozoan order Scleractinia (hard corals); whereas on coral reefs of the Caribbean members of the anthozoan subclass Alcyonaria (soft corals) are much more prominent. Aside from being the most numerous and covering the greatest area of any animals on the reef, the corals structure their environment, even after death. Soft corals contribute greatly to reef construction by the cementing action of the skeletal debris (spicules), filling in spaces between hard coral skeletons.

      Soft-bodied anthozoans are similarly dominant in other seas. Temperate rocky intertidal zones in many parts of the world are carpeted with sea anemones. They sequester the space that is therefore made unavailable to other organisms, thus having a profound impact on community structure. The curious hemispherical anemone Liponema is the most abundant benthic invertebrate in the Gulf of Alaska, in terms of numbers and biomass. Parts of the Antarctic seabed are covered by anemones, and they occur near the deep-sea hot vents.

Importance

      Prominent among organisms that foul water-borne vessels are sedentary cnidarians, especially hydroids. The muscles that make scyphomedusae strong swimmers are dried for human consumption in Asia. Sea anemones are eaten in some areas of Asia and North America.

      Throughout the tropics where reefs are accessible, coral skeletons are used as building material, either in blocks or slaked to create cement. Another use for cnidarian skeletons is in jewelry. The pink colour known as “coral” is the hue of the skeleton of a species of hydrocoral. Other hydrocorals have purplish skeletons. Skeletons vary in hue, and those considered most desirable command a high price. The core of some sea fans (sea fan), sea whips, and black corals are cut or bent into beads, bracelets, and cameos.

      All cnidarians have the potential to affect human physiology owing to the toxicity of their nematocysts. Most are not harmful to humans, but some can impart a painful sting—such as Physalia, the Portuguese man-of-war, and sea anemones of the genus Actinodendron. These, and even normally innocuous species, can be deadly in a massive dose or to a sensitive person, but the only cnidarians commonly fatal to humans are the cubomedusae, or box jellyfish. anaphylaxis (hypersensitivity due to prior exposure and subsequent sensitization) was discovered with experiments on Physalia toxin. Extracts of many cnidarians, mostly anthozoans, have heart-stimulant, antitumour, and anti-inflammatory properties.

Natural history

Reproduction and life cycles

      All cnidarian species are capable of sexual reproduction, which occurs in only one phase of the life cycle, usually the medusa. Many cnidarians also reproduce asexually, which may occur in both phases. In asexual reproduction, new individuals arise from bits of tissue that are budded off from a parent, or by a parent dividing lengthwise or crosswise into two smaller individuals. Polyps that remain physically attached to one another or embedded in a common mass of tissue constitute a colony. In some colonies, polyps share a common coelenteron through which food captured by any member is distributed to others. Hydrozoan polyp colonies, called hydroids, are prostrate, bushy, or feathery in form. Examples of other colonies are anthozoan soft corals and most reef-forming hard corals. Polyps that are produced asexually and then physically separate are called clones, or ramets. In this way, a single genotype can be represented by many separate “individuals.”

      Although genetically identical, colony members of many hydrozoans and some anthozoans are polymorphic (polymorphism), differing in morphology (form and structure) and/or physiology. Each zooid within the colony has a specific function and varies somewhat in form. For example, gastrozooids bear tentacles and are specialized for feeding. Some colonies possess dactylozooids, tentacleless polyps heavily armed with nematocysts that seem primarily concerned with defense. Gonozooids develop reproductive structures called gonophores. Members of the order Siphonophora, free-floating colonial hydrozoans, display an even greater variety of polymorphs. These include gas-filled floats called pneumatophores, pulsating, locomotory structures called nectophores, and flattened, protective individuals called bracts or phyllozooids.

      Although the medusa stage is absent in anthozoans, polyps produce additional polyps sexually and, in some species, asexually as well. Hydromedusae are budded from polyps that, in some colonial species, are specialized for this function; each polyp produces numerous medusae. The major distinguishing feature of the cubozoans is that each polyp transforms entirely into a medusa. Before this metamorphosis occurs, however, each cubozoan polyp may divide asexually to produce numerous genetically identical polyps, and each of these subsequent polyps can then produce a medusa. In most scyphozoans, a scyphistoma (scyphopolyp) produces immature medusae (ephyrae) by asexual fission at its oral end. This process, called strobilation, results in eight-armed, free-swimming ephyrae.

      Gametes (gamete) differentiate in parts of the body referred to as gonads, despite the fact that cnidarians cannot be said to have true ovaries and testes because they lack organs. In anthozoans, cubozoans, and scyphozoans, gametes develop in the endoderm, whereas in hydrozoans they ripen in the ectoderm, although they do not necessarily originate there. Sexes are commonly separate, but hermaphroditism is known. Some hermaphroditic species are capable of self-fertilization. Gametes are generally shed into the sea, where the eggs are fertilized. Cleavage produces a ciliated ball of cells that elongates and develops a tuft of cilia at one end to become a planula larva, which may be free-swimming and planktonic, or crawling and benthic. Its ciliated tuft, which may have sensory abilities, is directed forward in locomotion. After a motile period, the planula attaches by its forward end to a solid object and develops tentacles around its posterior end, thereby transforming into a polyp. In some anthozoans and a few scyphomedusae, eggs are fertilized without being released. Embryonic development passes either partly or entirely within the mother's coelenteron or, as in the case of some anemones and some members of the anthozoan subclass Alcyonaria (octocorals), attached to the outside of her body. In some species of hydroids that lack a free medusa stage, eggs are fertilized and the embryo develops in specialized zooids that are essentially attached medusae. Such brooding species may release offspring as very advanced larvae or as miniature adults, so that a planktonic stage is absent from the life cycle.

Ecology and habitats

      Most polyps require solid substrata for attachment, although a few burrow into soft sediments, extending only their tentacular crowns above the surface. Polyps are abundant in shallow waters, but sea anemones can also occur in the deepest parts of the oceans. Medusae maintain a favoured depth in the water and are carried about by currents. Most hydromedusae and scyphomedusae live in surface waters, generally in bays and along coasts, but certain species are abundant in the open ocean.

      The nematocysts of cnidarians restrict potential predators to a limited array of specialists. Nematocysts also allow cnidarians to prey on a variety of would-be competitors for space and food. Most species that are capable of monopolizing space reproduce predominantly asexually. A coral, for example, can cover an area rapidly and commonly has the ability to overgrow other organisms, including corals of other species. Clone-forming sea anemones of several species actively compete for space by killing others, primarily those of their own species. When members of one clone encounter those of another, the two combatants inflate and slap one another with nematocyst-studded fighting structures (acrorhagi) located below the tentacles. Attacks may result in the death of one of the anemones, or both may retreat. Tentacle touching is involved in the recognition of non-clonemates, which presumably is chemically mediated.

      Cnidarians are not immune from predation. Hydroids are victimized by nudibranchs (nudibranch) that bite through the chitinous skeleton or crawl into its openings. The crown of thorns starfish, Acanthaster planci (crown-of-thorns starfish), extrudes its stomach over a coral colony, releases digestive enzymes, and then absorbs the liquified tissue. Butterfly and parrot fishes eat corals, being insensitive to the effects of nematocysts, which is also true of marine turtles that feed on pelagic scyphomedusae. To combat these types of threats, many cnidarian species have evolved unique chemical defenses that effectively deter predation by fish and others.

Locomotion

      Medusae swim by jet propulsion (see below Tissues and muscles (cnidarian)); however, most do so weakly and are carried passively by currents over long distances. Polyps are generally sedentary. Pennatulacean (sea pen) colonies move slowly across soft substrata by action of their inflatable peduncle (a stalk that attaches to the strata in the lower end and to the polyp body on the higher end). Sea anemones that are attached to firm substrata can creep slowly on their pedal disks or detach altogether, often in response to unfavourable physical conditions or to attack by predators. When provoked by certain starfish and nudibranchs, individuals of a few anemone species swim by paddling their tentacles or flexing their columns.

Food and feeding

      All cnidarians are carnivores. Most use their cnidae and associated toxin to capture food, although none is known actually to pursue prey. Sessile polyps depend for food on organisms that come into contact with their tentacles. Some, such as colonial corals with minute polyps, feed on particulate material gathered in mucus impelled to the mouth by cilia (cilium) (microscopic hairlike projections of cells capable of beating or waving). A hydromedusa alternately swims upward and sinks: on the upward course, its trailing tentacles are not apt to encounter food organisms, but in sinking, the extended tentacles “fish” through the water, capturing food. Once a food item is captured, tentacles move it to the mouth, either by bending in that direction or by passing it to tentacles nearer the mouth. The mouth opens, the lips grasp the food, and muscular actions complete swallowing.

      The edges of the mouths of some scyphomedusae are elaborated into mouth arms that trail behind the slowly swimming jellyfish, presenting huge surfaces for food gathering. The mouth of a scyphomedusa of the order Rhizostomae is subdivided into thousands of minute pores that lead by tubes to the coelenteron. Each pore is associated with an external ciliated gutter that collects minute organisms and detrital material as the medusa rests mouth-upward on the sea bottom.

      Pink, orange, red, and brown cnidarians are commonly pigmented by carotenoids derived from crustaceans in their diet. Endodermal cells of some corals, medusae, hydras, and sea anemones contain single-celled golden-brown algae (dinoflagellates), called zooxanthellae (zooxanthella), or green algae, called zoochlorellae (zoochlorella). The carnivorous cnidarians cannot digest these algae but do derive a variety of nutrients from them, including glucose and oxygen. Carbon dioxide produced in respiration may be used by the algae in photosynthesis.

Associations (symbiosis)

      Cnidarians enter into complex associations with a variety of other organisms, including unicellular algae, fishes, and crustaceans. Many of these relationships, such as those with zooxanthellae and zoochlorellae, are mutualistic symbioses—i.e., relationships benefiting both partners. Reef-forming corals, which possess zooxanthellae, form more substantial skeletons than do non-reef-forming corals, which lack zooxanthellae, for reasons that are not understood but are related to the algae. Many corals are so dependent on zooxanthellae that they cannot live in prolonged darkness, which is why coral reefs develop only in shallow, well-illuminated waters.

      There are species of sea anemones that live on gastropod shells inhabited by hermit crabs (hermit crab), a type of crustacean that must change shells as it grows. Some hermit crabs (crab) move the anemones with them from the old shells to the new. In other cases, the anemones take the initiative, somersaulting from a now-empty shell onto the newly obtained one. A few deep-sea anemones form the shells in which their crabs dwell. This adaptation eliminates the need to change shells, but the death of one partner probably results in the death of the other. Certain true crabs carry anemones on their backs and legs, or even in their claws. These associations benefit the anemone by providing it with transport, and sometimes it can steal food from its crustacean partner. In turn, the sea anemone protects its host from predators such as octopuses and other crabs.

      Gastropods (of the phylum Mollusca) also associate with cnidarians. Among the most remarkable are the nudibranchs (nudibranch) that eat anemones and hydroids and then sequester certain types of immature, undischarged cnidae from the prey. Once the cnidae have matured within the nudibranch, they can be used in its own defense.

      One of the best-known cnidarian symbioses is the mutualism between 10 species of tropical anemones and 26 species of anemone fish (such as the clown fish). These fishes live within the protective field of anemone tentacles, where they take refuge when a predator threatens. Immunity of the fishes to the stings of the nematocytes results from the thin layer of mucus that covers their bodies. It is unclear whether the mucous is made by the fishes themselves, or acquired by contact with the anemone's tentacles. Without its mucus, the clown fish, like any other small fish, may be stung to death and eaten by the anemone. Anemone fishes serve their hosts by driving away fishes that prey on anemones. Other fishes have a similar association with large medusae.

Form and function

Tissues and muscles

      Cnidarians consist of two cell layers: an outer ectoderm and an inner endoderm (the gastrodermis) that lines the coelenteron. Between these is sandwiched the mesoglea, a largely noncellular layer composed of a jellylike material permeated by a complex network of supporting fibres that may be microscopically thin or very thick. The fibres and jelly are elastic. In medusae, mesoglea comprises the bulk of the animal and forms a resilient skeleton. In polyps, the water-filled coelenteron acts as a hydrostatic skeleton, which, in concert with the mesoglea, maintains the form of these animals.

      Muscles in cnidarians are extensions of the bases of ectodermal and endodermal cells. Individual muscle cells are relatively long and may occur in dense tracts in jellyfish or sea anemones. Most cnidarian muscles, however, are thin sheets at the base of ectodermal and endodermal layers.

      In polyps, ectodermal muscles are oriented lengthwise along the cylindrical body and tentacles; endodermal ones are usually circular. Contraction of circular muscles against coelenteric fluid causes the polyp's body to elongate; contraction of longitudinal muscles causes it to shorten. Similar layers of muscles extend and contract the tentacles. Bending results from unequal contractions of longitudinal muscles on opposite sides of the body.

      In medusae, all muscles are ectodermal, restricted to the concave oral surface (subumbrellar surface), and organized into circular and radial tracts. Contraction of circular muscles squeezes the subumbrellar space, forcing out contained water and causing the medusa to move by jet propulsion. Recovery of the elastic mesoglea re-extends contracted muscle fibres. Radial muscle contraction distorts the bell, directing the water jet at angles to the longitudinal axis of the bell, allowing the medusa to steer.

Support mechanisms and skeletons

      Most members of what is usually considered a soft-bodied group have some sort of skeleton aside from the hydrostatic system described above. Both external and internal skeletons occur in the phylum, but only among polyps.

      Most hydroid polyps secrete a horny, chitinous external skeleton that is essentially a tube around the polyp and the network of stolons that interconnect members of a colony. As well as being protective, it confers stiffness for support and has joints for flexibility. A few scyphozoan polyps have comparable chitinous skeletons. Unlike those of hydroids, hydrocoral skeletons are composed of calcium carbonate and are internal by virtue of being shallowly penetrated by channels of living tissue. Hydrocorals, which include the order Milleporina (millepores), commonly called fire coral, and the precious red coral used for jewelry, form encrusting or branching skeletons similar to those of anthozoan corals.

      An anthozoan coral polyp, which resembles a sea anemone, can nearly completely retract into the calcareous cup it secrets around itself. This external skeleton underlies a continuous, superficial layer of tissue. Non-reef-forming corals typically are solitary or form small, rather delicately branched colonies, their polyps being relatively large and widely spaced. In some species of reef-forming corals, polyps are so tightly packed that their individual units share common walls. Skeletons may be encrusting, massive, or arborescent (treelike). The latter type of skeleton is delicate and typical of quiet waters at depth or in lagoons, while the former two predominate where water motion is strong. Skeleton is laid down in massive corals (coral reef) at a rate of about one centimetre per year; branching corals may grow considerably more rapidly. The largest corals represent cooperative efforts of up to 1,000,000 tiny individuals precipitating calcium carbonate over centuries. Few attain such proportions, however, and even the largest are eventually broken down by boring organisms such as algae, worms, sponges, and barnacles, as well as by physical processes.

      The last major category of cnidarian skeletons, formed by the anthozoan subclass Alcyonaria and the order Antipatharia, are internal. sea fan and sea whip skeletons consist of the horny protein gorgonin with calcareous spicules fused to form a solid or jointed central rod. Soft coral spicules are discrete, mostly microscopic objects of diverse shapes that vary from needle-like to club- and anchor-shaped. Located in the ectoderm, spicules stiffen the colony. In some species the several spicules that form a protective cup around each polyp may be several millimetres long. The alcyonarian Tubipora is known as the organ-pipe coral after the form of its red calcareous skeleton. Blue corals (the order Helioporacea) have skeletons of crystalline calcareous fibres fused into sheets, which are used for jewelry. Colonies of black coral resemble bushes and may stand more than three metres tall. Their skeletons, made entirely of proteinaceous material similar to gorgonin, are likewise used for jewelry.

      Sea anemones do not produce hard skeletons, although their close relatives in the order Zoanthinaria (zoanthid) incorporate foreign objects (sand grains, sponge spicules) into their body walls, which gives them rigidity and toughness. Small anemones that live high in the intertidal zone commonly inhabit abandoned barnacle tests (shells), thereby acquiring some of the benefits of a skeleton.

Nervous system and organs of sensation

      Medusae have a more highly developed nerve net than do polyps, a feature that is associated with the more active way of life of medusae. Swimming is coordinated by the nervous system. Nervous systems that are capable of conducting nerve impulses both quickly and slowly give these animals considerable behavioral responsiveness and flexibility. Ganglia or other accumulations of nerve cell bodies are not found in cnidarians, but there are gap junctions between neurons and between neurons and effectors, which allow the transmission of nerve impulses. Statocysts, located between the tentacles or near the tentacular base, inform the animal of its orientation with respect to gravitational forces. Light-sensitive ocelli (external patches of pigment and photoreceptor cells organized in either a flat disk or a pit) occur in some medusae of each of the three classes that possess this stage. Such sensory structures are closely associated with a nerve net.

      In the past nematocysts were considered independent effectors; that is, they were thought to fire upon appropriate stimulation, without mediation by the nervous system. Evidence, however, favours there being some organismal control over their firing, which may consist only of adjusting the threshold for firing, or the selectivity. Some scientists believe that nervous complexes associated with batteries (groups) of cnidae are the mechanism of control.

Digestion, respiration, and excretion

      Food is taken in and wastes are discharged through the mouth. Extracellular digestion occurs in the coelenteron, which has, in all except hydrozoans and some tiny members of the other classes, radial projections of the wall into the coelenteron that increase the surface area. Ingested material is broken down somewhat in the coelenteron and then taken up by endodermal cells for final intracellular digestion.

      Respiration and excretion in cnidarians are carried on by individual cells that obtain their oxygen directly from water—either that in the coelenteron or that of the environment—and return metabolic wastes to it. Thus, all physiological functions are carried out at no more than the tissue level of differentiation.

Defense and aggression: nematocysts

      Cnidae range from only about 10 to 100 micrometres (0.0004 to 0.004 inch) long, but they are among the most complex intracellular secretion products known. Each consists of a spherical or cigar-shaped capsule with an eversible, hollow tubule extending from one end. In the unfired state, the tubule is coiled within the capsule. When a cnidarian contacts a predator or prey item, the capsule opens and the tubule everts. The tubule may be adhesive, or it may entangle the object. Both types serve to hold food items. A third type of tubule is armed with spines that penetrate predator or prey. Toxins contained in the capsule are injected through the tubule into the object being held. Each cnida can be fired only once. Undifferentiated interstitial cells of the ectoderm and endoderm appear to be the source of the cnidoblasts (cells that produce cnidae).

Evolution

      The exact relationships between the different cnidarian groups are unknown. Among theories proposed on the evolution of the phylum Cnidaria, most treat the radial symmetry and tissue level of organization as evidence that the group is primitive (that is, it evolved before the evolution of bilateral symmetry) and hold that the medusa is the original body form, being the sexually reproductive phase of the life cycle. Another theory is that the original cnidarian was a planula-like organism that preceded both polyp and medusa. In either case, Hydrozoa is considered to be the most ancient of cnidarian classes, and Trachylina is thought to be the most primitive extant order of that group. An alternative view is that anthozoans are the stem of the phylum, which evolved from bilateral flatworms and is secondarily simplified. A corollary to this theory is that the polyp is the ancestral body form.

      Speculations about the origin of the phylum are not easily resolved, for preservable skeletal structures developed relatively late in cnidarian evolution. The oldest fossilized cnidarians were soft-bodied. Representatives of all four modern classes have been identified in Ediacaran fauna of the Precambrian Period (about 542 million years ago) known from more than 20 sites worldwide. As much as 70 percent of Ediacaran species have been considered to be cnidarians. Curiously, there are few fossil cnidarians of the Cambrian Period (542 million to 488 million years ago). The Conulariida, which existed from the Cambrian Period to the Triassic Period (251 million to 200 million years ago) are considered by some scientists to be skeletal remains of scyphopolyps, either ancestral to the coronates or without modern derivatives. Presumed fossil sea anemones are found in the early Cambrian Period. Colonies of Stromatoporoidea, considered to be an order of the class Hydrozoa that extended from the mid-Cambrian Period to the Cretaceous Period (about 146 million to 65 million years ago), produced massive skeletons. Although there were two groups of Paleozoic corals, neither of which has modern descendants, they were not great reef-builders during that era. Scleractinians arose in the mid-Triassic Period; blue corals, gorgonians, millepores, and hydrocorals have records from the Jurassic Period (about 200 million to 146 million years ago) or the Cretaceous Period to the present. Most other cnidarians are known only from the Holocene Epoch (within the last 10,000 years).

Classification

Annotated classification

      The following classification, limited to living cnidarians, generally follows that used by D.G. Fautin in S.P. Parker (ed.), Synopsis and Classification of Living Organisms, vol. 1 (1982), and L.H. Hyman, The Invertebrates, vol. 1, Protozoa Through Ctenophora (1940).

Phylum Cnidaria (cnidarian) (Coelenterata)

 Nematocyst-bearing, radial metazoans without organs. Have a cellular inner endoderm and outer ectoderm, separated by noncellular mesoglea. Polyp and medusa forms; either or both may be present in one life history. Most polyps have tentacles around mouth; tentacles of medusae at bell margin. One internal cavity, the coelenteron, has 1 opening to exterior, the mouth. About 9,000 species.

      Class Anthozoa

 Exclusively polypoid with biradial symmetry. Oral end a disk with central mouth and hollow tentacles arising at margin and/or on surface. Mouth leads to coelenteron via stomodaeum that has ciliated troughs (siphonoglyphs) for water transport into and out of coelenteron. Coelenteron divided by radial mesenteries that extend inward and insert on the stomodaeum (complete mesenteries) or not (incomplete mesenteries). About 6,000 species.

      Subclass Alcyonaria

 Octocorals. Polyps with 8 pinnately branched tentacles, 8 mesenteries, and a single siphonoglyph. Nearly all colonial with internal skeletons.

      Order Stolonifera

 Polyps of colony connected by stolons. Skeletons of spicules or horny external cuticle. Shallow tropical and temperate seas.

      Order Telestacea

 Long axial polyps bear lateral polyps. Skeleton of spicules fused with a horny material. Tropical.

      Order Gorgonacea

 Sea fans and sea whips. Colonies commonly arborescent with axial skeleton of gorgonin and/or calcareous spicules. Polyps rarely dimorphic. Tropical and subtropical.

      Order Alcyonacea

 Soft corals. Small to massive colonial forms. Lower parts of polyps fused into a fleshy mass; oral ends protrude. Internal skeleton of isolated calcareous spicules. Primarily tropical.

      Order Helioporacea (Coenothecalia)

 Blue coral. Massive lobed calcareous skeleton. Tropical; 1 Caribbean and 1 Indo-West Pacific species.

      Order Pennatulacea

 Sea pens (sea pen) and sea pansies. Fleshy, always dimorphic, unbranched colonies, with 1 axial polyp and many lateral ones. Polyp-free peduncle burrows into soft sediments; polyp-bearing distal end of the polyp (rachis) extends into water and may be completely retractile. Central skeleton a calcified axial rod; polyps and rachis have isolated calcareous spicules.

      Subclass Ceriantipatharia

 Black corals and tube anemones.

      Order Antipatharia

 Black coral. Large bushy colonies with thorny, hornlike axial skeleton formed by small polyps with 6 simple tentacles and 1 siphonoglyph. Mostly tropical and subtropical.

      Order Ceriantharia

 Tube anemones. Solitary polyps with 2 sets of tentacles (oral and marginal) that form feltlike tubes of specialized cnidae (ptychocysts) and burrow in soft sediments. Shallow waters worldwide.

      Subclass Zoantharia

 Sea anemones and corals. Six (or multiples of 6) tentacles (rarely branched). Mesenteries commonly arranged hexamerously. Solitary or colonial. Skeletons non-spicular calcareous, horny, or lacking. Usually 2 siphonoglyphs.

      Order Actiniaria

 Sea anemones (sea anemone). Solitary or clonal, never colonial; lacking skeleton; with or without basilar muscles. Mostly littoral or benthic, commonly attached to firm substrata but some burrow in soft sediments. Worldwide.

      Order Corallimorpharia

 Sea-anemone-like solitary or aggregated polyps lacking basilar muscles and skeleton. Coral-like muscles and nematocysts. Mostly tropical.

      Order Ptychodactiaria

 Sea-anemone-like, lacking ciliated tract on edge of mesenteries and basilar muscles. Both poles.

      Order Scleractinia (Madreporaria)

 True or stony corals. Mostly colonial; calcareous external skeleton; no basilar muscles or siphonoglyphs. Mostly tropical and subtropical.

      Order Zoanthinaria (Zoanthidea (zoanthid))

 Solitary, clonal, or colonial polyps resembling sea anemones. Lack skeleton but may incorporate debris into body wall, commonly epizoic. One complete and 1 incomplete mesentery per pair. Mostly tropical.

      Class Cubozoa

 Tropical, cuboidal medusae that swim strongly; box jellyfishes. Margin simple with single or grouped tentacles arising above the 4 corners. Polypoid stage of most species unknown. Fiercely stinging members can cause human fatalities. Contains 1 order, Cubomedusae.

      Class Hydrozoa

 Life histories may involve both polypoid and medusoid stages, but either may be suppressed or absent. Tetramerous or radially symmetrical medusae small, with shelf of tissue (velum) across lower part of bell, which reduces diameter of subumbrellar aperture (condition known as craspedote). Colonial forms commonly polymorphic. Coelenteron undivided. Gametes ripen in ectoderm. Only class with some freshwater members. 2,700 species.

      Order Actinulida

 Curious groups of solitary, motile cnidarians with features of both polyps and medusae. Europe; in marine sand.

      Order Chondrophora

 Floating polymorphic colonies supported by chitinous skeleton. Free medusae are produced; includes Velella. Oceanic; worldwide.

      Order Hydroida (hydroid)

 Hydroids. Usually colonial and polymorphic; release free medusae or retain modified medusoid reproductive structures on polyp colony. Polyps usually have a chitinous exoskeleton. Includes naked, solitary freshwater polyp Hydra. Largest order of Hydrozoa.

      Suborder Anthomedusae

 Medusae bell-shaped, with gonads on the stomach or sides of manubrium. Sensory structures consist of pigmented eyespots (ocelli). Skeleton, if present, lacks cup (hydrotheca) into which polyp may withdraw (a condition known as gymnoblastic); few species with calcareous exoskeleton. Most abundant in bays and shallow coastal waters.

      Suborder Leptomedusae

 Medusae saucer-shaped, but lacking in many species. Gonads on radial canals. Sensory structures usually statocysts. Hydroids with hydrothecae (condition known as calyptoblastic). All shallow marine waters.

      Suborder Limnomedusae

 Small medusae with gonads on stomach walls or radial canals. Polyps solitary or colonial, commonly with 1 or 2 tentacles, and no skeleton. Mostly freshwater.

      Order Milleporina

 Fire coral. Colonial forms producing massive calcareous skeletons. Gastrozooids and dactylozooids project through pores in surface of skeleton. Reduced, acraspedote (lacking a velum) nonfeeding medusae are released. Tropical.

      Order Siphonophora

 Pelagic polypoid colonies with greatest degree of polymorphism in phylum; lack medusae. Oceanic; worldwide. Includes Portuguese man-of-war, Physalia.

      Order Stylasterina

 Hydrocorals. Resembling millepores; colonies erect and branching or prostrate. Commonly yellow, red, or purple. Reduced medusae not freed; develop and produce gametes in cavities of skeleton (ampullae). Worldwide; includes precious red coral, Corallium.

      Order Trachylina

 Medusa dominant; reduced or no polyp stage. Statocysts and special sensory structures (tentaculocysts). Differ from other hydromedusae by having tentacles inserted above umbrellar margin. Oceanic, mostly warmer waters.

      Suborder Laingiomedusae

 Medusae with features of both Narcomedusae and Trachymedusae. Polyp unknown.

      Suborder Narcomedusae

 Scalloped margin; gonads on stomach walls. Manubrium lacking.

      Suborder Trachymedusae

 Smooth bell margin; gonads on radial canals arising from the stomach. Polyp and asexual reproduction absent.

      Class Scyphozoa

 Exclusively marine group in which acraspedote medusae predominate. Life histories commonly involve alternation of a very small polyp, the scyphistoma, with a medusa, which develops from an ephyra released by the polyp. Coelenteron of both divided by 4 longitudinal septa producing tetramerous radial symmetry. Gonads endodermal. Marginal sensory structures (rhopalia) with statocysts and/or ocelli. Most abundant in coastal waters, but oceanic species exist. About 200 species.

      Order Coronatae

 Large medusae that are conical, dome-shaped, or flattened, with furrow around bell above scalloped margin. Some species have scyphistoma stage with external chitinous skeleton. Oceanic, some species living at great depths.

      Order Rhizostomae

 Medusae like those of Semaeostomeae but with mouth subdivided into minute pores that connect with coelenteron. Mostly tropical. Deep-water species may lack polypoid stage.

      Order Semaeostomeae

 Most common and best known jellyfishes. Full alternation of polyp and medusa stages. Bell domed or flattened, with the margin scalloped into 8 or more sections. Edges of single mouth drawn out into 4 long arms. Most species in warm, coastal waters, a few in frigid waters; some oceanic. Includes the giant Cyanea arctica, which may attain 2 m in diameter.

      Order Stauromedusae

 Sessile jellyfish that are vase-, goblet-, or trumpet-shaped, and usually bear 8 groups of tentacles. No more than 2–3 cm long. Apparently lacking polypoid stage. Temperate and cold temperate waters worldwide.

Critical appraisal

      The classification of living cnidarians is relatively stable and generally accepted. One unanswered question relates to their evolutionary position among the lower Metazoa (a division of the animal kingdom that includes all phyla except the Protozoa). Members of the small phylum Ctenophora (ctenophore) (comb jellies, sea walnuts) are superficially similar to medusae, having a gelatinous body with one opening, and tentacles. The one species of Ctenophora possessing nematocysts had been thought to link the phyla. Mills and Miller have shown that members of the phylum Ctenophora obtain their cnidae by preying on a medusa, and the similarity in body form is considered convergence due to the pelagic way of life.

      Anthozoa is a well-defined, coherent group, but relationships among its components are poorly understood, and the ranking of some of them is disputed. Some regard corallimorpharians as scleractinians that lack a skeleton. Similarity of larval ceriantharians to antipatharian polyps is the rationale for subclass Ceriantipatharia. Morphology of antipatharians is, however, in some ways, nearer that of alcyonarians than of zoantharians, and alternative schemes place Antipatharia in subclass Alcyonaria. Ceriantharia, too, sits uncomfortably in Zoantharia, although it bears no special relationship to the Alcyonaria. It is generally viewed as a divergent, early offshoot of the anthozoan line. Treatment of both Antipatharia and Ceriantharia as distinct subclasses of Anthozoa not particularly closely related to one another might better express their evolutionary relationships.

      Many cnidarian biologists continue to regard Cubozoa as an order of Scyphozoa. Cubozoans have features of both Scyphozoa and Hydrozoa, but the complete metamorphosis of polyp into medusa supports its placement in a class intermediate between the other two.

      Hydrozoan suborder Limnomedusae is not accepted by some workers, and the groups assigned to it are treated as members of the other suborders. In substantial ways, characters of many Limnomedusae bridge the differences between the Anthomedusae and the Leptomedusae. The hydrozoan order Chondrophora has morphological characteristics suggesting that the group might be better treated as part of Anthomedusae than as a discrete order.

Additional Reading

The standard technical treatment of the Cnidaria is Libbie Henrietta Hyman, The Invertebrates, vol. 1, Protozoa Through Ctenophora (1940). The history of the phylum as deduced from the fossil record is covered in Raymond C. Moore (ed.), Treatise on Invertebrate Paleontology, part F, Coelenterata (1956, reprinted 1967). C.E. Mills and R.L. Miller, “Ingestion of a Medusa (Aegina citrea) by the Nematocyst-Containing Ctenophore Haeckelia rubra (formerly Euchlora rubra): Phylogenetic Implications,” Marine Biology, 78(2):215–221 (1984), offers information on the relationship between Ctenophora and Cnidaria. The following proceedings of international symposia on cnidarians provide useful information: W.J. Rees (ed.), The Cnidaria and Their Evolution (1966); Takasi Tokioka and Saburō Nishimura (eds.), Recent Trends in Research in Coelenterate Biology (1973); G.O. Mackie (ed.), Coelenterate Ecology and Behavior (1976); and P. Tardent and R. Tardent (eds.), Developmental and Cellular Biology of Coelenterates (1980). See also the following proceedings of international coral reef symposia: Proceedings of the Third International Coral Reef Symposium, 2 vol. (1977); The Reef and Man: Proceedings of the Fourth International Coral Reef Symposium, 2 vol. (1982); and French Polynesian Coral Reefs: Proceedings of the Fifth International Coral Reef Congress, 6 vol. (1985). The major justification of recognizing class Cubozoa is B. Werner, “Bau und Lebensgeschichte des Polypen von Tripedalia cystophora (Cubozoa, class. nov., Carybdeidae) und seine Bedeutung für die Evolution der Cnidaria,” Helgoländer wissenschaftliche Meeresuntersuchungen, 27:461–504 (1975).Cadet H. Hand, Jr. Daphne Gail Fautin

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