Class Anthozoa

The class Anthozoa (“flower animals”) includes sea anemones (Figure 1), sea pens, and corals, with an estimated number of 6,100 described species. Sea anemones are usually brightly colored and can attain a size of 1.8 to 10 cm in diameter. Individual animals are cylindrical in shape and are attached directly to a substrate.

Figure 1. The sea anemone is shown (a) photographed and (b) in a diagram illustrating its morphology. (credit a: modification of work by “Dancing With Ghosts”/Flickr; credit b: modification of work by NOAA)

The mouth of a sea anemone is surrounded by tentacles that bear cnidocytes. The slit-like mouth opening and flattened pharynx are lined with ectoderm. This structure of the pharynx makes anemones bilaterally symmetrical. A ciliated groove called a siphonoglyph is found on two opposite sides of the pharynx and directs water into it. The pharynx is the muscular part of the digestive system that serves to ingest as well as egest food, and may extend for up to two-thirds the length of the body before opening into the gastrovascular cavity. This cavity is divided into several chambers by longitudinal septa called mesenteries. Each mesentery consists of a fold of gastrodermal tissue with a layer of mesoglea between the sheets of gastrodermis. Mesenteries do not divide the gastrovascular cavity completely, and the smaller cavities coalesce at the pharyngeal opening. The adaptive benefit of the mesenteries appears to be an increase in surface area for absorption of nutrients and gas exchange, as well as additional mechanical support for the body of the anemone.

Sea anemones feed on small fish and shrimp, usually by immobilizing their prey with nematocysts. Some sea anemones establish a mutualistic relationship with hermit crabs when the crab seizes and attaches them to their shell. In this relationship, the anemone gets food particles from prey caught by the crab, and the crab is protected from the predators by the stinging cells of the anemone. Some species of anemone fish, or clownfish, are also able to live with sea anemones because they build up an acquired immunity to the toxins contained within the nematocysts and also secrete a protective mucus that prevents them from being stung.

The structure of coral polyps is similar to that of anemones, although the individual polyps are usually smaller and part of a colony, some of which are massive and the size of small buildings. Coral polyps feed on smaller planktonic organisms, including algae, bacteria, and invertebrate larvae. Some anthozoans have symbiotic associations with dinoflagellate algae called zooxanthellae. The mutually beneficial relationship between zooxanthellae and modern corals—which provides the algae with shelter—gives coral reefs their colors and supplies both organisms with nutrients. This complex mutualistic association began more than 210 million years ago, according to a new study by an international team of scientists. That this symbiotic relationship arose during a time of massive worldwide coral-reef expansion suggests that the interconnection of algae and coral is crucial for the health of coral reefs, which provide habitat for roughly one-fourth of all marine life. Reefs are threatened by a trend in ocean warming that has caused corals to expel their zooxanthellae algae and turn white, a process called coral bleaching.

Anthozoans remain polypoid (note that this term is easily confused with “polyploid”) throughout their lives and can reproduce asexually by budding or fragmentation, or sexually by producing gametes. Male or female gametes produced by a polyp fuse to give rise to a free-swimming planula larva. The larva settles on a suitable substratum and develops into a sessile polyp.

Class Scyphozoa

Class Scyphozoa (“cup animals”) includes all (and only) the marine jellies, with about 200 known species. The medusa is the prominent stage in the life cycle, although there is a polyp stage in the life cycle of most species. Most jellies range from 2 to 40 cm in length but the largest scyphozoan species, Cyanea capillata, can reach a size of two meters in diameter. Scyphozoans display a characteristic bell-like morphology (Figure 2).

Figure 2. A jelly is shown (a) photographed and (b) in a diagram illustrating its morphology. (credit a: modification of work by “Jimg944″/Flickr; credit b: modification of work by Mariana Ruiz Villareal)

In the sea jelly, a mouth opening is present on the underside of the animal, surrounded by hollow tentacles bearing nematocysts. Scyphozoans live most of their life cycle as free-swimming, solitary carnivores. The mouth leads to the gastrovascular cavity, which may be sectioned into four interconnected sacs, called diverticuli. In some species, the digestive system may branch further into radial canals. Like the septa in anthozoans, the branched gastrovascular cells serve two functions: to increase the surface area for nutrient absorption and diffusion, and to support the body of the animal.

In scyphozoans, nerve cells are organized in a nerve net that extends over the entire body, with a nerve ring around the edge of the bell. Clusters of sensory organs called rhopalia may be present in pockets in the edge of the bell. Jellies have a ring of muscles lining the dome of the body, which provides the contractile force required to swim through water, as well as to draw in food from the water as they swim. Scyphozoans have separate sexes. The gonads are formed from the gastrodermis and gametes are expelled through the mouth. Planula larvae are formed by external fertilization; they settle on a substratum in a polypoid form. These polyps may bud to form additional polyps or begin immediately to produce medusa buds. In a few species, the planula larva may develop directly into the medusa. The life cycle (Figure 3) of most scyphozoans includes both sexual medusoid and asexual polypoid body forms.

Figure 3. Scyphozoan life cycle. The lifecycle of most jellyfish includes two stages: the medusa stage and the polyp stage. The polyp reproduces asexually by budding, and the medusa reproduces sexually. (credit “medusa”: modification of work by Francesco Crippa)

Class Cubozoa

This class includes jellies that have a box-shaped medusa, or a bell that is square in cross-section, and are colloquially known as “box jellyfish.” These species may achieve sizes of 15 to 25 cm, but typically members of the Cubozoa are not as large as those of the Scyphozoa. However, cubozoans display overall morphological and anatomical characteristics that are similar to those of the scyphozoans. A prominent difference between the two classes is the arrangement of tentacles. The cubozoans contain muscular pads called pedalia at the corners of the square bell canopy, with one or more tentacles attached to each pedalium. In some cases, the digestive system may extend into the pedalia. Nematocysts may be arranged in a spiral configuration along the tentacles; this arrangement helps to effectively subdue and capture prey. Cubozoans include the most venomous of all the cnidarians (Figure 4).

These animals are unusual in having image-forming eyes, including a cornea, lens, and retina. Because these structures are made from a number of interactive tissues, they can be called true organs. Eyes are located in four clusters between each pair of pedalia. Each cluster consists of four simple eye spots plus two image-forming eyes oriented in different directions. How images formed by these very complex eyes are processed remains a mystery, since cubozoans have extensive nerve nets but no distinct brain. Nontheless, the presence of eyes helps the cubozoans to be active and effective hunters of small marine animals like worms, arthropods, and fish.

Cubozoans have separate sexes and fertilization occurs inside the female. Planula larvae may develop inside the female or be released, depending on species. Each planula develops into a polyp. These polyps may bud to form more polyps to create a colony; each polyp then transforms into a single medusa.

Figure 4. The (a) tiny cubazoan jelly Malo kingi is thimble shaped and, like all cubozoan jellies, (b) has four muscular pedalia to which the tentacles attach. M. kingi is one of two species of jellies known to cause Irukandji syndrome, a condition characterized by excruciating muscle pain, vomiting, increased heart rate, and psychological symptoms. Two people in Australia, where Irukandji jellies are most commonly found, are believed to have died from Irukandji stings. (c) A sign on a beach in northern Australia warns swimmers of the danger. (credit c: modification of work by Peter Shanks)

Class Hydrozoa

Hydrozoa is a diverse group that includes nearly 3,200 species; most are marine, although some freshwater species are known (Figure 5). Most species exhibit both polypoid and medusoid forms in their lifecycles, although the familiar Hydra has only the polyp form. The medusoid form has a muscular veil or velum below the margin of the bell and for this reason is called a hydromedusa. In contrast, the medusoid form of Scyphozoa lacks a velum and is termed a scyphomedusa.

The polyp form in these animals often shows a cylindrical morphology with a central gastrovascular cavity lined by the gastrodermis. The gastrodermis and epidermis have a simple layer of mesoglea sandwiched between them. A mouth opening, surrounded by tentacles, is present at the oral end of the animal. Many hydrozoans form sessile, branched colonies of specialized polyps that share a common, branching gastrovascular cavity (coenosarc), such as is found in the colonial hydroid Obelia.

Free-floating colonial species called siphonophores contain both medusoid and polypoid individuals that are specialized for feeding, defense, or reproduction. The distinctive rainbow-hued float of the Portuguese man o’ war (Physalia physalis) creates a pneumatophore with which it regulates buoyancy by filling and expelling carbon monoxide gas. At first glance, these complex superorganisms appear to be a single organism; but the reality is that even the tentacles are actually composed of zooids laden with nematocysts. Thus, although it superficially resembles a typical medusozoan jellyfish, P. physalis is a free-floating hydrozoan colony; each specimen is made up of many hundreds of organisms, each specialized for a certain function, including motility and buoyancy, feeding, reproduction and defense. Although they are carnivorous and feed on many soft bodied marine animals, P. physalis lack stomachs and instead have specialized polyps called gastrozooids that they use to digest their prey in the open water.

Physalia has male and female colonies, which release their gametes into the water. The zygote develops into a single individual, which then buds asexually to form a new colony. Siphonophores include the largest known floating cnidarian colonies such as Praya dubia, whose chain of zoids can get up to 50 meters (165 feet) long. Other hydrozoan species are solitary polyps (Hydra) or solitary hydromedusae (Gonionemus). One defining characteristic shared by the hydrozoans is that their gonads are derived from epidermal tissue, whereas in all other cnidarians they are derived from gastrodermal tissue.

Figure 5. Hydrozoans. The polyp colony Obelia (a), siphonophore colonies Physalia (b) physalis, known as the Portuguese man o‘ war and Velella bae (c), and the solitary polyp Hydra (d) have different body shapes but all belong to the family Hydrozoa. (credit b: modification of work by NOAA; scale-bar data from Matt Russell)

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