Introduction to Cubozoa: The Box Jellies!

Cubozoans are marvelous animals.

They look like your basic jellyfish, but they can swim pretty fast, maneuver around things, and see fairly well despite not having a brain. Believe it or not. In general, box jellies are similar in form to the "true" jellyfish, known as scyphozoans. However, it is relatively easy to tell the two groups apart. Cubozoans have a square shape when viewed from above. (Gee, maybe that's how they got their name.) They also have four evenly spaced out tentacles or bunches of tentacles and well-developed eyes. Not surprisingly, given their squishy nature, there are not many fossil cubozoans known. Today, there are about 20 known species found in tropical and semitropical waters. The Australian stinger Chironex fleckeri is among the deadliest creatures in the world, having caused human fatalities.

If you're curious, you can learn a whole lot more about cubozoans.
Fossil Record Anthracomedusa turnbulli Life History and Ecology Activity and Agility Life Cycle Phylogeny, Systematics and Classification The Test: Carybdeid or Chirodropid? Carukia barnesi Carybdea sivickisi Carybdeid from Darwin, Australia Chironex fleckeri Chiropsalmus sp. Phylogenetic Position of Cubozoa General Morphology Manubrium and Mouth Velarium and Velarial Canals

Rhopalia and Eyes Nematocysts Planula and Polyp

The Cubozoan Anthracomedusa turnbulli

This is a really cool fossil. Think of how rare it must be for an animal without a hard shell or skeleton to be preserved as a fossil. Unfortunately, this is the only specimen of Anthracomedusa that we have in our collections at UCMP. Other specimens from Mazon Creek better reveal the square shape of the bell, as well as the four bunches of tentacles.

Cubozoa: Life History and Ecology

Cubozoans are voracious predators.

Many jellyfish and other gelatinous animals go unnoticed by those who visit the ocean. Like this "herd" of cubozoans in the picture above, gelatinous animals are often difficult to see. However, they are important members of marine ecosystems. For instance, box jellies are known to eat fish, worms, and crustacean arthropods. When the tentacles, which can stretch to ten or more times the height of the bell, come into contact with prey, nematocysts fire into the prey's skin. Tiny barbs of the nematocysts hold onto and transfer venom to the prey. As the prey is immobilized by the venom, the tentacle contracts and pulls the prey near to the bell. Then, an amazing thing occurs. The muscular pedalium bends and pushes the tentacle and prey into the bell of the medusa. The manubrium then reaches out for the prey and the mouth expands and engulfs it. The entire feeding process is relatively fast, taking place in as little as one minute. Cubozoans are not quite at the top of the food web, as they are sometimes eaten by large fish and sea turtles.

Cubozoans are active and agile swimmers.

Jellyfish are typically referred to as planktonic, meaning that their swimming is not strong enough to prevent them from drifting with currents. As a consequence, many jellyfish wash up on beaches. However, box jellies are very rarely found on beaches. This is because they are unusually strong swimmers. It has been reported that large specimens of Chironex fleckeri can swim as fast as two meters in one second. However, that speed has not been confirmed. Cubozoans (of various sizes) have commonly been observed to swim a meter in five to ten seconds. Presumably, cubozoans use strong muscles, as well as the Velarium, to achieve these speeds. Not only are cubozoans fast, but they are remarkably agile. They have been reported to maneuver around the pilings of peers and to flee would be human collectors. These behaviours can only be achieved because cubozoans have vision. Few experiments

have been tried in order to determine how well cubozoans can see, but it is clear that they can see well enough to avoid swimming into things and being captured. It is also likely that they can see eachother, which might help explain some interesting observations about their mating.

Cubozoan Life Cycle

Some species of cubozoans appear to pair up, male with female, in order to mate. The male puts his tentacles into the bell of the female and appears to pass packets of sperm. At least one other species has been observed in large mating aggregations. Mating appears to occur once every year. Fertilization takes place inside the females. In some species the fertilized eggs are released into the water column where they develop into planulae, while in others development into planulae occurs inside the female. Planulae swim in the water column for a few days and then settle on to the substrate. After settlement, the planulae grow into polyps. The polyps can move around, and they frequently bud off additional polyps. After a few months of feeding, the polyps are mature. Then, each polyp metamorphoses into a single juvenile medusa (above right). During metamorphosis, the polyp tentacles are resorbed and four new tentacles and four rhopalia are formed. With a couple of contractions, the entire individual becomes detached and swims away as a juvenile medusa.

Cubozoa: Systematics
Systematic Relationships within Cubozoa
Cubozoans have yet to be the subjects of a published phylogenetic analysis. However, UCMP graduate student Lisa-ann Gershwin is working on just such an analysis as part of her dissertation. Until she finishes this project, we are presenting a phylogenetic representation of the current classification scheme for Cubozoa.

There are two main groups of cubozoans, Chirodropidae and Carybdeidae. The two groups are relatively easy to tell apart. In carybdeids, each tentacle is connected to a single pedalium. Usually there are four pedalia each with a tentacle, however, in Tripedalia species, each corner of the bell has two or three tentacles each connected to a single pedalium. Chirodropids always have four pedalia, one at each corner, with multiple tentacles. Are you ready for the test?

The Phylogenetic Position of Cubozoa

The phylogenetic position of Cubozoa within Cnidaria has been the subject of a great deal of debate. Classically, cubozoans were viewed as a subgroup of the "true jellyfish" or Scyphozoa (as shown to the left below).

However, when it was observed that the cubozoan polyp and life cycle were rather different than those of scyphozoans, Cubozoa was placed on its own. Some scientists have proposed that cubozoans are the closest living relatives of scyphozoans, as shown in the middle above. Still others have argued that Cubozoa is more closely related to Hydrozoa (as shown to the right). Post-doctoral researcher and former UCMP graduate student, Allen G. Collins is presently working on this question and its evolutionary implications.

The

Cubozoan -- Carukia barnesi

A slight sting from Carukia barnesi, which is always less than 2 centimeters tall (unless it is misidentified), can be a terrible experience, called Irukandji Syndrome, for a human. The name Irukandji derives from the aboriginal tribe that formerly lived in the Cairns region of Australia where most stings occur. At first, the sting is mild. About ten minutes later, the skin where the sting occurred begins to sweat intensely. Within roughly 20 to 30 minutes, there are intense pains in the stomach, limbs, back and head. Breathing is difficult; vomiting or coughing may occur. It is probably lethal, although this has not been definitively demonstrated.

The Cubozoan Carybdea sivickisi

If you go out onto the reef flats at night in Guam with a flashlight, you might run into this little guy. If you do see it, be careful. They appear to be attracted to lights and they can deliver a painful little sting. According to Gustav Paulay, this cubozoan spends its days attached by its bell to the undersides of rocks. In the lab, they have been observed to stick themselves to a variety of substrates, including glass, dead coral, and live algae. To attach themselves, these box jellies have "sticky pads". The larger white patches near the top of the bell of the specimen above might be the sticky pads. Another interesting tidbit of information about this cubozoan is that it comes out in droves during the once-a-year coral spawning event on the Great Barrier Reef, Australia. Carybdea sivickisi has a bell that is about 1 centimeter tall in a fully grown individual.

A Cubozoan -- Carybdeid from Darwin, Australia

UCMP graduate student Lisa-ann Gershwin is in the process of giving this beautiful critter a formal name and scientific description.

The Cubozoan -- Chironex fleckeri

Be careful handling this critter from Northern Australia! Chironex fleckeri grows to about the size of a human head, and has tentacles up to three meters long. A big sting from this guy can easily kill a human, with death occurring in as little as three minutes. There have been roughly 100 deaths due to Chironex stings during the past 100 years in northern Australia. However, many people have been stung and not been killed. It is likely that these cubozoans swim away when they come into contact with something as large as a human. According to Phil Alderslade, contact with six to eight meters of tentacle is necessary to deliver enough venom to kill a person. The bell of Chironex fleckeri does not have nematocysts. Fortunately, these box jellies are in the business of catching and eating fish and crustaceans. Chironex individuals do not inhabit reef environments. Instead, the only polyps that have ever been observed in nature were living attached to the undersides of stones in an estuary of a river in northern Australia. The polyps metamorphose into juvenile medusae beginning in the austral spring (September) and continuing until the first large summer rains (usually in January). Medusae are then flushed out into the near shore waters.

The Cubozoan -- Chiropsalmus sp.

This species of Chiropsalmus is found in northern Australia. It is sometimes confused with the deadly Chironex fleckeri, but this box jelly only delivers a painful sting.

Cubozoa: More on Morphology

General Form of the Medusa

The cubozoan bell is square in horizontal cross section. Inside the bell are the manubrium and mouth. A flap of tissue called the velarium is located along the underside of the bell. Muscular fleshy pads called pedalia are located at the corners of the bell. One or more tentacles are connected to each pedalium. Actually, that's the way to tell apart the two main groups of cubozoans (Chirodropidae and Carybdeidae). Carybdeids always have one tentacle per pedalium, while chirodropids have multiple tentacles attached to each pedalium. Take the test? Like other cnidarians, cubozoans are composed of two layers of cells, ectoderm and endoderm. The gut, or stomach cavity, is partitioned by septa and extends to the tentacles through pedalial canals. On the bell, located midway between the pedalia, are four sensory structures called rhopalia.

Cubozoan Sensory Structures

If you take a close look inside the rhopaliar niche of a cubozoan (left), you'll see a remarkable thing . . . . looking back at you. That's right, cubozoans have eyes, and surprisingly complex ones at that. In this close-up of a rhopalium (right), you can see six reddish spots, all of which are sensitive to light. The four smallest spots are relatively simple. However, the two larger regions actually contain lenses, corneas, and retinas, not so unlike those in your eyes. It is still unclear how the images created by these lenses are interpreted by cubozoans since they do not have brains. A cubozoan can look both inward towards its mouth and manubrium and outward since each rhopalium dangles by a muscular stalk. Inside each rhopalium, located below the eyes, is an organ called a statocyst. Inside each statocyst is a hard nodule composed of calcium sulfate, the statolith. Statoliths appear to have daily growth rings. The statocysts are sensitive to orientation, and thus allow cubozoans to sense whether they are upside-down, sideways, or rightside-up.

Cubozoan Stinging Cells

Like all cnidarians, cubozoans are endowed with nematocysts, cells that fire a barb and transfer venom. As you can see to the right, the barb is coiled up inside a capsule. When a nematocyst touches something that might be prey or predator, the barb uncoils and fires from the capsule along with venom. Nematocysts are concentrated in rings (right) on the tentacles of cubozoans. This makes sense. When the tentacles capture a prey item, they contract, and the rings allow for maximum contact and transfer of venom between nematocyst and prey. In some cubozoans, such as Chironex fleckeri, nematocysts are absent from the bell.

The Cubozoan Larva and Polyp

Unfortunately, we do not have (yet) any good pictures of cubozoan larvae or polyps. These stages are not very well known in cubozoans. In fact, it was not until about 1970 that the complete life cycle of a cubozoan was observed. The cnidarian larva is called a planula. Cubozoan planulae are pear-shaped (that's "pyriform" in scientist lingo), have pigment spots which may be sensitive to light, and swim for a few days using cilia. After a planula settles, it grows into a polyp. The cubozoan polyp can crawl around like an inchworm and bud off more polyps. Its form is relatively simple. It has a mouth surrounded by up to 24 tentacles. The polyps do not closely resemble the polyps of scyphozoans. These differences may indicate that cubozoans have a separate evolutionary origin from scyphozoans. Finally, it is worth mentioning in case you go on to read more about cubozoans (or any other cnidarians for that matter) that the plains of symmetry in a cubozoan have special names. The interradii are marked by the pedalia; rhopalia occur along the perradii.

The Mouth and Manubrium of a Cubozoan

If the prey of a cubozoan is still alive at this point, this could be the last thing it sees. During feeding, the pedalium pushes the prey inside the bell of the medusa, the manubrium (which is the tube that connects the stomach on one end to the the fourparted mouth on the other) reaches out and finds the prey. The mouth then expands and engulfs the prey.

The Velarium of a Cubozoan

No, that is not some weird fish. That is just an odd angle view into the bell of a box jelly. What it shows really nicely (besides the rhopalia and eyes is the velarium, a ring of tissue that extends around the bottom inside of the bell. When the medusa contracts during swimming, increased thrust is created because the velarium narrows the opening at the bottom of the bell. The velarium increases the efficiency of the jet propulsion. Presumably, the velarium is at least partly responsible for the great speeds with which cubozoans are able to swim. In the close-up below, you can see velarial canals. These canals are extensions of the gut and they are very important in telling one species of cubozoan from another.