Cnidaria

Cnidaria (/nadri/[4] ) is a phylum containing over cnidarians, as well as the position of cnidarians as the
10,000[5] species of animals found exclusively in aquatic sister group of bilaterians.[7] Fossil cnidarians have been
(freshwater and marine) environments: they are predom- found in rocks formed about 580 million years ago, and
inantly marine species. Their distinguishing feature is other fossils show that corals may have been present
cnidocytes, specialized cells that they use mainly for cap- shortly before 490 million years ago and diversied a few
turing prey. Their bodies consist of mesoglea, a non- million years later. However, molecular clock analysis
living jelly-like substance, sandwiched between two lay- of mitochondrial genes suggests a much older age for the
ers of epithelium that are mostly one cell thick. They crown group of cnidarians, estimated around 741 million
have two basic body forms: swimming medusae and years ago, almost 200 million years before the Cambrian
sessile polyps, both of which are radially symmetrical period as well as any fossils.[8]
with mouths surrounded by tentacles that bear cnido-
cytes. Both forms have a single orice and body cavity
that are used for digestion and respiration. Many cnidar-
ian species produce colonies that are single organisms
composed of medusa-like or polyp-like zooids, or both
(hence they are trimorphic). Cnidarians activities are 1 Distinguishing features
coordinated by a decentralized nerve net and simple re-
ceptors. Several free-swimming species of Cubozoa and
Further information: Sponge, Ctenophore, and Bilateria
Scyphozoa possess balance-sensing statocysts, and some
have simple eyes. Not all cnidarians reproduce sexually,
with many species having complex life cycles of asexual Cnidarians form an animal phylum that are more complex
polyp stages and sexual medusae. Some, however, omit than sponges, about as complex as ctenophores (comb
either the polyp or the medusa stage. jellies), and less complex than bilaterians, which in-
clude almost all other animals. However, both cnidarians
Cnidarians were formerly grouped with ctenophores in
and ctenophores are more complex than sponges as they
the phylum Coelenterata, but increasing awareness of
have: cells bound by inter-cell connections and carpet-
their dierences caused them to be placed in separate
like basement membranes; muscles; nervous systems; and
phyla. Cnidarians are classied into four main groups:
some have sensory organs. Cnidarians are distinguished
the almost wholly sessile Anthozoa (sea anemones, corals,
from all other animals by having cnidocytes that re like
sea pens); swimming Scyphozoa (jellysh); Cubozoa
harpoons and are used mainly to capture prey. In some
(box jellies); and Hydrozoa, a diverse group that in-
species, cnidocytes can also be used as anchors.[9]
cludes all the freshwater cnidarians as well as many ma-
rine forms, and has both sessile members, such as Hydra, Like sponges and ctenophores, cnidarians have two main
and colonial swimmers, such as the Portuguese Man o' layers of cells that sandwich a middle layer of jelly-
War. Staurozoa have recently been recognised as a class like material, which is called the mesoglea in cnidarians;
in their own right rather than a sub-group of Scyphozoa, more complex animals have three main cell layers and
and Myxozoa and Polypodiozoa were only rmly recog- no intermediate jelly-like layer. Hence, cnidarians and
nized as cnidarians in 2007.[6] ctenophores have traditionally been labelled diploblastic,
along with sponges.[9][10] However, both cnidarians and
Most cnidarians prey on organisms ranging in size from
ctenophores have a type of muscle that, in more com-
plankton to animals several times larger than them-
plex animals, arises from the middle cell layer.[11] As
selves, but many obtain much of their nutrition from
a result, some recent text books classify ctenophores as
dinoagellates, and a few are parasites. Many are preyed
triploblastic,[12] and it has been suggested that cnidarians
on by other animals including starsh, sea slugs, sh and
evolved from triploblastic ancestors.[11]
turtles. Many scleractinian coralswhich form the struc-
tural foundation for coral reefspossess polyps that are
lled with zooxanthellae. While reef-forming corals are
almost entirely restricted to warm and shallow marine wa-
ters, other cnidarians can be found at great depths, in
polar regions, and in freshwater. 2 Description
Recent phylogenetic analyses support monophyly of

1
2 2 DESCRIPTION

2.1 Basic body forms 2.2 Skeletons

In medusae the only supporting structure is the mesoglea.
Hydra and most sea anemones close their mouths when
they are not feeding, and the water in the digestive
cavity then acts as a hydrostatic skeleton, rather like a
water-lled balloon. Other polyps such as Tubularia use
columns of water-lled cells for support. Sea pens stien
the mesoglea with calcium carbonate spicules and tough
brous proteins, rather like sponges.[10]
In some colonial polyps, a chitinous periderm gives sup-
Aboral port and some protection to the connecting sections and
end to the lower parts of individual polyps. Stony corals se-
Oral end crete massive calcium carbonate exoskeletons. A few
Mouth polyps collect materials such as sand grains and shell frag-
Oral end ments, which they attach to their outsides. Some colo-
Aboral end nial sea anemones stien the mesoglea with sediment
Exoderm particles.[10]
Gastroderm (Endoderm)
Mesoglea
Digestive cavity 2.3 Main cell layers
[10]
Medusa (left) and polyp (right)
Adult cnidarians appear as either swimming medusae or Cnidaria are diploblastic animals; in other words, they
have two main cell layers, while more complex animals
are triploblasts having three main layers. The two main
cell layers of cnidarians form epithelia that are mostly one
cell thick, and are attached to a brous basement mem-
brane, which they secrete. They also secrete the jelly-like
mesoglea that separates the layers. The layer that faces
outwards, known as the ectoderm (outside skin), gen-
erally contains the following types of cells:[9]

Epitheliomuscular cells whose bodies form part of

Oral end of actinodiscus polyp, with close-up of the mouth
sessile polyps, and many hydrozoan species are known
to alternate between the two forms. Both are radially
symmetrical , like a wheel and a tube respectively. Since
these animals have no heads, their ends are described as
oral (nearest the mouth) and aboral (furthest from the
mouth). Most have fringes of tentacles equipped with
cnidocytes around their edges, and medusae generally
have an inner ring of tentacles around the mouth. Some
hydroids may consist of colonies of zooids that serve dif-
ferent purposes, such as defense, reproduction and catch-
ing prey. The mesoglea of polyps is usually thin and often
soft, but that of medusae is usually thick and springy, so
that it returns to its original shape after muscles around In addition to epitheliomuscular, nerve and interstitial
the edge have contracted to squeeze water out, enabling cells, the inward-facing gastroderm (stomach skin) con-
medusae to swim by a sort of jet propulsion.[10]
the epithelium but whose bases extend to form
muscle bers in parallel rows.[16] The bers of the
outward-facing cell layer generally run at right an-
gles to the bers of the inward-facing one. In
Anthozoa (anemones, corals, etc.) and Scyphozoa
(jellysh), the mesoglea also contains some muscle
cells.[10]
Cnidocytes, the harpoon-like nettle cells that give
the phylum Cnidaria its name. These appear be-
tween or sometimes on top of the muscle cells.[9]
Nerve cells. Sensory cells appear between or some-
times on top of the muscle cells,[9] and communi-
cate via synapses (gaps across which chemical sig-
nals ow) with motor nerve cells, which lie mostly
between the bases of the muscle cells.[10]
Interstitial cells, which are unspecialized and can
replace lost or damaged cells by transforming into
the appropriate types. These are found between the
bases of muscle cells.[9]
tains gland cells that secrete digestive enzymes. In some
2.5 Cnidocytes 3

species it also contains low concentrations of cnidocytes, Most species have nematocysts.[9]
which are used to subdue prey that is still struggling.[9][10]
Spirocysts do not penetrate the victim or inject
The mesoglea contains small numbers of amoeba-like venom, but entangle it by means of small sticky hairs
cells,[10] and muscle cells in some species.[9] However, the on the thread.
number of middle-layer cells and types are much lower
than in sponges.[10] Ptychocysts are not used for prey capture instead
the threads of discharged ptychocysts are used for
building protective tubes in which their owners live.
2.4 Polymorphism Ptychocysts are found only in the order Ceriantharia,
tube anemones.[10]
Polymorphism refers to the occurrence of structurally and
functionally more than two dierent types of individu-
[9][10]
als within the same organism. It is a characteristic fea- The main components of a cnidocyte are:
ture of Cnidarians, particularly the polyp and medusa
forms, or of zooids within colonial organisms like those in
Hydrozoa.[17] In Hydrozoans, colonial individuals arising
from individuals zooids will take on separate tasks.[18] For
example, in Obelia there are feeding individuals, the gas-
trozooids; the individuals capable of asexual reproduc-
tion only, the gonozooids, blastostyles and free-living or
sexually reproducing individuals, the medusae.

2.5 Cnidocytes
These nettle cells function as harpoons, since their
A hydra's nematocyst, before ring.
payloads remain connected to the bodies of the cells by trigger cilium[10]
[9][10]
threads. Three types of cnidocytes are known:

A cilium (ne hair) which projects above the surface

and acts as a trigger. Spirocysts do not have cilia.

A tough capsule, the cnida, which houses the thread,

its payload and a mixture of chemicals that may
include venom or adhesives or both. (cnida is
derived from the Greek word , which means
nettle[19] )

A tube-like extension of the wall of the cnida that

points into the cnida, like the nger of a rubber glove
pushed inwards. When a cnidocyte res, the nger
pops out. If the cell is a venomous nematocyte, the
nger"'s tip reveals a set of barbs that anchor it in
the prey.

The thread, which is an extension of the nger and

coils round it until the cnidocyte res. The thread is
usually hollow and delivers chemicals from the cnida
to the target.
Firing sequence of the cnida in a hydras nematocyst[10]
Operculum (lid) An operculum (lid) over the end of the cnida. The
Finger that turns inside out lid may be a single hinged ap or three aps arranged
/ / / Barbs like slices of pie.
Venom
Victims skin The cell body, which produces all the other parts.
Victims tissues
It is dicult to study the ring mechanisms of cnidocytes
Nematocysts inject venom into prey, and usually as these structures are small but very complex. At least
have barbs to keep them embedded in the victims. four hypotheses have been proposed:[9]
4 2 DESCRIPTION

Rapid contraction of bers round the cnida may in- Hydras and some sea anemones can move slowly over
crease its internal pressure. rocks and sea or stream beds by various means: creeping
like snails, crawling like inchworms, or by somersaulting.
The thread may be like a coiled spring that extends A few can swim clumsily by waggling their bases.[10]
rapidly when released.

In the case of Chironex (the sea wasp), chemical 2.7 Nervous system and senses
changes in the cnidas contents may cause them to
expand rapidly by polymerization. Cnidarians are generally thought to have no brains or even
central nervous systems. However, they do have integra-
Chemical changes in the liquid in the cnida make it tive areas of neural tissue that could be considered some
a much more concentrated solution, so that osmotic form of centralization. Most of their bodies are inner-
pressure forces water in very rapidly to dilute it. This vated by decentralized nerve nets that control their swim-
mechanism has been observed in nematocysts of the ming musculature and connect with sensory structures,
class Hydrozoa, sometimes producing pressures as though clade has slightly dierent structures.[20] These
high as 140 atmospheres, similar to that of scuba air sensory structures, usually called rhopalia, can generate
tanks, and fully extending the thread in as little as 2 signals in response to various types of stimuli such as
milliseconds (0.002 second).[10] light, pressure, and much more. Medusa usually have sev-
eral of them around the margin of the bell that work to-
Cnidocytes can only re once, and about 25% of a hydras gether to control the motor nerve net, that directly inner-
nematocysts are lost from its tentacles when capturing a vates the swimming muscles. Most Cnidarians also have
brine shrimp. Used cnidocytes have to be replaced, which a parallel system. In scyphozoans, this takes the form of
takes about 48 hours. To minimise wasteful ring, two a diuse nerve net, which has modulatory eects on the
types of stimulus are generally required to trigger cnido- nervous system.[21] As well as forming the signal cables
cytes: nearby sensory cells detect chemicals in the water, between sensory neurons and motoneurons, intermediate
and their cilia respond to contact. This combination pre- neurons in the nerve net can also form ganglia that act
vents them from ring at distant or non-living objects. as local coordination centers. Communication between
Groups of cnidocytes are usually connected by nerves nerve cells can occur by chemical synapses or gap junc-
and, if one res, the rest of the group requires a weaker tions in hydrozoans, though gap junctions are not present
minimum stimulus than the cells that re rst.[9][10] in all groups. Cnidarians have many of the same neuro-
transmitters as many animals, including chemicals such
as glutamate, GABA, and acetylcholine.[22]
2.6 Locomotion This structure ensures that the musculature is excited
rapidly and simultaneously, and can be directly stimulated
from any point on the body, and it also is better able to
recover after injury.[20][21]
Medusae and complex swimming colonies such as
siphonophores and chondrophores sense tilt and acceler-
ation by means of statocysts, chambers lined with hairs
which detect the movements of internal mineral grains
called statoliths. If the body tilts in the wrong direc-
tion, the animal rights itself by increasing the strength
of the swimming movements on the side that is too
low. Most species have ocelli (simple eyes), which
can detect sources of light. However the agile Box Jel-
lysh are unique among Medusae because they possess
four kinds of true eyes that have retinas, corneas and
lenses.[23] Although the eyes probably do not form im-
A swimming sea nettle known as the purple-striped jelly ages, Cubozoa can clearly distinguish the direction from
(Chrysaora colorata) which light is coming as well as negotiate around solid-
colored objects.[9][23]
Medusae swim by a form of jet propulsion: muscles,
especially inside the rim of the bell, squeeze water out
of the cavity inside the bell, and the springiness of the 2.8 Feeding and excretion
mesoglea powers the recovery stroke. Since the tissue
layers are very thin, they provide too little power to swim Cnidarians feed in several ways: predation, absorbing
against currents and just enough to control movement dissolved organic chemicals, ltering food particles out
within currents.[10] of the water, obtaining nutrients from symbiotic algae
2.10 Regeneration 5

within their cells, and parasitism. Most obtain the ma- the opposite problem, an excess of oxygen, which may
jority of their food from predation but some, includ- prove toxic. The animals produce large quantities of
ing the corals Hetroxenia and Leptogorgia, depend almost antioxidants to neutralize the excess oxygen.[9]
completely on their endosymbionts and on absorbing dis-
solved nutrients.[9] Cnidaria give their symbiotic algae
carbon dioxide, some nutrients, a place in the sun and
protection against predators.[10] 2.10 Regeneration
Predatory species use their cnidocytes to poison or entan-
gle prey, and those with venomous nematocysts may start All cnidarians can regenerate, allowing them to recover
digestion by injecting digestive enzymes. The smell from injury and to reproduce asexually. Medusae have
of uids from wounded prey makes the tentacles fold in- limited ability to regenerate, but polyps can do so from
wards and wipe the prey o into the mouth. In medusae small pieces or even collections of separated cells. This
the tentacles round the edge of the bell are often short and enables corals to recover even after apparently being de-
most of the prey capture is done by oral arms, which stroyed by predators.[9]
are extensions of the edge of the mouth and are often
frilled and sometimes branched to increase their surface
area. Medusae often trap prey or suspended food parti-
cles by swimming upwards, spreading their tentacles and
oral arms and then sinking. In species for which sus-
3 Reproduction
pended food particles are important, the tentacles and
oral arms often have rows of cilia whose beating creates
currents that ow towards the mouth, and some produce
nets of mucus to trap particles.[9] Their digestion is both
intra and extracellular.
Once the food is in the digestive cavity, gland cells in the
gastroderm release enzymes that reduce the prey to slurry,
usually within a few hours. This circulates through the di-
gestive cavity and, in colonial cnidarians, through the con-
necting tunnels, so that gastroderm cells can absorb the
nutrients. Absorption may take a few hours, and digestion
within the cells may take a few days. The circulation of
nutrients is driven by water currents produced by cilia in
the gastroderm or by muscular movements or both, so that
nutrients reach all parts of the digestive cavity.[10] Nutri-
ents reach the outer cell layer by diusion or, for animals
or zooids such as medusae which have thick mesogleas,
are transported by mobile cells in the mesoglea.[9]
Indigestible remains of prey are expelled through the 1
mouth. The main waste product of cells internal pro- 2
cesses is ammonia, which is removed by the external and 3
internal water currents.[10] 4
5
6
2.9 Respiration 7
8
There are no respiratory organs, and both cell layers ab- 9
sorb oxygen from and expel carbon dioxide into the sur- 10
rounding water. When the water in the digestive cav- 11
ity becomes stale it must be replaced, and nutrients that 12
have not been absorbed will be expelled with it. Some 13
Anthozoa have ciliated grooves on their tentacles, allow- 14
ing them to pump water out of and into the digestive cav- Life cycle of a jellysh:[9][10]
ity without opening the mouth. This improves respiration 13 Larva searches for site
after feeding and allows these animals, which use the cav- 48 Polyp grows
ity as a hydrostatic skeleton, to control the water pressure 911 Polyp strobilates
in the cavity without expelling undigested food.[9] 1214 Medusa grows
Cnidaria that carry photosynthetic symbionts may have
6 4 CLASSIFICATION

3.1 Sexual in cnidarians the depression forms at the end further

from the yolk (at the animal pole), while in bilaterians
it forms at the other end (vegetal pole).[10] The larvae,
Cnidarian sexual reproduction often involves a complex called planulae, swim or crawl by means of cilia.[9] They
life cycle with both polyp and medusa stages. For exam- are cigar-shaped but slightly broader at the front end,
ple, in Scyphozoa (jellysh) and Cubozoa (box jellies) a which is the aboral, vegetal-pole end and eventually at-
larva swims until it nds a good site, and then becomes taches to a substrate if the species has a polyp stage.[10]
a polyp. This grows normally but then absorbs its ten-
Anthozoan larvae either have large yolks or are ca-
tacles and splits horizontally into a series of disks that
pable of feeding on plankton, and some already have
become juvenile medusae, a process called strobilation.
endosymbiotic algae that help to feed them. Since
The juveniles swim o and slowly grow to maturity, while
the parents are immobile, these feeding capabilities ex-
the polyp re-grows and may continue strobilating peri-
tend the larvaes range and avoid overcrowding of sites.
odically. The adults have gonads in the gastroderm, and
Scyphozoan and hydrozoan larvae have little yolk and
these release ova and sperm into the water in the breeding
most lack endosymbiotic algae, and therefore have to set-
season.[9][10]
tle quickly and metamorphose into polyps. Instead, these
This phenomenon of succession of dierently organized species rely on their medusae to extend their ranges.[10]
generations (one asexually reproducing, sessile polyp, and
one sexually reproducing, free-swimming medusa or ses-
sile polyp)[24] is sometimes called Alternation of asex- 3.2 Asexual
ual and sexual phases or metagenesis, but should not be
confused with the alternation of generations as found All known cnidaria can reproduce asexually by var-
in plants, characterized by an alternation of a multicel- ious means, in addition to regenerating after being
lular spore-producing form and a multicellular gamete- fragmented. Hydrozoan polyps only bud, while the
producing form. medusae of some hydrozoans can divide down the mid-
dle. Scyphozoan polyps can both bud and split down the
Shortened forms of this life cycle are common, for exam-
middle. In addition to both of these methods, Anthozoa
ple some oceanic scyphozoans omit the polyp stage com-
can split horizontally just above the base. Asexual re-
pletely, and cubozoan polyps produce only one medusa.
production makes the daughter cnidarian a clone of the
Hydrozoa have a variety of life cycles. Some have no
adult.[9][10]
polyp stages and some (e.g. hydra) have no medusae. In
some species, the medusae remain attached to the polyp
and are responsible for sexual reproduction; in extreme
cases these reproductive zooids may not look much like 4 Classication
medusae. Meanwhile, life cycle reversal, in which polyps
are formed directly from medusae without the involve- Cnidarians were for a long time grouped with
ment of sexual reproduction process, was observed in Ctenophores in the phylum Coelenterata, but in-
both Hydrozoa (Turritopsis dohrnii[25] and Laodicea un- creasing awareness of their dierences caused them
dulata[26] ) and Scyphozoa (Aurelia sp.1[27] ). Anthozoa to be placed in separate phyla. Modern cnidarians are
have no medusa stage at all and the polyps are responsi- generally classied into four main classes:[9] sessile
ble for sexual reproduction.[9] Anthozoa (sea anemones, corals, sea pens); swimming
Spawning is generally driven by environmental factors Scyphozoa (jellysh) and Cubozoa (box jellies); and
Hydrozoa, a diverse group that includes all the freshwater
such as changes in the water temperature, and their re-
lease is triggered by lighting conditions such as sun- cnidarians as well as many marine forms, and has both
sessile members such as Hydra and colonial swimmers
rise, sunset or the phase of the moon. Many species of
Cnidaria may spawn simultaneously in the same location, such as the Portuguese Man o' War. Staurozoa have
so that there are too many ova and sperm for predators to recently been recognised as a class in their own right
eat more than a tiny percentage one famous example rather than a sub-group of Scyphozoa, and the parasitic
is the Great Barrier Reef, where at least 110 corals and a Myxozoa and Polypodiozoa are now recognized as highly
few non-cnidarian invertebrates produce enough gametes derived cnidarians[6]
rather than more closely related to the
to turn the water cloudy. These mass spawnings may pro- bilaterians.
duce hybrids, some of which can settle and form polyps, Stauromedusae, small sessile cnidarians with stalks and
but it is not known how long these can survive. In some no medusa stage, have traditionally been classied as
species the ova release chemicals that attract sperm of the members of the Scyphozoa, but recent research suggests
same species.[9] they should be regarded as a separate class, Staurozoa.[28]
The fertilized eggs develop into larvae by dividing until The Myxozoa, microscopic parasites, were rst classied
there are enough cells to form a hollow sphere (blastula) as protozoans,[29] but recently as heavily modied cnidar-
and then a depression forms at one end (gastrulation) ians, and more closely related to Hydrozoa and Scypho-
and eventually become the digestive cavity. However, zoa than to Anthozoa.[30] However other recent research
 7

suggests that Polypodium hydriforme, a parasite within the ciently. In addition, reefs provide complex and varied
egg cells of sturgeon, is closely related to the Myxozoa habitats that support a wide range of other organisms.[39]
and that both Polypodium and the Myxozoa are interme- Fringing reefs just below low-tide level also have a mutu-
diate between cnidarians and bilaterian animals.[31] ally benecial relationship with mangrove forests at high-
Some researchers classify the extinct conulariids as tide level and sea grass meadows in between: the reefs
cnidarians, while others propose that they form a com- protect the mangroves and seagrass from strong currents
pletely separate phylum.[32] and waves that would damage them or erode the sedi-
ments in which they are rooted, while the mangroves and
seagrass protect the coral from large inuxes of silt, fresh
water and pollutants. This additional level of variety in
5 Ecology the environment is benecial to many types of coral reef
animals, which for example may feed in the sea grass and
use the reefs for protection or breeding.[40]
Many cnidarians are limited to shallow waters because
they depend on endosymbiotic algae for much of their
nutrients. The life cycles of most have polyp stages,
which are limited to locations that oer stable substrates.
6 Evolutionary history
Nevertheless, major cnidarian groups contain species that
have escaped these limitations. Hydrozoans have a world-
wide range: some, such as Hydra, live in freshwater;
Obelia appears in the coastal waters of all the oceans; and
Liriope can form large shoals near the surface in mid-
ocean. Among anthozoans, a few scleractinian corals,
sea pens and sea fans live in deep, cold waters, and
some sea anemones inhabit polar seabeds while others
live near hydrothermal vents over 10 km (6.2 mi) be-
low sea-level. Reef-building corals are limited to tropi-
cal seas between 30N and 30S with a maximum depth
of 46 m (151 ft), temperatures between 20 C (68 F)
and 28 C (82 F) high salinity and low carbon diox-
ide levels. Stauromedusae, although usually classied as
jellysh, are stalked, sessile animals that live in cool to
Arctic waters.[33] Cnidarians range in size from Hydra,
520 mm (0.200.79 in) long,[34] to the Lions mane jel-
lysh, which may exceed 2 m (6.6 ft) in diameter and 75
m (246 ft) in length.[35] The fossil coral Cladocora from Pliocene rocks in Cyprus
Prey of cnidarians ranges from plankton to animals sev-
eral times larger than themselves.[33][36] Some cnidarians
are parasites, mainly on jellysh but a few are major pests 6.1 Fossil record
of sh.[33] Others obtain most of their nourishment from
endosymbiotic algae or dissolved nutrients.[9] Predators The earliest widely accepted animal fossils are rather
of cnidarians include: sea slugs, which can incorporate modern-looking cnidarians, possibly from around 580
nematocysts into their own bodies for self-defense;[37] million years ago, although fossils from the Doushantuo
starsh, notably the crown of thorns starsh, which can Formation can only be dated approximately.[41] The iden-
devastate corals;[33] buttery sh and parrot sh, which tication of some of these as embryos of animals has
eat corals;[38] and marine turtles, which eat jellysh.[35] been contested, but other fossils from these rocks strongly
Some sea anemones and jellysh have a symbiotic re- resemble tubes and other mineralized structures made
lationship with some sh; for example clown sh live by corals.[42] Their presence implies that the cnidarian
among the tentacles of sea anemones, and each partner and bilaterian lineages had already diverged.[43] Although
protects the other against predators.[33] the Ediacaran fossil Charnia used to be classied as a
Coral reefs form some of the worlds most productive jellysh or sea pen,[44] more recent study of growth pat-
ecosystems. Common coral reef cnidarians include both terns in Charnia and modern cnidarians has cast doubt on
Anthozoans (hard corals, octocorals, anemones) and Hy- this hypothesis,[45][46] leaving only the Canadian polyp,
drozoans (re corals, lace corals). The endosymbiotic al- Haootia, as the only bona-de cnidarian body fossil from
gae of many cnidarian species are very eective primary the Ediacaran. Few fossils of cnidarians without mineral-
producers, in other words converters of inorganic chem- ized skeletons are known from more recent rocks, except
icals into organic ones that other organisms can use, and in lagersttten that preserved soft-bodied animals.[47]
their coral hosts use these organic chemicals very e- A few mineralized fossils that resemble corals have been
8 7 INTERACTION WITH HUMANS

found in rocks from the Cambrian period, and corals di- from the tip of the polyp. The traditional grouping of
versied in the Early Ordovician.[47] These corals, which Scyphozoa included the Staurozoa, but morphology and
were wiped out in the Permian-Triassic extinction about molecular phylogenetics indicate that Staurozoa are more
251 million years ago,[47] did not dominate reef construc- closely related to Cubozoa (box jellies) than to other
tion since sponges and algae also played a major part.[48] Scyphozoa. Similarities in the double body walls of
During the Mesozoic era rudist bivalves were the main Staurozoa and the extinct Conulariida suggest that they
reef-builders, but they were wiped out in the Cretaceous are closely related. The position of Anthozoa nearest the
Paleogene extinction event 65 million years ago,[49] and beginning of the cnidarian family tree also implies that
since then the main reef-builders have been scleractinian Anthozoa are the cnidarians most closely related to Bila-
corals.[47] teria, and this is supported by the fact that Anthozoa and
Bilateria share some genes that determine the main axes
of the body.[3][56]
6.2 Family tree
However, in 2005 Katja Seipel and Volker Schmid sug-
gested that cnidarians and ctenophores are simplied de-
Further information: Phylogeny
scendants of triploblastic animals, since ctenophores and
the medusa stage of some cnidarians have striated mus-
Family tree of Cnidaria and the origins of cle, which in bilaterians arises from the mesoderm. They
animals[3][50][51][52][6] did not commit themselves on whether bilaterians evolved
It is dicult to reconstruct the early stages in the from early cnidarians or from the hypothesized triploblas-
[11]
evolutionary family tree of animals using only tic ancestors of cnidarians.
morphology (their shapes and structures), because the In molecular phylogenetics analyses from 2005 onwards,
large dierences between Porifera (sponges), Cnidaria important groups of developmental genes show the same
plus Ctenophora (comb jellies), Placozoa and Bilateria variety in cnidarians as in chordates.[57] In fact cnidari-
(all the more complex animals) make comparisons ans, and especially anthozoans (sea anemones and corals),
dicult. Hence reconstructions now rely largely or retain some genes that are present in bacteria, protists,
entirely on molecular phylogenetics, which groups plants and fungi but not in bilaterians.[58]
organisms according to similarities and dierences in
The mitochondrial genome in the medusozoan cnidari-
their biochemistry, usually in their DNA or RNA.[53]
ans, unlike those in other animals, is linear with frag-
It is now generally thought that the Calcarea (sponges mented genes.[59] The reason for this dierence is un-
with calcium carbonate spicules) are more closely re- known.
lated to Cnidaria, Ctenophora (comb jellies) and Bilateria
(all the more complex animals) than they are to the
other groups of sponges.[50][54][55] In 1866 it was pro-
posed that Cnidaria and Ctenophora were more closely 7 Interaction with humans
related to each other than to Bilateria and formed a group
called Coelenterata (hollow guts), because Cnidaria and
Ctenophora both rely on the ow of water in and out of
a single cavity for feeding, excretion and respiration. In
1881, it was proposed that Ctenophora and Bilateria were
more closely related to each other, since they shared fea-
tures that Cnidaria lack, for example muscles in the mid-
dle layer (mesoglea in Ctenophora, mesoderm in Bilate-
ria). However more recent analyses indicate that these
similarities are rather vague, and the current view, based
on molecular phylogenetics, is that Cnidaria and Bilate-
ria are more closely related to each other than either is
to Ctenophora. This grouping of Cnidaria and Bilateria
has been labelled "Planulozoa" because it suggests that
the earliest Bilateria were similar to the planula larvae of
Cnidaria.[3][51]
Within the Cnidaria, the Anthozoa (sea anemones and
corals) are regarded as the sister-group of the rest, which The dangerous sea wasp
suggests that the earliest cnidarians were sessile polyps Chironex eckeri
with no medusa stage. However, it is unclear how the
other groups acquired the medusa stage, since Hydrozoa Jellysh stings killed about 1,500 people in the 20th
form medusae by budding from the side of the polyp century,[60] and cubozoans are particularly dangerous.
while the other Medusozoa do so by splitting them o On the other hand, some large jellysh are considered

a delicacy in East and Southeast Asia. Coral reefs have
long been economically important as providers of sh-
ing grounds, protectors of shore buildings against cur-
rents and tides, and more recently as centers of tourism.
However, they are vulnerable to over-shing, mining for
construction materials, pollution, and damage caused by
tourism.
Beaches protected from tides and storms by coral reefs
are often the best places for housing in tropical countries.
Reefs are an important food source for low-technology
shing, both on the reefs themselves and in the adja-
cent seas.[61] However, despite their great productivity,
reefs are vulnerable to over-shing, because much of the
organic carbon they produce is exhaled as carbon diox-
ide by organisms at the middle levels of the food chain
and never reaches the larger species that are of interest to
shermen.[39] Tourism centered on reefs provides much
of the income of some tropical islands, attracting pho-
tographers, divers and sports shermen. However, hu-
man activities damage reefs in several ways: mining for
construction materials; pollution, including large inuxes
of fresh water from storm drains; commercial shing, in-
cluding the use of dynamite to stun sh and the capture of
young sh for aquariums; and tourist damage caused by
boat anchors and the cumulative eect of walking on the
reefs.[61] Coral, mainly from the Pacic Ocean has long
been used in jewellery, and demand rose sharply in the
1980s.[62]
The dangerous Carukia barnesi, one of the known species of box
jellysh which can cause Irukandji syndrome.
Some large jellysh species of the Rhizostomae order
are commonly consumed in Japan, Korea and Southeast
Asia.[63][64][65] In parts of the range, shing industry is re-
stricted to daylight hours and calm conditions in two short
seasons, from March to May and August to November.[65]
The commercial value of jellysh food products depends
on the skill with which they are prepared, and Jellysh [10] Ruppert, E.E.; Fox, R.S. & Barnes, R.D. (2004). Inverte-
Masters guard their trade secrets carefully. Jellysh is
very low in cholesterol and sugars, but cheap preparation
can introduce undesirable amounts of heavy metals.[66]
The sea wasp Chironex eckeri has been described as
the worlds most venomous jellysh and is held respon-
sible for 67 deaths, although it is dicult to identify the
9
animal as it is almost transparent. Most stingings by C.
eckeri cause only mild symptoms.[67] Seven other box
jellies can cause a set of symptoms called Irukandji syn-
drome,[68] which takes about 30 minutes to develop,[69]
and from a few hours to two weeks to disappear.[70] Hos-
pital treatment is usually required, and there have been a
few deaths.[68]
8 Notes
[1] Subphyla Anthozoa and Medusozoa based on The Tax-
onomicon Taxon: Phylum Cnidaria. Universal Taxo-
nomic Services. Archived from the original on 2007-09-
29. Retrieved 2007-07-10.
[2] Classes in Medusozoa based on The Taxonomicon
Taxon: Subphylum Medusozoa. Universal Taxonomic
Services. Retrieved 2009-01-26.
[3] Collins, A.G. (May 2002). Phylogeny of Medu-
sozoa and the Evolution of Cnidarian Life Cycles
(PDF). Journal of Evolutionary Biology. 15 (3): 418
432. doi:10.1046/j.1420-9101.2002.00403.x. Retrieved
2008-11-27.
[4] Dictionary.com Unabridged. Random House, Inc.
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[5] Zhang, Z.-Q. (2011). Animal biodiversity: An introduc-
tion to higher-level classication and taxonomic richness
(PDF). Zootaxa. 3148: 712.
[6] E. Jmenez-Guri; et al. (July 2007). "Buddenbrockia
is a cnidarian worm. Science. 317 (116): 116118.
doi:10.1126/science.1142024. PMID 17615357.
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S, Church SH, et al. (2015). Phylogenomic
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[8] Park E, Hwang D, Lee J, Song J, Seo T, Won Y (2012).
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9 Further reading
9.1 Books
Arai, M.N. (1997). A Functional Biology of Scypho-
zoa. London: Chapman & Hall [p. 316]. ISBN
0-412-45110-7.
Ax, P. (1999). Das System der Metazoa I. Ein
Lehrbuch der phylogenetischen Systematik. Gustav
Fischer, Stuttgart-Jena: Gustav Fischer. ISBN 3-
437-30803-3.
Barnes, R.S.K., P. Calow, P. J. W. Olive, D. W.
Golding & J. I. Spicer (2001). The invertebratesa
synthesis. Oxford: Blackwell. 3rd edition [chapter
3.4.2, p. 54]. ISBN 0-632-04761-5.
Brusca, R.C., G.J. Brusca (2003). Invertebrates.
Sunderland, Mass.: Sinauer Associates. 2nd edition
[chapter 8, p. 219]. ISBN 0-87893-097-3.
Dalby, A. (2003). Food in the Ancient World: from
A to Z. London: Routledge.
Moore, J.(2001). An Introduction to the Inverte-
brates. Cambridge: Cambridge University Press
[chapter 4, p. 30]. ISBN 0-521-77914-6.
Schfer, W. (1997). Cnidaria, Nesseltiere. In
Rieger, W. (ed.) Spezielle Zoologie. Teil 1. Einzeller
und Wirbellose Tiere. Stuttgart-Jena: Gustav Fis-
cher. Spektrum Akademischer Verl., Heidelberg,
2004. ISBN 3-8274-1482-2.
10 EXTERNAL LINKS
Werner, B. 4. Stamm Cnidaria. In: V. Gruner
(ed.) Lehrbuch der speziellen Zoologie. Begr. von
Kaestner. 2 Bde. Stuttgart-Jena: Gustav Fischer,
Stuttgart-Jena. 1954, 1980, 1984, Spektrum Akad.
Verl., Heidelberg-Berlin, 1993. 5th edition. ISBN
3-334-60474-8.
9.2 Journal articles
D. Bridge, B. Schierwater, C. W. Cunningham, R.
DeSalle R, L. W. Buss: Mitochondrial DNA struc-
ture and the molecular phylogeny of recent cnidaria
classes. in: Proceedings of the Academy of Nat-
ural Sciences of Philadelphia. Philadelphia USA
89.1992, p. 8750. ISSN 0097-3157
D. Bridge, C. W. Cunningham, R. DeSalle, L.
W. Buss: Class-level relationships in the phylum
CnidariaMolecular and morphological evidence.
in: Molecular biology and evolution. Oxford Uni-
versity Press, Oxford 12.1995, p. 679. ISSN 0737-
4038
D. G. Fautin: Reproduction of Cnidaria. in: Cana-
dian Journal of Zoology. Ottawa Ont. 80.2002, p.
1735. (PDF, online) ISSN 0008-4301
G. O. Mackie: Whats new in cnidarian biology?
in: Canadian Journal of Zoology. Ottawa Ont.
80.2002, p. 1649. (PDF, online) ISSN 0008-4301
P. Schuchert: Phylogenetic analysis of the Cnidaria.
in: Zeitschrift fr zoologische Systematik und Evolu-
tionsforschung. Paray, Hamburg-Berlin 31.1993, p.
161. ISSN 0044-3808
G. Kass-Simon, A. A. Scappaticci Jr.: The behav-
ioral and developmental physiology of nematocysts.
in: Canadian Journal of Zoology. Ottawa Ont.
80.2002, p. 1772. (PDF, online) ISSN 0044-3808
J. Zrzav (2001). The interrelationships of meta-
zoan parasites: a review of phylum- and higher-level
hypotheses from recent morphological and molec-
ular phylogenetic analyses (PDF). Folia Parasito-
logica. 48 (2): 81103. doi:10.14411/fp.2001.013.
PMID 11437135. Archived from the original (PDF)
on 2007-10-25. Retrieved 2009-01-26.
10 External links
YouTube: Nematocysts Firing
YouTube:My Anemone Eat Meat Defensive and
feeding behaviour of sea anemone
Cnidaria - Guide to the Marine Zooplankton of
south eastern Australia, Tasmanian Aquaculture &
Fisheries Institute
 13

A Cnidaria homepage maintained by University of

California, Irvine
Cnidaria page at Tree of Life

Fossil Gallery: Cnidarians

The Hydrozoa Directory

Hexacorallians of the World

14 11 TEXT AND IMAGE SOURCES, CONTRIBUTORS, AND LICENSES

11 Text and image sources, contributors, and licenses

11.1 Text
Cnidaria Source: https://en.wikipedia.org/wiki/Cnidaria?oldid=775089483 Contributors: AxelBoldt, Mav, Malcolm Farmer, Josh Grosse,
PierreAbbat, William Avery, Azhyd, Heron, Zimriel, Hephaestos, Ubiquity, DopeshJustin, Dominus, Menchi, Tomi, Pcb21, Mdebets,
Ahoerstemeier, TUF-KAT, Angela, Glenn, Tristanb, Emperorbma, Andrewman327, Marshman, Itai, Taxman, Thue, Wiwaxia, Chl, Puz-
zletChung, Robbot, Altenmann, Romanm, Naddy, Chris Roy, Fenkys, Mirv, ShArky, Xpinki96x, Lisap, Hadal, UtherSRG, Tea2min, Marc
Venot, DocWatson42, Eran, Netoholic, Paul Pogonyshev, Michael Devore, Horatio, Chowbok, Gadum, Utcursch, Andycjp, Gdr, Knu-
tux, Williamb, Kaldari, DragonySixtyseven, Bumm13, Phil1988, Boojum, Gscshoyru, Ichigo20, Mdot2226, Neale Monks, Inniteim-
prob42, MattKingston, DanielCD, Discospinster, Rich Farmbrough, Vsmith, HeikoEvermann, Mani1, Joepearson, Dmr2, Bender235,
ESkog, Eric Forste, Kwamikagami, Mauler, Edward Z. Yang, Causa sui, Bobo192, Robotje, Smalljim, Arcadian, Ivansanchez, Van-
ished user 19794758563875, Jumbuck, Alansohn, Polarscribe, Lord Pistachio, Axl, B kimmel, Pippu d'Angelo, Aranae, BRW, Iustinus,
Adrian.benko, Stemonitis, Issk, Gmaxwell, 2004-12-29T22:45Z, LOL, PoccilScript, Sburke, Tylerni7, Prashanthns, JohnJohn, Rjwilmsi,
PinchasC, Quietust, DoubleBlue, FlaBot, Eubot, Margosbot~enwiki, Vandal B, RexNL, Shao, TheDJ, D.brodale, Alphachimp, Shark-
face217, Gdrbot, Dj Capricorn, Fenoxielo, Gwernol, YurikBot, Wavelength, Borgx, Wikipedia., RussBot, Spaully, Lovesick, Anoma-
locaris, K.C. Tang, NawlinWiki, IAMTHEEGGMAN, Wiki alf, Keithonearth, Aeusoes1, Apokryltaros, Peter Delmonte, Epipelagic,
Bota47, Derek.cashman, Nlu, Wknight94, Tigershrike, Mike Serfas, Lt-wiki-bot, Donald Albury, BorgQueen, BridgetDS, JLaTondre,
Asterion, Samuel Blanning, Mejor Los Indios, Luk, Twilight Realm, Attilios, RupertMillard, SmackBot, Jokl, Tigerghost, Reedy, Knowl-
edgeOfSelf, Bomac, KocjoBot~enwiki, Delldot, Timeforfun12345, Ga, Yamaguchi , Gilliam, Ohnoitsjamie, Skizzik, Kurykh, Ray-
lam0108, Hibernian, J. Spencer, Baa, Can't sleep, clown will eat me, Jere, Abyssal, Glloq, Rrburke, VMS Mosaic, Addshore, Sorchah,
BigBurkey, TheLimbicOne, Richard001, Paul H., Dvorak729, KeithB, Cephal-odd, Kukini, Thor Dockweiler, SashatoBot, JzG, Kuru,
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Falconus, JForget, CmdrObot, Tanthalas39, Causantin, Ricick, Lmcelhiney, TheAdventMaster, Maxxicum, Kupirijo, TheMarioManiac,
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pianist~enwiki, Thijs!bot, Epbr123, Isorhiza~enwiki, Kilva, Dogaroon, Sagaciousuk, N5iln, Mojo Hand, Headbomb, A3RO, James086,
Elhector, Michael A. White, Dawnseeker2000, AntiVandalBot, Mr Bungle, Seaphoto, Herald Alberich, Paste, Jj137, Danger, Bondolo,
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DrMicro, Thedjatclubrock, Blahah, A.Ou, Kurosa~enwiki, Lears Fool, Kyle the bot, Barneca, Cacolantern, Oshwah, Crustaceanguy,
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Insanity Incarnate, AlleborgoBot, Lyinginbedmon, Daveh4h, Steven Weston, The Random Editor, Tiddly Tom, Yintan, Keilana, Samak47,
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Kasmeister, Um, Materialscientist, The High Fin Sperm Whale, Citation bot, Xqbot, Maxcyber10, Friendoove, J04n, GrouchoBot,
Videovideo, Omnipaedista, RibotBOT, Doulos Christos, Penguinlord3, FrescoBot, Juwanna Mann, Renaynay329, Gouerouz, Finalius,
Drew R. Smith, Citation bot 1, Pinethicket, Jonesey95, Kristog, Serols, Kibi78704, Reconsider the static, Trappist the monk, 2ocean7,
Animalparty, Vrenator, RjwilmsiBot, TjBot, Bento00, Ripchip Bot, Mithril, Skamecrazy123, EmausBot, Ajraddatz, Clubeenoo, Cher-
mundy, Winner 42, Wikipelli, Dcirovic, Eternal.Pretzel, ZroBot, Jamie158100, rico, H3llBot, Korichi12, AManWithNoPlan, NGPriest,
Tolly4bolly, Lucasbrouwers, Brandmeister, Usb10, Mwwmenger, Lilacelder, ILKhanate, DASHBotAV, ClueBot NG, Gareth Grith-
Jones, Frerin, Mesoderm, Widr, Helpful Pixie Bot, Hurrderrderp, Titodutta, TruthWA, Bibcode Bot, Plantdrew, BG19bot, Wasbeer,
Acsmars, Silvrous, Glevum, Gug01, Patrick4124, Rhinopias, BattyBot, Jorg Ag, Mike1399is ninja, Mobydickforeal101, Ducknish, JYBot,
Dexbot, Mogism, Forgerz, Lugia2453, ComfyKem, Piaractus, Lord joel, Leafofprosperity, Epicgenius, Zorahia, LieutenantLatvia, Just-
Berry, Monkbot, Filedelinkerbot, Charles N. Gorichanaz, Septate, PotatoNinja, JAS Calvin Augustine, Alexinis10, Clara fcn, PaulTimmers,
Caftaric, BU Rob13, CodeBadger, Ishrat kaur, InternetArchiveBot, WikiCnidaria, Enoch1357, Af Brika1, Prahlad balaji, CCevol2016,
Ernestothomasta and Anonymous: 622

11.2 Images
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11.3 Content license 15

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