Echinoderm is the common name given to any member of the phylum Echinodermata (from

Ancient Greek, ??????, echinos � "hedgehog" and d??�a, derma � "skin")[2] of marine
animals. The adults are recognizable by their (usually five-point) radial symmetry,
and include such well-known animals as sea stars, sea urchins, sand dollars, and
sea cucumbers, as well as the sea lilies or "stone lilies".[3] Echinoderms are
found at every ocean depth, from the intertidal zone to the abyssal zone. The
phylum contains about 7000 living species,[4] making it the second-largest grouping
of deuterostomes (a superphylum), after the chordates (which include the
vertebrates, such as birds, fishes, mammals, and reptiles). Echinoderms are also
the largest phylum that has no freshwater or terrestrial (land-based)
representatives.

Aside from the hard-to-classify Arkarua (a Precambrian animal with echinoderm-like

pentamerous radial symmetry), the first definitive members of the phylum appeared
near the start of the Cambrian. One group of Cambrian echinoderms, the cinctans
(Homalozoa), which are close to the base of the echinoderm origin, have been found
to possess external gills used for filter feeding, like chordata and hemichordata.
[5]

The echinoderms are important both ecologically and geologically. Ecologically,

there are few other groupings so abundant in the biotic desert of the deep sea, as
well as shallower oceans. Most echinoderms are able to reproduce asexually and
regenerate tissue, organs, and limbs; in some cases, they can undergo complete
regeneration from a single limb. Geologically, the value of echinoderms is in their
ossified skeletons, which are major contributors to many limestone formations, and
can provide valuable clues as to the geological environment. They were the most
used species in regenerative research in the 19th and 20th centuries. Further, it
is held by some scientists that the radiation of echinoderms was responsible for
the Mesozoic Marine Revolution.

Contents [hide]
1 Taxonomy and evolution
2 Anatomy and physiology 2.1 Skin and skeleton
2.2 The water vascular system
2.3 Other organs

3 Regeneration
4 Reproduction 4.1 Sexual reproduction
4.2 Asexual reproduction

5 Larval development
6 Distribution and habitat
7 Mode of life 7.1 Locomotion
7.2 Feeding
7.3 Defense mechanisms

8 Ecology
9 Use by humans
10 See also
11 References
12 Cited texts
13 Further reading
14 External links

Taxonomy and evolution[edit]

See also: List of echinodermata orders

Along with the chordates and hemichordates, echinoderms are deuterostomes, one of
the two major divisions of the bilaterians, the other being the protostomes. During
the early development of the embryo, in deuterostomes, the blastopore (the first
opening to form) becomes the anus whereas in the protostomes, it becomes the mouth.
In deuterostomes, the mouth develops at a later stage, at the opposite end of the
blastula from the blastopore, and a gut forms connecting the two.[6] The larvae of
echinoderms have bilateral symmetry but this is lost during metamorphosis when
their bodies are reorganised and develop the characteristic radial symmetry of the
echinoderm, typically pentamerism.[7] The characteristics of adult echinoderms are
the possession of a water vascular system with external tube feet and a calcareous
endoskeleton consisting of ossicles connected by a mesh of collagen fibres.[8] A
2014 analysis of 219 genes from all classes of echinoderms gives the following
phylogenetic tree.[9]

Bilateria

Xenacoelomorpha Proporus sp.png

Nephrozoa
Deuterostomia

Chordata and allies Cyprinus carpio3.jpg

Echinodermata

Echinozoa

Holothuroidea Holothuroidea.JPG

Echinoidea S. variolaris.jpg

Asterozoa

Ophiuroidea Ophiura ophiura.jpg

Asteroidea Portugal 20140812-DSC01434 (21371237591).jpg

Crinoidea Crinoid on the reef of Batu Moncho Island.JPG

Protostomia

Ecdysozoa Long nosed weevil edit.jpg

Spiralia Grapevinesnail 01.jpg

610 mya

650 mya

The Ordovician cystoid Echinosphaerites from northeastern Estonia

There are a total of about 7,000 extant species of echinoderm as well as about
13,000 extinct species.[8] They are found in habitats ranging from shallow
intertidal areas to abyssal depths. Two main subdivisions are traditionally
recognised: the more familiar motile Eleutherozoa, which encompasses the Asteroidea
(starfish, 1,745 recent species), Ophiuroidea (brittle stars, 2,300 species),
Echinoidea (sea urchins and sand dollars, 900 species) and Holothuroidea (sea
cucumbers, 1,430 species); and the Pelmatozoa, some of which are sessile while
others move around. These consist of the Crinoidea (feather stars and sea lilies,
580 species) and the extinct blastoids and Paracrinoids.[10] A fifth class of
Eleutherozoa consisting of just three species, the Concentricycloidea (sea
daisies), were recently merged into the Asteroidea.[11] The fossil record includes
a large number of other classes which do not appear to fall into any extant crown
group.

Fossil crinoid crowns

All echinoderms are marine and nearly all are benthic.[12] The oldest known
echinoderm fossil may be Arkarua from the Precambrian of Australia. It is a disc-
like fossil with radial ridges on the rim and a five-pointed central depression
marked with radial lines. However, no stereom or internal structure showing a water
vascular system is present and the identification is inconclusive.[13]

The first universally accepted echinoderms appear in the Lower Cambrian period,
asterozoans appeared in the Ordovician and the crinoids were a dominant group in
the Paleozoic.[12] Echinoderms left behind an extensive fossil record.[12] It is
hypothesised that the ancestor of all echinoderms was a simple, motile, bilaterally
symmetrical animal with a mouth, gut and anus. This ancestral stock adopted an
attached mode of life and suspension feeding, and developed radial symmetry as this
was more advantageous for such an existence. The larvae of all echinoderms are even
now bilaterally symmetrical and all develop radial symmetry at metamorphosis. The
starfish and crinoids still attach themselves to the seabed while changing to their
adult form.[14]

The first echinoderms later gave rise to free-moving groups. The evolution of
endoskeletal plates with stereom structure and of external ciliary grooves for
feeding were early echinoderm developments.[15] The Paleozoic echinoderms were
globular, attached to the substrate and were orientated with their oral surfaces
upwards. The fossil echinoderms had ambulacral grooves extending down the side of
the body, fringed on either side by brachioles, structures very similar to the
pinnules of a modern crinoid. It seems probable that the mouth-upward orientation
is the primitive state and that at some stage, all the classes of echinoderms
except the crinoids reversed this to become mouth-downward. Before this happened,
the podia probably had a feeding function as they do in the crinoids today. Their
locomotor function came later, after the re-orientation of the mouth when the podia
were in contact with the substrate for the first time.[14]

Anatomy and physiology[edit]

Echinoderms evolved from animals with bilateral symmetry. Although adult

echinoderms possess pentaradial, or five-sided, symmetry, echinoderm larvae are
ciliated, free-swimming organisms that organize in bilateral symmetry which makes
them look like embryonic chordates. Later, the left side of the body grows at the
expense of the right side, which is eventually absorbed. The left side then grows
in a pentaradially symmetric fashion, in which the body is arranged in five parts
around a central axis.[16] Within the Asterozoa, there can be a few exceptions from
the rule. The starfish genus Leptasterias normally have six arms, although five
armed individuals can occur. Also the Brisingida have six armed species. Amongst
the brittle stars, six armed species such as Ophiothela danae, Ophiactis savignyi
and Ophionotus hexactis exists, and Ophiacantha vivipara often has more than six.
[17]

Echinoderms exhibit secondary radial symmetry in portions of their body at some

stage of life. This, however, is an adaptation to their sessile existence. They
developed from other members of the Bilateria and exhibit bilateral symmetry in
their larval stage. Many crinoids and some seastars exhibit symmetry in multiples
of the basic five, with starfish such as Labidiaster annulatus known to possess up
to fifty arms, and the sea-lily Comaster schlegelii having two hundred.[18]
Skin and skeleton[edit]

A brittle star, Ophionereis reticulata

A sea cucumber from Malaysia

Starfish exhibit a wide range of colours

Strongylocentrotus purpuratus, a well-armoured sea urchin

Crinoid on a coral reef

Echinoderms have a mesodermal skeleton composed of calcareous plates or ossicles.
Each one of these, even the articulating spine of a sea urchin, is composed
mineralogically of a crystal of calcite. If solid, these would form a heavy
skeleton, so they have a sponge-like porous structure known as stereom.[19]
Ossicles may be fused together, as in the test of sea urchins, or may articulate
with each other as in the arms of sea stars, brittle stars and crinoids. The
ossicles may be flat plates or bear external projections in the form of spines,
granules or warts and they are supported by a tough epidermis (skin). Skeletal
elements are also deployed in some specialized ways, such as the "Aristotle's
lantern" mouthparts of sea urchins used for grinding, the supportive stalks of
crinoids and the structural "lime ring" of sea cucumbers.[16]

Despite the robustness of the individual skeletal modules complete skeletons of

starfish, brittle stars and crinoids are rare in the fossil record. This is because
they quickly disarticulate (disconnect from each other) once the encompassing skin
rots away, and in the absence of tissue there is nothing to hold the plates
together. The modular construction is a result of the growth system employed by
echinoderms, which adds new segments at the centre of the radial limbs, pushing the
existing plates outwards and lengthening the arms. Sea urchins on the other hand
are often well preserved in chalk beds or limestone. During fossilization, the
cavities in the stereom are filled in with calcite that is in crystalline
continuity with the surrounding material.