Brittle star

Brittle stars, serpent stars, or ophiuroids (from Latin

ophiurus  'brittle star'; from Ancient Greek ὄφις Brittle star
(óphis) 'serpent', and οὐρά (ourá) 'tail'; referring to the serpent- Temporal range: Ordovician to
like arms of the brittle star) are echinoderms in the class Present
Ophiuroidea, closely related to starfish. They crawl across the
sea floor using their flexible arms for locomotion. The
ophiuroids generally have five long, slender, whip-like arms
which may reach up to 60  cm (24  in) in length on the largest
specimens.

The Ophiuroidea contain two large clades, Ophiurida (brittle

stars) and Euryalida (basket stars). Over 2,000 species of brittle
stars live today.[1] More than 1,200 of these species are found in
deep waters, greater than 200 m deep.[1]
Common brittlestar (Ophiura ophiura)

Scientific classification
Contents Kingdom: Animalia

Range Phylum: Echinodermata

Anatomy Superclass: Asterozoa
Water-vascular system
Class: Ophiuroidea

Nervous system
Gray, 1840
Digestion
Respiration Type species
Musculoskeletal system Ophiura ophiura

Reproduction Linnaeus, 1758

Fission
Reorganized Orders
Life span
Regeneration Subclass Myophiuroidea
Locomotion Matsumoto, 1915

Bioluminescence
Superorder Euryophiurida
Ecology
O'Hara, Hugall, Thuy, Stöhr &
Parasites
Martynov, 2017
Diversity and taxonomy
Fossil record Euryalida Lamarck, 1816
Human relations Ophiurida Müller & Troschel,
Aquaria
1840 sensu O'Hara et al., 2017
References
Range Superorder Ophintegrida
O'Hara, Hugall, Thuy, Stöhr &
The ophiuroids diverged in the Early Ordovician, about 500 Martynov, 2017
million years ago.[2] Ophiuroids can be found today in all of the
major marine provinces, from the poles to the tropics.[3] Basket Amphilepidida O'Hara,
stars are usually confined to the deeper parts of this range; Hugall, Thuy, Stöhr &
Ophiuroids are known even from abyssal (>6,000 m) depths.[4]
Martynov, 2017
However, brittle stars are also common members of reef
communities, where they hide under rocks and even within Ophiacanthida O'Hara,
other living organisms. A few ophiuroid species can even Hugall, Thuy, Stöhr &
tolerate brackish water, an ability otherwise almost unknown
Martynov, 2017
among echinoderms.[5] A brittle star's skeleton is made up of
embedded ossicles. Ophioleucida O'Hara,
Hugall, Thuy, Stöhr &
Anatomy Martynov, 2017

Ophioscolecida O'Hara,
Of all echinoderms, the Ophiuroidea may have the strongest
tendency toward five-segment radial (pentaradial) symmetry. Hugall, Thuy, Stöhr &
The body outline is similar to that of starfish, in that ophiuroids Martynov, 2017
have five arms joined to a central body disk. However, in
ophiuroids, the central body disk is sharply marked off from the
arms.[6]

The disk contains all of the viscera. That is, the internal organs of
digestion and reproduction never enter the arms, as they do in the
Asteroidea. The underside of the disk contains the mouth, which
has five toothed jaws formed from skeletal plates. The madreporite
is usually located within one of the jaw plates, and not on the
upper side of the animal as it is in starfish.[6]
Brittle star in Kona, Hawaii
The ophiuroid coelom is strongly reduced, particularly in
comparison to other echinoderms.

Water-vascular system

The vessels of the water vascular system end in tube feet. The
water vascular system generally has one madreporite. Others, such
as certain Euryalina, have one per arm on the aboral surface. Still
other forms have no madreporite at all. Suckers and ampullae are Ophiopetra lithographica from the
absent from the tube feet. Lower Hienheim Beds (Lower
Tithonian, Upper Jurassic) near
Regensburg, Germany
Nervous system

The nervous system consists of a main nerve ring which runs around the central disk. At the base of
each arm, the ring attaches to a radial nerve which runs to the end of the limb. The nerves in each limb
run through a canal at the base of the vertebral ossicles.[6]
Most ophiuroids have no eyes, or other specialised sense organs.
However, they have several types of sensitive nerve endings in
their epidermis, and are able to sense chemicals in the water,
touch, and even the presence or absence of light.[6] Moreover, tube
feet may sense light as well as odors. These are especially found at
the ends of their arms, detecting light and retreating into crevices.

Digestion
Asteriacites, a trace fossil of an
The mouth is rimmed with five jaws, and serves as an anus ophiuroid; Carmel Formation
(egestion) as well as a mouth (ingestion). Behind the jaws is a (Middle Jurassic), near Gunlock,
short esophagus and a stomach cavity which occupies much of the Utah; scale bar is 10 mm.
dorsal half of the disk. Digestion occurs within 10 pouches or
infolds of the stomach, which are essentially ceca, but unlike in sea
stars, almost never extend into the arms.[6] The stomach wall
contains glandular hepatic cells.

Ophiuroids are generally scavengers or detritivores. Small organic

particles are moved into the mouth by the tube feet. Ophiuroids
may also prey on small crustaceans or worms. Basket stars in
particular may be capable of suspension feeding, using the mucus
coating on their arms to trap plankton and bacteria. They extend
one arm out and use the other four as anchors. Brittle stars will eat
small suspended organisms if available. In large, crowded areas, Green brittle star - Ophiarachna
brittle stars eat suspended matter from prevailing seafloor incrassata
currents.

In basket stars, the arms are used to sweep food rhythmically to the mouth. Pectinura consumes
beech pollen in the New Zealand fjords (since those trees hang over the water). Eurylina clings to
coral branches to browse on the polyps.

Respiration

Gas exchange and excretion occur through cilia-lined sacs called bursae; each opens between the arm
bases on the underside of the disk. Typically ten bursae are found, and each fits between two stomach
digestive pouches. Water flows through the bursae by means of cilia or muscular contraction. Oxygen
is transported through the body by the hemal system, a series of sinuses and vessels distinct from
the water vascular system.[6]

The bursae are probably also the main organs of excretion, with phagocytic "coelomocytes" collecting
waste products in the body cavity and then migrating to the bursae for expulsion from the body.[6]

Musculoskeletal system

Like all echinoderms, the Ophiuroidea possess a skeleton of calcium carbonate in the form of calcite.
In ophiuroids, the calcite ossicles are fused to form armor plates which are known collectively as the
test. The plates are covered by the epidermis, which consists of a smooth syncytium. In most species,
the joints between the ossicles and superficial plates allow the arm to bend to the side, but cannot
bend upwards. However, in the basket stars, the arms are flexible in all directions.[6]
Both the Ophiurida and Euryalida (the basket stars) have
five long, slender, flexible, whip-like arms, up to 60  cm in
length. They are supported by an internal skeleton of calcium
carbonate plates referred to as vertebral ossicles. These
"vertebrae" articulate by means of ball-and-socket joints, and
are controlled by muscles. They are essentially fused plates
which correspond to the parallel ambulacral plates in sea
stars and five Paleozoic families of ophiuroids. In modern
forms, the vertebrae occur along the median of the arm.

The ossicles are surrounded by a relatively thin ring of soft A field of the soft coral Callogorgia sp. with
tissue, and then by four series of jointed plates, one each on its brittle star symbionts
the upper, lower, and lateral surfaces of the arm. The two
lateral plates often have a number of elongated spines
projecting outwards; these help to provide traction against the substrate while the animal is moving.
The spines, in ophiuroids, compose a rigid border to the arm edges, whereas in euryalids they are
transformed into downward-facing clubs or hooklets. Euryalids are similar to ophiurids, if larger, but
their arms are forked and branched. Ophiuroid podia generally function as sensory organs. They are
not usually used for feeding, as in Asteroidea. In the Paleozoic era, brittle stars had open ambulacral
grooves, but in modern forms, these are turned inward.

In living ophiuroids, the vertebrae are linked by well-structured longitudinal muscles. Ophiuroida
move horizontally, and Euryalina species move vertically. The latter have bigger vertebrae and
smaller muscles. They are less spasmodic, but can coil their arms around objects, holding on even
after death. These movement patterns are distinct to the taxa, separating them. Ophiuroida move
quickly when disturbed. One arm presses ahead, whereas the other four act as two pairs of opposite
levers, thrusting the body in a series of rapid jerks. Although adults do not use their tube feet for
locomotion, very young stages use them as stilts and even serve as an adhesive structure.

Reproduction
The sexes are separate in most species, though a few are hermaphroditic or protandric. The gonads
are located in the disk, and open into pouches between the arms, called genital bursae. Fertilisation is
external in most species, with the gametes being shed into the surrounding water through the bursal
sacs.[6] An exception is the Ophiocanopidae, in which the gonads do not open into bursae and are
instead paired in a chain along the basal arm joints.

Many species brood developing larvae in the bursae, effectively giving birth to live young. A few, such
as Amphipholus squamata, are truly viviparous, with the embryo receiving nourishment from the
mother through the wall of the bursa. However, some species do not brood their young, and instead
have a free-swimming larval stage. Referred to as an ophiopluteus, these larvae have four pairs of
rigid arms lined with cilia. They develop directly into an adult, without the attachment stage found in
most starfish larvae.[6] The number of species exhibiting ophiopluteus larvae are fewer than those that
directly develop.

In a few species, the female carries a dwarf male, clinging to it with the mouth.[6]

Fission
Some brittle stars, such as the six-armed members of the family Ophiactidae, exhibit fissiparity
(division through fission), with the disk splitting in half. Regrowth of both the lost part of the disk and
the arms occur[7] which yields an animal with three large arms and three small arms during the period
of growth.

The West Indian brittle star, Ophiocomella ophiactoides, frequently undergoes asexual reproduction
by fission of the disk with subsequent regeneration of the arms. In both summer and winter, large
numbers of individuals with three long arms and three short arms can be found. Other individuals
have half a disk and only three arms. A study of the age range of the population indicates little
recruitment and fission is the primary means of reproduction in this species.[8]

In this species, fission appears to start with the softening of one side of the disk and the initiation of a
furrow. This deepens and widens until it extends across the disk and the animal splits in two. New
arms begin to grow before the fission is complete, thus minimizing the time between possible
successive divisions. The plane of fission varies so that some newly formed individuals have existing
arms of different lengths. The time period between successive divisions is 89 days, so theoretically,
each brittle star can produce 15 new individuals during the course of a year.[8]

Life span

Brittle stars generally sexually mature in two to three years, become full grown in three to four years,
and live up to five years.[9] Members of Euryalina, such as Gorgonocephalus, may live much longer.

Regeneration

Ophiuroids can readily regenerate lost arms or arm segments

unless all arms are lost. Ophiuroids use this ability to escape
predators, in a way similar to lizards which deliberately shed the
distal part of their tails to confuse pursuers. Moreover, the
Amphiuridae can regenerate gut and gonad fragments lost along
with the arms. Discarded arms have not been shown to have the
ability to regenerate.

Locomotion An Ophiothrix fragilis brittle star with

missing arm segments from Póvoa
Brittle stars use their arms for locomotion. Brittle stars move fairly de Varzim, Portugal
rapidly by wriggling their arms which are highly flexible and
enable the animals to make either snake-like or rowing
movements. However, they tend to attach themselves to the sea floor or to sponges or cnidarians, such
as coral. They move as if they were bilaterally symmetrical, with an arbitrary leg selected as the
symmetry axis and the other four used in propulsion. The axial leg may be facing or trailing the
direction of motion, and due to the radially symmetrical nervous system, can be changed whenever a
change in direction is necessary.[10]

Bioluminescence
Over 60 species of brittle stars are known to be bioluminescent.[11]
Most of these produce light in the green wavelengths, although a
few blue-emitting species have also been discovered. Both shallow-
water and deep-sea species of brittle stars are known to produce
light. Presumably, this light is used to deter predators.

Ecology
Brittle stars live in areas from the low-tide level downwards. Six
families live at least 2 m deep; the genera Ophiura,
Micro brittle starfish and Caulerpa
Amphiophiura, and Ophiacantha range below 4 m. Shallow
racemosa
species live among sponges, stones, or coral, or under the sand or
mud, with only their arms protruding. Two of the best-known
shallow species are the green brittle star (Ophioderma
brevispina), found from Massachusetts to Brazil, and the common
European brittle star (Ophiothrix fragilis). Deep-water species
tend to live in or on the sea floor or adhere to coral, urchins, or
xenophyophores. The most widespread species is the long-armed
brittle star (Amphipholis squamata), a grayish or bluish, strongly
luminescent species.
0:31

Parasites Brittle star in motion

The main parasite to enter the digestive tract or genitals are

protozoans. Crustaceans, nematodes, trematodes, and polychaete annelids also serve as parasites.
Algal parasites such as Coccomyxa ophiurae cause spinal malformation. Unlike in sea stars and sea
urchins, annelids are not typical parasites.

Diversity and taxonomy

Between 2,064[12] and 2,122 species of brittle stars are currently known,
but the total number of modern species may be over 3,000.[13] This
makes brittle stars the most abundant group of current echinoderms
(before sea stars). Around 270 genera are known, these are distributed in
16 families,[1] which makes them at the same time a relatively poorly
diversified group structurally, compared with the other echinoderms.[12]
For example, 467 species belong to the sole family of Amphiuridae (frail
brittle stars which live buried in the sediment leaving only their arms in
the stream to capture the plankton). There are also 344 species in the
family of Ophiuridae.[12]

List of families according to the World Register of Marine Species,

following O'Hara 2017:
Plate of brittle stars from
the Kunstformen der Natur
subclass Myophiuroidea Matsumoto, 1915
from Ernst Haeckel (1904)
infra-class Metophiurida Matsumoto, 1913
super-order Euryophiurida O'Hara, Hugall, Thuy, Stöhr & Martynov, 2017
 order Euryalida Lamarck, 1816
family Asteronychidae Ljungman, 1867
family Euryalidae Gray, 1840
family Gorgonocephalidae Ljungman, 1867
order Ophiurida Müller & Troschel, 1840 sensu O'Hara et al., 2017
suborder Ophiomusina O'Hara et al., 2017
family Ophiomusaidae (O'Hara, Stöhr, Hugall, Thuy,
Martynov, 2018)
family Ophiosphalmidae (O'Hara, Stöhr, Hugall, Thuy &
Martynov, 2018)
Ophiomusina incertae sedis
Second plate of brittle stars
suborder Ophiurina Müller & Troschel, 1840 sensu O'Hara et
from Haeckel
al., 2017

family Astrophiuridae Sladen, 1879

family Ophiopyrgidae Perrier, 1893
family Ophiuridae Müller & Troschel, 1840
Ophiurina incertae sedis
Ophiurida incertae sedis
super-order Ophintegrida O'Hara, Hugall, Thuy, Stöhr &
Martynov, 2017

order Amphilepidida O'Hara, Hugall, Thuy, Stöhr &

Martynov, 2017

suborder Gnathophiurina Matsumoto, 1915

super-family Amphiuroidea Ljungman, 1867
family Amphiuridae Ljungman, 1867
1852 drawing of ophioderma
family Amphilepididae Matsumoto, 1915
super-family Ophiactoidea Ljungman, 1867
family Ophiactidae Matsumoto, 1915
family Ophiopholidae O'Hara, Stöhr, Hugall, Thuy & Martynov, 2018
family Ophiothamnidae O'Hara, Stöhr, Hugall, Thuy & Martynov, 2018
family Ophiotrichidae Ljungman, 1867
suborder Ophionereidina O'Hara, Hugall, Thuy, Stöhr & Martynov, 2017
super-family Ophiolepidoidea Ljungman, 1867
family Hemieuryalidae Verrill, 1899
family Ophiolepididae Ljungman, 1867 (restricted)
super-family Ophionereidoidea Ljungman, 1867
family Amphilimnidae O'Hara, Stöhr, Hugall, Thuy & Martynov, 2018
family Ophionereididae Ljungman, 1867
suborder Ophiopsilina Matsumoto, 1915
 super-family Ophiopsiloidea Matsumoto, 1915
family Ophiopsilidae Matsumoto, 1915
order Ophiacanthida O'Hara, Hugall, Thuy, Stöhr & Martynov, 2017
suborder Ophiacanthina O'Hara, Hugall, Thuy, Stöhr & Martynov, 2017
family Clarkcomidae O'Hara, Stöhr, Hugall, Thuy & Martynov, 2018
family Ophiacanthidae Ljungman, 1867
family Ophiobyrsidae Matsumoto, 1915
family Ophiocamacidae (O'Hara, Stöhr, Hugall, Thuy, Martynov, 2018)
family Ophiopteridae O'Hara, Stöhr, Hugall, Thuy & Martynov, 2018
family Ophiotomidae Paterson, 1985
family Ophiojuridae O'Hara, Thuy & Hugall, 2021
suborder Ophiodermatina Ljungman, 1867
super-family Ophiocomoidea Ljungman, 1867
family Ophiocomidae Ljungman, 1867
super-family Ophiodermatoidea Ljungman, 1867
family Ophiodermatidae Ljungman, 1867
family Ophiomyxidae Ljungman, 1867
family Ophiopezidae O'Hara, Stöhr, Hugall, Thuy & Martynov, 2018
Ophiacanthida incertae sedis
order Ophioleucida O'Hara, Hugall, Thuy, Stöhr & Martynov, 2017
family Ophiernidae O'Hara, Stöhr, Hugall, Thuy & Martynov, 2018
family Ophioleucidae Matsumoto, 1915
order Ophioscolecida O'Hara, Hugall, Thuy, Stöhr & Martynov, 2017
family Ophiohelidae Perrier, 1893
family Ophioscolecidae Lütken, 1869
Ophiuroidea incertae sedis


Asteronyx loveni, Une Squamophis Euryale aspera, an

an Asteroschematidae albozosteres, an Euryalidae
Asteronychidae Astrocharidae

Astrocladus Ophiomusa Ophiomusium Ophiomisidium

euryale, a acufera, an lymani, an crosnieri, an
Gorgonocephalida Ophiomusaidae Ophiosphalmidae Astrophiuridae
e

Spinophiura Ophiura Amphioplus Ophiactis savignyi, Ophiocoma pica,

jolliveti, an ophiura, an thrombodes, an Ophiactidae an Ophiocomidae
Ophiopyrgidae Ophiuridae an
Amphiuridae

Ophionereis Ophiothri Clarkcoma Ophiacantha Ophiocamax

reticulata, an x canaliculata, an alternata, an fasciculata, an
Ophionereididae suensonii Clarkcomidae Ophiacanthidae Ophiocamacidae
, an
Ophiotric
hidae


Ophiopteris Ophiocomina Ophioderma Ophioplocus

antipodum, an nigra, an brevispina, an bispinosa, an
Ophiopteridae Ophiotomidae Ophiodermatidae Hemieuryalidae

Ophiolepis Ophiomyxa australis, an Ophiopeza cf. fallax, an Ophiernus adspersus,

superba, an Ophiomyxidae Ophiopezidae an Ophiernidae
Ophiolepididae

Ophioleuce Ophiomyces
seminudum, an frutectosus, an
Ophioleucidae Ophiohelidae

Fossil record

The first known brittle stars date from Early Ordovician.[14] Study of past distribution and evolution of
brittle stars has been hampered by the tendency of dead brittle stars to disarticulate and scatter,
providing poor brittle star fossils.[14] Until discoveries in the Agrio Formation of Neuquén Basin in the
2010s no fossil brittle star was known in the Southern Hemisphere, nor was any brittle star of
Cretaceous age known.[14]

Human relations
Brittle stars are not used as food, though they are not toxic, because of their strong skeleton.
Even if some species have blunt spines, no brittlestar is known to
be dangerous, nor venomous. There is no harm evidence towards
humans, and even with their predators, brittlestars' only means of
defense is escaping or discarding an arm.

Aquaria

Brittle stars are a moderately popular invertebrate in fishkeeping.

They can easily thrive in marine tanks; in fact, the micro brittle Tropical black brittlestar
star is a common "hitchhiker" that will propagate and become (Ophiocoma erinaceus) held in hand
common in almost any saltwater tank, if one happens to come at Reunion island.
along on some live rock.

Larger brittle stars are popular because, unlike Asteroidea, they

are not generally seen as a threat to coral, and are also faster-moving and more active than their more
archetypical cousins.

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