Comatulid
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Comatulida is an
order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...
of
crinoids Crinoids are marine invertebrates that make up the class Crinoidea. Crinoids that remain attached to the sea floor by a stalk in their adult form are commonly called sea lilies, while the unstalked forms, called feather stars or comatulids, are ...
. Members of this order are known as feather stars and mostly do not have a stalk as adults. The oral surface with the mouth is facing upwards and is surrounded by five, often divided rays with feathery pinnules. Comatulids live on the seabed and on reefs in tropical and temperate waters.


Taxonomy

Bourgueticrinida, the sea lilies, has traditionally been viewed as an order of Articulata and a sister
taxon In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
to Comatulida. A study published in 2011 suggested that it should be renamed Bourgueticrinina and viewed as a suborder of Comatulida.


Characteristics

Like other echinoderms, comatulids have
pentamer A pentamer is an entity composed of five subunits. In chemistry, it applies to molecules made of five monomers. In biochemistry, it applies to macromolecules, particularly pentameric proteins, made of five protein sub-units. In microbiology, a ...
ous symmetry (five sided) as adults though the larvae have
bilateral symmetry Symmetry in biology refers to the symmetry observed in organisms, including plants, animals, fungi, and bacteria. External symmetry can be easily seen by just looking at an organism. For example, the face of a human being has a plane of symme ...
. Late in their development, the larvae are attached to the seabed by a stalk, but this is broken at
metamorphosis Metamorphosis is a biological process by which an animal physically develops including birth transformation or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and different ...
and the juvenile crinoid is free living. The body has an endoskeleton made from a number of articulated calcareous plates known as ossicles covered by a thin epidermis. It is in the shape of a cup (the calyx) with a lid (the tegmen) which has a central mouth and an anus near the edge, the gut being U-shaped. There is a ring of clawlike appendages (the cirri) near the base of the aboral underside; these grip the
substrate Substrate may refer to: Physical layers *Substrate (biology), the natural environment in which an organism lives, or the surface or medium on which an organism grows or is attached ** Substrate (aquatic environment), the earthy material that exi ...
to keep the feather star in place. There are five long, often branched, rays attached round the edge of the tegmen. Each of these is further subdivided into
branchlet A branch, also called a ramus in botany, is a stem that grows off from another stem, or when structures like veins in leaves are divided into smaller veins. History and etymology In Old English, there are numerous words for branch, includ ...
s (the pinnules). Most comatulids originally have 10 arms, each ray being subdivided once. The arms are fragile, and if one is broken off, at least two grow in its place; in this way the number of arms can increase. The arms are composed of articulating ossicles held together by ligaments, and the pinnules have a similar structure. The arms are very flexible and can be spread widely or coiled up. An ambulacral groove starts on each pinnule and joins with others to form grooves on the arms all leading to grooves on the tegmen ending at the mouth. These food-collecting grooves are overhung by calcareous plates (the lappets) and have a lining of fine cilia.


Behavior

Many comatulids live in crevices, under corals or inside sponges, the only visible part being some of the arms. Some come out at night and perch themselves on eminences to feed. Many species can locomote across the seabed, raising their body on their arms. Many can also swim with their arms but most are largely sedentary, seldom moving far from their chosen place of concealment.


Feeding

Comatulids are
suspension feeder Filter feeders are aquatic animals that acquire nutrients by feeding on organic matters, food particles or smaller organisms (bacteria, microalgae and zooplanktons) suspended in water, typically by having the water pass over or through a spec ...
s. The arms are extended and held in such a position as to maximise the feeding surface with regard to the current. At each junction of the ossicles in the pinnules there are a group of three suckerless
tube feet Tube or tubes may refer to: * Tube (2003 film), ''Tube'' (2003 film), a 2003 Korean film * "Tubes" (Peter Dale), performer on the Soccer AM#Tubes, Soccer AM television show * Tube (band), a Japanese rock band * Tube & Berger, the alias of dance/e ...
. The longest of these searches for
plankton Plankton are the diverse collection of organisms that drift in Hydrosphere, water (or atmosphere, air) but are unable to actively propel themselves against ocean current, currents (or wind). The individual organisms constituting plankton are ca ...
in the surrounding water. When a particle is found, it is gathered in and thrust into the ambulacral groove by all three tube feet. Here it is formed into a bolus with mucus and moved down to the mouth by the actions of the cilia, being retained in the groove by the lappets.


Reproduction

Comatulids are
dioecious Dioecy ( ; ; adj. dioecious, ) is a characteristic of certain species that have distinct unisexual individuals, each producing either male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproduction is ...
, each individual being either male or female. The
gamete A gamete ( ) is a Ploidy#Haploid and monoploid, haploid cell that fuses with another haploid cell during fertilization in organisms that Sexual reproduction, reproduce sexually. Gametes are an organism's reproductive cells, also referred to as s ...
s are produced in specialised pinnules at the base of the arms, and fertilisation is external. The larvae are planktonic and drift with the water flow. After several larval stages they settle on the seabed and anchor themselves with a stalk. At metamorphosis, the stalk breaks and the juveniles can move around.


Ecology

Comatulids consist of 80%
calcium carbonate Calcium carbonate is a chemical compound with the chemical formula . It is a common substance found in Rock (geology), rocks as the minerals calcite and aragonite, most notably in chalk and limestone, eggshells, gastropod shells, shellfish skel ...
and are unappetising to most predators. A number of species of fish are known to feed on them, usually pulling off a single arm or the visceral mass, both of which can be regenerated. 47% of specimens seen by one researcher were lacking one or more arms or had regenerating limbs, so sub-lethal predation is probably low. Many other invertebrates live as
commensals Commensalism is a long-term biological interaction (symbiosis) in which members of one species gain benefits while those of the other species neither benefit nor are harmed. This is in contrast with mutualism, in which both organisms benefit fro ...
among the rays of crinoids and it may be these succulent morsels that are the principal objective of most predators. The comatulid '' Florometra serratissima'', in the north east Pacific, has been reported as being preyed on by the graceful
decorator crab Decorator crabs are crabs of several different species, belonging to the superfamily Majoidea (not all of which are decorators), that use materials from their environment to hide from, or ward off, predators. They decorate themselves by stickin ...
''
Oregonia gracilis ''Oregonia gracilis'', commonly known as the graceful decorator crab, is a species of crab belonging to the family Oregoniidae The Oregoniidae are a family of crabs, formerly included in the family Majidae. It contains the four genera ''Chiono ...
'' and the sunflower seastar ''
Pycnopodia helianthoides ''Pycnopodia helianthoides'', commonly known as the sunflower sea star, is a large starfish, sea star found in the northeastern Pacific Ocean. The only species of its genus, it is among the largest sea stars in the world, with a maximum arm span ...
''. The loss of the arms may be due to
autotomy Autotomy (from the Greek ''auto-'', "self-" and ''tome'', "severing", αὐτοτομία) or 'self-amputation', is the behaviour whereby an animal sheds or discards an appendage, usually as a self-defense mechanism to elude a predator's grasp ...
, the shedding of an arm to save the rest of the organism. A arm was found to be fully regenerated in nine months in this species.


Order Comatulida

The World Register of Marine Species includes the following suborders, superfamilies and families in Comatulida: *Suborder Bourgueticrinina **Family Atelecrinidae **Family Bathycrinidae **Family
Phrynocrinidae Phrynocrinidae is a family of echinoderms belonging to the order Comatulida Comatulida is an order of crinoids Crinoids are marine invertebrates that make up the class Crinoidea. Crinoids that remain attached to the sea floor by a stalk i ...
**Family Rhizocrinidae *Superfamily Antedonoidea **Family
Antedonidae Antedonidae is a Family (biology), family of crinoids or feather stars in the phylum Echinodermata. Members of the family are unstalked and have ten feathery arms. They can move about freely and have clawed Cirrus (biology), cirri to attach them ...
**Family
Pentametrocrinidae Pentametrocrinidae is a family of echinoderms belonging to the order Comatulida Comatulida is an order of crinoids Crinoids are marine invertebrates that make up the class Crinoidea. Crinoids that remain attached to the sea floor by a sta ...
**Family Zenometridae *Superfamily
Comatuloidea Comatulidae is a family of comatulid crinoids. Since 2015, it replaces the family Comasteridae. Description and characteristics This family is of recent restoration, and still has no consensual description. However the description of the fam ...
**Family
Comatulidae Comatulidae is a family of comatulid crinoids. Since 2015, it replaces the family Comasteridae. Description and characteristics This family is of recent restoration, and still has no consensual description. However the description of the fam ...
*Superfamily Himerometroidea **Family
Colobometridae ''Colobometridae'' is a family of crinoids belonging to the order Comatulida. Members of this order are known as feather stars. Genera Genera within this family include: * '' Alisometra'' A.H. Clark, 1947 * '' Analcidometra'' A.H. Clark, 1918 * ...
**Family
Himerometridae Himerometridae is a family of echinoderms belonging to the order Comatulida Comatulida is an order of crinoids Crinoids are marine invertebrates that make up the class Crinoidea. Crinoids that remain attached to the sea floor by a stalk i ...
**Family
Mariametridae Mariametridae is a family of echinoderms belonging to the order Comatulida Comatulida is an order of crinoids Crinoids are marine invertebrates that make up the class Crinoidea. Crinoids that remain attached to the sea floor by a stalk in ...
**Family
Zygometridae ''Zygometridae'' is a family of echinoderms belonging to the order Comatulida Comatulida is an order of crinoids Crinoids are marine invertebrates that make up the class Crinoidea. Crinoids that remain attached to the sea floor by a stalk ...
*Superfamily Notocrinoidea **Family
Aporometridae Aporometridae is a monotypic family of crinoids, the only genus being ''Aporometra'', which contains three species, all endemic to the seas around Australia. Description Members of this family have five arms which subdivide near the base givin ...
**Family Notocrinidae *Superfamily Tropiometroidea **Family Asterometridae **Family Calometridae **Family Charitometridae **Family Ptilometridae **Family Thalassometridae **Family Tropiometridae *Superfamily ''
incertae sedis or is a term used for a taxonomy (biology), taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...
'' within Comatulida **Family Atopocrinidae **Family Bourgueticrinidae **Family
Eudiocrinidae Eudiocrinidae is a family of echinoderms belonging to the order Comatulida Comatulida is an order of crinoids Crinoids are marine invertebrates that make up the class Crinoidea. Crinoids that remain attached to the sea floor by a stalk in ...
**Family Guillecrinidae **Family Septocrinidae


References


External links


Tree of Life Project: Crinoidea
{{Taxonbar, from=Q1028507 Articulata (Crinoidea) Echinoderm orders