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Crinoid
Crinoids are marine animals that make up the class Crinoidea, one of the
classes of the phylum Echinodermata, which also includes the starfish,
Crinoids
brittle stars, sea urchins and sea cucumbers.[3] Those crinoids which in their Temporal range: Darriwilian–recent[1]
adult form are attached to the sea bottom by a stalk are commonly called sea
lilies, while the unstalked forms are called feather stars or comatulids, PreЄ Є O S D C P T J K PgN
being members of the largest crinoid order Comatulida.

Adult crinoids are characterised by having the mouth located on the upper
surface. This is surrounded by feeding arms, and is linked to a U-shaped gut,
with the anus being located on the oral disc near the mouth. Although the
basic echinoderm pattern of fivefold symmetry can be recognised, in most
crinoids the five arms are subdivided into ten or more. These have feathery
pinnules and are spread wide to gather planktonic particles from the water.
At some stage in their life, most crinoids have a stem used to attach
themselves to the substrate, but many live attached only as juveniles and
become free-swimming as adults.

There are only about 600 living species of crinoid,[4] but the class was much
more abundant and diverse in the past. Some thick limestone beds dating to
the mid- to late-Paleozoic era are almost entirely made up of disarticulated
crinoid fragments.
Crinoid on the reef of Batu Moncho
Island, Indonesia

Scientific classification
Contents
Kingdom: Animalia
Etymology
Morphology
Phylum: Echinodermata
Biology Subphylum: Crinozoa
Feeding
Class: Crinoidea
Predation
Water vascular system Miller, 1821[2]
Nervous system Subclasses
Reproduction and life cycle
Locomotion Articulata (540 species)
†Flexibilia
Evolution
Origins †Camerata
Diversity †Disparida
Fossils
Taxonomy
Phylogeny
In culture
Fossil crinoids
References

Etymology
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The name "Crinoidea" comes from the Greek word κρίνος, "a lily", with the suffix –oid meaning "like".[5][6] They live in
both shallow water[7] and in depths as great as 9,000 meters (30,000 ft).[8] Those crinoids which in their adult form are
attached to the sea bottom by a stalk are commonly called sea lilies.[9] The unstalked forms are called feather stars[10] or
comatulids, being members of the largest crinoid order, Comatulida.[11]

Morphology
The basic body form of a crinoid is a stem (not present in adult
feather stars) and a crown consisting of a cup-like central body
known as the theca, and a set of five rays or arms, usually branched
and feathery. The mouth and anus are both located on the upper side
of the theca, making the dorsal (upper) surface the oral surface,
unlike in the other echinoderm groups such as the sea urchins,
starfish and brittle stars where the mouth is on the underside.[12]
The numerous calcareous plates make up the bulk of the crinoid,
with only a small percentage of soft tissue. These ossicles fossilise
well and there are beds of limestone dating from the Lower
Carboniferous around Clitheroe, England, formed almost exclusively
from a diverse fauna of crinoid fossils.[13]

Anatomy of a stalked crinoid

The stem of sea lilies is composed of a column of highly porous ossicles which
are connected by ligamentary tissue. It attaches to the substrate with a
flattened holdfast or with whorls of jointed, root-like structures known as cirri.
Further cirri may occur higher up the stem. In crinoids that attach to hard
surfaces, the cirri may be robust and curved, resembling birds' feet, but when
crinoids live on soft sediment, the cirri may be slender and rod-like. Juvenile
feather stars have a stem, but this is later lost, with many species retaining a
few cirri at the base of the crown. The majority of living crinoids are free-
swimming and have only a vestigial stalk. In those deep-sea species that still
Stalked crinoid drawn by Ernst retain a stalk, it may reach up to 1 m (3 ft) in length (although usually much
Haeckel smaller), and fossil species are known with 20 m (66 ft) stems.[14]

The theca is pentamerous (has five-part symmetry) and is homologous with the
body or disc of other echinoderms. The base of the theca is formed from a cup-shaped set of ossicles (bony plates), the
calyx, while the upper surface is formed by the weakly-calcified tegmen, a mebranous disc. The tegmen is divided into five
"ambulacral areas", including a deep groove from which the tube feet project, and five "interambulacral areas" between
them. The mouth is near the centre or on the margin of the tegmen, and ambulacral grooves lead from the base of the arms
to the mouth. The anus is also located on the tegmen, often on a small elevated cone, in an interambulacral area. The theca
is relatively small and contains the crinoid's digestive organs.[14]

The arms are supported by a series of articulating ossicles similar to those in the stalk. Primitively, crinoids had only five
arms, but in most modern forms these are divided into two at ossicle II, giving ten arms in total. In most living species,
especially the free-swimming feather stars, the arms branch several more times, producing up to two hundred branches in
total. Being jointed, the arms can curl up. They are lined, on either side alternately, by smaller jointed appendages known
as "pinnules" which give them their feather-like appearance. Both arms and pinnules have tube feet along the margins of

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the ambulacral grooves. The tube feet come in groups of three of different size; they have no suction pads and are used to
hold and manipulate food particles. The grooves are equipped with cilia which facilitate feeding by moving the organic
particles along the arm and into the mouth.[14]

Stem, theca and arms of a "true" Oxycomanthus bennetti Tegmen of a

(stalked) crinoid (family (comatulid) Lamprometra palmata.
Isselicrinidae) The mouth is located at
the center of the 5
feeding grooves, and
the anus at the top of
the column.

Close-up on the cirri that Close-up on the pinnules of a

allow comatulids to walk and Tropiometra carinata (with
attach themselves parasites Myzostoma
fuscomaculatum)

Biology

Feeding
Crinoids are passive suspension feeders, filtering plankton and small particles of
detritus from the sea water flowing past them with their feather-like arms. The
arms are raised to form a fan-shape which is held perpendicular to the current.
Mobile crinoids move to perch on rocks, coral heads or other eminences to
maximise their feeding opportunities. The food particles are caught by the primary
(longest) tube feet, which are fully extended and held erect from the pinnules,
forming a food-trapping mesh, while the secondary and tertiary tube feet are
involved in manipulating anything encountered.[14]

The tube feet are covered with sticky mucus that traps any particles which come in
contact. Once they have caught a particle of food, the tube feet flick it into the
ambulacral groove, where the cilia propel the mucus and food particles towards
the mouth. Lappets at the side of the groove help keep the mucus stream in place.
The total length of the food-trapping surface may be very large; the 56 arms of a
Two arms with pinnules and tube
Japanese sea lily with 24 cm (9 in) arms, have a total length of 80 m (260 ft)
feet outstretched

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including the pinnules. Generally speaking, crinoids living in environments with relatively little plankton have longer and
more highly branched arms than those living in food-rich environments.[14]

The mouth descends into a short oesophagus. There is no true stomach, so the oesophagus connects directly to the
intestine, which runs in a single loop right around the inside of the calyx. The intestine often includes numerous
diverticulae, some of which may be long or branched. The end of the intestine opens into a short muscular rectum. This
ascends towards the anus, which projects from a small conical protuberance at the edge of the tegmen. Faecal matter is
formed into large, mucous-cemented pellets which fall onto the tegmen and thence the substrate.[14]

Predation
Specimens of the sea urchin Calocidaris micans found in the vicinity of the crinoid Endoxocrinus parrae, have been
shown to contain large quantities of stem portions in their guts. These consist of articulated ossicles with soft tissue,
whereas the local sediment contained only disarticulated ossicles without soft tissue. This makes it highly likely that these
sea urchins are predators of the crinoids, and that the crinoids flee, offering part of their stem in the process.[15]

Various crinoid fossils hint at possible prehistoric predators. Coprolites of both fish and cephalopods have been found
containing ossicles of various crinoids, such as the pelagic crinoid Saccocoma, from the Jurassic lagerstatten
Solnhofen,[16] while damaged crinoid stems with bite marks matching the toothplates of coccosteid placoderms have been
found in Late Devonian Poland.[17] The calyxes of several Devonian to Carboniferous-aged crinoids have the shells of a
snail, Platyceras, intimately associated with them.[18] Some have the snail situated over the anus, suggesting that
Platyceras was a coprophagous commensal, while others have the animal directly situated over a borehole, suggesting a
more pernicious relationship.[19]

Water vascular system

Like other echinoderms, crinoids possess a water vascular system that maintains hydraulic pressure in the tube feet. This
is not connected to external sea water via a madreporite, as in other echinoderms, but only connected through a large
numbers of pores to the coelom (body cavity). The main fluid reservoir is the muscular-walled ring canal which is
connected to the coelom by stone canals lined with calcareous material. The coelom is divided into a number of
interconnecting spaces by mesenteries. It surrounds the viscera in the disc and has branches within the stalk and arms,
with smaller branches extending into the pinnules. It is the contraction of the ring canal that extends the tube feet. Three
narrow branches of the coelom enter each arm, two on the oral side and one aborally, and pinnules. The action of cilia
cause there to be a slow flow of fluid (1mm per second) in these canals, outward in the oral branches and inward in the
aboral ones, and this is the main means of transport of nutrients and waste products. There is no heart and separate
circulatory system but at the base of the disc there is a large blood vessel known as the axial organ, containing some
slender blind-ended tubes of unknown function, which extends into the stalk.[14]

These various fluid-filled spaces, in addition to transporting nutrients around the body, also function as both a respiratory
and an excretory system. Oxygen is absorbed primarily through the tube feet, which are the most thin-walled parts of the
body, with further gas exchange taking place over the large surface area of the arms. There are no specialised organs for
excretion while waste is collected by phagocytic coelomocytes.[14]

Nervous system
The crinoid nervous system is divided into three parts, with numerous connections between them. The oral or uppermost
portion is the only one homologous with the nervous systems of other echinoderms. It consists of a central nerve ring
surrounding the mouth, and radial nerves branching into the arms and is sensory in function. Below this lies an
intermediate nerve ring, giving off radial nerves supplying the arms and pinnules. These nerves are motor in nature, and
control the musculature of the tube feet. The third portion of the nervous system lies aborally, and is responsible for the
flexing and movement actions of the arms, pinnules and cirri. This is centred on a mass of neural tissue near the base of
the calyx, and provides a single nerve to each arm and a number of nerves to the stalk.[14]

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Reproduction and life cycle

Crinoids are not capable of clonal reproduction as are some starfish and brittle stars, but are capable of regenerating lost
body parts. Arms torn off by predators or damaged by adverse environmental conditions can regrow, and even the visceral
mass can regenerate over the course of a few weeks. This regeneration may be vital in surviving attacks by predatory
fish.[14]

Crinoids are dioecious, with individuals being either male or female. In most species, the gonads are located in the
pinnules but in a few, they are located in the arms. Not all the pinnules are reproductive, just those closest to the crown.
The gametes are produced in genital canals enclosed in genital coeloms. The pinnules eventually rupture to release the
sperm and eggs into the surrounding sea water. In certain genera, such as Antedon, the fertilised eggs are cemented to the
arms with secretions from epidermal glands; in others, especially cold water species from Antarctica, the eggs are brooded
in specialised sacs on the arms or pinnules.[14]

The fertilised eggs hatch to release free-swimming vitellaria larvae. The bilaterally symmetrical larva is barrel-shaped with
rings of cilia running round the body, and a tuft of sensory hairs at the upper pole. While both feeding and non-feeding
larvae exist among the four other extant echinoderm classes, all present day crinoids appear to be descendants from a
surviving clade that went through a bottleneck after the Permian extinction, at that time losing the feeding larval stage.[20]
The larva's free-swimming period lasts for only a few days before it settles on the bottom and attaches itself to the
underlying surface using an adhesive gland on its underside. The larva then undergoes an extended period of
metamorphoses into a stalked juvenile, becoming radially symmetric in the process. Even the free-swimming feather stars
go through this stage, with the adult eventually breaking away from the stalk.[14]

Locomotion
Most modern crinoids, i.e., the feather stars, are free-moving and lack a
stem as adults. Examples of fossil crinoids that have been interpreted as
free-swimming include Marsupitsa, Saccocoma and Uintacrinus.[21] In
general, crinoids move to new locations by crawling, using the cirri as
legs. Such a movement may be induced in relation to a change in current
direction, the need to climb to an elevated perch to feed, or because of
an agonistic behaviour by an encountered individual.[22] Crinoids can
also swim. They do this by co-ordinated, repeated sequential
A stalked crinoid (white) and a comatulid
movements of the arms in three groups. At first the direction of travel is
(red) in deep sea, showing the differences
upwards but soon becomes horizontal, travelling at about 7 cm (2.8 in)
between these two sister groups
per second with the oral surface in front. Swimming usually takes place
as short bursts of activity lasting up to half a minute, and in the
comatulid Florometra serratissima at least, only takes place after mechanical stimulation or as an escape response evoked
by a predator.[22]

In 2005, a stalked crinoid was recorded pulling itself along the sea floor off the Grand Bahama Island. While it has been
known that stalked crinoids could move, before this recording the fastest motion known for a stalked crinoid was 0.6
metres (2 feet) per hour. The 2005 recording showed one of these moving across the seabed at the much faster rate of 4 to
5 cm (1.6 to 2.0 in) per second (144 to 180 metres per hour).[23]

Evolution

Origins
If one ignores the enigmatic Echmatocrinus of the Burgess Shale, the earliest known unequivocal crinoid groups date back
to the Ordovician. There are two competing hypotheses pertaining to the origin of the group: the traditional viewpoint
holds that crinoids evolved from within the blastozoans (the eocrinoids and their derived descendants, the blastoids and

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the cystoids), whereas the most popular alternative suggests that the crinoids
split early from among the edrioasteroids.[24] The debate is difficult to settle, in
part because all three candidate ancestors share many characteristics,
including radial symmetry, calcareous plates, and stalked or direct attachment
to the substrate.[24]

Diversity
Echinoderms with mineralized skeletons entered the fossil record in the early
Cambrian (540 mya), and during the next 100 million years, the crinoids and
blastoids (also stalked filter-feeders) were dominant.[25] At that time, the
Echinodermata included twenty taxa of class rank, only five of which survived Agaricocrinus americanus, a fossil
the mass extinction events that followed. The long and varied geological history crinoid from the Carboniferous of
of the crinoids demonstrates how well the echinoderms had adapted to filter- Indiana
feeding.[3]

The crinoids underwent two periods of abrupt adaptive radiation, the first
during the Ordovician (485 to 444 mya), and the other during the early Triassic
(around 230 mya).[26] This Triassic radiation resulted in forms possessing
flexible arms becoming widespread; motility, predominantly a response to
predation pressure, also became far more prevalent than sessility.[27] This
radiation occurred somewhat earlier than the Mesozoic marine revolution,
possibly because it was mainly prompted by increases in benthic predation,
specifically of echinoids.[28] There then followed a selective mass extinction at
Middle Jurassic (Callovian)
the end of the Permian period, during which all blastoids and most crinoids Apiocrinites crinoid pluricolumnals
became extinct.[26] After the end-Permian extinction, crinoids never regained from the Matmor Formation in
the morphological diversity and dominant position they enjoyed in the southern Israel
Paleozoic; they employed a different suite of ecological strategies open to them
from those that had proven so successful in the Paleozoic.[26]

Fossils
Some fossil crinoids, such as Pentacrinites, seem to have lived attached to floating driftwood and complete colonies are
often found. Sometimes this driftwood would become waterlogged and sink to the bottom, taking the attached crinoids
with it. The stem of Pentacrinites can be several metres long. Modern relatives of Pentacrinites live in gentle currents
attached to rocks by the end of their stem. The largest fossil crinoid on record had a stem 40 m (130 ft) in length.[29]

In 2012, three geologists reported they had isolated complex organic molecules from 340-million-year-old (Mississippian)
fossils of multiple species of crinoids. Identified as "resembl[ing ...] aromatic or polyaromatic quinones", these are the
oldest molecules to be definitively associated with particular individual fossils, as they are believed to have been sealed
inside ossicle pores by precipitated calcite during the fossilization process.[30]

Taxonomy
Crinoidea has been accepted as a distinct clade of echinoderms since the definition of the group by Miller in 1821.[31]
According to the World Register of Marine Species, Articulata, the only extant subclass of Crinoidea, includes the following
families:-[32]

order Comatulida Clark, 1908

super-family Antedonoidea Norman, 1865

family Antedonidae Norman, 1865

family Pentametrocrinidae AH Clark, 1908

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family Zenometridae AH Clark, 1909

super-family Atelecrinoidea Bather, 1899

family Atelecrinidae Bather, 1899

super-family Comatuloidea Fleming, 1828

family Comatulidae Fleming, 1828

super-family Himerometroidea AH Clark, 1908

family Colobometridae AH Clark, 1909

family Eudiocrinidae AH Clark, 1907
family Himerometridae AH Clark, 1907
family Mariametridae AH Clark, 1909
family Zygometridae AH Clark, 1908
super-family Notocrinoidea Mortensen, 1918

family Aporometridae HL Clark, 1938

family Notocrinidae Mortensen, 1918
super-family Paracomatuloidea Hess, 1951 †
super-family Tropiometroidea AH Clark, 1908
Colorful crinoids in shallow waters in
family Asterometridae Gislén, 1924 Indonesia
family Calometridae AH Clark, 1911
family Charitometridae AH Clark, 1909
family Ptilometridae AH Clark, 1914
family Thalassometridae AH Clark, 1908
family Tropiometridae AH Clark, 1908
Comatulida incertae sedis

family Atopocrinidae Messing, 2011 (in Hess & Messing, 2011)

family Bathycrinidae Bather, 1899
family Bourgueticrinidae Loriol, 1882
family Guillecrinidae Mironov & Sorokina, 1998
family Phrynocrinidae AH Clark, 1907
family Septocrinidae Mironov, 2000
order Cyrtocrinida

Sub-order Cyrtocrinina

family Sclerocrinidae Jaekel, 1918

Sub-order Holopodina

family Eudesicrinidae Bather, 1899

family Holopodidae Zittel, 1879 Multiple crinoids on a reef in Indonesia
order Encrinida †
order Hyocrinida

family Hyocrinidae Carpenter, 1884

order Isocrinida

Sub-order Isocrinina

family Cainocrinidae Simms, 1988

family Isocrinidae Gislén, 1924
family Isselicrinidae Klikushkin, 1977
family Proisocrinidae Rasmussen, 1978
Sub-order Pentacrinitina †

family Pentacrinitidae Gray, 1842 †

order Millericrinida †

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Phylogeny
The phylogeny, geologic history, and classification of the Crinoidea was discussed by Wright et al. (2017).[33] These
authors presented new phylogeny-based and rank-based classifications based on results of recent phylogenetic
analyses.[31][34][35][36] Their rank-based classification of crinoid higher taxa (down to Order), not fully resolved and with
numerous groups incertae sedis (of uncertain placement), is illustrated in the cladogram.

Crinoidea   † Protocrinoidea (incertae sedis)

   
  † Diplobathrida
 

† Camerata † Eucamerata
      † Monobathrida
 

Pentacrinoidea Inadunata   Eustenocrinida

     
  Maennilicrinida
 
  Tetragonocrinida
 
Disparida   Calceocrinida
   
  ‘Homocrinida’ (incertae sedis)
 
  ‘Myelodactyla’ (incertae sedis)
 
  ‘Pisocrinoidea’ (incertae sedis)
 
Cladida   Porocrinida
 
Porocrinoidea  
    Hybocrinida
 
  Taxocrinida
Flexibilia  
    Sagenocrinida
 
Eucladida
    ‘Cyathocrinida’ (incertae
  sedis)

  ‘Dendrocrinida’ (incertae
  sedis)
Cyathoformes
 
  ‘Poteriocrinida’ (incertae
  sedis)

  † ‘Ampelocrinida’ (incertae
  sedis)
Articulata   Holocrinida
   
 
 
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† Encrinida

  Millericrinida
 

  Uintacrinida
 
  Roveacrinida
 

  Cyrtocrinida
 

  Hyocrinida
 

  Isocrinida
 

  Comatulida
 

In culture
Fossilised crinoid columnal segments extracted from limestone quarried on Lindisfarne, or found washed up along the
foreshore, were threaded into necklaces or rosaries, and became known as St. Cuthbert's beads in the Middle Ages.[37]
Similarly, in the Midwestern United States, fossilized segments of the columns of crinoids are sometimes known as Indian
beads.[38] Crinoids are the state fossil of Missouri.[39]

Fossil crinoids

Fossil from Germany showing the 330 million year old crinoids fossils Crinoid holdfasts and bryozoans
stem, calyx, and arms with from Iowa on an Upper Ordovician cobble
pinnules from northern Kentucky

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Seirocrinus subangularis from the Crinoid columnals (Isocrinus Root-like crinoid holdfast from the
Early Jurassic Posidonia Shale at nicoleti) from the Middle Jurassic Upper Ordovician, southern Ohio
Holzmaden, Germany Carmel Formation at Mount
Carmel Junction, Utah

Internal mold of crinoid stem Fossils of Seirocrinus

lumen (and external mold of stem) subsingularis from the Jurassic
from Lower Carboniferous, Ohio Holzmaden Black Shale
Formation, Germany

References
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106). Proceedings of the Royal Society B: Biological Sciences. 3: e1450. doi:10.7717/peerj.1450 (https://doi.org/10.77
17%2Fpeerj.1450). PMC 4675106 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4675106). PMID 26664800 (https://
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