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Sea cucumbers are
echinoderm An echinoderm () is any member of the phylum Echinodermata (). The adults are recognisable by their (usually five-point) radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the sea ...
s from the
class Class or The Class may refer to: Common uses not otherwise categorized * Class (biology), a taxonomic rank * Class (knowledge representation), a collection of individuals or objects * Class (philosophy), an analytical concept used differentl ...
Holothuroidea (). They are
marine animal Marine life, sea life, or ocean life is the plants, animals and other organisms that live in the salt water of seas or oceans, or the brackish water of coastal estuaries. At a fundamental level, marine life affects the nature of the planet. M ...
s with a leathery skin and an elongated body containing a single, branched
gonad A gonad, sex gland, or reproductive gland is a mixed gland that produces the gametes and sex hormones of an organism. Female reproductive cells are egg cells, and male reproductive cells are sperm. The male gonad, the testicle, produces sper ...
. Sea cucumbers are found on the sea floor worldwide. The number of holothurian () species worldwide is about 1,717, with the greatest number being in the
Asia-Pacific Asia-Pacific (APAC) is the part of the world near the western Pacific Ocean. The Asia-Pacific region varies in area depending on context, but it generally includes East Asia, Russian Far East, South Asia, Southeast Asia, Australia and Pacific Isla ...
region. Many of these are gathered for human consumption and some species are cultivated in aquaculture systems. The harvested product is variously referred to as '' trepang'', ''namako'', ''bêche-de-mer'', or ''balate''. Sea cucumbers serve a useful role in the
marine ecosystem Marine ecosystems are the largest of Earth's aquatic ecosystems and exist in waters that have a high salt content. These systems contrast with freshwater ecosystems, which have a lower salt content. Marine waters cover more than 70% of the surf ...
as they help recycle nutrients, breaking down
detritus In biology, detritus () is dead particulate organic material, as distinguished from dissolved organic material. Detritus typically includes the bodies or fragments of bodies of dead organisms, and fecal material. Detritus typically hosts commun ...
and other
organic matter Organic matter, organic material, or natural organic matter refers to the large source of carbon-based compounds found within natural and engineered, terrestrial, and aquatic environments. It is matter composed of organic compounds that have c ...
, after which
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...
can continue the
decomposition Decomposition or rot is the process by which dead organic substances are broken down into simpler organic or inorganic matter such as carbon dioxide, water, simple sugars and mineral salts. The process is a part of the nutrient cycle and is e ...
process. Like all
echinoderm An echinoderm () is any member of the phylum Echinodermata (). The adults are recognisable by their (usually five-point) radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the sea ...
s, sea cucumbers have an
endoskeleton An endoskeleton (From Greek ἔνδον, éndon = "within", "inner" + σκελετός, skeletos = "skeleton") is an internal support structure of an animal, composed of mineralized tissue. Overview An endoskeleton is a skeleton that is on the ...
just below the skin, calcified structures that are usually reduced to isolated microscopic ossicles (or sclerietes) joined by connective tissue. In some species these can sometimes be enlarged to flattened plates, forming an armour. In
pelagic The pelagic zone consists of the water column of the open ocean, and can be further divided into regions by depth (as illustrated on the right). The word ''pelagic'' is derived . The pelagic zone can be thought of as an imaginary cylinder or wa ...
species such as ''
Pelagothuria natatrix ''Pelagothuria'' is a genus of sea cucumbers in the family Pelagothuriidae. It is monotypic, being represented by the single species ''Pelagothuria natatrix''. Characteristics This sea cucumber is somewhat unusual in appearance in comparison ...
'' (order
Elasipodida Elasipodida is an order of sea cucumbers. They have numerous appendages, including conical papillae and leaf-like tentacles. Although many species are benthic, a number are pelagic, and may have their appendages modified to form sails or fins. ...
, family
Pelagothuriidae Pelagothuriidae is a family of deep-sea swimming sea cucumbers. They are somewhat unusual in appearance, in comparison with other sea cucumbers, having numerous appendages, including conical papillae and leaf-like tentacles. Most of them are bent ...
), the skeleton is absent and there is no calcareous ring. Sea cucumbers are named for their resemblance to the fruit of the
cucumber Cucumber (''Cucumis sativus'') is a widely-cultivated Vine#Horticultural climbing plants, creeping vine plant in the Cucurbitaceae family that bears usually cylindrical Fruit, fruits, which are used as culinary vegetables.
plant.


Overview

Most sea cucumbers, as their name suggests, have a soft and cylindrical body, more or less lengthened, rounded off and occasionally fat in the extremities, and generally without solid appendages. Their shape ranges from almost spherical for "sea apples" (genus ''
Pseudocolochirus Sea apple is the common name for the colorful and somewhat round sea cucumbers of the genus ''Pseudocolochirus'', found in Indo-Pacific waters. Sea apples are filter feeders with tentacles, ovate bodies, and tube-like feet. As with many othe ...
'') to serpent-like for
Apodida Apodida is an order of littoral to deep-sea, largely infaunal holothurians, sea cucumbers. This order comprises three families, 32 genera and about 270 known species, called apodids, "without feet". Characteristics These sea cucumbers are vagile ...
or the classic sausage-shape, while others resemble caterpillars. The mouth is surrounded by tentacles, which can be pulled back inside the animal. Holothurians measure generally between 10 and 30 centimetres long, with extremes of some millimetres for '' Rhabdomolgus ruber'' and up to more than 3 metres for ''
Synapta maculata ''Synapta maculata'', the snake sea cucumber, is a species of sea cucumber in the family Synaptidae. It is found in shallow waters in the tropical Indo-Pacific Ocean. Sometimes growing as long as , it is one of the longest sea cucumbers in the w ...
''. The largest
American American(s) may refer to: * American, something of, from, or related to the United States of America, commonly known as the "United States" or "America" ** Americans, citizens and nationals of the United States of America ** American ancestry, pe ...
species, '' Holothuria floridana'', which abounds just below low-water mark on the
Florida Florida is a state located in the Southeastern region of the United States. Florida is bordered to the west by the Gulf of Mexico, to the northwest by Alabama, to the north by Georgia, to the east by the Bahamas and Atlantic Ocean, and to ...
reefs, has a volume of well over , and long. Most possess five rows of tube feet (called " podia"), but
Apodida Apodida is an order of littoral to deep-sea, largely infaunal holothurians, sea cucumbers. This order comprises three families, 32 genera and about 270 known species, called apodids, "without feet". Characteristics These sea cucumbers are vagile ...
lacks these and moves by crawling; the podia can be of smooth aspect or provided with fleshy appendages (like ''
Thelenota ananas ''Thelenota ananas'', also known as pineapple sea cucumber, oloturia ananas, tripang, prickly skin cucumber, pointed teat sea cucumber, armoured sea cucumber, giant sea cucumber, sand fish or prickly redfish, is a species of sea cucumber found i ...
''). The podia on the dorsal surface generally have no locomotive role, and are transformed into papillae. At one of the extremities opens a rounded mouth, generally surrounded with a crown of tentacles which can be very complex in some species (they are in fact modified podia); the anus is postero-dorsal. Holothurians do not look like other echinoderms at first glance, because of their tubular body, without visible skeleton nor hard appendixes. Furthermore, the fivefold symmetry, classical for echinoderms, although preserved structurally, is doubled here by a bilateral symmetry which makes them look like
chordate A chordate () is an animal of the phylum Chordata (). All chordates possess, at some point during their larval or adult stages, five synapomorphies, or primary physical characteristics, that distinguish them from all the other taxa. These fiv ...
s. However, a central symmetry is still visible in some species through five 'radii', which extend from the mouth to the anus (just like for sea urchins), on which the tube feet are attached. There is thus no "oral" or "aboral" face as for sea stars and other echinoderms, but the animal stands on one of its sides, and this face is called ''trivium'' (with three rows of tube feet), while the dorsal face is named ''bivium''. A remarkable feature of these animals is the "catch"
collagen Collagen () is the main structural protein in the extracellular matrix found in the body's various connective tissues. As the main component of connective tissue, it is the most abundant protein in mammals, making up from 25% to 35% of the whole ...
that forms their body wall."Catch" collagen has two states, soft and stiff, that are under neurological contro
Jose del Castillo and David S. Smith. (1996) "We Still Invoke Friction and Occam's Razor to Explain Catch in the Spines of ''Eucidaris Tribuloides''." ''Biological Bulletin'' 190:243-244
/ref> This can be loosened and tightened at will, and if the animal wants to squeeze through a small gap, it can essentially liquefy its body and pour into the space. To keep itself safe in these crevices and cracks, the sea cucumber will hook up all its collagen fibers to make its body firm again. The most common way to separate the subclasses is by looking at their oral tentacles. Order
Apodida Apodida is an order of littoral to deep-sea, largely infaunal holothurians, sea cucumbers. This order comprises three families, 32 genera and about 270 known species, called apodids, "without feet". Characteristics These sea cucumbers are vagile ...
have a slender and elongate body lacking tube feet, with up to 25 simple or
pinnate Pinnation (also called pennation) is the arrangement of feather-like or multi-divided features arising from both sides of a common axis. Pinnation occurs in biological morphology, in crystals, such as some forms of ice or metal crystals, and in ...
oral tentacles. Aspidochirotida are the most common sea cucumbers encountered, with a strong body and 10–30 leaflike or shield-like oral tentacles.
Dendrochirotida Dendrochirotida are an order of sea cucumbers. Members of this order have branched tentacles and are suspension feeders. Examples include '' Thyonella'' and ''Cucumaria''. Characteristics Holothurians in this order are characterised by ten to thi ...
are filter-feeders, with plump bodies and 8–30 branched oral tentacles (which can be extremely long and complex). Image:Apostichopus californicus.004 - Aquarium Finisterrae.jpg, Details of the mouth with its tentacles. Image:Synaptula lamberti.JPG, '' Synaptula lamperti'' lives on sponges (here in
Indonesia Indonesia, officially the Republic of Indonesia, is a country in Southeast Asia and Oceania between the Indian and Pacific oceans. It consists of over 17,000 islands, including Sumatra, Java, Sulawesi, and parts of Borneo and New Guine ...
). Image:Reef1118 - Flickr - NOAA Photo Library.jpg, ''
Synapta maculata ''Synapta maculata'', the snake sea cucumber, is a species of sea cucumber in the family Synaptidae. It is found in shallow waters in the tropical Indo-Pacific Ocean. Sometimes growing as long as , it is one of the longest sea cucumbers in the w ...
'', the longest known sea cucumber (
Apodida Apodida is an order of littoral to deep-sea, largely infaunal holothurians, sea cucumbers. This order comprises three families, 32 genera and about 270 known species, called apodids, "without feet". Characteristics These sea cucumbers are vagile ...
). Image:Thelenota anax géante.JPG, The king sea cucumber ('' Thelenota anax'', family
Stichopodidae The Stichopodidae are a family of sea cucumbers, part of the order Synallactida. Description Members of this family are mostly large or medium-sized holothuroids with a squarish cross-section, a flat ventral surface, and large, fleshy, cone-sha ...
) is one of the heaviest known holothurians. Image:Chiridota heheva.jpg, ''
Chiridota heheva ''Chiridota heheva'' is a species of sea cucumber in the family Chiridotidae. The species is found in the Western Atlantic Ocean, in deeper regions and cold seeps. It was described by Pawson and Vance in 2004. References Chiridotidae A ...
'', abyssal species. Image:Cucumaria big.jpg, '' Cucumaria miniata'', a filter-feeding sea cucumber. Image:Pseudocolochirus axiologus.jpg, ''
Pseudocolochirus Sea apple is the common name for the colorful and somewhat round sea cucumbers of the genus ''Pseudocolochirus'', found in Indo-Pacific waters. Sea apples are filter feeders with tentacles, ovate bodies, and tube-like feet. As with many othe ...
'' ("sea apple"). Image:Holothuria leucospilota.jpg, ''
Holothuria leucospilota ''Holothuria leucospilota'', commonly known as the black sea cucumber or black tarzan, is a species of marine invertebrate in the family Holothuriidae. It is placed in the subgenus ''Mertensiothuria'' making its full scientific name ''Holothuria ...
'' Image:Isostichopus badionotus.jpg, '' Isostichopus badionotus'' Image:Treasure chest candycane sea cucumber 2.jpg, '' Thelenota rubralineata'' Image:Holothuria fuscopunctata.jpg, '' Holothuria fuscopunctata'' Image:Bohadschia argus.jpg, '' Bohadschia argus''


Anatomy

Sea cucumbers are typically in length, although the smallest known species are just long, and the largest can reach . The body ranges from almost spherical to worm-like, and lacks the arms found in many other echinoderms, such as
starfish Starfish or sea stars are star-shaped echinoderms belonging to the class Asteroidea (). Common usage frequently finds these names being also applied to ophiuroids, which are correctly referred to as brittle stars or basket stars. Starfish ...
. The anterior end of the animal, containing the mouth, corresponds to the oral pole of other echinoderms (which, in most cases, is the underside), while the posterior end, containing the anus, corresponds to the aboral pole. Thus, compared with other echinoderms, sea cucumbers can be said to be lying on their side.


Body plan

The body of a holothurian is roughly cylindrical. It is radially symmetrical along its longitudinal axis, and has weak bilateral symmetry transversely with a dorsal and a ventral surface. As in other
Echinozoa Echinozoa is a subphylum of free-living echinoderms in which the body is or originally was a modified globe with meridional symmetry. Echinozoans lack arms, brachioles, or other appendages, and do not at any time exhibit pinnate structure. Their t ...
ns, there are five
ambulacra Ambulacrum is an architectural word that denotes an atrium, courtyard, or parvise in front of a basilica or church that is surrounded by arcades or colonnades, or trees, and which often contains a fountain A fountain, from the Latin "f ...
separated by five ambulacral grooves, the interambulacra. The ambulacral grooves bear four rows of tube feet but these are diminished in size or absent in some holothurians, especially on the dorsal surface. The two dorsal ambulacra make up the bivium while the three ventral ones are known as the trivium.''Cucumaria frondosa''
Invertebrate anatomy online. ed2012-02-12.
At the anterior end, the mouth is surrounded by a ring of tentacles which are usually retractable into the mouth. These are modified tube feet and may be simple, branched or arborescent. They are known as the introvert and posterior to them there is an internal ring of large calcareous ossicles. Attached to this are five bands of muscle running internally longitudinally along the ambulacra. There are also circular muscles, contraction of which cause the animal to elongate and the introvert to extend. Anterior to the ossicles lie further muscles, contraction of which cause the introvert to retract. The body wall consists of an epidermis and a dermis and contains smaller calcareous ossicles, the types of which are characteristics which help to identify different species. Inside the body wall is the coelom which is divided by three longitudinal
mesenteries In zoology, a mesentery is a membrane inside the body cavity of an animal. The term identifies different structures in different phyla: in vertebrates it is a double fold of the peritoneum enclosing the intestines; in other organisms it forms com ...
which surround and support the internal organs.


Digestive system

A
pharynx The pharynx (plural: pharynges) is the part of the throat behind the mouth and nasal cavity, and above the oesophagus and trachea (the tubes going down to the stomach and the lungs). It is found in vertebrates and invertebrates, though its struc ...
lies behind the mouth and is surrounded by a ring of ten
calcareous Calcareous () is an adjective meaning "mostly or partly composed of calcium carbonate", in other words, containing lime or being chalky. The term is used in a wide variety of scientific disciplines. In zoology ''Calcareous'' is used as an adje ...
plates. In most sea cucumbers, this is the only substantial part of the skeleton, and it forms the point of attachment for muscles that can retract the tentacles into the body for safety as for the main muscles of the body wall. Many species possess an
oesophagus The esophagus (American English) or oesophagus (British English; both ), non-technically known also as the food pipe or gullet, is an organ in vertebrates through which food passes, aided by peristaltic contractions, from the pharynx to the ...
and
stomach The stomach is a muscular, hollow organ in the gastrointestinal tract of humans and many other animals, including several invertebrates. The stomach has a dilated structure and functions as a vital organ in the digestive system. The stomach i ...
, but in some the pharynx opens directly into the
intestine The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The GI tract contains all the major organs of the digestive system, in humans ...
. The intestine is typically long and coiled, and loops through the body three times before terminating in a
cloaca In animal anatomy, a cloaca ( ), plural cloacae ( or ), is the posterior orifice that serves as the only opening for the digestive, reproductive, and urinary tracts (if present) of many vertebrate animals. All amphibians, reptiles and birds, a ...
l chamber, or directly as the
anus The anus (Latin, 'ring' or 'circle') is an opening at the opposite end of an animal's digestive tract from the mouth. Its function is to control the expulsion of feces, the residual semi-solid waste that remains after food digestion, which, d ...
.


Nervous system

Sea cucumbers have no true
brain A brain is an organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It is located in the head, usually close to the sensory organs for senses such as vision. It is the most complex organ in a v ...
. A ring of neural tissue surrounds the oral cavity, and sends nerves to the tentacles and the
pharynx The pharynx (plural: pharynges) is the part of the throat behind the mouth and nasal cavity, and above the oesophagus and trachea (the tubes going down to the stomach and the lungs). It is found in vertebrates and invertebrates, though its struc ...
. The animal is, however, quite capable of functioning and moving about if the nerve ring is surgically removed, demonstrating that it does not have a central role in nervous coordination. In addition, five major nerves run from the nerve ring down the length of the body beneath each of the ambulacral areas. Most sea cucumbers have no distinct sensory organs, although there are various nerve endings scattered through the skin, giving the animal a sense of touch and a sensitivity to the presence of light. There are, however, a few exceptions: members of the Apodida order are known to possess
statocyst The statocyst is a balance sensory receptor present in some aquatic invertebrates, including bivalves, cnidarians, ctenophorans, echinoderms, cephalopods, and crustaceans. A similar structure is also found in '' Xenoturbella''. The statocyst c ...
s, while some species possess small eye-spots near the bases of their tentacles.


Respiratory system

Sea cucumbers extract oxygen from water in a pair of "respiratory trees" that branch in the
cloaca In animal anatomy, a cloaca ( ), plural cloacae ( or ), is the posterior orifice that serves as the only opening for the digestive, reproductive, and urinary tracts (if present) of many vertebrate animals. All amphibians, reptiles and birds, a ...
just inside the
anus The anus (Latin, 'ring' or 'circle') is an opening at the opposite end of an animal's digestive tract from the mouth. Its function is to control the expulsion of feces, the residual semi-solid waste that remains after food digestion, which, d ...
, so that they "breathe" by drawing water in through the anus and then expelling it. The trees consist of a series of narrow tubules branching from a common duct, and lie on either side of the digestive tract. Gas exchange occurs across the thin walls of the tubules, to and from the fluid of the main body cavity. Together with the intestine, the respiratory trees also act as excretory organs, with nitrogenous waste diffusing across the tubule walls in the form of
ammonia Ammonia is an inorganic compound of nitrogen and hydrogen with the formula . A stable binary hydride, and the simplest pnictogen hydride, ammonia is a colourless gas with a distinct pungent smell. Biologically, it is a common nitrogenous was ...
and
phagocytic Phagocytosis () is the process by which a cell uses its plasma membrane to engulf a large particle (≥ 0.5 μm), giving rise to an internal compartment called the phagosome. It is one type of endocytosis. A cell that performs phagocytosis is ...
coelomocyte A coelomocyte () is a phagocytic leukocyte that appears in the bodies of animals that have a coelom. In most, it attacks and digests invading organisms such as bacteria and viruses through encapsulation and phagocytosis, though in some animals (e.g. ...
s depositing particulate waste.


Circulatory systems

Like all echinoderms, sea cucumbers possess both a
water vascular system The water vascular system is a hydraulic system used by echinoderms, such as sea stars and sea urchins, for locomotion, food and waste transportation, and respiration. The system is composed of canals connecting numerous tube feet. Echinoderms m ...
that provides hydraulic pressure to the tentacles and tube feet, allowing them to move, and a ''haemal system''. The latter is more complex than that in other echinoderms, and consists of well-developed vessels as well as open
sinuses Paranasal sinuses are a group of four paired air-filled spaces that surround the nasal cavity. The maxillary sinuses are located under the eyes; the frontal sinuses are above the eyes; the ethmoidal sinuses are between the eyes and the sphenoid ...
. A central haemal ring surrounds the pharynx next to the ring canal of the water vascular system, and sends off additional vessels along the radial canals beneath the ambulacral areas. In the larger species, additional vessels run above and below the intestine and are connected by over a hundred small muscular ampullae, acting as miniature hearts to pump blood around the haemal system. Additional vessels surround the respiratory trees, although they contact them only indirectly, via the coelomic fluid. Indeed, the blood itself is essentially identical with the coelomic fluid that bathes the organs directly, and also fills the water vascular system. Phagocytic coelomocytes, somewhat similar in function to the
white blood cell White blood cells, also called leukocytes or leucocytes, are the cell (biology), cells of the immune system that are involved in protecting the body against both infectious disease and foreign invaders. All white blood cells are produced and de ...
s of
vertebrates Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
, are formed within the haemal vessels, and travel throughout the body cavity as well as both circulatory systems. An additional form of coelomocyte, not found in other echinoderms, has a flattened discoid shape, and contains
hemoglobin Hemoglobin (haemoglobin BrE) (from the Greek word αἷμα, ''haîma'' 'blood' + Latin ''globus'' 'ball, sphere' + ''-in'') (), abbreviated Hb or Hgb, is the iron-containing oxygen-transport metalloprotein present in red blood cells (erythrocyte ...
. As a result, in many (though not all) species, both the blood and the coelomic fluid are red in colour.
Vanadium Vanadium is a chemical element with the symbol V and atomic number 23. It is a hard, silvery-grey, malleable transition metal. The elemental metal is rarely found in nature, but once isolated artificially, the formation of an oxide layer ( pas ...
has been reported in high concentrations in holothurian blood, however researchers have been unable to reproduce these results.


Locomotive organs

Like all echinoderms, sea cucumbers possess
pentaradial 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, take the face of a human being which has a pla ...
, with their bodies divided into five nearly identical parts around a central axis. However, because of their posture, they have secondarily evolved a degree of bilateral symmetry. For example, because one side of the body is typically pressed against the substratum, and the other is not, there is usually some difference between the two surfaces (except for
Apodida Apodida is an order of littoral to deep-sea, largely infaunal holothurians, sea cucumbers. This order comprises three families, 32 genera and about 270 known species, called apodids, "without feet". Characteristics These sea cucumbers are vagile ...
). Like
sea urchin Sea urchins () are spiny, globular echinoderms in the class Echinoidea. About 950 species of sea urchin live on the seabed of every ocean and inhabit every depth zone from the intertidal seashore down to . The spherical, hard shells (tests) of ...
s, most sea cucumbers have five strip-like ambulacral areas running along the length of the body from the mouth to the anus. The three on the lower surface have numerous
tube feet Tube feet (technically podia) are small active tubular projections on the oral face of an echinoderm, whether the arms of a starfish, or the undersides of sea urchins, sand dollars and sea cucumbers; they are more discreet though present on br ...
, often with suckers, that allow the animal to crawl along; they are called ''trivium''. The two on the upper surface have under-developed or vestigial tube feet, and some species lack tube feet altogether; this face is called ''bivium''. In some species, the ambulacral areas can no longer be distinguished, with tube feet spread over a much wider area of the body. Those of the order
Apodida Apodida is an order of littoral to deep-sea, largely infaunal holothurians, sea cucumbers. This order comprises three families, 32 genera and about 270 known species, called apodids, "without feet". Characteristics These sea cucumbers are vagile ...
have no tube feet or ambulacral areas at all, and burrow through sediment with muscular contractions of their body similar to that of worms, however five radial lines are generally still obvious along their body. Even in those sea cucumbers that lack regular tube feet, those that are immediately around the mouth are always present. These are highly modified into retractile
tentacle In zoology, a tentacle is a flexible, mobile, and elongated organ present in some species of animals, most of them invertebrates. In animal anatomy, tentacles usually occur in one or more pairs. Anatomically, the tentacles of animals work main ...
s, much larger than the locomotive tube feet. Depending on the species, sea cucumbers have between ten and thirty such tentacles and these can have a wide variety of shapes depending on the diet of the animal and other conditions. Many sea cucumbers have papillae, conical fleshy projections of the body wall with sensory tube feet at their apices. These can even evolve into long antennae-like structures, especially on the abyssal genus ''
Scotoplanes ''Scotoplanes'' is a genus of deep-sea sea cucumbers of the family Elpidiidae. Its species are commonly known as sea pigs. Locomotion Members of the Elpidiidae have particularly enlarged tube "feet" that have taken on a leg-like appearance, usi ...
''.


Endoskeleton

Echinoderms typically possess an internal skeleton composed of plates of calcium carbonate. In most sea cucumbers, however, these have become reduced to microscopic ossicles embedded beneath the skin. A few genera, such as '' Sphaerothuria'', retain relatively large plates, giving them a scaly armour.


Life history and behaviour


Habitat

Sea cucumbers can be found in great numbers on the deep seafloor, where they often make up the majority of the animal biomass. At depths deeper than , sea cucumbers comprise 90% of the total mass of the macrofauna. Sea cucumbers form large herds that move across the bathygraphic features of the ocean, hunting food. The body of some deep water holothurians, such as ''Enypniastes eximia'', ''Peniagone leander'' and ''Paelopatides confundens'', is made of a tough gelatinous tissue with unique properties that makes the animals able to control their own buoyancy, making it possible for them to either live on the ocean floor or to actively swim or float over it in order to move to new locations, in a manner similar to how the group
Torquaratoridae Torquaratoridae (Latin for "neck plow") is a family of acorn worms ( Hemichordata) that lives in deep waters between 350 to 4000 meters (the species Tergivelum baldwinae has been found 4100 meters below the surface). They can grow up to three fee ...
floats through water. Holothurians appear to be the echinoderms best adapted to extreme depths, and are still very diversified beyond 5,000 m deep: several species from the family
Elpidiidae Elpidiidae is a family of deep-sea sea cucumbers. They have a translucent body with long and stout podia acting like legs. The mouth is surrounded by thick and short oral tentacles, and the dorsal part often shows pairs of elongated podia as we ...
("sea pigs") can be found deeper than 9,500 m, and the record seems to be some species of the genus '' Myriotrochus'' (in particular '' Myriotrochus bruuni''), identified down to 10,687 meters deep. In more shallow waters, sea cucumbers can form dense populations. The strawberry sea cucumber (''Squamocnus brevidentis'') of
New Zealand New Zealand ( mi, Aotearoa ) is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and over 700 smaller islands. It is the sixth-largest island count ...
lives on rocky walls around the southern coast of the South Island where populations sometimes reach densities of 1,000 animals per square meter (93 /sq ft). For this reason, one such area in
Fiordland Fiordland is a geographical region of New Zealand in the south-western corner of the South Island, comprising the westernmost third of Southland. Most of Fiordland is dominated by the steep sides of the snow-capped Southern Alps, deep lakes, ...
is called the strawberry fields.


Locomotion

Some abyssal species in the abyssal order
Elasipodida Elasipodida is an order of sea cucumbers. They have numerous appendages, including conical papillae and leaf-like tentacles. Although many species are benthic, a number are pelagic, and may have their appendages modified to form sails or fins. ...
have evolved to a "benthopelagic" behaviour: their body is nearly the same density as the water around them, so they can make long jumps (up to high), before falling slowly back to the ocean floor. Most of them have specific swimming appendages, such as some kind of umbrella (like ''
Enypniastes ''Enypniastes'' is a genus of deep-sea sea cucumber. It is monotypic, being represented by the single species ''Enypniastes eximia''. Due to its unique appearance, the species has been dubbed the headless chicken fish, headless chicken monster, ...
''), or a long lobe on top of the body (''
Psychropotes ''Psychropotes'' is a genus of sea cucumbers in the family Psychropotidae. The members of this genus possess the ability to swim, although this is only facultative. Description As adults, species of ''Psychropotes'' are roughly cylindrical in sh ...
''). Only one species is known as a true completely
pelagic The pelagic zone consists of the water column of the open ocean, and can be further divided into regions by depth (as illustrated on the right). The word ''pelagic'' is derived . The pelagic zone can be thought of as an imaginary cylinder or wa ...
species, that never comes close to the bottom: ''
Pelagothuria natatrix ''Pelagothuria'' is a genus of sea cucumbers in the family Pelagothuriidae. It is monotypic, being represented by the single species ''Pelagothuria natatrix''. Characteristics This sea cucumber is somewhat unusual in appearance in comparison ...
''.


Diet

Holothuroidea are generally
scavenger Scavengers are animals that consume dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a herbivorous feeding b ...
s, feeding on debris in the
benthic zone The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from ancient Greek, βένθος (bénthos), meaning " ...
of the ocean. Exceptions include some
pelagic The pelagic zone consists of the water column of the open ocean, and can be further divided into regions by depth (as illustrated on the right). The word ''pelagic'' is derived . The pelagic zone can be thought of as an imaginary cylinder or wa ...
cucumbers and the species ''Rynkatorpa pawsoni'', which has a
commensal 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 ...
relationship with deep-sea
anglerfish The anglerfish are fish of the teleost order Lophiiformes (). They are bony fish named for their characteristic mode of predation, in which a modified luminescent fin ray (the esca or illicium) acts as a lure for other fish. The luminescence ...
. The
diet Diet may refer to: Food * Diet (nutrition), the sum of the food consumed by an organism or group * Dieting, the deliberate selection of food to control body weight or nutrient intake ** Diet food, foods that aid in creating a diet for weight loss ...
of most cucumbers consists of
plankton Plankton are the diverse collection of organisms found in Hydrosphere, water (or atmosphere, air) that are unable to propel themselves against a Ocean current, current (or wind). The individual organisms constituting plankton are called plankt ...
and decaying organic matter found in the sea. Some sea cucumbers position themselves in
currents Currents, Current or The Current may refer to: Science and technology * Current (fluid), the flow of a liquid or a gas ** Air current, a flow of air ** Ocean current, a current in the ocean *** Rip current, a kind of water current ** Current (stre ...
and catch food that flows by with their open tentacles. They also sift through the bottom
sediment Sediment is a naturally occurring material that is broken down by processes of weathering and erosion, and is subsequently transported by the action of wind, water, or ice or by the force of gravity acting on the particles. For example, sand an ...
s using their tentacles. Other species can dig into bottom silt or sand until they are completely buried. They then extrude their feeding tentacles, ready to withdraw at any hint of danger. In the South Pacific sea cucumbers may be found in densities of 40 individuals per square meter (33 /sq yd). These populations can process 19 kilograms of sediment per square meter (34 lb /sq yd) per year. The shape of the tentacles is generally adapted to the diet, and to the size of the particles to be ingested: the filter-feeding species mostly have complex arborescent tentacles, intended to maximize the surface area available for filtering, while the species feeding on the substratum will more often need digitate tentacles to sort out the nutritional material; the detritivore species living on fine sand or mud more often need shorter "peltate" tentacles, shaped like shovels. A single specimen can swallow more than 45 kg of sediment a year, and their excellent digestive capacities allow them to reject a finer, purer and homogeneous sediment. Therefore, sea cucumbers play a major role in the biological processing of the sea bed (bioturbation, purge, homogenization of the substratum etc.). Image:Euapta godeffroyi, détail.jpg, The mouth of an ''
Euapta godeffroyi ''Euapta godeffroyi'', the sticky snake sea cucumber, is a species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest ...
'', showing pinnate tentacles. Image:Apostichopus californicus.004 - Aquarium Finisterrae.jpg, Mouth of a ''
Holothuria ''Holothuria'' is the type genus of the marine animal family Holothuriidae, part of the class Holothuroidea, commonly known as sea cucumbers. Members of the genus are found in coastal waters in tropical and temperate regions. They are soft bodie ...
sp.'', showing peltate tentacles. Image:Cucumaria main.jpg, Mouth of a '' Cucumaria miniata'', with dendritic tentacles, for filtering the water. Image:Holothurie sp..jpg, Faeces of an holothurian. This participates in sediment homogenization and purification.


Communication and sociability


Reproduction

Most sea cucumbers reproduce by releasing
sperm Sperm is the male reproductive cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm with a tail known as a flagellum, whi ...
and
ova , abbreviated as OVA and sometimes as OAV (original animation video), are Japanese animated films and series made specially for release in home video formats without prior showings on television or in theaters, though the first part of an OVA s ...
into the ocean water. Depending on conditions, one organism can produce thousands of
gamete A gamete (; , ultimately ) is a haploid cell that fuses with another haploid cell during fertilization in organisms that reproduce sexually. Gametes are an organism's reproductive cells, also referred to as sex cells. In species that produce t ...
s. Sea cucumbers are typically
dioecious Dioecy (; ; adj. dioecious , ) is a characteristic of a species, meaning that it has distinct individual organisms (unisexual) that produce male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproductio ...
, with separate male and female individuals, but some species are protandric. The reproductive system consists of a single
gonad A gonad, sex gland, or reproductive gland is a mixed gland that produces the gametes and sex hormones of an organism. Female reproductive cells are egg cells, and male reproductive cells are sperm. The male gonad, the testicle, produces sper ...
, consisting of a cluster of tubules emptying into a single duct that opens on the upper surface of the animal, close to the tentacles. At least 30 species, including the red-chested sea cucumber ('' Pseudocnella insolens''), fertilize their eggs internally and then pick up the fertilized zygote with one of their feeding tentacles. The egg is then inserted into a pouch on the adult's body, where it develops and eventually hatches from the pouch as a juvenile sea cucumber. A few species are known to brood their young inside the body cavity, giving birth through a small rupture in the body wall close to the anus.


Development

In all other species, the egg develops into a free-swimming
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
, typically after around three days of development. The first stage of larval development is known as an auricularia, and is only around in length. This larva swims by means of a long band of cilia wrapped around its body, and somewhat resembles the
bipinnaria A bipinnaria is the first stage in the larval development of most starfish, and is usually followed by a brachiolaria stage. Movement and feeding is accomplished by the bands of cilia. Starfish that brood their young generally lack a bipinnaria st ...
larva of starfish. As the larva grows it transforms into the doliolaria, with a barrel-shaped body and three to five separate rings of cilia. The pentacularia is the third larval stage of sea cucumber, where the tentacles appear. The tentacles are usually the first adult features to appear, before the regular tube feet.


Symbiosis and commensalism

Numerous small animals can live in
symbiosis Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasit ...
or
commensalism 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 ...
with sea cucumbers, as well as some parasites. Some cleaner shrimps can live on the tegument of holothurians, in particular several species of the genus ''
Periclimenes ''Periclimenes'', commonly known as glass shrimp or cleaner shrimp, is a commensal and often symbiotic genus of semi-transparent shrimp within the family Palaemonidae. Species of this large genus feature a wide variety of coloration and patterns, ...
'' (genus which is specialized in echinoderms), in particular ''
Periclimenes imperator ''Periclimenes imperator'', known as the emperor shrimp, is a species of shrimp with a wide distribution across the Indo-Pacific. It lives commensally on a number of hosts, including the sea slug ''Hexabranchus''. A. J. Bruce first described ...
''.. A variety of fish, most commonly
pearl fish Pearlfish are marine (ocean), marine fish in the Actinopterygii, ray-finned fish Family (biology), family Carapidae. Pearlfishes inhabit the Tropics, tropical waters of the Atlantic Ocean, Atlantic, Indian Ocean, Indian, and Pacific Oceans at d ...
, have evolved a commensalistic symbiotic relationship with sea cucumbers in which the pearl fish will live in sea cucumber's cloaca using it for protection from predation, a source of food (the nutrients passing in and out of the anus from the water), and to develop into their adult stage of life. Many
polychaete Polychaeta () is a paraphyletic class of generally marine annelid worms, commonly called bristle worms or polychaetes (). Each body segment has a pair of fleshy protrusions called parapodia that bear many bristles, called chaetae, which are made ...
worms (family
Polynoidae Polynoidae is a family of marine Polychaete worms known as "scale worms" due to the scale-like elytra on the dorsal surface. Almost 900 species are currently recognised belonging to 9 subfamilies and 167 genera. They are active hunters, but gene ...
) and
crab Crabs are decapod crustaceans of the infraorder Brachyura, which typically have a very short projecting "tail" (abdomen) ( el, βραχύς , translit=brachys = short, / = tail), usually hidden entirely under the thorax. They live in all the ...
s (like '' Lissocarcinus orbicularis'') have also specialized to use the mouth or the cloacal respiratory trees for protection by living inside the sea cucumber. Nevertheless, holothurians species of the genus '' Actinopyga'' have anal teeth that prevent visitors from penetrating their anus. Sea cucumbers can also shelter bivalvia as endocommensals, such as '' Entovalva sp''. Image:Swimming Crab - Lissocarcinus Orbicularis on Sea Cucumber.jpg, '' Lissocarcinus orbicularis'', a symbiotic crab. Image:Shrimp on Sea Cucumber.jpg, ''
Periclimenes imperator ''Periclimenes imperator'', known as the emperor shrimp, is a species of shrimp with a wide distribution across the Indo-Pacific. It lives commensally on a number of hosts, including the sea slug ''Hexabranchus''. A. J. Bruce first described ...
'', a symbiotic shrimp. Image:Polychètes sur T. anax.JPG, Polynoid worms on a king sea cucumber.


Predators and defensive systems

Sea cucumbers are often ignored by most of the marine predators because of the toxins they contain (in particular
holothurin The holothurins are a group of toxins originally isolated from the sea cucumber '' Actinopyga agassizii''. They are contained within clusters of sticky threads called Cuvierian tubules which are expelled from the sea cucumber as a mode of sel ...
) and because of their often spectacular defensive systems. However, they remain a prey for some highly specialized predators which are not affected by their toxins, such as the big mollusks '' Tonna galea'' and ''
Tonna perdix ''Tonna perdix'', common name the partridge tun, is a species of very large sea snail, a marine (ocean), marine gastropod mollusc in the family (biology), family Tonnidae, the tun Gastropod shell, shells.Vos, C. (2010)Tonna perdix (Linnaeus, 175 ...
'', which paralyzes them using powerful poison before swallowing them completely. Some other less specialized and opportunist predators can also prey on sea cucumbers sometimes when they cannot find any better food, such as certain species of fish (
triggerfish Triggerfish are about 40 species of often brightly colored fish of the family Balistidae. Often marked by lines and spots, they inhabit tropical and subtropical oceans throughout the world, with the greatest species richness in the Indo-Pacifi ...
, pufferfish) and crustaceans (crabs, lobsters,
hermit crab Hermit crabs are anomuran decapod crustaceans of the superfamily Paguroidea that have adapted to occupy empty scavenged mollusc shells to protect their fragile exoskeletons. There are over 800 species of hermit crab, most of which possess an as ...
s). Some species of coral-reef sea cucumbers within the order Aspidochirotida can defend themselves by expelling their sticky
cuvierian tubules Cuvierian tubules are clusters of fine tubes located at the base of the respiratory tree in some sea cucumbers in the genera ''Bohadschia'', ''Holothuria'' and ''Pearsonothuria'', all of which are included in the family Holothuriidae. The tubules ...
(enlargements of the respiratory tree that float freely in the coelom) to entangle potential predators. When startled, these cucumbers may expel some of them through a tear in the wall of the
cloaca In animal anatomy, a cloaca ( ), plural cloacae ( or ), is the posterior orifice that serves as the only opening for the digestive, reproductive, and urinary tracts (if present) of many vertebrate animals. All amphibians, reptiles and birds, a ...
in an autotomic process known as evisceration. Replacement tubules grow back in one and a half to five weeks, depending on the species. The release of these tubules can also be accompanied by the discharge of a toxic chemical known as
holothurin The holothurins are a group of toxins originally isolated from the sea cucumber '' Actinopyga agassizii''. They are contained within clusters of sticky threads called Cuvierian tubules which are expelled from the sea cucumber as a mode of sel ...
, which has similar properties to soap. This chemical can kill animals in the vicinity and is one more method by which these sedentary animals can defend themselves.


Estivation

If the water temperature becomes too high, some species of sea cucumber from temperate seas can aestivate. While they are in this state of dormancy they stop feeding, their gut atrophies, their metabolism slows down and they lose weight. The body returns to its normal state when conditions improve.


Phylogeny and classification

Holothuroidea (sea cucumbers) are one of five extant classes that make up the phylum Echinodermata. This is one of the most distinctive and diverse phyla, ranging from starfish to urchins to sea cucumbers and many other organisms. The echinoderms are mainly distinguished from other phyla by their body plan and organization. While the organisms in this phylum may not all look the same from the outside, their make-up is another story. The earlliest sea cucumbers are known from the middle
Ordovician The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start ...
, over 450 million years ago. The apodida is the sister group to the other orders of sea cucumbers. All echinoderms share three main characteristics. When mature, echinoderms have a pentamerous radial symmetry. While this can easily be seen in a sea star or brittle star, in the sea cucumber it is less distinct and seen in their five primary tentacles. The pentamerous radial symmetry can also be seen in their five ambulacral canals. The ambulacral canals are used in their water vascular system which is another characteristic that binds this phylum together. The water vascular system develops from their middle coelom or hydrocoel. Echinoderms use this system for many things including movement by pushing water in and out of their podia or "tube feet". Echinoderms tube feet (including sea cucumbers) can be seen aligned along the side of their axes. While echinoderms are invertebrates, meaning they do not have a spine, they do all have an endoskeleton that is secreted by the
mesenchyme Mesenchyme () is a type of loosely organized animal embryonic connective tissue of undifferentiated cells that give rise to most tissues, such as skin, blood or bone. The interactions between mesenchyme and epithelium help to form nearly every o ...
. This endoskeleton is composed of plates called ossicles. They are always internal but may only be covered by a thin epidermal layer like in sea urchin's spines. In the sea cucumber the ossicles are only found in the dermis, making them a very supple organism. For most echinoderms, their ossicles are found in units making up a three dimensional structure. However, in sea cucumbers the ossicles are found in a two-dimensional network. All echinoderms also possess anatomical feature(s) called
mutable collagenous tissues Catch connective tissue (also called mutable collagenous tissue) is a kind of connective tissue found in echinoderms (such as starfish and sea cucumbers) which can change its mechanical properties in a few seconds or minutes through nervous control ...
, or MCTs. Such tissues can rapidly change their passive mechanical properties from soft to stiff under the control of the nervous system and coordinated with muscle activity. Different echinoderm classes use MCTs in different ways. The asteroids, sea stars, can detach limbs for self-defense and then regenerate them. The Crinoidea, sea fans, can go from stiff to limp depending on the current for optimal filter feeding. The Echinoidea, sand dollars, use MCTs to grow and replace their rows of teeth when they need new ones. The Holothuroidea, sea cucumbers, use MCTs to eviscerate their gut as a self-defense response. MCTs can be used in many ways but are all similar at the cellular level and in mechanics of function. A common trend in the uses of MCTs is that they are generally used for self-defense mechanisms and in regeneration. Holothurian classification is complex and their paleontological phylogeny relies on a limited number of well-preserved specimens. The modern taxonomy is based first of all on the presence or the shape of certain soft parts (podia, lungs, tentacles, peripharingal crown) to determine the main orders, and secondarily on the microscopic examination of ossicles to determine the genus and the species. Contemporary genetic methods have been helpful in clarifying their classification. Taxonomic classification according to
World Register of Marine Species The World Register of Marine Species (WoRMS) is a taxonomic database that aims to provide an authoritative and comprehensive list of names of marine organisms. Content The content of the registry is edited and maintained by scientific specialist ...
: * subclass Actinopoda Ludwig, 1891 ** order
Dendrochirotida Dendrochirotida are an order of sea cucumbers. Members of this order have branched tentacles and are suspension feeders. Examples include '' Thyonella'' and ''Cucumaria''. Characteristics Holothurians in this order are characterised by ten to thi ...
Grube, 1840 *** family
Cucumariidae Cucumariidae is a family of sea cucumbers, marine animals with elongated bodies, leathery skins and tentacles that are found on the sea bed. Description Members of the family Cucumariidae are small to medium-sized sea cucumbers, characterised by ...
Ludwig, 1894 *** family
Cucumellidae ''Cucumella'' is a genus of sea cucumbers belonging to the monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies ...
Thandar & Arumugam, 2011 *** family
Heterothyonidae Heterothyonidae is a family of sea cucumbers belonging to the order Dendrochirotida Dendrochirotida are an order of sea cucumbers. Members of this order have branched tentacles and are suspension feeders. Examples include '' Thyonella'' and ''C ...
Pawson, 1970 *** family † Monilipsolidae Smith & Gallemí, 1991 *** family
Paracucumidae Paracucumidae is a family of sea cucumbers belonging to the order Dendrochirotida Dendrochirotida are an order of sea cucumbers. Members of this order have branched tentacles and are suspension feeders. Examples include '' Thyonella'' and ''Cucu ...
Pawson & Fell, 1965 *** family Phyllophoridae Östergren, 1907 *** family
Placothuriidae ''Placothuria'' is a genus of sea cucumbers belonging to the monotypic family Placothuriidae. The species of this genus are found in Australia and New Zealand. Species: *''Placothuria huttoni'' *''Placothuria molpadioides'' *''Placothuria squ ...
Pawson & Fell, 1965 *** family
Psolidae Psolidae is a family of sea cucumbers, marine animals with elongated bodies, leathery skins and tentacles that are found on the sea bed. Description Members of the family Psolidae are small and inconspicuous sea cucumbers found in crevices and u ...
Burmeister, 1837 *** family
Rhopalodinidae Rhopalodinidae is a family of sea cucumbers belonging to the order Dendrochirotida Dendrochirotida are an order of sea cucumbers. Members of this order have branched tentacles and are suspension feeders. Examples include '' Thyonella'' and ''Cuc ...
Théel, 1886 *** family
Sclerodactylidae Sclerodactylidae is a family of sea cucumbers Sea cucumbers are echinoderms from the class Holothuroidea (). They are marine animals with a leathery skin and an elongated body containing a single, branched gonad. Sea cucumbers are found on t ...
Panning, 1949 *** family
Vaneyellidae Vaneyellidae is a family of sea cucumbers belonging to the order Dendrochirotida Dendrochirotida are an order of sea cucumbers. Members of this order have branched tentacles and are suspension feeders. Examples include '' Thyonella'' and ''Cucum ...
Pawson & Fell, 1965 *** family
Ypsilothuriidae Ypsilothuriidae is a family of sea cucumbers belonging to the order Dendrochirotida Dendrochirotida are an order of sea cucumbers. Members of this order have branched tentacles and are suspension feeders. Examples include '' Thyonella'' and ''Cu ...
Heding, 1942 ** order
Elasipodida Elasipodida is an order of sea cucumbers. They have numerous appendages, including conical papillae and leaf-like tentacles. Although many species are benthic, a number are pelagic, and may have their appendages modified to form sails or fins. ...
Théel, 1882 *** family
Elpidiidae Elpidiidae is a family of deep-sea sea cucumbers. They have a translucent body with long and stout podia acting like legs. The mouth is surrounded by thick and short oral tentacles, and the dorsal part often shows pairs of elongated podia as we ...
Théel, 1882 *** family
Laetmogonidae Laetmogonidae is a family of sea cucumbers Sea cucumbers are echinoderms from the class Holothuroidea (). They are marine animals with a leathery skin and an elongated body containing a single, branched gonad. Sea cucumbers are found on the ...
Ekman, 1926 *** family † Palaeolaetmogonidae Reich, 2012 *** family
Pelagothuriidae Pelagothuriidae is a family of deep-sea swimming sea cucumbers. They are somewhat unusual in appearance, in comparison with other sea cucumbers, having numerous appendages, including conical papillae and leaf-like tentacles. Most of them are bent ...
Ludwig, 1893 *** family
Psychropotidae Psychropotidae is a family of deep-sea swimming sea cucumbers. The geographic range of some psychropotids is very extensive at abyssal depths, whereas other species are found within more restricted ranges.P. A. Tyler & D. S. M. Billett (1988) Th ...
Théel, 1882 ** order
Holothuriida Holothuriida is an order of sea cucumbers. Taxa within the order Holothuriida were previously classified in the order Aspidochirotida, which was determined to be polyphyletic in 2017. Some taxa were also reclassified into the clades Synallactida ...
Miller, Kerr, Paulay, Reich, Wilson, Carvajal & Rouse, 2017 *** family
Holothuriidae Holothuriidae is a family of sea cucumbers, a type of echinoderm. Description Members of the family Holothuriidae have thick fleshy bodies and several rows of tube feet which are used for moving around and for adhering to the surface. The body ...
Burmeister, 1837 *** family Mesothuriidae Smirnov, 2012 ** order
Molpadida Molpadida is an order of sea cucumbers. The body shape is fusiform and unlike other sea cucumbers, their hind body is narrowed to form a distinct tail. Although they possess tentacles around the mouth derived from the water vascular system, they ...
Haeckel, 1896 *** family
Caudinidae Caudinidae is a family of sea cucumbers, marine animals with elongated bodies, leathery skins and tentacles that are found on the sea floor. Description Members of the family Caudinidae are fairly small, plump sea cucumbers with a thin body wal ...
Heding, 1931 *** family Eupyrgidae Semper, 1867 *** family Gephyrothuriidae Koehler & Vaney, 1905 *** family
Molpadiidae Molpadiidae is a family of sea cucumbers, marine invertebrates with elongated bodies, leathery skins and tentacles that are found on or burrowing in the sea floor. Description Members of Molpadiidae are fairly small, plump sea cucumbers with a s ...
Müller, 1850 ** order
Persiculida Persiculida is an order of sea cucumbers. Taxa within the order Persiculida were previously classified in an order called Aspidochirotida, which was determined to be polyphyletic A polyphyletic group is an assemblage of organisms or other ...
Miller, Kerr, Paulay, Reich, Wilson, Carvajal & Rouse, 2017 *** family Gephyrothuriidae Koehler & Vaney, 1905 *** family
Molpadiodemidae ''Molpadiodemas'' is a genus of sea cucumbers belonging to the monotypic family Molpadiodemidae. The genus has an almost cosmopolitan distribution. Species The following species are recognised in the genus ''Molpadiodemas'': *''Molpadiodemas atla ...
Miller, Kerr, Paulay, Reich, Wilson, Carvajal & Rouse, 2017 *** family Pseudostichopodidae Miller, Kerr, Paulay, Reich, Wilson, Carvajal & Rouse, 2017 ** order Synallactida Miller, Kerr, Paulay, Reich, Wilson, Carvajal & Rouse, 2017 *** family Deimatidae Théel, 1882 *** family
Stichopodidae The Stichopodidae are a family of sea cucumbers, part of the order Synallactida. Description Members of this family are mostly large or medium-sized holothuroids with a squarish cross-section, a flat ventral surface, and large, fleshy, cone-sha ...
Haeckel, 1896 *** family
Synallactidae The Synallactidae are a family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being o ...
Ludwig, 1894 * subclass †ArthrochirotaceaSmirnov, 2012 ** order † Arthrochirotida Brandt, 1835 *** family † Palaeocucumariidae Frizzell & Exline, 1966 * subclass Paractinopoda Ludwig, 1891 ** order
Apodida Apodida is an order of littoral to deep-sea, largely infaunal holothurians, sea cucumbers. This order comprises three families, 32 genera and about 270 known species, called apodids, "without feet". Characteristics These sea cucumbers are vagile ...
Brandt, 1835Small text *** family
Chiridotidae Chiridotidae is a family of sea cucumbers found in the order Apodida. Within the family, there are 16 recognized genera all with different ranges of body types and functions. Sea cucumbers play a fundamental role in many marine ecosystems. Des ...
Östergren, 1898 *** family
Myriotrochidae Myriotrochidae is a family of sea cucumbers. Description and characteristics Members of this family have digitate tentacles, bearing 2-8 digits on each side. They lack podia, radial canals and respiratory tree. The soft body wall is supporte ...
Théel, 1877 *** family
Synaptidae Synaptidae is a family of sea cucumbers that have no tube feet, tentacle ampullae, retractor muscles, respiratory trees, or cuvierian tubules. They also lack radial canals of the water-vascular system, with only the circumoral ring present. S ...
Burmeister, 1837


Relation to humans


Food

To supply the markets of
Southern China South China () is a geographical and cultural region that covers the southernmost part of China. Its precise meaning varies with context. A notable feature of South China in comparison to the rest of China is that most of its citizens are not n ...
,
Makassar Makassar (, mak, ᨆᨀᨔᨑ, Mangkasara’, ) is the capital of the Indonesian province of South Sulawesi. It is the largest city in the region of Eastern Indonesia and the country's fifth-largest urban center after Jakarta, Surabaya, Med ...
trepanger Trepanging is the act of collection or harvesting of sea cucumbers, known in Indonesian as ''trepang'', Malay těripang, and used as food. The collector, or fisher, of ''trepang'' is a trepanger. Trepanging is comparable to clamming, crabbin ...
s traded with the
Indigenous Australians Indigenous Australians or Australian First Nations are people with familial heritage from, and membership in, the ethnic groups that lived in Australia before British colonisation. They consist of two distinct groups: the Aboriginal peoples ...
of
Arnhem Land Arnhem Land is a historical region of the Northern Territory of Australia, with the term still in use. It is located in the north-eastern corner of the territory and is around from the territory capital, Darwin. In 1623, Dutch East India Compan ...
from at least the 18th century and probably earlier. This is the first recorded example of trade between the inhabitants of the Australian continent and their Asian neighbours. There are many commercially important species of sea cucumber that are harvested and dried for export for use in
Chinese cuisine Chinese cuisine encompasses the numerous cuisines originating from China, as well as overseas cuisines created by the Chinese diaspora. Because of the Chinese diaspora and historical power of the country, Chinese cuisine has influenced many ot ...
as ''hoisam''. Some of the more commonly found species in markets include: *''
Holothuria nobilis ''Holothuria'' (''Microthele'') ''nobilis'', the black teatfish, is a species of sea cucumber in the genus ''Holothuria ''Holothuria'' is the type genus of the marine animal family Holothuriidae, part of the class Holothuroidea, commonly kno ...
'' *''
Thelenota ananas ''Thelenota ananas'', also known as pineapple sea cucumber, oloturia ananas, tripang, prickly skin cucumber, pointed teat sea cucumber, armoured sea cucumber, giant sea cucumber, sand fish or prickly redfish, is a species of sea cucumber found i ...
'' *''
Actinopyga echinites ''Actinopyga echinites'', commonly known as the brownfish or deep water redfish, is a species of sea cucumber in the family Holothuriidae. It is native to the tropical Indo-Pacific region and is harvested for food. Description ''Actinopyga echi ...
'' *''
Actinopyga palauensis ''Actinopyga palauensis'', the Panning's blackfish, is a species of sea cucumber within the family Holothuriidae. The species distribution is in the western Pacific near areas such as Palau, the Federated States of Micronesia, Tonga, Niue, eastern ...
'' *''
Holothuria scabra ''Holothuria scabra'', or sandfish, is a species of sea cucumber in the family Holothuriidae. It was placed in the subgenus ''Metriatyla'' by Rowe in 1969 and is the type species of the subgenus. Sandfish are harvested and processed into "beche- ...
'' *''
Holothuria fuscogilva ''Holothuria (Microthele) fuscogilva'', also known as the white teatfish or white teeth, is a species of sea cucumber in the genus '' Holothuria'', subgenus ''Microthele''. The cucumber is found in the tropical waters of the Indo-Pacific ocean. Th ...
'' *''
Actinopyga mauritiana ''Actinopyga mauritiana'', commonly known as the surf redfish, is a species of sea cucumber in the family Holothuriidae. It is native to the tropical West Indo-Pacific region and is harvested for food. Description ''Actinopyga mauritiana'' grow ...
'' *'' Stichius japonicus'' *'' Apostichopus californicus'' *''
Acaudina molpadioides ''Acaudina molpadioides'' is a species of sea cucumber in the genus ''Acaudina''. The species is found in the tropical waters of India, Thailand, Malaysia, Indonesia, Philippines and Australia. The cucumber is harvested for use in traditional med ...
'' *'' Isostichopus fuscus''


Medicine

According to the
American Cancer Society The American Cancer Society (ACS) is a nationwide voluntary health organization dedicated to eliminating cancer. Established in 1913, the society is organized into six geographical regions of both medical and lay volunteers operating in more than ...
, although it has been used in traditional Asian folk medicine for a variety of ailments, "there is little reliable scientific evidence to support claims that sea cucumber is effective in treating cancer, arthritis, and other diseases" but research is examining "whether some compounds made by sea cucumbers may be helpful against cancer". Various pharmaceutical companies emphasize '' gamat'', the traditional medicinal usage of this animal. Extracts are prepared and made into oil, cream or cosmetics. Some products are intended to be taken internally. A review article found that
chondroitin sulfate Chondroitin sulfate is a sulfated glycosaminoglycan (GAG) composed of a chain of alternating sugars ( N-acetylgalactosamine and glucuronic acid). It is usually found attached to proteins as part of a proteoglycan. A chondroitin chain can have ov ...
and related compounds found in sea cucumbers can help in treating joint-pain, and that dried sea cucumber is "medicinally effective in suppressing
arthralgia Arthralgia (from Greek ''arthro-'', joint + ''-algos'', pain) literally means ''joint pain''. Specifically, arthralgia is a symptom of injury, infection, illness (in particular arthritis), or an allergic reaction to medication. According to MeSH, ...
". Another study suggested that sea cucumbers contain all the fatty acids necessary to play a potentially active role in tissue repair. Sea cucumbers are under investigation for use in treating ailments including
colorectal cancer Colorectal cancer (CRC), also known as bowel cancer, colon cancer, or rectal cancer, is the development of cancer from the colon or rectum (parts of the large intestine). Signs and symptoms may include blood in the stool, a change in bowel m ...
. Surgical probes made of nanocomposite material based on the sea cucumber have been shown to reduce brain scarring. One study found that a lectin from '' Cucumaria echinata'' impaired the development of the malaria parasite when produced by transgenic
mosquitoes Mosquitoes (or mosquitos) are members of a group of almost 3,600 species of small flies within the family Culicidae (from the Latin ''culex'' meaning "gnat"). The word "mosquito" (formed by ''mosca'' and diminutive ''-ito'') is Spanish for "litt ...
. File:Dried sea cucumber.jpg, "Teripang" in a market in Asia. File:Sea cucumber dish 2.jpg, Sea cucumber in sauce in China. File:Haisom Cah Jamur 1.JPG, ''Haisom cah jamur'', Chinese Indonesian sea cucumber with mushroom. File:Sea cucumber dish 3.jpg,
Deep fried Deep frying (also referred to as deep fat frying) is a cooking method in which food is submerged in hot fat, traditionally lard but today most commonly oil, as opposed to the shallow oil used in conventional frying done in a frying pan. Norma ...
sea cucumbers. File:Keripik teripang 1.JPG, ''
Kripik ''Kripik'' or ''keripik'' are Indonesian chips or crisps, bite-size snack crackers that can be savoury or sweet. They are made from various dried fruits, tubers, vegetables, and fish that have undergone a deep frying process in hot vegetable o ...
teripang'', Indonesian sea cucumber cracker.


Procurement

Sea cucumbers are harvested from the environment, both legally and illegally, and are increasingly farmed via
aquaculture Aquaculture (less commonly spelled aquiculture), also known as aquafarming, is the controlled cultivation ("farming") of aquatic organisms such as fish, crustaceans, mollusks, algae and other organisms of value such as aquatic plants (e.g. lot ...
. The harvested animals are normally dried for resale. In 2016, prices on Alibaba ranged up to $1,000/kg.


Commercial harvest

In recent years, the sea cucumber industry in Alaska has increased due to increased demand for the skins and muscles to China. Wild sea cucumbers are caught by divers. Wild Alaskan sea cucumbers have higher nutritional value and are larger than farmed Chinese sea cucumbers. Larger size and higher nutritional value has allowed the Alaskan fisheries to continue to compete for market share. One of Australia's oldest fisheries is the collection of sea cucumber, harvested by divers from throughout the
Coral Sea The Coral Sea () is a marginal sea of the South Pacific off the northeast coast of Australia, and classified as an interim Australian bioregion. The Coral Sea extends down the Australian northeast coast. Most of it is protected by the Fre ...
in far
North Queensland North Queensland or the Northern Region is the northern part of the Australian state of Queensland that lies just south of Far North Queensland. Queensland is a massive state, larger than many countries, and its tropical northern part has been ...
,
Torres Straits The Torres Strait (), also known as Zenadh Kes, is a strait between Australia and the Melanesian island of New Guinea. It is wide at its narrowest extent. To the south is Cape York Peninsula, the northernmost extremity of the Australian mai ...
and
Western Australia Western Australia (commonly abbreviated as WA) is a state of Australia occupying the western percent of the land area of Australia excluding external territories. It is bounded by the Indian Ocean to the north and west, the Southern Ocean to th ...
. In the late 1800s as many as 400 divers operated from Cook Town, Queensland.
Overfishing Overfishing is the removal of a species of fish (i.e. fishing) from a body of water at a rate greater than that the species can replenish its population naturally (i.e. the overexploitation of the fishery's existing fish stock), resulting in th ...
of sea cucumbers in the Great Barrier Reef is threatening their population. Their popularity as luxury seafood in
East Asia East Asia is the eastern region of Asia, which is defined in both geographical and ethno-cultural terms. The modern states of East Asia include China, Japan, Mongolia, North Korea, South Korea, and Taiwan. China, North Korea, South Korea and ...
n countries poses a serious threat.


Black market

As of 2013 a thriving
black market A black market, underground economy, or shadow economy is a clandestine market or series of transactions that has some aspect of illegality or is characterized by noncompliance with an institutional set of rules. If the rule defines the se ...
was driven by demand in China where at its peak might have sold for the equivalent of US$300 and a single sea cucumber for about US$160. A crackdown by governments both in and out of China reduced both prices and consumption, particularly among government officials who had been known to eat (and were able to afford purchasing) the most expensive and rare species. In the
Caribbean Sea The Caribbean Sea ( es, Mar Caribe; french: Mer des Caraïbes; ht, Lanmè Karayib; jam, Kiaribiyan Sii; nl, Caraïbische Zee; pap, Laman Karibe) is a sea of the Atlantic Ocean in the tropics of the Western Hemisphere. It is bounded by Mexico ...
off the shores of the
Yucatán Peninsula The Yucatán Peninsula (, also , ; es, Península de Yucatán ) is a large peninsula in southeastern Mexico and adjacent portions of Belize and Guatemala. The peninsula extends towards the northeast, separating the Gulf of Mexico to the north ...
near fishing ports such as Dzilam de Bravo, illegal harvesting had devastated the population and resulted in conflict as rival gangs struggled to control the harvest.


Aquaculture

Overexploitation Overexploitation, also called overharvesting, refers to harvesting a renewable resource to the point of diminishing returns. Continued overexploitation can lead to the destruction of the resource, as it will be unable to replenish. The term app ...
of sea cucumber stocks in many parts of the world provided motivation for the development of sea cucumber
aquaculture Aquaculture (less commonly spelled aquiculture), also known as aquafarming, is the controlled cultivation ("farming") of aquatic organisms such as fish, crustaceans, mollusks, algae and other organisms of value such as aquatic plants (e.g. lot ...
in the early 1980s. The Chinese and Japanese were the first to develop successful hatchery technology on ''
Apostichopus japonicus ''Apostichopus japonicus'' is a species of sea cucumber in the family Stichopodidae. It is found in shallow temperate waters along the coasts of south east Asia and is commonly known as the Japanese spiky sea cucumber or the Japanese sea cucumbe ...
'', prized for its high meat content and success in commercial hatcheries.James, B. D. (2004
Captive breeding of the sea cucumber, ''Holothuria scabra'', from India
In Lovatelli, A. (comp./ed.); Conand, C.; Purcell, S.; Uthicke, S.; Hamel, J.-F.; Mercier, A. (eds.) ''Advances in sea cucumber aquaculture and management''. FAO Fisheries Technical Paper. No. 463. Rome, FAO. 2004.425p.
Using techniques pioneered by the Chinese and Japanese, a second species, ''
Holothuria scabra ''Holothuria scabra'', or sandfish, is a species of sea cucumber in the family Holothuriidae. It was placed in the subgenus ''Metriatyla'' by Rowe in 1969 and is the type species of the subgenus. Sandfish are harvested and processed into "beche- ...
'', was cultured for the first time in India in 1988.James, D. B., Gandhi, A. D., Palaniswamy, N., & Rodrigo, J. X. (1994
Techniques and Culture of the Sea-cucumber ''Holothuria scabra''
CMFRI Special Publication(57),1-40.
In recent years Australia, Indonesia, New Caledonia, Maldives, Solomon Islands and Vietnam have successfully cultured ''H. scabra'' using the same technology, and now culture other species.


Conservation

In 2020, the Indian government created the world's first sea cucumber conservation area, the
Dr KK Mohammed Koya Sea Cucumber Conservation Reserve The Dr KK Mohammed Koya Sea Cucumber Conservation Reserve is the first sea cucumber conservation area in the world, announced following reports of smuggling of sea cucumbers and other vulnerable species for trade in East Asia. It is located in the ...
, to protect the sea cucumber species. In India, the commercial harvesting and transportation of sea cucumbers is banned.


In popular culture

*
Edgar Allan Poe Edgar Allan Poe (; Edgar Poe; January 19, 1809 – October 7, 1849) was an American writer, poet, editor, and literary critic. Poe is best known for his poetry and short stories, particularly his tales of mystery and the macabre. He is wide ...
's only novel, ''
The Narrative of Arthur Gordon Pym of Nantucket ''The Narrative of Arthur Gordon Pym of Nantucket'' (1838) is the only complete novel written by American writer Edgar Allan Poe. The work relates the tale of the young Arthur Gordon Pym, who stows away aboard a whaling ship called the ''Grampus' ...
'' (1838), includes in its 20th chapter a long, detailed description of sea cucumbers, which the narrator calls ''biche de mer''. *In
Kir Bulychov Kir Bulychev (russian: Кир Булычёв ''Kir Bulychyov 18 October 1934 – 5 September 2003) is a pen name of Igor Vsevolodovich Mozheiko (И́горь Все́володович Може́йко), a Soviet Russian science fiction writer, c ...
's science-fiction novella, "Half a Life", a human being kidnapped by alien machines described her fellow alien prisoners as "trepangs". *The first movement of French composer
Erik Satie Eric Alfred Leslie Satie (, ; ; 17 May 18661 July 1925), who signed his name Erik Satie after 1884, was a French composer and pianist. He was the son of a French father and a British mother. He studied at the Paris Conservatoire, but was an und ...
's Embryons desséchés is titled "D'Holothournie". It is said to emulate the "
purring A purr is a tonal fluttering sound made by some species of felids and two species of genets. It varies in loudness and tone among species and in the same animal. Felids are a family of mammals that belong to the order Carnivora and are info ...
" of the Holothourian. *Sea cucumbers have inspired thousands of
haiku is a type of short form poetry originally from Japan. Traditional Japanese haiku consist of three phrases that contain a ''kireji'', or "cutting word", 17 '' on'' (phonetic units similar to syllables) in a 5, 7, 5 pattern, and a ''kigo'', or se ...
in
Japan Japan ( ja, 日本, or , and formally , ''Nihonkoku'') is an island country in East Asia. It is situated in the northwest Pacific Ocean, and is bordered on the west by the Sea of Japan, while extending from the Sea of Okhotsk in the north ...
, where they are called ''namako'' (海鼠), written with characters that can be translated as "sea mice" (an example of
gikun are the logographic Chinese characters taken from the Chinese script and used in the writing of Japanese. They were made a major part of the Japanese writing system during the time of Old Japanese and are still used, along with the subsequent ...
). In English translations of these haiku, they are usually called "sea slugs". According to the ''
Oxford English Dictionary The ''Oxford English Dictionary'' (''OED'') is the first and foundational historical dictionary of the English language, published by Oxford University Press (OUP). It traces the historical development of the English language, providing a com ...
'', the English term "sea slug" was originally applied to holothurians during the 18th century. The term is now applied to several groups of
sea snail Sea snail is a common name for slow-moving marine gastropod molluscs, usually with visible external shells, such as whelk or abalone. They share the taxonomic class Gastropoda with slugs, which are distinguished from snails primarily by the ...
s, marine
gastropod The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. T ...
mollusk Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is e ...
s that have no shell or only a very reduced shell, including the
nudibranch Nudibranchs () are a group of soft-bodied marine gastropod molluscs which shed their shells after their larval stage. They are noted for their often extraordinary colours and striking forms, and they have been given colourful nicknames to matc ...
s. Almost 1,000 Japanese holothurian haiku translated into English appear in the book ''Rise, Ye Sea Slugs!'' by
Robin D. Gill Robin Dallas Gill, born in 1951 at Miami Beach, Florida, USA, and brought up on the island of Key Biscayne in the Florida Keys, is a bilingual author in Japanese and English. He first wrote extensively on stereotypes of Japanese identity bef ...
.Gill, Robin
''Rise, Ye Sea Slugs!''
Paraverse Press, 2003,
*Nobel laureate poet
Wisława Szymborska Maria Wisława Anna SzymborskaVioletta Szosta gazeta.pl, 9 February 2012. ostęp 2012-02-11 (; 2 July 1923 – 1 February 2012) was a Polish poet, essayist, translator, and recipient of the 1996 Nobel Prize in Literature. Born in Prowent ( ...
wrote a poem which mentions holothurians, titled "Autotomy". *In the book ''
John Dies at the End ''John Dies at the End'' is a Comic novel, comic horror fiction, horror novel written by David Wong (writer), David Wong that was first published online as a webserial beginning in 2001, then as an edited ebook, manuscript in 2004, and a printe ...
'', the character Amy Sullivan was nicknamed "Cucumber" by the narrator/author when the two were children. This is assumed by other characters to have a sexual connotation but is actually a reference to her frequent nausea. The name is in reference to a sea cucumber's use of vomiting as a method of self-defense. *A descriptive passage in American novelist
Cormac McCarthy Cormac McCarthy (born Charles Joseph McCarthy Jr., July 20, 1933) is an American writer who has written twelve novels, two plays, five screenplays and three short stories, spanning the Western and post-apocalyptic genres. He is known for his gr ...
's 1985 anti-Western ''
Blood Meridian ''Blood Meridian or the Evening Redness in the West'' is a 1985 in literature, 1985 Epic (genre), epic novel by American author Cormac McCarthy, classified under the Western (genre), Western, or sometimes the Revisionist Western, anti-Western, g ...
'' likens cactus embers to holothurians: "In the thorn forest through which they'd passed the little desert wolves yapped and on the dry plain before them others answered and the wind fanned the coals that he watched. The bones of cholla that glowed there in their incandescent basketry pulsed like burning holothurians in the phosphorous dark of the sea's deeps."


See also

* Gamat *
Trepanging Trepanging is the act of collection or harvesting of sea cucumbers, known in Indonesian as ''trepang'', Malay těripang, and used as food. The collector, or fisher, of ''trepang'' is a trepanger. Trepanging is comparable to clamming, crabbing, ...


References

Informational notes Citations


External links

* {{Taxonbar, from=Q127470 Extant Ordovician first appearances Edible shellfish Commercial echinoderms