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Starfish or sea stars are star-shaped
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 s ...
s belonging to the class Asteroidea (). Common usage frequently finds these names being also applied to ophiuroids, which are correctly referred to as
brittle star Brittle stars, serpent stars, or ophiuroids (; ; referring to the serpent-like arms of the brittle star) are echinoderms in the class Ophiuroidea, closely related to starfish. They crawl across the sea floor using their flexible arms for locomot ...
s or basket stars. Starfish are also known as asteroids due to being in the class Asteroidea. About 1,900 species of starfish live on the
seabed The seabed (also known as the seafloor, sea floor, ocean floor, and ocean bottom) is the bottom of the ocean. All floors of the ocean are known as 'seabeds'. The structure of the seabed of the global ocean is governed by plate tectonics. Most of ...
in all the world's
ocean The ocean (also the sea or the world ocean) is the body of salt water that covers approximately 70.8% of the surface of Earth and contains 97% of Earth's water. An ocean can also refer to any of the large bodies of water into which the wo ...
s, from warm, tropical zones to frigid,
polar regions The polar regions, also called the frigid zones or polar zones, of Earth are the regions of the planet that surround its geographical poles (the North and South Poles), lying within the polar circles. These high latitudes are dominated by floa ...
. They are found from the intertidal zone down to abyssal depths, at below the surface. Starfish are
marine invertebrates Marine invertebrates are the invertebrates that live in marine habitats. Invertebrate is a blanket term that includes all animals apart from the vertebrate members of the chordate phylum. Invertebrates lack a vertebral column, and some have ev ...
. They typically have a central disc and usually five arms, though some species have a larger number of arms. The aboral or upper surface may be smooth, granular or spiny, and is covered with overlapping plates. Many species are brightly coloured in various shades of red or orange, while others are blue, grey or brown. Starfish have
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 ...
operated by a
hydraulic system Hydraulics (from Greek: Υδραυλική) is a technology and applied science using engineering, chemistry, and other sciences involving the mechanical properties and use of liquids. At a very basic level, hydraulics is the liquid coun ...
and a mouth at the centre of the oral or lower surface. They are
opportunistic Opportunism is the practice of taking advantage of circumstances – with little regard for principles or with what the consequences are for others. Opportunist actions are expedient actions guided primarily by self-interested motives. The term ...
feeders and are mostly predators on benthic invertebrates. Several species have specialized feeding behaviours including eversion of their stomachs and
suspension feeding Filter feeders are a sub-group of suspension feeding animals that feed by straining suspended matter and food particles from water, typically by passing the water over a specialized filtering structure. Some animals that use this method of feedin ...
. They have complex life cycles and can reproduce both sexually and asexually. Most can regenerate damaged parts or lost arms and they can shed arms as a means of defense. The Asteroidea occupy several significant ecological roles. Starfish, such as the
ochre sea star ''Pisaster ochraceus'', generally known as the purple sea star, ochre sea star, or ochre starfish, is a common seastar found among the waters of the Pacific Ocean. Identified as a keystone species, ''P. ochraceus'' is considered an important ind ...
(''Pisaster ochraceus'') and the reef sea star (''Stichaster australis''), have become widely known as examples of the keystone species concept in ecology. The tropical crown-of-thorns starfish (''Acanthaster planci'') is a voracious predator of
coral Corals are marine invertebrates within the class Anthozoa of the phylum Cnidaria. They typically form compact colonies of many identical individual polyps. Coral species include the important reef builders that inhabit tropical oceans and ...
throughout the Indo-Pacific region, and the northern Pacific sea star is considered to be one of the world's 100 worst invasive species. The
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
record for starfish is ancient, dating back to the
Ordovician The Ordovician ( ) is a geologic period and system, the second of six periods of the Paleozoic Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start of the Silurian Period Mya. T ...
around 450 million years ago, but it is rather sparse, as starfish tend to disintegrate after death. Only the ossicles and spines of the animal are likely to be preserved, making remains hard to locate. With their appealing symmetrical shape, starfish have played a part in literature, legend, design and popular culture. They are sometimes collected as curios, used in design or as logos, and in some cultures, despite possible toxicity, they are eaten.


Anatomy

Most starfish have five arms that radiate from a central disc, but the number varies with the group. Some species have six or seven arms and others have 10–15 arms. The Antarctic '' Labidiaster annulatus'' can have over fifty.


Body wall

The body wall consists of a thin cuticle, an epidermis consisting of a single layer of cells, a thick dermis formed of connective tissue and a thin coelomic myoepithelial layer, which provides the longitudinal and circular musculature. The dermis contains 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 ...
of calcium carbonate components known as ossicles. These are honeycombed structures composed of calcite microcrystals arranged in a lattice.Ruppert et al., 2004. p. 876 They vary in form, with some bearing external granules, tubercles and spines, but most are tabular plates that fit neatly together in a tessellated manner and form the main covering of the aboral surface. Some are specialised structures such as the
madreporite The madreporite is a light colored calcareous opening used to filter water into the water vascular system of echinoderms. It acts like a pressure-equalizing valve. It is visible as a small red or yellow button-like structure, looking like a sma ...
(the entrance to the water vascular system),
pedicellaria A pedicellaria (plural: pedicellariae) is a small wrench- or claw-shaped appendage with movable jaws, called valves, commonly found on echinoderms (phylum Echinodermata), particularly in sea stars (class Asteroidea) and sea urchins (class Echi ...
e and paxillae. Pedicellariae are compound ossicles with forceps-like jaws. They remove debris from the body surface and wave around on flexible stalks in response to physical or chemical stimuli while continually making biting movements. They often form clusters surrounding spines. Paxillae are umbrella-like structures found on starfish that live buried in sediment. The edges of adjacent paxillae meet to form a false cuticle with a water cavity beneath in which the madreporite and delicate gill structures are protected. All the ossicles, including those projecting externally, are covered by the epidermal layer. Several groups of starfish, including
Valvatida The Valvatida are an order of starfish in the class Asteroidea, which contains 695 species in 172 genera in 17 families. Description The order encompasses both tiny species, which are only a few millimetres in diameter, like those in the genu ...
and
Forcipulatida The Forcipulatida are an order of sea stars, containing three families and 49 genera. Description Forcipulatids share with the brisingid sea stars distinctive pedicellariae, consisting of a short stalk with three skeletal ossicles. Unlike that ...
, possess
pedicellariae A pedicellaria (plural: pedicellariae) is a small wrench- or claw-shaped appendage with movable jaws, called valves, commonly found on echinoderms (phylum Echinodermata), particularly in sea stars (class Asteroidea) and sea urchins (class Echinoi ...
. In Forcipulatida, such as ''
Asterias ''Asterias'' is a genus of the Asteriidae family of sea stars. It includes several of the best-known species of sea stars, including the (Atlantic) common starfish, ''Asterias rubens'', and the northern Pacific seastar, ''Asterias amurensis''. Th ...
'' and '' Pisaster'', they occur in
pompom A pom-pom – also spelled pom-pon, pompom or pompon – is a decorative ball or tuft of fibrous material. The term may refer to large tufts used by cheerleaders, or a small, tighter ball attached to the top of a hat, also known as a ...
-like tufts at the base of each spine, whereas in the Goniasteridae, such as '' Hippasteria phrygiana'', the pedicellariae are scattered over the body surface. Some are thought to assist in defence, while others aid in feeding or in the removal of organisms attempting to settle on the starfish's surface. Some species like ''Labidiaster annulatus'', '' Rathbunaster californicus'' and '' Novodinia antillensis'' use their large pedicellariae to capture small fish and crustaceans. There may also be
papula Papulae (sing. papula; also occasionally papulla, papullae), also known as dermal branchiae or skin gills, are projections of the coelom of Asteroidea that serve in respiration and waste removal. Papulae are soft, covered externally with the epid ...
e, thin-walled protrusions of the body cavity that reach through the body wall and extend into the surrounding water. These serve a
respiratory The respiratory system (also respiratory apparatus, ventilatory system) is a biological system consisting of specific organs and structures used for gas exchange in animals and plants. The anatomy and physiology that make this happen varies gre ...
function. The structures are supported by collagen fibres set at right angles to each other and arranged in a three-dimensional web with the ossicles and papulae in the
interstices An interstitial space or interstice is a space between structures or objects. In particular, interstitial may refer to: Biology * Interstitial cell tumor * Interstitial cell, any cell that lies between other cells * Interstitial collagenase, ...
. This arrangement enables both easy flexion of the arms by the starfish and the rapid onset of stiffness and rigidity required for actions performed under stress.


Water vascular system

The water
vascular The blood vessels are the components of the circulatory system that transport blood throughout the human body. These vessels transport blood cells, nutrients, and oxygen to the tissues of the body. They also take waste and carbon dioxide away f ...
system of the starfish is a
hydraulic system Hydraulics (from Greek: Υδραυλική) is a technology and applied science using engineering, chemistry, and other sciences involving the mechanical properties and use of liquids. At a very basic level, hydraulics is the liquid coun ...
made up of a network of fluid-filled canals and is concerned with locomotion, adhesion, food manipulation and
gas exchange Gas exchange is the physical process by which gases move passively by diffusion across a surface. For example, this surface might be the air/water interface of a water body, the surface of a gas bubble in a liquid, a gas-permeable membrane, or a ...
. Water enters the system through the
madreporite The madreporite is a light colored calcareous opening used to filter water into the water vascular system of echinoderms. It acts like a pressure-equalizing valve. It is visible as a small red or yellow button-like structure, looking like a sma ...
, a porous, often conspicuous, sieve-like ossicle on the aboral surface. It is linked through a stone canal, often lined with calcareous material, to a ring canal around the mouth opening. A set of radial canals leads off this; one radial canal runs along the
ambulacral Ambulacral is a term typically used in the context of anatomical parts of the phylum Echinodermata or class Asteroidea and Edrioasteroidea. Echinoderms can have ambulacral parts that include ossicles, plates, spines, and suckers. For example, sea ...
groove in each arm. There are short lateral canals branching off alternately to either side of the radial canal, each ending in an ampulla. These bulb-shaped organs are joined to tube feet (podia) on the exterior of the animal by short linking canals that pass through ossicles in the ambulacral groove. There are usually two rows of tube feet but in some species, the lateral canals are alternately long and short and there appear to be four rows. The interior of the whole canal system is lined with cilia.Ruppert et al., 2004. pp. 879–883 When longitudinal muscles in the ampullae contract, valves in the lateral canals close and water is forced into the tube feet. These extend to contact the substrate. Although the tube feet resemble suction cups in appearance, the gripping action is a function of adhesive chemicals rather than suction. Other chemicals and relaxation of the ampullae allow for release from the substrate. The tube feet latch on to surfaces and move in a wave, with one arm section attaching to the surface as another releases. Some starfish turn up the tips of their arms while moving which gives maximum exposure of the sensory tube feet and the eyespot to external stimuli. Having descended from
bilateral Bilateral may refer to any concept including two sides, in particular: *Bilateria, bilateral animals *Bilateralism, the political and cultural relations between two states *Bilateral, occurring on both sides of an organism ( Anatomical terms of l ...
organisms, starfish may move in a bilateral fashion, particularly when hunting or in danger. When crawling, certain arms act as the leading arms, while others trail behind. Most starfish cannot move quickly, a typical speed being that of the leather star (''Dermasterias imbricata''), which can manage just in a minute. Some burrowing species from the genera ''
Astropecten ''Astropecten'' is a genus of sea stars of the family Astropectinidae. Identification These sea stars are similar one to each other and it can be difficult to determine with certainty the species only from a photograph. To have a certain d ...
'' and '' Luidia'' have points rather than suckers on their long
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 ...
and are capable of much more rapid motion, "gliding" across the ocean floor. The sand star ('' Luidia foliolata'') can travel at a speed of per minute. When a starfish finds itself upside down, two adjacent arms are bent backwards to provide support, the opposite arm is used to stamp the ground while the two remaining arms are raised on either side; finally the stamping arm is released as the starfish turns itself over and recovers its normal stance. Apart from their function in locomotion, the tube feet act as accessory gills. The water vascular system serves to transport
oxygen Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements as ...
from, and carbon dioxide to, the tube feet and also nutrients from the gut to the muscles involved in locomotion. Fluid movement is bidirectional and initiated by cilia. Gas exchange also takes place through other
gills A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they are ...
known as papulae, which are thin-walled bulges on the aboral surface of the disc and arms. Oxygen is transferred from these to the coelomic fluid, which acts as the transport medium for gasses. Oxygen dissolved in the water is distributed through the body mainly by the fluid in the main body cavity; the circulatory system may also play a minor role.


Digestive system and excretion

The gut of a starfish occupies most of the disc and extends into the arms. The mouth is located in the centre of the oral surface, where it is surrounded by a tough peristomial membrane and closed with a
sphincter A sphincter is a circular muscle that normally maintains constriction of a natural body passage or orifice and which relaxes as required by normal physiological functioning. Sphincters are found in many animals. There are over 60 types in the h ...
. The mouth opens through a short
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 th ...
into a
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 ...
divided by a constriction into a larger, eversible cardiac portion and a smaller pyloric portion. The cardiac stomach is glandular and pouched, and is supported by ligaments attached to ossicles in the arms so it can be pulled back into position after it has been everted. The pyloric stomach has two extensions into each arm: the pyloric caeca. These are elongated, branched hollow tubes that are lined by a series of glands, which secrete digestive
enzyme Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as products ...
s and absorb nutrients from the food. A short intestine and rectum run from the pyloric stomach to open at a small anus at the apex of the aboral surface of the disc.Ruppert et al., 2004. p. 885 Primitive starfish, such as ''Astropecten'' and ''Luidia'', swallow their
prey Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill ...
whole, and start to digest it in their cardiac stomachs. Shell valves and other inedible materials are ejected through their mouths. The semi-digested fluid is passed into their pyloric stomachs and caeca where digestion continues and absorption ensues. In more advanced species of starfish, the cardiac stomach can be everted from the organism's body to engulf and digest food. When the prey is a clam or other bivalve, the starfish pulls with its tube feet to separate the two valves slightly, and inserts a small section of its stomach, which releases enzymes to digest the prey. The stomach and the partially digested prey are later retracted into the disc. Here the food is passed on to the pyloric stomach, which always remains inside the disc. The retraction and contraction of the cardiac stomach is activated by a
neuropeptide Neuropeptides are chemical messengers made up of small chains of amino acids that are synthesized and released by neurons. Neuropeptides typically bind to G protein-coupled receptors (GPCRs) to modulate neural activity and other tissues like the ...
known as NGFFYamide. Because of this ability to digest food outside the body, starfish can hunt prey much larger than their mouths. Their diets include clams and
oyster Oyster is the common name for a number of different families of salt-water bivalve molluscs that live in marine or brackish habitats. In some species, the valves are highly calcified, and many are somewhat irregular in shape. Many, but not ...
s,
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
s, small
fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of ...
and gastropod molluscs. Some starfish are not pure
carnivore A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other s ...
s, supplementing their diets with algae or organic detritus. Some of these species are grazers, but others trap food particles from the water in sticky
mucus Mucus ( ) is a slippery aqueous secretion produced by, and covering, mucous membranes. It is typically produced from cells found in mucous glands, although it may also originate from mixed glands, which contain both serous and mucous cells. It ...
strands that are swept towards the mouth along ciliated grooves. The main nitrogenous waste product is
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 wa ...
. Starfish have no distinct excretory organs; waste ammonia is removed by diffusion through the tube feet and papulae. The body fluid contains
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 c ...
cells called coelomocytes, which are also found within the hemal and water vascular systems. These cells engulf waste material, and eventually migrate to the tips of the papulae, where a portion of body wall is nipped off and ejected into the surrounding water. Some waste may also be excreted by the pyloric glands and voided with the
faeces Feces ( or faeces), known colloquially and in slang as poo and poop, are the solid or semi-solid remains of food that was not digested in the small intestine, and has been broken down by bacteria in the large intestine. Feces contain a relati ...
.Ruppert et al., 2004. pp. 886–887 Starfish do not appear to have any mechanisms for
osmoregulation Osmoregulation is the active regulation of the osmotic pressure of an organism's body fluids, detected by osmoreceptors, to maintain the homeostasis of the organism's water content; that is, it maintains the fluid balance and the concentration o ...
, and keep their body fluids at the same salt concentration as the surrounding water. Although some species can tolerate relatively low salinity, the lack of an osmoregulation system probably explains why starfish are not found in fresh water or even in many estuarine environments.


Sensory and nervous systems

Although starfish do not have many well-defined sense organs, they are sensitive to touch, light, temperature, orientation and the status of the water around them. The tube feet, spines and pedicellariae are sensitive to touch. The tube feet, especially those at the tips of the rays, are also sensitive to chemicals, enabling the starfish to detect odour sources such as food. There are eyespots at the ends of the arms, each one made of 80–200 simple
ocelli A simple eye (sometimes called a pigment pit) refers to a form of eye or an optical arrangement composed of a single lens and without an elaborate retina such as occurs in most vertebrates. In this sense "simple eye" is distinct from a multi-l ...
. These are composed of
pigment A pigment is a colored material that is completely or nearly insoluble in water. In contrast, dyes are typically soluble, at least at some stage in their use. Generally dyes are often organic compounds whereas pigments are often inorganic compou ...
ed epithelial cells that respond to light and are covered by a thick, transparent cuticle that both protects the ocelli and acts to focus light. Many starfish also possess individual
photoreceptor cells A photoreceptor cell is a specialized type of neuroepithelial cell found in the retina that is capable of visual phototransduction. The great biological importance of photoreceptors is that they convert light (visible electromagnetic radiatio ...
in other parts of their bodies and respond to light even when their eyespots are covered. Whether they advance or retreat depends on the species.Ruppert et al., 2004. pp. 883–884 While a starfish lacks a centralized brain, it has a complex
nervous system In biology, the nervous system is the highly complex part of an animal that coordinates its actions and sensory information by transmitting signals to and from different parts of its body. The nervous system detects environmental changes ...
with a nerve ring around the mouth and a radial nerve running along the ambulacral region of each arm parallel to the radial canal. The peripheral nerve system consists of two nerve nets: a sensory system in the epidermis and a motor system in the lining of the coelomic cavity. Neurons passing through the dermis connect the two. The ring nerves and radial nerves have sensory and motor components and coordinate the starfish's balance and directional systems. The sensory component receives input from the sensory organs while the motor nerves control the tube feet and musculature. The starfish does not have the capacity to plan its actions. If one arm detects an attractive odour, it becomes dominant and temporarily over-rides the other arms to initiate movement towards the prey. The mechanism for this is not fully understood.


Circulatory system

The body cavity contains the circulatory or haemal system. The vessels form three rings: one around the mouth (the hyponeural haemal ring), another around the digestive system (the gastric ring) and the third near the aboral surface (the genital ring). The heart beats about six times a minute and is at the apex of a vertical channel (the axial vessel) that connects the three rings. At the base of each arm are paired gonads; a lateral vessel extends from the genital ring past the gonads to the tip of the arm. This vessel has a blind end and there is no continuous circulation of the fluid within it. This liquid does not contain a pigment and has little or no respiratory function but is probably used to transport nutrients around the body.Ruppert et al., 2004. p. 886


Secondary metabolites

Starfish produce a large number of
secondary metabolite Secondary metabolites, also called specialised metabolites, toxins, secondary products, or natural products, are organic compounds produced by any lifeform, e.g. bacteria, fungi, animals, or plants, which are not directly involved in the norma ...
s in the form of
lipid Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids includ ...
s, including steroidal derivatives of
cholesterol Cholesterol is any of a class of certain organic molecules called lipids. It is a sterol (or modified steroid), a type of lipid. Cholesterol is biosynthesized by all animal cells and is an essential structural component of animal cell mem ...
, and
fatty acid In chemistry, particularly in biochemistry, a fatty acid is a carboxylic acid with an aliphatic chain, which is either saturated or unsaturated. Most naturally occurring fatty acids have an unbranched chain of an even number of carbon atoms, ...
amide In organic chemistry, an amide, also known as an organic amide or a carboxamide, is a compound with the general formula , where R, R', and R″ represent organic groups or hydrogen atoms. The amide group is called a peptide bond when it i ...
s of
sphingosine Sphingosine (2-amino-4-trans-octadecene-1,3-diol) is an 18-carbon amino alcohol with an unsaturated hydrocarbon chain, which forms a primary part of sphingolipids, a class of cell membrane lipids that include sphingomyelin, an important phos ...
. The steroids are mostly saponins, known as asterosaponins, and their sulphated derivatives. They vary between species and are typically formed from up to six sugar molecules (usually
glucose Glucose is a simple sugar with the molecular formula . Glucose is overall the most abundant monosaccharide, a subcategory of carbohydrates. Glucose is mainly made by plants and most algae during photosynthesis from water and carbon dioxide, u ...
and
galactose Galactose (, '' galacto-'' + ''-ose'', "milk sugar"), sometimes abbreviated Gal, is a monosaccharide sugar that is about as sweet as glucose, and about 65% as sweet as sucrose. It is an aldohexose and a C-4 epimer of glucose. A galactose molecu ...
) connected by up to three
glycosidic A glycosidic bond or glycosidic linkage is a type of covalent bond that joins a carbohydrate (sugar) molecule to another group, which may or may not be another carbohydrate. A glycosidic bond is formed between the hemiacetal or hemiketal group ...
chains. Long-chain fatty acid amides of sphingosine occur frequently and some of them have known
pharmacological activity In pharmacology, biological activity or pharmacological activity describes the beneficial or adverse effects of a drug on living matter. When a drug is a complex chemical mixture, this activity is exerted by the substance's active ingredient or ...
. Various
ceramide Ceramides are a family of waxy lipid molecules. A ceramide is composed of N-acetyl sphingosine and a fatty acid. Ceramides are found in high concentrations within the cell membrane of eukaryotic cells, since they are component lipids that make ...
s are also known from starfish and a small number of
alkaloid Alkaloids are a class of basic, naturally occurring organic compounds that contain at least one nitrogen atom. This group also includes some related compounds with neutral and even weakly acidic properties. Some synthetic compounds of similar ...
s have also been identified. The functions of these chemicals in the starfish have not been fully investigated but most have roles in defence and communication. Some are feeding deterrents used by the starfish to discourage predation. Others are antifoulants and supplement the pedicellariae to prevent other organisms from settling on the starfish's aboral surface. Some are alarm
pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
s and escape-eliciting chemicals, the release of which trigger responses in
conspecific Biological specificity is the tendency of a characteristic such as a behavior or a biochemical variation to occur in a particular species. Biochemist Linus Pauling stated that "Biological specificity is the set of characteristics of living organis ...
starfish but often produce escape responses in potential prey. Research into the efficacy of these compounds for possible pharmacological or industrial use occurs worldwide.


Life cycle


Sexual reproduction

Most species of starfish are gonochorous, there being separate male and female individuals. These are usually not distinguishable externally as the gonads cannot be seen, but their sex is apparent when they
spawn Spawn or spawning may refer to: * Spawn (biology), the eggs and sperm of aquatic animals Arts, entertainment, and media * Spawn (character), a fictional character in the comic series of the same name and in the associated franchise ** '' Spawn: ...
. Some species are simultaneous hermaphrodites, producing eggs and sperm at the same time and in a few of these, the same gonad, called an
ovotestis An ovotestis is a gonad with both testicular and ovarian aspects. In humans, ovotestes are an infrequent anatomical variation associated with gonadal dysgenesis. The only mammals where ovotestes are not symptomatic of an intersex variation are mole ...
, produces both eggs and sperm. Other starfish are sequential hermaphrodites. Protandrous individuals of species like '' Asterina gibbosa'' start life as males before changing sex into females as they grow older. In some species such as ''
Nepanthia belcheri ''Nepanthia belcheri'' is a species of starfish in the family Asterinidae. It is found in shallow water in Southeast Asia and northeastern Australia. It is an unusual species in that it can reproduce sexually or can split in two by fission to fo ...
'', a large female can split in half and the resulting offspring are males. When these grow large enough they change back into females. Each starfish arm contains two gonads that release
gametes 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 ...
through openings called gonoducts, located on the central disc between the arms.
Fertilization Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a new individual organism or offspring and initiate its development. Proce ...
is generally external but in a few species, internal fertilization takes place. In most species, the buoyant eggs and sperm are simply released into the water (free spawning) and the resulting embryos and
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. ...
e live as part of the
plankton Plankton are the diverse collection of organisms found in water (or air) that are unable to propel themselves against a current (or wind). The individual organisms constituting plankton are called plankters. In the ocean, they provide a crucia ...
. In others, the eggs may be stuck to the undersides of rocks. In certain species of starfish, the females
brood Brood may refer to: Nature * Brood, a collective term for offspring * Brooding, the incubation of bird eggs by their parents * Bee brood, the young of a beehive * Individual broods of North American Periodical Cicadas: ** Brood X, the largest b ...
their eggs – either by simply enveloping them or by holding them in specialised structures. Brooding may be done in pockets on the starfish's aboral surface, inside the pyloric stomach ('' Leptasterias tenera'') or even in the interior of the gonads themselves. Those starfish that brood their eggs by "sitting" on them usually assume a humped posture with their discs raised off the substrate. '' Pteraster militaris'' broods a few of its young and disperses the remaining eggs, that are too numerous to fit into its pouch. In these brooding species, the eggs are relatively large, and supplied with
yolk Among animals which produce eggs, the yolk (; also known as the vitellus) is the nutrient-bearing portion of the egg whose primary function is to supply food for the development of the embryo. Some types of egg contain no yolk, for example ...
, and they generally develop directly into miniature starfish without an intervening larval stage. The developing young are called lecithotrophic because they obtain their nutrition from the yolk as opposed to "planktotrophic" larvae that feed in the
water column A water column is a conceptual column of water from the surface of a sea, river or lake to the bottom sediment.Munson, B.H., Axler, R., Hagley C., Host G., Merrick G., Richards C. (2004).Glossary. ''Water on the Web''. University of Minnesota-D ...
. In '' Parvulastra parvivipara'', an intragonadal brooder, the young starfish obtain nutrients by eating other eggs and embryos in the brood pouch. Brooding is especially common in polar and deep-sea species that live in environments unfavourable for larval developmentRuppert et al., 2004. pp. 887–888 and in smaller species that produce just a few eggs. In the tropics, a plentiful supply of phytoplankton is continuously available for starfish larvae to feed on. Spawning takes place at any time of year, each species having its own characteristic breeding season. In temperate regions, the spring and summer brings an increase in food supplies. The first individual of a species to spawn may release a
pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
that serves to attract other starfish to aggregate and to release their gametes synchronously. In other species, a male and female may come together and form a pair. This behaviour is called
pseudocopulation Pseudocopulation describes behaviors similar to copulation that serve a reproductive function for one or both participants but do not involve actual sexual union between the individuals. It is most generally applied to a pollinator attempting to co ...
and the male climbs on top, placing his arms between those of the female. When she releases eggs into the water, he is induced to spawn. Starfish may use environmental signals to coordinate the time of spawning (day length to indicate the correct time of the year, dawn or dusk to indicate the correct time of day), and chemical signals to indicate their readiness to breed. In some species, mature females produce chemicals to attract sperm in the sea water.


Larval development

Most starfish embryos hatch at the
blastula Blastulation is the stage in early animal embryonic development that produces the blastula. In mammalian development the blastula develops into the blastocyst with a differentiated inner cell mass and an outer trophectoderm. The blastula (f ...
stage. The original ball of cells develops a lateral pouch, the
archenteron The primary gut that forms during gastrulation in a developing embryo is known as the archenteron, the gastrocoel or the primitive digestive tube. It develops into the endoderm and mesoderm of an animal. Formation in sea urchins As primary mesen ...
. The entrance to this is known as the blastopore and it will later develop into the anus—together with
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 fi ...
s,
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 s ...
s are deuterostomes, meaning the second (''deutero'') invagination becomes the mouth (''stome''); members of all other phyla are
protostome Protostomia () is the clade of animals once thought to be characterized by the formation of the organism's mouth before its anus during embryogenesis, embryonic development. This nature has since been discovered to be extremely variable among Pro ...
s, and their first invagination becomes the mouth. Another invagination of the surface will fuse with the tip of the archenteron as the mouth while the interior section will become the gut. At the same time, a band of cilia develops on the exterior. This enlarges and extends around the surface and eventually onto two developing arm-like outgrowths. At this stage the larva is known as a
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 ...
. The cilia are used for locomotion and feeding, their rhythmic beat wafting phytoplankton towards the mouth. The next stage in development is a brachiolaria larva and involves the growth of three short, additional arms. These are at the anterior end, surround a sucker and have adhesive cells at their tips. Both bipinnaria and brachiolaria larvae are bilaterally symmetrical. When fully developed, the brachiolaria settles on the seabed and attaches itself with a short stalk formed from the ventral arms and sucker. Metamorphosis now takes place with a radical rearrangement of tissues. The left side of the larval body becomes the oral surface of the juvenile and the right side the aboral surface. Part of the gut is retained, but the mouth and anus move to new positions. Some of the body cavities degenerate but others become the water vascular system and the visceral coelom. The starfish is now pentaradially symmetrical. It casts off its stalk and becomes a free-living juvenile starfish about in diameter. Starfish of the order Paxillosida have no brachiolaria stage, with the bipinnaria larvae settling on the seabed and developing directly into juveniles.


Asexual reproduction

Some species of starfish are able to reproduce asexually as adults either by fission of their central discs or by
autotomy Autotomy (from the Greek ''auto-'', "self-" and ''tome'', "severing", αὐτοτομία) or self-amputation, is the behaviour whereby an animal sheds or discards one or more of its own appendages, usually as a self-defense mechanism to elude ...
of one or more of their arms. Which of these processes occurs depends on the genus. Among starfish that are able to regenerate their whole body from a single arm, some can do so even from fragments just long. Single arms that regenerate a whole individual are called comet forms. The division of the starfish, either across its disc or at the base of the arm, is usually accompanied by a weakness in the structure that provides a fracture zone. The larvae of several species of starfish can reproduce asexually before they reach maturity. They do this by autotomising some parts of their bodies or by
budding Budding or blastogenesis is a type of asexual reproduction in which a new organism develops from an outgrowth or bud due to cell division at one particular site. For example, the small bulb-like projection coming out from the yeast cell is kno ...
. When such a larva senses that food is plentiful, it takes the path of asexual reproduction rather than normal development. Though this costs it time and energy and delays maturity, it allows a single larva to give rise to multiple adults when the conditions are appropriate.


Regeneration

Some species of starfish have the ability to regenerate lost arms and can regrow an entire new limb given time. A few can regrow a complete new disc from a single arm, while others need at least part of the central disc to be attached to the detached part. Regrowth can take several months or years, and starfish are vulnerable to infections during the early stages after the loss of an arm. A separated limb lives off stored nutrients until it regrows a disc and mouth and is able to feed again. Other than fragmentation carried out for the purpose of reproduction, the division of the body may happen inadvertently due to part being detached by a predator, or part may be actively shed by the starfish in an escape response. The loss of parts of the body is achieved by the rapid softening of a special type of connective tissue in response to nervous signals. This type of tissue is called catch connective tissue and is found in most echinoderms. An autotomy-promoting factor has been identified which, when injected into another starfish, causes rapid shedding of arms.


Lifespan

The lifespan of a starfish varies considerably between species, generally being longer in larger forms and in those with planktonic larvae. For example, '' Leptasterias hexactis'' broods a small number of large-yolked eggs. It has an adult weight of , reaches sexual maturity in two years and lives for about ten years. '' Pisaster ochraceus'' releases a large number of eggs into the sea each year and has an adult weight of up to . It reaches maturity in five years and has a maximum recorded lifespan of 34 years.


Ecology


Distribution and habitat

Echinoderms, including starfish, maintain a delicate internal electrolyte balance that is in equilibrium with sea water, making it impossible for them to live in a freshwater habitat. Starfish species inhabit all of the world's oceans. Habitats range from tropical
coral reef A coral reef is an underwater ecosystem characterized by reef-building corals. Reefs are formed of colonies of coral polyps held together by calcium carbonate. Most coral reefs are built from stony corals, whose polyps cluster in groups. C ...
s, rocky shores,
tidal pool A tide pool or rock pool is a shallow pool of seawater that forms on the rocky intertidal shore. Many of these pools exist as separate bodies of water only at low tide. Many tide pool habitats are home to especially adaptable animals that ...
s, mud, and sand to
kelp forest Kelp forests are underwater areas with a high density of kelp, which covers a large part of the world's coastlines. Smaller areas of anchored kelp are called kelp beds. They are recognized as one of the most productive and dynamic ecosystems on Ea ...
s,
seagrass meadow A seagrass meadow or seagrass bed is an underwater ecosystem formed by seagrasses. Seagrasses are marine (saltwater) plants found in shallow coastal waters and in the brackish waters of estuaries. Seagrasses are flowering plants with stems and ...
s and the deep-sea floor down to at least . The greatest diversity of species occurs in coastal areas.


Diet

Most species are generalist predators, eating microalgae,
sponge Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate throug ...
s, bivalves,
snail A snail is, in loose terms, a shelled gastropod. The name is most often applied to land snails, terrestrial pulmonate gastropod molluscs. However, the common name ''snail'' is also used for most of the members of the molluscan class G ...
s and other small animals. The crown-of-thorns starfish consumes
coral Corals are marine invertebrates within the class Anthozoa of the phylum Cnidaria. They typically form compact colonies of many identical individual polyps. Coral species include the important reef builders that inhabit tropical oceans and ...
polyps, while other species are detritivores, feeding on decomposing organic material and faecal matter. in Lawrence (2013) A few are suspension feeders, gathering in phytoplankton; ''
Henricia ''Henricia'' is a large genus of slender-armed sea stars belonging to the family Echinasteridae. It contains about fifty species. The sea stars from this genus are ciliary suspension-feeders, filtering phytoplankton. Species According to the Wo ...
'' and '' Echinaster'' often occur in association with sponges, benefiting from the water current they produce. Various species have been shown to be able to absorb organic nutrients from the surrounding water, and this may form a significant portion of their diet. The processes of feeding and capture may be aided by special parts; ''
Pisaster brevispinus ''Pisaster brevispinus'', commonly called the pink sea star, giant pink sea star, or short-spined sea star, is a species of sea star in the northeast Pacific Ocean. It was first described to science by William Stimson in 1857. The type specime ...
'', the short-spined pisaster from the West Coast of America, can use a set of specialized tube feet to dig itself deep into the soft substrate to extract prey (usually
clam Clam is a common name for several kinds of bivalve molluscs. The word is often applied only to those that are edible and live as infauna, spending most of their lives halfway buried in the sand of the seafloor or riverbeds. Clams have two shel ...
s). Grasping the shellfish, the starfish slowly pries open the prey's shell by wearing out its adductor muscle, and then inserts its everted stomach into the crack to digest the soft tissues. The gap between the valves need only be a fraction of a millimetre wide for the stomach to gain entry. Cannibalism has been observed in juvenile sea stars as early as four days after metamorphosis.


Ecological impact

Starfish are keystone species in their respective marine communities. Their relatively large sizes, diverse diets and ability to adapt to different environments makes them ecologically important. The term "keystone species" was in fact first used by Robert Paine in 1966 to describe a starfish, ''Pisaster ochraceus''. When studying the low intertidal coasts of
Washington state Washington (), officially the State of Washington, is a state in the Pacific Northwest region of the Western United States. Named for George Washington—the first U.S. president—the state was formed from the western part of the Washington ...
, Paine found that predation by ''P. ochraceus'' was a major factor in the diversity of species. Experimental removals of this top predator from a stretch of shoreline resulted in lower species diversity and the eventual domination of '' Mytilus'' mussels, which were able to outcompete other organisms for space and resources. Similar results were found in a 1971 study of '' Stichaster australis'' on the intertidal coast of the South Island 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 ...
. ''S. australis'' was found to have removed most of a batch of transplanted mussels within two or three months of their placement, while in an area from which ''S. australis'' had been removed, the mussels increased in number dramatically, overwhelming the area and threatening
biodiversity Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic (''genetic variability''), species (''species diversity''), and ecosystem (''ecosystem diversity'') l ...
. The feeding activity of the
omnivorous An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nut ...
starfish '' Oreaster reticulatus'' on sandy and seagrass bottoms in the
Virgin Islands The Virgin Islands ( es, Islas Vírgenes) are an archipelago in the Caribbean Sea. They are geologically and biogeographically the easternmost part of the Greater Antilles, the northern islands belonging to the Puerto Rico Trench and St. Cro ...
appears to regulate the diversity, distribution and abundance of microorganisms. These starfish engulf piles of sediment removing the surface films and algae adhering to the particles. Organisms that dislike this disturbance are replaced by others better able to rapidly recolonise "clean" sediment. In addition, foraging by these migratory starfish creates diverse patches of organic matter, which may play a role in the distribution and abundance of organisms such as fish, crabs and sea urchins that feed on the sediment. Starfish sometimes have negative effects on ecosystems. Outbreaks of crown-of-thorns starfish have caused damage to coral reefs in Northeast Australia and French Polynesia. A study in Polynesia found that coral cover declined drastically with the arrival of migratory starfish in 2006, dropping from 50% to under 5% in three years. This had a cascading effect on the whole benthic community and reef-feeding fish. '' Asterias amurensis'' is one of a few echinoderm invasive species. Its larvae likely arrived in
Tasmania ) , nickname = , image_map = Tasmania in Australia.svg , map_caption = Location of Tasmania in AustraliaCoordinates: , subdivision_type = Country , subdi ...
from central Japan via water discharged from ships in the 1980s. The species has since grown in numbers to the point where they threaten commercially important bivalve populations. As such, they are considered pests, in Lawrence (2013) and are on the Invasive Species Specialist Group's
list of the world's 100 worst invasive species 100 of the World's Worst Invasive Alien Species is a list of invasive species compiled in 2014 by the Global Invasive Species Database, a database of invasive species around the world. The database is run by the Invasive Species Specialist Group ...
. Sea Stars (starfish) are the main predators of kelp-eating sea urchins. Satellite imagery shows that sea urchin populations have exploded due to starfish mass deaths, and that by 2021, sea urchins have destroyed 95% of California's kelp forests.


Threats

Starfish may be preyed on by conspecifics, sea anemones, other starfish species, tritons, crabs, fish,
gull Gulls, or colloquially seagulls, are seabirds of the family Laridae in the suborder Lari. They are most closely related to the terns and skimmers and only distantly related to auks, and even more distantly to waders. Until the 21st century ...
s and sea otters. in Lawrence (2013) in Lawrence (2013) in Lawrence (2013) Their first lines of defence are the saponins present in their body walls, which have unpleasant flavours. Some starfish such as '' Astropecten polyacanthus'' also include powerful toxins such as
tetrodotoxin Tetrodotoxin (TTX) is a potent neurotoxin. Its name derives from Tetraodontiformes, an order that includes pufferfish, porcupinefish, ocean sunfish, and triggerfish; several of these species carry the toxin. Although tetrodotoxin was discovere ...
among their chemical armoury, and the slime star can ooze out large quantities of repellent mucus. They also have body armour in the form of hard plates and spines. The crown-of-thorns starfish is particularly unattractive to potential predators, being heavily defended by sharp spines, laced with toxins and sometimes with bright warning colours. Other species protect their vulnerable tube feet and arm tips by lining their ambulacral grooves with spines and heavily plating their extremities. Several species sometimes suffer from a
wasting In medicine, wasting, also known as wasting syndrome, refers to the process by which a debilitating disease causes muscle and fat tissue to "waste" away. Wasting is sometimes referred to as "acute malnutrition" because it is believed that epis ...
condition caused by bacteria in the genus ''
Vibrio ''Vibrio'' is a genus of Gram-negative bacteria, possessing a curved-rod (comma) shape, several species of which can cause foodborne infection, usually associated with eating undercooked seafood. Being highly salt tolerant and unable to survive ...
''; however, a more widespread
wasting disease In medicine, wasting, also known as wasting syndrome, refers to the process by which a debilitating disease causes muscle and fat tissue to "waste" away. Wasting is sometimes referred to as "acute malnutrition" because it is believed that epis ...
, causing mass mortalities among starfish, appears sporadically. A paper published in November 2014 revealed the most likely cause of this disease to be a densovirus the authors named sea star-associated densovirus (SSaDV). The protozoan '' Orchitophrya stellarum'' is known to infect the gonads of starfish and damage tissue. Starfish are vulnerable to high temperatures. Experiments have shown that the feeding and growth rates of ''P. ochraceus'' reduce greatly when their body temperatures rise above and that they die when their temperature rises to . This species has a unique ability to absorb seawater to keep itself cool when it is exposed to sunlight by a receding tide. It also appears to rely on its arms to absorb heat, so as to protect the central disc and vital organs like the stomach. Starfish and other echinoderms are sensitive to marine pollution. The common starfish is considered to be a
bioindicator A bioindicator is any species (an indicator species) or group of species whose function, population, or status can reveal the qualitative status of the environment. The most common indicator species are animals. For example, copepods and other sma ...
for marine ecosystems. A 2009 study found that ''P. ochraceus'' is unlikely to be affected by
ocean acidification Ocean acidification is the reduction in the pH value of the Earth’s ocean. Between 1751 and 2021, the average pH value of the ocean surface has decreased from approximately 8.25 to 8.14. The root cause of ocean acidification is carbon dioxid ...
as severely as other marine animals with
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 ad ...
skeletons. In other groups, structures made of calcium carbonate are vulnerable to dissolution when the pH is lowered. Researchers found that when ''P. ochraceus'' were exposed to and 770  ppm
carbon dioxide Carbon dioxide ( chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is trans ...
(beyond rises expected in the next century), they were relatively unaffected. Their survival is likely due to the nodular nature of their skeletons, which are able to compensate for a shortage of carbonate by growing more fleshy tissue.


Evolution


Fossil record

Echinoderms first appeared in the
fossil record A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
in the Cambrian. The first known asterozoans were the Somasteroidea, which exhibit characteristics of both groups. Starfish are infrequently found as fossils, possibly because their hard skeletal components separate as the animal decays. Despite this, there are a few places where accumulations of complete skeletal structures occur, fossilized in place in
Lagerstätte A Lagerstätte (, from ''Lager'' 'storage, lair' '' Stätte'' 'place'; plural ''Lagerstätten'') is a sedimentary deposit that exhibits extraordinary fossils with exceptional preservation—sometimes including preserved soft tissues. These f ...
n – so-called "starfish beds". By the late
Paleozoic The Paleozoic (or Palaeozoic) Era is the earliest of three geologic eras of the Phanerozoic Eon. The name ''Paleozoic'' ( ;) was coined by the British geologist Adam Sedgwick in 1838 by combining the Greek words ''palaiós'' (, "old") and ' ...
, the
crinoids Crinoids are marine animals that make up the class Crinoidea. Crinoids that are attached to the sea bottom by a stalk in their adult form are commonly called sea lilies, while the unstalked forms are called feather stars or comatulids, which are ...
and blastoids were the predominant echinoderms, and some limestones from this period are made almost entirely from fragments from these groups. In the two major extinction events that occurred during the late Devonian and late
Permian The Permian ( ) is a geologic period and System (stratigraphy), stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last ...
, the blastoids were wiped out and only a few species of crinoids survived. Many starfish species also became extinct in these events, but afterwards the surviving few species diversified rapidly within about sixty million years during the
Early Jurassic The Early Jurassic Epoch (geology), Epoch (in chronostratigraphy corresponding to the Lower Jurassic series (stratigraphy), Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic-J ...
and the beginning of the Middle Jurassic. A 2012 study found that speciation in starfish can occur rapidly. During the last 6,000 years, divergence in the larval development of '' Cryptasterina hystera'' and '' Cryptasterina pentagona'' has taken place, the former adopting internal fertilization and brooding and the latter remaining a broadcast spawner.


Diversity

The scientific name Asteroidea was given to starfish by the French zoologist de Blainville in 1830. It is derived from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
''aster'', ἀστήρ (a star) and the Greek ''eidos'', εἶδος (form, likeness, appearance). The class Asteroidea belongs to the phylum
Echinodermata 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 s ...
. As well as the starfish, the echinoderms include sea urchins,
sand dollar Sand dollars (also known as a sea cookie or snapper biscuit in New Zealand, or pansy shell in South Africa) are species of flat, burrowing sea urchins belonging to the order Clypeasteroida. Some species within the order, not quite as flat, are k ...
s,
brittle A material is brittle if, when subjected to stress, it fractures with little elastic deformation and without significant plastic deformation. Brittle materials absorb relatively little energy prior to fracture, even those of high strength. Br ...
and
basket star The Euryalida are an order of brittle stars, which includes large species with either branching arms (called "basket stars") or long and curling arms (called "snake stars"). Characteristics Many of the species in this order have characteristi ...
s, sea cucumbers and
crinoid Crinoids are marine animals that make up the class Crinoidea. Crinoids that are attached to the sea bottom by a stalk in their adult form are commonly called sea lilies, while the unstalked forms are called feather stars or comatulids, which are ...
s. The larvae of echinoderms have bilateral symmetry, but during metamorphosis this is replaced with
radial 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 ...
, typically
pentameric A pentamer is an entity composed of five sub-units. In chemistry, it applies to molecules made of five monomers. In biochemistry, it applies to macromolecules, in particular to pentameric proteins, made of five proteic sub-units. In microbiolo ...
. Adult echinoderms are characterized by having 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 ...
with external
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 ...
and a
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 ad ...
endoskeleton consisting of ossicles connected by a mesh of collagen fibres. Starfish are included in the subphylum Asterozoa, the characteristics of which include a flattened, star-shaped body as adults consisting of a central disc and multiple radiating arms. The subphylum includes the two classes of Asteroidea, the starfish, and
Ophiuroidea Brittle stars, serpent stars, or ophiuroids (; ; referring to the serpent-like arms of the brittle star) are echinoderms in the class Ophiuroidea, closely related to starfish. They crawl across the sea floor using their flexible arms for locomot ...
, the brittle stars and basket stars. Asteroids have broad-based arms with skeletal support provided by calcareous plates in the body wall while ophiuroids have clearly demarcated slender arms strengthened by paired fused ossicles forming jointed "vertebrae". The starfish are a large and diverse class with over 1,900 living species. There are seven extant orders,
Brisingida The Brisingids are deep-sea-dwelling starfish in the order Brisingida. Description These starfish have between 6 to 18 long, attenuated arms which they use for suspension feeding. Other characteristics include a single series of marginals, a fu ...
,
Forcipulatida The Forcipulatida are an order of sea stars, containing three families and 49 genera. Description Forcipulatids share with the brisingid sea stars distinctive pedicellariae, consisting of a short stalk with three skeletal ossicles. Unlike that ...
, Notomyotida,
Paxillosida The Paxillosida are a large order of sea stars. Characteristics Paxillosida adults lack an anus and have no suckers on their tube feet. They do not develop the brachiolaria stage in their early development.Matsubara, M., Komatsu, M., Araki, T., ...
,
Spinulosida The Spinulosida are an order of sea stars containing at least 120 species in seven genera and one family. Spinulosids completely lack pedicellariae and have a delicate skeletal arrangement. Their name comes from the presence of numerous low spi ...
,
Valvatida The Valvatida are an order of starfish in the class Asteroidea, which contains 695 species in 172 genera in 17 families. Description The order encompasses both tiny species, which are only a few millimetres in diameter, like those in the genu ...
and
Velatida The Velatida are an order of sea stars containing about 200 species in five families. These sea stars normally have thick bodies with large discs. Description and characteristics This order contains mostly deep or cold seas sea stars, often wi ...
and two extinct ones, Calliasterellidae and Trichasteropsida. Living asteroids, the Neoasteroidea, are morphologically distinct from their forerunners in the Paleozoic. The taxonomy of the group is relatively stable but there is ongoing debate about the status of the
Paxillosida The Paxillosida are a large order of sea stars. Characteristics Paxillosida adults lack an anus and have no suckers on their tube feet. They do not develop the brachiolaria stage in their early development.Matsubara, M., Komatsu, M., Araki, T., ...
, and the deep-water sea daisies, though clearly Asteroidea and currently included in
Velatida The Velatida are an order of sea stars containing about 200 species in five families. These sea stars normally have thick bodies with large discs. Description and characteristics This order contains mostly deep or cold seas sea stars, often wi ...
, do not fit easily in any accepted lineage. Phylogenetic data suggests that they may be a sister group, the Concentricycloidea, to the Neoasteroidea, or that the Velatida themselves may be a sister group.


Living groups

;
Brisingida The Brisingids are deep-sea-dwelling starfish in the order Brisingida. Description These starfish have between 6 to 18 long, attenuated arms which they use for suspension feeding. Other characteristics include a single series of marginals, a fu ...
(2 families, 17 genera, 111 species) :Species in this order have a small, inflexible disc and 6–20 long, thin arms, which they use for suspension feeding. They have a single series of marginal plates, a fused ring of disc plates, a reduced number of aboral plates, crossed pedicellariae, and several series of long spines on the arms. They live almost exclusively in deep-sea habitats, although a few live in shallow waters in the Antarctic. In some species, the tube feet have rounded tips and lack suckers. ;
Forcipulatida The Forcipulatida are an order of sea stars, containing three families and 49 genera. Description Forcipulatids share with the brisingid sea stars distinctive pedicellariae, consisting of a short stalk with three skeletal ossicles. Unlike that ...
(6 families, 63 genera, 269 species) :Species in this order have distinctive pedicellariae, consisting of a short stalk with three skeletal ossicles. They tend to have robust bodies and have tube feet with flat-tipped suckers usually arranged in four rows. The order includes well-known species from temperate regions, including the common starfish of North Atlantic coasts and rock pools, as well as cold-water and abyssal species. ; Notomyotida (1 family, 8 genera, 75 species) :These starfish are deep-sea dwelling and have particularly flexible arms. The inner dorso-lateral surfaces of the arms contain characteristic longitudinal muscle bands. In some species, the tube feet lack suckers. ;
Paxillosida The Paxillosida are a large order of sea stars. Characteristics Paxillosida adults lack an anus and have no suckers on their tube feet. They do not develop the brachiolaria stage in their early development.Matsubara, M., Komatsu, M., Araki, T., ...
(7 families, 48 genera, 372 species) :This is a primitive order and members do not extrude their stomach when feeding, lack an anus and have no suckers on their tube feet. Papulae are plentiful on their aboral surface and they possess marginal plates and paxillae. They mostly inhabit soft-bottomed areas of sand or mud.Ruppert et al, 2004. pp. 888–889 There is no brachiolaria stage in their larval development. The comb starfish ('' Astropecten polyacanthus'') is a member of this order. ;
Spinulosida The Spinulosida are an order of sea stars containing at least 120 species in seven genera and one family. Spinulosids completely lack pedicellariae and have a delicate skeletal arrangement. Their name comes from the presence of numerous low spi ...
(1 family, 8 genera, 121 species) :Most species in this order lack pedicellariae and all have a delicate skeletal arrangement with small or no marginal plates on the disc and arms. They have numerous groups of short spines on the aboral surface. This group includes the red starfish ''
Echinaster sepositus ''Echinaster sepositus'', the Mediterranean red sea star (sometimes only red sea star, but this name is also used for other species), is a species of starfish from the East Atlantic, including the Mediterranean Sea.Villamor, A.; and M. A. Becerr ...
''. ;
Valvatida The Valvatida are an order of starfish in the class Asteroidea, which contains 695 species in 172 genera in 17 families. Description The order encompasses both tiny species, which are only a few millimetres in diameter, like those in the genu ...
(16 families, 172 genera, 695 species) :Most species in this order have five arms and two rows of tube feet with suckers. There are conspicuous marginal plates on the arms and disc. Some species have paxillae and in some, the main pedicellariae are clamp-like and recessed into the skeletal plates. This group includes the cushion stars, the leather star and the sea daisies. ;
Velatida The Velatida are an order of sea stars containing about 200 species in five families. These sea stars normally have thick bodies with large discs. Description and characteristics This order contains mostly deep or cold seas sea stars, often wi ...
(4 families, 16 genera, 138 species) :This order of starfish consists mostly of deep-sea and other cold-water starfish often with a global distribution. The shape is pentagonal or star-shaped with five to fifteen arms. They mostly have poorly developed skeletons with papulae widely distributed on the aboral surface and often spiny pedicellariae. This group includes the slime star.


Extinct groups

Extinct groups within the Asteroidea include: * † Calliasterellidae, with the type genus '' Calliasterella'' from the Devonian and Carboniferous * † Palastericus, a Devonian genus * † Trichasteropsida, with the
Triassic The Triassic ( ) is a geologic period and system (stratigraphy), system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago (Year#Abbreviations yr and ya, Mya), to the beginning of the Jurassic Period 201.36 ...
genus '' Trichasteropsis'' (at least 2 species)


Phylogeny


External

Starfish are deuterostome animals, like the
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 fi ...
s. A 2014 analysis of 219 genes from all classes of echinoderms gives the following phylogenetic tree. The times at which the clades diverged is shown under the labels in millions of years ago (mya).


Internal

The phylogeny of the Asteroidea has been difficult to resolve, with visible (morphological) features proving inadequate, and the question of whether traditional
taxa In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
are clades in doubt. The phylogeny proposed by Gale in 1987 is: The phylogeny proposed by Blake in 1987 is: Later work making use of molecular evidence, with or without the use of morphological evidence, had by 2000 failed to resolve the argument. In 2011, on further molecular evidence, Janies and colleagues noted that the phylogeny of the echinoderms "has proven difficult", and that "the overall phylogeny of extant echinoderms remains sensitive to the choice of analytical methods". They presented a phylogenetic tree for the living Asteroidea only; using the traditional names of starfish orders where possible, and indicating "part of" otherwise, the phylogeny is shown below. The Solasteridae are split from the Velatida, and the old Spinulosida is broken up.


Human relations


In research

Starfish are deuterostomes, closely related, together with all other echinoderms, to
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 fi ...
s, and are used in reproductive and developmental studies. Female starfish produce large numbers of
oocyte An oocyte (, ), oöcyte, or ovocyte is a female gametocyte or germ cell involved in reproduction. In other words, it is an immature ovum, or egg cell. An oocyte is produced in a female fetus in the ovary during female gametogenesis. The femal ...
s that are easily isolated; these can be stored in a pre-meiosis phase and stimulated to complete division by the use of 1-methyladenine. Starfish oocytes are well suited for this research as they are large and easy to handle, transparent, simple to maintain in sea water at room temperature, and they develop rapidly. '' Asterina pectinifera'', used as a model organism for this purpose, is resilient and easy to breed and maintain in the laboratory. Another area of research is the ability of starfish to regenerate lost body parts. The stem cells of adult humans are incapable of much differentiation and understanding the regrowth, repair and cloning processes in starfish may have implications for human medicine. Starfish also have an unusual ability to expel foreign objects from their bodies, which makes them difficult to tag for research tracking purposes.


In legend and culture

An
aboriginal Australian Aboriginal Australians are the various Indigenous peoples of the Australian mainland and many of its islands, such as Tasmania, Fraser Island, Hinchinbrook Island, the Tiwi Islands, and Groote Eylandt, but excluding the Torres Strait Isl ...
fable retold by the Welsh school headmaster William Jenkyn Thomas (1870–1959) tells how some animals needed a canoe to cross the ocean. Whale had one but refused to lend it, so Starfish kept him busy, telling him stories and grooming him to remove parasites, while the others stole the canoe. When Whale realized the trick he beat Starfish ragged, which is how Starfish still is today. In 1900, the scholar
Edward Tregear Edward Robert Tregear , Ordre des Palmes académiques (1846–1931) was a New Zealand public servant and scholar. He was an architect of New Zealand's advanced social reforms and progressive labour legislation during the 1890s. Biography He was ...
documented ''The Creation Song'', which he describes as "an ancient prayer for the dedication of a high chief" of
Hawaii Hawaii ( ; haw, Hawaii or ) is a state in the Western United States, located in the Pacific Ocean about from the U.S. mainland. It is the only U.S. state outside North America, the only state that is an archipelago, and the only state ...
. Among the "uncreated gods" described early in the song are the male Kumulipo ("Creation") and the female Poele, both born in the night, a coral insect, the earthworm, and the starfish.
Georg Eberhard Rumpf Georg Eberhard Rumphius (originally: Rumpf; baptized c. 1 November 1627 – 15 June 1702) was a German-born botanist employed by the Dutch East India Company in what is now eastern Indonesia, and is best known for his work ''Herbarium Amboinense' ...
's 1705 ''The Ambonese Curiosity Cabinet'' describes the tropical varieties of ''Stella Marina'' or ''Bintang Laut'', "Sea Star", in Latin and Malay respectively, known in the waters around
Ambon Ambon may refer to: Places * Ambon Island, an island in Indonesia ** Ambon, Maluku, a city on Ambon Island, the capital of Maluku province ** Governorate of Ambon, a colony of the Dutch East India Company from 1605 to 1796 * Ambon, Morbihan, a c ...
. He writes that the ''Histoire des Antilles'' reports that when the sea stars "see thunder storms approaching,
hey Hey or Hey! may refer to: Music * Hey (band), a Polish rock band Albums * ''Hey'' (Andreas Bourani album) or the title song (see below), 2014 * ''Hey!'' (Julio Iglesias album) or the title song, 1980 * ''Hey!'' (Jullie album) or the title s ...
grab hold of many small stones with their little legs, looking to ... hold themselves down as if with anchors". ''Starfish'' is the title of novels by Peter Watts and Jennie Orbell, and in 2012, Alice Addison wrote a non-fiction book titled "Starfish - A year in the life of bereavement and depression". '' The Starfish and the Spider'' is a 2006
business management Business administration, also known as business management, is the administration of a commercial enterprise. It includes all aspects of overseeing and supervising the business operations of an organization. From the point of view of managemen ...
book by Ori Brafman and Rod Beckstrom; its title alludes to the ability of the starfish to regenerate itself because of its decentralized nervous system, and the book suggests ways that a decentralized organisation may flourish. In the
Nickelodeon Nickelodeon (often shortened to Nick) is an American pay television channel which launched on April 1, 1979, as the first cable channel for children. It is run by Paramount Global through its networks division's Kids and Family Group. It ...
animated
television series A television show – or simply TV show – is any content produced for viewing on a television set which can be broadcast via over-the-air, satellite, or cable, excluding breaking news, advertisements, or trailers that are typically placed be ...
''
SpongeBob SquarePants ''SpongeBob SquarePants'' (or simply ''SpongeBob'') is an American animated comedy television series created by marine science educator and animator Stephen Hillenburg for Nickelodeon. It chronicles the adventures of the title character ...
'', the
eponymous An eponym is a person, a place, or a thing after whom or which someone or something is, or is believed to be, named. The adjectives which are derived from the word eponym include ''eponymous'' and ''eponymic''. Usage of the word The term ''epon ...
character's best friend is a dim-witted starfish,
Patrick Star Patrick Star is a fictional character in the American animated television series ''SpongeBob SquarePants''. He is voiced by actor Bill Fagerbakke and was created and designed by marine biology, marine biologist and animator Stephen Hillenburg ...
.


As food

Starfish are widespread in the oceans, but are only occasionally used as food. There may be good reason for this: the bodies of numerous species are dominated by bony ossicles, and the body wall of many species contains saponins, which have an unpleasant taste, and others contain
tetrodotoxin Tetrodotoxin (TTX) is a potent neurotoxin. Its name derives from Tetraodontiformes, an order that includes pufferfish, porcupinefish, ocean sunfish, and triggerfish; several of these species carry the toxin. Although tetrodotoxin was discovere ...
s which are poisonous. Some species that prey on
bivalve molluscs Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, bival ...
can transmit
paralytic shellfish poisoning Paralytic shellfish poisoning (PSP) is one of the four recognized syndromes of shellfish poisoning, which share some common features and are primarily associated with bivalve mollusks (such as mussels, clams, oysters and scallops). These shellfi ...
. Georg Eberhard Rumpf found few starfish being used for food in the
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 ...
n archipelago, other than as bait in fish traps, but on the island of "Huamobel" the people cut them up, squeeze out the "black blood" and cook them with sour
tamarind Tamarind (''Tamarindus indica'') is a leguminous tree bearing edible fruit that is probably indigenous to tropical Africa. The genus ''Tamarindus'' is monotypic, meaning that it contains only this species. It belongs to the family Fabaceae ...
leaves; after resting the pieces for a day or two, they remove the outer skin and cook them in coconut milk. Starfish are sometimes eaten in China, Japan and in Micronesia.


As collectables

Starfish are in some cases taken from their habitat and sold to tourists as
souvenir A souvenir (), memento, keepsake, or token of remembrance is an object a person acquires for the memories the owner associates with it. A souvenir can be any object that can be collected or purchased and transported home by the traveler as a m ...
s,
ornaments An ornament is something used for decoration. Ornament may also refer to: Decoration *Ornament (art), any purely decorative element in architecture and the decorative arts *Biological ornament, a characteristic of animals that appear to serve on ...
, curios or for display in aquariums. In particular, ''Oreaster reticulatus'', with its easily accessed habitat and conspicuous coloration, is widely collected in the Caribbean. In the early to mid 20th century, this species was common along the coasts of the West Indies, but collection and trade have severely reduced its numbers. In the
State of 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 th ...
, ''O. reticulatus'' is listed as
endangered An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching and in ...
and its collection is illegal. Nevertheless, it is still sold throughout its range and beyond. A similar phenomenon exists in the Indo-Pacific for species such as '' Protoreaster nodosus''.


In industry and military history

With its multiple arms, the starfish provides a popular metaphor for
computer network A computer network is a set of computers sharing resources located on or provided by network nodes. The computers use common communication protocols over digital interconnections to communicate with each other. These interconnections are ...
s, companies and
software Software is a set of computer programs and associated software documentation, documentation and data (computing), data. This is in contrast to Computer hardware, hardware, from which the system is built and which actually performs the work. ...
tools. It is also the name of a seabed imaging system and company. Starfish has repeatedly been chosen as a name in military history. Three ships of the
Royal Navy The Royal Navy (RN) is the United Kingdom's naval warfare force. Although warships were used by English and Scottish kings from the early medieval period, the first major maritime engagements were fought in the Hundred Years' War against ...
have borne the name HMS ''Starfish'': an A-class destroyer launched in 1894; an R-class destroyer launched in 1916; and an S-class submarine launched in 1933 and lost in 1940. In the
World War II World War II or the Second World War, often abbreviated as WWII or WW2, was a world war that lasted from 1939 to 1945. It involved the vast majority of the world's countries—including all of the great powers—forming two opposing ...
, Starfish sites were large-scale night-time
decoys A decoy (derived from the Dutch ''de'' ''kooi'', literally "the cage" or possibly ''ende kooi'', " duck cage") is usually a person, device, or event which resembles what an individual or a group might be looking for, but it is only meant to lu ...
created during
The Blitz The Blitz was a German bombing campaign against the United Kingdom in 1940 and 1941, during the Second World War. The term was first used by the British press and originated from the term , the German word meaning 'lightning war'. The Germa ...
to simulate burning British cities. Starfish Prime was a high-altitude nuclear test conducted by the
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territori ...
on 9 July 1962.


References


Bibliography

* *


External links

* , a blog about sea stars by a passionate and professional specialist. {{featured article Articles containing video clips Late Ordovician first appearances Extant Ordovician first appearances Taxa named by Henri Marie Ducrotay de Blainville