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An octopus ( : octopuses or octopodes, see below for variants) is a soft-bodied, eight- limbed
mollusc 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 est ...
of the
order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of ...
Octopoda (, ). The order consists of some 300
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 group of organisms in which any two individuals of the appropriat ...
and is grouped within 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 differently ...
Cephalopod A cephalopod is any member of the molluscan class Cephalopoda ( Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head ...
a with
squid True squid are molluscs with an elongated soft body, large eyes, eight arms, and two tentacles in the superorder Decapodiformes, though many other molluscs within the broader Neocoleoidea are also called squid despite not strictly fittin ...
s,
cuttlefish Cuttlefish or cuttles are marine molluscs of the order Sepiida. They belong to the class Cephalopoda which also includes squid, octopuses, and nautiluses. Cuttlefish have a unique internal shell, the cuttlebone, which is used for control of ...
, and nautiloids. Like other cephalopods, an octopus is
bilaterally symmetric 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 pl ...
with two eyes and a
beak The beak, bill, or rostrum is an external anatomical structure found mostly in birds, but also in turtles, non-avian dinosaurs and a few mammals. A beak is used for eating, preening, manipulating objects, killing prey, fighting, probing for fo ...
ed mouth at the center point of the eight limbs. The soft body can radically alter its shape, enabling octopuses to squeeze through small gaps. They trail their eight appendages behind them as they swim. The siphon is used both for respiration and for locomotion, by expelling a jet of water. Octopuses have a complex nervous system and excellent sight, and are among the most intelligent and behaviourally diverse of all
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chorda ...
s. Octopuses inhabit various regions of the
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 wor ...
, including
coral reef A coral reef is an underwater ecosystem characterized by reef-building corals. Reefs are formed of Colony (biology), colonies of coral polyp (zoology), polyps held together by calcium carbonate. Most coral reefs are built from stony corals, wh ...
s,
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 w ...
waters, and 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 o ...
; some live in the intertidal zone and others at abyssal depths. Most species grow quickly, mature early, and are short-lived. In most species, the male uses a specially adapted arm to deliver a bundle of sperm directly into the female's mantle cavity, after which he becomes senescent and dies, while the female deposits fertilised eggs in a den and cares for them until they hatch, after which she also dies. Strategies to defend themselves against predators include the expulsion of ink, the use of
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...
and threat displays, the ability to jet quickly through the water and hide, and even deceit. All octopuses are venomous, but only the blue-ringed octopuses are known to be deadly to humans. Octopuses appear in mythology as sea monsters like the Kraken of Norway and the
Akkorokamui The is a gigantic octopus-like monster from Ainu folklore, similar to the Nordic Kraken, which supposedly lurks in Uchiura Bay The or is a bay southeast of the island of Hokkaido in northern Japan. It has also been known as Iburi Bay and ...
of the Ainu, and probably the Gorgon of
ancient Greece Ancient Greece ( el, Ἑλλάς, Hellás) was a northeastern Mediterranean civilization, existing from the Greek Dark Ages of the 12th–9th centuries BC to the end of classical antiquity ( AD 600), that comprised a loose collection of cu ...
. A battle with an octopus appears in
Victor Hugo Victor-Marie Hugo (; 26 February 1802 – 22 May 1885) was a French Romantic writer and politician. During a literary career that spanned more than sixty years, he wrote in a variety of genres and forms. He is considered to be one of the great ...
's book '' Toilers of the Sea'', inspiring other works such as
Ian Fleming Ian Lancaster Fleming (28 May 1908 – 12 August 1964) was a British writer who is best known for his postwar ''James Bond'' series of spy novels. Fleming came from a wealthy family connected to the merchant bank Robert Fleming & Co., an ...
's ''
Octopussy ''Octopussy'' is a 1983 spy film and the thirteenth in the ''James Bond'' series produced by Eon Productions. It is the sixth to star Roger Moore as the MI6 agent James Bond. It was directed by John Glen and the screenplay was written by ...
''. Octopuses appear in Japanese erotic art, ''
shunga is a type of Japanese erotic art typically executed as a kind of ukiyo-e, often in woodblock print format. While rare, there are also extant erotic painted handscrolls which predate ukiyo-e. Translated literally, the Japanese word ''shunga' ...
''. They are eaten and considered a delicacy by humans in many parts of the world, especially the
Mediterranean The Mediterranean Sea is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean Basin and almost completely enclosed by land: on the north by Western and Southern Europe and Anatolia, on the south by North Africa, and on ...
and the Asian seas.


Etymology and pluralisation

The scientific Latin term ''octopus'' was derived from
Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic pe ...
ὀκτώπους, a compound form of ὀκτώ (''oktō'', "eight") and πούς (''pous'', "foot"), itself a variant form of ὀκτάπους, a word used for example by Alexander of Tralles (c. 525–c. 605) for the common octopus. The standard
plural The plural (sometimes list of glossing abbreviations, abbreviated pl., pl, or ), in many languages, is one of the values of the grammatical number, grammatical category of number. The plural of a noun typically denotes a quantity greater than the ...
ised form of "octopus" in English is "octopuses"; the Ancient Greek plural ὀκτώποδες, "octopodes" (), has also been used historically. The alternative plural "octopi" is considered grammatically incorrect because it wrongly assumes that ''octopus'' is a
Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through ...
second declension " -us" noun or adjective when, in either Greek or Latin, it is a
third declension {{No footnotes, date=February 2021 The third declension is a category of nouns in Latin and Greek with broadly similar case formation — diverse stems, but similar endings. Sanskrit also has a corresponding class (although not commonly ter ...
noun. Historically, the first plural to commonly appear in English language sources, in the early 19th century, is the latinate form "octopi", followed by the English form "octopuses" in the latter half of the same century. The Hellenic plural is roughly contemporary in usage, although it is also the rarest. '' Fowler's Modern English Usage'' states that the only acceptable plural in English is "octopuses", that "octopi" is misconceived, and "octopodes" pedantic; the last is nonetheless used frequently enough to be acknowledged by the descriptivist ''Merriam-Webster 11th Collegiate Dictionary'' and ''Webster's New World College Dictionary''. 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 c ...
'' lists "octopuses", "octopi", and "octopodes", in that order, reflecting frequency of use, calling "octopodes" rare and noting that "octopi" is based on a misunderstanding. The '' New Oxford American Dictionary'' (3rd Edition, 2010) lists "octopuses" as the only acceptable pluralisation, and indicates that "octopodes" is still occasionally used, but that "octopi" is incorrect.


Anatomy and physiology


Size

The giant Pacific octopus ''(Enteroctopus dofleini)'' is often cited as the largest known octopus species. Adults usually weigh around 15 kg (33 lb), with an arm span of up to 4.3 m (14 ft). The largest specimen of this species to be scientifically documented was an animal with a live mass of 71 kg (156.5 lb). Much larger sizes have been claimed for the giant Pacific octopus:Norman, M. 2000. ''Cephalopods: A World Guide''. ConchBooks, Hackenheim. p. 214. one specimen was recorded as 272 kg (600 lb) with an arm span of 9 m (30 ft). A carcass of the seven-arm octopus, ''Haliphron atlanticus'', weighed 61 kg (134 lb) and was estimated to have had a live mass of 75 kg (165 lb). The smallest species is ''
Octopus wolfi ''Octopus wolfi'', the star-sucker pygmy octopus, is the smallest known octopus An octopus ( : octopuses or octopodes, see below for variants) is a soft-bodied, eight- limbed mollusc of the order Octopoda (, ). The order consists of s ...
'', which is around and weighs less than .


External characteristics

The octopus is
bilaterally symmetrical 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 pl ...
along its dorso-ventral (back to belly) axis; the head and foot are at one end of an elongated body and function as the anterior (front) of the animal. The head includes the mouth and brain. The foot has evolved into a set of flexible, prehensile appendages, known as "arms", that surround the mouth and are attached to each other near their base by a webbed structure. The arms can be described based on side and sequence position (such as L1, R1, L2, R2) and divided into four pairs. The two rear appendages are generally used to walk on the sea floor, while the other six are used to forage for food. The bulbous and hollow
mantle A mantle is a piece of clothing, a type of cloak. Several other meanings are derived from that. Mantle may refer to: *Mantle (clothing), a cloak-like garment worn mainly by women as fashionable outerwear **Mantle (vesture), an Eastern Orthodox ve ...
is fused to the back of the head and is known as the visceral hump; it contains most of the vital organs. The mantle cavity has muscular walls and contains the gills; it is connected to the exterior by a funnel or siphon. The mouth of an octopus, located underneath the arms, has a sharp hard
beak The beak, bill, or rostrum is an external anatomical structure found mostly in birds, but also in turtles, non-avian dinosaurs and a few mammals. A beak is used for eating, preening, manipulating objects, killing prey, fighting, probing for fo ...
. The skin consists of a thin outer epidermis with mucous cells and sensory cells, and a connective tissue
dermis The dermis or corium is a layer of skin between the epidermis (with which it makes up the cutis) and subcutaneous tissues, that primarily consists of dense irregular connective tissue and cushions the body from stress and strain. It is divided ...
consisting largely of
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 whol ...
fibres and various cells allowing colour change. Most of the body is made of soft tissue allowing it to lengthen, contract, and contort itself. The octopus can squeeze through tiny gaps; even the larger species can pass through an opening close to in diameter. Lacking skeletal support, the arms work as muscular hydrostats and contain longitudinal, transverse and circular muscles around a central axial nerve. They can extend and contract, twist to left or right, bend at any place in any direction or be held rigid. The interior surfaces of the arms are covered with circular, adhesive suckers. The suckers allow the octopus to anchor itself or to manipulate objects. Each sucker is usually circular and bowl-like and has two distinct parts: an outer shallow cavity called an infundibulum and a central hollow cavity called an
acetabulum The acetabulum (), also called the cotyloid cavity, is a concave surface of the pelvis. The head of the femur meets with the pelvis at the acetabulum, forming the hip joint. Structure There are three bones of the ''os coxae'' (hip bone) that c ...
, both of which are thick muscles covered in a protective chitinous cuticle. When a sucker attaches to a surface, the orifice between the two structures is sealed. The infundibulum provides adhesion while the acetabulum remains free, and muscle contractions allow for attachment and detachment. Each of the eight arms senses and responds to light, allowing the octopus to control the limbs even if its head is obscured. The eyes of the octopus are large and at the top of the head. They are similar in structure to those of a fish, and are enclosed in a cartilaginous capsule fused to the cranium. The
cornea The cornea is the transparent front part of the eye that covers the iris, pupil, and anterior chamber. Along with the anterior chamber and lens, the cornea refracts light, accounting for approximately two-thirds of the eye's total optical ...
is formed from a
translucent In the field of optics, transparency (also called pellucidity or diaphaneity) is the physical property of allowing light to pass through the material without appreciable scattering of light. On a macroscopic scale (one in which the dimensions ...
epidermal layer; the slit-shaped pupil forms a hole in the iris just behind the cornea. The lens is suspended behind the pupil; photoreceptive retinal cells cover the back of the eye. The pupil can be adjusted in size; a retinal pigment screens incident light in bright conditions. Some species differ in form from the typical octopus body shape. Basal species, the Cirrina, have stout gelatinous bodies with webbing that reaches near the tip of their arms, and two large fins above the eyes, supported by an internal shell. Fleshy papillae or cirri are found along the bottom of the arms, and the eyes are more developed.


Circulatory system

Octopuses have a closed
circulatory system The blood circulatory system is a system of organs that includes the heart, blood vessels, and blood which is circulated throughout the entire body of a human or other vertebrate. It includes the cardiovascular system, or vascular system, tha ...
, in which the blood remains inside blood vessels. Octopuses have three hearts; a systemic or main heart that circulates blood around the body and two branchial or gill hearts that pump it through each of the two gills. The systemic heart is inactive when the animal is swimming and thus it tires quickly and prefers to crawl. Octopus blood contains the
copper Copper is a chemical element with the symbol Cu (from la, cuprum) and atomic number 29. It is a soft, malleable, and ductile metal with very high thermal and electrical conductivity. A freshly exposed surface of pure copper has a pink ...
-rich protein haemocyanin to transport oxygen. This makes the blood very
viscous The viscosity of a fluid is a measure of its resistance to deformation at a given rate. For liquids, it corresponds to the informal concept of "thickness": for example, syrup has a higher viscosity than water. Viscosity quantifies the in ...
and it requires considerable pressure to pump it around the body; octopuses'
blood pressure Blood pressure (BP) is the pressure of circulating blood against the walls of blood vessels. Most of this pressure results from the heart pumping blood through the circulatory system. When used without qualification, the term "blood pressure ...
s can exceed . In cold conditions with low oxygen levels, haemocyanin transports oxygen more efficiently than haemoglobin. The haemocyanin is dissolved in the plasma instead of being carried within blood cells, and gives the blood a bluish colour. The systemic heart has muscular contractile walls and consists of a single ventricle and two atria, one for each side of the body. The blood vessels consist of arteries, capillaries and veins and are lined with a cellular
endothelium The endothelium is a single layer of squamous endothelial cells that line the interior surface of blood vessels and lymphatic vessels. The endothelium forms an interface between circulating blood or lymph in the lumen and the rest of the ve ...
which is quite unlike that of most other
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chorda ...
s. The blood circulates through the aorta and capillary system, to the vena cavae, after which the blood is pumped through the gills by the branchial hearts and back to the main heart. Much of the venous system is contractile, which helps circulate the blood.


Respiration

Respiration involves drawing water into the mantle cavity through an aperture, passing it through the gills, and expelling it through the siphon. The ingress of water is achieved by contraction of radial muscles in the mantle wall, and flapper valves shut when strong circular muscles force the water out through the siphon. Extensive connective tissue lattices support the respiratory muscles and allow them to expand the respiratory chamber. The lamella structure of the gills allows for a high oxygen uptake, up to 65% in water at . Water flow over the gills correlates with locomotion, and an octopus can propel its body when it expels water out of its siphon. The thin skin of the octopus absorbs additional oxygen. When resting, around 41% of an octopus's oxygen absorption is through the skin. This decreases to 33% when it swims, as more water flows over the gills; skin oxygen uptake also increases. When it is resting after a meal, absorption through the skin can drop to 3% of its total oxygen uptake.


Digestion and excretion

The digestive system of the octopus begins with the
buccal mass The digestive system of gastropods has evolved to suit almost every kind of diet and feeding behavior. Gastropods (snails and slugs) as the largest taxonomic class of the mollusca are very diverse: the group includes carnivores, herbivores, scaven ...
which consists of the mouth with its
chitin Chitin ( C8 H13 O5 N)n ( ) is a long-chain polymer of ''N''-acetylglucosamine, an amide derivative of glucose. Chitin is probably the second most abundant polysaccharide in nature (behind only cellulose); an estimated 1 billion tons of chit ...
ous beak, the pharynx,
radula The radula (, ; plural radulae or radulas) is an anatomical structure used by molluscs for feeding, sometimes compared to a tongue. It is a minutely toothed, chitinous ribbon, which is typically used for scraping or cutting food before the food ...
and salivary glands. The radula is a spiked, muscular tongue-like organ with multiple rows of tiny teeth. Food is broken down and is forced into the oesophagus by two lateral extensions of the esophageal side walls in addition to the radula. From there it is transferred to the gastrointestinal tract, which is mostly suspended from the roof of the mantle cavity by numerous membranes. The tract consists of a
crop A crop is a plant that can be grown and harvested extensively for profit or subsistence. When the plants of the same kind are cultivated at one place on a large scale, it is called a crop. Most crops are cultivated in agriculture or hydropon ...
, where the food is stored; a stomach, where food is ground down; a caecum where the now sludgy food is sorted into fluids and particles and which plays an important role in absorption; the digestive gland, where liver cells break down and absorb the fluid and become "brown bodies"; and the intestine, where the accumulated waste is turned into faecal ropes by secretions and blown out of the funnel via the rectum. During osmoregulation, fluid is added to the pericardia of the branchial hearts. The octopus has two nephridia (equivalent to vertebrate kidneys) which are associated with the branchial hearts; these and their associated ducts connect the pericardial cavities with the mantle cavity. Before reaching the branchial heart, each branch of the
vena cava In anatomy, the venae cavae (; singular: vena cava ; ) are two large veins (great vessels) that return deoxygenated blood from the body into the heart. In humans they are the superior vena cava and the inferior vena cava, and both empty into th ...
expands to form renal appendages which are in direct contact with the thin-walled nephridium. The urine is first formed in the pericardial cavity, and is modified by excretion, chiefly of ammonia, and selective absorption from the renal appendages, as it is passed along the associated duct and through the nephridiopore into the mantle cavity.


Nervous system and senses

Octopuses (along with
cuttlefish Cuttlefish or cuttles are marine molluscs of the order Sepiida. They belong to the class Cephalopoda which also includes squid, octopuses, and nautiluses. Cuttlefish have a unique internal shell, the cuttlebone, which is used for control of ...
) have the highest brain-to-body mass ratios of all invertebrates; this is greater than that of many vertebrates. Octopuses have the same
jumping genes A transposable element (TE, transposon, or jumping gene) is a nucleic acid sequence in DNA that can change its position within a genome, sometimes creating or reversing mutations and altering the cell's genetic identity and genome size. Tra ...
that are active in the human brain, implying an
evolutionary convergence Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
at molecular level. The
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 ...
is complex, only part of which is localised in its brain, which is contained in a cartilaginous capsule. Two-thirds of an octopus's
neuron A neuron, neurone, or nerve cell is an electrically excitable cell that communicates with other cells via specialized connections called synapses. The neuron is the main component of nervous tissue in all animals except sponges and placozoa ...
s are in the nerve cords of its arms; these are capable of complex reflex actions without input from the brain. Unlike vertebrates, the complex motor skills of octopuses are not organised in their brains via internal somatotopic maps of their bodies. Like other cephalopods, octopuses have camera-like eyes, and can distinguish the polarisation of light.
Colour vision Color vision, a feature of visual perception, is an ability to perceive differences between light composed of different wavelengths (i.e., different spectral power distributions) independently of light intensity. Color perception is a part of ...
appears to vary from species to species, for example being present in ''O. aegina'' but absent in ''O. vulgaris''.
Opsin Animal opsins are G-protein-coupled receptors and a group of proteins made light-sensitive via a chromophore, typically retinal. When bound to retinal, opsins become Retinylidene proteins, but are usually still called opsins regardless. Most ...
s in the skin respond to different wavelengths of light and help the animals choose a coloration that camouflages them; the chromatophores in the skin can respond to light independently of the eyes. An alternative hypothesis is that cephalopod eyes in species which only have a single
photoreceptor protein Photoreceptor proteins are light-sensitive proteins involved in the sensing and response to light in a variety of organisms. Some examples are rhodopsin in the photoreceptor cells of the vertebrate retina, phytochrome in plants, and bacteriorhod ...
may use
chromatic aberration In optics, chromatic aberration (CA), also called chromatic distortion and spherochromatism, is a failure of a lens to focus all colors to the same point. It is caused by dispersion: the refractive index of the lens elements varies with the w ...
to turn monochromatic vision into colour vision, though this sacrifices image quality. This would explain pupils shaped like the letter U, the letter W, or a dumbbell, as well as explaining the need for colourful mating displays. Attached to the brain are two organs called statocysts (sac-like structures containing a mineralised mass and sensitive hairs), that allow the octopus to sense the orientation of its body. They provide information on the position of the body relative to gravity and can detect angular acceleration. An autonomic response keeps the octopus's eyes oriented so that the pupil is always horizontal. Octopuses may also use the statocyst to hear sound. The common octopus can hear sounds between 400 Hz and 1000 Hz, and hears best at 600 Hz. Octopuses have an excellent
somatosensory system In physiology, the somatosensory system is the network of neural structures in the brain and body that produce the perception of touch (haptic perception), as well as temperature (thermoception), body position (proprioception), and pain. It is ...
. Their suction cups are equipped with
chemoreceptors A chemoreceptor, also known as chemosensor, is a specialized sensory receptor which transduces a chemical substance (endogenous or induced) to generate a biological signal. This signal may be in the form of an action potential, if the chemorecept ...
so they can taste what they touch. Octopus arms move easily because the sensors recognise octopus skin and prevent self-attachment. Octopuses appear to have poor proprioceptive sense and must observe the arms visually to keep track of their position.


Ink sac

The ink sac of an octopus is located under the digestive gland. A gland attached to the sac produces the ink, and the sac stores it. The sac is close enough to the funnel for the octopus to shoot out the ink with a water jet. Before it leaves the funnel, the ink passes through glands which mix it with mucus, creating a thick, dark blob which allows the animal to escape from a predator. The main pigment in the ink is
melanin Melanin (; from el, μέλας, melas, black, dark) is a broad term for a group of natural pigments found in most organisms. Eumelanin is produced through a multistage chemical process known as melanogenesis, where the oxidation of the amino ...
, which gives it its black colour. Cirrate octopuses usually lack the ink sac.


Lifecycle


Reproduction

Octopuses are gonochoric and have a single, posteriorly-located gonad which is associated with the
coelom The coelom (or celom) is the main body cavity in most animals and is positioned inside the body to surround and contain the digestive tract and other organs. In some animals, it is lined with mesothelium. In other animals, such as molluscs, ...
. The testis in males and the
ovary The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. ...
in females bulges into the gonocoel and the
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 ...
s are released here. The gonocoel is connected by the gonoduct to the mantle cavity, which it enters at the gonopore. An optic gland creates hormones that cause the octopus to mature and age and stimulate gamete production. The gland may be triggered by environmental conditions such as temperature, light and nutrition, which thus control the timing of reproduction and lifespan. When octopuses reproduce, the male uses a specialised arm called a hectocotylus to transfer
spermatophore A spermatophore or sperm ampulla is a capsule or mass containing spermatozoa created by males of various animal species, especially salamanders and arthropods, and transferred in entirety to the female's ovipore during reproduction. Spermatophore ...
s (packets of sperm) from the terminal organ of the reproductive tract (the cephalopod "penis") into the female's mantle cavity. The hectocotylus in
benthic 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 " ...
octopuses is usually the third right arm, which has a spoon-shaped depression and modified suckers near the tip. In most species, fertilisation occurs in the mantle cavity. The reproduction of octopuses has been studied in only a few species. One such species is the giant Pacific octopus, in which courtship is accompanied, especially in the male, by changes in skin texture and colour. The male may cling to the top or side of the female or position himself beside her. There is some speculation that he may first use his hectocotylus to remove any spermatophore or sperm already present in the female. He picks up a spermatophore from his spermatophoric sac with the hectocotylus, inserts it into the female's mantle cavity, and deposits it in the correct location for the species, which in the giant Pacific octopus is the opening of the oviduct. Two spermatophores are transferred in this way; these are about one metre (yard) long, and the empty ends may protrude from the female's mantle. A complex hydraulic mechanism releases the sperm from the spermatophore, and it is stored internally by the female. About forty days after mating, the female giant Pacific octopus attaches strings of small fertilised eggs (10,000 to 70,000 in total) to rocks in a crevice or under an overhang. Here she guards and cares for them for about five months (160 days) until they hatch. In colder waters, such as those off
Alaska Alaska ( ; russian: Аляска, Alyaska; ale, Alax̂sxax̂; ; ems, Alas'kaaq; Yup'ik: ''Alaskaq''; tli, Anáaski) is a state located in the Western United States on the northwest extremity of North America. A semi-exclave of the U ...
, it may take up to ten months for the eggs to completely develop. The female aerates them and keeps them clean; if left untended, many will die. She does not feed during this time and dies soon after. Males become senescent and die a few weeks after mating. The eggs have large yolks; cleavage (division) is superficial and a germinal disc develops at the pole. During gastrulation, the margins of this grow down and surround the yolk, forming a yolk sac, which eventually forms part of the gut. The dorsal side of the disc grows upward and forms the embryo, with a shell gland on its dorsal surface, gills, mantle and eyes. The arms and funnel develop as part of the foot on the ventral side of the disc. The arms later migrate upward, coming to form a ring around the funnel and mouth. The yolk is gradually absorbed as the embryo develops. Most young octopuses hatch as
paralarva Paralarvae (singular: ''paralarva'') are young cephalopods in the planktonic stages between hatchling and subadult. This stage differs from the larval stage of animals that undergo true metamorphosis. Paralarvae have been observed only in mem ...
e and are
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 cruc ...
ic for weeks to months, depending on the species and water temperature. They feed on copepods,
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 ...
larvae and other
zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
, eventually settling on the ocean floor and developing directly into adults with no distinct
metamorphoses The ''Metamorphoses'' ( la, Metamorphōsēs, from grc, μεταμορφώσεις: "Transformations") is a Latin narrative poem from 8 CE by the Roman poet Ovid. It is considered his '' magnum opus''. The poem chronicles the history of the ...
that are present in other groups of
mollusc 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 est ...
larvae. Octopus species that produce larger eggs – including the southern blue-ringed, Caribbean reef, California two-spot, ''
Eledone moschata ''Eledone moschata'', the musky octopus, is a species of octopus belonging to the family Octopodidae. Taxonomy The skin of the single specimen of '' Eledone microsicya'' is very similar to the skin of ''Eledone moschata'' and some authorities ta ...
'' and deep sea octopuses – instead hatch as benthic animals similar to the adults. In the argonaut (paper nautilus), the female secretes a fine, fluted, papery shell in which the eggs are deposited and in which she also resides while floating in mid-ocean. In this she broods the young, and it also serves as a buoyancy aid allowing her to adjust her depth. The male argonaut is minute by comparison and has no shell.


Lifespan

Octopuses have a relatively short lifespan; some species live for as little as six months. The Giant Pacific octopus, one of the two largest species of octopus, may live for as much as five years. Octopus lifespan is limited by reproduction. For most octopuses the last stage of their life is called senescence. It is the breakdown of cellular function without repair or replacement. For males, this typically begins after mating. Senescence may last from weeks to a few months, at most. For females, it begins when they lay a clutch of eggs. Females will spend all their time aerating and protecting their eggs until they are ready to hatch. During senescence, an octopus does not feed and quickly weakens. Lesions begin to form and the octopus literally degenerates. Unable to defend themselves, octopuses often fall prey to predators. The
larger Pacific striped octopus The larger Pacific striped octopus (LPSO), or Harlequin octopus, is a species of octopus known for its intelligence and gregarious nature. The species was first documented in the 1970s and, being fairly new to scientific observation, has yet to b ...
(LPSO) is an exception, as it can reproduce repeatedly over a life of around two years. Octopus reproductive organs mature due to the hormonal influence of the optic gland but result in the inactivation of their digestive glands. Unable to feed, the octopus typically dies of starvation. Experimental removal of both optic glands after spawning was found to result in the cessation of broodiness, the resumption of feeding, increased growth, and greatly extended lifespans. It has been proposed that the naturally short lifespan may be functional to prevent rapid overpopulation.


Distribution and habitat

Octopuses live in every ocean, and different species have adapted to different marine habitats. As juveniles, common octopuses inhabit shallow tide pools. The Hawaiian day octopus (''
Octopus cyanea ''Octopus cyanea'', also known as the big blue octopus or day octopus, is an octopus in the family Octopodidae. It occurs in both the Pacific and Indian Oceans, from Hawaii to the eastern coast of Africa.Norman, M.D. 2000. ''Cephalopods: A Worl ...
'') lives on coral reefs;
argonauts The Argonauts (; Ancient Greek: ) were a band of heroes in Greek mythology, who in the years before the Trojan War (around 1300 BC) accompanied Jason to Colchis in his quest to find the Golden Fleece. Their name comes from their ship, ''Argo ...
drift in pelagic waters. '' Abdopus aculeatus'' mostly lives in near-shore
seagrass Seagrasses are the only flowering plants which grow in marine environments. There are about 60 species of fully marine seagrasses which belong to four families ( Posidoniaceae, Zosteraceae, Hydrocharitaceae and Cymodoceaceae), all in the ...
beds. Some species are adapted to the cold, ocean depths. The spoon-armed octopus ('' Bathypolypus arcticus'') is found at depths of , and '' Vulcanoctopus hydrothermalis'' lives near hydrothermal vents at . The cirrate species are often free-swimming and live in deep-water habitats. Although several species are known to live at bathyal and abyssal depths, there is only a single indisputable record of an octopus in the hadal zone; a species of '' Grimpoteuthis'' (dumbo octopus) photographed at . No species are known to live in fresh water.


Behaviour and ecology

Most species are solitary when not mating, though a few are known to occur in high densities and with frequent interactions, signaling, mate defending and eviction of individuals from dens. This is likely the result of abundant food supplies combined with limited den sites. The LPSO has been described as particularly social, living in groups of up to 40 individuals. Octopuses hide in dens, which are typically crevices in rocky outcrops or other hard structures, though some species burrow into sand or mud. Octopuses are not
territorial A territory is an area of land, sea, or space, particularly belonging or connected to a country, person, or animal. In international politics, a territory is usually either the total area from which a state may extract power resources or an ...
but generally remain in a home range; they may leave in search of food. They can
navigate Navigation is a field of study that focuses on the process of monitoring and controlling the movement of a craft or vehicle from one place to another.Bowditch, 2003:799. The field of navigation includes four general categories: land navigation, ...
back to a den without having to retrace their outward route. They are not migratory. Octopuses bring captured prey to the den, where they can eat it safely. Sometimes the octopus catches more prey than it can eat, and the den is often surrounded by a
midden A midden (also kitchen midden or shell heap) is an old dump for domestic waste which may consist of animal bone, human excrement, botanical material, mollusc shells, potsherds, lithics (especially debitage), and other artifacts and eco ...
of dead and uneaten food items. Other creatures, such as fish,
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 th ...
s, molluscs and
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, often share the den with the octopus, either because they have arrived as scavengers, or because they have survived capture. On rare occasions, octopuses hunt cooperatively with other species, with fish as their partners. They regulate the species composition of the hunting and the behavior of their by punching them.


Feeding

Nearly all octopuses are predatory; bottom-dwelling octopuses eat mainly
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapoda, decapods, ostracoda, seed shrimp, branchiopoda, branchiopods, argulidae, fish lice, krill, remipedes, isopoda, isopods, barnacles, copepods, ...
s, polychaete worms, and other molluscs such as whelks and clams; open-ocean octopuses eat mainly prawns, fish and other cephalopods. Major items in the diet of the giant Pacific octopus include bivalve molluscs such as the cockle ''
Clinocardium nuttallii ''Clinocardium '' is a genus of marine bivalve molluscs in the family Cardiidae, the cockles. It is monotypic, being represented by the single species ''Clinocardium nuttallii'', also known as the basket cockle or Nuttall's cockle, a large edi ...
'', clams and scallops and crustaceans such as crabs and
spider crabs The Majoidea are a superfamily of crabs which includes the various spider crabs. Taxonomy In "''A classification of living and fossil genera of decapod crustaceans''" De Grave and colleagues divided Majoidea into six families: * Family Epialt ...
. Prey that it is likely to reject include moon snails because they are too large and limpets, rock scallops, chitons and
abalone Abalone ( or ; via Spanish , from Rumsen ''aulón'') is a common name for any of a group of small to very large marine gastropod molluscs in the family Haliotidae. Other common names are ear shells, sea ears, and, rarely, muttonfish or mutto ...
, because they are too securely fixed to the rock. A benthic (bottom-dwelling) octopus typically moves among the rocks and feels through the crevices. The creature may make a jet-propelled pounce on prey and pull it toward the mouth with its arms, the suckers restraining it. Small prey may be completely trapped by the webbed structure. Octopuses usually inject crustaceans like crabs with a paralysing
saliva Saliva (commonly referred to as spit) is an extracellular fluid produced and secreted by salivary glands in the mouth. In humans, saliva is around 99% water, plus electrolytes, mucus, white blood cells, epithelial cells (from which DNA can ...
then dismember them with their beaks. Octopuses feed on shelled molluscs either by forcing the valves apart, or by drilling a hole in the shell to inject a nerve toxin. It used to be thought that the hole was drilled by the radula, but it has now been shown that minute teeth at the tip of the salivary papilla are involved, and an enzyme in the toxic saliva is used to dissolve the calcium carbonate of the shell. It takes about three hours for ''O. vulgaris'' to create a hole. Once the shell is penetrated, the prey dies almost instantaneously, its muscles relax, and the soft tissues are easy for the octopus to remove. Crabs may also be treated in this way; tough-shelled species are more likely to be drilled, and soft-shelled crabs are torn apart. Some species have other modes of feeding. '' Grimpoteuthis'' has a reduced or non-existent radula and swallows prey whole. In the deep-sea genus ''
Stauroteuthis ''Stauroteuthis'' is a genus of deepwater octopus, a cephalopod mollusk. This is the only genus in the family Stauroteuthidae, and only two species have been described in this genus. The organisms live below water depth; although sometimes foun ...
'', some of the muscle cells that control the suckers in most species have been replaced with photophores which are believed to fool prey by directing them to the mouth, making them one of the few bioluminescent octopuses.


Locomotion

Octopuses mainly move about by relatively slow crawling with some swimming in a head-first position. Jet propulsion or backward swimming, is their fastest means of locomotion, followed by swimming and crawling. When in no hurry, they usually crawl on either solid or soft surfaces. Several arms are extended forward, some of the suckers adhere to the substrate and the animal hauls itself forward with its powerful arm muscles, while other arms may push rather than pull. As progress is made, other arms move ahead to repeat these actions and the original suckers detach. During crawling, the heart rate nearly doubles, and the animal requires ten or fifteen minutes to recover from relatively minor exercise. Most octopuses swim by expelling a jet of water from the mantle through the siphon into the sea. The physical principle behind this is that the force required to accelerate the water through the orifice produces a reaction that propels the octopus in the opposite direction. The direction of travel depends on the orientation of the siphon. When swimming, the head is at the front and the siphon is pointed backward but, when jetting, the visceral hump leads, the siphon points at the head and the arms trail behind, with the animal presenting a fusiform appearance. In an alternative method of swimming, some species flatten themselves dorso-ventrally, and swim with the arms held out sideways, and this may provide lift and be faster than normal swimming. Jetting is used to escape from danger, but is physiologically inefficient, requiring a mantle pressure so high as to stop the heart from beating, resulting in a progressive oxygen deficit. Cirrate octopuses cannot produce jet propulsion and rely on their fins for swimming. They have neutral buoyancy and drift through the water with the fins extended. They can also contract their arms and surrounding web to make sudden moves known as "take-offs". Another form of locomotion is "pumping", which involves symmetrical contractions of muscles in their webs producing peristaltic waves. This moves the body slowly. In 2005, '' Adopus aculeatus'' and veined octopus ('' Amphioctopus marginatus'') were found to walk on two arms, while at the same time mimicking plant matter. This form of locomotion allows these octopuses to move quickly away from a potential predator without being recognised. Some species of octopus can crawl out of the water briefly, which they may do between tide pools. "Stilt walking" is used by the veined octopus when carrying stacked coconut shells. The octopus carries the shells underneath it with two arms, and progresses with an ungainly gait supported by its remaining arms held rigid.


Intelligence

Octopuses are highly intelligent.
Maze A maze is a path or collection of paths, typically from an entrance to a goal. The word is used to refer both to branching tour puzzles through which the solver must find a route, and to simpler non-branching ("unicursal") patterns that le ...
and
problem-solving Problem solving is the process of achieving a goal by overcoming obstacles, a frequent part of most activities. Problems in need of solutions range from simple personal tasks (e.g. how to turn on an appliance) to complex issues in business an ...
experiments have shown evidence of a memory system that can store both short- and
long-term memory Long-term memory (LTM) is the stage of the Atkinson–Shiffrin memory model in which informative knowledge is held indefinitely. It is defined in contrast to short-term and working memory, which persist for only about 18 to 30 seconds. Long-t ...
. Young octopuses learn nothing from their parents, as adults provide no parental care beyond tending to their eggs until the young octopuses hatch. In laboratory experiments, octopuses can readily be trained to distinguish between different shapes and patterns. They have been reported to practise observational learning, although the validity of these findings is contested. Octopuses have also been observed in what has been described as play: repeatedly releasing bottles or toys into a circular current in their aquariums and then catching them. Octopuses often break out of their aquariums and sometimes into others in search of food. The
veined octopus ''Amphioctopus marginatus'', also known as the coconut octopus and veined octopus, is a medium-sized cephalopod belonging to the genus '' Amphioctopus''. It is found in tropical waters of the western Pacific Ocean. It commonly preys upon shrimp, ...
collects discarded coconut shells, then uses them to build a shelter, an example of
tool use Tool use by animals is a phenomenon in which an animal uses any kind of tool in order to achieve a goal such as acquiring food and water, grooming, defence, communication, recreation or construction. Originally thought to be a skill possessed o ...
.


Camouflage and colour change

Octopuses use
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...
when hunting and to avoid predators. To do this they use specialised skin cells which change the appearance of the skin by adjusting its colour, opacity, or reflectivity. Chromatophores contain yellow, orange, red, brown, or black pigments; most species have three of these colours, while some have two or four. Other colour-changing cells are reflective iridophores and white leucophores. This colour-changing ability is also used to communicate with or warn other octopuses. Octopuses can create distracting patterns with waves of dark coloration across the body, a display known as the "passing cloud". Muscles in the skin change the texture of the mantle to achieve greater camouflage. In some species, the mantle can take on the spiky appearance of algae; in others, skin anatomy is limited to relatively uniform shades of one colour with limited skin texture. Octopuses that are diurnal and live in shallow water have evolved more complex skin than their nocturnal and deep-sea counterparts. A "moving rock" trick involves the octopus mimicking a rock and then inching across the open space with a speed matching that of the surrounding water.


Defence

Aside from humans, octopuses may be preyed on by fishes,
seabird Seabirds (also known as marine birds) are birds that are adapted to life within the marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent evolution, as the same envir ...
s, sea otters,
pinniped Pinnipeds (pronounced ), commonly known as seals, are a widely distributed and diverse clade of carnivorous, fin-footed, semiaquatic, mostly marine mammals. They comprise the extant families Odobenidae (whose only living member is the ...
s, cetaceans, and other cephalopods. Octopuses typically hide or disguise themselves by camouflage and
mimicry In evolutionary biology, mimicry is an evolved resemblance between an organism and another object, often an organism of another species. Mimicry may evolve between different species, or between individuals of the same species. Often, mimicry f ...
; some have conspicuous warning coloration (aposematism) or
deimatic behaviour Deimatic behaviour or startle display means any pattern of bluffing behaviour in an animal that lacks strong defences, such as suddenly displaying conspicuous eyespots, to scare off or momentarily distract a predator, thus giving the prey anima ...
. An octopus may spend 40% of its time hidden away in its den. When the octopus is approached, it may extend an arm to investigate. 66% of ''Enteroctopus dofleini'' in one study had scars, with 50% having amputated arms. The blue rings of the highly venomous blue-ringed octopus are hidden in muscular skin folds which contract when the animal is threatened, exposing the iridescent warning. The
Atlantic white-spotted octopus ''Callistoctopus macropus'', also known as the Atlantic white-spotted octopus, white-spotted octopus,Norman, M.D. 2000. ''Cephalopods: A World Guide''. ConchBooks.Bouchet, P. (2014). Callistoctopus macropus (Risso, 1826). Accessed through: World ...
(''Callistoctopus macropus'') turns bright brownish red with oval white spots all over in a high contrast display. Displays are often reinforced by stretching out the animal's arms, fins or web to make it look as big and threatening as possible. Once they have been seen by a predator, they commonly try to escape but can also use distraction with an ink cloud ejected from the ink sac. The ink is thought to reduce the efficiency of olfactory organs, which would aid evasion from predators that employ smell for hunting, such as
shark Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachi ...
s. Ink clouds of some species might act as
pseudomorphs In mineralogy, a pseudomorph is a mineral or mineral compound that appears in an atypical form ( crystal system), resulting from a substitution process in which the appearance and dimensions remain constant, but the original mineral is replaced b ...
, or decoys that the predator attacks instead. When under attack, some octopuses can perform arm
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 ...
, in a manner similar to the way skinks and other lizards detach their tails. The crawling arm may distract would-be predators. Such severed arms remain sensitive to stimuli and move away from unpleasant sensations. Octopuses can replace lost limbs. Some octopuses, such as the mimic octopus, can combine their highly flexible bodies with their colour-changing ability to mimic other, more dangerous animals, such as lionfish, sea snakes, and eels.


Pathogens and parasites

The diseases and parasites that affect octopuses have been little studied, but cephalopods are known to be the intermediate or final
hosts A host is a person responsible for guests at an event or for providing hospitality during it. Host may also refer to: Places * Host, Pennsylvania, a village in Berks County People *Jim Host (born 1937), American businessman *Michel Host ...
of various parasitic cestodes,
nematode The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant- parasitic nematodes also known as eelworms. They are a diverse animal phylum inhabiting a bro ...
s and copepods; 150 species of
protist A protist () is any eukaryotic organism (that is, an organism whose cells contain a cell nucleus) that is not an animal, plant, or fungus. While it is likely that protists share a common ancestor (the last eukaryotic common ancestor), the e ...
an and
metazoa Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage in ...
n parasites have been recognised. The
Dicyemidae The Dicyemidae is a family of tiny parasites that live in the renal appendages of cephalopods. It contains the following genera and species: * ''Dicyema'' von Kolliker, 1849 * '' Dicyemennea'' Whitman, 1883 * '' Dicyemodeca'' ** '' Dicyemodeca ...
are a family of tiny worms that are found in the renal appendages of many species; it is unclear whether they are parasitic or endosymbionts.
Coccidia Coccidia (Coccidiasina) are a subclass of microscopic, spore-forming, single-celled obligate intracellular parasites belonging to the apicomplexan class Conoidasida. As obligate intracellular parasites, they must live and reproduce within an a ...
ns in the genus '' Aggregata'' living in the gut cause severe disease to the host. Octopuses have an innate immune system; their haemocytes respond to infection by phagocytosis, encapsulation, infiltration, or cytotoxic activities to destroy or isolate the pathogens. The haemocytes play an important role in the recognition and elimination of foreign bodies and wound repair. Captive animals are more susceptible to pathogens than wild ones. A gram-negative bacterium, '' Vibrio lentus'', can cause skin lesions, exposure of muscle and sometimes death.


Evolution

The scientific name Octopoda was first coined and given as the order of octopuses in 1818 by English biologist
William Elford Leach William Elford Leach FRS (2 February 1791 – 25 August 1836) was an English zoologist and marine biologist. Life and work Elford Leach was born at Hoe Gate, Plymouth, the son of an attorney. At the age of twelve he began a medical appre ...
, who classified them as Octopoida the previous year. The Octopoda consists of around 300 known species and were historically divided into two suborders, the Incirrina and the Cirrina. More recent evidence suggests Cirrina is merely the most basal species, not a unique
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English ter ...
. The incirrate octopuses (the majority of species) lack the cirri and paired swimming fins of the cirrates. In addition, the internal shell of incirrates is either present as a pair of stylets or absent altogether.


Fossil history and phylogeny

The Cephalopoda evolved from a mollusc resembling the
Monoplacophora Monoplacophora , meaning "bearing one plate", is a polyphyletic superclass of molluscs with a cap-like shell inhabiting deep sea environments . Extant representatives were not recognized as such until 1952; previously they were known only from ...
in the
Cambrian The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago ...
some 530 million years ago. The Coleoidea diverged from the nautiloids in the
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, wh ...
some 416 million years ago. In turn, the coleoids (including the squids and octopods) brought their shells inside the body and some 276 million years ago, during the
Permian The Permian ( ) is a geologic period and 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 period of the Paleo ...
, split into the Vampyropoda and the Decabrachia. The octopuses arose from the
Muensterelloidea Muensterelloidea is a superfamily (or clade) of stem- octopod cephalopods from the Early Jurassic to Late Cretaceous. Two families are currently identified, Muensterellidae, and Patelloctopodidae. The clade is the ancestral group from which mod ...
within the Vampyropoda in the
Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
. The earliest octopus likely lived near the sea floor (
benthic 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 " ...
to
demersal The demersal zone is the part of the sea or ocean (or deep lake) consisting of the part of the water column near to (and significantly affected by) the seabed and the benthos. The demersal zone is just above the benthic zone and forms a layer ...
) in shallow marine environments. Octopuses consist mostly of soft tissue, and so fossils are relatively rare. As soft-bodied cephalopods, they lack the external shell of most molluscs, including other cephalopods like the nautiloids and the extinct Ammonoidea. They have eight limbs like other Coleoidea, but lack the extra specialised feeding appendages known as
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 which are longer and thinner with suckers only at their club-like ends. The vampire squid ('' Vampyroteuthis'') also lacks tentacles but has sensory filaments. The cladograms are based on Sanchez et al., 2018, who created a molecular phylogeny based on
mitochondria A mitochondrion (; ) is an organelle found in the cells of most Eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is used ...
l and nuclear DNA marker sequences. The position of the Eledonidae is from Ibáñez et al., 2020, with a similar methodology. Dates of divergence are from Kröger et al., 2011 and Fuchs et al, 2019. The molecular analysis of the octopods shows that the suborder Cirrina (Cirromorphida) and the superfamily Argonautoidea are
paraphyletic In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and most of its descendants, excluding a few monophyletic subgroups. The group is said to be paraphyletic ''with respect to'' the excluded subgroups. In ...
and are broken up; these names are shown in quotation marks and italics on the cladogram.


RNA editing and the genome

Octopuses, like other coleoid cephalopods but unlike more basal cephalopods or other molluscs, are capable of greater RNA editing, changing the nucleic acid sequence of the primary transcript of RNA molecules, than any other organisms. Editing is concentrated in the nervous system, and affects proteins involved in neural excitability and neuronal morphology. More than 60% of RNA transcripts for coleoid brains are recoded by editing, compared to less than 1% for a human or fruit fly. Coleoids rely mostly on ADAR enzymes for RNA editing, which requires large double-stranded RNA structures to flank the editing sites. Both the structures and editing sites are conserved in the coleoid genome and the mutation rates for the sites are severely hampered. Hence, greater transcriptome plasticity has come at the cost of slower genome evolution. The octopus genome is unremarkably
bilaterian The Bilateria or bilaterians are animals with bilateral symmetry as an embryo, i.e. having a left and a right side that are mirror images of each other. This also means they have a head and a tail (anterior-posterior axis) as well as a belly an ...
except for large developments of two gene families:
protocadherin Protocadherins (Pcdhs) are the largest mammalian subgroup of the cadherin superfamily of homophilic cell-adhesion proteins. They were discovered by Shintaro Suzuki's group, when they used PCR to find new members of the cadherin family. The PCR fra ...
s, which regulate the development of neurons; and the
C2H2 zinc-finger A zinc finger is a small protein structural motif that is characterized by the coordination of one or more zinc ions (Zn2+) in order to stabilize the fold. It was originally coined to describe the finger-like appearance of a hypothesized struct ...
transcription factors. Many genes specific to cephalopods are expressed in the animals' skin, suckers, and nervous system.


Relationship to humans


In culture

Ancient seafaring people were aware of the octopus, as evidenced by artworks and designs. For example, a stone carving found in the archaeological recovery from Bronze Age Minoan Crete at
Knossos Knossos (also Cnossos, both pronounced ; grc, Κνωσός, Knōsós, ; Linear B: ''Ko-no-so'') is the largest Bronze Age archaeological site on Crete and has been called Europe's oldest city. Settled as early as the Neolithic period, the na ...
(1900–1100 BC) depicts a fisherman carrying an octopus. The terrifyingly powerful Gorgon of
Greek mythology A major branch of classical mythology, Greek mythology is the body of myths originally told by the ancient Greeks, and a genre of Ancient Greek folklore. These stories concern the origin and nature of the world, the lives and activities o ...
may have been inspired by the octopus or squid, the octopus itself representing the severed head of Medusa, the beak as the protruding tongue and fangs, and its tentacles as the snakes. The Kraken are legendary sea monsters of giant proportions said to dwell off the coasts of Norway and Greenland, usually portrayed in art as giant octopuses attacking ships. Linnaeus included it in the first edition of his 1735 ''
Systema Naturae ' (originally in Latin written ' with the ligature æ) is one of the major works of the Swedish botanist, zoologist and physician Carl Linnaeus (1707–1778) and introduced the Linnaean taxonomy. Although the system, now known as binomial ...
''. One translation of the Hawaiian
creation myth A creation myth (or cosmogonic myth) is a symbolic narrative of how the world began and how people first came to inhabit it., "Creation myths are symbolic stories describing how the universe and its inhabitants came to be. Creation myths develo ...
the Kumulipo suggests that the octopus is the lone survivor of a previous age. The
Akkorokamui The is a gigantic octopus-like monster from Ainu folklore, similar to the Nordic Kraken, which supposedly lurks in Uchiura Bay The or is a bay southeast of the island of Hokkaido in northern Japan. It has also been known as Iburi Bay and ...
is a gigantic octopus-like monster from Ainu folklore, worshipped in
Shinto Shinto () is a religion from Japan. Classified as an East Asian religion by scholars of religion, its practitioners often regard it as Japan's indigenous religion and as a nature religion. Scholars sometimes call its practitioners ''Shint ...
. A battle with an octopus plays a significant role in
Victor Hugo Victor-Marie Hugo (; 26 February 1802 – 22 May 1885) was a French Romantic writer and politician. During a literary career that spanned more than sixty years, he wrote in a variety of genres and forms. He is considered to be one of the great ...
's 1866 book ''Travailleurs de la mer'' ('' Toilers of the Sea'').
Ian Fleming Ian Lancaster Fleming (28 May 1908 – 12 August 1964) was a British writer who is best known for his postwar ''James Bond'' series of spy novels. Fleming came from a wealthy family connected to the merchant bank Robert Fleming & Co., an ...
's 1966 short story collection ''
Octopussy and The Living Daylights ''Octopussy and The Living Daylights'' (sometimes published as ''Octopussy'') is the 14th and final James Bond book written by Ian Fleming in the Bond series. The book is a collection of short stories published posthumously in the United Kin ...
'', and the 1983 ''James Bond'' film were partly inspired by Hugo's book. Japanese erotic art, ''
shunga is a type of Japanese erotic art typically executed as a kind of ukiyo-e, often in woodblock print format. While rare, there are also extant erotic painted handscrolls which predate ukiyo-e. Translated literally, the Japanese word ''shunga' ...
'', includes
ukiyo-e Ukiyo-e is a genre of Japanese art which flourished from the 17th through 19th centuries. Its artists produced woodblock prints and paintings of such subjects as female beauties; kabuki actors and sumo wrestlers; scenes from history and folk ta ...
woodblock prints such as
Katsushika Hokusai , known simply as Hokusai, was a Japanese ukiyo-e artist of the Edo period, active as a painter and printmaker. He is best known for the woodblock print series '' Thirty-Six Views of Mount Fuji'', which includes the iconic print '' The Grea ...
's 1814 print ''Tako to ama'' ('' The Dream of the Fisherman's Wife''), in which an
ama diver are Japanese divers famous for collecting pearls, though traditionally their main catch is seafood. The vast majority of are women. Terminology There are several sea occupations that are pronounced "ama" and several words that refer to sea occ ...
is sexually intertwined with a large and a small octopus. The print is a forerunner of tentacle erotica. The biologist P. Z. Myers noted in his science blog, '' Pharyngula'', that octopuses appear in "extraordinary" graphic illustrations involving women, tentacles, and bare breasts. Since it has numerous arms emanating from a common centre, the octopus is often used as a symbol for a powerful and manipulative organisation, company, or country.


Danger

Octopuses generally avoid humans, but incidents have been verified. For example, a Pacific octopus, said to be nearly perfectly camouflaged, "lunged" at a diver and "wrangled" over his camera before it let go. Another diver recorded the encounter on video. All species are venomous, but only blue-ringed octopuses have venom that is lethal to humans. Bites are reported each year across the animals' range from Australia to the eastern Indo-Pacific Ocean. They bite only when provoked or accidentally stepped upon; bites are small and usually painless. The venom appears to be able to penetrate the skin without a puncture, given prolonged contact. It contains tetrodotoxin, which causes paralysis by blocking the transmission of
nerve impulse An action potential occurs when the membrane potential of a specific cell location rapidly rises and falls. This depolarization then causes adjacent locations to similarly depolarize. Action potentials occur in several types of animal cells, c ...
s to the muscles. This causes death by respiratory failure leading to cerebral anoxia. No antidote is known, but if breathing can be kept going artificially, patients recover within 24 hours. Bites have been recorded from captive octopuses of other species; they leave swellings which disappear in a day or two.


Fisheries

Octopus
fisheries Fishery can mean either the enterprise of raising or harvesting fish and other aquatic life; or more commonly, the site where such enterprise takes place ( a.k.a. fishing ground). Commercial fisheries include wild fisheries and fish farms, ...
exist around the world with total catches varying between 245,320 and 322,999 metric tons from 1986 to 1995. The world catch peaked in 2007 at 380,000 tons, and had fallen by a tenth by 2012. Methods to capture octopuses include pots, traps, trawls, snares, drift fishing, spearing, hooking and hand collection. Octopus is eaten in many cultures, such as on the Mediterranean and Asian coasts. The arms and sometimes other body parts are prepared in various ways, often varying by species or geography. Live octopuses are eaten in several countries around the world, including the US. Animal welfare groups have objected to this practice on the basis that octopuses can experience pain. Octopuses have a food conversion efficiency greater than that of chickens, making octopus aquaculture a possibility. Octopuses compete with human fisheries targeting other species, and even rob traps and nets for their catch; they may, themselves, be caught as
bycatch Bycatch (or by-catch), in the fishing industry, is a fish or other marine species that is caught unintentionally while fishing for specific species or sizes of wildlife. Bycatch is either the wrong species, the wrong sex, or is undersized or juve ...
if they cannot get away.


In science and technology

In classical Greece,
Aristotle Aristotle (; grc-gre, Ἀριστοτέλης ''Aristotélēs'', ; 384–322 BC) was a Greek philosopher and polymath during the Classical period in Ancient Greece. Taught by Plato, he was the founder of the Peripatetic school of ...
(384–322 BC) commented on the colour-changing abilities of the octopus, both for camouflage and for signalling, in his '' Historia animalium'': "The octopus ... seeks its prey by so changing its colour as to render it like the colour of the stones adjacent to it; it does so also when alarmed." Aristotle noted that the octopus had a hectocotyl arm and suggested it might be used in sexual reproduction. This claim was widely disbelieved until the 19th century. It was described in 1829 by the French zoologist
Georges Cuvier Jean Léopold Nicolas Frédéric, Baron Cuvier (; 23 August 1769 – 13 May 1832), known as Georges Cuvier, was a French naturalist and zoologist, sometimes referred to as the "founding father of paleontology". Cuvier was a major figure in na ...
, who supposed it to be a parasitic worm, naming it as a new species, ''Hectocotylus octopodis''. Other zoologists thought it a spermatophore; the German zoologist
Heinrich Müller Heinrich Müller may refer to: * Heinrich Müller (cyclist) (born 1926), Swiss cyclist * Heinrich Müller (footballer, born 1888) (1888–1957), Swiss football player and manager * Heinrich Müller (footballer, born 1909) (1909–2000), Austrian ...
believed it was "designed" to detach during copulation. In 1856 the Danish zoologist
Japetus Steenstrup Johannes Japetus Smith Steenstrup FRS(For) HFRSE (8 March 1813 – 20 June 1897) was a Danish zoologist, biologist, and professor. Life Born in Vang, Thy on 8 March 1813, he held a lectorate in mineralogy in Sorø until 1845 when he became ...
demonstrated that it is used to transfer sperm, and only rarely detaches. Octopuses offer many possibilities in biological research, including their ability to regenerate limbs, change the colour of their skin, behave intelligently with a distributed nervous system, and make use of 168 kinds of
protocadherin Protocadherins (Pcdhs) are the largest mammalian subgroup of the cadherin superfamily of homophilic cell-adhesion proteins. They were discovered by Shintaro Suzuki's group, when they used PCR to find new members of the cadherin family. The PCR fra ...
s (humans have 58), the proteins that guide the connections neurons make with each other. The California two-spot octopus has had its genome sequenced, allowing exploration of its molecular adaptations. Having independently evolved mammal-like intelligence, octopuses have been compared by the philosopher
Peter Godfrey-Smith Peter Godfrey-Smith (born 1965) is an Australian philosopher of science and writer, who is currently Professor of History and Philosophy of Science at the University of Sydney. He works primarily in philosophy of biology and philosophy of mind, ...
, who has studied the nature of intelligence, to hypothetical intelligent extraterrestrials. Their problem-solving skills, along with their mobility and lack of rigid structure enable them to escape from supposedly secure tanks in laboratories and public aquariums. Due to their intelligence, octopuses are listed in some countries as experimental animals on which surgery may not be performed without
anesthesia Anesthesia is a state of controlled, temporary loss of sensation or awareness that is induced for medical or veterinary purposes. It may include some or all of analgesia (relief from or prevention of pain), paralysis (muscle relaxation), ...
, a protection usually extended only to vertebrates. In the UK from 1993 to 2012, the common octopus (''Octopus vulgaris'') was the only invertebrate protected under the Animals (Scientific Procedures) Act 1986. In 2012, this legislation was extended to include all cephalopods in accordance with a general EU directive. Some
robotics Robotics is an interdisciplinary branch of computer science and engineering. Robotics involves design, construction, operation, and use of robots. The goal of robotics is to design machines that can help and assist humans. Robotics integrat ...
research is exploring biomimicry of octopus features. Octopus arms can move and sense largely autonomously without intervention from the animal's central nervous system. In 2015 a team in Italy built soft-bodied robots able to crawl and swim, requiring only minimal computation. In 2017 a German company made an arm with a soft pneumatically controlled
silicone A silicone or polysiloxane is a polymer made up of siloxane (−R2Si−O−SiR2−, where R = organic group). They are typically colorless oils or rubber-like substances. Silicones are used in sealants, adhesives, lubricants, medicine, cookin ...
gripper fitted with two rows of suckers. It is able to grasp objects such as a metal tube, a magazine, or a ball, and to fill a glass by pouring water from a bottle.


See also

*


Notes


References


Bibliography

* * *


Further reading

* *


External links


Octopuses – Overview
at the '' Encyclopedia of Life''
Octopoda
at the Tree of Life Web Project {{Authority control Articles containing video clips Commercial molluscs Extant Pennsylvanian first appearances Tool-using animals