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Common Octopus

Octopus vulgaris Cuvier 1797

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These animals have a life span of 12 to 24 months.

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Case, R. 1999. "Octopus vulgaris" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Octopus_vulgaris.html
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Robin J. Case, University of Michigan-Ann Arbor
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Conservation Status

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There is the potential for the overfishing of these animals, which threatens their proliferation. However, at this time, they are not at any specific risk.

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Case, R. 1999. "Octopus vulgaris" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Octopus_vulgaris.html
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Life Cycle

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The duration of embryonic development is related to temperature, as it is in all cephalopods, and it also depends on the size of the egg.

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Case, R. 1999. "Octopus vulgaris" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Octopus_vulgaris.html
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Benefits

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In 1975, some 121,000 tons of O. vulgaris were caught by fisheries. In 1976, the number was 137,000 tons.

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Case, R. 1999. "Octopus vulgaris" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Octopus_vulgaris.html
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Robin J. Case, University of Michigan-Ann Arbor
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Trophic Strategy

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Octopus vulgaris are active predators that feed primarily on gastropods and bivalves. Small hatchlings typically spend several weeks as active predators in the plankton before they settle down to the benthic mode of life at a size of about 0.2 grams.

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Case, R. 1999. "Octopus vulgaris" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Octopus_vulgaris.html
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Robin J. Case, University of Michigan-Ann Arbor
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Distribution

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This species has a world-wide distribution. It is abundant in the Mediterranean Sea, the Eastern Atlantic Ocean, and in Japanese waters.

Biogeographic Regions: indian ocean (Native ); atlantic ocean (Native ); pacific ocean (Native )

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Case, R. 1999. "Octopus vulgaris" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Octopus_vulgaris.html
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Robin J. Case, University of Michigan-Ann Arbor
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Habitat

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Octopus vulgaris is found in tropical, subtropical, and temperate waters between the surface and a depth of 100 to 150 meters. . It is not found in polar or subpolar regions. It lives in costal waters and the upper part of the continental shelf.

Habitat Regions: temperate ; tropical

Aquatic Biomes: benthic ; reef ; coastal

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Case, R. 1999. "Octopus vulgaris" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Octopus_vulgaris.html
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Robin J. Case, University of Michigan-Ann Arbor
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Morphology

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Reach 1-3 feet in length including arms. The skin is smooth. Like other octopuses, members of this species have 8 arms that are lined with suckers, and they lack any internal shell.

Other Physical Features: ectothermic ; bilateral symmetry

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Case, R. 1999. "Octopus vulgaris" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Octopus_vulgaris.html
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Robin J. Case, University of Michigan-Ann Arbor
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Reproduction

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Octopus vulgaris has individuals of both sexes. During mating, the male approaches the female, who fends him off for a while, but then accepts him. He sits next to her or mounts her, inserting the hectocotylus in her mantle cavity to pass the spermatophores. They may copulate for several hours. The same pair often repeat mating over a period of a week or so, but a male copulates with other females and a female accepts other males. Mating often occurs when the females are immature. Only females ready to lay eggs consistently fend off the males.

Mating System: polygynandrous (promiscuous)

Females become restless and search for a sheltered place where they can lay and brood the eggs without disturbance. The spermatophores are placed in the oviducts and empty cases are discarded. Fertilization takes place in the oviductal glands as the mature eggs pass through them on thir way out of the oviducts. Two secretions from the oviductal glands, together with the mucus, are used to stick the egg stalks together in strings and attach these to a substrate. Eggs are laid in shallow water. They are always attached to a substrate. On rocky shores, females find a hole, a crevice or sheltered place and they often protect their homes with shells, stones and other solid objects that they gather. Coral reefs provide suitable shelter. On sandy or muddy bottom, eggs are laid in empty mollusc shells or in man-made objects such as cans, tins, bottles, tires, boots, and amphorae . In tropical and subtropical waters, eggs are laid throughout the year. The total number of eggs laid by a female varies from 100,000 to 500,000. During egg laying and subsequent brooding, the female rarely leaves the egg mass. She usually does not feed during the entire period of spawning and brooding, which can be as long as 4-5 months at low temperatures. Egg care includes cleaning the eggs with the arm tips and directing jets of water from the funnel through the strings. Intruders, including potential prey, are pushed away, although crabs left overnight may occasionally be eaten. As a rule, females die shortly after the hatching of the last embryos after losing one-third of their pre-spawning weight.

Range number of offspring: 100000 to 500000.

Key Reproductive Features: semelparous ; seasonal breeding ; year-round breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; fertilization (Internal ); oviparous

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Case, R. 1999. "Octopus vulgaris" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Octopus_vulgaris.html
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Robin J. Case, University of Michigan-Ann Arbor
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Biology

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Like all cephalopods, the common octopus is an intelligent active predator (4). They have modified salivary glands that produce venom used to incapacitate prey. It is often easy to identify what a common octopus has been feeding on, as they leave piles of debris known as 'middens' around the entrance of the protective lair in which they live. These middens consist of debris from a range of species and often include mollusc shells and the carapaces of crabs and other crustaceans (5). All cephalopods are good swimmers, and are able to move rapidly by jet propulsion when threatened; water is rapidly expelled through a funnel which causes the octopus to be propelled away rapidly (3). Cephalopods are also able to mask themselves as they escape with a cloud of ink released into the water (2).
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Conservation

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Conservation action has not been targeted at this common species.
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Description

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The cephalopods (meaning 'head-footed) are a group of molluscs that contain the octopuses, squid and cuttlefish, and are probably the most intelligent of all invertebrates. They have well-developed heads, with large complex eyes and mouths that feature beak-like jaws. All octopuses have eight tentacle-like arms; indeed 'octopus' derives from the Greek for 'eight-footed' (3). The common octopus usually measures around 60 centimetres in length, but it can grow up to 1 metre (2). It is able to change its colour depending on its mood and situation, but individuals are usually greyish-yellow or brownish-green with extensive mottling. They are often very well camouflaged (2). The body is warty, and the thick arms bear two rows of suckers (4).
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Habitat

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Occurs along rocky coasts in the shallow sublittoral zone (4).
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Range

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This octopus is found from the southern North Sea down to South Africa. It also occurs in the Mediterranean (2). It reaches the north-eastern extreme of its range in Britain where it is found only around the coasts of the south and south west (4).
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Status

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Common and widespread (2).
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Threats

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Not currently threatened.
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Diagnostic Description

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Medium to large sized; animal chunky in appearance.

Arms stout, of about equal length and thickness, dorsal pair of arms slightly shorter; shortened right arm III of males hectocotylized by modification of tip into a very small, spoon-shaped ligula; ligula index (length of ligula expressed as percentage of length of hectocotylized arm) less than 2.5;

7 to 11 gill lamellae on outer side of of gill, including terminal lamella.

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FAO Species catalogue VOL. 3. Cephalopods of the world An Annotated and Illustrated Catalogue of Species of Interest to FisheriesClyde F.E. Roper Michael J. Sweeney Cornelia E. Nauen 1984. FAO Fisheries Synopsis No. 125, Volume 3
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Distribution

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Worldwide in temperate and tropical waters; limits unknown.
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FAO Species catalogue VOL. 3. Cephalopods of the world An Annotated and Illustrated Catalogue of Species of Interest to FisheriesClyde F.E. Roper Michael J. Sweeney Cornelia E. Nauen 1984. FAO Fisheries Synopsis No. 125, Volume 3
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Size

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Maximum total length 1.2 m in females and to 1.3 m in males; maximum weight 10 kg; common to 3 kg. In the western Mediterranean, mantle length at first maturity is about 9.5 cm in males, 13.5 cm in females.
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FAO Species catalogue VOL. 3. Cephalopods of the world An Annotated and Illustrated Catalogue of Species of Interest to FisheriesClyde F.E. Roper Michael J. Sweeney Cornelia E. Nauen 1984. FAO Fisheries Synopsis No. 125, Volume 3
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Brief Summary

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A benthic, neritic species occurring from the coastline to the outer edge of the continental shelf(in depths from 0 to 200 m), where it is found in a variety of habitats, such as rocks, coral reefs, and grass beds.It is inactive in waters of 7°C and colder.

Throughout its distribution range, this species is known to undertake limited seasonal migrations, usually overwintering in deeper waters and occurring in shallower waters during summer. In the western Mediterranean, large mature or maturing individuals migrate inshore in early spring, followed later on by smaller, immature individuals. These two groups begin their retreat into deeper waters by August/September and November/ December respectively. Similar migration patterns are found in other sea areas. Two spawning peaks per year can be observed for this species throughout its distributional range: in the Mediterranean and the Inland Sea of Japan, the first occurs in April/ May corresonding to the group migrating in shore in spring (most important in the Mediterranean) and the second in October, corresponding to the group migrating in autumn (most important in Japan); off West Africa, around Cape Blanc, the first spawning peak occurs in May/June and the second (more important) in September. Females may produce between 120 000 and 400 000 eggs little longer than 2 mm, which they deposit in strings in crevices or holes, usually in shallow waters. Spawning may extend up to 1 month. During the brooding period (25 to 65 days), females almost cease feeding and many die after the hatching of the larvae. The hatchlings are pelagic, but settle to benthic life after about 40 days at a minimum size of approximately 12 mm.

In the Inland Sea of Japan, common octopus reaches about 1 kg weight in 4 months; in the western Mediterranean it grows from 3 to about 20 cm in 17 months. A von Bertalanffy growth expression is given for the Mediterranean and eastern Atlantic populations by Guerra (1979). Food consists of bivalves and crustaceans.Larvae and juveniles are preyed upon by albacore (Thunnus alalunga) etc., and adults by benthic finfishes.

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FAO Species catalogue VOL. 3. Cephalopods of the world An Annotated and Illustrated Catalogue of Species of Interest to FisheriesClyde F.E. Roper Michael J. Sweeney Cornelia E. Nauen 1984. FAO Fisheries Synopsis No. 125, Volume 3
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Benefits

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World catches attributed to O. vulgaris declined from peaks in the late sixties (more than 100 000 t per year) to 20 000 to 30 000 t in recent years (FAO, 1983). Part of the nonidentified world catches of Octopus oscillating between 120 000 and 160 000 t annually, also pertain to this species (including part of the nearly 45 000 t caught by Japan in the Inland Sea, Fishing Area 61, and most of the catches (between 40 000 and 50 000 t) taken by Spanish vessels on the Sahara Banks off West Africa, Fishing Area 34). The species is highly desirable and commands high prices throughout its distributional range and supports artisanal as well as industrial fisheries. It is taken mainly with lures, hooks and lines, pots, spears and otter trawls. In the Inland Sea of Japan, some 12 000 to 17 000 pots are laid out annually to provide shelter and enhance egg conservation as a measure to safeguard recruitment of the fishable stock. This species is marketed fresh, frozen and dried salted, mostly for human consumption.The total catch reported for this species to FAO for 1999 was 34 262 t. The countries with the largest catches were Mexico (19 081 t) and Italy (8 844 t).
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FAO Species catalogue VOL. 3. Cephalopods of the world An Annotated and Illustrated Catalogue of Species of Interest to FisheriesClyde F.E. Roper Michael J. Sweeney Cornelia E. Nauen 1984. FAO Fisheries Synopsis No. 125, Volume 3
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Comprehensive Description

provided by Smithsonian Contributions to Zoology
Octopus vulgaris Cuvier,

Octopus vulgare Cuvier, [1797]:381 [coast of France].—Hemming, 1954:278, 288, 291–293.

Octopus vulgaris Lamarck, 1798:130.—Stearns, 1893:339.—Robson, 1929:57–62.—Adam, 1952:117–125.—Pickford, 1955:156.—Lane, 1957:221.

Octopus occidentalis Hoyle, 1886:77.

PREVIOUS ASCENSION RECORDS.—Hoyle (1886); dredged in 20–30 fathoms, according to Stearns(1893).

PRESENT MATERIAL.—3 (ML 22, 28, 80 mm) ASC 5; 2 (ML 11, 42 mm) ASC 11, 21; 1 (ML 28 mm) ASC 23, R. B. Manning, May 1971.

DISTRIBUTION.—North and South Atlantic (east and west), Mediterranean Sea, North Sea (Adam, 1952).
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Rosewater, Joseph. 1975. "An annotated list of the marine mollusks of Ascension Island, South Atlantic Ocean." Smithsonian Contributions to Zoology. 1-41. https://doi.org/10.5479/si.00810282.189

Comprehensive Description

provided by Smithsonian Contributions to Zoology
Octopus vulgaris Cuvier, 1797

Octopus vulgaris Cuvier, 1797:380, pl. 9: fig. 2.

Sepia rugosa Bosc, 1792:24, pl. 5: figs. 1, 2.

Octopus granulatus Lamarck, 1798:130.

Octopus vulgaris Lamarck, 1798:130.

Octopus tuberculatus Risso, 1826:3.

Octopus Cassiopeia Gray, 1849:9.

Octopus troscheli Targioni-Tozzetti, 1869:17.

Octopus rugosus.—Robson, 1929:63, figs. 8, 9, pl. 2: fig. 3.

A full synonymy is presented in Hochberg and Mangold (MS).

DIAGNOSIS.—Mediterranean: Animals of medium to large size (males 20–250 mm ML, females 20–200 mm ML). Mantle broadly oval to saccular, widest in posterior (MWI 62.2–80.0). Neck slightly constricted. Head distinctly narrower than mantle (HWI 32.5–53.5). Mantle aperture wide. Funnel elongate, opening relatively narrow (FLI 30.0–48.2); funnel organ W-shaped, lateral limbs usually, but not always, slightly shorter than median limbs. Arms long, robust, tapering to narrow rounded tips (ALI 76.3–88.7; MAI 18.0–25.2). Lateral arms distinctly longer than median arms; arm formula III > II > IV > I or III = II > IV > I. Suckers very mobile, medium-sized (SIn males 12.5–13.5, females 9.7–10.3); 2 to 3 enlarged suckers (among 15th to 19th) on lateral arms in both sexes (SIe males 18.2–21.1, females 10.5–11.2). Right arm III of male hectocotylized (HALI 56.0–72.2), shorter than opposite arm (OAI 75.2–82.0), bearing 140–180 suckers. Ligula small to minute (LLI 1.2–2.1), tip narrow, transverse striations faint; calamus distinct, relatively long (CLI 47–52). Web moderately shallow (WDI 16.5–18.5); web formula C > D > B > E > A, often variable but sector A always shallowest. Ink sac present, superficial or partially embedded in digestive gland. Gills with 9–11 lamellae per outer demibranch. Mature ova 2.2 mm long, 1.0 mm wide; stalk about 2.5 times ova length. Penis moderately long (PLI 15–21), with small, rounded diverticulum. Spermatophores of moderate size (SpLI 31–81). Radula with A2 to As seriation of rachidian. Skin firm, covered with papillae or smooth. Color in life variable, brown yellow, red brown, dark brown, or greyish.

ORIGINAL DESCRIPTION.—Cuvier, 1797:380, pl. 9: fig. 2.

TYPE LOCALITY.—Not stated in original description, presumed to be the western Mediterranean Sea.

TYPE.—Not designated, presumed to be not extant (Lu et al., 1995).

DISTRIBUTION.—Mediterranean Sea: Western and eastern basins, Adriatic Sea. Eastern Atlantic Ocean: South coast of England; west, south, and southeast coast of Africa; Azores, Canary Islands, Cape Verde Islands, and St. Helena Islands. Western Atlantic Ocean: Described from numerous localities.

This is a shallow-water species that lives on the continental shelf from the surface to 100 (150) m, rarely deeper, on sandy, rocky, or muddy bottoms.
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Voss, N. A. and Sweeney, M. J. 1998. "Systematics and Biogeography of cephalopods. Volume II." Smithsonian Contributions to Zoology. 277-599. https://doi.org/10.5479/si.00810282.586.277

Common octopus

provided by wikipedia EN

The common octopus (Octopus vulgaris) is a mollusc belonging to the class Cephalopoda. Octopus vulgaris is the most studied of all octopus species. It is cosmopolitan, that is, a global species, which ranges from the eastern Atlantic, extends from the Mediterranean Sea and the southern coast of England, to the southern coast of South Africa. It also occurs off the Azores, Canary Islands, and Cape Verde Islands. The species is also common in the Western Atlantic. The common octopus hunts at dusk. Crabs, crayfish, and bivalve molluscs (two-shelled, such as cockles) are preferred, although the octopus eats almost anything it can catch. It is able to change colour to blend in with its surroundings, and is able to jump upon any unwary prey that strays across its path. Using its beak, it is able to break into the shells of shelled molluscs. Training experiments have shown the common octopus can distinguish the brightness, size, shape, and horizontal or vertical orientation of objects.

Characteristics

Octopus vulgaris Merculiano.jpg
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Common octopus in the Aquarium of Seville, Spain
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O. vulgaris from the Mediterranean Sea
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Common octopus, Staatliches Museum für Naturkunde Karlsruhe, Germany
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Common octopus of Croatia
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Common octopus near Crete

Octopus vulgaris grows to 25 cm (10 inches) in mantle length with arms up to 1 m (3.3 feet) long.[2] It lives for a couple of years and may weigh up to 9 kg (20 pounds).[3][4] Mating may become cannibalistic.[5] O. vulgaris is caught by bottom trawls on a huge scale off the northwestern coast of Africa. More than 20,000 tonnes (22,000 short tons) are harvested annually.[2]

The common octopus hunts at dusk. Crabs, crayfish, and bivalve molluscs (such as cockles) are preferred, although the octopus eats almost anything it can catch. It is able to change colour to blend in with its surroundings, and is able to jump upon any unwary prey that strays across its path. Using its beak, it is able to break into the shells of shelled molluscs. It also possesses venom to subdue its prey.[6]

They have evolved to have large nervous systems and brains. An individual has about 500 million neurons in its body, almost comparable to dogs. They are intelligent enough to distinguish brightness, navigate mazes, recognize individual people, learn how to unscrew a jar or raid lobster traps.[7][8][9] O. vulgaris was the first invertebrate animal protected by the Animals (Scientific Procedures) Act 1986 in the UK.[10]

Physiology

Habitat and demands

The common octopus has world wide distribution in tropical, subtropical and temperate waters throughout the world. [11][12][13] They prefer the floor of relatively shallow, rocky, coastal waters, often no deeper than 200 m (660 feet).[13] Although they prefer around 36 grams per liter (0.0013 lb/cu in), salinity throughout their global habitat is found to be between roughly 30 and 45 grams per liter (0.0011 and 0.0016 lb/cu in).[14] They are exposed to a wide variety of temperatures in their environments, but their preferred temperature ranges from about 15 to 16 °C (59 to 61 °F).[14] In especially warm seasons, the octopus can often be found deeper than usual to escape the warmer layers of water.[15] In moving vertically throughout the water, the octopus is subjected to various pressures and temperatures, which affect the concentration of oxygen available in the water.[14] This can be understood through Henry's law, which states that the concentration of a gas in a substance is proportional to pressure and solubility, which is influenced by temperature. These various discrepancies in oxygen availability introduce a requirement for regulation methods.[16]

Primarily, the octopus situates itself in a shelter where a minimal amount of its body is presented to the external water.[17] When it does move, most of the time it is along the ocean or sea floor, in which case the underside of the octopus is still obscured.[17] This crawling increases metabolic demands greatly, requiring they increase their oxygen intake by roughly 2.4 times the amount required for a resting octopus.[18] This increased demand is met by an increase in the stroke volume of the octopus' heart.[19]

The octopus does sometimes swim throughout the water, exposing itself completely.[14] In doing so, it uses a jet mechanism that involves creating a much higher pressure in its mantle cavity that allows it to propel itself through the water.[19] As the common octopus' heart and gills are located within its mantle, this high pressure also constricts and puts constraints on the various vessels that are returning blood to the heart.[19] Ultimately, this creates circulation issues and is not a sustainable form of transportation, as the octopus cannot attain an oxygen intake that can balance the metabolic demands of maximum exertion.[19]

Respiration

The octopus uses gills as its respiratory surface. The gill is composed of branchial ganglia and a series of folded lamellae. Primary lamellae extend out to form demibranches and are further folded to form the secondary free folded lamellae, which are only attached at their tops and bottoms.[20] The tertiary lamellae are formed by folding the secondary lamellae in a fan-like shape.[20] Water moves slowly in one direction over the gills and lamellae, into the mantle cavity and out of the octopus' funnel.[21]

The structure of the octopus' gills allows for a high amount of oxygen uptake; up to 65% in water at 20 °C (68 °F).[21] The thin skin of the octopus accounts for a large portion of in-vitro oxygen uptake; estimates suggest around 41% of all oxygen absorption is through the skin when at rest.[17] This number is affected by the activity of the animal – the oxygen uptake increases when the octopus is exercising due to its entire body being constantly exposed to water, but the total amount of oxygen absorption through skin is actually decreased to 33% as a result of the metabolic cost of swimming.[17] When the animal is curled up after eating, its absorption through its skin can drop to 3% of its total oxygen uptake.[17] The octopus' respiratory pigment, hemocyanin, also assists in increasing oxygen uptake.[16] Octopuses can maintain a constant oxygen uptake even when oxygen concentrations in the water decrease to around 3.5 kPa (0.51 psi)[21] or 31.6% saturation (standard deviation 8.3%).[16] If oxygen saturation in sea water drops to about 1–10% it can be fatal for Octopus vulgaris depending on the weight of the animal and the water temperature.[16] Ventilation may increase to pump more water carrying oxygen across the gills but due to receptors found on the gills the energy use and oxygen uptake remains at a stable rate.[21] The high percent of oxygen extraction allows for energy saving and benefits for living in an area of low oxygen concentration.[20]

Water is pumped into the mantle cavity of the octopus, where it comes into contact with the internal gills. The water has a high concentration of oxygen compared to the blood returning from the veins, so oxygen diffuses into the blood. The tissues and muscles of the octopus use oxygen and release carbon dioxide when breaking down glucose in the Krebs cycle. The carbon dioxide then dissolves into the blood or combines with water to form carbonic acid, which decreases blood pH. The Bohr effect explains why oxygen concentrations are lower in venous blood than arterial blood and why oxygen diffuses into the bloodstream. The rate of diffusion is affected by the distance the oxygen has to travel from the water to the bloodstream as indicated by Fick's laws of diffusion. Fick's laws explain why the gills of the octopus contain many small folds that are highly vascularised. They increase surface area, thus also increase the rate of diffusion. The capillaries that line the folds of the gill epithelium have a very thin tissue barrier (10 µm), which allows for fast, easy diffusion of the oxygen into the blood.[22] In situations where the partial pressure of oxygen in the water is low, diffusion of oxygen into the blood is reduced,[23] Henry's law can explain this phenomenon. The law states that at equilibrium, the partial pressure of oxygen in water will be equal to that in air; but the concentrations will differ due to the differing solubility. This law explains why O. vulgaris has to alter the amount of water cycled through its mantle cavity as the oxygen concentration in water changes.[21]

The gills are in direct contact with water – carrying more oxygen than the blood – that has been brought into the mantle cavity of the octopus. Gill capillaries are quite small and abundant, which creates an increased surface area that water can come into contact with, thus resulting in enhanced diffusion of oxygen into the blood. Some evidence indicates that lamellae and vessels within the lamellae on the gills contract to aid in propelling blood through the capillaries.[24]

Circulation

The octopus has three hearts, one main two-chambered heart charged with sending oxygenated blood to the body and two smaller branchial hearts, one next to each set of gills. The circulatory circuit sends oxygenated blood from the gills to the atrium of the systemic heart, then to its ventricle which pumps this blood to the rest of the body. Deoxygenated blood from the body goes to the branchial hearts which pump the blood across the gills to oxygenate it, and then the blood flows back to the systemic atrium for the process to begin again.[25] Three aortae leave the systemic heart, two minor ones (the abdominal aorta and the gonadal aorta) and one major one, the dorsal aorta which services most of the body.[26] The octopus also has large blood sinuses around its gut and behind its eyes that function as reserves in times of physiologic stress.[27]

The octopus' heart rate does not change significantly with exercise, though temporary cardiac arrest of the systemic heart can be induced by oxygen debt, almost any sudden stimulus, or mantle pressure during jet propulsion.[28] Its only compensation for exertion is through an increase in stroke volume of up to three times by the systemic heart,[28] which means it suffers an oxygen debt with almost any rapid movement.[28][29] The octopus is, however, able to control how much oxygen it pulls out of the water with each breath using receptors on its gills,[21] allowing it to keep its oxygen uptake constant over a range of oxygen pressures in the surrounding water.[28] The three hearts are also temperature and oxygen dependent and the beat rhythm of the three hearts are generally in phase with the two branchial hearts beating together followed by the systemic heart.[25] The Frank–Starling law also contributes to overall heart function, through contractility and stroke volume, since the total volume of blood vessels must be maintained, and must be kept relatively constant within the system for the heart to function properly.[30]

The blood of the octopus is composed of copper-rich hemocyanin, which is less efficient than the iron-rich hemoglobin of vertebrates, thus does not increase oxygen affinity to the same degree.[31] Oxygenated hemocyanin in the arteries binds to CO
2
, which is then released when the blood in the veins is deoxygenated. The release of CO
2
into the blood causes it to acidify by forming carbonic acid.[32] The Bohr effect explains that carbon dioxide concentrations affect the blood pH and the release or intake of oxygen. The Krebs cycle uses the oxygen from the blood to break down glucose in active tissues or muscles and releases carbon dioxide as a waste product, which leads to more oxygen being released. Oxygen released into the tissues or muscles creates deoxygenated blood, which returns to the gills in veins. The two brachial hearts of the octopus pump blood from the veins through the gill capillaries. The newly oxygenated blood drains from the gill capillaries into the systemic heart, where it is then pumped back throughout the body.[25]

Blood volume in the octopus' body is about 3.5% of its body weight[27] but the blood's oxygen-carrying capacity is only about 4 volume percent.[28] This contributes to their susceptibility to the oxygen debt mentioned before. Shadwick and Nilsson[29] concluded that the octopus circulatory system is "fundamentally unsuitable for high physiologic performance". Since the binding agent is found within the plasma and not the blood cells, a limit exists to the oxygen uptake that the octopus can experience. If it were to increase the hemocyanin within its blood stream, the fluid would become too viscous for the myogenic[33] hearts to pump.[30] Poiseuille's law explains the rate of flow of the bulk fluid throughout the entire circulatory system through the differences of blood pressure and vascular resistance.[30]

Like those of vertebrates, octopus blood vessels are very elastic, with a resilience of 70% at physiologic pressures. They are primarily made of an elastic fibre called octopus arterial elastomer, with stiffer collagen fibres recruited at high pressure to help the vessel maintain its shape without over-stretching.[34] Shadwick and Nilsson[29] theorized that all octopus blood vessels may use smooth-muscle contractions to help move blood through the body, which would make sense in the context of them living under water with the attendant pressure.

The elasticity and contractile nature of the octopus aorta serves to smooth out the pulsing nature of blood flow from the heart as the pulses travel the length of the vessel, while the vena cava serves in an energy-storage capacity.[29] Stroke volume of the systemic heart changes inversely with the difference between the input blood pressure through the vena cava and the output back pressure through the aorta.

Osmoregulation

 src=
Common octopus in Santander, Spain.

The hemolymph, pericardial fluid and urine of cephalopods, including the common octopus, are all isosmotic with each other, as well as with the surrounding sea water.[35] It has been suggested that cephalopods do not osmoregulate, which would indicate that they are conformers.[35] This means that they adapt to match the osmotic pressure of their environment, and because there is no osmotic gradient, there is no net movement of water from the organism to the seawater, or from the seawater into the organism.[35] Octopuses have an average minimum salinity requirement of 27 g/L (0.00098 lb/cu in), and that any disturbance introducing significant amounts of fresh water into their environment can prove fatal.[36]

In terms of ions, however, a discrepancy does seem to occur between ionic concentrations found in the seawater and those found within cephalopods.[35] In general, they seem to maintain hypoionic concentrations of sodium, calcium, and chloride in contrast to the salt water.[35] Sulfate and potassium exist in a hypoionic state, as well, with the exception of the excretory systems of cephalopods, where the urine is hyperionic.[35] These ions are free to diffuse, and because they exist in hypoionic concentrations within the organism, they would be moving into the organism from the seawater.[35] The fact that the organism can maintain hypoionic concentrations suggests not only that a form of ionic regulation exists within cephalopods, but also that they also actively excrete certain ions such as potassium and sulfate to maintain homeostasis.[35]

O. vulgaris has a mollusc-style kidney system, which is very different from mammals. The system is built around an appendage of each branchial heart, which is essentially an extension of its pericardium.[35] These long, ciliated ducts filter the blood into a pair of kidney sacs, while actively reabsorbing glucose and amino acids into the bloodstream.[35] The renal sacs actively adjust the ionic concentrations of the urine, and actively add nitrogenous compounds and other metabolic waste products to the urine.[35] Once filtration and reabsorption are complete, the urine is emptied into O. vulgaris' mantle cavity via a pair of renal papillae, one from each renal sac.[35]

Temperature and body size directly affect the oxygen consumption of O. vulgaris, which alters the rate of metabolism.[18] When oxygen consumption decreases, the amount of ammonia excretion also decreases due to the slowed metabolic rate.[18] O. vulgaris has four different fluids found within its body: blood, pericardial fluid, urine, and renal fluid. The urine and renal fluid have high concentrations of potassium and sulphate, but low concentrations of chloride. The urine has low calcium concentrations, which suggests it has been actively removed. The renal fluid has similar calcium concentrations to the blood. Chloride concentrations are high in the blood, while sodium varies. The pericardial fluid has concentrations of sodium, potassium, chlorine and calcium similar to that of the salt water supporting the idea that O. vulgaris does not osmoregulate, but conforms. However, it has lower sulphate concentrations.[35] The pericardial duct contains an ultrafiltrate of the blood known as the pericardial fluid, and the rate of filtration is partly controlled by the muscle- and nerve-rich branchial hearts.[35] The renal appendages move nitrogenous and other waste products from the blood to the renal sacs, but do not add volume. The renal fluid has a higher concentration of ammonia than the urine or the blood, thus the renal sacs are kept acidic to help draw the ammonia from the renal appendages. The ammonia diffuses down its concentration gradient into the urine or into the blood, where it gets pumped through the branchial hearts and diffuses out the gills.[35] The excretion of ammonia by O. vulgaris makes them ammonotelic organisms. Aside from ammonia, a few other nitrogenous waste products have been found to be excreted by O. vulgaris such as urea, uric acid, purines, and some free amino acids, but in smaller amounts.[35]

Within the renal sacs, two recognized and specific cells are responsible for the regulation of ions. The two kinds of cells are the lacuna-forming cells and the epithelial cells that are typical to kidney tubules. The epithelia cells are ciliated, cylindrical, and polarized with three distinct regions. These three regions are apical, middle cytoplasmic, and basal lamina. The middle cytoplasmic region is the most active of the three due to the concentration of multiple organelles within, such as mitochondria and smooth and rough endoplasmic reticulum, among others. The increase of activity is due to the interlocking labyrinth of the basal lamina creating a crosscurrent activity similar to the mitochondrial-rich cells found in teleost marine fish. The lacuna-forming cells are characterized by contact to the basal lamina, but not reaching the apical rim of the associated epithelial cells and are located in the branchial heart epithelium. The shape varies widely and are occasionally more electron-dense than the epithelial cells, seen as a "diffused kidney" regulating ion concentrations.[37]

One adaptation that O. vulgaris has is some direct control over its kidneys.[35] It is able to switch at will between the right or left kidney doing the bulk of the filtration, and can also regulate the filtration rate so that the rate does not increase when the animal's blood pressure goes up due to stress or exercise.[35] Some species of octopuses, including O. vulgaris, also have a duct that runs from the gonadal space into the branchial pericardium.[35] Wells[35] theorized that this duct, which is highly vascularized and innervated, may enable the reabsorption of important metabolites from the ovisac fluid of pregnant females by directing this fluid into the renal appendages.

Thermoregulation

As an oceanic organism, O. vulgaris experiences a temperature variance due to many factors, such as season, geographical location, and depth.[38] For example, octopuses living around Naples may experience a temperature of 25 °C (77 °F) in the summer and 15 °C (59 °F) in the winter.[38] These changes would occur quite gradually, however, and thus would not require any extreme regulation.

The common octopus is a poikilothermic, eurythermic ectotherm, meaning that it conforms to the ambient temperature.[39] This implies that no real temperature gradient is seen between the organism and its environment, and the two are quickly equalized. If the octopus swims to a warmer locale, it gains heat from the surrounding water, and if it swims to colder surroundings, it loses heat in a similar fashion.

O. vulgaris can apply behavioral changes to manage wide varieties of environmental temperatures. Respiration rate in octopods is temperature-sensitive – respiration increases with temperature.[40] Its oxygen consumption increases when in water temperatures between 16 and 28 °C (61 and 82 °F), reaches a maximum at 28 °C (82 °F), and then begins to drop at 32 °C (90 °F).[40] The optimum temperature for metabolism and oxygen consumption is between 18 and 24 °C (64 and 75 °F).[40] Variations in temperature can also induce a change in hemolymph protein levels along oxygen consumption.[40] As temperature increases, protein concentrations increase in order to accommodate the temperature. Also the cooperativity of hemocyanin increases, but the affinity decreases.[41] Conversely, a decrease in temperature results in a decrease in respiratory pigment cooperativity and increase in affinity.[41] The slight rise in P50 that occurs with temperature change allows oxygen pressure to remain high in the capillaries, allowing for elevated diffusion of oxygen into the mitochondria during periods of high oxygen consumption.[41] The increase in temperature results in higher enzyme activity, yet the decrease in hemocyanin affinity allows enzyme activity to remain constant and maintain homeostasis. The highest hemolymph protein concentrations are seen at 32 °C (90 °F) and then drop at temperatures above this.[40] Oxygen affinity in the blood decreases by 0.20 kPa/°C (0.016 psi/°F) at a pH of 7.4.[41] The octopod's thermal tolerance is limited by its ability to consume oxygen, and when it fails to provide enough oxygen to circulate at extreme temperatures the effects can be fatal.[40] O. vulgaris has a pH-independent venous reserve that represents the amount of oxygen that remains bound to the respiratory pigment at constant pressure of oxygen. This reserve allows the octopus to tolerate a wide range of pH related to temperature.[41]

As a temperature conformer,[42] O. vulgaris does not have any specific organ or structure dedicated to heat production or heat exchange. Like all animals, they produce heat as a result of ordinary metabolic processes such as digestion of food,[38] but take no special means to keep their body temperature within a certain range. Their preferred temperature directly reflects the temperature to which they are acclimated.[42] They have an acceptable ambient temperature range of 13–28 °C (55–82 °F),[42] with their optimum for maximum metabolic efficiency being about 20 °C (68 °F).[39]

As ectothermal animals, common octopuses are highly influenced by changes in temperature. All species have a thermal preference where they can function at their basal metabolic rate.[42] The low metabolic rate allows for rapid growth, thus these cephalopods mate as the water becomes closest to the preferential zone. Increasing temperatures cause an increase in oxygen consumption by O. vulgaris.[18] Increased oxygen consumption can be directly related to the metabolic rate, because the breakdown of molecules such as glucose requires an input of oxygen, as explained by the Krebs cycle. The amount of ammonia excreted conversely decreases with increasing temperature.[18] The decrease in ammonia being excreted is also related to the metabolism of the octopus due to its need to spend more energy as the temperature increases. Octopus vulgaris will reduce the amount of ammonia excreted in order to use the excess solutes that it would have otherwise excreted due to the increased metabolic rate. Octopuses do not regulate their internal temperatures until it reaches a threshold where they must begin to regulate to prevent death.[18] The increase in metabolic rate shown with increasing temperatures is likely due to the octopus swimming to shallower or deeper depths to stay within its preferential temperature zone.

See also

References

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  33. ^ Excitation-contraction coupling
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Common octopus: Brief Summary

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The common octopus (Octopus vulgaris) is a mollusc belonging to the class Cephalopoda. Octopus vulgaris is the most studied of all octopus species. It is cosmopolitan, that is, a global species, which ranges from the eastern Atlantic, extends from the Mediterranean Sea and the southern coast of England, to the southern coast of South Africa. It also occurs off the Azores, Canary Islands, and Cape Verde Islands. The species is also common in the Western Atlantic. The common octopus hunts at dusk. Crabs, crayfish, and bivalve molluscs (two-shelled, such as cockles) are preferred, although the octopus eats almost anything it can catch. It is able to change colour to blend in with its surroundings, and is able to jump upon any unwary prey that strays across its path. Using its beak, it is able to break into the shells of shelled molluscs. Training experiments have shown the common octopus can distinguish the brightness, size, shape, and horizontal or vertical orientation of objects.

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Distribution

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circum-global
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bibliographic citation
Check List of European Marine Mollusca (CLEMAM). Backeljau, T. (1986). Lijst van de recente mariene mollusken van België [List of the recent marine molluscs of Belgium]. Koninklijk Belgisch Instituut voor Natuurwetenschappen: Brussels, Belgium. 106 pp. Stocks, K. 2009. Seamounts Online: an online information system for seamount biology. Version 2009-1. World Wide Web electronic publication. van der Land, J. (ed). (2008). UNESCO-IOC Register of Marine Organisms (URMO). North-West Atlantic Ocean species (NWARMS) van der Land, J. (ed). (2008). UNESCO-IOC Register of Marine Organisms (URMO).
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Jacob van der Land [email]

Distribution

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Western Atlantic: Occurs primarily south of Cape Hatteras, but it is distributed as far north as Connecticut.
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Check List of European Marine Mollusca (CLEMAM). Backeljau, T. (1986). Lijst van de recente mariene mollusken van België [List of the recent marine molluscs of Belgium]. Koninklijk Belgisch Instituut voor Natuurwetenschappen: Brussels, Belgium. 106 pp. Stocks, K. 2009. Seamounts Online: an online information system for seamount biology. Version 2009-1. World Wide Web electronic publication. van der Land, J. (ed). (2008). UNESCO-IOC Register of Marine Organisms (URMO). North-West Atlantic Ocean species (NWARMS) van der Land, J. (ed). (2008). UNESCO-IOC Register of Marine Organisms (URMO).
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Mary Kennedy [email]

Habitat

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coastal to shelf
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Check List of European Marine Mollusca (CLEMAM). Backeljau, T. (1986). Lijst van de recente mariene mollusken van België [List of the recent marine molluscs of Belgium]. Koninklijk Belgisch Instituut voor Natuurwetenschappen: Brussels, Belgium. 106 pp. Stocks, K. 2009. Seamounts Online: an online information system for seamount biology. Version 2009-1. World Wide Web electronic publication. van der Land, J. (ed). (2008). UNESCO-IOC Register of Marine Organisms (URMO). North-West Atlantic Ocean species (NWARMS) van der Land, J. (ed). (2008). UNESCO-IOC Register of Marine Organisms (URMO).
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Jacob van der Land [email]

Habitat

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Known from seamounts and knolls
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bibliographic citation
Check List of European Marine Mollusca (CLEMAM). Backeljau, T. (1986). Lijst van de recente mariene mollusken van België [List of the recent marine molluscs of Belgium]. Koninklijk Belgisch Instituut voor Natuurwetenschappen: Brussels, Belgium. 106 pp. Stocks, K. 2009. Seamounts Online: an online information system for seamount biology. Version 2009-1. World Wide Web electronic publication. van der Land, J. (ed). (2008). UNESCO-IOC Register of Marine Organisms (URMO). North-West Atlantic Ocean species (NWARMS) van der Land, J. (ed). (2008). UNESCO-IOC Register of Marine Organisms (URMO).
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[email]