A squid (: squid) is a
mollusc
Mollusca is a phylum of protostome, protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000 extant taxon, extant species of molluscs are recognized, making it the second-largest animal phylum ...
with an elongated soft body, large eyes, eight
arms
Arms or ARMS may refer to:
*Arm or arms, the upper limbs of the body
Arm, Arms, or ARMS may also refer to:
People
* Ida A. T. Arms (1856–1931), American missionary-educator, temperance leader
Coat of arms or weapons
*Armaments or weapons
**Fi ...
, and two
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 in the orders
Myopsida
Myopsida is one of the four orders of squid. It consists of two families: the monotypic '' Australiteuthis'' and the diverse and commercially important Loliginidae (~50 species). Some taxonomists classify this taxon as a suborder of the order T ...
,
Oegopsida
Oegopsida is one of the four orders of squid in the superorder Decapodiformes, in the class Cephalopoda. Together with the Myopsina, it was formerly considered to be a suborder of the order Teuthida, in which case it was known as Oegopsina. Thi ...
, and
Bathyteuthida
Bathyteuthida is an order of cephalopods, consisting of small, mesopelagic to bathypelagic squid that in some ways resemble myopsid squid, such as ''Loligo'' and in others the pelagic oegopsid squid. Its two families, the Bathyteuthidae and ...
(though many other molluscs within the broader
Neocoleoidea
Neocoleoidea is a large group of marine cephalopods. This cohort contains two extant groups: Decapodiformes (squid, cuttlefish, and relatives) and Octopodiformes (octopuses and the vampire squid). Species within this group exist in all major ...
are also called ''squid'' despite not strictly fitting these criteria). Like all other
cephalopods
A cephalopod is any member of the molluscan Taxonomic rank, class Cephalopoda (Greek language, Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral symm ...
, squid have a distinct head,
bilateral symmetry
Symmetry in biology refers to the symmetry observed in organisms, including plants, animals, fungi, and bacteria. External symmetry can be easily seen by just looking at an organism. For example, the face of a human being has a plane of symme ...
, and a
mantle. They are mainly soft-bodied, like
octopus
An octopus (: octopuses or octopodes) is a soft-bodied, eight-limbed mollusc of the order Octopoda (, ). The order consists of some 300 species and is grouped within the class Cephalopoda with squids, cuttlefish, and nautiloids. Like oth ...
es, but have a small internal skeleton in the form of a rod-like
gladius
''Gladius'' () is a Latin word properly referring to the type of sword that was used by Ancient Rome, ancient Roman foot soldiers starting from the 3rd century BC and until the 3rd century AD. Linguistically, within Latin, the word also came t ...
or pen, made of
chitin
Chitin (carbon, C8hydrogen, H13oxygen, O5nitrogen, N)n ( ) is a long-chain polymer of N-Acetylglucosamine, ''N''-acetylglucosamine, an amide derivative of glucose. Chitin is the second most abundant polysaccharide in nature (behind only cell ...
.
Squid diverged from other cephalopods during the
Jurassic
The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately 143.1 Mya. ...
and occupy a similar
role
A role (also rôle or social role) is a set of connected behaviors, rights, obligations, beliefs, and norms as conceptualized by people in a social situation. It is an
expected or free or continuously changing behavior and may have a given indi ...
to
teleost
Teleostei (; Ancient Greek, Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts (), is, by far, the largest group of ray-finned fishes (class Actinopterygii), with 96% of all neontology, extant species of f ...
fish as open-water
predator
Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
s of similar size and behaviour. They play an important role in the open-water food web. The two long tentacles are used to grab prey and the eight arms to hold and control it. The beak then cuts the food into suitable size chunks for swallowing. Squid are rapid swimmers, moving by
jet propulsion, and largely locate their prey by sight. They are among the most intelligent of
invertebrates
Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordate subphylum ...
, with groups of
Humboldt squid
The Humboldt squid (''Dosidicus gigas''), also known as jumbo squid or jumbo flying squid, is a large, predatory squid living in the eastern Pacific Ocean. It is the Monotypic taxon, only known species of the genus ''Dosidicus'' of the subfamily ...
having been observed
hunting cooperatively. They are preyed on by
shark
Sharks are a group of elasmobranch cartilaginous fish characterized by a ribless endoskeleton, dermal denticles, five to seven gill slits on each side, and pectoral fins that are not fused to the head. Modern sharks are classified within the ...
s, other fish, sea birds,
seals and
cetacea
Cetacea (; , ) is an infraorder of aquatic mammals belonging to the order Artiodactyla that includes whales, dolphins and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively c ...
ns, particularly
sperm whale
The sperm whale or cachalot (''Physeter macrocephalus'') is the largest of the toothed whales and the largest toothed predator. It is the only living member of the Genus (biology), genus ''Physeter'' and one of three extant species in the s ...
s.
Squid can change colour for
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
signalling
A signal is both the process and the result of transmission of data over some media accomplished by embedding some variation. Signals are important in multiple subject fields including signal processing, information theory and biology.
In ...
. Some species are
bioluminescent
Bioluminescence is the emission of light during a chemiluminescence reaction by living organisms. Bioluminescence occurs in multifarious organisms ranging from marine vertebrates and invertebrates, as well as in some fungi, microorganisms inc ...
, using their light for
counter-illumination
Counter-illumination is a method of active camouflage seen in marine animals such as firefly squid and midshipman fish, and in military prototypes, producing light to match their backgrounds in both brightness and wavelength.
Marine animals of ...
camouflage, while many species can eject a cloud of
ink to distract predators.
Squid are used for human consumption with commercial fisheries in Japan, the Mediterranean, the southwestern Atlantic, the eastern Pacific and elsewhere. They are used in cuisines around the world, often known as "
calamari". Squid have featured in literature since classical times, especially in tales of giant squid and
sea monster
Sea monsters are beings from folklore believed to dwell in the sea and are often imagined to be of immense size. Marine monsters can take many forms, including sea dragons, sea serpents, or tentacled beasts. They can be slimy and scaly and are of ...
s.
Taxonomy and phylogeny
Squid are members of the class
Cephalopod
A cephalopod is any member of the molluscan Taxonomic rank, class Cephalopoda (Greek language, Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral symm ...
a, subclass
Coleoidea
Coleoidea
or Dibranchiata is one of the two subclasses of cephalopod molluscs containing all the various taxa popularly thought of as "soft-bodied" or "shell-less" (i.e. octopus, squid and cuttlefish). Unlike its extant sister group Nauti ...
. The squid
orders
Order, ORDER or Orders may refer to:
* A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica
* Categorization, the process in which ideas and objects are recognized, differentiated, and understood
* H ...
Myopsida
Myopsida is one of the four orders of squid. It consists of two families: the monotypic '' Australiteuthis'' and the diverse and commercially important Loliginidae (~50 species). Some taxonomists classify this taxon as a suborder of the order T ...
and
Oegopsida
Oegopsida is one of the four orders of squid in the superorder Decapodiformes, in the class Cephalopoda. Together with the Myopsina, it was formerly considered to be a suborder of the order Teuthida, in which case it was known as Oegopsina. Thi ...
are in the superorder
Decapodiformes (from the
Greek
Greek may refer to:
Anything of, from, or related to Greece, a country in Southern Europe:
*Greeks, an ethnic group
*Greek language, a branch of the Indo-European language family
**Proto-Greek language, the assumed last common ancestor of all kno ...
for "ten-legged"). Two other orders of decapodiform cephalopods are also called squid, although they are
taxonomically distinct from squids and differ recognizably in their gross anatomical features. They are the
bobtail squid
Bobtail squid (order Sepiolida) are a group of cephalopods closely related to cuttlefish. Bobtail squid tend to have a rounder mantle (mollusc), mantle than cuttlefish and have no cuttlebone. They have eight suckered arms and two tentacles and ar ...
of order
Sepiolida
Bobtail squid (order Sepiolida) are a group of cephalopods closely related to cuttlefish. Bobtail squid tend to have a rounder mantle than cuttlefish and have no cuttlebone. They have eight suckered arms and two tentacles and are generally quit ...
and the
ram's horn squid of the monotypic order
Spirulida. The
vampire squid (''
Vampyroteuthis infernalis''), however, is more closely related to the octopus than to any squid.
The
cladogram
A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
, not fully resolved, is based on Sanchez et al., 2018.
Their
molecular phylogeny
Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
used
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 us ...
l and
nuclear DNA
Nuclear DNA (nDNA), or nuclear deoxyribonucleic acid, is the DNA contained within each cell nucleus of a eukaryotic organism. It encodes for the majority of the genome in eukaryotes, with mitochondrial DNA and plastid DNA coding for the rest. ...
marker sequences; they comment that a robust
phylogeny
A phylogenetic tree or phylogeny is a graphical representation which shows the evolutionary history between a set of species or Taxon, taxa during a specific time.Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, M ...
"has proven very challenging to obtain". If it is accepted that
Sepiidae cuttlefish are a kind of squid, then the squids, excluding the vampire squid, form a
clade
In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
as illustrated.
Orders are shown in boldface; all the families not included in those orders are in the
paraphyletic
Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
order "Oegopsida", except Sepiadariidae and Sepiidae that are in the polyphyletic order "Sepiida",
Evolution
Crown
A crown is a traditional form of head adornment, or hat, worn by monarchs as a symbol of their power and dignity. A crown is often, by extension, a symbol of the monarch's government or items endorsed by it. The word itself is used, parti ...
coleoids (the common ancestor of octopuses and squid) diverged in the late
Paleozoic
The Paleozoic ( , , ; or Palaeozoic) Era is the first of three Era (geology), geological eras of the Phanerozoic Eon. Beginning 538.8 million years ago (Ma), it succeeds the Neoproterozoic (the last era of the Proterozoic Eon) and ends 251.9 Ma a ...
(
Mississippian), according to fossils of ''
Syllipsimopodi'', an early relative of vampire squids and octopuses. True squid diverged during the Jurassic, but many squid families appeared in or after the
Cretaceous
The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
.
Both the coleoids and the teleost fish were involved in much adaptive radiation at this time, and the two modern groups resemble each other in size, ecology, habitat, morphology and behaviour, however some fish moved into fresh water while the coleoids remained in marine environments.
[
The ancestral coleoid was probably nautiloid-like with a strait septate shell that became immersed in the mantle and was used for buoyancy control. Four lines diverged from this, Spirulida (with one living member), the ]cuttlefish
Cuttlefish, or cuttles, are Marine (ocean), marine Mollusca, molluscs of the order (biology), suborder Sepiina. They belong to the class (biology), class Cephalopoda which also includes squid, octopuses, and nautiluses. Cuttlefish have a unique ...
es, the squids and the octopus
An octopus (: octopuses or octopodes) is a soft-bodied, eight-limbed mollusc of the order Octopoda (, ). The order consists of some 300 species and is grouped within the class Cephalopoda with squids, cuttlefish, and nautiloids. Like oth ...
es. Squid have differentiated from the ancestral mollusc
Mollusca is a phylum of protostome, protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000 extant taxon, extant species of molluscs are recognized, making it the second-largest animal phylum ...
such that the body plan has been condensed antero-posteriorly and extended dorso-ventrally. What may have been the foot
The foot (: feet) is an anatomical structure found in many vertebrates. It is the terminal portion of a limb which bears weight and allows locomotion. In many animals with feet, the foot is an organ at the terminal part of the leg made up o ...
of the ancestor is modified into a complex set of appendages around the mouth. The sense organs are highly developed and include advanced eyes similar to those of vertebrate
Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain.
The vertebrates make up the subphylum Vertebra ...
s.
The ancestral shell has been lost, with only an internal gladius
''Gladius'' () is a Latin word properly referring to the type of sword that was used by Ancient Rome, ancient Roman foot soldiers starting from the 3rd century BC and until the 3rd century AD. Linguistically, within Latin, the word also came t ...
, or pen, remaining. The pen, made of a chitin-like material, is a feather-shaped internal structure that supports the squid's mantle and serves as a site for muscle attachment. The cuttlebone
Cuttlebone, also known as cuttlefish bone, is a hard, brittle internal structure (an internal Mollusc shell, shell) found in all members of the family (biology), family Sepiidae, commonly known as cuttlefish, within the cephalopods. In othe ...
or sepion of the Sepiidae is calcareous
Calcareous () is an adjective meaning "mostly or partly composed of calcium carbonate", in other words, containing lime (mineral), lime or being chalky. The term is used in a wide variety of Science, scientific disciplines.
In zoology
''Calcare ...
and appears to have evolved afresh in the Tertiary
Tertiary (from Latin, meaning 'third' or 'of the third degree/order..') may refer to:
* Tertiary period, an obsolete geologic period spanning from 66 to 2.6 million years ago
* Tertiary (chemistry), a term describing bonding patterns in organic ch ...
.
File:Rhomboteuthis lehmani (cropped).jpg, Fossil '' Rhomboteuthis'' from the Lower Callovian (, middle Jurassic) of La Voulte-sur-Rhône, France
File:Plesioteuthis prisca 01.jpg, Fossil '' Plesioteuthis'' from the Tithonian
In the geological timescale, the Tithonian is the latest age (geology), age of the Late Jurassic Epoch and the uppermost stage (stratigraphy), stage of the Upper Jurassic Series. It spans the time between 149.2 ±0.7 annum, Ma and 143.1 ±0.6 (mi ...
(, upper Jurassic), Solnhofen
Solnhofen is a municipality in the district of Weißenburg-Gunzenhausen in the region of Middle Franconia in the ' of Bavaria in Germany. It is in the Altmühl valley.
The local area is famous in geology and palaeontology for Solnhofen lime ...
, Germany
Description
Squid are soft-bodied molluscs whose forms evolved to adopt an active predatory lifestyle. The head and foot of the squid are at one end of a long body, and this end is functionally anterior
Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pro ...
, leading the animal as it moves through the water. A set of eight arms and two distinctive tentacles surround the mouth; each appendage takes the form of a muscular hydrostat
A muscular hydrostat is a biological structure found in animals. It is used to manipulate items (including food) or to move its host about and consists mainly of muscles with no skeletal support. It performs its hydraulic movement without fluid ...
and is flexible and prehensile, usually bearing disc-like suckers.[
The suckers may lie directly on the arm or be stalked. Their rims are stiffened with chitin and may contain minute toothlike denticles. These features, as well as strong musculature, and a small ganglion beneath each sucker to allow individual control, provide a very powerful adhesion to grip prey. Hooks are present on the arms and tentacles in some species, but their function is unclear.][ The two tentacles are much longer than the arms and are retractile. Suckers are limited to the spatulate tip of the tentacle, known as the manus.][
In the mature male, the outer half of one of the left arms is ]hectocotylised
A hectocotylus (: hectocotyli) is one of the cephalopod limb, arms of male cephalopods that is specialized to store and transfer spermatophores to the female. Structurally, hectocotyli are muscular hydrostats. Depending on the species, the male ma ...
'' ''– and ends in a copulatory pad rather than suckers. This is used for depositing a spermatophore
A spermatophore, from Ancient Greek σπέρμα (''spérma''), meaning "seed", and -φόρος (''-phóros''), meaning "bearing", or sperm ampulla is a capsule or mass containing spermatozoa created by males of various animal species, especiall ...
inside the mantle cavity of a female. A ventral part of the foot has been converted into a funnel
A funnel is a tube or pipe that is wide at the top and narrow at the bottom, used for guiding liquid or powder into a small opening.
Funnels are usually made of stainless steel, aluminium, glass, or plastic. The material used in its constructi ...
through which water exits the mantle cavity.[
The main body mass is enclosed in the mantle, which has a swimming fin along each side. These fins are not the main source of locomotion in most species. The mantle wall is heavily muscled and internal. The visceral mass, which is covered by a thin, membranous ]epidermis
The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and Subcutaneous tissue, hypodermis. The epidermal layer provides a barrier to infection from environmental pathogens and regulates the ...
, forms a cone-shaped posterior region known as the "visceral hump". The mollusc shell is reduced to an internal, longitudinal chitinous "pen" in the functionally dorsal part of the animal; the pen acts to stiffen the squid and provides attachments for muscles.[
On the functionally ventral part of the body is an opening to the mantle cavity, which contains the ]gill
A gill () is a respiration organ, respiratory organ that many aquatic ecosystem, aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow r ...
s (ctenidia) and openings from the excretory, digestive and reproductive system
The reproductive system of an organism, also known as the genital system, is the biological system made up of all the anatomical organs involved in sexual reproduction. Many non-living substances such as fluids, hormones, and pheromones are al ...
s. An inhalant siphon behind the funnel draws water into the mantle cavity via a valve. The squid uses the funnel for locomotion via precise jet propulsion. In this form of locomotion, water is sucked into the mantle cavity and expelled out of the funnel in a fast, strong jet. The direction of travel is varied by the orientation of the funnel.[ Squid are strong swimmers and certain species can "fly" for short distances out of the water.
]
Camouflage
Squid make use of different kinds of camouflage, namely active camouflage
Active camouflage or adaptive camouflage is camouflage that adapts, often rapidly, to the surroundings of an object such as an animal or military vehicle. In theory, active camouflage could provide perfect concealment from visual detection.
Acti ...
for background matching (in shallow water) and counter-illumination. This helps to protect them from their predators and allows them to approach their prey.
The skin is covered in controllable chromatophore
Chromatophores are cells that produce color, of which many types are pigment-containing cells, or groups of cells, found in a wide range of animals including amphibians, fish, reptiles, crustaceans and cephalopod
A cephalopod is any member o ...
s of different colours, enabling the squid to match its coloration to its surroundings. The play of colours may in addition distract prey from the squid's approaching tentacles. The skin also contains light reflectors called iridophores and leucophores that, when activated, in millisecond
A millisecond (from '' milli-'' and second; symbol: ms) is a unit of time in the International System of Units equal to one thousandth (0.001 or 10−3 or 1/1000) of a second or 1000 microseconds.
A millisecond is to one second, as one second i ...
s create changeable skin patterns of polarized light. Such skin camouflage may serve various functions, such as communication with nearby squid, prey detection, navigation, and orientation during hunting or seeking shelter. Neural control of the iridophores enabling rapid changes in skin iridescence
Iridescence (also known as goniochromism) is the phenomenon of certain surfaces that appear gradually to change colour as the angle of view or the angle of illumination changes. Iridescence is caused by wave interference of light in microstru ...
appears to be regulated by a cholinergic
Cholinergic agents are compounds which mimic the action of acetylcholine and/or butyrylcholine. In general, the word " choline" describes the various quaternary ammonium salts containing the ''N'',''N'',''N''-trimethylethanolammonium cation ...
process affecting reflectin protein
Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residue (biochemistry), residues. Proteins perform a vast array of functions within organisms, including Enzyme catalysis, catalysing metab ...
s.
Some mesopelagic
The mesopelagic zone (Greek language, Greek μέσον, middle), also known as the middle pelagic or twilight zone, is the part of the pelagic zone that lies between the photic epipelagic and the aphotic bathypelagic zones. It is defined by light ...
squid such as the firefly squid (''Watasenia scintillans'') and the midwater squid (''Abralia veranyi'') use counter-illumination camouflage, generating light to match the downwelling light from the ocean surface. This creates the effect of countershading
Countershading, or Thayer's law, is a method of camouflage in which animal coloration, an animal's coloration is darker on the top or upper side and lighter on the underside of the body. This pattern is found in many species of mammals, reptile ...
, making the underside lighter than the upperside.
Counter-illumination is also used by the Hawaiian bobtail squid (''Euprymna scolopes''), which has symbiotic bacteria (''Aliivibrio fischeri'') that produce light to help the squid avoid nocturnal predators. This light shines through the squid's skin on its underside and is generated by a large and complex two-lobed light organ inside the squid's mantle cavity. From there, it escapes downwards, some of it travelling directly, some coming off a reflector at the top of the organ (dorsal side). Below there is a kind of iris, which has branches (diverticula) of its ink sac
An ink sac is an anatomical feature that is found in many cephalopod mollusks used to produce the defensive cephalopod ink. With the exception of nocturnal and very deep water cephalopods, all Coleoidea (squid, octopus and cuttlefish) which dwell ...
, with a lens below that; both the reflector and lens are derived from mesoderm
The mesoderm is the middle layer of the three germ layers that develops during gastrulation in the very early development of the embryo of most animals. The outer layer is the ectoderm, and the inner layer is the endoderm.Langman's Medical ...
. The squid controls light production by changing the shape of its iris or adjusting the strength of yellow filters on its underside, which presumably change the balance of wavelengths emitted. Light production shows a correlation with intensity of down-welling light, but it is about one third as bright; the squid can track repeated changes in brightness. Because the Hawaiian bobtail squid hides in sand during the day to avoid predators, it does not use counter-illumination during daylight hours.
File:Chromatophores.jpg, upright=1.25, Controllable chromatophore
Chromatophores are cells that produce color, of which many types are pigment-containing cells, or groups of cells, found in a wide range of animals including amphibians, fish, reptiles, crustaceans and cephalopod
A cephalopod is any member o ...
s of different colours in the skin of a squid allow it to change its coloration and patterns rapidly, whether for camouflage or signalling.
File:Squid Counterillumination.png, Principle of counter-illumination
Counter-illumination is a method of active camouflage seen in marine animals such as firefly squid and midshipman fish, and in military prototypes, producing light to match their backgrounds in both brightness and wavelength.
Marine animals of ...
camouflage of the firefly squid, '' Watasenia scintillans''. When seen from below by a predator
Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
, the animal's light helps to match its brightness and colour to the sea surface above.
Predator distraction with ink
Squid distract attacking predators by ejecting a cloud of ink, giving themselves an opportunity to escape. The ink gland and its associated ink sac empties into the rectum close to the anus, allowing the squid to rapidly discharge black ink into the mantle cavity and surrounding water. The ink is a suspension of melanin
Melanin (; ) is a family of biomolecules organized as oligomers or polymers, which among other functions provide the pigments of many organisms. Melanin pigments are produced in a specialized group of cells known as melanocytes.
There are ...
particles and quickly disperses to form a dark cloud that obscures the escape manoeuvres of the squid. Predatory fish may also be deterred by the alkaloid nature of the discharge which may interfere with their chemoreceptor
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 chemorece ...
s.[
]
Nervous system and sense organs
Cephalopods have the most highly developed nervous systems among invertebrate
Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordata, chordate s ...
s. Squids have a complex brain in the form of a nerve ring encircling the oesophagus
The esophagus (American English), oesophagus (British English), or œsophagus ( archaic spelling) ( see spelling difference) all ; : ((o)e)(œ)sophagi or ((o)e)(œ)sophaguses), colloquially known also as the food pipe, food tube, or gullet, ...
, enclosed in a cartilaginous cranium
The skull, or cranium, is typically a bony enclosure around the brain of a vertebrate. In some fish, and amphibians, the skull is of cartilage. The skull is at the head end of the vertebrate.
In the human, the skull comprises two prominent ...
. Paired cerebral ganglia
A ganglion (: ganglia) is a group of neuron cell bodies in the peripheral nervous system. In the somatic nervous system, this includes dorsal root ganglia and trigeminal ganglia among a few others. In the autonomic nervous system, there a ...
above the oesophagus receive sensory information from the eyes and statocyst
The statocyst is a balance sensory receptor present in some aquatic invertebrates, including bivalves, cnidarians, ctenophorans, echinoderms, cephalopods, crustaceans, and gastropods, A similar structure is also found in '' Xenoturbella''. T ...
s, and further ganglia below control the muscles of the mouth, foot, mantle and viscera. Giant axons up to in diameter convey nerve messages with great rapidity to the circular muscles of the mantle wall, allowing a synchronous, powerful contraction and maximum speed in the jet propulsion system.[
The paired eyes, on either side of the head, are housed in capsules fused to the cranium. Their structure is very similar to that of a fish eye, with a globular ]lens
A lens is a transmissive optical device that focuses or disperses a light beam by means of refraction. A simple lens consists of a single piece of transparent material, while a compound lens consists of several simple lenses (''elements'') ...
that has a depth of focus from to infinity. The image is focused by changing the position of the lens, as in a camera or telescope
A telescope is a device used to observe distant objects by their emission, Absorption (electromagnetic radiation), absorption, or Reflection (physics), reflection of electromagnetic radiation. Originally, it was an optical instrument using len ...
, rather than changing the shape of the lens, as in the human eye
The human eye is a sensory organ in the visual system that reacts to light, visible light allowing eyesight. Other functions include maintaining the circadian rhythm, and Balance (ability), keeping balance.
The eye can be considered as a living ...
. Squid adjust to changes in light intensity by expanding and contracting the slit-shaped pupil
The pupil is a hole located in the center of the iris of the eye that allows light to strike the retina.Cassin, B. and Solomon, S. (1990) ''Dictionary of Eye Terminology''. Gainesville, Florida: Triad Publishing Company. It appears black becau ...
.[ Deep sea squids in the family Histioteuthidae have eyes of two different types and orientation. The large left eye is tubular in shape and looks upwards, presumably searching for the silhouettes of animals higher in the ]water column
The (oceanic) water column is a concept used in oceanography to describe the physical (temperature, salinity, light penetration) and chemical ( pH, dissolved oxygen, nutrient salts) characteristics of seawater at different depths for a defined ...
. The normally-shaped right eye points forwards and downwards to detect prey.
The statocysts are involved in maintaining balance and are analogous to the inner ear of fish. They are housed in cartilaginous capsules on either side of the cranium. They provide the squid with information on its body position in relation to gravity, its orientation, acceleration and rotation, and are able to perceive incoming vibrations. Without the statocysts, the squid cannot maintain equilibrium.[ Squid appear to have limited hearing, but the head and arms bear lines of hair-cells that are weakly sensitive to water movements and changes in pressure, and are analogous in function to the ]lateral line
The lateral line, also called the lateral line organ (LLO), is a system of sensory organs found in fish, used to detect movement, vibration, and pressure gradients in the surrounding water. The sensory ability is achieved via modified epithelia ...
system of fish.[
]
Reproductive system
The sexes are separate in squid, with a single gonad
A gonad, sex gland, or reproductive gland is a Heterocrine gland, mixed gland and sex organ that produces the gametes and sex hormones of an organism. Female reproductive cells are egg cells, and male reproductive cells are sperm. The male gon ...
in the posterior part of the body. Fertilisation is external and usually takes place in the mantle cavity of the female. The male has a testis
A testicle or testis ( testes) is the gonad in all male bilaterians, including humans, and is Homology (biology), homologous to the ovary in females. Its primary functions are the production of sperm and the secretion of Androgen, androgens, p ...
from which sperm pass into a single gonoduct where they are rolled together into a long bundle, or spermatophore. The gonoduct is elongated into a "penis" that extends into the mantle cavity and through which spermatophores are ejected. In shallow water species, the penis is short, and the spermatophore is removed from the mantle cavity by a tentacle of the male, which is specially adapted for the purpose and known as a hectocotylus
A hectocotylus (: hectocotyli) is one of the arms of male cephalopods that is specialized to store and transfer spermatophores to the female. Structurally, hectocotyli are muscular hydrostats. Depending on the species, the male may use it merel ...
, and placed inside the mantle cavity of the female during mating.[
]
The female has a large translucent ovary
The ovary () is a gonad in the female reproductive system that produces ova; when released, an ovum travels through the fallopian tube/ oviduct into the uterus. There is an ovary on the left and the right side of the body. The ovaries are end ...
, situated towards the posterior of the visceral mass. From here, eggs travel along the gonocoel, where there are a pair of white nidamental gland
Nidamental glands are internal organs found in some elasmobranchs and certain molluscs, including cephalopods (specifically Decapodiformes and nautiluses) and gastropod
Gastropods (), commonly known as slugs and snails, belong to a large Ta ...
s, which lie anterior to the gills. Also present are red-spotted accessory nidamental glands containing symbiotic
Symbiosis (Ancient Greek : living with, companionship < : together; and ''bíōsis'': living) is any type of a close and long-term biolo ...
bacteria; both organs are associated with nutrient manufacture and forming shells for the eggs. The gonocoel enters the mantle cavity at the gonopore
A gonopore, sometimes called a gonadopore, is a genital pore in many invertebrates. Hexapods, including insects, have a single common gonopore, except mayflies, which have a pair of gonopores. More specifically, in the unmodified female, it is ...
, and in some species, receptacles for storing spermatophores are located nearby, in the mantle wall.[
In shallow-water species of the ]continental shelf
A continental shelf is a portion of a continent that is submerged under an area of relatively shallow water, known as a shelf sea. Much of these shelves were exposed by drops in sea level during glacial periods. The shelf surrounding an islan ...
and epipelagic
The photic zone (or euphotic zone, epipelagic zone, or sunlight zone) is the uppermost layer of a body of water that receives sunlight, allowing phytoplankton to perform photosynthesis. It undergoes a series of physical, chemical, and biological ...
or mesopelagic zone
The mesopelagic zone (Greek μέσον, middle), also known as the middle pelagic or twilight zone, is the part of the pelagic zone that lies between the photic epipelagic and the aphotic bathypelagic zones. It is defined by light, and begins ...
s, it is frequently one or both of arm pair IV of males that are modified into hectocotyli. However, most deep-sea squid lack hectocotyl arms and have longer penises; Ancistrocheiridae and Cranchiinae are exceptions. Giant squid of the genus ''Architeuthis'' are unusual in that they possess both a large penis and modified arm tips, although whether the latter are used for spermatophore transfer is uncertain. Penis elongation has been observed in the deep-water species ''Onykia ingens
''Moroteuthopsis ingens'', the greater hooked squid, is a species of squid in the family Onychoteuthidae. It occurs worldwide in subantarctic oceans.
Taxonomy
''M. ingens'' was long attributed to the genus ''Moroteuthis'', which became a junior ...
''; when erect, the penis may be as long as the mantle, head, and arms combined. As such, deep-water squid have the greatest known penis length relative to body size of all mobile animals, second in the entire animal kingdom only to certain sessile barnacle
Barnacles are arthropods of the subclass (taxonomy), subclass Cirripedia in the subphylum Crustacean, Crustacea. They are related to crabs and lobsters, with similar Nauplius (larva), nauplius larvae. Barnacles are exclusively marine invertebra ...
s.
Digestive system
Like all cephalopods, squids are predators and have complex digestive systems. The mouth is equipped with a sharp, horny 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 pecking, grasping, and holding (in probing for food, eating, manipulating and ...
mainly made of chitin and cross-link
In chemistry and biology, a cross-link is a bond or a short sequence of bonds that links one polymer chain to another. These links may take the form of covalent bonds or ionic bonds and the polymers can be either synthetic polymers or natural ...
ed proteins, which is used to kill and tear prey into manageable pieces. The beak is very robust, but does not contain minerals, unlike the teeth and jaws of many other organisms; the cross-linked proteins are histidine- and glycine-rich and give the beak a stiffness and hardness greater than most equivalent synthetic organic materials. The stomachs of captured whales often have indigestible squid beaks inside. The mouth contains the radula
The radula (; : radulae or radulas) is an anatomical structure used by mollusks 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 enters ...
, the rough tongue common to all molluscs
Mollusca is a phylum of protostome, protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000 extant taxon, extant species of molluscs are recognized, making it the second-largest animal phylum ...
except bivalvia
Bivalvia () or bivalves, in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class (biology), class of aquatic animal, aquatic molluscs (marine and freshwater) that have laterally compressed soft bodies enclosed b ...
, which is equipped with multiple rows of teeth.[ In some species, toxic ]saliva
Saliva (commonly referred as spit or drool) 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 ...
helps to control large prey; when subdued, the food can be torn in pieces by the beak, moved to the oesophagus by the radula, and swallowed.
The food bolus is moved along the gut by waves of muscular contractions (peristalsis
Peristalsis ( , ) is a type of intestinal motility, characterized by symmetry in biology#Radial symmetry, radially symmetrical contraction and relaxation of muscles that propagate in a wave down a tube, in an wikt:anterograde, anterograde dir ...
). The long oesophagus leads to a muscular stomach
The stomach is a muscular, hollow organ in the upper gastrointestinal tract of Human, humans and many other animals, including several invertebrates. The Ancient Greek name for the stomach is ''gaster'' which is used as ''gastric'' in medical t ...
roughly in the middle of the visceral mass. The digestive gland
The hepatopancreas, digestive gland or midgut gland is an organ of the digestive tract of arthropods and molluscs. It provides the functions which in mammals are provided separately by the liver and pancreas, including the production of digestive ...
, which is equivalent to a vertebrate liver, diverticulates here, as does the pancreas
The pancreas (plural pancreases, or pancreata) is an Organ (anatomy), organ of the Digestion, digestive system and endocrine system of vertebrates. In humans, it is located in the abdominal cavity, abdomen behind the stomach and functions as a ...
, and both of these empty into the caecum
The cecum ( caecum, ; plural ceca or caeca, ) is a pouch within the peritoneum that is considered to be the beginning of the large intestine. It is typically located on the right side of the body (the same side of the body as the appendix, ...
, a pouch-shaped sac where most of the absorption of nutrients takes place.[ Indigestible food can be passed directly from the stomach to the ]rectum
The rectum (: rectums or recta) is the final straight portion of the large intestine in humans and some other mammals, and the gut in others. Before expulsion through the anus or cloaca, the rectum stores the feces temporarily. The adult ...
where it joins the flow from the caecum and is voided through the anus
In mammals, invertebrates and most fish, the anus (: anuses or ani; from Latin, 'ring' or 'circle') is the external body orifice at the ''exit'' end of the digestive tract (bowel), i.e. the opposite end from the mouth. Its function is to facil ...
into the mantle cavity.[ Cephalopods are short-lived, and in mature squid, priority is given to reproduction; the female '' Onychoteuthis banksii'' for example, sheds its feeding tentacles on reaching maturity, and becomes flaccid and weak after spawning.
]
Cardiovascular and excretory systems
The squid mantle cavity is a seawater-filled sac containing three hearts and other organs supporting circulation, respiration, and excretion
Excretion is elimination of metabolic waste, which is an essential process in all organisms. In vertebrates, this is primarily carried out by the lungs, Kidney (vertebrates), kidneys, and skin. This is in contrast with secretion, where the substa ...
. Squid have a main systemic heart that pumps blood around the body as part of the general circulatory system
In vertebrates, the circulatory system is a system of organs that includes the heart, blood vessels, and blood which is circulated throughout the body. It includes the cardiovascular system, or vascular system, that consists of the heart ...
, and two branchial hearts. The systemic heart consists of three chambers, a lower ventricle and two upper atria, all of which can contract to propel the blood. The branchial hearts pump blood specifically to the gills for oxygenation, before returning it to the systemic heart.[ The blood contains the ]copper
Copper is a chemical element; it has symbol Cu (from Latin ) 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 pinkish-orang ...
-rich protein hemocyanin
Hemocyanins (also spelled haemocyanins and abbreviated Hc) are proteins that transport oxygen throughout the bodies of some invertebrate animals. These metalloproteins contain two copper atoms that reversibly bind a single oxygen molecule (O2 ...
, which is used for oxygen transport at low ocean temperatures and low oxygen concentrations, and makes the oxygenated blood a deep, blue color.[ As systemic blood returns via two ]venae cavae
In anatomy, the ''venae cavae'' (; ''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 the ...
to the branchial hearts, excretion of urine
Urine is a liquid by-product of metabolism in humans and many other animals. In placental mammals, urine flows from the Kidney (vertebrates), kidneys through the ureters to the urinary bladder and exits the urethra through the penile meatus (mal ...
, carbon dioxide
Carbon dioxide is a chemical compound with the chemical formula . It is made up of molecules that each have one carbon atom covalent bond, covalently double bonded to two oxygen atoms. It is found in a gas state at room temperature and at norma ...
, and waste solute
In chemistry, a solution is defined by IUPAC as "A liquid or solid phase containing more than one substance, when for convenience one (or more) substance, which is called the solvent, is treated differently from the other substances, which are ...
s occurs through outpockets (called nephridial appendages) in the venae cavae walls that enable gas exchange and excretion via the mantle cavity seawater.[
]
Buoyancy
Unlike nautiloids and cuttlefish which have gas-filled chambers inside their shells which provide buoyancy, and octopuses which live near and rest on the seabed and do not require to be buoyant, many squid have a fluid-filled receptacle, equivalent to the swim bladder
The swim bladder, gas bladder, fish maw, or air bladder is an internal gas-filled organ (anatomy), organ in bony fish that functions to modulate buoyancy, and thus allowing the fish to stay at desired water depth without having to maintain lift ...
of a fish, in the coelom
The coelom (or celom) is the main body cavity in many 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, i ...
or connective tissue
Connective tissue is one of the four primary types of animal tissue, a group of cells that are similar in structure, along with epithelial tissue, muscle tissue, and nervous tissue. It develops mostly from the mesenchyme, derived from the mesod ...
. This reservoir acts as a chemical buoyancy chamber, with the heavy metallic cations typical of seawater replaced by low molecular-weight ammonium
Ammonium is a modified form of ammonia that has an extra hydrogen atom. It is a positively charged (cationic) polyatomic ion, molecular ion with the chemical formula or . It is formed by the protonation, addition of a proton (a hydrogen nucleu ...
ions, a product of excretion. The small difference in density provides a small contribution to buoyancy per unit volume, so the mechanism requires a large buoyancy chamber to be effective. Since the chamber is filled with liquid, it has the advantage over a swim bladder of not changing significantly in volume with pressure. Glass squids in the family Cranchiidae for example, have an enormous transparent coelom containing ammonium ions and occupying about two-thirds the volume of the animal, allowing it to float at the required depth. About half of the 28 families of squid use this mechanism to solve their buoyancy issues.[ The family Bathyteuthidae get their buoyancy from an oily substance found in their liver and around their mantle and head.
]
Largest and smallest
The majority of squid are no more than long, although the giant squid may reach . The smallest species are probably the benthic pygmy squids '' Idiosepius'', which grow to a mantle length of , and have short bodies and stubby arms.
In 1978, sharp, curved claws on the suction cups of squid tentacles cut up the rubber coating on the hull of the USS ''Stein''. The size suggested the largest squid known at the time.
In 2003, a large specimen of an abundant but poorly understood species, ''Mesonychoteuthis hamiltoni'' (the colossal squid), was discovered. This species may grow to in length, making it the largest invertebrate. In February 2007, a New Zealand fishing vessel caught the largest squid ever documented, weighing and measuring around off the coast of Antarctica. Dissection showed that the eyes, used to detect prey in the deep Southern Ocean, exceeded the size of footballs; these may be among the largest eyes ever to exist in the animal kingdom.
Development
The eggs of squid are large for a mollusc, containing a large amount of yolk to nourish the embryo as it develops directly, without an intervening veliger
A veliger is the planktonic larva of many kinds of sea snails and freshwater snails, as well as most bivalve molluscs (clams) and tusk shells.
Description
The veliger is the characteristic larva of the gastropod, bivalve and scaphopod taxono ...
larval stage. The embryo grows as a disc of cells on top of the yolk. During the gastrulation
Gastrulation is the stage in the early embryonic development of most animals, during which the blastula (a single-layered hollow sphere of cells), or in mammals, the blastocyst, is reorganized into a two-layered or three-layered embryo known as ...
stage, the margins of the disc grow to surround the yolk, forming a yolk sac, which eventually forms part of the animal's gut. The dorsal side of the disc grows upwards 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 upwards, coming to form a ring around the funnel and mouth. The yolk is gradually absorbed as the embryo grows. Some juvenile squid live higher in the water column than do adults. Squids tend to be short-lived; ''Loligo
''Loligo'' is a genus of squid and one of the most representative and widely distributed groups of myopsid squid.
The genus was first described by Jean Baptiste Lamarck in 1798. However, the name had been used earlier than Lamarck ( Schneider, ...
'' for example lives from one to three years according to species, typically dying soon after spawning.[
]
In a well-studied bioluminescent species, the Hawaiian bobtail squid, a special light organ in the squid's mantle is rapidly colonized with '' Aliivibrio fischeri'' bacteria within hours of hatching. This light-organ colonization requires this particular bacterial species for a symbiotic relationship; no colonization occurs in the absence of ''A. fischeri''. Colonization occurs in a horizontal manner, such that the hosts acquires its bacterial partners from the environment. The symbiosis is obligate
{{wiktionary, obligate
As an adjective, obligate means "by necessity" (antonym '' facultative'') and is used mainly in biology in phrases such as:
* Obligate aerobe, an organism that cannot survive without oxygen
* Obligate anaerobe, an organism ...
for the squid, but facultative for the bacteria. Once the bacteria enter the squid, they colonize interior epithelial cells
Epithelium or epithelial tissue is a thin, continuous, protective layer of cells with little extracellular matrix. An example is the epidermis, the outermost layer of the skin. Epithelial ( mesothelial) tissues line the outer surfaces of man ...
in the light organ, living in crypts with complex microvilli
Microvilli (: microvillus) are microscopic cellular membrane protrusions that increase the surface area for diffusion and minimize any increase in volume, and are involved in a wide variety of functions, including absorption, secretion, cellula ...
protrusions. The bacteria also interact with hemocyte
A blood cell (also called a hematopoietic cell, hemocyte, or hematocyte) is a cell produced through hematopoiesis and found mainly in the blood. Major types of blood cells include red blood cells (erythrocytes), white blood cells (leukocytes), ...
s, macrophage-like blood cells that migrate between epithelial cells, but the mechanism and function of this process is not well understood. Bioluminescence reaches its highest levels during the early evening hours and bottoms out before dawn; this occurs because at the end of each day, the contents of the squid's crypts are expelled into the surrounding environment. Approximately 95% of the bacteria are voided each morning before the bacterial population builds up again by nightfall.
Behaviour
Locomotion
Squid can move about in several different ways. Slow movement is achieved by a gentle undulation of the muscular lateral fins on either side of the trunk which drives the animal forward. A more common means of locomotion providing sustained movement is achieved using jetting, during which contraction of the muscular wall of the mantle cavity provides jet propulsion.[
Slow jetting is used for ordinary locomotion, and ventilation of the gills is achieved at the same time. The circular muscles in the mantle wall contract; this causes the inhalant valve to close, the exhalant valve to open and the mantle edge to lock tightly around the head. Water is forced out through the funnel which is pointed in the opposite direction to the required direction of travel. The inhalant phase is initiated by the relaxation of the circular muscles causes them to stretch, the connective tissue in the mantle wall recoils elastically, the mantle cavity expands causing the inhalant valve to open, the exhalant valve to close and water to flow into the cavity. This cycle of exhalation and inhalation is repeated to provide continuous locomotion.][
Fast jetting is an escape response. In this form of locomotion, radial muscles in the mantle wall are involved as well as circular ones, making it possible to hyper-inflate the mantle cavity with a larger volume of water than during slow jetting. On contraction, water flows out with great force, the funnel always being pointed anteriorly, and travel is backwards. During this means of locomotion, some squid exit the water in a similar way to ]flying fish
The Exocoetidae are a family (biology), family of Saltwater fish, marine Actinopterygii, ray-finned fish in the order (biology), order Beloniformes, known colloquially as flying fish or flying cod. About 64 species are grouped in seven genus, ge ...
, gliding through the air for up to , and occasionally ending up on the decks of ships.[
]
Feeding
Squid are carnivore
A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant
Plants are the eukaryotes that form the Kingdom (biology), kingdom Plantae; they ar ...
s, and, with their strong arms and suckers, can overwhelm relatively large animals efficiently. Prey is identified by sight or by touch, grabbed by the tentacles which can be shot out with great rapidity, brought back to within reach of the arms, and held by the hooks and suckers on their surface.[ In some species, the squid's saliva contains ]toxin
A toxin is a naturally occurring poison produced by metabolic activities of living cells or organisms. They occur especially as proteins, often conjugated. The term was first used by organic chemist Ludwig Brieger (1849–1919), derived ...
s which act to subdue the prey. These are injected into its bloodstream when the prey is bitten, along with vasodilators and chemicals to stimulate the heart, and quickly circulate to all parts of its body.[ The deep sea squid '' Taningia danae'' has been filmed releasing blinding flashes of light from large photophores on its arms to illuminate and disorientate potential prey.
]
Although squid can catch large prey, the mouth is relatively small, and the food must be cut into pieces by the chitinous beak with its powerful muscles before being swallowed. The radula is located in the buccal cavity and has multiple rows of tiny teeth that draw the food backwards and grind it in pieces.[ The deep sea squid '']Mastigoteuthis
''Mastigoteuthis'' is a genus of whip-lash squid containing at least seven valid species. Some teuthologists consider ''Idioteuthis'' synonymous with this taxon.
The genus contains bioluminescent species.
Species
*Genus ''Mastigoteuthis''
**''M ...
'' has the whole length of its whip-like tentacles covered with tiny suckers; it probably catches small organisms in the same way that flypaper traps flies. The tentacles of some bathypelagic
The bathypelagic zone or bathyal zone (from Greek βαθύς (bathýs), deep) is the part of the open ocean that extends from a depth of below the ocean surface. It lies between the mesopelagic above and the abyssopelagic below. The bathypela ...
squids bear photophore
A photophore is a specialized anatomical structure found in a variety of organisms that emits light through the process of boluminescence. This light may be produced endogenously by the organism itself (symbiotic) or generated through a mut ...
s which may bring food within its reach by attracting prey.
Squid are among the most intelligent invertebrates. For example, groups of Humboldt squid
The Humboldt squid (''Dosidicus gigas''), also known as jumbo squid or jumbo flying squid, is a large, predatory squid living in the eastern Pacific Ocean. It is the Monotypic taxon, only known species of the genus ''Dosidicus'' of the subfamily ...
hunt cooperatively, spiralling up through the water at night and coordinating their vertical and horizontal movements while foraging.
Reproduction
Courtship in squid takes place in the open water and involves the male selecting a female, the female responding, and the transfer by the male of spermatophores to the female. In many instances, the male may display to identify himself to the female and drive off any potential competitors. Elaborate changes in body patterning take place in some species in both agonistic and courtship behaviour. The Caribbean reef squid
The Caribbean reef squid (''Sepioteuthis sepioidea''), commonly called the reef squid, is a species of small, torpedo-shaped squid with undulating fins that extend nearly the entire length of the body, approximately 20 cm (8 in) in length. They ...
(''Sepioteuthis sepioidea''), for example, employs a complex array of colour changes during courtship and social interactions and has a range of about 16 body patterns in its repertoire.
The pair adopt a head-to-head position, and "jaw locking" may take place, in a similar manner to that adopted by some cichlid
Cichlids ()
are a large, diverse, and widespread family of percomorph fish in the family Cichlidae, order Cichliformes. At least 1,760 species have been scientifically described, making it one of the largest vertebrate families, with on ...
fish. The hectocotylus of the male is used to transfer the spermatophore and deposit it in the female's mantle cavity in the position appropriate for the species; this may be adjacent to the gonopore or in a seminal receptacle.[
The sperm may be used immediately or may be stored. As the eggs pass down the oviduct, they are wrapped in a gelatinous coating, before continuing to the mantle cavity, where they are fertilised. In '']Loligo
''Loligo'' is a genus of squid and one of the most representative and widely distributed groups of myopsid squid.
The genus was first described by Jean Baptiste Lamarck in 1798. However, the name had been used earlier than Lamarck ( Schneider, ...
'', further coatings are added by the nidimental glands in the walls of the cavity and the eggs leave through a funnel formed by the arms. The female attaches them to the substrate in strings or groups, the coating layers swelling and hardening after contact with sea water. ''Loligo'' sometimes forms breeding aggregations which may create a "community pile" of egg strings. Some pelagic and deep sea squid do not attach their egg masses, which float freely.[
]
Ecology
Squid mostly have an annual life cycle, growing fast and dying soon after spawning. The diet changes as they grow but mostly consists of large zooplankton
Zooplankton are the heterotrophic component of the planktonic community (the " zoo-" prefix comes from ), having to consume other organisms to thrive. Plankton are aquatic organisms that are unable to swim effectively against currents. Consequent ...
and small nekton
Nekton or necton (from the ) is any aquatic organism that can actively and persistently propel itself through a water column (i.e. swimming) without touching the bottom. Nektons generally have powerful tails and appendages (e.g. fins, pleopods, ...
. In Antarctica for example, krill
Krill ''(Euphausiids)'' (: krill) are small and exclusively marine crustaceans of the order (biology), order Euphausiacea, found in all of the world's oceans. The name "krill" comes from the Norwegian language, Norwegian word ', meaning "small ...
is the main constituent of the diet, with other food items being amphipods
Amphipoda () is an order (biology), order of malacostracan crustaceans with no carapace and generally with laterally compressed bodies. Amphipods () range in size from and are mostly detritivores or scavengers. There are more than 10,700 amphip ...
, other small crustacean
Crustaceans (from Latin meaning: "those with shells" or "crusted ones") are invertebrate animals that constitute one group of arthropods that are traditionally a part of the subphylum Crustacea (), a large, diverse group of mainly aquatic arthrop ...
s, and large arrow worms. Fish are also eaten, and some squid are cannibalistic.
Different species of squid vary wildly in size, and even giant squid young are rice-grain-sized at hatching. Throughout their life cycle, they can serve as a food source for many sizes of predator. Juvenile squid provide part of the diet for worms
The World Register of Marine Species (WoRMS) is a taxonomic database that aims to provide an authoritative and comprehensive catalogue and list of names of marine organisms.
Content
The content of the registry is edited and maintained by scien ...
and small fish. Larger squid are food for larger predators, including sharks, sea birds, seals and whales.
When researchers studied the contents of the stomachs of elephant seals in South Georgia, they found 96% squid by weight.[ In a single day, a sperm whale can eat 700 to 800 squid,] and a Risso's dolphin
Risso's dolphin (''Grampus griseus'') is a marine mammal and dolphin, the only species of the genus ''Grampus''. Some of the most closely related species to these dolphins include: pilot whales (''Globicephala'' spp.), pygmy killer whales (''Fere ...
entangled in a net in the Mediterranean was found to have eaten angel clubhook squid, umbrella squid, reverse jewel squid and European flying squid
The European flying squid (''Todarodes sagittatus'') is a species of squid from the continental slope and oceanic waters of the eastern Atlantic Ocean and the Mediterranean Sea. It is the type species of the genus ''Todarodes'', the type genus o ...
, all identifiable from their indigestible beaks.
'' Ornithoteuthis volatilis'', a common squid from the tropical Indo-Pacific, is predated by yellowfin tuna
The yellowfin tuna (''Thunnus albacares'') is a species of tuna found in pelagic waters of tropical and subtropical oceans worldwide.
Yellowfin is often marketed as ahi, from the Hawaiian , a name also used there for the closely related bigeye ...
, longnose lancetfish, common dolphinfish and swordfish
The swordfish (''Xiphias gladius''), also known as the broadbill in some countries, are large, highly migratory predatory fish characterized by a long, flat, pointed bill. They are the sole member of the Family (biology), family Xiphiidae. They ...
, the tiger shark
The tiger shark (''Galeocerdo cuvier'') is a species of ground shark, and the only extant member of the genus '' Galeocerdo'' and family Galeocerdonidae. It is a large apex predator, with females capable of attaining a length of over . Popula ...
, the scalloped hammerhead shark and the smooth hammerhead shark. Sperm whales also hunt this species extensively as does the brown fur seal
The brown fur seal (''Arctocephalus pusillus''), also known as the Cape fur seal, and Afro-Australian fur seal, is a species of fur seal.
Description
The brown fur seal is the largest and most robust member of the fur seals. It has a large an ...
. In the Southern Ocean
The Southern Ocean, also known as the Antarctic Ocean, comprises the southernmost waters of the world ocean, generally taken to be south of 60th parallel south, 60° S latitude and encircling Antarctica. With a size of , it is the seco ...
, penguin
Penguins are a group of aquatic flightless birds from the family Spheniscidae () of the order Sphenisciformes (). They live almost exclusively in the Southern Hemisphere. Only one species, the Galápagos penguin, is equatorial, with a sm ...
s and wandering albatross
The snowy albatross (''Diomedea exulans''), also known as the wandering albatross, white-winged albatross, or goonie, is a large seabird from the family Diomedeidae
Albatrosses, of the biological family (biology), family Diomedeidae, are la ...
es are major predators of '' Gonatus antarcticus''.
Human uses
In literature and art
Giant squid have featured as monsters of the deep since classical times. Giant squid were described by Aristotle (4th century BC) in his ''History of Animals
''History of Animals'' (, ''Ton peri ta zoia historion'', "Inquiries on Animals"; , "History of Animals") is one of the major texts on biology by the ancient Greek philosopher Aristotle. It was written in sometime between the mid-fourth centur ...
'' and Pliny the Elder
Gaius Plinius Secundus (AD 23/24 79), known in English as Pliny the Elder ( ), was a Roman Empire, Roman author, Natural history, naturalist, and naval and army commander of the early Roman Empire, and a friend of the Roman emperor, emperor Vesp ...
(1st century AD) in his ''Natural History
Natural history is a domain of inquiry involving organisms, including animals, fungi, and plants, in their natural environment, leaning more towards observational than experimental methods of study. A person who studies natural history is cal ...
''. The Gorgon
The Gorgons ( ; ), in Greek mythology, are three monstrous sisters, Stheno, Euryale, and Medusa, said to be the daughters of Phorcys and Ceto. They lived near their sisters the Graeae, and were able to turn anyone who looked at them to sto ...
of Greek mythology
Greek mythology is the body of myths originally told by the Ancient Greece, ancient Greeks, and a genre of ancient Greek folklore, today absorbed alongside Roman mythology into the broader designation of classical mythology. These stories conc ...
may have been inspired by squid or octopus, the animal itself representing the severed head of Medusa
In Greek mythology, Medusa (; ), also called Gorgo () or the Gorgon, was one of the three Gorgons. Medusa is generally described as a woman with living snakes in place of hair; her appearance was so hideous that anyone who looked upon her wa ...
, the beak as the protruding tongue and fangs, and its tentacles as the snakes. The six-headed sea monster of the ''Odyssey
The ''Odyssey'' (; ) is one of two major epics of ancient Greek literature attributed to Homer. It is one of the oldest surviving works of literature and remains popular with modern audiences. Like the ''Iliad'', the ''Odyssey'' is divi ...
'', Scylla
In Greek mythology, Scylla ( ; , ) is a legendary, man-eating monster that lives on one side of a narrow channel of water, opposite her counterpart, the sea-swallowing monster Charybdis. The two sides of the strait are within an arrow's range o ...
, may have had a similar origin. The Nordic legend of the kraken may also have derived from sightings of large cephalopods.
In literature, H. G. Wells
Herbert George Wells (21 September 1866 – 13 August 1946) was an English writer, prolific in many genres. He wrote more than fifty novels and dozens of short stories. His non-fiction output included works of social commentary, politics, hist ...
' short story " The Sea Raiders" featured a man-eating squid species ''Haploteuthis ferox''. The science fiction
Science fiction (often shortened to sci-fi or abbreviated SF) is a genre of speculative fiction that deals with imaginative and futuristic concepts. These concepts may include information technology and robotics, biological manipulations, space ...
writer Jules Verne
Jules Gabriel Verne (;''Longman Pronunciation Dictionary''. ; 8 February 1828 – 24 March 1905) was a French novelist, poet and playwright.
His collaboration with the publisher Pierre-Jules Hetzel led to the creation of the ''Voyages extraor ...
told a tale of a kraken-like monster in his 1870 novel ''Twenty Thousand Leagues Under the Seas
''Twenty Thousand Leagues Under the Seas'' () is a science fiction
Science fiction (often shortened to sci-fi or abbreviated SF) is a genre of speculative fiction that deals with imaginative and futuristic concepts. These concepts may inclu ...
''.[
]
As food
Squid form a major food resource and are used in cuisines around the world, notably in Japan where it is eaten as ika sōmen IKA may refer to
* IATA code and common English abbreviation of Tehran Imam Khomeini International Airport
* Industrias Kaiser Argentina, former Argentine motor company
* Informativna katolička agencija, a Croatian Catholic news agency
* ''Interna ...
, sliced into vermicelli-like strips; as sashimi
is a Japanese cuisine, Japanese delicacy consisting of fresh raw fish or Raw meat, meat sliced into thin pieces and often eaten with soy sauce.
Origin
The word ''sashimi'' means 'pierced body', i.e., "wikt:刺身, 刺身" = ''sashimi'', whe ...
; and as tempura
is a typical Japanese dish that usually consists of seafood and vegetables that have been coated in a thin batter and deep-fried. Tempura originated in the 16th century, when Portuguese Jesuits brought the Western-style cooking method of ...
. Three species of ''Loligo'' are used in large quantities: '' L. vulgaris'' in the Mediterranean (known as in Spanish, in Italian); '' L. forbesii'' in the Northeast Atlantic; and '' L. pealei'' on the American East Coast. Among the Ommastrephidae, '' Todarodes pacificus'' is the main commercial species, harvested in large quantities across the North Pacific in Canada, Japan and China.
In English-speaking countries, squid as food is often called '' calamari'', adopted from Italian into English in the 17th century. Squid are found abundantly in certain areas, and provide large catches for 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 grounds). Commercial fisheries include wild fisheries and fish farm ...
. The body can be stuffed whole, cut into flat pieces, or sliced into rings. The arms, tentacles, and ink are also edible; the only parts not eaten are the beak and gladius (pen). Squid is a good food source for zinc
Zinc is a chemical element; it has symbol Zn and atomic number 30. It is a slightly brittle metal at room temperature and has a shiny-greyish appearance when oxidation is removed. It is the first element in group 12 (IIB) of the periodic tabl ...
and manganese
Manganese is a chemical element; it has Symbol (chemistry), symbol Mn and atomic number 25. It is a hard, brittle, silvery metal, often found in minerals in combination with iron. Manganese was first isolated in the 1770s. It is a transition m ...
, and high in copper, selenium
Selenium is a chemical element; it has symbol (chemistry), symbol Se and atomic number 34. It has various physical appearances, including a brick-red powder, a vitreous black solid, and a grey metallic-looking form. It seldom occurs in this elem ...
, vitamin B12, and riboflavin
Riboflavin, also known as vitamin B2, is a vitamin found in food and sold as a dietary supplement. It is essential to the formation of two major coenzymes, flavin mononucleotide and flavin adenine dinucleotide. These coenzymes are involved in ...
.
Commercial fishing
According to the FAO
The Food and Agriculture Organization of the United Nations; . (FAO) is a List of specialized agencies of the United Nations, specialized agency of the United Nations that leads international efforts to defeat hunger and improve nutrition ...
, the cephalopod catch for 2002 was . Of this, 2,189,206 tonnes, or 75.8 percent, was squid. The following table lists squid species fishery catches that exceeded in 2002.
In biomimicry
Prototype chromatophores that mimic
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. In the simples ...
the squid's adaptive camouflage have been made by Bristol University
The University of Bristol is a public research university in Bristol, England. It received its royal charter in 1909, although it can trace its roots to a Merchant Venturers' school founded in 1595 and University College, Bristol, which had ...
researchers using an electroactive dielectric
In electromagnetism, a dielectric (or dielectric medium) is an Insulator (electricity), electrical insulator that can be Polarisability, polarised by an applied electric field. When a dielectric material is placed in an electric field, electric ...
elastomer
An elastomer is a polymer with viscoelasticity (i.e. both viscosity and elasticity) and with weak intermolecular forces, generally low Young's modulus (E) and high failure strain compared with other materials. The term, a portmanteau of ''ela ...
, a flexible "smart" material that changes its colour and texture in response to electrical signals. The researchers state that their goal is to create an artificial skin that provides rapid active camouflage.
The squid giant axon inspired Otto Schmitt
Otto Herbert Schmitt (April 6, 1913 – January 6, 1998) was an American inventor, engineer, and biophysicist known for his scientific contributions to biophysics and for establishing the field of biomedical engineering. Schmitt also coined ...
to develop a comparator circuit with hysteresis
Hysteresis is the dependence of the state of a system on its history. For example, a magnet may have more than one possible magnetic moment in a given magnetic field, depending on how the field changed in the past. Plots of a single component of ...
now called the Schmitt trigger
In electronics, a Schmitt trigger is a comparator circuit with hysteresis implemented by applying positive feedback to the noninverting input of a comparator or differential amplifier. It is an passivity (engineering), active circuit which con ...
, replicating the axon's propagation of nerve impulse
An action potential (also known as a nerve impulse or "spike" when in a neuron) is a series of quick changes in voltage across a cell membrane. An action potential occurs when the membrane potential of a specific Cell (biology), cell rapidly ri ...
s.
See also
* Cephalopod
A cephalopod is any member of the molluscan Taxonomic rank, class Cephalopoda (Greek language, Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral symm ...
* Paralarva
Notes
References
Sources
*
External links
CephBase: Teuthida
Colossal Squid at the Museum of New Zealand Te Papa Tongarewa
Market squid mating, laying eggs (video)
Scientific American – Giant Squid
The Cephalopod Page
The Octopus News Magazine Online
{{Authority control
Commercial molluscs
Cenozoic cephalopods
Cretaceous cephalopods
Extant Devonian first appearances
Mollusc common names