Cephalopod

A cephalopod (/ˈsɛfələpɒd, ˈkɛf-/) is any

member of the molluscan class
Cephalopoda (Greek plural κεφαλόποδα,
kephalópoda; "head-feet") such as a
squid, octopus, or nautilus. These
exclusively marine animals are
characterized by bilateral body
symmetry, a prominent head, and a set of
arms or tentacles (muscular hydrostats)
modified from the primitive molluscan
foot. Fishermen sometimes call them
inkfish, referring to their common ability
to squirt ink. The study of cephalopods is
a branch of malacology known as
teuthology.
 Cephalopod
Temporal range: Late Cambrian – Present[1]
PreЄ Є O S D C P T J K PN
g

Bigfin reef squid

(Sepioteuthis lessoniana)

Scientific classification

Kingdom: Animalia

Phylum: Mollusca

Class: Cephalopoda
Cuvier, 1797

Subclasses
 Orthoceratoidea †
Nautiloidea
Ammonoidea †
Coleoidea

Cephalopods became dominant during

the Ordovician period, represented by
primitive nautiloids. The class now
contains two, only distantly related,
extant subclasses: Coleoidea, which
includes octopuses, squid, and cuttlefish;
and Nautiloidea, represented by Nautilus
and Allonautilus. In the Coleoidea, the
molluscan shell has been internalized or
is absent, whereas in the Nautiloidea, the
external shell remains. About 800 living
species of cephalopods have been
identified. Two important extinct taxa are
the Ammonoidea (ammonites) and
Belemnoidea (belemnites).

Distribution

Left: A pair of Sepia officinalis in shallow water

Right: Benthoctopus sp. on the Davidson Seamount
at 2,422 m depth

There are over 800 extant species of

cephalopod,[2] although new species
continue to be described. An estimated
11,000 extinct taxa have been described,
although the soft-bodied nature of
cephalopods means they are not easily
fossilised.[3]

Cephalopods are found in all the oceans

of Earth. None of them can tolerate
freshwater, but the brief squid,
Lolliguncula brevis, found in Chesapeake
Bay, is a notable partial exception in that
it tolerates brackish water.[4]
Cephalopods are thought to be unable to
live in freshwater due to multiple
biochemical constraints, and in their
+400 million year existence have never
ventured into fully freshwater habitats.[5]
Cephalopods occupy most of the depth
of the ocean, from the abyssal plain to
the sea surface. Their diversity is
greatest near the equator (~40 species
retrieved in nets at 11°N by a diversity
study) and decreases towards the poles
(~5 species captured at 60°N).[6]

Biology
Nervous system and behavior

L ft A t i t i ith
Left: An octopus opening a container with a screw
cap
Right: Hawaiian bobtail squid, Euprymna scolopes,
burying itself in the sand, leaving only the eyes
exposed

Cephalopods are widely regarded as the

most intelligent of the invertebrates, and
have well developed senses and large
brains (larger than those of
gastropods).[7] The nervous system of
cephalopods is the most complex of the
invertebrates[8] and their brain-to-body-
mass ratio falls between that of
endothermic and ectothermic
vertebrates.[6]:14 Captive cephalopods
have also been known to climb out of
their aquaria, maneuver a distance of the
lab floor, enter another aquarium to feed
on the crabs, and return to their own
aquarium.[9]

The brain is protected in a cartilaginous

cranium. The giant nerve fibers of the
cephalopod mantle have been widely
used for many years as experimental
material in neurophysiology; their large
diameter (due to lack of myelination)
makes them relatively easy to study
compared with other animals.[10]

Many cephalopods are social creatures;

when isolated from their own kind, some
species have been observed shoaling
with fish.[11]
Some cephalopods are able to fly
through the air for distances of up to
50 m. While cephalopods are not
particularly aerodynamic, they achieve
these impressive ranges by jet-
propulsion; water continues to be
expelled from the funnel while the
organism is in the air.[12] The animals
spread their fins and tentacles to form
wings and actively control lift force with
body posture.[13] One species, Todarodes
pacificus, has been observed spreading
tentacles in a flat fan shape with a
mucus film between the individual
tentacles[13][14] while another,
Sepioteuthis sepioidea, has been
observed putting the tentacles in a
circular arrangement.[15]

Senses

Cephalopods have advanced vision, can

detect gravity with statocysts, and have a
variety of chemical sense organs.[6]:34
Octopuses use their arms to explore their
environment and can use them for depth
perception.[6]

Vision
The primitive nautilus eye functions similarly to a
pinhole camera.

Play media
The W-shaped pupil of the cuttlefish expanding when
the lights are turned off

Most cephalopods rely on vision to

detect predators and prey, and to
communicate with one another.[16]
Consequently, cephalopod vision is
acute: training experiments have shown
that the common octopus can
distinguish the brightness, size, shape,
and horizontal or vertical orientation of
objects. The morphological construction
gives cephalopod eyes the same
performance as sharks'; however, their
construction differs, as cephalopods lack
a cornea, and have an everted retina.[16]
Cephalopods' eyes are also sensitive to
the plane of polarization of light.[17]
Unlike many other cephalopods,
nautiluses do not have good vision; their
eye structure is highly developed, but
lacks a solid lens. They have a simple
"pinhole" eye through which water can
pass. Instead of vision, the animal is
thought to use olfaction as the primary
sense for foraging, as well as locating or
identifying potential mates.
A cuttlefish with W-shaped pupils which may help
them discriminate colors.

Surprisingly, given their ability to change

color, all octopodes[18] and most
cephalopods[19] are considered to be
color blind. Coleoid cephalopods
(octopus, squid, cuttlefish) have a single
photoreceptor type and lack the ability to
determine color by comparing detected
photon intensity across multiple spectral
channels. When camouflaging
themselves, they use their
chromatophores to change brightness
and pattern according to the background
they see, but their ability to match the
specific color of a background may come
from cells such as iridophores and
leucophores that reflect light from the
environment.[20] They also produce visual
pigments throughout their body, and may
sense light levels directly from their
body.[21] Evidence of color vision has
been found in the sparkling enope squid
(Watasenia scintillans),[19][22] which
achieves color vision by the use of three
distinct retinal molecules (A1, sensitive
to red; A2, to purple, and A4, to yellow?)
which bind to its opsin.[23]
In 2015, a novel mechanism for spectral
discrimination in cephalopods was
described. This relies on the exploitation
of chromatic aberration (wavelength-
dependence of focal length). Numerical
modeling shows that chromatic
aberration can yield useful chromatic
information through the dependence of
image acuity on accommodation. The
unusual off-axis slit and annular pupil
shapes in cephalopods enhance this
ability.[24]

Photoreception

In 2015, molecular evidence was

published indicating that cephalopod
chromatophores are photosensitive;
reverse transcription polymerase chain
reactions (RT-PCR) revealed transcripts
encoding rhodopsin and retinochrome
within the retinas and skin of the longfin
inshore squid (Doryteuthis pealeii), and
the common cuttlefish (Sepia officinalis)
and broadclub cuttlefish (Sepia
latimanus). The authors claim this is the
first evidence that cephalopod dermal
tissues may possess the required
combination of molecules to respond to
light.[25]

Hearing

Some squids have been shown to detect

sound using their statocysts.[26]
Use of light

This broadclub cuttlefish (Sepia latimanus) can

change from camouflage tans and browns (top) to
yellow with dark highlights (bottom) in less than a
second.

Most cephalopods possess an

assemblage of skin components that
interact with light. These may include
iridophores, leucophores,
chromatophores and (in some species)
photophores. Chromatophores are
colored pigment cells that expand and
contract in accordance to produce color
and pattern which they can use in a
startling array of fashions.[6][25] As well
as providing camouflage with their
background, some cephalopods
bioluminesce, shining light downwards to
disguise their shadows from any
predators that may lurk below.[6] The
bioluminescence is produced by
bacterial symbionts; the host cephalopod
is able to detect the light produced by
these organisms.[27] Bioluminescence
may also be used to entice prey, and
some species use colorful displays to
impress mates, startle predators, or even
communicate with one another.[6]

Coloration

Cephalopods can change their colors

and patterns in milliseconds, whether for
signalling (both within the species and
for warning) or active camouflage,[6] as
their chromatophores are expanded or
contracted.[28] Although color changes
appear to rely primarily on vision input,
there is evidence that skin cells,
specifically chromatophores, can detect
light and adjust to light conditions
independently of the eyes.[29] Coloration
is typically stronger in near-shore species
than those living in the open ocean,
whose functions tend to be restricted to
disruptive camouflage.[6]:2 Most
octopuses mimic select structures in
their field of view rather than becoming a
composite color of their full
background.[30]

Evidence of original coloration has been

detected in cephalopod fossils dating as
far back as the Silurian; these orthoconic
individuals bore concentric stripes, which
are thought to have served as
camouflage.[31] Devonian cephalopods
bear more complex color patterns, of
unknown function.[32]
Ink

With the exception of the Nautilidae and

the species of octopus belonging to the
suborder Cirrina,[33] all known
cephalopods have an ink sac, which can
be used to expel a cloud of dark ink to
confuse predators.[18] This sac is a
muscular bag which originated as an
extension of the hindgut. It lies beneath
the gut and opens into the anus, into
which its contents – almost pure melanin
– can be squirted; its proximity to the
base of the funnel means the ink can be
distributed by ejected water as the
cephalopod uses its jet propulsion.[18]
The ejected cloud of melanin is usually
mixed, upon expulsion, with mucus,
produced elsewhere in the mantle, and
therefore forms a thick cloud, resulting in
visual (and possibly chemosensory)
impairment of the predator, like a
smokescreen. However, a more
sophisticated behavior has been
observed, in which the cephalopod
releases a cloud, with a greater mucus
content, that approximately resembles
the cephalopod that released it (this
decoy is referred to as a Pseudomorph).
This strategy often results in the predator
attacking the pseudomorph, rather than
its rapidly departing prey.[18] For more
information, see Inking behaviors.
The ink sac of cephalopods has led to a
common name of "inkfish",[34] formerly
the pen-and-ink fish.[35]

Viscera of
Chtenopteryx sicula

Viscera of Ocythoe
tuberculata

Circulatory system
Cephalopods are the only mollusks with
a closed circulatory system. Coleoids
have two gill hearts (also known as
branchial hearts) that move blood
through the capillaries of the gills. A
single systemic heart then pumps the
oxygenated blood through the rest of the
body.[36]

Like most molluscs, cephalopods use

hemocyanin, a copper-containing protein,
rather than hemoglobin, to transport
oxygen. As a result, their blood is
colorless when deoxygenated and turns
blue when exposed to air.[37]

Respiration
Cephalopods exchange gases with the
seawater by forcing water through their
gills, which are attached to the roof of the
organism.[38]:488[39] Water enters the
mantle cavity on the outside of the gills,
and the entrance of the mantle cavity
closes. When the mantle contracts, water
is forced through the gills, which lie
between the mantle cavity and the
funnel. The water's expulsion through the
funnel can be used to power jet
propulsion. The gills, which are much
more efficient than those of other
mollusks, are attached to the ventral
surface of the mantle cavity.[39] There is a
trade-off with gill size regarding lifestyle.
To achieve fast speeds, gills need to be
small – water will be passed through
them quickly when energy is needed,
compensating for their small size.
However, organisms which spend most
of their time moving slowly along the
bottom do not naturally pass much water
through their cavity for locomotion; thus
they have larger gills, along with complex
systems to ensure that water is
constantly washing through their gills,
even when the organism is stationary.[38]
The water flow is controlled by
contractions of the radial and circular
mantle cavity muscles.[40]

The gills of cephalopods are supported

by a skeleton of robust fibrous proteins;
the lack of mucopolysaccharides
distinguishes this matrix from
cartilage.[41][42] The gills are also thought
to be involved in excretion, with NH4+
being swapped with K+ from the
seawater.[39]

Locomotion and buoyancy

Octopuses swim headfirst, with arms trailing behind

While most cephalopods can move by jet

propulsion, this is a very energy-
consuming way to travel compared to the
tail propulsion used by fish.[43] The
efficiency of a propellor-driven waterjet
(i.e. Froude efficiency) is greater than a
rocket.[44] The relative efficiency of jet
propulsion decreases further as animal
size increases; paralarvae are far more
efficient than juvenile and adult
individuals.[45] Since the Paleozoic era, as
competition with fish produced an
environment where efficient motion was
crucial to survival, jet propulsion has
taken a back role, with fins and tentacles
used to maintain a steady velocity.[3]
Whilst jet propulsion is never the sole
mode of locomotion,[3]:208 the stop-start
motion provided by the jets continues to
be useful for providing bursts of high
speed – not least when capturing prey or
avoiding predators.[3] Indeed, it makes
cephalopods the fastest marine
invertebrates,[6]:Preface and they can out-
accelerate most fish.[38] The jet is
supplemented with fin motion; in the
squid, the fins flap each time that a jet is
released, amplifying the thrust; they are
then extended between jets (presumably
to avoid sinking).[45] Oxygenated water is
taken into the mantle cavity to the gills
and through muscular contraction of this
cavity, the spent water is expelled
through the hyponome, created by a fold
in the mantle. The size difference
between the posterior and anterior ends
of this organ control the speed of the jet
the organism can produce.[46] The
velocity of the organism can be
accurately predicted for a given mass
and morphology of animal.[47] Motion of
the cephalopods is usually backward as
water is forced out anteriorly through the
hyponome, but direction can be
controlled somewhat by pointing it in
different directions.[48] Some
cephalopods accompany this expulsion
of water with a gunshot-like popping
noise, thought to function to frighten
away potential predators.[49]

Cephalopods employ a similar method of

propulsion despite their increasing size
(as they grow) changing the dynamics of
the water in which they find themselves.
Thus their paralarvae do not extensively
use their fins (which are less efficient at
low Reynolds numbers) and primarily use
their jets to propel themselves upwards,
whereas large adult cephalopods tend to
swim less efficiently and with more
reliance on their fins.[45]

Nautilus belauensis seen from the front, showing the

opening of the hyponome
Early cephalopods are thought to have
produced jets by drawing their body into
their shells, as Nautilus does today.[50]
Nautilus is also capable of creating a jet
by undulations of its funnel; this slower
flow of water is more suited to the
extraction of oxygen from the water.[50]
The jet velocity in Nautilus is much
slower than in coleoids, but less
musculature and energy is involved in its
production.[51] Jet thrust in cephalopods
is controlled primarily by the maximum
diameter of the funnel orifice (or,
perhaps, the average diameter of the
funnel)[52]:440 and the diameter of the
mantle cavity.[53] Changes in the size of
the orifice are used most at intermediate
velocities.[52] The absolute velocity
achieved is limited by the cephalopod's
requirement to inhale water for
expulsion; this intake limits the maximum
velocity to eight body-lengths per second,
a speed which most cephalopods can
attain after two funnel-blows.[52] Water
refills the cavity by entering not only
through the orifices, but also through the
funnel.[52] Squid can expel up to 94% of
the fluid within their cavity in a single jet
thrust.[44] To accommodate the rapid
changes in water intake and expulsion,
the orifices are highly flexible and can
change their size by a factor of twenty;
the funnel radius, conversely, changes
only by a factor of around 1.5.[52]
Some octopus species are also able to
walk along the seabed. Squids and
cuttlefish can move short distances in
any direction by rippling of a flap of
muscle around the mantle.

While most cephalopods float (i.e. are

neutrally buoyant or nearly so; in fact
most cephalopods are about 2–3%
denser than seawater[11]), they achieve
this in different ways.[43] Some, such as
Nautilus, allow gas to diffuse into the gap
between the mantle and the shell; others
allow purer water to ooze from their
kidneys, forcing out denser salt water
from the body cavity;[43] others, like some
fish, accumulate oils in the liver;[43] and
some octopuses have a gelatinous body
with lighter chlorine ions replacing
sulfate in the body chemistry.[43]

The Macrotritopus defilippi, or the sand-

dwelling octopus, was seen mimicking
both the coloration and the swimming
movements of the sand-dwelling
flounder Bothus lunatus to avoid
predators. The octopuses were able to
flatten their bodies and put their arms
back to appear the same as the
flounders as well as move with the same
speed and movements.[54]

Females of two species, Ocythoe

tuberculata and Haliphron atlanticus,
have evolved a true swim bladder.[55]
Shell

Cross section of
Spirula spirula,
showing the
position of the
shell inside the
mantle

Cuttlebone of
Sepia officinalis

Gladius of
Sepioteuthis
 lessoniana

Nautiluses are the only extant

cephalopods with a true external shell.
However, all molluscan shells are formed
from the ectoderm (outer layer of the
embryo); in cuttlefish (Sepia spp.), for
example, an invagination of the ectoderm
forms during the embryonic period,
resulting in a shell (cuttlebone) that is
internal in the adult.[56] The same is true
of the chitinous gladius of squid[56] and
octopuses.[57] Cirrate octopods have
arch-shaped cartilaginous fin
supports,[58] which are sometimes
referred to as a "shell vestige" or
"gladius".[59] The Incirrina have either a
pair of rod-shaped stylets or no vestige
of an internal shell,[60] and some squid
also lack a gladius.[61] The shelled
coleoids do not form a clade or even a
paraphyletic group.[62] The Spirula shell
begins as an organic structure, and is
then very rapidly mineralized.[63] Shells
that are "lost" may be lost by resorption
of the calcium carbonate component.[64]

Females of the octopus genus Argonauta

secrete a specialized paper-thin egg case
in which they reside, and this is popularly
regarded as a "shell", although it is not
attached to the body of the animal and
has a separate evolutionary origin.
The largest group of shelled
cephalopods, the ammonites, are extinct,
but their shells are very common as
fossils.

The deposition of carbonate, leading to a

mineralized shell, appears to be related
to the acidity of the organic shell matrix
(see Mollusc shell); shell-forming
cephalopods have an acidic matrix,
whereas the gladius of squid has a basic
matrix.[65] The basic arrangement of the
cephalopod outer wall is: an outer
(spherulitic) prismatic layer, a laminar
(nacreous) layer and an inner prismatic
layer. The thickness of every layer
depends on the taxa.[66] In modern
cephalopods, the Ca carbonate is
aragonite. As for other mollusc shells or
coral skeletons, the smallest visible units
are irregular rounded granules.[67]

Left: A giant squid found in Logy Bay, Newfoundland,

in 1873. The two long feeding tentacles are visible
on the extreme left and right.
Right: Detail of the tentacular club of Abraliopsis
morisi
Head appendages

Cephalopods, as the name implies, have

muscular appendages extending from
their heads and surrounding their
mouths. These are used in feeding,
mobility, and even reproduction. In
coleoids they number eight or ten.
Decapods such as cuttlefish and squid
have five pairs. The longer two, termed
tentacles, are actively involved in
capturing prey;[1]:225 they can lengthen
rapidly (in as little as 15
milliseconds[1]:225). In giant squid they
may reach a length of 8 metres. They
may terminate in a broadened, sucker-
coated club.[1]:225 The shorter four pairs
are termed arms, and are involved in
holding and manipulating the captured
organism.[1]:225 They too have suckers,
on the side closest to the mouth; these
help to hold onto the prey.[1]:226 Octopods
only have four pairs of sucker-coated
arms, as the name suggests, though
developmental abnormalities can modify
the number of arms expressed.[68]

The tentacle consists of a thick central

nerve cord (which must be thick to allow
each sucker to be controlled
independently)[69] surrounded by circular
and radial muscles. Because the volume
of the tentacle remains constant,
contracting the circular muscles
decreases the radius and permits the
rapid increase in length. Typically a 70%
lengthening is achieved by decreasing
the width by 23%.[1]:227 The shorter arms
lack this capability.

The size of the tentacle is related to the

size of the buccal cavity; larger, stronger
tentacles can hold prey as small bites are
taken from it; with more numerous,
smaller tentacles, prey is swallowed
whole, so the mouth cavity must be
larger.[70]

Externally shelled nautilids (Nautilus and

Allonautilus) have on the order of 90
finger-like appendages, termed tentacles,
which lack suckers but are sticky instead,
and are partly retractable.

Feeding

The two-part beak of the giant squid, Architeuthis sp.

All living cephalopods have a two-part

beak;[6]:7 most have a radula, although it
is reduced in most octopus and absent
altogether in Spirula.[6]:7[71]:110 They feed
by capturing prey with their tentacles,
drawing it into their mouth and taking
bites from it.[18] They have a mixture of
toxic digestive juices, some of which are
manufactured by symbiotic algae, which
they eject from their salivary glands onto
their captured prey held in their mouths.
These juices separate the flesh of their
prey from the bone or shell.[18] The
salivary gland has a small tooth at its end
which can be poked into an organism to
digest it from within.[18]

The digestive gland itself is rather

short.[18] It has four elements, with food
passing through the crop, stomach and
caecum before entering the intestine.
Most digestion, as well as the absorption
of nutrients, occurs in the digestive
gland, sometimes called the liver.
Nutrients and waste materials are
exchanged between the gut and the
digestive gland through a pair of
connections linking the gland to the
junction of the stomach and caecum.[18]
Cells in the digestive gland directly
release pigmented excretory chemicals
into the lumen of the gut, which are then
bound with mucus passed through the
anus as long dark strings, ejected with
the aid of exhaled water from the
funnel.[18] Cephalopods tend to
concentrate ingested heavy metals in
their body tissue.[72]

Radula
Amphioctopus marginatus eating a crab

The cephalopod radula consists of

multiple symmetrical rows of up to nine
teeth[73] – thirteen in fossil classes.[74]
The organ is reduced or even vestigial in
certain octopus species and is absent in
Spirula.[74] The teeth may be homodont
(i.e. similar in form across a row),
heterodont (otherwise), or ctenodont
(comb-like).[74] Their height, width and
number of cusps is variable between
species.[74] The pattern of teeth repeats,
but each row may not be identical to the
last; in the octopus, for instance, the
sequence repeats every five rows.[74]:79

Cephalopod radulae are known from

fossil deposits dating back to the
Ordovician.[75] They are usually preserved
within the cephalopod's body chamber,
commonly in conjunction with the
mandibles; but this need not always be
the case;[76] many radulae are preserved
in a range of settings in the Mason
Creek.[77] Radulae are usually difficult to
detect, even when they are preserved in
fossils, as the rock must weather and
crack in exactly the right fashion to
expose them; for instance, radulae have
only been found in nine of the 43
ammonite genera,[78] and they are rarer
still in non-ammonoid forms: only three
pre-Mesozoic species possess one.[75]

Excretory system

Most cephalopods possess a single pair

of large nephridia. Filtered nitrogenous
waste is produced in the pericardial
cavity of the branchial hearts, each of
which is connected to a nephridium by a
narrow canal. The canal delivers the
excreta to a bladder-like renal sac, and
also resorbs excess water from the
filtrate. Several outgrowths of the lateral
vena cava project into the renal sac,
continuously inflating and deflating as
the branchial hearts beat. This action
helps to pump the secreted waste into
the sacs, to be released into the mantle
cavity through a pore.[79]

Nautilus, unusually, possesses four

nephridia, none of which are connected
to the pericardial cavities.

The incorporation of ammonia is

important for shell formation in
terrestrial molluscs and other non-
molluscan lineages.[80] Because protein
(i.e. flesh) is a major constituent of the
cephalopod diet, large amounts of
ammonium ions are produced as waste.
The main organs involved with the
release of this excess ammonium are the
gills.[81] The rate of release is lowest in
the shelled cephalopods Nautilus and
Sepia as a result of their using nitrogen
to fill their shells with gas to increase
buoyancy.[81] Other cephalopods use
ammonium in a similar way, storing the
ions (as ammonium chloride) to reduce
their overall density and increase
buoyancy.[81]

Reproduction and life cycle

Female Argonauta argo with eggcase and eggs

Detail of the hectocotylus of Ocythoe tuberculata

A dissected male specimen of Onykia ingens,

showing a non-erect penis (the white tubular
structure located below most of the other organs)

A i f th i hibiti
A specimen of the same species exhibiting an
elongation of the penis to 67 cm in length

Cephalopods are a diverse group of

species, but share common life history
traits, for example, they have a rapid
growth rate and short life spans.[82]
Stearns (1992) suggested that in order to
produce the largest possible number of
viable offspring, spawning events depend
on the ecological environmental factors
of the organism. The majority of
cephalopods do not provide parental
care to their offspring, except, for
example, octopus, which helps this
organism increase the survival rate of
their offspring.[82] Marine species' life
cycles are affected by various
environmental conditions.[83] The
development of a cephalopod embryo
can be greatly affected by temperature,
oxygen saturation, pollution, light
intensity, and salinity.[82] These factors
are important to the rate of embryonic
development and the success of
hatching of the embryos. Food
availability also plays an important role in
the reproductive cycle of cephalopods. A
limitation of food influences the timing of
spawning along with their function and
growth.[83] Spawning time and spawning
vary among marine species; it's
correlated with temperature, though
cephalopods in shallow water spawn in
cold months so that the offspring would
hatch at warmer temperatures. Breeding
can last from several days to a month.[82]

Sexual maturity

Cephalopods that are sexually mature

and of adult size begin spawning and
reproducing. After the transfer of genetic
material to the following generation, the
adult cephalopods then die.[82] Sexual
maturation in male and female
cephalopods can be observed internally
by the enlargement of gonads and
accessory glands.[84] Mating would be a
poor indicator of sexual maturation in
females; they can receive sperm when
not fully reproductively mature and store
them until they are ready to fertilize the
eggs.[83] Males are more aggressive in
their pre-mating competition when in the
presence of immature females than
when competing for a sexually mature
female.[85] Most cephalopod males
develop a hectocotylus, an arm tip which
is capable of transferring their
spermatozoa into the female mantel
cavity. Though not all species use a
hectocotylus; for example, the adult
nautilus releases a spadix.[86] An
indication of sexual maturity of females
is the development of brachial
photophores to attract mates.[87]

Fertilization
Cephalopods are not broadcast
spawners. During the process of
fertilization, the females use sperm
provided by the male via external
fertilization. Internal fertilization is seen
only in octopodes.[84] The initiation of
copulation begins when the male
catches a female and wraps his arm
around her, either in a "male to female
neck" position or mouth to mouth
position, depending on the species. The
males then initiate the process of
fertilization by contracting their mantle
several times to release the
spermatozoa.[88] Cephalopods often
mate several times, which influences
males to mate longer with females that
have previously, nearly tripling the
number of contractions of the mantle.[88]
To ensure the fertilization of the eggs,
female cephalopods release a sperm-
attracting peptide through the gelatinous
layers of the egg to direct the
spermatozoa. Female cephalopods lay
eggs in clutches; each egg is composed
of a protective coat to ensure the safety
of the developing embryo when released
into the water column. Reproductive
strategies differ between cephalopod
species. In giant Pacific octopus, large
eggs are laid in a den; it will often take
several days to lay all of them.[84] Once
the eggs are released and attached to a
sheltered substrate, the females then
die,[84] making them semelparous. In
some species of cephalopods, egg
clutches are anchored to substrates by a
mucilaginous adhesive substance. These
eggs are swelled with perivitelline fluid
(PVF), a hypertonic fluid that prevents
premature hatching.[89] Fertilized egg
clusters are neutrally buoyant depending
on the depth that they were laid, but can
also be found in substrates such as sand,
a matrix of corals, or seaweed.[83]
Because these species do not provide
parental care for their offspring, egg
capsules can be injected with ink by the
female in order to camouflage the
embryos from predators.[83]
Male–male competition

Most cephalopods engage in aggressive

sex: a protein in the male capsule sheath
stimulates this behavior. They also
engage in male–male aggression, where
larger males tend to win the
interactions.[82] When a female is near,
the males charge one another
continuously and flail their arms. If
neither male backs away, the arms
extend to the back, exposing the mouth,
followed by the biting of arm tips.[90]
During mate competition males also
participate in a technique called flushing.
This technique is used by the second
male attempting to mate with a female.
Flushing removes spermatophores in the
buccal cavity that was placed there by
the first mate by forcing water into the
cavity.[82] Another behavior that males
engage in is sneaker mating or mimicry –
smaller males adjust their behavior to
that of a female in order to reduce
aggression. By using this technique, they
are able to fertilize the eggs while the
larger male is distracted by a different
male.[90] During this process, the sneaker
males quickly insert drop-like sperm into
the seminal receptacle.[91]

Mate choice

Mate choice is seen in cuttlefish species,

where females prefer some males over
others, though characteristics of the
preferred males are unknown.[82] A
hypothesis states that females reject
males by olfactory cues rather than
visual cues.[82] Several cephalopod
species are polyandrous- accepting and
storing multiple male spermatophores,
which has been identified by DNA
fingerprinting.[88] Females are no longer
receptive to mating attempts when
holding their eggs in their arms. Females
can store sperm in two places (1) the
buccal cavity where recently mated
males place their spermatophores, and
(2) the internal sperm-storage
receptacles where sperm packages from
previous males are stored.[82]
Spermatophore storage results in sperm
competition; which states that the female
controls which mate fertilizes the eggs.
In order to reduce this sort of
competition, males develop agonistic
behaviors like mate guarding and
flushing.[82] The Hapalochlaena lunulata,
or the blue-ringed octopus, is unable to
distinguish between males and females
and readily mates with both.[92]

Sexual dimorphism

In a variety of marine organisms, it is

seen that females are larger in size
compared to the males in some closely
related species. In some lineages, such
as the blanket octopus, males become
structurally smaller and smaller
resembling a term, "dwarfism" dwarf
males usually occurs at low densities.[93]
The blanket octopus male is an example
of sexual-evolutionary dwarfism; females
grow 10,000 to 40,000 times larger than
the males and the sex ratio between
males and females can be distinguished
right after hatching of the eggs.[93]

Egg cases laid by a female squid

Embryology
Cephalopod eggs span a large range of
sizes, from 1 to 30 mm in diameter.[94]
The fertilised ovum initially divides to
produce a disc of germinal cells at one
pole, with the yolk remaining at the
opposite pole. The germinal disc grows
to envelop and eventually absorb the
yolk, forming the embryo. The tentacles
and arms first appear at the hind part of
the body, where the foot would be in
other molluscs, and only later migrate
towards the head.[79][95]

The funnel of cephalopods develops on

the top of their head, whereas the mouth
develops on the opposite surface.[96]:86
The early embryological stages are
reminiscent of ancestral gastropods and
extant Monoplacophora.[95]

The shells develop from the ectoderm as

an organic framework which is
subsequently mineralized.[56] In Sepia,
which has an internal shell, the ectoderm
forms an invagination whose pore is
sealed off before this organic framework
is deposited.[56]

Development

Chtenopteryx sicula paralarvae. Left: Two very

young paralarvae. The circular tentacular clubs bear
approximately 20 irregularly arranged suckers. Two
chromatophores are present on each side of the
mantle. Centre: Ventral, dorsal and side views of a
more advanced paralarva. An equatorial circulet of
seven large yellow-brown chromatophores is present
on the mantle. Posteriorly the expanded vanes of the
gladius are visible in the dorsal view. Right: Ventral
and dorsal views of a very advanced paralarva.

Left: Immature specimens of Chiroteuthis veranyi. In

this paralarval form, known as the doratopsis stage,
the pen is longer than the mantle and 'neck'
combined
Right: A mature Chiroteuthis veranyi. This species
has some of the longest tentacles in proportion to
its size of any known cephalopod.

The length of time before hatching is

highly variable; smaller eggs in warmer
waters are the fastest to hatch, and
newborns can emerge after as little as a
few days. Larger eggs in colder waters
can develop for over a year before
hatching.[94]

The process from spawning to hatching

follows a similar trajectory in all species,
the main variable being the amount of
yolk available to the young and when it is
absorbed by the embryo.[94]
Unlike most other mollusks, cephalopods
do not have a morphologically distinct
larval stage. Instead, the juveniles are
known as paralarvae. They quickly learn
how to hunt, using encounters with prey
to refine their strategies.[94]

Growth in juveniles is usually allometric,

whilst adult growth is isometric.[97]

Evolution
The traditional view of cephalopod
evolution holds that they evolved in the
Late Cambrian from a monoplacophoran-
like ancestor[98] with a curved, tapering
shell,[99] which was closely related to the
gastropods (snails).[100] The similarity of
the early shelled cephalopod
Plectronoceras to some gastropods was
used in support of this view. The
development of a siphuncle would have
allowed the shells of these early forms to
become gas-filled (thus buoyant) in order
to support them and keep the shells
upright while the animal crawled along
the floor, and separated the true
cephalopods from putative ancestors
such as Knightoconus, which lacked a
siphuncle.[100] Neutral or positive
buoyancy (i.e. the ability to float) would
have come later, followed by swimming
in the Plectronocerida and eventually jet
propulsion in more derived
cephalopods.[101]
However, some morphological evidence
is difficult to reconcile with this view, and
the redescription of Nectocaris pteryx,
which did not have a shell and appeared
to possess jet propulsion in the manner
of "derived" cephalopods, complicated
the question of the order in which
cephalopod features developed –
provided Nectocaris is a cephalopod at
all.[102]

Early cephalopods were likely predators

near the top of the food chain.[18] After
the late Cambrian extinction led to the
disappearance of many
Anomalocaridids, predatory niches
became available for other animals.[103]
During the Ordovician period the primitive
cephalopods underwent pulses of
diversification[104] to become diverse and
dominant in the Paleozoic and Mesozoic
seas.[105]

In the Early Palaeozoic, their range was

far more restricted than today; they were
mainly constrained to sublittoral regions
of shallow shelves of the low latitudes,
and usually occurred in association with
thrombolites.[106] A more pelagic habit
was gradually adopted as the Ordovician
progressed.[106] Deep-water
cephalopods, whilst rare, have been
found in the Lower Ordovician – but only
in high-latitude waters.[106] The mid-
Ordovician saw the first cephalopods
with septa strong enough to cope with
the pressures associated with deeper
water, and could inhabit depths greater
than 100–200 m.[104] The direction of
shell coiling would prove to be crucial to
the future success of the lineages;
endogastric coiling would only permit
large size to be attained with a straight
shell, whereas exogastric coiling –
initially rather rare – permitted the spirals
familiar from the fossil record to develop,
with their corresponding large size and
diversity.[107] (Endogastric mean the shell
is curved so as the ventral or lower side
is longitudinally concave (belly in);
exogastric means the shell is curved so
as the ventral side is longitudinally
convex (belly out) allowing the funnel to
be pointed backward beneath the
shell.)[107]

An ammonoid with the body chamber missing,

showing the septal surface (especially at right) with
its undulating lobes and saddles

The ancestors of coleoids (including

most modern cephalopods) and the
ancestors of the modern nautilus, had
diverged by the Floian Age of the Early
Ordovician Period, over 470 million years
ago.[106][108] The Bactritida, a Silurian–
Triassic group of orthocones, are widely
held to be paraphyletic to the coleoids
and ammonoids, that is, the latter groups
arose from within the Bactritida.[109]:393
An increase in the diversity of the
coleoids and ammonoids is observed
around the start of the Devonian period
and corresponds with a profound
increase in fish diversity. This could
represent the origin of the two derived
groups.[109]

Unlike most modern cephalopods, most

ancient varieties had protective shells.
These shells at first were conical but
later developed into curved nautiloid
shapes seen in modern nautilus species.
Competitive pressure from fish is
thought to have forced the shelled forms
into deeper water, which provided an
evolutionary pressure towards shell loss
and gave rise to the modern coleoids, a
change which led to greater metabolic
costs associated with the loss of
buoyancy, but which allowed them to
recolonize shallow waters.[100]:36
However, some of the straight-shelled
nautiloids evolved into belemnites, out of
which some evolved into squid and
cuttlefish. The loss of the shell may also
have resulted from evolutionary pressure
to increase maneuverability, resulting in a
more fish-like habit.[1]:289
There has been debate on the
embryological origin of cephalopod
appendages.[110] Until the mid-twentieth
century, the "Arms as Head" hypothesis
was widely recognized. In this theory, the
arms and tentacles of cephalopods look
similar to the head appendages of
gastropods, suggesting that they might
be homologous structures. Cephalopod
appendages surround the mouth, so
logically they could be derived from
embryonic head tissues.[111] However,
the "Arms as Foot" hypothesis, proposed
by Adolf Naef in 1928, has increasingly
been favoured;[110] for example, fate
mapping of limb buds in the chambered
nautilus indicates that limb buds
originate from "foot" embryonic
tissues.[112]

Phylogeny

The approximate consensus of extant

cephalopod phylogeny, after Strugnell et
al. 2007, is shown in the cladogram.[62]
Mineralized taxa are in bold. The
attachment of the clade including Sepia
and Spirula is unclear; either of the points
marked with an asterisk may represent
the root of this clade.
  Nautiloids Nautilus
   
Coleoids Basal octopods (e.g.
   
  Argonauta)
   
    Vampyroteuthis
  Heteroteuthis
 
  (bobtail squid)
 

* Sepia
    (cuttlefish)
   
 
  Idiosepius
 
  Sepioteuthis
   
 
   
    Spirula
 Certain
squid (e.g.
* Bathyteuthis)
 

The internal phylogeny of the

cephalopods is difficult to constrain;
many molecular techniques have been
adopted, but the results produced are
conflicting.[62][113] Nautilus tends to be
considered an outgroup, with
Vampyroteuthis forming an outgroup to
other squid; however in one analysis the
nautiloids, octopus and teuthids plot as a
polytomy.[62] Some molecular
phylogenies do not recover the
mineralized coleoids (Spirula, Sepia, and
Metasepia) as a clade; however, others
do recover this more parsimonious-
seeming clade, with Spirula as a sister
group to Sepia and Metasepia in a clade
that had probably diverged before the
end of the Triassic.[114][115]

Molecular estimates for clade divergence

vary. One 'statistically robust' estimate
has Nautilus diverging from Octopus at
415 ± 24 million years ago.[116]

Taxonomy
Chambered nautilus (Nautilus pompilius)

Common cuttlefish (Sepia officinalis)

Atlantic bobtail (Sepiola atlantica)

European squid (Loligo vulgaris)

Common octopus (Octopus vulgaris)

The classification presented here, for

recent cephalopods, follows largely from
Current Classification of Recent
Cephalopoda (May 2001), for fossil
cephalopods takes from Arkell et al.
1957, Teichert and Moore 1964, Teichert
1988, and others. The three subclasses
are traditional, corresponding to the three
orders of cephalopods recognized by
Bather.[117]

Class Cephalopoda († indicates extinct

groups)

Subclass Nautiloidea: Fundamental

ectocochliate cephalopods that
provided the source for the
Ammonoidea and Coleoidea.
Order † Plectronocerida: the
ancestral cephalopods from the
Cambrian Period
Order † Ellesmerocerida
(500 to 470 Ma)
Order † Endocerida
(485 to 430 Ma)
Order † Actinocerida
(480 to 312 Ma)
Order † Discosorida
(482 to 392 Ma)
Order † Pseudorthocerida
(432 to 272 Ma)
Order † Tarphycerida
(485 to 386 Ma)
Order † Oncocerida
(478.5 to 324 Ma)
Order Nautilida (extant; 410.5 Ma
to present)
Order † Orthocerida
(482.5 to 211.5 Ma)
 Order † Ascocerida
(478 to 412 Ma)
Order † Bactritida
(418.1 to 260.5 Ma)
Subclass † Ammonoidea: Ammonites
(479 to 66 Ma)
Order † Goniatitida
(388.5 to 252 Ma)
Order † Ceratitida (254 to 200 Ma)
Order † Ammonitida
(215 to 66 Ma)
Subclass Coleoidea (410.0 Ma-Rec)
Cohort † Belemnoidea: Belemnites
and kin
Genus † Jeletzkya
 Order † Aulacocerida
(265 to 183 Ma)
Order † Phragmoteuthida
(189.6 to 183 Ma)
Order † Hematitida
(339.4 to 318.1 Ma)
Order † Belemnitida
(339.4 to 66 Ma)
Genus † Belemnoteuthis
(189.6 to 183 Ma)
Cohort Neocoleoidea
Superorder Decapodiformes
(also known as Decabrachia
or Decembranchiata)
Order Spirulida: Ram's
horn squid
 Order Sepiida: cuttlefish
Order Sepiolida: pygmy,
bobtail and bottletail
squid
Order Teuthida: squid
Superorder Octopodiformes
(also known as
Vampyropoda)
Family †
Trachyteuthididae
Order Vampyromorphida:
Vampire squid
Order Octopoda: octopus
Superorder †
Palaeoteuthomorpha
Order † Boletzkyida
Other classifications differ, primarily in
how the various decapod orders are
related, and whether they should be
orders or families.

Suprafamilial classification of the

Treatise

This is the older classification that

combines those found in parts K and L of
the Treatise on Invertebrate Paleontology,
which forms the basis for and is retained
in large part by classifications that have
come later.

Nautiloids in general (Teichert and

Moore, 1964) sequence as given.
Subclass † Endoceratoidea. Not used
by Flower, e.g. Flower and Kummel
1950, interjocerids included in the
Endocerida.
Order † Endocerida
Order † Intejocerida
Subclass † Actinoceratoidea Not used
by Flower, ibid
Order † Actinocerida
Subclass Nautiloidea Nautiloidea in
the restricted sense.
Order † Ellesmerocerida
Plectronocerida subsequently split
off as separate order.
Order † Orthocerida Includes
orthocerids and pseudorthocerids
Order † Ascocerida
 Order † Oncocerida
Order † Discosorida
Order † Tarphycerida
Order † Barrandeocerida A
polyphyletic group now included in
the Tarphycerida
Order Nautilida
Subclass † Bactritoidea
Order † Bactritida

Paleozoic Ammonoidea (Miller, Furnish

and Schindewolf, 1957)

Suborder † Anarcestina
Suborder † Clymeniina
Suborder † Goniatitina
Suborder † Prolecanitina
Mesozoic Ammonoidea (Arkel et al.,
1957)

Suborder † Ceratitina
Suborder † Phylloceratina
Suborder † Lytoceratina
Suborder † Ammonitina

Subsequent revisions include the

establishment of three Upper Cambrian
orders, the Plectronocerida,
Protactinocerida, and Yanhecerida;
separation of the pseudorthocerids as
the Pseudorthocerida, and elevating
orthoceratid as the Subclass
Orthoceratoidea.

Shevyrev classification
Shevyrev (2005) suggested a division
into eight subclasses, mostly comprising
the more diverse and numerous fossil
forms,[118][119] although this classification
has been criticized as arbitrary.[120]

Various species of ammonites

Holotype of Ostenoteuthis siroi from family

Ostenoteuthidae.
A fossilised belemnite

Class Cephalopoda

Subclass † Ellesmeroceratoidea
Order † Plectronocerida
(501 to 490 Ma)
Order † Protactinocerida
Order † Yanhecerida
Order † Ellesmerocerida
(500 to 470 Ma)
Subclass † Endoceratoidea
(485 to 430 Ma)
 Order † Endocerida
(485 to 430 Ma)
Order † Intejocerida
(485 to 480 Ma)
Subclass † Actinoceratoidea
Order † Actinocerida
(480 to 312 Ma)
Subclass Nautiloidea (490.0 Ma- Rec)
Order † Basslerocerida
(490 to 480 Ma)
Order † Tarphycerida
(485 to 386 Ma)
Order † Lituitida (485 to 480 Ma)
Order † Discosorida
(482 to 392 Ma)
 Order † Oncocerida
(478.5 to 324 Ma)
Order Nautilida (410.5 Ma-Rec)
Subclass † Orthoceratoidea
(482.5 to 211.5 Ma)
Order † Orthocerida
(482.5 to 211.5 Ma)
Order † Ascocerida
(478 to 412 Ma)
Order † Dissidocerida
(479 to 457.5 Ma)
Order † Bajkalocerida
Subclass † Bactritoidea
(422 to 252 Ma)
Subclass † Ammonoidea
(410 to 66 Ma)
 Subclass Coleoidea (410.0 Ma-rec)[121]

Cladistic classification

Pyritized fossil of Vampyronassa rhodanica, a

vampyromorphid from the Lower Callovian
(166.1 million years ago)

Another recent system divides all

cephalopods into two clades. One
includes nautilus and most fossil
nautiloids. The other clade
(Neocephalopoda or Angusteradulata) is
closer to modern coleoids, and includes
belemnoids, ammonoids, and many
orthocerid families. There are also stem
group cephalopods of the traditional
Ellesmerocerida that belong to neither
clade.[122][123]

The coleoids, despite some doubts,[1]:289

appear from molecular data to be
monophyletic.[124]

In culture

Pen and wash drawing of an imagined colossal

octopus attacking a ship, by the malacologist Pierre
de Montfort, 1801

Ancient seafaring people were aware of

cephalopods, as evidenced by artworks
such as a stone carving found in the
archaeological recovery from Bronze Age
Minoan Crete at Knossos (1900 – 1100
BC) has a depiction of a fisherman
carrying an octopus.[125] The terrifyingly
powerful Gorgon of Greek mythology
may have been inspired by the octopus
or squid, the octopus's body representing
the severed head of Medusa, the beak as
the protruding tongue and fangs, and its
tentacles as the snakes.[126]
The NROL-39 mission patch, depicting the National
Reconnaissance Office as an octopus with a long
reach

The Kraken are legendary sea monsters

of giant proportions said to dwell off the
coasts of Norway and Greenland, usually
portrayed in art as giant cephalopods
attacking ships. Linnaeus included it in
the first edition of his 1735 Systema
Naturae.[127][128] A Hawaiian creation
myth says that the present cosmos is the
last of a series which arose in stages
from the ruins of the previous universe.
In this account, the octopus is the lone
survivor of the previous, alien
universe.[129] The Akkorokamui is a
gigantic tentacled monster from Ainu
folklore.[130]

A battle with an octopus plays a

significant role in Victor Hugo's book
Travailleurs de la mer (Toilers of the Sea),
relating to his time in exile on
Guernsey.[131]Ian Fleming's 1966 short
story collection Octopussy and The Living
Daylights, and the 1983 James Bond film
were partly inspired by Hugo's book.[132]

Japanese erotic art, shunga, includes

ukiyo-e woodblock prints such as
Katsushika Hokusai's 1814 print Tako to
ama (The Dream of the Fisherman's
Wife), in which an ama diver is sexually
intertwined with a large and a small
octopus.[133][134] The print is a forerunner
of tentacle erotica.[135] The biologist P. Z.
Myers noted in his science blog,
Pharyngula, that octopuses appear in
"extraordinary" graphic illustrations
involving women, tentacles, and bare
breasts.[136][137]

Since it has numerous arms emanating

from a common center, the octopus is
often used as a symbol for a powerful
and manipulative organization, usually
negatively.[138]
See also

Book: Cephalopoda

Cephalopod size
Cephalopod eye
Cephalopod intelligence
Pain in cephalopods
Kraken
List of nautiloids
List of ammonites

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Further reading
Barskov, I. S.; Boiko, M. S.; Konovalova,
V. A.; Leonova, T. B.; Nikolaeva, S. V.
(2008). "Cephalopods in the marine
ecosystems of the Paleozoic".
Paleontological Journal. 42 (11):
1167–1284.
doi:10.1134/S0031030108110014 . A
comprehensive overview of Paleozoic
cephalopods.
Campbell, Neil A.; Reece, Jane B.;
Mitchell, Lawrence G. (1999). Biology,
fifth edition. Menlo Park, California:
Addison Wesley Longman, Inc.
ISBN 978-0-8053-6566-5.
Felley, J., Vecchione, M., Roper, C. F. E.,
Sweeney, M. & Christensen, T., 2001–
2003: Current Classification of Recent
Cephalopoda. National Museum of
Natural History: Department of
Systematic Biology: Invertebrate
Zoology: Cephalopods
N. Joan Abbott, Roddy Williamson,
Linda Maddock. Cephalopod
Neurobiology. Oxford University Press,
1995. ISBN 0-19-854790-0
Marion Nixon & John Z. Young. The
brains and lives of Cephalopods.
Oxford University Press, 2003. ISBN 0-
19-852761-6
Hanlon, Roger T. & John B. Messenger.
Cephalopod Behaviour . Cambridge
University Press, 1996. ISBN 0-521-
42083-0
Martin Stevens & Sami Merilaita.
Animal camouflage: mechanisms and
function. Cambridge University Press,
2011. ISBN 0-521-19911-5
Rodhouse, P. G.; Nigmatullin, Ch. M.
(1996). "Role as Consumers".
Philosophical Transactions of the Royal
Society B: Biological Sciences. 351
(1343): 1003–1022.
doi:10.1098/rstb.1996.0090 .
Classification key to modern
cephalopods:
[Link]
0e/[Link]

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