Collection

Cephalopoda
An overview
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Contents
Articles
Cephalopods 1
Cephalopod 1
Octopus 22
Squid 33
Cuttlefish 40
Nautiloid 46
Nautilus 51
Ammonite 58
Belemnoidea 67
Argonaut 71
Anatomy and behaviour 78

Cephalopod intelligence 78
Cephalopod size 81
Cephalopod ink 89
Ink sac 91
Cephalopod arm 91
Hectocotylus 93
Tentacle 94
Dactylus 97
Cephalopod eye 97
Chromatophore 99
Mantle 107
Nidamental gland 110
Siphon 110
Squid giant axon 116
Cephalopod shells 117

Cuttlebone 117
Septum 118
Aptychus 119
Orthocone 120
Phragmocone 120
Siphuncle 121
Body chamber 122
References
Article Sources and Contributors 123
Image Sources, Licenses and Contributors 126
Article Licenses
License 130
1
Cephalopods
Cephalopod
Cephalopods
Bigfin reef squid (Sepioteuthis

lessoniana)
Scientific classification [ e ]
Kingdom: Animalia
Phylum: Mollusca
Class: Cephalopoda
Cuvier, 1797
Subclasses and orders
A cephalopod is any member of the molluscan class Cephalopoda (Greek plural Κεφαλόποδα (kephalópoda);
"head-feet"). 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.
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).
Cephalopod 2
Distribution
There are around 800 extant species of cephalopod,[1] although new species continue to be described. It is estimated
that around 11,000 extinct taxa have been described, although the soft-bodied nature of cephalopods means that they
are not easily fossilised.[2]
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 may be a notable exception in that it tolerates brackish water which
has a low salinity.[3]
Cephalopods occupy most of the depth of the ocean, from the abyssal plane 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).[4]
Nervous system and behaviour
Cephalopods are widely regarded as the most intelligent of the invertebrates and have well developed senses and
large brains; larger than the brains of gastropods. The nervous system of cephalopods is the most complex of the
invertebrates,[5] and their brain to body mass ratio falls between that of warm and cold blooded vertebrates.[4] :14 The
giant nerve fibers of the cephalopod mantle have frequently been used as an experimental material of
neurophysiologists for many years; their large diameter (due to lack of myelination) makes them easier to study.[6]
Cephalopods are social creatures; when isolated from their own kind, they will take to shoaling with fish.[7]
Cephalopod 3
Some cephalopods are able to fly distances up to 50 m. While the organisms are not particularly aerodynamic, they
achieve these rather impressive ranges by use of jet-propulsion; water continues to be expelled from the funnel while
the organism is in flight.[8]
Senses
Cephalopods have advanced vision, can detect gravity with statocysts, and have a variety of chemical sense
organs.[4] :34 Octopuses use their tentacles to explore their environment and can use them for depth perception.[4]
Vision
Most cephalopods rely on vision to detect predators and prey, and to
communicate with one another.[9] 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
The primitive nautilus eye functions similarly to a construction differs as cephalopods lack a cornea, and have an everted
[9]
pinhole camera. retina. Cephalopods' eyes are also sensitive to the plane of
polarization of light. Surprisingly—given their ability to change
[10]
color—all octopuses and most cephalopods[11] are color blind. 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.[12] They also produce visual pigments throughout their body, and may sense light levels directly from
their body.[13] Evidence of color vision has been found in the Sparkling Enope Squid (Watasenia scintillans),[11] [14]
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.[15]
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.
Cephalopod 4
Hearing
Cephalopods can use their statocyst to detect sound.[16]
Use of light
Most cephalopods possess chromatophores - that is, coloured pigments
- which they can use in a startling array of fashions.[4] 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.[4] The bioluminescence is produced
by bacterial symbionts; the host cephalopod is able to detect the light
produced by these organisms.[17] Bioluminescence may also be used to
entice prey, and some species use colourful displays to impress mates,
startle predators, or even communicate with one another.[4] It is not
certain whether bioluminescence is actually of epithelial origin or if it
is a bacterial production.[18]
Colouration
Colouration can be changed in milliseconds as they adapt to their
environment,[4] and the pigment cells are expandable by muscular
contraction.[18] Colouration is typically more pronounced in near-shore
species than those living in the open ocean, whose functions tend to be This Broadclub Cuttlefish (Sepia latimanus) can
restricted to camouflage by breaking their outline.[4] :2 go from camouflage tans and browns (top) to
yellow with dark highlights (bottom) in less than
Evidence of original colouration has been detected in cephalopod a second.
fossils dating as far back as the Silurian; these orthoconic individuals
bore concentric stripes, which are thought to have served as camouflage.[19] Devonian cephalopods bear more
complex colour patterns, whose function may be more complex.[20]
Ink
With the exception of the Nautilidae and the species of octopus belonging to the suborder Cirrina,[21] all known
cephalopods have an ink sac, which can be used to expel a cloud of dark ink to confuse predators.[10] This sac is a
muscular bag which originated as an extension of the hind gut. 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 that the ink
can be distributed by ejected water as the cephalopod uses its jet propulsion.[10] 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 behaviour 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.[10] For more
information, see Inking behaviors.
The inking behaviour of cephalopods has led to a common name of "inkfish", primarily used in fisheries science and
the fishing industry, paralleling the terms white fish, oily fish, and shellfish.
[[file:Chtenopteryx Viscera of Chtenopteryx sicula]][[file:Ocythoe tuberculata Viscera of Ocythoe

sicula2.jpg|thumb|left viscera.jpg|thumb|left tuberculata]]
Cephalopod 5
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.[22]
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.[23]
Respiration
Cephalopods exchange gasses with the seawater by forcing water through their gills, which are attached to the roof
of the organism.[24] :488[25] 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 molluscs, are attached to the ventral surface of the mantle cavity.[25] 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.[24] The water flow is controlled by contractions of the
radial and circular mantle cavity muscles.[26]
The gills of cephalopods are supported by a skeleton of robust fibrous proteins; the lack of mucopolysaccharides
distinguishes this matrix from cartilage.[27] [28] The gills are also thought to be involved in excretion, with NH4+
being swapped with K+ from the seawater.[25]
Locomotion and buoyancy

While all cephalopods can move by jet propulsion, this is a very
energy-consuming way to travel compared to the tail propulsion used
by fish.[29] The relative efficiency of jet propulsion decreases further as
animal size increases; paralarvae are far more efficient than juvenile
and adult individuals.[30] 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.[2] Whilst jet propulsion is never the
sole mode of locomotion,[2] :208 the stop-start motion provided by the
jets continues to be useful for providing bursts of high speed - not least Octopuses swim headfirst, with arms trailing
when capturing prey or avoiding predators.[2] Indeed, it makes behind
cephalopods the fastest marine invertebrates,[4] :Preface and they can
out-accelerate most fish.[24] 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).[31] 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.[32] The velocity of the organism can
be accurately predicted for a given mass and morhpology of animal.[33] 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.[34] Some cephalopods accompany this expulsion of water with a gunshot-like
popping noise, thought to function to frighten away potential predators.[35]
Cephalopod 6
Early cephalopods are thought to have produced jets by drawing their

body into their shells, as Nautilus does today.[36] 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.[36]
The jet velocity in Nautilus is much slower than in coleoids, but less
musculature and energy is involved in its production.[37] Jet thrust in
cephalopods is controlled primarily by the maximum diameter of the
funnel orifice (or, perhaps, the average diameter of the funnel)[38] :440
and the diameter of the mantle cavity.[39] Changes in the size of the
Nautilus belauensis seen from the front, showing
orifice are used most at intermediate velocities.[38] The absolute
the opening of the hyponome
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.[38] Water refills the
cavity by entering not only through the orifices, but also though the funnel.[38] 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.[38]
Some octopus species are also able to walk along the sea bed. 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[7] ), they achieve this in different ways.[29] 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;[29] others, like some fish, accumulate oils in the liver;[29] and some octopuses have a
gelatinous body with lighter chlorine ions replacing sulfate in the body chemistry.[29]
Shell
[[file:Spirula Cross section of Spirula spirula, showing the position Cuttlebone of Sepia Gladius of
spirula.jpg|thumb|right of the shell inside the mantle]][[file:Herklots 1859 I 2 officinalis]][[file:Sepioteuthis lessoniana Sepioteuthis
Sepia officinalis - schelp.jpg|thumb|right gladius.jpg|thumb|right lessoniana]]
Nautiluses are the only extant cephalopods with an 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 that is internal in the adult.[40] The same is true of the chitinous
gladius of squid[40] and octopus.[41] Cirrate octopuses have cartilaginous fin supports,[42] which are sometimes
referred to as a "shell vestige" or "gladius".[43] : The Incirrina have no vestige of an internal shell, and some squid
also lack a gladius.[44] Interestingly, the shelled coleoids do not form a clade or even a paraphyletic group.[45] The
Spirula shell begins as an organic structure, and is then very rapidly mineralized.[46] Shells that are "lost" may be lost
by resorption of the calcium carbonate component.[47]
Females of the octopus genus Argonauta secrete a specialised paper-thin eggcase in which they reside, and this is
popularly regarded as a "shell", although it is not attached to the body of the animal.
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.[48]
Cephalopod 7
Head appendages
Cuttlefish and squid have five pairs of muscular appendages surrounding their mouths. The longer two, termed
tentacles, are actively involved in capturing prey;[49] :225 they can lengthen rapidly (in as little as 15 milliseconds[49]
:225
). In giant squid they may reach a length of 8 metres. They may terminate by broadening into a sucker-coated
club.[49] :225 The shorter four pairs are termed arms, and are involved in holding and manipulating the captured
organism.[49] :225 They too have suckers, on the side closest to the mouth; these help to hold onto the prey.[49] :226
The tentacle consists of a thick central nerve cord (which must be thick to allow each sucker to be controlled
independently)[50] 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%.[49] :227
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.[51]
Feeding
All living cephalopods have a two-part beak;[4] :7 most have a radula,
although it is reduced in most octopus and absent altogether in
Spirula.[4] :7[52] :110 They feed by capturing prey with their tentacles,
drawing it in to their mouth and taking bites from it.[10] 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 mouth. These juices separate the flesh of
their prey from the bone or shell.[10] The salivary gland has a small The two-part beak of the giant squid, Architeuthis
sp.
tooth at its end which can be poked into an organism to digest it from
within.[10]
The digestive gland itself is rather short.[10] 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
Cephalopod 8
gland through a pair of connections linking the gland to the junction of the stomach and caecum.[10] 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.[10]
Radula
The cephalopod radula consists of multiple symmetrical rows of up to
nine teeth[53] – thirteen in fossil classes.[54] The organ is reduced or
even vestigial in certain octopus species and is absent in Spirula.[54]
The teeth may be homodont (i.e. similar in form across a row),
heterodont (otherwise), or ctenodont (comb-like).[54] Their height,
width and number of cusps is variable between species.[54] 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.[54] :79
Cephalopod radulae are known from fossil deposits dating back to the Amphioctopus marginatus eating a crab
Silurian.[54] They are usually preserved within the cephalopod's body
chamber, commonly in conjunction with the mandibles; but this need not always be the case;[55] many radulae are
preserved in a range of settings in the Mason Creek.[56] 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.[57]
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.[58]
Nautilus, unusually, possesses four nephridia, none of which are connected to the pericardial cavities.
Ammonium
The handling of ammonia is thought to be important in shell formation in terrestrial molluscs, and in other
non-molluscan lineages.[59]
Because protein (i.e. flesh) is a major constituent of the cephalopod diet, large amounts of ammonium are produced
as waste. The main organs involved with the release of this excess ammonium are the gills.[60]
The rate of this release is the lowest in the shelled cephalopods Nautilus and Sepia, probably as a result of their use
of nitrogen to fill their shells with gas, in order to produce buoyancy.[60] Other cephalopods use ammonium in a
similar way, storing the ions (as ammonium chloride) themselves in order to reduce their overall density and thus
become more buoyant.[60]
Cephalopod 9
Reproduction and life cycle

[[file:Papierboot Argonauta Female Argonauta argo with eggcase and eggs]][[file:Ocythoe Detail of the hectocotylus of
200705181139.jpg|thumb|right tuberculata hectocotylus.jpg|thumb|right Ocythoe tuberculata]]
[[file:Onykia ingens with A dissected male specimen of Onykia ingens, showing a non-erect penis (the A specimen of the same species
non-erect penis.jpg|thumb|right white tubular structure located below most of the other exhibiting elongation of the penis
organs)]][[file:Onykia ingens with erect penis.jpg|thumb|right to 67 cm in length]]
With a few exceptions, Coleoidea live short lives with rapid growth. Most of the energy extracted from their food is
used for growing. The penis in most male Coleoidea is a long and muscular end of the gonoduct used to transfer
spermatophores to a modified arm called a hectocotylus. That in turn is used to transfer the spermatophores to the
female. In species where the hectocotylus is missing, the penis is long and able to extend beyond the mantle cavity
and transfers the spermatophores directly to the female. 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 barnacles.[61]
Penis elongation in Onykia ingens may result in a penis that is as long as the mantle, head and arms combined.[61]
[62]
Most cephalopods tend towards a semelparous reproduction strategy; they lay many small eggs in one batch and die
afterwards. The Nautiloidea, on the other hand, stick to iteroparity; they produce a few large eggs in each batch and
live for a long time.
External sexual characteristics are lacking in cephalopods, so cephalopods use colour communication. A courting
male will approach a likely looking opposite number flashing his brightest colours, often in rippling displays. If the
other cephalopod is female and receptive, her skin will change colour to become pale, and mating will occur. If the
other cephalopod remains brightly coloured, it is taken as a warning.[63]
The male has a sperm-carrying arm, known as the hectocotylous arm, with which to impregnate the female. In many
cephalopods, mating occurs head to head and the male may simply transfer sperm to the female. Others may detach
the sperm-carrying arm and leave it attached to the female. In the paper nautilus, this arm remains active and
wriggling for some time, prompting the zoologists who discovered it to conclude it was some sort of worm-like
parasite. It was duly given a genus name Hectocotylus, which held for some time until the mistake was
discovered.[63] :227
The eggs may be brooded: female paper nautilus construct a shelter for the young, while Gonatiid squid carry a
larva-laden membrane from the hooks on their arms.[64] Other cephalopods deposit their young under rocks and
aerate them with their tentacles hatching. Often, though, the eggs are left to their own devices; many squid lay
sausage-like bunches of eggs in crevices or occasionally on the sea floor. Cuttlefish lay their eggs separately in cases
and attach them to coral or algal fronds.[65] Fossilised egg clutches show that ammonites also laid clutches of
eggs.[66]
Cephalopods are occasionally long-lived, especially in the deep water or polar forms, but most of the group live fast
and die young, maturing rapidly to their adult size. Some may gain as much as 12% of their body mass each day.[10]
Most live for one to two years,[10] reproducing and then dying shortly thereafter.[67]
In order to free up resources for reproduction, many squid are known to resorb the muscle tissue of their mantle and
tentacles, breaking down the tissue and using the energy contained therein to produce more gametes.[68]
Cephalopod 10
Embryology
Unlike most other molluscs, cephalopods do not have a distinct larval
stage. 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.[58] [69]
Egg cases laid by a female squid
The funnel of cephalopods develops on the top of their head, whereas
the mouth develops on the opposite surface.[70] :86 The early
embryological stages are reminiscent of ancestral gastropods and extant Monoplacophora.[69]
The shells develop from the ectoderm as an organic framework which is subsequently mineralised.[40] In Sepia,
which has an internal shell, the ectoderm forms an invagination whose pore is sealed off before this organic
framework is deposited.[40]
The gene engrailed is expressed first in the arms, funnel and optic vesicles, and is only later present in the tentacles
and eyelids.[40] It is expressed in embryonic stages 17–19 in all arm buds, and subsequently in the future-tentacles in
stages 24–5, suggesting that it may serve a role in the differential development of tentacles. Sequential expression of
Hox genes is also observed in cephalopod arms.[40]
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.
Cephalopod 11
Cephalopod eggs span a large range of sizes, from 1 to 30 mm in diameter.[71] 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.[71]
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.[71]
Young do not pass through a larval stage sensu stricto (in the narrowest sense). They quickly learn how to hunt,
using encounters with prey to refine their strategies.[71]
Growth in juveniles is usually allometric, whilst adult growth is isometric.[72]
Evolution
The traditional view of cephalopod evolution holds that they evolved in the Late Cambrian from a
monoplacophoran-like ancestor[73] with a curved, tapering shell,[74] which was closely related to the gastropods
(snails).[75] 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.[75] 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.[76]
However, some morphological evidence is difficult to reconcile with this view, and the re-description 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.[77] Their position within the Mollusca is currently wide open to interpretation - see Mollusca#Phylogeny.
Early cephalopods were likely predators near the top of the food chain.[10] They underwent pulses of diversification
during the Ordovician period[78] to become diverse and dominant in the Paleozoic and Mesozoic seas.[79] In the
Early Palaeozoic, their range was far more restricted than today; they were mainly constrained to sub-littoral regions
of shallow shelves of the low latitudes, and usually occur in association with thrombolites.[80] A more pelagic habit
was gradually adopted as the Ordovician progressed.[80] Deep-water cephalopods, whilst rare, have been found in the
Lower Ordovician - but only in high-latitude waters.[80] 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.[78] 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.[81] (Endogastric mean the shell is curved so as the ventral or lower side is longitudinally concave
(belly in); exogastric means the shell is curve so as the ventral side is longitudinally convex (belly out) allowing the
funnel to be pointed backwards beneath the shell.)[81]
Cephalopod 12
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.[80] [82] It is widely
held that the Bactritida, an Silurian–Triassic group of
orthocones, are paraphyletic to the coleoids and
An ammonitic ammonoid with the body chamber missing, showing ammonoids – that is, the latter groups arose from
the septal surface (especially at right) with its undulating lobes and within the Bactritida.[83] :393 An increase in the
saddles. 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.[83]
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. It is thought that competitive pressure
from fish 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 recolonise shallow waters.[75] :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 manoeuvrability, resulting in a more fish-like habit.[49] :289
Phylogeny
Cephalopod phylogeny
Nautiloids Nautilus
Basal Octopods (e.g. Argonauta)
Vampyroteuthis
Heteroteuthis (bobtail squid)
Sepia (cuttlefish)
Idiosepius
Coleoids
* Sepioteuthis
Spirula
* Certain squid (e.g. Bathyteuthis)

Cephalopod 13
[45]
Approximate consensus of extant cephalopod phylogeny, after Strugnell et al. 2007 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.
The internal phylogeny of the cephalopods is difficult to constrain; many molecular techniques have been adopted,
but the results produced are conflicting.[45] [84] 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.[45] 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.[85] [86]
Molecular estimates for clade divergence vary. One 'statistically robust' estimate has Nautilus diverging from
Octopus at 415 [87] ± 24 million years ago.[88]
Taxonomy
The classification presented here, for recent cephalopods,
follows largely from Current Classification of Recent
Cephalopoda [89] (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.[90]
Class Cephalopoda († indicates extinct groups)

• Subclass Nautiloidea: Fundamental ectocochliate
cephalopods that provided the source for the
Ammonoidea and Coleoidea. Chambered Nautilus (Nautilus pompilius)
• Order † Plectronocerida: the ancestral cephalopods
from the Cambrian Period
• Order † Ellesmerocerida (500 to 470.0 [91] Ma)
• Order † Endocerida (485 to 430 [92] Ma)
• Order † Actinocerida (480 to 312 [93] Ma)
• Order † Discosorida (482.0 to 392 [94] Ma)
• Order † Pseudorthocerida (432 to 272 [95] Ma)
• Order † Tarphycerida (485 to 386 [96] Ma)
• Order † Oncocerida (478.5 to 324 [97] Ma)
• Order Nautilida (extant; 410.5 to 0 [98] Ma)
• Order † Orthocerida (482.5 to 211.5 [99] Ma)
• Order † Ascocerida (478.0 to 412 [100] Ma)
• Order † Bactritida (418.1 to 260.5 [101] Ma) Common Cuttlefish (Sepia officinalis)
[102]
• Subclass † Ammonoidea: Ammonites (479 to 65
Ma)
• Order † Goniatitida (388.5 to 252 [103] Ma)
• Order † Ceratitida (254 to 200 [104] Ma)
• Order † Ammonitida (215 to 66 [105] Ma)
• Subclass Coleoidea (410.0 Ma-Rec)
• Cohort † Belemnoidea: Belemnites and kin
Cephalopod 14
• Genus † Jeletzkya
• Order † Aulacocerida (265 to 183 [106] Ma)
• Order † Phragmoteuthida (189.6 to 183 [107] Ma)
• Order † Hematitida (339.4 to 318.1 [108] Ma)
• Order † Belemnitida (339.4 to 65.5 [109] Ma)
• Genus † Belemnoteuthis (189.6 to 183 [107] Ma)
• Cohort Neocoleoidea
• Superorder Decapodiformes (also known as
Decabrachia or Decembranchiata)
• ?Order † Boletzkyida
• Order Spirulida: Ram's Horn Squid Atlantic Bobtail (Sepiola atlantica)
• 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
Other classifications differ, primarily in how the various
decapod orders are related, and whether they should be
orders or families.
European Squid (Loligo vulgaris)
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 Common Octopus (Octopus vulgaris)
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
Cephalopod 15
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
orthoceritoids as the Subclass Orthoceratoidea.
Shevyrev classification
Shevyrev (2005) suggested a division into eight subclasses, mostly comprising the more diverse and numerous fossil
forms,[110] [111] although this classification has been criticized as arbitrary.[112]
Class Cephalopoda
• Subclass † Ellesmeroceratoidea
• Order † Plectronocerida (501 to 490.0 [113] Ma)
• Order † Protactinocerida
• Order † Yanhecerida
• Order † Ellesmerocerida (500 to 470.0 [91] Ma)
• Subclass † Endoceratoidea (485 to 430 [92] Ma)
• Order † Endocerida (485 to 430 [92] Ma)
• Order † Intejocerida (485 to 480 [114] Ma)
Various species of ammonites
• Subclass † Actinoceratoidea
• Order † Actinocerida (480 to 312 [93] Ma)
• Subclass Nautiloidea (490.0 Ma- Rec)
• Order † Basslerocerida (490.0 to 480 [115] Ma)
• Order † Tarphycerida (485 to 386 [96] Ma)
• Order † Lituitida (485 to 480 [114] Ma)
• Order † Discosorida (482.0 to 392 [94] Ma)
• Order † Oncocerida (478.5 to 324 [97] Ma)
• Order Nautilida (410.5 Ma-Rec)
• Subclass † Orthoceratoidea (482.5 to 211.5 [99] Ma)
Cephalopod 16
• Order † Orthocerida (482.5 to 211.5 [99] Ma)

• Order † Ascocerida (478.0 to 412 [100] Ma)
• Order † Dissidocerida (479 to 457.5 [116] Ma)
• Order † Bajkalocerida
• Subclass † Bactritoidea (422 to 252 [117] Ma)
• Subclass † Ammonoidea (410 to 66 [118] Ma)
• Subclass Coleoidea (410.0 Ma-rec)[119]
Holotype of Ostenoteuthis siroi from family
Ostenoteuthidae.
A fossilised belemnite
Cladistic classification
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.[121] [122]
Monophyly of coeloids Pyritized fossil of Vampyronassa rhodanica, a vampyromorphid

from the Lower Callovian (unknown operator: u'callovianround'
[120]
The coeloids have been thought to possibly represent a million years ago)
polyphyletic group,[49] :289 although this has not been
supported by the rising body of molecular data.[123]
Post-mortem decay
After death, if undisturbed, cephalopods[124] decay relatively quickly. Their muscle softens within a couple of days,
and may swell; egg sacs can swell so much that they rip through the mantle. Subsequently, the organs shrink again;
at this point the organism may start to break up into fragments. The eyes retain their size while the head shrinks
around them. The gills may remain swollen at this point. After around a week, the carcass collapses in on itself and
begins to disintegrate. The ink sac solidifies around this point. After a fortnight little is left but a blob with eyes,
arms and ink sac visible. After a couple of months, these are only recognisable as flattened dark stains - although in
some cases the eye lenses can remain intact for up to a year.[68]
Cephalopod 17
Notes
[1] [updated 13-Jun-2003] [cit. 27-Feb-2005] http:/ / www. cephbase. utmb. edu/ spdb/ allsp. cfm
[2] Wilbur, Karl M.; Clarke, M.R.; Trueman, E.R., eds. (1985), The Mollusca, 12. Paleontology and neontology of Cephalopods, New York:
Academic Press, ISBN 0-12-728702-7
[3] Bartol, I.K., R. Mann & M. Vecchione (2002). Distribution of the euryhaline squid Lolliguncula brevis in Chesapeake Bay: effects of selected
abiotic factors. Mar. Ecol. Prog. Ser. 226: 235–247.
[4] Marion Nixon; J.Z. Young (2003). The brains and lives of cephalopods. New York: Oxford University Press. ISBN 0-19-852761-6.
[5] http:/ / books. google. com/ books?id=dW5e6FHOH-4C& lpg=PA115& ots=y5Thv-b0Vf&
dq=nervous%20system%20most%20complex%20invertebrates& lr& pg=PA115#v=onepage&
q=nervous%20system%20most%20complex%20invertebrates& f=false
[6] http:/ / www. pnas. org/ content/ 50/ 4/ 619. full. pdf
[7] Packard, A. (1972). "Cephalopods and Fish: the Limits of Convergence". Biological Reviews 47: 241–307.
doi:10.1111/j.1469-185X.1972.tb00975.x.
[8] http:/ / mollus. oxfordjournals. org/ cgi/ pdf_extract/ 70/ 3/ 297
[9] Serb, J. M.; Eernisse, D. J. (2008). "Charting Evolution’s Trajectory: Using Molluscan Eye Diversity to Understand Parallel and Convergent
Evolution". Evolution Education and Outreach 1: 439–447. doi:10.1007/s12052-008-0084-1.
[10] Boyle, Peter; Rodhouse, Paul (2004). Cephalopods : ecology and fisheries (http:/ / books. google. com/ ?id=4UtCi2B4VnoC). Ames, Iowa:
Blackwell. doi:10.1002/9780470995310.ch2. ISBN 0632060484. .
[11] Messenger, John B.; Roger T. Hanlon (1998). Cephalopod Behaviour. Cambridge: Cambridge University Press. pp. 17–21.
ISBN 0-521-64583-2.
[12] Hanlon and Messenger, 68.
[13] Mäthger, L.; Roberts, S.; Hanlon, R. (2010). "Evidence for distributed light sensing in the skin of cuttlefish, Sepia officinalis" (http:/ / www.
pubmedcentral. nih. gov/ articlerender. fcgi?tool=pmcentrez& artid=2936158). Biology letters 6 (5): 600–603. doi:10.1098/rsbl.2010.0223.
PMID 20392722. PMC 2936158.
[14] Michinomae, M; Masuda, H; Seidou, M; Kito, Y (1994). "Structural basis for wavelength discrimination in the banked retina of the firefly
squid Watasenia Scintillans". The Journal of experimental biology 193 (1): 1–12. PMID 9317205.
[15] Seidou, M.; Sugahara, M.; Uchiyama, H.; Hiraki, K.; Hamanaka, T.; Michinomae, M.; Yoshihara, K.; Kito, Y. (1990). "On the three visual
pigments in the retina of the firefly squid, Watasenia scintillans". Journal of Comparative Physiology A 166. doi:10.1007/BF00187321.
[16] "The cephalopods can hear you" (http:/ / news. bbc. co. uk/ earth/ hi/ earth_news/ newsid_8095000/ 8095977. stm). BBC News. 2009-06-15.
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[17] Tong, D.; Rozas, S.; Oakley, H.; Mitchell, J.; Colley, J.; Mcfall-Ngai, J. (Jun 2009). "Evidence for light perception in a bioluminescent
organ" (http:/ / www. pubmedcentral. nih. gov/ articlerender. fcgi?tool=pmcentrez& artid=2700988). Proceedings of the National Academy of
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[18] "integument (mollusks)."Encyclopædia Britannica. 2009. Encyclopædia Britannica 2006 Ultimate Reference Suite DVD
[19] Štěpán Manda and Vojtěch Turek Acta Palaeontologica Polonica 54 (3), 2009: 503-512. Minute Silurian oncocerid nautiloids with unusual
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[20] Colour patterns in Early Devonian cephalopods from the Barrandian Area: Taphonomy and taxonomy Vojtěch Turek Acta Palaeontologica
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[21] Roger T. Hanlon, John B. Messenger: Cephalopod Behaviour, page 2. Cambridge University Press, 1999, ISBN 0521645832
[22] Wells, M.J. (1 April 1980). "Nervous control of the heartbeat in octopus" (http:/ / jeb. biologists. org/ cgi/ content/ abstract/ 85/ 1/ 111).
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Cephalopod 19
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[74] Wingstrand, KG (1985). "On the anatomy and relationships of Recent Monoplacophora" (http:/ / www. zmuc. dk/ inverweb/ Galathea/
Galathea_p5. html) (Link to free full text + plates). Galathea Rep. 16: 7–94. .
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[77] Smith, Martin R and Caron, Jean-Bernard (2010). "Primitive soft-bodied cephalopods from the Cambrian". Nature 465 (7297): 427–428.
doi:10.1038/465427a. PMID 20505713
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[79] Dzik, J. (1981). "Origin of the cephalopoda" (http:/ / www. paleo. pan. pl/ people/ Dzik/ Publications/ Cephalopoda. pdf) (PDF). Acta
Palaeontologica Polonica 26 (2): 161–191. .
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[81] Holland, C. H. (1987). "The nautiloid cephalopods: a strange success: President's anniversary address 1986". Journal of the Geological
Society 144: 1–0. doi:10.1144/gsjgs.144.1.0001.
[82] Kröger, Björn (2006). "Early growth-stages and classification of orthoceridan Cephalopods of the Darriwillian (Middle Ordovician) of
Baltoscandia" (http:/ / www3. interscience. wiley. com/ journal/ 119918421/ abstract). Lethaia 39 (2): 129–139.
doi:10.1080/00241160600623749. .
[83] Young, R.E.; Vecchione, M.; Donovan, D.T.. "The evolution of coleoid cephalopods and their present biodivesity and ecology". In Payne,
AIL; Lipin'ski, M.R.; Clarke, M.R.; Roeleveld, M.A.C. Cephalopod biodiversity, ecology & evolution. South Afriocan journal of Marine
Sciences. 20. pp. 393–420.
[84] Strugnell, J.; Norman, M.; Jackson, J.; Drummond, A.; Cooper, A. (2005). "Molecular phylogeny of coleoid cephalopods (Mollusca:
Cephalopoda) using a multigene approach; the effect of data partitioning on resolving phylogenies in a Bayesian framework". Molecular
phylogenetics and evolution 37 (2): 426–441. doi:10.1016/j.ympev.2005.03.020. PMID 15935706.
[85] Strugnell, J.; Jackson, J.; Drummond, A. J.; Cooper, A. (2006). "Divergence time estimates for major cephalopod groups: evidence from
multiple genes". Cladistics 22: 89–96. doi:10.1111/j.1096-0031.2006.00086.x.
Cephalopod 20
[86] Carlini, DB; Reece, KS; Graves, JE (2000). "Actin gene family evolution and the phylogeny of coleoid cephalopods (Mollusca:
Cephalopoda)". Molecular biology and evolution 17 (9): 1353–70. PMID 10958852.
[87] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. php?Ma=415
[88] Bergmann, S.; Lieb, B.; Ruth, P.; Markl, J. (2006). "The hemocyanin from a living fossil, the cephalopod Nautilus pompilius: protein
structure, gene organization, and evolution". Journal of molecular evolution 62 (3): 362–374. doi:10.1007/s00239-005-0160-x.
PMID 16501879.
[89] http:/ / www. mnh. si. edu/ cephs/ newclass. pdf
[90] Bather, F.A. (1888b). "Professor Blake and Shell-Growth in Cephalopoda.". Annals and Magazine of Natural History Series 6, Vol. 1:
421–426.
[91] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. php?Ma=500–470
[97] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. php?Ma=478. 5–324. 0
[98] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. php?Ma=410. 5–0
[110] Shevyrev, A.A. (2005). "The Cephalopod Macrosystem: A Historical Review, the Present State of Knowledge, and Unsolved Problems: 1.
Major Features and Overall Classification of Cephalopod Mollusks.". 'Paleontological Journal 39 (6): 606–614. "Translated from
Paleontologicheskii Zhurnal No. 6, 2005, 33-42.".
[111] Shevyrev, A. A. (2006). "The cephalopod macrosystem; a historical review, the present state of knowledge, and unsolved problems; 2,
Classification of nautiloid cephalopods". Paleontological Journal 40 (1): 46–54. doi:10.1134/S0031030106010059.
[112] Kroger, B.. "Peer review in the Russian “Paleontological Journal”" (http:/ / www. tiefes-leben. de/ 2009/ 03/
peer-review-in-the-russian-paleontological-journal). .
[119] Bather, F.A. (1888a). "Shell-growth in Cephalopoda (Siphonopoda).". Annals and Magazine of Natural History Series 6, Vol. 1: 298–310.
[120] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. php?Ma=Callovian
[121] Berthold, Thomas, & Engeser, Theo (1987). "Phylogenetic analysis and systematization of the Cephalopoda (Mollusca)". Verhandlungen
Naturwissenschaftlichen Vereins in Hamburg. (NF) 29: 187–220.
[122] Engeser (1997). "Fossil Nautiloidea Page" (http:/ / web. archive. org/ web/ 20060925110442/ http:/ / userpage. fu-berlin. de/ ~palaeont/
fossilnautiloidea/ fossnautcontent. htm). Archived from the original (http:/ / userpage. fu-berlin. de/ ~palaeont/ fossilnautiloidea/
fossnautcontent. htm) on 2006-09-25. .
[123] Lindgren, A. R.; Giribet, G.; Nishiguchi, M. K. (2004). "A combined approach to the phylogeny of Cephalopoda (Mollusca)". Cladistics
20: 454. doi:10.1111/j.1096-0031.2004.00032.x.
[124] Experiments were performed on a variety of squid
Cephalopod 21
References
Further reading
• Barskov, I.S., M.S. Boiko, V.A. Konovalova, T.B. Leonova & S.V. Nikolaeva (2008). "Cephalopods in the
marine ecosystems of the Paleozoic". Paleontological Journal 42: 1167–1284. doi:10.1134/S0031030108110014.
A comprehensive overview of Paleozoic cephalopods.
• Campbell, Neil A., Reece, Jane B., and Mitchell, Lawrence G.: Biology, fifth edition. Addison Wesley Longman,
Inc. Menlo Park, California. 1999. ISBN 0-8053-6566-4.
• Felley, J., Vecchione, M., Roper, C. F. E., Sweeney, M. & Christensen, T., 2001-2003: Current Classification of
Recent Cephalopoda. Internet: National Museum of Natural History: Department of Systematic Biology:
Invertebrate Zoology: http://www.mnh.si.edu/cephs/
External links
• CephBase - cephalopod database (http://www.cephbase.utmb.edu/)
• TONMO.COM - The Octopus News Magazine Online - cephalopod articles and discussion (http://www.tonmo.
com/)
• Tree of Life Web Project - Cephalopoda (http://tolweb.org/Cephalopoda)
• Mikko's Phylogeny Tree (http://www.fmnh.helsinki.fi/users/haaramo/Metazoa/Protostoma/Mollusca/
Cephalopoda/Coleoidea.htm)
• Fish vs. Cephalopods (http://www.cco.caltech.edu/~brokawc/Bi11/cephalopods.html)
• Will Fast Growing Squid Replace Slow Growing Fish? (http://www.atse.org.au/index.php?sectionid=329)
• Biomineralisation in modern and fossil cephalopods (http://biomin.geol.u-psud.fr/ydweb/cephalo/index.htm)
• Scientific American: Can a Squid Fly Out of the Water? (http://www.scientificamerican.com/article.
cfm?id=can-squid-fly)
Octopus 22
Octopus
{{{name}}}
The Common Octopus, Octopus

vulgaris.
Kingdom: Animalia
Phylum: Mollusca
Class: Cephalopoda
Superorder: Octopodiformes
Order: Octopoda
[1]
Leach, 1818
Suborders
• Cirrina
• Incirrina
Synonyms
• Octopoida
[2]
Leach, 1817
The octopus is a cephalopod mollusc of the order Octopoda. Octopuses have two eyes and four pairs of arms, and
like other cephalopods they are bilaterally symmetric. An octopus has a hard beak, with its mouth at the center point
of the arms. Octopuses have no internal or external skeleton (although some species have a vestigial remnant of a
shell inside their mantle), allowing them to squeeze through tight places. Octopuses are among the most intelligent
and behaviorally flexible of all invertebrates.
The octopus inhabits many diverse regions of the ocean, including coral reefs, pelagic waters, and the ocean floor.
They have numerous strategies for defending themselves against predators, including the expulsion of ink, the use of
camouflage and deimatic displays, their ability to jet quickly through the water, and their ability to hide. An octopus
trails its eight arms behind it as it swims. All octopuses are venomous, but only one group, the blue-ringed
octopuses, is known to be deadly to humans.[3]
In the larger sense, there are around 300 recognized octopus species, which is over one-third of the total number of
known cephalopod species. The term octopus may also be used to refer only to those creatures in the genus Octopus.
Octopus 23
Biology
Octopuses are characterized by their eight arms, usually bearing suction cups. The arms of octopuses are often
distinguished from the pair of feeding tentacles found in squid and cuttlefish.[4] Both types of limbs are muscular
hydrostats. Unlike most other cephalopods, the majority of octopuses – those in the suborder most commonly
known, Incirrina – have almost entirely soft bodies with no internal skeleton. They have neither a protective outer
shell like the nautilus, nor any vestige of an internal shell or bones, like cuttlefish or squid. A beak, similar in shape
to a parrot's beak, is the only hard part of their body. This enables them to squeeze through very narrow slits between
underwater rocks, which is very helpful when they are fleeing from morays or other predatory fish. The octopuses in
the less familiar Cirrina suborder have two fins and an internal shell, generally reducing their ability to squeeze into
small spaces. These cirrate species are often free-swimming and live in deep-water habitats, while incirrate octopus
species are found in reefs and other shallower seafloor habitats.
Octopuses have a relatively short life expectancy, and some species
live for as little as six months. Larger species, such as the North Pacific
Giant Octopus, may live for up to five years under suitable
circumstances. However, reproduction is a cause of death: males can
only live for a few months after mating, and females die shortly after
their eggs hatch. They neglect to eat during the (roughly) one month
period spent taking care of their unhatched eggs, but they do not die of
starvation. Endocrine secretions from the two optic glands are the
cause of genetically programmed death (and if these glands are
An octopus moving between tide pools during
surgically removed, the octopus may live many months beyond
low tide
reproduction, until she finally starves).
Octopuses have three hearts. Two branchial hearts pump blood through
each of the two gills, while the third pumps blood through the body.
Octopus blood contains the copper-rich protein hemocyanin for
transporting oxygen. Although less efficient under normal conditions
than the iron-rich hemoglobin of vertebrates, in cold conditions with
low oxygen pressure, hemocyanin oxygen transportation is more
efficient than hemoglobin oxygen transportation. The hemocyanin is
dissolved in the plasma instead of being carried within red blood cells
and gives the blood a bluish color. Octopuses draw water into their
Grimpoteuthis discoveryi, a finned octopus of the
mantle cavity where it passes through its gills. As mollusks, octopuses
suborder Cirrina
have gills that are finely divided and vascularized outgrowths of either
the outer or the inner body surface.
Intelligence
Octopuses are highly intelligent, likely more so than any other order of invertebrates. The exact extent of their
intelligence and learning capability is much debated among biologists,[5] [6] [7] [8] but maze and problem-solving
experiments have shown that they show evidence of a memory system that can store both short- and long-term
memory. It is not known precisely what contribution learning makes to adult octopus behavior. Young octopuses
learn almost no behaviors from their parents, with whom they have very little contact.
Octopus 24
An octopus has a highly complex nervous system, only part of which is

localized in its brain. Two-thirds of an octopus's neurons are found in
the nerve cords of its arms, which have limited functional autonomy.
Octopus arms show a variety of complex reflex actions that persist
even when they have no input from the brain.[9] Unlike vertebrates, the
complex motor skills of octopuses are not organized in their brain
using an internal somatotopic map of its body, as is the motor system
in vertebrates[10] Some octopuses, such as the mimic octopus, will
move their arms in ways that emulate the movements of other sea
creatures.
In laboratory experiments, octopuses can be readily trained to

distinguish between different shapes and patterns. They have been
reported to practice observational learning,[11] although the validity of
these findings is widely contested on a number of grounds.[5] [6]
Octopuses have also been observed in what some have described as
play: repeatedly releasing bottles or toys into a circular current in their
An octopus opening a container with a screw cap aquariums and then catching them.[12] Octopuses often break out of
their aquariums and sometimes into others in search of food. They
have even boarded fishing boats and opened holds to eat crabs.[7]
In some countries, octopuses are on the list of experimental animals on which surgery may not be performed without
anesthesia. In the UK, cephalopods such as octopuses are regarded as honorary vertebrates under the Animals
(Scientific Procedures) Act 1986 and other cruelty to animals legislation, extending to them protections not normally
afforded to invertebrates.[13]
The octopus is the only invertebrate which has been shown to use tools. At least four specimens of the Veined
Octopus (Amphioctopus marginatus) have been witnessed retrieving discarded coconut shells, manipulating them,
and then reassembling them to use as shelter. This discovery was documented in the journal Current Biology and has
also been caught on video.[14] [15]
Octopus 25
Defense
An octopus's main (primary) defense is to hide, either not to be seen at
all, or not to be detected as an octopus.[16] Octopuses have several
secondary defenses (defenses they use once they have been seen by a
predator). The most common secondary defense is fast escape. Other
defenses include the use of ink sacs, camouflage, and autotomising
limbs.
Most octopuses can eject a thick blackish ink in a large cloud to aid in
escaping from predators. The main coloring agent of the ink is
melanin, which is the same chemical that gives humans their hair and
skin color. This ink cloud is thought to reduce the efficiency of
olfactory organs, which would aid an octopus's evasion from predators
that employ smell for hunting, such as sharks. Ink clouds of some
species might serve as pseudomorphs, or decoys that the predator
attacks instead.[17]
Greater Blue-ringed Octopus (Hapalochlaena
lunulata)
An octopus's camouflage is aided by certain specialized skin cells

which can change the apparent color, opacity, and reflectiveness of the
epidermis. Chromatophores contain yellow, orange, red, brown, or
black pigments; most species have three of these colors, while some
have two or four. Other color-changing cells are reflective iridophores,
and leucophores (white).[18] This color-changing ability can also be
used to communicate with or warn other octopuses. The very
venomous blue-ringed octopus becomes bright yellow with blue rings
when it is provoked. Octopuses can use muscles in the skin to change
Amphioctopus marginatus travels with shells it
the texture of their mantle to achieve a greater camouflage. In some
has collected for protection
species the mantle can take on the spiky appearance of seaweed, or the
scraggly, bumpy texture of a rock, among other disguises. However in
some species skin anatomy is limited to relatively patternless shades of one color, and limited skin texture. It is
thought that octopuses that are day-active and/or live in complex habitats such as coral reefs have evolved more
complex skin than their nocturnal and/or sand-dwelling relatives.[16]
When under attack, some octopuses can perform arm autotomy, in a similar manner to the way skinks and other
lizards detach their tails. The crawling arm serves as a distraction to would-be predators.
A few species, such as the Mimic Octopus, have a fourth defense mechanism. They can combine their highly flexible
bodies with their color changing ability to accurately mimic other, more dangerous animals such as lionfish, sea
snakes, and eels.[19] [20]
Reproduction
When octopuses reproduce, males use a specialized arm called a hectocotylus to insert spermatophores (packets of
sperm) into the female's mantle cavity. The hectocotylus in benthic octopuses is usually the third right arm. Males
die within a few months of mating. In some species, the female octopus can keep the sperm alive inside her for
weeks until her eggs are mature. After they have been fertilized, the female lays about 200,000 eggs (this figure
dramatically varies between families, genera, species and also individuals). The female hangs these eggs in strings
Octopus 26
from the ceiling of her lair, or individually attaches them to the substrate depending on the species. The female cares
for the eggs, guarding them against predators, and gently blowing currents of water over them so that they get
enough oxygen. The female does not hunt during the roughly one-month period spent taking care of the unhatched
eggs and may ingest some of her own arms for sustenance. At around the time the eggs hatch, the mother leaves the
lair and is too weak to defend herself from predators like cod, often succumbing to their attacks. The young larval
octopuses spend a period of time drifting in clouds of plankton, where they feed on copepods, larval crabs and larval
starfish until they are ready to descend to the ocean bottom, where the cycle repeats. This is a dangerous time for the
larval octopuses; in the plankton cloud they are vulnerable to plankton eaters. In some deeper dwelling species, the
young do not go through this period.
Sensation
Octopuses have keen eyesight. Octopuses, like other cephalopods, can
distinguish the polarization of light. Color vision appears to vary from
species to species, being present in Octopus aegina but absent in
Octopus vulgaris.[21] Attached to the brain are two special organs,
called statocysts, that allow the octopus to sense the orientation of its
body relative to horizontal. An autonomic response keeps the octopus's
eyes oriented so that the pupil slit is always horizontal.
Octopuses also have an excellent sense of touch. An octopus's suction

cups are equipped with chemoreceptors so that the octopus can taste Eye of Octopus vulgaris
what it is touching. The arms contain tension sensors so that the
octopus knows whether its arms are stretched out. However, the octopus has a very poor proprioceptive sense. The
tension receptors are not sufficient for the octopus brain to determine the position of the octopus's body or arms. (It is
not clear that the octopus brain would be capable of processing the large amount of information that this would
require; the flexibility of an octopus's arms is much greater than that of the limbs of vertebrates, which devote large
areas of cerebral cortex to the processing of proprioceptive inputs.) As a result, the octopus does not possess
stereognosis; that is, it does not form a mental image of the overall shape of the object it is handling. It can detect
local texture variations, but cannot integrate the information into a larger picture.[22]
The neurological autonomy of the arms means that the octopus has great difficulty learning about the detailed effects
of its motions. The brain may issue a high-level command to the arms, but the nerve cords in the arms execute the
details. There is no neurological path for the brain to receive feedback about just how its command was executed by
the arms; the only way it knows just what motions were made is by observing the arms visually.[22]
Octopuses appear to have limited hearing.[23]
Octopuses swim headfirst, with arms trailing

behind
Octopus 27
Locomotion
Octopuses move about by crawling or swimming. Their
main means of slow travel is crawling, with some
swimming. Jet propulsion is their fastest means of
locomotion, followed by swimming and walking.[24]
They crawl by walking on their arms, usually on many at
once, on both solid and soft surfaces, while supported in
water. In 2005 it was reported that some octopuses
(Adopus aculeatus and Amphioctopus marginatus under
current taxonomy) can walk on two arms, while at the
same time resembling plant matter.[25] This form of
locomotion allows these octopuses to move quickly away
from a potential predator while possibly not triggering Video of an octopus in its natural habitat
that predator's search image for octopus (food).[24]
Octopuses swim by expelling a jet of water from a contractile mantle, and aiming it via a muscular siphon.
Size
The North Pacific Giant Octopus, Enteroctopus dofleini, is often cited
as the largest octopus species. Adults usually weigh around 15 kg
(33 lb), with an arm span of up to 4.3 m (14 ft).[26] The largest
specimen of this species to be scientifically documented was an animal
with a live mass of 71 kg (156.5 lb).[27] The alternative contender is the
Seven-arm Octopus, Haliphron atlanticus, based on a 61 kg (134 lb)
carcass estimated to have a live mass of 75 kg (165 lb).[28] [29]
However, there are a number of questionable size records that would
suggest E. dofleini is the largest of all octopus species by a
An adult North Pacific Giant Octopus,
considerable margin;[30] one such record is of a specimen weighing
Enteroctopus dofleini
272 kg (600 lb) and having an arm span of 9 m (30 ft).[31]
Terminology
The term octopus, pronounced /ˈɒktəpʊs/, is from Greek ὀκτάπους (oktapous), "eight-footed",[32] [33] with plural
forms: octopuses /ˈɒktəpʊsɪz/, octopi /ˈɒktəpaɪ/, or octopodes /ɒkˈtɒpədiːz/. Currently, octopuses is the most
common form in both the US and the UK; octopodes is rare, and octopi is often objectionable.[34]
The plural form octopi is often described as a hypercorrection. The Oxford English Dictionary (2008 Draft
Revision)[35] lists octopuses, octopi and octopodes (in that order); it labels octopodes "rare", although the correct
Greek plural form, and notes that octopi derives from the "apprehension" that octōpūs is a second declension Latin
noun, though it is not. It is a Latinization of Greek third-declension masculine oktṓpous (ὀκτώπους, 'eight-foot'),
plural oktṓpodes (ὀκτώποδες). If the word were native to Latin, it would be octōpēs, plural octōpedes, after the
pattern of pēs ('foot'), plural pedēs, analogous to "centipede".[36] The actual Latin word for octopus and other similar
species is polypus, from Greek polýpous (πολύπους, 'many-foot'); usually the inaccurate plural polypī is used instead
of polypodēs.
In modern Greek, the word is khtapódi (χταπόδι), plural khtapódia (χταπόδια), from Medieval oktapódion
(ὀκταπόδιον), equivalent to Classical oktápous (ὀκτάπους), variant of oktṓpous.
Octopus 28
Chambers 21st Century Dictionary[37] and the Compact Oxford Dictionary[38] list only octopuses, although the latter
notes that octopodes is "still occasionally used"; the British National Corpus has 29 instances of octopuses, 11 of
octopi and 4 of octopodes. Merriam-Webster 11th Collegiate Dictionary lists octopuses and octopi, in that order;
Webster's New World College Dictionary lists octopuses, octopi and octopodes (in that order).
Fowler's Modern English Usage states that "the only acceptable plural in English is octopuses," and that octopi is
misconceived and octopodes pedantic.
The term octopod (plural octopods or octopodes) is taken from the taxonomic order Octopoda but has no classical
equivalent. The collective form octopus is usually reserved for animals consumed for food.
Relationship to humans
Ancient peoples of the Mediterranean were aware of the octopus, as evidenced by certain artworks and designs of
prehistory. For example, a stone carving found in the archaeological recovery from Bronze Age Minoan Crete at
Knossos has a depiction of a fisherman carrying an octopus.[39]
Octopuses were often depicted in the art of the Moche people of ancient Peru, who worshipped the sea and its
animals.[40]
Octopus 29
In mythology
The Hawaiian creation myth relates that the present cosmos is only the last of a series, having arisen in stages from
the wreck of the previous universe. In this account, the octopus is the lone survivor of the previous, alien
universe.[41]
In literature
The octopus has a significant role in Victor Hugo's book Travailleurs de la mer (Toilers of the Sea).[42]
As food
Humans eat octopus in many cultures. The arms and sometimes other
body parts are prepared in various ways, often varying by species.
Octopus is a common ingredient in Japanese cuisine, including sushi,
takoyaki, and Akashiyaki. Some small species are sometimes eaten
alive as a novelty food. Similarly, a live octopus may be sliced up and
the legs eaten while still squirming, which continues for some minutes.
Octopus at Tsukiji fish market
Octopus is eaten regularly in Hawaii, since many popular dishes are Asian in
origin. Locally known by their Hawaiian or Japanese names ("he'e" and "tako"
respectively), octopus is also a popular fish bait.
Octopus is a common food in Mediterranean cuisine and Portuguese cuisine. In
Galicia, polbo á feira (market fair style octopus) is a local delicacy. Restaurants
which specialize or serve this dish are known as pulperías. On the Tunisian
island of Djerba, local people catch octopuses by taking advantage of the
animals' habit of hiding in safe places during the night. In the evening they put
grey ceramic pots on the sea bed. The morning of the following day they check
Octopuses are "tickled" out of their
them for octopuses that sheltered there. A common scene in the Greek islands is holes‎ in the Hawaiian Islands with
octopuses hanging in the sunlight from a rope, just like laundry from a 3-pronged polespears
clothesline. They are often caught by spear fishing close to the shore. The
fisherman brings his prey to land and tenderizes the flesh by pounding the carcass against a stone surface. Thus
treated they are hung out to dry, and later will be served grilled either hot, or chilled in a salad. They are considered a
superb meze, especially alongside ouzo.
According to the USDA Nutrient Database (2007), cooked octopus contains approximately 139 calories per three
ounce portion, and is a source of vitamin B3, B12, potassium, phosphorus, and selenium.[43]
Care must be taken to boil the octopus properly, to rid it of slime, smell, and residual ink.
Octopus 30
As pets
Though octopuses can be difficult to keep in captivity, some people keep them as pets. Octopuses often escape even
from supposedly secure tanks, due to their problem-solving skills, mobility and lack of rigid structure.
The variation in size and life span among octopus species makes it difficult to know how long a new specimen can
naturally be expected to live. That is, a small octopus may be just born or may be an adult, depending on its species.
By selecting a well-known species, such as the California Two-spot Octopus, one can choose a small octopus
(around the size of a tennis ball) and be confident that it is young with a full life ahead of it.
Octopuses are also quite strong for their size. Octopuses kept as pets have been known to open the covers of their
aquariums and survive for a time in the air in order to get to a nearby feeder tank and gorge themselves on the fish
there. Large octopuses have also been known to catch and kill some species of sharks.[44]
Classification
• Class CEPHALOPODA
• Subclass Nautiloidea: nautilus
• Subclass Coleoidea
• Superorder Decapodiformes: squid, cuttlefish
• Superorder Octopodiformes
• Family †Trachyteuthididae (incertae sedis)
• Order Vampyromorphida: Vampire Squid
• Order Octopoda
• Genus †Keuppia (incertae sedis)
• Genus †Palaeoctopus (incertae sedis)
• Genus †Paleocirroteuthis (incertae sedis)
• Genus †Pohlsepia (incertae sedis)
• Genus †Proteroctopus (incertae sedis)
• Genus †Styletoctopus (incertae sedis)
• Suborder Cirrina: finned deep-sea octopus
Cirrothauma murrayi
• Family Opisthoteuthidae: umbrella octopus
• Family Cirroteuthidae
• Family Stauroteuthidae
• Suborder Incirrina
• Family Amphitretidae: telescope octopus
• Family Bolitaenidae: gelatinous octopus
• Family Octopodidae: benthic octopus
Amphitretus pelagicus
• Family Vitreledonellidae: Glass Octopus
• Superfamily Argonautoida
• Family Alloposidae: Seven-arm Octopus
• Family Argonautidae: argonauts
• Family Ocythoidae: Tuberculate Pelagic Octopus
• Family Tremoctopodidae: blanket octopus
Octopus 31
See also
• Octopus wrestling
• Legend of the Octopus
• Henry the Hexapus, a six-armed octopus
• Paul the Octopus, an octopus famous for predicting Germany national football team results
References
[1] ITIS Report: Octopoda Leach, 1818 (http:/ / www. itis. gov/ servlet/ SingleRpt/ SingleRpt?search_topic=TSN& search_value=82589)
[2] Helsinki.fi (http:/ / www. helsinki. fi/ ~mhaaramo/ metazoa/ protostoma/ mollusca/ Cephalopoda/ Coleoidea. htm), Mikko's Phylogeny
Archive: Coleoidea – Recent cephalopods
[3] Unimelb.edu.au (http:/ / uninews. unimelb. edu. au/ view. php?articleID=5755), Tentacles of venom: new study reveals all octopuses are
venomous, University of Melbourne, Media Release, Wednesday 15 April 2009
[4] Norman, M. 2000. Cephalopods: A World Guide. ConchBooks, Hackenheim. p. 15. "There is some confusion around the terms arms versus
tentacles. The numerous limbs of nautiluses are called tentacles. The ring of eight limbs around the mouth in cuttlefish, squids and octopuses
are called arms. Cuttlefish and squid also have a pair of specialized limbs attached between the bases of the third and fourth arm pairs [...].
These are known as feeding tentacles and are used to shoot out and grab prey."
[5] What is this octopus thinking? (http:/ / www. fortunecity. com/ emachines/ e11/ 86/ cephpod. html). By Garry Hamilton.
[6] NFW.org? (http:/ / www. nwf. org/ nationalwildlife/ article. cfm?articleId=604& issueId=53), Is the octopus really the invertebrate intellect of
the sea, by Doug Stewart. In: National Wildlife. Feb/Mar 1997, vol.35 no.2.
[7] Giant Octopus—Mighty but Secretive Denizen of the Deep (http:/ / web. archive. org/ web/ 20080102230427/ http:/ / nationalzoo. si. edu/
Support/ AdoptSpecies/ AnimalInfo/ GiantOctopus/ default. cfm)
[8] Slate.com (http:/ / www. slate. com/ id/ 2192211/ ), How Smart is the Octopus?
[9] Yoram Yekutieli, Roni Sagiv-Zohar1, Ranit Aharonov, Yaakov Enge, Binyamin Hochner and Tamar Flash (2005). Dynamic Model of the
Octopus Arm. I. Biomechanics of the Octopus Reaching Movement J. Neurophysiology. 94:1443-1458. PMID 15829594
[10] Zullo L, Sumbre G, Agnisola C, Flash T, Hochner B. (2009). Nonsomatotopic organization of the higher motor centers in octopus. Curr
Biol. 19(19):1632-6. PMID 19765993
[11] Octopus intelligence: jar opening (http:/ / news. bbc. co. uk/ 2/ hi/ europe/ 2796607. stm)
[12] What behavior can we expect of octopuses? (http:/ / www. thecephalopodpage. org/ behavior. php). By Dr. Jennifer Mather, Department of
Psychology and Neuroscience, University of Lethbridge and Roland C. Anderson, The Seattle Aquarium.
[13] United Kingdom Animals (Scientific Procedures) act of 1986 (http:/ / www. archive. official-documents. co. uk/ document/ hoc/ 321/
321-xa. htm)
[14] "Octopus snatches coconut and runs" (http:/ / news. bbc. co. uk/ 1/ hi/ sci/ tech/ 8408233. stm). BBC News. 2009-12-14. . Retrieved
2010-05-20.
[15] http:/ / www. edutube. org/ video/ coconut-shelter-evidence-tool-use-octopuses
[16] Hanlon, R.T. & J.B. Messenger 1996. Cephalopod Behaviour. Cambridge University Press, Cambridge.
[17] Caldwell, R. L. (2005). "An Observation of Inking Behavior Protecting Adult Octopus bocki from Predation by Green Turtle (Chelonia
mydas) Hatchlings". Pacific Science 59 (1): 69–72.
[18] Meyers, Nadia. "Tales from the Cryptic: The Common Atlantic Octopus" (http:/ / www. dnr. sc. gov/ marine/ sertc/ species_month. htm).
Southeastern Regional Taxonomic Center. . Retrieved 2006-07-27.
[19] Norman, M.D., J. Finn & T. Tregenza (2001). Dynamic mimicry in an Indo-Malayan octopus. (http:/ / marinebio. org/ upload/ files/ mimic.
pdf)PDF (312 KB) Proceedings of the Royal Society 268: 1755–1758.
[20] Norman, M.D. & F.G.Hochberg (2005). The "Mimic Octopus" (Thaumoctopus mimicus n. gen. et sp.), a new octopus from the tropical
Indo-West Pacific (Cephalopoda: Octopodidae). Molluscan Research 25: 57–70. Abstract (http:/ / www. mapress. com/ mr/ content/ v25/
2005f/ n2p070. htm)
[21] Kawamura, G., et al. (2001). Color Discrimination Conditioning in Two Octopus Octopus aegina and O. vulgaris. (http:/ / rms1. agsearch.
agropedia. affrc. go. jp/ contents/ JASI/ pdf/ society/ 62-1620. pdf)PDF (453 KB) . Nippon Suisan Gakkashi 67(1): 35–39.
[22] Wells. Martin John. Octopus: physiology and behaviour of an advanced invertebrate. London : Chapman and Hall ; New York : distributed
in the U.S.A. by Halsted Press, 1978.
[23] Matt Walker (15 June 2009). "The cephalopods can hear you" (http:/ / news. bbc. co. uk/ earth/ hi/ earth_news/ newsid_8095000/ 8095977.
stm). BBC. . Retrieved 2010-04-02.
[24] Locomotion by Abdopus aculeatus, C.L. Huffard 2006 (http:/ / jeb. biologists. org/ cgi/ content/ abstract/ 209/ 19/ 3697)
[25] Science, vol. 307, p. 1927 (http:/ / www. sciencemag. org/ cgi/ content/ full/ 307/ 5717/ 1927)
[26] Smithsonian National Zoological Park: Giant Pacific Octopus (http:/ / nationalzoo. si. edu/ Animals/ Invertebrates/ Facts/ cephalopods/
FactSheets/ Pacificoctopus. cfm)
[27] Cosgrove, J.A. 1987. Aspects of the Natural History of Octopus dofleini, the Giant Pacific Octopus. M.Sc. Thesis. Department of Biology,
University of Victoria (Canada), 101 pp.
Octopus 32
[28] O'Shea, S. 2004. The giant octopus Haliphron atlanticus (Mollusca : Octopoda) in New Zealand waters. New Zealand Journal of Zoology
31(1): 7–13.
[29] O'Shea, S. 2002. Haliphron atlanticus — a giant gelatinous octopus. Biodiversity Update 5: 1.
[30] Norman, M. 2000. Cephalopods: A World Guide. ConchBooks, Hackenheim. p. 214.
[31] High, W.L. (1976). "The giant Pacific octopus". U.S. National Marine Fisheries Service, Marine Fisheries Review 38 (9): 17–22.
[32] Oktapous (http:/ / www. perseus. tufts. edu/ cgi-bin/ ptext?doc=Perseus:text:1999. 04. 0057:entry=#72829), Henry George Liddell, Robert
Scott, A Greek-English Lexicon, at Perseus
[33] Scientific Latin from Greek ὀκτώποδ-, ὀκτώπους (also ὀκτάποδ- ὀκτάπους) "eight-footed" > ὀκτώ- or ὀκτά- [combination form of ὀκτώ
"eight"] and πόδ-, πούς "foot". Cf. Modern Greek χταπόδι < οκταπόδι < οκταπόδιον < ὀκτάπους.
[34] Peters, Pam (2004). The Cambridge Guide to English Usage. Cambridge: Cambridge University Press. ISBN 0-521-62181-X, p. 388.
[35] OED.com (http:/ / dictionary. oed. com/ cgi/ entry/ 00330051?single=1& query_type=word& queryword=octopus& first=1&
max_to_show=10) (subscription required). Retrieved February 1, 2010.
[36] " centipede (http:/ / oed. com/ search?searchType=dictionary& q=centipede)". Oxford English Dictionary. Oxford University Press. 2nd ed.
1989.
[37] Chambersharrap.co.uk (http:/ / www. chambersharrap. co. uk/ chambers/ features/ chref/ chref. py/ main?title=21st& query=octopus),
Retrieved October 19, 2007.
[38] Askoxford.com (http:/ / www. askoxford. com/ concise_oed/ octopus), Retrieved October 19, 2007.
[39] C. Michael Hogan. 2007 Knossos fieldnotes, The Modern Antiquarian (http:/ / www. themodernantiquarian. com/ site/ 10854/ knossos.
html#fieldnotes)
[40] Berrin, Katherine & Larco Museum. The Spirit of Ancient Peru:Treasures from the Museo Arqueológico Rafael Larco Herrera. New York:
Thames and Hudson, 199 7.
[41] Dixon, Roland Burrage (1916). The Mythology of All Races. 9. Marshall Jones. p. 15.
[42] This article incorporates text from a publication now in the public domain: Chisholm, Hugh, ed (1911). Encyclopædia Britannica (Eleventh
ed.). Cambridge University Press.
[43] Octopus Calories And Nutrition (http:/ / www. annecollins. com/ calories/ calories-octopus-squid. htm)
[44] Archived Google video of an octopus catching a shark (http:/ / video. google. com/ videoplay?docid=-7004909622962894202), from The
Octopus Show by Mike deGruy
External links
• CephBase: Octopoda (http://www.cephbase.dal.ca/spdb/ordergroup.cfm?Orderr=Octopoda)
• TONMO.COM – The Octopus News Magazine Online (http://www.tonmo.com/)
• Tree of Life website: information about cephalopods along with pictures and videos (http://tolweb.org/
tree?group=Cephalopoda)
• Discussion about the plural (http://itre.cis.upenn.edu/~myl/languagelog/archives/000813.html)
• An octopus's shark encounter (http://video.pbs.org:8080/ramgen/wnet/nature/octopus/sharkT1.rm) –
footage of an octopus eating a shark ( also in Quicktime format (http://www.archive.org/download/
Octopus_eats_a_shark_1/octopus.mov))
• Camouflage in action (http://www.mbl.edu/mrc/hanlon/video.html)
• Video showing an Octopus escaping through a 1 inch hole (http://video.google.com/
videoplay?docid=4007016107763801953)
• Bipedal Octopuses- Video, Information, Original paper (http://ist-socrates.berkeley.edu/~chuffard/index_files/
Bipedal_octopuses.htm)
• Why Cephalopods Change Color (http://www.thecephalopodpage.org/cephschool/
WhyCephalopodsChangeColor.pdf)PDF (359 KB)
• Video of walking octopuses (http://berkeley.edu/news/media/releases/2005/03/24_octopus.shtml)
Squid 33
Squid
{{{name}}}
Bigfin Reef Squid, Sepioteuthis

lessoniana
Kingdom: Animalia
Phylum: Mollusca
Class: Cephalopoda
Superorder: Decapodiformes
Order: Teuthida
A. Naef, 1916
Suborders
†Plesioteuthididae (incertae sedis)

Myopsina
Oegopsina
Squid are marine cephalopods of the order Teuthida, which comprises around 300 species. Like all other
cephalopods, squid have a distinct head, bilateral symmetry, a mantle, and arms. Squid, like cuttlefish, have eight
arms arranged in pairs and two, usually longer, tentacles. Squid are strong swimmers and certain species can 'fly' for
short distances out of the water.[1]
Modification from ancestral forms

Squid have differentiated from their ancestral molluscs such that the body plan has been condensed
antero-posteriorly and extended dorso-ventrally. What before may have been the foot of the ancestor is modified into
a complex set of tentacles and highly developed sense organs, including advanced eyes similar to those of
vertebrates.
The ancestral shell has been lost, with only an internal gladius, or pen, remaining. The pen is a feather-shaped
internal structure that supports the squid's mantle and serves as a site for muscle attachment. It is made of a
chitin-like substance.
Squid 34
Anatomy
The main body mass is enclosed in the mantle, which
has a swimming fin along each side. These fins, unlike
in other marine organisms, are not the main source of
locomotion in most species.
The skin is covered in chromatophores, which enable
the squid to change color to suit its surroundings,
making it effectively invisible. The underside is also
almost always lighter than the topside, to provide
camouflage from both prey and predator.
Under the body are openings to the mantle cavity,
which contains the gills (ctenidia) and openings to the
excretory and reproductive systems. At the front of the European Squid (Loligo vulgaris)
mantle cavity lies the siphon, which the squid uses for
locomotion via precise jet propulsion. In this form of locomotion, water is sucked into the mantle cavity and expelled
out of the siphon in a fast, strong jet. The direction of the siphon can be changed, to suit the direction of travel.
Inside the mantle cavity, beyond the siphon, lies the visceral mass, which is covered by a thin, membranous
epidermis. Under this are all the major internal organs.
Nervous system
The giant axon, which may be up to 1 mm (0.04 inches) in diameter in some larger species, innervates the mantle
and controls part of the jet propulsion system.
As cephalopods, squid exhibit relatively high intelligence among invertebrates. For example, groups of Humboldt
squid hunt cooperatively, using active communication. (See Cephalopod intelligence.)
Reproductive system
[[file:Onykia ingens with A dissected male specimen of Onykia ingens, showing a non-erect penis (the A specimen of the same species
non-erect penis.jpg|thumb|right white tubular structure located below most of the other exhibiting elongation of the penis
organs)]][[file:Onykia ingens with erect penis.jpg|thumb|right to 67 cm in length]]
In females the ink sac is hidden from view by a pair of white nidamental glands, which lie anterior to the gills. There
are also red-spotted accessory nidamental glands. Both organs are associated with food manufacture and shells for
the eggs. Females also have a large translucent ovary, situated towards the posterior of the visceral mass.
Males do not possess these organs, but instead have a large testis in place of the ovary, and a spermatophoric gland
and sac. In mature males, this sac may contain spermatophores, which are placed inside the female's mantle during
mating.
Shallow water species of the continental shelf and epipelagic/mesopelagic zones are characterised by the presence of
hectocotyli, specially modified arms used to fertilise the female's eggs.[2] Most deep sea squid lack hectocotyli and
have longer penises; Ancistrocheiridae and Cranchiinae are exceptions.[2] Giant squid of the genus Architeuthis are
unusual in that they possess both a large penis and modified arm tips, although it is uncertain whether the latter are
used for spermatophore transfer.[2] Penis elongation has been observed in the deep water species Onykia ingens;
when erect, the penis may be as long as the mantle, head and arms combined.[2] [3] 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 barnacles.[2]
Squid 35
Digestive system
Like all cephalopods, squid have complex digestive systems. The muscular stomach is found roughly in the midpoint
of the visceral mass. From there, the bolus moves into the caecum for digestion. The caecum, a long, white organ, is
found next to the ovary or testis. In mature squid, more priority is given to reproduction such that the stomach and
caecum often shrivel up during the later life stages. Finally, food goes to the liver (or digestive gland), found at the
siphon end, for absorption. Solid waste is passed out of the rectum. Beside the rectum is the ink sac, which allows a
squid to rapidly discharge black ink into the mantle cavity.
Cardiovascular system
Squid have three hearts. Two branchial hearts feed the
gills, each surrounding the larger systemic heart that
pumps blood around the body. Squid blood contains the
copper-rich protein hemocyanin for transporting
oxygen. The faintly greenish hearts are surrounded by
the renal sacs - the main excretory system. The kidneys
Ventral view of the viscera of the female Chtenopteryx sicula are difficult to identify and stretch from the hearts
(located at the posterior side of the ink sac) to the liver.
The systemic heart is made of three chambers, a lower ventricle and two upper auricles.
Head
The head end bears 8 arms and 2 tentacles, each a form of muscular hydrostat containing many suckers along the
edge. These tentacles do not grow back if severed. In the mature male, one basal half of the left ventral tentacle is
hectocotylised — and ends in a copulatory pad rather than suckers. It is used for intercourse.
The mouth is equipped with a sharp horny beak mainly made of chitin[4] and cross-linked proteins, and 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, including marine species.[5] Captured whales often have indigestible squid beaks in
their stomachs. The mouth contains the radula (the rough tongue common to all molluscs except bivalvia and
aplacophora).
The eyes, on either side of the head, each contain a hard lens. The lens is focused through movement, much like the
lens of a camera or telescope, rather than changing shape as the lens in the human eye does.
Squids appear to have limited hearing.[6]
Size
The majority are no more than
60 centimetres (24 in) long, although
the giant squid may reach 13 metres
(43 ft).[7]
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 Giant squid in Melbourne Aquarium
largest squid known at the time.[8]
In 2003, a large specimen of an abundant[9] but poorly understood species, Mesonychoteuthis hamiltoni (the Colossal
Squid), was discovered. This species may grow to 14 metres (46 ft) in length, making it the largest invertebrate.[10]
Squid 36
Squid have the largest eyes in the animal kingdom. Giant squid are featured in literature and folklore with a
frightening connotation. The Kraken is a legendary tentacled monster possibly based on sightings of real giant squid.
In February 2007, a New Zealand fishing vessel caught a Colossal Squid weighing 495 kilograms (1090 lb) and
measuring around 10 metres (33 ft) off the coast of Antarctica.[11] This specimen represents the largest cephalopod
to ever be scientifically documented.
Classification
Squid are members of the class Cephalopoda, subclass Coleoidea,
order Teuthida, of which there are two major suborders, Myopsina
and Oegopsina (including giant squids like Architeuthis dux). Teuthida
is the largest cephalopod order with around 300 species classified into
29 families.
The order Teuthida is a member of the superorder Decapodiformes
(from the Greek for "ten legs"). Two other orders of decapodiform
cephalopods are also called squid, although they are taxonomically
distinct from Teuthida and differ recognizably in their gross anatomical Bathyteuthis abyssicola
features. They are the bobtail squid of order Sepiolida and the ram's
horn squid of the monotypic order Spirulida. The vampire squid,
however, is more closely related to the octopuses than to any squid.
• CLASS CEPHALOPODA
• Subclass Coleoidea: squid, octopus, cuttlefish
Grimalditeuthis bonplandi
• Superorder Decapodiformes
• ?Order †Boletzkyida
• Order Spirulida: Ram's Horn Squid
• Order Sepiolida: bobtail squid
• Family †Plesioteuthididae (incertae sedis)
• Suborder Myopsina
• Family Australiteuthidae
• Family Loliginidae: inshore, calamari, and grass
squid
• Suborder Oegopsina
• Family Ancistrocheiridae: Sharpear Enope Squid
• Family Architeuthidae: giant squid
• Family Bathyteuthidae
• Family Batoteuthidae: Bush-club Squid
• Family Brachioteuthidae
Histioteuthis reversa
• Family Chiroteuthidae
• Family Chtenopterygidae: comb-finned squid
• Family Cranchiidae: glass squid
• Family Cycloteuthidae
Squid 37
• Family Enoploteuthidae
• Family Gonatidae: armhook squid
• Family Histioteuthidae: jewel squid
• Family Joubiniteuthidae: Joubin's Squid
• Family Lepidoteuthidae: Grimaldi Scaled Squid
• Family Lycoteuthidae Mastigoteuthis flammea
• Family Magnapinnidae: bigfin squid
• Family Mastigoteuthidae: whip-lash squid
• Family Neoteuthidae
• Family Octopoteuthidae
• Family Ommastrephidae: flying squid
• Family Onychoteuthidae: hooked squid
• Family Pholidoteuthidae
• Family Promachoteuthidae
• Family Psychroteuthidae: Glacial Squid
• Family Pyroteuthidae: fire squid
• Family Thysanoteuthidae: rhomboid squid
Onychoteuthis banksii
• Family Walvisteuthidae
• Parateuthis tunicata (incertae sedis)
Commercial fishing
According to the FAO, the cephalopod catch for 2002 was
3173272 tonnes (6.995867×109 lb). Of this, 2,189,206 tonnes, or 75.8
percent, was squid.[12] The following table lists the squid species
fishery catches which exceeded 10000 tonnes ( lb) in 2002.
Pterygioteuthis giardi
Squid 38
[12]
World squid catch in 2002
Species Family Common name Catch Percent

tonnes
Loligo gahi Loliginidae Patagonian squid 24,976 1.1
Loligo pealei Loliginidae Longfin squid 16,684 0.8
[13] Loliginidae 225,958 10.3

Common squids nei
Ommastrephes bartramii Ommastrephidae Neon flying squid 22,483 1.0
Illex argentinus Ommastrephidae Argentine shortfin squid 511,087 23.3
Dosidicus gigas Ommastrephidae Jumbo flying squid 406,356 18.6
Todarodes pacificus Ommastrephidae Japanese flying squid 504,438 23.0
Nototoda russloani Ommastrephidae Wellington Flying Squid 62,234 2.8
[13] Various 414,990 18.6

Squids nei
Total squid 2,189,206 100.0
As food
Many species are popular as food in cuisines as diverse as Chinese,
Greek, Turkish, Japanese, Portuguese, Italian, Spanish, Korean, Indian,
and Filipino.
In English-speaking countries, squid as food is often marketed using
the Italian word calamari. Squid are found abundantly in certain areas,
and provide large catches for fisheries. The body can be stuffed whole,
cut into flat pieces or sliced into rings. The arms, tentacles and ink are
also edible; in fact, the only parts that are not eaten are the beak and
gladius (pen). Squid is a good food source for zinc, manganese and
Fried calamari: breaded, deep-fried squid
high in the recommended daily intake of copper,[14] selenium, vitamin
B12, and riboflavin.[15]
References
[1] Jabr, F. 2010. Fact or Fiction: Can a Squid Fly Out of the Water? (http:/ / www. scientificamerican. com/ article. cfm?id=can-squid-fly&
sc=WR_20100804) Scientific American, August 2, 2010.
[2] Arkhipkin, A.I. & V.V. Laptikhovsky 2010. Observation of penis elongation in Onykia ingens: implications for spermatophore transfer in
deep-water squid. Journal Molluscan Studies, published online on June 30, 2010. doi:10.1093/mollus/eyq019
[3] Walker, M. 2010. Super squid sex organ discovered (http:/ / news. bbc. co. uk/ earth/ hi/ earth_news/ newsid_8792000/ 8792008. stm). BBC
Earth News, July 7, 2010.
[4] Clarke, M.R. (1986). A Handbook for the Identification of Cephalopod Beaks. Oxford: Clarendon Press. ISBN 0-19-857603-X.
[5] Miserez, A; Li, Y; Waite, H; Zok, F (2007). "Jumbo squid beaks: Inspiration for design of robust organic composites". Acta Biomaterialia 3:
139–149. doi:10.1016/j.actbio.2006.09.004.
[6] Matt Walker (15 June 2009). "The cephalopods can hear you" (http:/ / news. bbc. co. uk/ earth/ hi/ earth_news/ newsid_8095000/ 8095977.
stm). BBC. . Retrieved 2010-04-02.
[7] O'Shea, S. (2003.). "Giant Squid and Colossal Squid Fact Sheet" (http:/ / www. tonmo. com/ science/ public/ giantsquidfacts. php). The
Octopus News Magazine Online.. .
[8] Johnson, C. Scott "Sea Creatures and the Problem of Equipment Damage" United States Naval Institute Proceedings August 1978 pp.106-107
[9] Xavier, J.C., P.G. Rodhouse, P.N. Trathan & A.G. Wood 1999. A Geographical Information System (GIS) Atlas of cephalopod distribution in
the Southern Ocean. (http:/ / www. journals. cambridge. org/ production/ action/ cjoGetFulltext?fulltextid=219642)PDF Antarctic Science
11:61-62. online version (http:/ / www. nerc-bas. ac. uk/ public/ mlsd/ squid-atlas/ )
Squid 39
[10] Anderton, H.J. (2007.). "Amazing specimen of world's largest squid in NZ" (http:/ / www. beehive. govt. nz/ ViewDocument.
aspx?DocumentID=28451). New Zealand Government website. .
[11] "Microwave plan for colossal squid" (http:/ / news. bbc. co. uk/ 2/ hi/ asia-pacific/ 6478801. stm). BBC News. March 22, 2007. .
[12] Rodhouse, Paul G (2005). "Review of the state of world marine fishery resources: Fisheries technical paper" (http:/ / www. fao. org/ docrep/
009/ y5852e/ Y5852E08. htm#ch3. 2). World squid resources (FAO) (447). ISBN 95-5-105267-0. .
[13] nei: not elsewhere included
[14] http:/ / www. foodmarketexchange. com/ datacenter/ product/ seafood/ squid/ detail/ dc_pi_sf_squid_0204. htm
[15] http:/ / www. nmfs. noaa. gov/ fishwatch/ species/ market_squid. htm
External links
• CephBase: Teuthida (http://www.cephbase.utmb.edu/spdb/squid.cfm)
• Colossal Squid at the Museum of New Zealand Te Papa Tongarewa (http://www.tepapa.govt.nz/TePapa/
English/CollectionsAndResearch/CollectionAreas/NaturalEnvironment/Molluscs/ColossalSquid/)
• Market squid mating, laying eggs (video) (http://diving.rogerbly.com/video/squid)
• Scientific American - Giant Squid (http://www.sciam.com/article.cfm?chanID=sa003&
articleID=00030326-0783-133B-878383414B7F0000)
• The Cephalopod Page (http://www.thecephalopodpage.org)
• The Octopus News Magazine Online (http://www.tonmo.com)
Cuttlefish 40
Cuttlefish
Cuttlefish
Sepia latimanus, East

Timor
Kingdom: Animalia
Phylum: Mollusca
Class: Cephalopoda
Superorder: Decapodiformes
Order: Sepiida
Zittel, 1895
Suborders and Families
• †Vasseuriina
• †Vasseuriidae
• †Belosepiellidae
• Sepiina
• †Belosaepiidae
• Sepiadariidae
• Sepiidae
Cuttlefish are marine animals of the order Sepiida. They belong to the class Cephalopoda (which also includes
squid, octopuses, and nautiluses). Despite their name, cuttlefish are not fish but molluscs. Recent studies indicate that
cuttlefish are among the most intelligent invertebrates.[1] Cuttlefish also have one of the largest brain-to-body size
ratios of all invertebrates.[1]
The origin of the word cuttlefish can be found in the old English term cudele, which derived in the 15th century from
the Norwegian koddi (cushion, testicle) and the Middle German kudel (pouch), a good description of the
cephalopod's shape. The Greco-Roman world valued the cephalopod as a source of the unique brown pigment that
the creature releases from its siphon when it is alarmed. The word for it in Greek and Latin, sepia (later seppia in
Italian), is used to refer to a brown pigment in English.
Cuttlefish have an internal shell (the cuttlebone), large W-shaped pupils, and eight arms and two tentacles furnished
with denticulated suckers, with which they secure their prey. They generally range in size from 15 cm (5.9 in) to
25 cm (9.8 in), with the largest species, Sepia apama, reaching 50 cm (20 in) in mantle length and over 10.5 kg (23
lb) in weight.[2]
Cuttlefish eat small molluscs, crabs, shrimp, fish, octopuses, worms, and other cuttlefish. Their predators include
dolphins, sharks, fish, seals and other cuttlefish. Their life expectancy is about one to two years.
Cuttlefish 41
Physiology
Cuttlebone
Cuttlefish possess an internal structure
called the cuttlebone, which is porous and is
made of aragonite. This provides the
cuttlefish with buoyancy. Buoyancy can be
regulated by changing the gas-to-liquid ratio
in the chambered cuttlebone via the ventral
siphuncle.[3] Each species has a distinct
shape, size, and pattern of ridges or texture
on the cuttlebone. The cuttlebone is unique
to cuttlefish, one of the features that
distinguishes them from their squid Video of a cuttlefish in its natural habitat
relatives. Jewelers and silversmiths
traditionally use cuttlebones as moulds for casting small objects[4] but they are probably better known as the tough
material given to parakeets and other caged birds as a source of dietary calcium.
This Broadclub Cuttlefish (Sepia

latimanus) can go from camouflage tans
and browns (top) to yellow with dark
highlights (bottom) in less than a second.
Cuttlefish 42
Skin
Cuttlefish are sometimes referred to as the chameleon
of the sea because of their remarkable ability to rapidly
alter their skin color at will. Cuttlefish change color and
light polarity to communicate to other cuttlefish and to
camouflage themselves from predators.
This color-changing function is produced by groups of
red, yellow, brown, and black pigmented
chromatophores above a layer of reflective iridophores
and leucophores, with up to 200 of these specialized
pigment cells per square millimeter, which corresponds
to about 359 DPI. The pigmented chromatophores have
a sac of pigment and a large membrane that is folded
An infant cuttlefish protects itself with camouflage
when retracted. There are 6-20 small muscle cells on
the sides which can contract to squash the elastic sac
into a disc against the skin. Yellow chromatophores (xanthophores) are closest to the surface of the skin, red and
orange are below (erythrophores), and brown or black are just above the iridophore layer (melanophores). The
iridophores reflect blue and green light. Iridophores are plates of chitin or protein, which can reflect the environment
around a cuttlefish. They are responsible for the metallic blues, greens, golds, and silvers often seen on cuttlefish. All
of these cells can be used in combinations. For example, orange is produced by red and yellow chromatophores,
while purple can be created by a red chromatophore and an iridophore. The cuttlefish can also use an iridophore and
a yellow chromatophore to produce a brighter green. As well as being able to influence the color of light as it reflects
off their skin, cuttlefish can also affect the light's polarization, which can be used to signal to other marine animals,
many of which can also sense polarization.[5]
Eyes
Cuttlefish eyes are among the most developed in the animal kingdom.
The organogenesis of cephalopod eyes differs fundamentally from that
of vertebrates like humans.[6] Superficial similarities between
cephalopod and vertebrate eyes are thought to be examples of
convergent evolution. The cuttlefish pupil is a smoothly-curving W
shape. Although they cannot see color,[7] they can perceive the
polarization of light, which enhances their perception of contrast. They
have two spots of concentrated sensor cells on their retina (known as
foveae), one to look more forward, and one to look more backwards.
The lenses, instead of being reshaped as they are in humans, are pulled
around by reshaping the entire eye to change focus. Unlike the
vertebrate eye, there is no blind spot as the optic nerve is positioned Close up of a cuttlefish eye
behind the retina.
Scientists have speculated that cuttlefish's eyes are fully developed before birth and start observing their
surroundings while still in the egg. One team of French researchers has additionally suggested that cuttlefish prefer
to hunt the prey they saw before hatching.[8]
Cuttlefish 43
Circulation
The blood of a cuttlefish is an unusual shade of green-blue because it uses the copper-containing protein hemocyanin
to carry oxygen instead of the red iron-containing protein hemoglobin that is found in vertebrates' blood. The blood
is pumped by three separate hearts: two branchial hearts pump blood to the cuttlefish's pair of gills (one heart for
each), and the third pumps blood around the rest of the body. Cuttlefish blood must flow more rapidly than most
other animals because hemocyanin carries substantially less oxygen than hemoglobin.
Ink
Cuttlefish have ink, like squid and octopuses, which they use to help evade predators.
Toxicity
Like octopuses and some squid, all cuttlefish have bacterially-produced neurotoxins in their saliva.[9]
The muscles of Pfeffer's Flamboyant Cuttlefish contain
a highly toxic compound that is yet to be identified.[1]
Mark Norman with Museum Victoria in Victoria,
Australia, has shown the toxin to be as lethal as that of
a fellow cephalopod, the blue-ringed octopus.[10]
Ecology
Diet
The preferred diet of the cuttlefish is crabs and fish.[11]
Pfeffer's Flamboyant Cuttlefish from Sipadan, Malaysia
The cuttlefish uses its camouflage to hunt and sneak up
on its prey. When it gets close enough, it opens its eight arms and shoots out two long feeding tentacles. On the end
of each is a pad covered in suckers that grabs and pulls prey toward its beak.[11]
Range and habitat

Family Sepiidae, which contains all cuttlefish, inhabit tropical/temperate ocean waters. They are mostly
shallow-water animals although they are known to go to depths of about 600 metres (2000 ft).[12] They have an
unusual biogeographic pattern: totally absent from the Americas, but present along the coasts of east and south Asia,
western Europe, the Mediterranean, as well as all coasts of Africa and Australia. By the time the family evolved,
ostensibly in the Old World, the north Atlantic possibly had become too cold and deep for these warm water species
to cross.[13]
Cuttlefish 44
Taxonomy
There are over 120 species of cuttlefish currently
recognised, grouped into 5 genera. Sepiadariidae
contains seven species and 2 genera; all the rest are in
Sepiidae.
• CLASS CEPHALOPODA
• Subclass Coleoidea: squid, octopus, cuttlefish
• Superorder Decapodiformes
• ?Order †Boletzkyida
• Order Spirulida: Ram's Horn Squid
Sepia officinalis from Turkish waters
• Suborder †Vasseuriina
• Family †Vasseuriidae
• Family †Belosepiellidae
• Suborder Sepiina
• Family †Belosaepiidae
• Family Sepiadariidae
• Family Sepiidae
• Order Sepiolida: bobtail squid
Relation to humans
Gastronomy
Cuttlefish are caught for food in the Mediterranean, East Asia, the English Channel and elsewhere. Although squid is
more popular as a restaurant dish all over the world, in East Asia dried, shredded cuttlefish is a popular snack food.
Cuttlefish is especially popular in Italy, where it is used in Risotto al
Nero di Seppia (literally black cuttlefish rice). The Croatian Crni Rižot
is virtually the same recipe, which probably originated in Venice and
then spread across both coasts of the Adriatic. "Nero" and "Crni" mean
black, the color the rice turns because of the cuttlefish ink. Spanish
cuisine, especially that of the coastal regions, uses cuttlefish and squid
ink for the marine flavor and smoothness it provides; it is included in
dishes such as rice, pasta and fish stews.
In Portugal, it is the regional dish of the city of Setúbal and Linguine with cuttlefish and ink sauce served at a
surrounding areas, where it is served as deep-fried strips or in a variant Venetian osteria
of feijoada, with red kidney beans.
Cultural significance
Eugenio Montale's ground-breaking debut collection of poetry Cuttlefish Bones (Ossi di seppia) was published in
Turin in 1925. Montale, who grew up in Liguria along the Mediterranean Sea, was awarded the Nobel Prize for
Cuttlefish 45
Literature in 1975, for his long and prolific career. Cuttlefish Bones remains one of the best-known and influential
collections of 20th-century Italian poetry.
Sepia
Cuttlefish ink was formerly an important dye, called sepia. Today artificial dyes have mostly replaced natural sepia.
However, recently some Jews have resumed using sepia for the techelet dye on their Tallit strings.
References
[1] NOVA, 2007. Cuttlefish: Kings of Camouflage. (http:/ / www. pbs. org/ wgbh/ nova/ camo/ ) (television program) NOVA, PBS, April 3,
2007.
[2] Reid, A., P. Jereb, & C.F.E. Roper 2005. Family Sepiidae. In: P. Jereb & C.F.E. Roper, eds. Cephalopods of the world. An annotated and
illustrated catalogue of species known to date. Volume 1. Chambered nautiluses and sepioids (Nautilidae, Sepiidae, Sepiolidae, Sepiadariidae,
Idiosepiidae and Spirulidae). FAO Species Catalogue for Fishery Purposes. No. 4, Vol. 1. Rome, FAO. pp. 57–152.
[3] Rexfort, A.; Mutterlose, J. (2006). "Stable isotope records from Sepia officinalis—a key to understanding the ecology of belemnites?". Earth
and Planetary Science Letters 247: 212–212. doi:10.1016/j.epsl.2006.04.025.
[4] Casting Silver with Cuttlefish (http:/ / www. silverstall. com/ casting-silver-jewellery. html)
[5] Mäthger, L. M., Shashar, N., and R.T. Hanlon. 2009. Do cephalopods communicate using polarized light reflections from their skin? Journal
of Experimental Biology 212: 2133–2140. doi:10.1242/jeb.020800
[6] Muller, Matthew. "Development of the Eye in Vertebrates and Cephalopods and Its Implications for Retinal Structure" (http:/ / www. bio.
davidson. edu/ people/ midorcas/ animalphysiology/ websites/ 2003/ Muller/ development of the cephalopod eye. htm). The Cephalopod Eye.
Davidson College Biology Department. . Retrieved 2007-04-06.
[7] Mäthger, Lydia M.. "Color blindness and contrast perception in cuttlefish (Sepia offcinalis) determined by a visual sensorimotor assay" (http:/
/ www. mbl. edu/ mrc/ hanlon/ pdfs/ mathger_et_al_visres_2006. pdf). Vision Research, Volume 46, Issue 11, May 2006. Elsevier Ltd.. .
Retrieved 2007-09-03.
[8] BBC News (2008-06-05). "Cuttlefish spot target prey early" (http:/ / news. bbc. co. uk/ 1/ hi/ sci/ tech/ 7435757. stm). . Retrieved
2008-05-06.
[9] http:/ / news. nationalgeographic. com/ news/ 2009/ 04/ 090417-octopus-venom. html
[10] Teacher's Guide (http:/ / www. pbs. org/ wgbh/ nova/ teachers/ viewing/ 3404_camo. html) to NOVA episode - Kings of Camouflage (http:/
/ www. pbs. org/ wgbh/ nova/ camo/ ) on PBS (After Watching: Activity 2).
[11] Cuttlefish Basics (http:/ / www. tonmo. com/ articles/ basiccuttlefish. php)
[12] Lu, C. C. and C. F. E. Roper. 1991. Aspects of the biology of Sepia cultrata from southeastern Australia. In: La Seiche, The Cuttlefish.
Boucaud-Camou, E. (Ed). Caen, France; Centre de Publications de l'Université de Caen: 192.
[13] Young, R. E., M. Vecchione and D. Donovan, 1998. The evolution of coleoid cephalopods and their present biodiversity and ecology. South
African Jour. Mar. Sci., 20: 393-420.
External links
• TONMO.com Community Forum - Keeping cuttlefish in the home aquarium (http://www.tonmo.com/forums/
forumdisplay.php?f=18)
• Scientific Database with photos and videos of cuttlefish and other cephalopods (http://www.cephbase.utmb.
edu/)
• YouTube video with examples of color and texture modulation. (http://www.youtube.com/
watch?v=sFYX9D2RQUM)
• Youtube video Cuttlefish changing colour and shape (http://au.youtube.com/watch?v=9XLObuvEryg)
• Amazing cuttlefish - Cephalopods with natural camouflage and sepia ink (http://www.users.on.net/
~jamesmosby/cuttlefish/index.html)
• PBS.org - Nova - Kings of Camouflage (http://www.pbs.org/wgbh/nova/camo/)
Nautiloid 46
Nautiloid
Nautiloids
Orthoceras
Kingdom: Animalia
Phylum: Mollusca
Class: Cephalopoda
Subclass: Nautiloidea
Agassiz, 1847
Orders
Palcephalopoda
• †Plectronocerida
• †Ellesmerocerida
• †Actinocerida
• †Pseudorthocerida
• †Ascocerida
• †Endocerida
• †Tarphycerida
• †Oncocerida
• †Discosorida
• Nautilida
Neocephalopoda (in part)
• †Orthocerida
• †Lituitida
• †Bactritida
Nautiloids are a large and diverse group of marine cephalopods (Mollusca) belonging to the subclass Nautiloidea
that began in the Late Cambrian and are represented today by the living Nautilus. Nautiloids flourished during the
early Paleozoic era, where they constituted the main predatory animals, and developed an extraordinary diversity of
shell shapes and forms. Some 2,500 species of fossil nautiloids are known, but only a handful of species survive to
the present day.
Taxonomic relationships
Nautiloids are among the group of animals known as cephalopods, an advanced class of mollusks which also
includes ammonoids, belemnites and modern coleoids such as octopus and squid. Other mollusks include gastropods,
scaphopods and pelecypods.
Traditionally, the most common classification of the cephalopods has been a three-fold division (by Bather, 1888),
into the nautiloids, ammonoids, and coleoids. This article is about nautiloids in that broad sense, sometimes called
Nautiloidea sensu lato.
Cladistically speaking, nautiloids are a paraphyletic assemblage united by shared primitive (plesiomorphic) features
not found in derived cephalopods. In other words, they are a grade group that is thought to have given rise to both
ammonoids and coleoids, and are defined by the exclusion of both those descendent groups. Both ammonoids and
Nautiloid 47
coleoids have traditionally been assumed to have descended from bactritids, which in turn arose from
straight-shelled orthocerid nautiloids.
The ammonoids (a group which includes the ammonites and the goniatites) are extinct cousins of the nautiloids that
evolved early in the Devonian period, some 400 million years ago.
Some workers apply the name Nautiloidea to a more exclusive group, called Nautiloidea sensu stricto. This taxon
consists only of those orders that are clearly related to the modern nautilus. The membership assigned varies
somewhat from author to author, but usually includes Tarphycerida, Oncocerida, and Nautilida.
Characteristics
The subclass nautiloidea, in the broad original sense, is distinguished by two main characters, simple concave septa,
concave in the forward direction, that produce generally simple sutures, and a siphuncle in which the septal necks
point to the rear, i.e. is retrosiphonate, throughout the ontogeny of the animal.
The septa between the chambers (camerae) of the phragmocone (the chambered part of the shell) are formed during
growth spurts of the animal. At that time the rear of the mantle secretes a new septum adding another chamber while
the more forward part adds on to the shell. The body of the animal, its viscera, continues to occupy the last chamber
of the shell – the living chamber.
The septa are perforated by the siphuncle, which runs through each of the internal chambers of the shell.
Surrounding the fleshy tube of the siphuncle are structures made of Aragonite (a polymorph of Calcium Carbonate –
which during fossilisation is converted to Calcite): septal necks and connecting rings. Some of the earlier nautiloids
deposited calcium carbonate in the empty chambers (called cameral deposits) or within the siphuncle
(endosiphuncular deposits), a process which may have been connected with controlling buoyancy. The nature of the
siphuncle and its position within the shell are important in classifying nautiloids.
Sutures (or suture lines) are visible as a series of narrow wavy lines on the surface of the shell, and they appear
where each septum contacts the wall of the outer shell. The sutures of the nautiloids are simple in shape, being either
straight or slightly curved. This is different from the "zigzag" sutures of the goniatites and the highly complex
sutures of the ammonites.
Modern nautiloids
Much of what is known about the extinct nautiloids is
based on what we know about the modern nautiluses,
such as the Chambered Nautilus which is found in the
south west Pacific Ocean, from Samoa to the
Philippines, and the in the Indian Ocean off of the coast
of Australia. It is not usually found in waters less than
100 meters deep and may be found as far down as 500
to 700 meters (2,300 ft).
Nautiluses are free swimming animals that possess a

head with two simple lens-free eyes and arms (or
tentacles). They each have a smooth shell, with a large
body chamber, which is divided into chambers that are
Nautilus belauensis
filled with an inert gas (similar to air but with more
nitrogen and less oxygen) making the animal buoyant
in the water. As many as 90 tentacles are arranged in two circles around their mouth. The animal has jaws which are
horny and beak-like, and it is a predator, feeding mainly on crustaceans.
Nautiloid 48
Empty nautilus shells may drift a considerable distance and have been reported from Japan, India and Africa.
Undoubtedy the same applies to the shells of fossil nautiloids, the gas inside the shell keeping it buoyant for some
time after the animal's death so that the empty shell was carried some distance from where the animal lived before it
finally sank to the sea-floor.
Nautiluses propel themselves by jet propulsion, expelling water from an elongated funnel called the hyponome,
which can be pointed in different directions to control their movement. They do not have an ink sac like that found in
belemnites and some of the other cephalopods, and there is no evidence to suggest that the extinct forms possessed
an ink sac either. Unlike the extinct ammonoids, the modern nautiluses lack any sort of plate for closing their shell.
With one exception, no such plate has been found in any of the extinct nautiloids either.
The coloration of the shell of the modern nautiluses is quite prominent, and, although it is somewhat rare, the shell
coloration has been known to be preserved in fossil nautiloids. They often show color patterns on the dorsal side
only, which suggests the living animals swam horizontally.
Fossil record
Nautiloids are often found as fossils in early Palaeozoic rocks (less so
in more recent strata). The shells of fossil nautiloids may be either
straight (i.e., orthoconic as in Orthoceras and Rayonnoceras), curved
(as in Cyrtoceras) coiled (as in Cenoceras), or rarely a helical coil (as
in Lorieroceras). Some species' shells—especially in the late Paleozoic
and early Mesozoic—are ornamented with spines and ribs, but most Fossil nautiloid Trilacinoceras from the
have a smooth shell. Ordovician of China.
The shells are formed of aragonite, although the cameral deposits may
consist of primary calcite.[1]
The rocks of the Ordovician period in the Baltic coast and parts of the
United States contain a variety of nautiloid fossils, and specimens such
as Discitoceras and Rayonnoceras may be found in the limestones of
the Carboniferous period in Ireland. The marine rocks of the Jurassic
period in Britain often yield specimens of Cenoceras, and nautiloids
such as Eutrephoceras are also found in the Pierre Shale formation of
the Cretaceous period in the north-central United States.
Fossil orthoconic nautiloid from the Ordovician
Specimens of the Ordovician nautiloid Endoceras have been recorded of Kentucky; an internal mold showing siphuncle
and half-filled camerae, both encrusted.
measuring up to 3.5 meters (13 ft) in length, and Cameroceras is
(somewhat doubtfully) estimated to have reached 11 meters (36 ft).
These large nautiloids must have been formidable predators of other marine animals at the time they lived.
In some localities, such as Scandinavia and Morocco, the fossils of orthoconic nautiloids accumulated in such large
numbers that they form Orthoceras limestones. Although the term Orthoceras now only refers to a Baltic coast
Ordovician genus, in prior times it was employed as a general name given to all straight-shelled nautiloids that lived
from the Ordovician to the Triassic periods (but were most common in the early Paleozoic era.
Nautiloid 49
Evolutionary history
Nautiloids are first known from the late Cambrian Fengshan Formation of northeastern China, where they seem to
have been quite diverse (at the time this was a warm shallow sea rich in marine life). However, although four orders
have been proposed from the 131 species named, there is no certainty that all of these are valid, and indeed it is
likely that these taxa are seriously oversplit.
Most of these early forms died out, but a single family, the Ellesmeroceratidae, survived to the early Ordovician,
where it ultimately gave rise to all subsequent cephalopods. In the Early and Middle Ordovician the nautiloids
underwent an evolutionary radiation.[2] [3] Some eight new orders appeared at this time, covering a great diversity of
shell types and structure, and ecological lifestyles.
Nautiloids remained at the height of their range of adaptations and variety of forms throughout the Ordovician,
Silurian, and Devonian periods, with various straight, curved and coiled shell forms coexisting at the same time.
Several of the early orders became extinct over that interval, but others rose to prominence.
Nautiloids began to decline in the Devonian, perhaps due to competition with their descendants and relatives the
Ammonoids and Coleoids, with only the Nautilida holding their own (and indeed increasing in diversity). Their
shells became increasingly tightly coiled, while both numbers and variety of non-Nautilid species continued to
decrease throughout the Carboniferous and Permian.
The massive extinctions at the end of the Permian were less damaging to nautiloids than to other taxa and a few
groups survived into the early Mesozoic, including pseudorthocerids, bactritids, nautilids and possibly orthocerids.
The last straight-shelled forms were long thought to have disappeared at the end of the Triassic, but a possible
orthocerid has been found in Cretaceous rocks. Apart from that exception, only a single nautiloid suborder, the
Nautilina, continued throughout the Mesozoic, where they co-existed quite happily with their more specialised
ammonoid cousins. Most of these forms differed only slightly from the modern nautilus. They had a brief resurgence
in the early Tertiary (perhaps filling the niches vacated by the ammonoids in the end Cretaceous extinction), and
maintained a worldwide distribution up until the middle of the Cenozoic Era. With the global cooling of the Miocene
and Pliocene, their geographic distribution shrank and these hardy and long-lived animals declined in diversity again.
Today there are only six living species, all belonging to two genera, Nautilus (the pearly nautilus), and Allonautilus.
Classification
Classifications vary and a subject to change as new information is found and in accordance with the perspective of
various workers. The taxonomy of the Taxo Box is one such scheme,Teichert's 1988 classification is another, that of
Teichert et al. 1964 in the Treatise Part K , still another.
Wade (1988) divided the subclass Nautiloidea into 6 superorders, combing orders that are phylogenetically related.
They are the:
• Plectronoceratoidea = Plectronocerida, Protactinocerida, Yanhecerida,and Ellesmerocerida.
• Endoceratoidea = Endocerida
• Orthoceratoidea: = Orthocerida, Ascocerida, and Pseudorthocerida (the Orthoceratoidea of Kroger 1007)
• Nautilitoidea = Tarphycerida, Oncocerida, and Nautilida.
• Actinoceratoidea = Actinocerida
• Discosoritoidea = Discosorida
Three of them are established as equivalent places to put the Endocerida, Actinocerida, and Discosorida. Three unite
related orders that share a common ancestor and form a branch of the nautiloid taxonomic tree; the
Plectronoceratoidea which are mostly small Cambrian forms that include the ancestors of subsequent stocks; the
Orthoceratoidea which unites different primarily orthoconic orders of which one is the source for the Bacritida and
Ammonoidea; and the Nautilitoidea which includes the first coilded cephalopods, the Tarphycerida, as well as the
Nautilida which includes the recent Nautilus
Nautiloid 50
Another order, the Bactritida, which are derived from the Orthocerida are sometimes included with the Nautiloidea,
sometimes with the ammonoidea, and sometimes are placed in a subclass of their own, the Bactritoidea.
Recently some workers in the field have come to recognize the Dissidocerida as a distinct order, along with the
Pseudorthocerida, both previously included in the Orthoceridaas subtaxa.
A more recent interpretation (Engeser 1997-1998) suggests that nautiloids, and indeed cephalopods in general, fall
into two main groups, the Palcephalopoda (including all the nautiloids except Orthocerida and Ascocerida) and the
Neocephalopoda (the rest of the cephalopods).
References
[1] Stehli, F. G. (8 June 1956). "Shell Mineralogy in Paleozoic Invertebrates" (http:/ / jstor. org/ stable/ 1750042). Science 123 (3206):
1031–1032. doi:10.1126/science.123.3206.1031. ISSN 00368075. PMID 17800970. .
[2] KrÖger, B. �R.; Landing, E. (2008). "Onset of the Ordovician cephalopod radiation – evidence from the Rochdale Formation (middle Early
Ordovician, Stairsian) in eastern New York". Geological Magazine 145. doi:10.1017/S0016756808004585.
[3] Kröger, B.; Yun-bai, Y. B. (2009). "Pulsed cephalopod diversification during the Ordovician". Palaeogeography Palaeoclimatology
Palaeoecology 273: 174–201. doi:10.1016/j.palaeo.2008.12.015.
• Doguzhaeva, Larisa. (1994) An Early Cretaceous orthocerid cephalopod from north-western Caucasus.
Palaeontology 37(4): 889-899.
• Engeser, T., (1997-1998) The Palcephalopoda/Neocephalopoda Hypothesis (http://userpage.fu-berlin.de/
~palaeont/fossilnautiloidea/fossnautpalneocephalopoda.htm)
• Teichert, C. (1988) "Main Features of Cephalopod Evolution", in The Mollusca vol.12, Paleontology and
Neontology of Cephalopods, ed. by M.R. Clarke & E.R. Trueman, Academic Press, Harcourt Brace Jovanovich,
External links
• Nautiloids: The First Cephalopods (TONMO.com) (http://www.tonmo.com/science/fossils/nautiloids.php)
• Palaeos (http://www.palaeos.com/Invertebrates/Molluscs/Cephalopoda/Nautiloidea.htm)
• CephBase: Nautiloidea (http://www.cephbase.utmb.edu/spdb/subclassgroup.cfm?Subclass=Nautiloidea)
Nautilus 51
Nautilus
Nautilus
Nautilus belauensis
Scientific classification
Kingdom: Animalia
Phylum: Mollusca
Class: Cephalopoda
Subclass: Nautiloidea
Order: Nautilida
Superfamily: Nautilaceae
Family: Nautilidae
Blainville, 1825
Genera
†Carinonautilus
†Cenoceras
†Eutrephoceras
†Pseudocenoceras
†Strionautilus
Allonautilus
Nautilus
Nautilus (from Greek ναυτίλος, 'sailor') is the common name of marine creatures of cephalopod family Nautilidae,
the sole extant family of the superfamily Nautilaceae and of its smaller but near equal suborder, Nautilina. It
comprises six living species in two genera, the type of which is the genus Nautilus. Though it more specifically
refers to species Nautilus pompilius, the name chambered nautilus is also used for any species of the Nautilidae.
Nautilidae, both extant and extinct, are characterized by involute or slightly evolute shells that are generally smooth,
with compressed or depressed whorl sections, straight to sinuous sutures, and a tubular, generally central
siphuncle.[1] Having survived relatively unchanged for millions of years, nautiluses represent the only living
members of the subclass Nautiloidea, and are often considered "living fossils."
The name "Nautilus" originally referred to the Argonauta, otherwise known as paper nautiluses, because the ancients
believed these animals used their two expanded arms as sails (cf. Aristotle Historia Animalium 622b). However, this
octopus is not closely related to the Nautiloidea.
Nautilus 52
Anatomy
The nautilus is similar in general form to other cephalopods, with a
prominent head and tentacles. Nautiluses typically have more tentacles
than other cephalopods, up to ninety. These tentacles are arranged into
two circles and, unlike the tentacles of other cephalopods, they have no
suckers, are undifferentiated and retractable. The radula is wide and
distinctively has nine teeth. There are two pairs of gills. These are the
only remnants of the ancestral metamerism to be visible in extant
cephalopods.[2] :56
Nautilus pompilius is the largest species in the genus. One form from
western Australia may reach 26.8 centimetres (10.6 in) in diameter.
However, most other nautilus species never exceed 20 centimetres (7.9
in). Nautilus macromphalus is the smallest species, usually measuring
only 16 centimetres (6.3 in).
Shell
Nautiluses are the sole living cephalopods whose bony body structure
The anatomy of Nautilus. The top figure is
is externalized as a shell. The animal can withdraw completely into its dissected; the bottom just has the shell removed.
shell and close the opening with a leathery hood formed from two
specially folded tentacles. The shell is coiled, aragonitic,[3] nacreous and pressure resistant, imploding at a depth of
about 800 metres (2600 ft). The nautilus shell is composed of 2 layers: a matte white outer layer, and a striking white
iridescent inner layer. The innermost portion of the shell is a pearlescent blue-gray. The osmena pearl, contrarily to
its name, is not a pearl, but a jewelry product derived from this part of the shell.
Internally, the shell divides into camerae

(chambers), the chambered section being
called the phragmocone. The divisions are
defined by septa, each of which is pierced in
the middle by a duct, the siphuncle. As the
nautilus matures it creates new, larger
camerae, and moves its growing body into
the larger space, sealing the vacated
chamber with a new septum. The camerae
increase in number from around four at the
Nautilus shells: N. macromphalus (left), A. scrobiculatus (centre), N. pompilius moment of hatching to thirty or more in
(right) adults.
The shell coloration also keeps the animal

cryptic in the water. When seen from above, the shell is darker in color and marked with irregular stripes, which
helps it blend into the dark water below. The underside is almost completely white, making the animal
indistinguishable from brighter waters near the surface. This mode of camouflage is named countershading.
The nautilus shell presents one of the finest natural examples of a logarithmic spiral, although it is not a golden
spiral. The use of nautilus shells in art and literature is covered at nautilus shell.
Nautilus 53
A nautilus shell viewed from above (left), from underneath (centre), and a hemishell showing the camerae in a logarithmic spiral (right)
Tentacles
Nautilus tentacles differ from those of other cephalopods. Lacking pads, the tentacles stick to prey by virtue of their
ridged surface.[4] Nautiloids have a powerful grip. Attempts to take an object already seized by a nautilus may tear
tentacles away from the creature, which remain firmly attached to the surface of the object.[4] Two pairs of tentacles
are separate from the other 90-ish, the pre-ocular and post-ocular, situated before and behind the eye. These are more
evidently grooved, with more pronounced ridges. They are extensively ciliated and serve an olfactory purpose.[4] [5]
[6]
Physiology
Buoyancy and movement

In order to swim, the nautilus draws water into and out of the living
chamber with its hyponome, which uses jet propulsion. While water is
inside the chamber, the siphuncle extracts salt from it and diffuses it
into the blood. The animal adjusts its buoyancy by osmotically
pumping gas and fluid into or out of the camerae along the siphuncles.
This limits them; they cannot operate under the extreme hydrostatic
pressures found at depths greater than approximately 800 metres (2600
ft).
In the wild, nautiluses usually inhabit depths of about 300 metres (980

ft), rising to around 100 metres (330 ft) at night to feed, mate and to lay
eggs.
Nautilus with extended tentacles and hyponome

visible
Nautilus 54
Senses
Unlike many other cephalopods, they do not have good vision; their
eye structure is highly developed but lacks a solid lens. They have a
simple "pinhole" eye open to the environment.
Instead of vision, the animal is thought to use olfaction as the primary
sense for foraging, locating or identifying potential mates.[7]
Reproduction and lifespan

Nautiluses reproduce by laying eggs. Gravid females attach the Head of Nautilus pompilius showing the eye
fertilized eggs to rocks in shallow waters, whereupon the eggs take

eight to twelve months to develop until the 30 millimetres (1.2 in) juveniles hatch. Females spawn once per year and
regenerate their gonads, making nautiluses the only cephalopods to present iteroparity or polycyclic spawning.
Nautiluses are sexually dimorphic, in that males have four tentacles modified into an organ, called the "spadix,"
which transfers sperm into the female's mantle during mating.
The lifespan of nautiluses may exceed 20 years, which is exceptionally lengthy for a cephalopod.[8]
Ecology
Diet
Nautiluses are predators that feed mainly on shrimp, small fish and crustaceans, which are captured by the tentacles.
Due to the limited energy they expend in swimming, nautiloids only need to eat once a month.
Range and habitat

Nautiluses are only found in the Indo-Pacific, from 30° N to 30° S latitude and 90° to 185° W longitude. They
inhabit the deep slopes of coral reefs.
Evolution
Fossil records indicate that nautiluses have not evolved much during the last 500 million years. Many were initially
straight-shelled, as in the extinct genus Lituites. They developed in the Cambrian period and became a significant sea
predator in the Ordovician period. Certain species reached over 2.5 metres (8 ft 2 in) in size. The other cephalopod
subclass, Coleoidea, diverged from the Nautilidae long ago and the nautilus has remained relatively unchanged
since. Nautiloids were much more extensive and varied 200 million years ago. Extinct relatives of the nautilus
include ammonites, such as the baculites and goniatites.
The Nautilidae has its origin in the Trigonocerataceae (Centroceratina), specifically in the Syringonautilidae of the
Late Triassic[1] and continues to this day with Nautilus, the type genus, and its close relative, Allonautilus.
Nautilus 55
Fossil genera
The Nautilidae begin with Cenoceras in the Late Triassic, a highly varied genus that makes up the Jurassic
Cenoceras complex. Cenoceras is evolute to involute, and globular to lentincular; with a suture that generally has a
shallow ventral and lateral lobe and a siphuncle that is variable in position but never extremely ventral or dorsal.
Cenoceras is not found above the Middle Jurassic and is followed by the Upper Jurassic-Miocene Eutrephoceras.
Eutrephoceras is generally subgobular, broadly rounded laterally and ventrally, with a small to occluded umbilicus,
broadly rounded hyponomic sinus, only slightly sinuous sutures, and a small siphuncle that is variable in position.
Next to appear is the Lower Cretaceous Strionautilus from India and the European ex-USSR, named by Shimankiy
in 1951. Strionautilus is compressed, involute, with fine longitudinal striations. Whorl sections are subrectangular,
sutures sinuous, the siphuncle subcentral.
Also from the Cretaceous is Pseudocenoceras, named by Spath in 1927. Pseudocenoceras is compressed, smooth,
with subrectangular whorl sections, flattened venter, and a deep umbilicus. The suture crosses the venter essentially
straight and has a broad, shallow, lateral lobe. The siphuncle is small and subcentral. Pseudocenoceras is found in
the Crimea and in Libya.
Carinonautilus is a genus from the Upper Cretaceous of India, named by Spengler in 1919. Carinonautilus is a very
involute form with high whorl section and flanks that converge on a narrow venter that bears a prominent rounded
keel. The umbilicus is small and shallow, the suture only slightly sinuous. The siphuncle is unknown.
Taxonomy
The family Nautilidae contains six
extant species and several extinct
species.
• Genus Allonautilus
• A. perforatus
• A. scrobiculatus
• Genus Nautilus
• N. belauensis
• †N. cookanum
• N. macromphalus
• N. pompilius (type)
• N. p. pompilius
• N. p. suluensis
• †N. praepompilius
• N. stenomphalus Shell characters of the genera Allonautilus and Nautilus
Dubious or uncertain taxa

The following taxa associated with the family Nautilidae are of uncertain taxonomic status:[9]
Nautilus 56
Binomial name and author Current systematic status Type locality Type repository
citation
N. alumnus Iredale, 1944 Species dubium [fide Saunders (1987:49)] Queensland, Not designated [fide Saunders
Australia (1987:49)]
N. ambiguus Sowerby, 1848 Species dubium [fide Saunders (1987:48)] Not designated Unresolved
N. beccarii Linne, 1758 Non-cephalopod; Foraminifera [fide Frizzell and Keen

(1949:106)]
N. calcar Linne, 1758 ?Non-cephalopod; Foraminifera Lenticulina Adriatic Sea Unresolved; Linnean Society of
London?
N. crispus Linne, 1758 Undetermined Mediterranean Sea Unresolved; Linnean Society of

London?
N. crista Linne, 1758 Non-cephalopod; Turbo [fide Dodge (1953:14)]
N. fascia Linne, 1758 Undetermined Adriatic Sea Unresolved; Linnean Society of

London?
N. granum Linne, 1758 Undetermined Mediterranean Sea Unresolved; Linnean Society of

London?
N. lacustris Lightfoot, 1786 Non-cephalopod; Helix [fide Dillwyn (1817:339)]
N. legumen Linne, 1758 Undetermined Adriatic Sea Unresolved; Linnean Society of

London?
N. micrombilicatus Joubin, 1888 Nomen nudum
N. obliquus Linne, 1758 Undetermined Adriatic Sea Unresolved; Linnean Society of

London?
N. pompilius marginalis Willey, Species dubium [fide Saunders (1987:50)] New Guinea Unresolved
1896
N. pompilius moretoni Willey, Species dubium [fide Saunders (1987:49)] New Guinea Unresolved
1896
N. pompilius perforatus Willey, Species dubium [fide Saunders (1987:49)] New Guinea Unresolved
1896
N. radicula Linne, 1758 ?Non-cephalopod; Foraminifera Nodosaria Adriatic Sea Unresolved; Linnean Society of
London?
N. raphanistrum Linne, 1758 Undetermined Mediterranean Sea Unresolved; Linnean Society of

London?
N. raphanus Linne, 1758 Undetermined Adriatic Sea Unresolved; Linnean Society of

London?
N. semi-lituus Linne, 1758 Undetermined Liburni, Adriatic Unresolved; Linnean Society of

Sea London?
N. sipunculus Linne, 1758 Undetermined "freto Siculo" Unresolved; Linnean Society of

London?
N. texturatus Gould, 1857 Nomen nudum
Octopodia nautilus Schneider, Rejected specific name [fide Opinion 233, ICZN
1784 (1954:278)]
Nautilus 57
References
[1] Kümmel,B. 1964. Nautiloidae-Nautilida, in the Treatise on Invertebrate Paleontology, Geological Society of America and Univ of Kansas
Press, Teichert and Moore eds.
[2] Wingstrand, KG (1985). "On the anatomy and relationships of Recent Monoplacophora" (http:/ / www. zmuc. dk/ inverweb/ Galathea/
Galathea_p5. html) (Link to free full text + plates). Galathea Rep. 16: 7–94. .
[3] "Diagenesis of aragonite from Upper Cretaceous ammonites: a geochemical case-study". Sedimentology 28: 423–438. 1981.
doi:10.1111/j.1365-3091.1981.tb01691.x.
[4] Willey, Arthur (1897). "The Pre-ocular and Post-ocular Tentacles and Osphradia of Nautilus" (http:/ / jcs. biologists. org/ content/ vols2-40/
issue157/ ). Quarterly Journal of Microscopical Science 40 (1): 197–201. .
[5] Fukuda, Y. 1987. Histology of the long digital tentacles. In: W.B. Saunders & N.H. Landman (eds.) Nautilus: The Biology and Paleobiology
of a Living Fossil. Springer Netherlands. pp. 249–256. doi:10.1007/978-90-481-3299-7_17
[6] Kier, W.M. 1987. The functional morphology of the tentacle musculature of Nautilus pompilius. (http:/ / biology. unc. edu/ faculty/ Kier/ lab/
pdf/ Kier_1987. pdf)PDF In: W.B. Saunders & N.H. Landman (eds.) Nautilus: The Biology and Paleobiology of a Living Fossil. Springer
Netherlands. pp. 257–269. doi:10.1007/978-90-481-3299-7_18
[7] Grasso, F.; Basil, J. (2009). "The evolution of flexible behavioral repertoires in cephalopod molluscs". Brain, Behavior and Evolution 74 (3):
231–245. doi:10.1159/000258669. PMID 20029186.
[8] Saunders WB (June 1984). "Nautilus Growth and Longevity: Evidence from Marked and Recaptured Animals". Science 224 (4652):
990–992. doi:10.1126/science.224.4652.990. PMID 17731999.
[9] Sweeney, M.J. 2002. Taxa Associated with the Family Nautilidae Blainville, 1825. (http:/ / tolweb. org/ accessory/
Nautilidae_Taxa?acc_id=2324) Tree of Life web project.
• Ward, P.D. 1988. In Search of Nautilus. Simon and Schuster.

• Nautilus: the biology and palaeontology of a living fossil (http://www.springerlink.com/content/
978-90-481-3298-0/#section=639864&page=10&locus=96). ISBN 978-90-481-3299-7.
• CephBase: Nautilidae (http://www.cephbase.utmb.edu/spdb/familygroup.cfm?Family=Nautilidae)
External links
• Nautilidae discussion forum (http://www.tonmo.com/forums/forumdisplay.php?f=49), tonmo.com
• Waikïkï Aquarium: Marine Life Profile: Chambered Nautilus (http://www.waquarium.org/MLP/root/pdf/
MarineLife/Invertebrates/Molluscs/Nautilus.pdf), waguarium.org
• A molecular and karyological approach to the taxonomy of Nautilus (http://www.cephbase.utmb.edu/refdb/
pdf/8036.pdf), utmb.edu
Ammonite 58
Ammonite
Ammonites
Artist's reconstruction of
Asteroceras
Kingdom: Animalia
Phylum: Mollusca
Class: Cephalopoda
Subclass: †Ammonoidea
Zittel, 1884
Orders and Suborders
Order Anarcestida
• Anarcestina
• Pharciceratina
• Prolobinina
Order Ammonitida
• Ammonitina
• Ancyloceratina
• Phylloceratina
• Lytoceratina
Order Ceratitida
• Ceratitina
• Otoceratina
• Noritacina
• Clydonitina
Order Clymeniida
• Clymeniina
• Gonioclymeniina
• Cyrtoclymeniina
Order Goniatitida
• Goniatitina
• Tornoceratina
Order Prolecanitida
• Prolecanitina
• Medlicottiana
Ammonites are an extinct group of marine invertebrate animals in the subclass Ammonoidea of the class
Cephalopoda. These molluscs are more closely related to living coleoids (i.e. octopuses, squid, and cuttlefish) than
Ammonite 59
they are to shelled nautiloids such as the living Nautilus species.

Ammonites are excellent index fossils, and it is often possible to link the rock layer in which they are found to
specific geological time periods. Their fossil shells usually take the form of planispirals, although there were some
helically-spiraled and non-spiraled forms (known as heteromorphs).
The name ammonite, from which the scientific term is derived, was inspired by the spiral shape of their fossilized
shells, which somewhat resemble tightly-coiled rams' horns. Pliny the Elder (d. 79 AD. near Pompeii) called fossils
of these animals ammonis cornua ("horns of Ammon") because the Egyptian god Ammon (Amun) was typically
depicted wearing ram's horns.[1] Often the name of an ammonite genus ends in -ceras, which is Greek (κέρας) for
"horn".
Classification
Originating from within the bactritoid nautiloids, the ammonoid cephalopods first appeared in the Devonian (circa
400 million years ago) and became extinct at the close of the Cretaceous (65.5 Ma) along with the dinosaurs. The
classification of ammonoids is based in part on the ornamentation and structure of the septa comprising their shells'
gas chambers; by these and other characteristics we can divide subclass Ammonoidea into three orders and eight
known suborders. While nearly all nautiloids show gently curving sutures, the ammonoid suture line (the intersection
of the septum with the outer shell) was folded, forming saddles (or peaks) and lobes (or valleys).
Suture patterns
Three major types of suture patterns in Ammonoidea have been noted:
• Goniatitic - numerous undivided lobes and saddles; typically 8 lobes around the conch. This pattern is
characteristic of the Paleozoic ammonoids.
• Ceratitic - lobes have subdivided tips, giving them a saw-toothed appearance, and rounded undivided saddles.
This suture pattern is characteristic of Triassic ammonoids and appears again in the Cretaceous "pseudoceratites".
• Ammonitic - lobes and saddles are much subdivided (fluted); subdivisions are usually rounded instead of
saw-toothed. Ammonoids of this type are the most important species from a biostratigraphical point of view. This
suture type is characteristic of Jurassic and Cretaceous ammonoids but extends back all the way to the Permian.
Orders and suborders

The Ammonoidea can be divided into eight orders, listed here starting
with the most primitive and going to the more derived.
• Anarcestida, Devonian
• Clymeniida, Upper Devonian
• Goniatitida, Middle Devonian - Upper Permian
An ammonitic ammonoid with the body chamber
• Prolecanitida, Upper Devonian - Upper Triassic missing, showing the septal surface (especially at
• Ceratitida, Permian - Triassic right) with its undulating lobes and saddles.
• Phylloceratida, Triassic - Cretaceous

• Lytoceratida, Jurassic - Cretaceous
• Ammonitida, Lower Jurassic - Upper Cretaceous
Note that in some classifications these are referred to as suborders, included in only three orders: Goniatitida,
Ceratitida, and Ammonitida.
Ammonite 60
Taxonomy of the Treatise

The Treatise on Invertebrate Paleontology (1964) includes the
Ammonitina, Lytoceratina, and Phylloceratina as separate suborders
within the subclass Ammonoidea, without the use of orders, and
divides them into superfamilies. In other, subsequent taxonomies the
Ammonitina, Lytoceratina, and Phylloceratina are placed within the
order, Ammonitida. The Ancyloceratina which is sometimes treated as
a separate suborder is treated as a superfamily, the Ancylocerataceae in Iridescent ancient ammonite fossil on display at
the Lytoceratina in the Treatise. the American Museum of Natural History, New
York City, around 2.5 feet in diameter.
According to the Treatise, the Ammonitina are derived from the Phyllocerarina and Lytoceratina beginning in the
Early Jurassic with the Psilocerataceae and ending with nine superfamilies, although not all extant at the same time.
These are the Acanthocerataceae, Desmocerataceae, Eoderocerataceae, Haploceratacea, Hildocerataceae,
Hoplitaceae, Perispinctaceae, Psilocerataceae, and Stephanocerataceae.
The Eoderocerataceae, Hildocerataceae, Psilocerataceae, and Stephanocerataceae are strictly Jurassic groups. The
Acanthocerataceae, Desmocerataceae, and Hoplitaceae are known only from the Cretaceous. But the
Haplocerataceae and Peripinctaceae extend from the Jurassic well into the Cretaceous.
Life
Because ammonites and their close relatives are extinct, little is known
about their way of life. Their soft body parts are very rarely preserved
in any detail. Nonetheless, much has been worked out by examining
ammonoid shells and by using models of these shells in water tanks.
Many ammonoids probably lived in the open water of ancient seas,
rather than at the sea bottom. This is suggested by the fact that their
fossils are often found in rocks that were laid down under conditions
where no bottom-dwelling life is found. Many of them (such as
Oxynoticeras) are thought to have been good swimmers with flattened,
discus-shaped, streamlined shells, although some ammonoids were less
effective swimmers and were likely to have been slow-swimming Jeletzkytes, a Cretaceous ammonite from the
bottom-dwellers. Synchrotron analysis of an aptychophoran ammonite USA
revealed remains of isopod and mollusc larva in its buccal cavity,
indicating that at least this kind of ammonite fed on plankton[2] . Fossilized ammonoids have been found showing
tooth marks from such attacks. They may have avoided predation by squirting ink, much like modern cephalopods;
ink is occasionally preserved in fossil specimens.[3]
The soft body of the creature occupied the largest segments of the shell at the end of the coil. The smaller earlier
segments were walled off and the animal could maintain its buoyancy by filling them with gas. Thus the smaller
sections of the coil would have floated above the larger sections.[4]
Ammonite 61
Asteroceras, a Jurassic ammonite from England
Shell anatomy and diversity
Basic shell anatomy

The chambered part of the ammonite shell is called a phragmocone.
The phragmocone contains a series of progressively larger chambers,
called camerae (sing. camera) that are divided by thin walls called
septa (sing. septum). Only the last and largest chamber, the body
chamber, was occupied by the living animal at any given moment. As
it grew, it added newer and larger chambers to the open end of the coil.
A thin living tube called a siphuncle passed through the septa,
extending from the ammonite's body into the empty shell chambers.
Through a hyperosmotic active transport process, the ammonite
emptied water out of these shell chambers. This enabled it to control
the buoyancy of the shell and thereby rise or descend in the water
column.
A primary difference between ammonites and nautiloids is that the

siphuncle of ammonites (excepting Clymeniina) runs along the ventral
periphery of the septa and camerae (i.e., the inner surface of the outer
A variety of ammonite forms, from Ernst
axis of the shell), while the siphuncle of nautiloids runs more or less
Haeckel's 1904 Kunstformen der Natur (Artforms through the center of the septa and camerae.
of Nature).
Ammonite 62
Sexual dimorphism
One feature found in shells of the modern Nautilus is the variation in
the shape and size of the shell according to the sex of the animal, the
shell of the male being slightly smaller and wider than that of the
female. This sexual dimorphism is thought to be an explanation for the
variation in size of certain ammonite shells of the same species, the
larger shell (called a macroconch) being female, and the smaller shell
(called a microconch) being male. This is thought to be because the
female required a larger body size for egg production. A good example
of this sexual variation is found in Bifericeras from the early part of the
Jurassic period of Europe.
It is only in relatively recent years that the sexual variation in the shells
Discoscaphites iris, Owl Creek Formation (Upper
of ammonites has been recognized. The macroconch and microconch
Cretaceous), Ripley, Mississippi.
of one species were often previously mistaken for two closely related
but different species occurring in the same rocks. However, these
"pairs" were so consistently found together that it became apparent that they were in fact sexual forms of the same
species.
Variations in shape
The majority of ammonite species feature a shell that is a planispiral flat coil, but other species feature a shell that is
nearly straight (as in baculites). Still other species' shells are coiled helically, superficially like that of a large
gastropod (as in Turrilites and Bostrychoceras). Some species' shells are even initially uncoiled, then partially coiled,
and finally straight at maturity (as in Australiceras). These partially uncoiled and totally uncoiled forms began to
diversify mainly during the early part of the Cretaceous and are known as heteromorphs.
Perhaps the most extreme and bizarre looking example of a heteromorph is Nipponites, which appears to be a tangle
of irregular whorls lacking any obvious symmetrical coiling. However, upon closer inspection the shell proves to be
a three-dimensional network of connected "U" shapes. Nipponites occurs in rocks of the upper part of the Cretaceous
in Japan and the USA.
Ammonites vary greatly in the ornamentation (surface relief) of their shells. Some may be smooth and relatively
featureless, except for growth lines, and resemble that of the modern Nautilus. In others various patterns of spiral
ridges and ribs or even spines are shown. This type of ornamentation of the shell is especially evident in the later
ammonites of the Cretaceous.
Ammonite 63
Aptychus
Some ammonites have been found in association with a single horny
plate or a pair of calcitic plates. In the past it was assumed that these
plates served to close the opening of the shell in much the same way as
an operculum, however more recently it has been postulated that they
were instead a jaw apparatus.[5] [6] [7] [8]
The plates are collectively termed the aptychus or aptychi in the case
of a pair of plates, and anaptychus in the case of a single plate. The
paired aptychi were symmetrical to one another and equal in size and
appearance.
A drawing of an aptychus named "Trigonellites
Anaptychi are relatively rare as fossils. They are found representing latus" from the Kimmeridge Clay Formation in
ammonites from the Devonian period through those of the Cretaceous England
period.
Calcified aptychi only occur in ammonites from the Mesozoic era. They are almost always found detached from the
shell, and are only very rarely preserved in place. Still, sufficient numbers have been found closing the apertures of
fossil ammonite shells as to leave no doubt as to their identity as part of the anatomy of an ammonite.
Large numbers of detached aptychi occur in certain beds of rock (such as those from the Mesozoic in the Alps).
These rocks are usually accumulated at great depths. The modern Nautilus lacks any calcitic plate for closing its
shell, and only one extinct nautiloid genus is known to have borne anything similar. Nautilus does, however, have a
leathery head shield (the hood) which it uses to cover the opening when it retreats inside.
There are many forms of aptychus, varying in shape and the sculpture of the inner and outer surfaces, but because
they are so rarely found in position within the shell of the ammonite it is often unclear to which species of ammonite
one kind of aptychus belongs. A number of aptychi have been given their own genus and even species names
independent of their unknown owners' genus and species, pending future discovery of verified occurrences within
ammonite shells.
Non-mineralized anatomy
Annomoids bore a radula and beak, a marginal siphuncle, and probably ten arms.[9] They operated by direct
development with sexual reproduction, were carnivorous and had a crop for food storage. It is unlikely that any
ammonoids dwelt in fresh or brackish water.[10]
Soft parts
Although ammonites do occur in exceptional lagerstatten such as the Solnhofen limestone, their soft part record is
surprisingly bleak - beyond a tentative ink sac and possible digestive organs, no soft parts are known at all.[11] It can
be tentatively assumed that they had numerous tentacles, each quite weak, and engulfed prey almost whole.[11]
Ammonite 64
Size
Few of the ammonites occurring in the lower and middle part of the
Jurassic period reach a size exceeding 23 centimetres (9 inches) in
diameter. Much larger forms are found in the later rocks of the upper
part of the Jurassic and the lower part of the Cretaceous, such as
Titanites from the Portland Stone of Jurassic of southern England,
which is often 53 centimetres (2 feet) in diameter, and Parapuzosia
seppenradensis of the Cretaceous period of Germany, which is one of
the largest known ammonites, sometimes reaching 2 metres (6.5 feet)
in diameter. The largest documented North American ammonite is 2-metre (6.5-foot) Parapuzosia seppenradensis
cast in Germany
Parapuzosia bradyi from the Cretaceous with specimens measuring
137 centimetres (4.5 feet) in diameter, although a new 2.3-metre
(7.5-foot) British Columbian specimen, if authentic, would appear to trump even the European champion.[12]
Distribution
Starting from the mid-Devonian, ammonoids were extremely abundant,
especially as ammonites during the Mesozoic era. Many genera
evolved and ran their course quickly, becoming extinct in a few million
years. Due to their rapid evolution and widespread distribution,
ammonoids are used by geologists and paleontologists for
biostratigraphy. They are excellent index fossils, and it is often
possible to link the rock layer in which they are found to specific
geological time periods.
Due to their free-swimming and/or free-floating habits, ammonites

often happened to live directly above seafloor waters so poor in oxygen
A specimen of Hoploscaphites from the Pierre
as to prevent the establishment of animal life on the seafloor. When
Shale of South Dakota. Much of the original upon death the ammonites fell to this seafloor and were gradually
shell, including the nacre, has survived. buried in accumulating sediment, bacterial decomposition of these
corpses often tipped the delicate balance of local redox conditions
sufficiently to lower the local solubility of minerals dissolved in the seawater, notably phosphates and carbonates.
The resulting spontaneous concentric precipitation of minerals around a fossil is called a concretion and is
responsible for the outstanding preservation of many ammonite fossils.
When ammonites are found in clays their original mother-of-pearl coating is often preserved. This type of
preservation is found in ammonites such as Hoplites from the Cretaceous Gault clay of Folkestone in Kent, England.
The Cretaceous Pierre Shale formation of the United States and Canada is well known for the abundant ammonite
fauna it yields, including Baculites, Placenticeras, Scaphites, Hoploscaphites, and Jeletzkytes, as well as many
uncoiled forms. Many of these also have much or all of the original shell, as well as the complete body chamber, still
intact. Many Pierre Shale ammonites, and indeed many ammonites throughout earth history, are found inside
concretions.
Ammonite 65
Other fossils, such as many found in Madagascar and Alberta

(Canada), display iridescence. These iridescent ammonites are often of
gem quality (ammolite) when polished. In no case would this
iridescence have been visible during the animal's life; additional shell
layers covered it.
The majority of ammonoid specimens, especially those of the
Paleozoic era, are preserved only as internal molds; that it to say, the
outer shell (composed of aragonite)[13] has been lost during the
fossilization process. It is only in these internal-mold specimens that
the suture lines can be observed; in life the sutures would have been
hidden by the outer shell.
The ammonoids as a group continued though several major extinction

An iridescent ammonite from Madagascar.
event, although it appears that often only a few species survived. Each
time, however, this handful of species diversified into a multitude of
forms. Ammonite fossils became less abundant during the latter part of the Mesozoic, with none surviving into the
Cenozoic era. The last surviving lineages disappeared, along with the dinosaurs, 65 million years ago in the
Cretaceous-Tertiary extinction event. The reason why no ammonites survived the extinction event at the end of the
Cretaceous, whereas some nautiloid cousins survived, might be due to differences in ontogeny. If their extinction
was due to a bolide strike, plankton around the globe could have been severely diminished, thereby dooming
ammonite reproduction during its planktonic stage.
Extinction
The extinction of the ammonites along with other marine animals and of course, non-avian dinosaurs, has been
attributed to a bolide impact, marking the end of the Cretaceous Period. Regardless of what effect an impact may
have had, many of these groups, including ammonoids, were already in serious decline. Previously ammonoid
cephalopods barely survived several earlier major extinction events, often with only a few species surviving from
which a multitude of forms diversified.
Eight or so species from only two families made it almost to the end of the Cretaceous, the order having gone
through a more or less steady decline since the middle of the period. Six other families made it well into the upper
Maastrichtian (uppermost stage of the Cretaceous) but were extinct well before the end. All told, 11 families entered
the Maastrichtian, a decline from the 19 families known from the Cenomanian in the middle of the Cretaceous.
One reason given for their demise is that Cretaceous ammonites, being closely related to coleoids, had a similar
reproductive strategy in which a huge number of eggs is laid in a single batch at the end of the life span. These, along
with juvenile ammonites, are thought to have been part of the plankton at the surface of the ocean where they were
killed off by the effects of an impact. Nautiloids, exemplified by modern nautiluses, are thought on the other hand to
have had a reproductive strategy in which eggs were laid in smaller batches many times during the life span and on
the sea floor well away from any direct effects of such a bolide strike, and thus survived.
Mythology
In medieval Europe, fossilised ammonites were thought to be petrified coiled snakes, and were called "snakestones"
or, more commonly in medieval England, "serpentstones". They were considered to be evidence for the actions of
saints such as Saint Hilda and Saint Patrick, and were held to have healing or oracular powers. Traders would
occasionally carve the head of a snake onto the empty, wide end of the ammonite fossil, and then sell them to the
public. In other cases the snake's head would be simply painted on.[14] Ammonites from the Gandaki river in Nepal
are known as saligrams, and are believed by Hindus to be a concrete manifestation of God or Vishnu.[15]
Ammonite 66
Terminological note
The words ammonite and ammonoid are both used quite loosely in common parlance to refer to any member of
subclass Ammonoidea. However, in stricter usage the term ammonite is reserved for members of suborder
Ammonitina (or sometimes even order Ammonitida).
References
[1] NH 37.40.167
[2] The Role of Ammonites in the Mesozoic Marine Food Web Revealed by Jaw Preservation, Isabelle Kruta, Neil Landman, Isabelle Rouget,
Fabrizio Cecca, Paul Tafforeau, SCIENCE, JANUARY 2011 VOL 331
[3] doi: 10.1007/978-1-4020-6806-5_11
wiki/ Template:cite_doi/ . 0910. 1007. 2f978-1-4020-6806-5_11?preload=Template:Cite_doi/ preload& editintro=Template:Cite_doi/
[4] "Introduction to Ammonoidea" (http:/ / www. bbm. me. uk/ portsdown/ PH_232_Ammonites. htm). The Geology of Portsdown Hill. .
Retrieved 2007-04-26.
[5] Morton, N. 1981. Aptychi: the myth of the ammonite operculum. Lethaia 14(1): 57–61. doi:10.1111/j.1502-3931.1981.tb01074.x
[6] Morton, N. & M. Nixon 1987. Size and function of ammonite aptychi in comparison with buccal masses of modem cephalopods. Lethaia
20(3): 231–238. doi:10.1111/j.1502-3931.1987.tb02043.x
[7] Lehmann, U. & C. Kulicki 1990. Double function of aptychi (Ammonoidea) as jaw elements and opercula. Lethaia 23: 325–331.
doi:10.1111/j.1502-3931.1990.tb01365.x
[8] Seilacher, A. 1993. Ammonite aptychi; how to transform a jaw into an operculum? American Journal of Science 293: 20–32.
doi:10.2475/ajs.293.A.20
[9] Landman, Neil H; Tanabe, Kazushige; Davis, Richard Arnold (1996). Ammonoid paleobiology (http:/ / books. google. ca/
books?id=hKbkB4MzUIkC& pg=PA14). ISBN 9780306452222. .
[10] Landman, Neil H; Tanabe, Kazushige; Davis, Richard Arnold (1996). Ammonoid paleobiology (http:/ / books. google. ca/
books?id=hKbkB4MzUIkC& pg=PA17). ISBN 9780306452222. .
[11] Wippich, M. G. E.; Lehmann, J. (2004). "Allocrioceras from the Cenomanian (mid-Cretaceous) of the Lebanon and its bearing on the
palaeobiological interpretation of heteromorphic ammonites". Palaeontology 47: 1093–1107. doi:10.1111/j.0031-0239.2004.00408.x.
[12] "Ammonites" (http:/ / web. archive. org/ web/ 20030210130400/ http:/ / www. hanmansfossils. com/ catalogs/ fossils/ ammonites/
ammonites. shtml). Hanman's Fossil Replicas and Minerals. Archived from the original (http:/ / www. hanmansfossils. com/ catalogs/ fossils/
ammonites/ ammonites. shtml) on 2003-02-10. .
[13] "Diagenesis of aragonite from Upper Cretaceous ammonites: a geochemical case-study". Sedimentology 28: 423–438. 1981.
doi:10.1111/j.1365-3091.1981.tb01691.x.
[14] Cadbury, D. the Dinosaur Hunters. (Fourth Estate, 2000) (ISBN 1-85702-963-1), p.7
[15] "Fossils: myths, mystery, and magic" (http:/ / news. independent. co. uk/ sci_tech/ article2259490. ece). The Independent (London).
2007-02-12. . Retrieved 2010-04-23.
• Neal L. Larson, Steven D Jorgensen, Robert A Farrar and Peter L Larson. Ammonites and the other Cephalopods
of the Pierre Seaway. Geoscience Press, 1997.
• Lehmann, Ulrich. The Ammonites: Their life and their world. Cambridge University Press, New York, 1981.
Translated from German by Janine Lettau.
• Monks, Neale and Palmer, Phil. Ammonites. Natural History Museum, 2002.
• Walker, Cyril and Ward, David. Fossils. Dorling, Kindersley Limited, London, 2002.
• A Broad Brush History of the Cephalopoda (http://is.dal.ca/~ceph/TCP/evolution.html) by Dr. Neale Monks,
from The Cephalopod Page.
• Ammonite maturity, pathology and old age (http://is.dal.ca/~ceph/TCP/ammonage.html) By Dr. Neale
Monks, from The Cephalopod Page. Essay about the life span of Ammonites.
• Cretaceous Fossils Taxonomic Index for Order Ammonoitida (http://www.cretaceousfossils.com/invertebrates/
ammonites/ammonites_index.htm)
• Deeply Buried Sediments Tell Story of Sudden Mass Extinction (http://www.nsf.gov/discoveries/disc_summ.
jsp?cntn_id=100280&org=NSF)
Ammonite 67
External links
• Descriptions and pictures of ammonite fossils (http://www.fossilmuseum.net/Fossil_Galleries/Ammonites.
htm)
• goniat.org, a palaezoic ammonoid database (http://www.goniat.org/)
• paleozoic.org: gallery of ammonite photographs (http://www.paleozoic.org/gallery.htm)
• photos of ammonites at Lyme Regis, UK (http://y2u.co.uk/&002_Images/Lyme Regis Fossils 01.htm)
• TaxonConcept's data on cretaceous ammonites (http://taxonconcept.stratigraphy.net/source_main.
php?doctaxid=618&doctaxname=Ammonoidea)
• The ammonites of Peacehaven - photos of giant cretaceous ammonites in Southern England (http://
thinkingwithpictures.blogspot.com/2008/06/ammonites-of-peacehaven.html)
• tonmo.com: The octopus news magazine online (http://www.tonmo.com/science/fossils/fossilsjump.php),
Cephalopod fossil articles.
Belemnoidea
Belemnites (or belemnoids) are an extinct group of marine cephalopod, very similar in many ways to the modern
squid and closely related[1] to the modern cuttlefish. Like them, the belemnites possessed an ink sac[2] , but, unlike
the squid, they possessed ten arms of roughly equal length, and no tentacles.[3] The name "belemnoid" comes from
the Greek word belemnon meaning "a dart or arrow" and the Greek word eidos meaning "form". [4]
Occurrence
Belemnites were numerous during the Jurassic and Cretaceous periods, and their fossils are abundant in Mesozoic
marine rocks, often accompanying their cousins the ammonites. The belemnites become extinct at the end of the
Cretaceous period along with the ammonites. The belemnites' origin lies within the bactritoid nautiloids, which date
from the Devonian period; well-formed belemnite guards can be found in rocks dating from the Mississippian (or
Early Carboniferous) onward through the Cretaceous. Other fossil cephalopods include baculites, nautiloids and
goniatites.
Anatomy
Belemnites possessed a central phragmocone made of aragonite and with negative buoyancy.[5] To the rear of the
creature was a heavy calcite guard whose main role appears to have been to counterbalance the front of the
organism; it positions the centre of mass below the centre of buoyancy, increasing the stability of the swimming
organism.[5] The guard would account for between a third and a fifth of the length of the complete organism, arms
included.[5]
Like some modern squid, belemnite arms carried a series of small hooks for grabbing prey. Belemnites were efficient
carnivores that caught small fish and other marine animals with their arms and ate them with their beak-like jaws. In
turn, belemnites appear to have formed part of the diet of marine reptiles such as Ichthyosaurs, whose fossilized
stomachs frequently contain phosphatic hooks from the arms of cephalopods.
Belemnoidea 68
Ecology
Belemnites were effectively neutrally buoyant, and swam in near-shore to mid-shelf oceans.[5] Their fins could be
used to their advantage in all water speeds; in a gentle current they could be flapped for propulsion; in a stronger
current they could be held erect to generate lift; and when swimming rapidly by jet propulsion they could be tucked
in to the body for streamlining.[5]
Preservation
Normally with fossil belemnites only the back part of the shell (called
the guard or rostrum) is found. The guard is elongated and
bullet-shaped, that is to say, cylindrical and pointed or rounded at one
end. The hollow region at the front of the guard is termed the alveolus,
and this houses a chambered conical-shaped part of the shell (called the
phragmocone). The phragmocone is usually only found with the better
A belemnite fossil from the Franconian Jura. preserved specimens. Projecting forwards from one side of the
phragmocone is the thin pro-ostracum.
While belemnite phragmocones are homologous with the shells of other cephalopods and are similarly composed of
aragonite, belemnite guards are evolutionarily novel and are composed of calcite, thus tending to preserve well.
Broken guards show a structure of radiating calcite fibers and may also display concentric growth rings.
The guard, phragmocone and pro-ostracum were all internal to the living creature, forming a skeleton which was
enclosed entirely by soft muscular tissue. The original living creature would have been larger than the fossilized
shell, with a long streamlined body and prominent eyes. The guard would have been in place toward the rear of the
creature, with the phragmocone behind the head and the pointed end of the guard facing backward.
The guard of the belemnite Megateuthis gigantea, which is found in
Europe and Asia, can measure up to 46 cm in length (18 inches),
giving the living animal an estimated length of 3 metres (10 feet).
Belemnites
Belemnoidea 69
Very exceptional belemnite specimens have been found showing the

preserved soft parts of the animal. Elsewhere in the fossil record,
bullet-shaped belemnite guards are locally found in such profusion that
such deposits are referred to semi-formally as "belemnite battlefields"
(cf. "orthocone orgies"). It remains unclear whether these deposits
represent post-mating mass death events, as are common among
modern cephalopods and other semelparous creatures.
Uses
Belemnite fossils at Bristol City Museum, Bristol,
The bulk geochemical signature contained within belemnite guards of England. Found in the Lower Lias strata,
Gloucestershire, England.
the Peedee Formation (Cretaceous, southeast USA) has long been used
as a global standard ("PDB") against which all other geochemical
samples are measured, for both carbon isotopes and oxygen isotopes.
Some belemnites (such as Belemnites) serve as index fossils, particularly in the Cretaceous Chalk Formation of
Europe, enabling geologists to date the age the rocks in which they are found.
Classification
Note: all families extinct
• Cohort Belemnoidea
• Basal and unresolved
• Genus Jeletzkya
• Genus Belemnoteuthis
• Order Aulacocerida
• Family Aulacoceratidae
• Family Dictyoconitidae
• Family Hematitidae
• Family Palaeobelemnopseidae Belemnite in the very top bedding plane of the
Zohar Formation (Jurassic) near Neve Atif, the
• Family Xiphoteuthididae
Golan. Note the central fold along the axis
• Order Belemnitida characteristic of some genera.
• Suborder Belemnitina
• Family Cylindroteuthididae
• Family Hastitidae
• Family Oxyteuthididae
• Family Passaloteuthididae
• Family Salpingoteuthididae
• Suborder Belemnopseina
• Family Belemnitellidae Fossilised belemnite
• Family Belemnopseidae
• Family Dicoelitidae
• Family Dimitobelidae
• Family Duvaliidae
• Suborder Belemnotheutina
Belemnoidea 70
• Family Belemnotheutididae
• Family Chitinobelidae
• Family Sueviteuthididae
• Order Diplobelida
• Family Chondroteuthididae
• Family Diplobelidae
• Order Phragmoteuthida
• Family Phragmoteuthididae
• Family Rhiphaeoteuthidae
See also
• Nautiloidea
• Ammonoidea
• List of belemnites
References
[1] Yancey, T. E.; Garvie, C. L.; Wicksten, M. (2010). "The Middle Eocene Belosaepia ungula (Cephalopoda: Coleoida) from Texas: Structure,
Ontogeny and Function" (http:/ / www. lakeneosho. org/ Belosaepia/ pdf/ Growth and skeleton characters of Belosaepia. pdf). Journal of
Paleontology 84: 267. doi:10.1666/09-018R.1. .
[2] Lehmann, U. 1981. The Ammonites: Their life and their world. London: Cambridge University Press.
[3] Doyle, P.; Shakides, E. V. (2004). "The Jurassic Belemnite Suborder Belemnotheutina". Palaeontology 47: 983–998.
doi:10.1111/j.0031-0239.2004.00395.x.
[4] Webster's New Universal Unabridged Dictionary. 2nd ed. 1979.
[5] "The function of the belemnite guard" (http:/ / www. springerlink. com/ content/ p0pw7246ht1v78n1/ ). Paläontologische Zeitschrift 70:
425–431. 1996. doi:10.1007/BF02988082 (inactive 2010-05-02). .
External links
• TONMO.com Cephalopod Fossils articles and discussion forums (http://www.tonmo.com/science/fossils/
fossilsjump.php)
Argonaut 71
Argonaut
For other uses, see Argonaut.
Argonauts
Female Argonauta argo with eggs

Scientific classification
Kingdom: Animalia
Phylum: Mollusca
Class: Cephalopoda
Order: Octopoda
Superfamily: Argonautoida
Family: Argonautidae
Genus: Argonauta
Linnaeus, 1758
Species
†Argonauta absyrtus
Argonauta argo (type)
Argonauta bottgeri
Argonauta cornuta*
Argonauta hians
†Argonauta itoigawai
†Argonauta joanneus
Argonauta nodosa
Argonauta nouryi
Argonauta pacifica*
†Argonauta tokunagai
*Species status questionable.
Synonyms
• Argonautarius
Dumeril, 1806
• Todarus nom. nud.
Rafinesque, 1815
• Todarus
Rafinesque, 1840
• Trichocephalus
Chiaje, 1827 in 1823-1831
Argonaut 72
The argonauts (genus Argonauta, the only extant genus in the Argonautidae family) are a group of pelagic
octopuses. They are also called paper nautiluses, referring to the paper-thin eggcase that females secrete. This
structure lacks the gas-filled chambers present in chambered nautilus shells and is not a true cephalopod shell, but
rather an evolutionary innovation unique to the genus Argonauta.[1] It is used as a brood chamber and for trapped
surface air to maintain buoyancy.
Argonauts are found in tropical and subtropical waters worldwide; they live in the open ocean, i.e. they are pelagic.
Like most octopuses, they have a rounded body, eight arms and no fins. However, unlike most octopuses, argonauts
live close to the sea surface rather than on the seabed. Argonauta species are characterised by very large eyes and
small distal webs. The mantle-funnel locking apparatus is a major diagnostic feature of this taxon. It consists of
knob-like cartilages in the mantle and corresponding depressions in the funnel. Unlike the closely allied genera
Ocythoe and Tremoctopus, Argonauta species lack water pores.
Of its names, "argonaut" means "sailor on the Argo"; "nautilus" is derived from the Greek ναυτίλος, meaning
"sailor", because it was formerly supposed that Argonauta used their shell-secreting arms as sails when they were at
the surface.
The chambered nautilus was later named after the argonaut, but belongs to a different order, the Nautilida.
Physical description
Sexual dimorphism and reproduction

Argonauts exhibit extreme sexual dimorphism in size and lifespan. Females grow up to 10 cm and make shells up to
30 cm, while males rarely surpass 2 cm. The males only mate once in their short lifetime, whereas the females are
iteroparous, capable of having offspring many times over the course of their lives. In addition, the females have been
known since ancient times, while the males were only described in the late 19th century.
The males lack the dorsal tentacles used by the females to create their eggcases. The males use a modified arm, the
hectocotylus, to transfer sperm to the female. For fertilization, the arm is inserted into the female's pallial cavity, then
is detached from the male. The hectocotylus when found in females was originally described as a parasitic worm.[2]
Mature female A. Juvenile female A. hians Immature male A. hians

nodosa
Eggcase
Female argonauts produce a laterally-compressed calcareous eggcase in which they reside. This "shell" has a double
keel fringed by two rows of alternating tubercles. The sides are ribbed with the centre either flat or having winged
protrusions. The eggcase curiously resembles the shells of extinct ammonites. It is secreted by the tips of the female's
two greatly expanded dorsal tentacles (third left arms) before egg laying. After she deposits her eggs in the floating
eggcase, the female takes shelter in it, often retaining the male's detached hectocotylus. She is usually found with her
head and tentacles protruding from the opening, but she retreats deeper inside if disturbed. These ornate curved white
eggcases are occasionally found floating on the sea, sometimes with the female argonaut clinging to it. It is not made
Argonaut 73
of aragonite as most other shells are, but of calcite, with a three-layered structure[3] and a higher proportion of
magnesium carbonate (7%) than other cephalopod shells.[4]
The eggcase contains a bubble of air that the animal captures at the surface of the water and uses for buoyancy, in a
manner similar to other shelled cephalopods, although it does not have a chambered phragmocone as do other
shelled cephalopods.[3] Once thought to contribute to occasional mass strandings on beaches, the air bubble is under
sophisticated control, evident from the behaviour of animals from which air has been removed under experimental
diving conditions.[5] [6] [7]
Most other octopuses lay eggs in caves; Neale Monks and C. Phil Palmer speculate that, before ammonites died out
during the Cretaceous–Tertiary extinction event, the argonauts may have evolved to use discarded ammonite shells
for their egg laying, eventually becoming able to mend the shells and perhaps make their own shells.[8] However,
this is uncertain and it is unknown whether this is the result of convergent evolution.
Argonauta argo is the largest species in the genus and also produces the largest eggcase, which may reach a length
of 300 mm.[9] [10] The smallest species is Argonauta bottgeri, with a maximum recorded size of 67 mm.[11] [12]
Female The eggcase of A. The eggcase The eggcase of A. hians

A. nodosa argo of A.
with its nodosa
eggcase
Beak
The beaks of Argonauta species are distinctive, being characterised by a very small rostrum and a fold that runs to
the lower edge or near the free corner. The rostrum is 'pinched in' at the sides, making it much narrower than in other
octopuses, with the exception of the closely allied monotypic genera Ocythoe and Vitreledonella. The jaw angle is
curved and indistinct. Beaks have a sharp shoulder, which may or may not have posterior and anterior parts at
different slopes. The hood lacks a notch and is very broad, flat, and low. The hood to crest ratio (f/g) is
approximately 2-2.4. The lateral wall of the beak has no notch near the wide crest. Argonaut beaks are most similar
to those of Ocythoe tuberculata and Vitreledonella richardi, but differ in 'leaning back' to a greater degree than the
former and having a more curved jaw angle than the latter.[12]
Feeding and defense

Feeding mostly occurs during the day. Argonauts use tentacles to grab prey and drag it toward the mouth. It then
bites the prey to inject it with poison from the salivary gland. They feed on small crustaceans, molluscs, jellyfish and
salps. If the prey is shelled, the argonaut uses its radula to drill into the organism, then inject the poison.
Argonauts are capable of altering their color. They can blend in with their surroundings to avoid predators. They also
produce ink, which is ejected when the animal is being attacked. This ink paralyzes the olfaction of the attacker,
providing time for the argonaut to escape. The female is also able to pull back the web covering of her shell, making
a silvery flash, which may deter a predator from attacking.
Argonauts are preyed upon by tunas, billfishes, and dolphins. Shells and remains of argonauts have been recorded
from the stomachs of Alepisaurus ferox and Coryphaena hippurus.[12]
Male argonauts have been observed residing inside salps, although little is known about this relationship.[13]
Argonaut 74
Classification
The genus Argonauta contains up to seven extant
species. Several extinct species are also known.
†Argonauta absyrtus
Argonauta argo (type)
Argonauta bottgeri
Argonauta cornuta*
Argonauta hians
†Argonauta itoigawai
†Argonauta joanneus
Argonauta nodosa
Argonauta nouryi
Argonauta pacifica*
†Argonauta tokunagai
*Species status questionable.

The extinct species Obinautilus awaensis was
originally assigned to Argonauta, but has since been
transferred to the genus Obinautilus.[14]
Shells of various Argonauta species
Dubious or uncertain taxa

The following taxa associated with the family Argonautidae are of uncertain taxonomic status:[15]
Binomial name and author Current systematic status Type locality Type repository
citation
Argonauta arctica Fabricius, Undetermined Unresolved; ?Tullukaurfak, Unresolved

1780 Greenland
Argonauta bibula Röding, Undetermined Unresolved Unresolved

1798
Argonauta compressa Undetermined Mer de Indes Unresolved; [other Blainville types at

Blainville, 1826 MNHN] [not reported by Lu et al.
(1995)]
Argonauta conradi Species of uncertain status [fide "New Nantucket, Pacific Ocean" Unresolved
Parkinson, 1856 Robson (1932:200)]
Argonauta cornu Gmelin, Undetermined Unresolved Unresolved; LS?

1791
Argonauta cymbium Linné, Non-cephalopod; foraminiferous

1758 shell [fide Von Martens (1867:103)
Argonauta fragilis Parkinson, Species of uncertain status [fide Not designated Unresolved
1856 Robson (1932:200)]
Argonauta geniculata Gould, Species of uncertain status [fide Near Sugarloaf Mountain, Rio de Type not extant [fide Johnson
1852 Robson (1932:200)] Janeiro, Brazil (1964:32)]
Argonauta maxima Dall, Nomen nudum

1871
Argonaut 75
Argonauta navicula Species dubium [fide Rehder Not designated Unresolved

Lightfoot, 1786 (1967:11)]
Argonauta rotunda Perry, Non-cephalopod; Carcinaria sp.

1811 [fide Robson (1932:201)]
Argonauta rufa Owen, 1836 Incertae sedis [fide Robson "Indian seas" ["South Pacific ocean" Unresolved; Museum of the Royal
(1932:181)] fide Owen (1842:114)] College of Surgeons? Holotype
Argonauta sulcata Lamarck, Nomen nudum

1801
Argonauta tuberculata f. Undetermined Unresolved ZMB

aurita Von Martens, 1867
Argonauta tuberculata f. Undetermined Coast of Brazil ZMB Holotype

mutica Von Martens, 1867
Argonauta tuberculata f. Undetermined Not designated ZMB Syntypes

obtusangula Von Martens,
1867
Argonauta vitreus Gmelin, Undetermined Not designated Unresolved; LS?

1791
Octopus (Ocythoe) Undetermined [Argonauta?] Not designated MNHN Holotype; specimen not
raricyathus Blainville, 1826 extant [fide Lu et al. (1995:323)]
Ocythoe punctata Say, 1819 Argonauta sp. [fide Robson Atlantic Ocean near the North Unresolved; ANSP? Holotype [not
(1929d:215)] American coast (from stomach of traced by Spamer and Bogan (1992)]
dolphin)
Tremoctopus hirondellei Argonauta or Ocythoe [fide Thomas 44°28′56″N 46°48′15″W (Atlantic MOM Holotype [station 151] [fide
Joubin, 1895 (1977:386)] Ocean) Belloc (1950:3)]
Argonaut 76
In design
The argonaut was inspiration for a number of classical and modern art and decorative forms including use on pottery
and architectural elements. Some early examples are found in Minoan art from Crete.[16] A variation known as the
double argonaut design was also found in Minoan jewelry.[17]
In literature and etymology

.
• Argonauts are featured in Twenty Thousand Leagues Under the Sea,
noted for their ability to use their tentacles as sails. There is no
evidence for this.
• A female argonaut is also described in Marianne Moore's poem
"The Paper Nautilus."
• "Argonauta" is the name of a chapter in Anne Morrow Lindbergh's
Gift from the Sea.
• Paper nautiluses were caught in the The Swiss Family Robinson
novel.[18]
• Argonauts gave their name to an Arabidopsis thaliana mutation and
by extension to Argonaute proteins.
References
[1] (German) Naef, A. (1923). "Die Cephalopoden, Systematik". Fauna Flora Golf.
Napoli (35) 1: 1–863. Argonauts surrounding the Nautilus, in Jules
[2] (Italian) Delle Chiaje, S. (1825). Memorie sulla storia e notomia degli animali. Verne's novel Twenty Thousand Leagues Under
Senza Vertebre del Regno di Napoli. I. the Sea
[3] Nixon, M. & J.Z. Young (2003). The Brains and Lives of Cephalopods. Oxford
University Press.
[4] Saul, L. & C. Stadum (2005). "Fossil Argonauts (Mollusca: Cephalopoda: Octopodida) From Late Miocene Siltstones Of The Los Angeles
Basin, California" (http:/ / apt. allenpress. com/ aptonline/ ?request=get-abstract& issn=0022-3360& volume=079& issue=03& page=0520).
Journal of Paleontology 79 (3): 520–531. doi:10.1666/0022-3360(2005)079<0520:FAMCOF>2.0.CO;2. .
[5] Finn, J.K. & M.D. Norman 2010. The argonaut shell: gas-mediated buoyancy control in a pelagic octopus. Proceedings of the Royal Society
B: Biological Sciences, published online May 19, 2010. doi:10.1098/rspb.2010.0155
[6] " Museum Victoria 'Argonaut buoyancy' video" (http:/ / museumvictoria. com. au/ about/ mv-news/ 2010/ argonaut-buoyancy/ )
museumvictoria.com.au. URL accessed on 19 May 2010.
[7] Pidcock, R. 2010. Ancient octopus mystery resolved (http:/ / news. bbc. co. uk/ 1/ hi/ science_and_environment/ 10127611. stm). BBC News,
May 19, 2010.
[8] Monks, N. & P. Palmer (2002). Ammonites. Smithsonian Institution Press, Washington D.C..
[9] Pisor, D. L. (2005). Registry of World Record Size Shells (4th ed.). Snail's Pace Productions and ConchBooks. pp. 12.
[10] (Russian) Nesis, K. N. 1982. Abridged key to the cephalopod mollusks of the world's ocean. Light and Food Industry Publishing House,
Moscow, 385+ii pp. [Translated into English by B. S. Levitov, ed. by L. A. Burgess (1987), Cephalopods of the world. T. F. H. Publications,
Neptune City, NJ, 351 pp.]
[11] Pisor, D. L. (2005). Registry of World Record Size Shells (4th ed.). Snail's Pace Productions and ConchBooks. pp. 12.
[12] Clarke, M. R. (1986). A Handbook for the Identification of Cephalopod Beaks. Oxford University Press. pp. 273 pp.
[13] Banas, P. T., D. E. Smith & D. C. Biggs (1982). "An association between a pelagic octopod, Argonauta sp. Linnaeus 1758, and aggregate
salps". Fish. Bull. U.S. 80: 648–650.
[14] Martill, D.M. & M.J. Barker (2006). A paper nautilus (Octopoda, Argonauta) from the Miocene Pakhna Formation of Cyprus. (http:/ / www.
blackwell-synergy. com/ doi/ abs/ 10. 1111/ j. 1475-4983. 2006. 00578. x?journalCode=pala) Palaeontology 49 (5): 1035-1041.
[15] Sweeney, M. J. Taxa Associated with the Family Argonautidae Tryon, 1879. (http:/ / www. tolweb. org/ accessory/
Argonautidae_Taxa?acc_id=2464) Tree of Life web project.
[16] Eleni M. Konstantinidi, Jewellery Revealed in the Burial Contexts of the Greek Bronze Age, 2001, Hadrian Books, 322 pages ISBN
1841711659
Argonaut 77
[17] C.Michael Hogan, Knossos Fieldnotes, The Modern Antiquarian (2007) (http:/ / themodernantiquarian. com/ site/ 10857/ phaistos.
html#fieldnotes)
[18] Johann David Wyss and Jenny H. Stickney, The Swiss Family Robinson, Ginn & Co., 1898, 364 pages
External links
• CephBase: Argonauta (http://www.cephbase.utmb.edu/spdb/genusgroup.cfm?Genus=Argonauta)
• Tree of Life web project: Argonauta (http://tolweb.org/tree?group=Argonauta)
78
Anatomy and behaviour
Cephalopod intelligence
Cephalopod intelligence has an important comparative aspect in the
understanding of intelligence, because it relies on a nervous system
fundamentally different from that of vertebrates.[1] The cephalopod
class of molluscs, particularly the Coleoidea subclass (cuttlefish, squid
and octopuses), are considered the most intelligent invertebrates and an
important example of advanced cognitive evolution in animals.
The scope of cephalopod intelligence is controversial, complicated by

the challenges of studying these elusive and fundamentally different
creatures. Classical conditioning of cephalopods has been reported, and An octopus in a zoo
one study (Fiorito and Scotto, 1992) even concluded that octopuses
practice observational learning.[2] However, the latter idea is strongly disputed, and doubt has been shed on some
other reported capabilities as well.[3] In any case, impressive spatial learning capacity, navigational abilities, and
predatory techniques remain beyond question.
Examples of intelligence
Predation techniques
Unlike most other molluscs, all cephalopods are active predators (with the possible exception of the bigfin squid).
Their requirement to locate and capture their prey has been a probable driving force behind the development of their
intelligence, uniquely advanced in their phylum.
The humboldt squid hunts schools of fish, showing extraordinary cooperation and communication in its hunting
techniques. This is the first observation of such behaviour in invertebrates.[4]
Crabs, the staple food source of most octopus species, present significant challenges with their powerful pincers and
their potential to exhaust the cephalopod's respiration system from a prolonged pursuit. In the face of these
challenges, octopuses will instead seek out lobster traps and steal the prize inside. They are also known to climb
aboard fishing boats and hide in the containers that hold dead or dying crabs.[5]
Dexterity
Dexterity, a trait essential for tool use and manipulation is also found in cephalopods. The highly sensitive suction
cups and prehensile arms of octopuses, squid, and cuttlefish are as effective at holding and manipulating objects as
the human hand. However, unlike vertebrates, the motor skills of octopuses do not seem to depend upon mapping
their body within their brains, as the ability to organize complex movements is not thought to be linked to particular
arms.[6]
One particularly clever octopus called Otto has been known to juggle his fellow tankmates around out of boredom,
as well as throwing rocks and smashing the aquarium glass. On more than one occasion he even caused short circuits
by crawling out of his tank and shooting a jet of water at the overhead lamp.[7]
Octopus opening a container with a screw cap
Communication
Another example of cephalopod intelligence is the communication that takes place between the more social species
of squid. Some cephalopods are capable of rapid changes in skin color and pattern through nervous control of
chromatophores.[8] This ability almost certainly evolved primarily for camouflage, but squids use color, patterns, and
flashing to communicate with one another in various courtship rituals. Caribbean Reef Squid can send one message
via color patterns to a squid on their right, while they send another message to a squid on their left.[9] [10]
Tool use
As of 2009, the octopus is the only invertebrate animal which has been conclusively shown to use tools. At least four
specimens of the Veined Octopus (Amphioctopus marginatus) have been witnessed retrieving discarded coconut
shells, manipulating them, transporting them some distance, and then reassembling them to use as a shelter. This
discovery was documented in the journal Current Biology and has been filmed on video.[11] [12] Most hermit crabs
use discarded shells of other species for habitation and other crabs choose sea anemones to cultivate on their
carapaces as camouflage; numerous insects use rocks, sand, leaves and so on as building materials, however none of
this behavior compares to the complexity of the octopus's fortress behavior, which involves picking up and carrying
a tool to use later on.
See also
• Paul the Octopus
• Paul II (octopus)
References
[1] "Cephalopod intelligence" (http:/ / www. daviddarling. info/ encyclopedia/ C/ cephalopodintel. html) in The Encyclopedia of Astrobiology,
Astronomy, and Spaceflight.
[2] "What is this octopus thinking?" (http:/ / www. fortunecity. com/ emachines/ e11/ 86/ cephpod. html) by Garry Hamilton
[3] Is the octopus really the invertebrate intellect of the sea? (http:/ / www. nwf. org/ nationalwildlife/ article. cfm?articleId=604& issueId=53)
by Doug Stewart. In: National Wildlife. Feb/Mar 1997, vol.35 no.2
[4] Behold the Humboldt squid (http:/ / outside. away. com/ outside/ features/ 200607/ sea-of-cortez-humboldt-squid-1. html). Tim Zimmermann,
Outside Magazine, July 2006.
[5] Cousteau, Jacques Yves (1978). Octopus and Squid: The Soft Intelligence
[6] Zullo L, Sumbre G, Agnisola C, Flash T, Hochner B. (2009). Nonsomatotopic organization of the higher motor centers in octopus. Curr Biol.
19(19):1632-6. PMID 19765993
[7] "Otto the Octopus wreaks havoc" (http:/ / www. telegraph. co. uk/ news/ newstopics/ howaboutthat/ 3328480/
Otto-the-octopus-wrecks-havoc. html)
[8] Cloney, R.A. & E. Florey 1968. Ultrastructure of cephalopod chromatophore organs. Z Zellforsch Mikrosk Anat 89: 250-280. PMID 5700268
[9] "''Sepioteuthis sepioidea'', Caribbean Reef squid" (http:/ / www. thecephalopodpage. org/ Ssepioidea. php). The Cephalopod Page. . Retrieved
2010-01-20.
[10] Byrne, R.A., U. Griebel, J.B. Wood & J.A. Mather 2003. Squids say it with skin: a graphic model for skin displays in Caribbean Reef Squid.
(http:/ / userpage. fu-berlin. de/ ~jevers/ data/ palbio3/ 05. pdf)PDF (3.86 MB) Berliner Geowissenschaftliche Abhandlungen 3: 29-35.
[11] Morelle, Rebecca (2009-12-14). "Octopus snatches coconut and runs" (http:/ / news. bbc. co. uk/ 1/ hi/ sci/ tech/ 8408233. stm). BBC News.
. Retrieved 2010-01-20.
[12] "Coconut shelter: evidence of tool use by octopuses | EduTube Educational Videos" (http:/ / www. edutube. org/ video/
coconut-shelter-evidence-tool-use-octopuses). Edutube.org. 2009-12-14. . Retrieved 2010-01-20.
Further reading
• What behavior can we expect of octopuses? (http://www.thecephalopodpage.org/behavior.php) by Dr.
Jennifer Mather, Department of Psychology and Neuroscience, University of Lethbridge and Roland C.
Anderson, The Seattle Aquarium.
• Is the octopus really the invertebrate intellect of the sea? (http://www.nwf.org/nationalwildlife/article.
cfm?articleId=604&issueId=53) by Doug Stewart. In: National Wildlife. Feb/Mar 1997, vol.35 no.2.
• Giant Octopus — Mighty but Secretive Denizen of the Deep (http://nationalzoo.si.edu/Support/AdoptSpecies/
AnimalInfo/GiantOctopus/default.cfm) from the Smithsonian National Zoological Park
• Living Fossils Have Long- And Short-term Memory Despite Lacking Brain Structures Of Modern Cephalopods
(http://www.sciencedaily.com/releases/2008/05/080531074905.htm)
• M.J. Wells (1962). Brain and Behaviour in Cephalopods. Heinemann.
• Roger T. Hanlon & John B. Messenger (1996). Cephalopod Behaviour. Cambridge University Press.
• Marion Nixon and John Z. Young (2003). The Brains and Livees of Cephalopods. Oxford University Press.
• Binyamin Hochner; Tal Shomrat & Graziano Fiorito (June 1, 2006). "The Octopus: A Model for a Comparative
Analysis of the Evolution of Learning and Memory Mechanisms" (http://www.biolbull.org/cgi/content/full/
210/3/308). The Biol. Bull. 210 (210): 308–817. PMID 16801504.
• Octopuses are Smart Suckers!? (http://www.thecephalopodpage.org/smarts.php) By Dr. Jennifer Mather,
Department of Psychology and Neuroscience, University of Lethbridge and Roland C. Anderson, The Seattle
Aquarium
• Through the Eye of an Octopus (http://discovermagazine.com/2003/oct/feateye), by Eric Scigliano, Discover
Magazine, October 1, 2003.
Cephalopod size 81
Cephalopod size
Size has been one of the most interesting
aspects of cephalopod science to the general
public. This article lists the largest
cephalopods from various groups, sorted in
order of mantle length, total length, weight,
and shell diameter. Extinct taxa are also
included.
The giant squid (Architeuthis sp.) was for a long time thought to be the largest
extant cephalopod. It is now known that the Colossal Squid (Mesonychoteuthis
hamiltoni) attains an even greater size.
Mantle length
North Pacific Giant Octopus, Enteroctopus dofleini
Octopoda (octopuses)
Species Maximum mantle References Notes

length
Haliphron atlanticus 0.69 m O'Shea (2004)
Enteroctopus dofleini 0.6 m Norman (2000:214)
Sepiida (cuttlefish)

length
Sepia apama 0.5 m Reid et al. (2005:68)
Sepia latimanus 0.5 m Reid et al. (2005:92)
Sepia officinalis 0.49 m Reid et al. (2005:99)
Sepia pharaonis 0.42 m Reid et al. (2005:107)
Sepiolida (bobtail squid)

Cephalopod size 82

length
Austrorossia antillensis 0.09 m Reid et al. (2005:192)
Rossia pacifica 0.09 m (female) Reid et al. (2005:185) Male grows to 45 mm in mantle length (Reid et al., 2005).
Rossia macrosoma 0.085 m Reid et al. (2005:184) More typically the mantle length is 20–60 mm (Reid et al., 2005).
Neorossia caroli 0.083 m (female) Reid et al. (2005:190) Male grows to 51 mm in mantle length (Reid et al., 2005).
Spirulida (Ram's Horn Squid) (only one extant species)

length
Spirula spirula rarely exceeds 0.045 Reid et al. (2005:211)

m
Teuthida (squid)

length
Mesonychoteuthis 4 m (estimate) O'Shea (2005a) Estimate based on largest known beak (LRL: 49 mm).
hamiltoni
Galiteuthis phyllura 2.65–2.75 m Nesis (1985) Estimate based on 0.4 m long arm and 1.15 m tentacle.
(estimate)
2.25 m O'Shea (2005a)

Architeuthis sp.†
Onykia robusta 2m Norman (2000:174) Kubodera et al. (1998) give maximum of at least 1.615 m.
Megalocranchia fisheri 1.8 m Tsuchiya & Okutani (1993)
Dosidicus gigas 1.75 m Glaubrecht & Norman (2000:165) gives maximum of 1.5 m.
Salcedo-Vargas (2004)
Taningia danae 1.7 m Nesis (1982)
Kondakovia longimana probably 1.15+ m O'Shea (2005b) Longest confirmed specimen measures 0.85 m (O'Shea, 2005b). Total
(estimate) length to at least 2.3 m (Carrington, 2000).
Thysanoteuthis 1m Roper et al. (1984) Commonly grows to a mantle length of 0.6 m (Roper et al., 1984).
rhombus
cf. Magnapinna ~1 m (estimate) Vecchione et al. (2001) Estimate based on specimen observed by ROV Tiburon in May 2001,
north of Oahu, Hawaii, at a depth of 3380 m.
Vampyromorphida (Vampire Squid) (only one extant species)

length
Vampyroteuthis 0.13 m Nesis (1982)

infernalis
†
The taxonomy of the giant squid has not been entirely resolved. Lumpers and splitters may propose as many as eight species or as few as one.
No genetic or physical basis for distinguishing between the named species has been proposed.
Cephalopod size 83
Total length
A long-arm squid (cf. Magnapinna) filmed in the Gulf of Mexico.
Not to be confused with armspan, which is approximately double the total length.
Species Maximum total References Notes

length
Enteroctopus dofleini > 6.1 m Cosgrove (1987)
Haliphron atlanticus 4 m (estimate) O'Shea (2004) Estimate based on incomplete 2.90 m specimen.
Teuthida (squid)
Total length including long feeding tentacles.
Species Maximum total References Notes

length
Mesonychoteuthis 14 m (estimate) O'Shea (2005a) Estimate based on largest known beak (LRL: 49 mm).
hamiltoni
Architeuthis sp. 13 m (female) O'Shea (2005a) Measured post mortem and relaxed. Older records were exaggerated by stretching
of the tentacles (O'Shea, 2005a).
cf. Magnapinna at least 8 m Bolstad (2003) Estimate based on video evidence.

(estimate)
Asperoteuthis 5.5 m Tsuchiya & Length of immature specimen measuring 0.45 m ML. Largest known specimen
acanthoderma Okutani (1993) (0.78 m ML) would presumably be longer if it were complete (Okutani, 1995).
Onykia robusta over 4 m Verrill (1876)
Galiteuthis phyllura over 4 m Nesis (1985) Estimate based on 0.40 m long arm and 1.15 m tentacle.
(estimate)
Cephalopod size 84
Weight
Sepia apama, the heaviest species of cuttlefish
Species Maximum References Notes

weight
Haliphron atlanticus 75 kg (estimate) O'Shea (2004) Estimate based on incomplete 61.0 kg specimen.
Enteroctopus dofleini 71 kg Cosgrove (1987) Weight of live specimen. There exists a highly dubious record of a 272 kg
specimen (High, 1976).
Sepiida (cuttlefish)

weight
Sepia apama in excess of Reid et al. (2005:68)

10.5 kg
Sepia latimanus 10 kg Reid et al. (2005:92)
Sepia pharaonis 5 kg Reid et al.

(2005:107)
Sepia officinalis 4 kg Reid et al. (2005:99)
Teuthida (squid)

weight
Mesonychoteuthis 495 kg [Anonymous] (2007) Weight of mature specimen caught in early 2007. Originally estimated to
hamiltoni weigh 450 kg (Anderton, 2007).
Architeuthis sp. 275 kg (female) O'Shea (2005a)
Taningia danae 61.4 kg Kubodera et al.

(2006)
Onykia robusta 50 kg Roper et al. (1984)
Dosidicus gigas 50 kg Nigmatullin et al.

(2001)
Thysanoteuthis rhombus 30 kg Miyahara et al.

(2006)
Cephalopod size 85
Shell diameter
Nautilus shells: N. macromphalus (left), A. scrobiculatus (centre),

N. pompilius (right). Argonauta hians shell, 121.5 mm in diameter.
Internal shell of Spirula spirula.
Nautilida (nautiluses) (all extant species listed)
Species Maximum shell References Notes

diameter
Nautilus pompilius 268 mm Pisor (2005:93) lists maximum shell diameter of 254.0 mm. Nautilus repertus is treated
[1]
pompilius here in synonymy with N. pompilius pompilius. Pisor (2005:93) lists 230.0 mm record for
N. repertus.
Nautilus belauensis 226 mm Jereb

(2005:54)
Allonautilus 215.0 mm Pisor

scrobiculatus (2005:93)
Nautilus 201.0 mm Pisor

stenomphalus (2005:93)
Allonautilus around 180 mm Jereb

perforatus (2005:55)
Nautilus 180.0 mm Pisor

macromphalus (2005:93)
Nautilus pompilius 148.0 mm Pisor

suluensis (2005:93)
Cephalopod size 86
Octopoda (octopuses) (all extant Argonauta species listed)

Females of the genus Argonauta produce a calcareous eggcase in which they reside.

diameter
Argonauta argo 300.0 mm Pisor

(2005:12)
Argonauta nodosa 292.0 mm Pisor

(2005:12)
220.0 mm Pisor
Argonauta pacifica‡
(2005:12)
Argonauta hians 112.6 mm Pisor

(2005:12)
Argonauta cornuta‡ 98.6 mm Pisor

(2005:12)
Argonauta nouryi 95.5 mm Pisor

(2005:12)
Argonauta bottgeri 67.0 mm Pisor

(2005:12)
Spirulida (Ram's Horn Squid) (only one extant species)

The Ram's Horn Squid possesses a chambered internal shell, which it uses for buoyancy control.

diameter
Spirula spirula 28.8 mm Pisor (2005:108) lists maximum shell diameter of 26.9 mm.
[2]
‡
Species status questionable.
Extinct taxa
Fossilised guards of Megateuthis gigantea (top

Cast of Parapuzosia seppenradensis and centre)
Ammonoidea (ammonites)

diameter
Parapuzosia 2.55 m (estimate) Kennedy & Kaplan Estimate based on 1.95 m diameter specimen with an incomplete
seppenradensis (1995) living chamber.
Belemnoidea (belemnites)
Cephalopod size 87
Species Maximum rostrum References Notes

length
Megateuthis gigantea 0.46 m Eyden (2003) The whole belemnite is estimated to have been 3–5 m long.
Nautiloidea (nautiloids)
Species Maximum shell length References Notes
Cameroceras sp. 11 m (estimate) Teichert & Kümmel

(1960)
Vampyromorphida (vampire squid)

length
Tusoteuthis longa over 1.8 m (estimate) Eyden (2004)
References
• [Anonymous] 2007. Colossal squid may be headed for oven [3]. Associated Press.
• Anderton, H.J. 2007. Amazing specimen of world's largest squid in NZ [4]. New Zealand Government website.
• Bolstad, K. 2003. Deep-Sea Cephalopods: An Introduction and Overview [5]. The Octopus News Magazine
Online.
• Carrington, D. 2000. Big squid breaks record [6]. BBC News, July 3, 2000.
• Cosgrove, J.A. 1987. Aspects of the Natural History of Octopus dofleini, the Giant Pacific Octopus. M.Sc. Thesis.
Department of Biology, University of Victoria (Canada), 101 pp.
• Eyden, P. 2003. Belemnites: A Quick Look [7]. The Octopus News Magazine Online.
• Eyden, P. 2004. Cretaceous Giant Squid [8]. The Octopus News Magazine Online.
• Glaubrecht, M. & M.A. Salcedo-Vargas 2004. The Humboldt squid Dosidicus gigas (Orbigny, 1835): History of
the Berlin specimen, with a reappraisal of other (bathy-)pelagic gigantic cephalopods (Mollusca,
Ommastrephidae, Architeuthidae). Zoosystematics and Evolution 80(1): 53–69. doi:10.1002/mmnz.20040800105
• High, W.L. 1976. The giant Pacific octopus. U.S. National Marine Fisheries Service, Marine Fisheries Review
38(9): 17-22.
• Jereb, P. 2005. Family Nautilidae. In: P. Jereb & C.F.E. Roper, eds. Cephalopods of the world. An annotated and
Illustrated catalogue of species known to date. Volume 1. Chambered nautiluses and sepioids (Nautilidae,
Sepiidae, Sepiolidae, Sepiadariidae, Idiosepiidae and Spirulidae). FAO Species Catalogue for Fishery Purposes.
No. 4, Vol. 1. Rome, FAO. pp. 51–55.
• (German) Kennedy, W.J. & U. Kaplan 1995. Parapuzosia (Parapuzosia) seppenradensis (LANDOIS) und die
Ammoniten fauna der Dülmener Schichten, Westfalen. Geol. Paläont. Westf. 33: 127 p., 43 pls.
• Kubodera, T., U. Piatkowski, T. Okutani & M.R. Clarke. 1998. Taxonomy and Zoogeography of the Family
Onychoteuthidae (Cephalopoda: Oegopsida). Smithsonian Contributions to Zoology 586: 277-291.
• Kubodera, T., Y. Koyama & K. Mori 2006. Observations of wild hunting behaviour and bioluminescence of a
large deep-sea, eight-armed squid, Taningia danae. [9]PDF (295 KB) Proceedings of the Royal Society B:
Biological Sciences 274(1613): 1029–1034. doi:10.1098/rspb.2006.0236
• Miyahara, K., K. Fukui, T. Ota & T. Minami 2006. Laboratory observations on the early life stages of the
diamond squid Thysanoteuthis rhombus. Journal of Molluscan Studies 72(2): 199–205.
doi:10.1093/mollus/eyi068
• (Russian) Nesis, K.N. 1982. Abridged key to the cephalopod mollusks of the world's ocean. Light and Food
Industry Publishing House, Moscow. 385+ii pp. [Translated into English by B. S. Levitov, ed. by L. A. Burgess
1987. Cephalopods of the world. T.F.H. Publications, Neptune City, NJ. 351pp.]
Cephalopod size 88
• (Russian) Nesis, K.N. 1985. A Giant Squid in the Sea of Okhotsk. Priroda 10: 112-113. [Translated from Russian
by Yuri Nektorenko.]
• Nigmatullin, C.M., K.N. Nesis & A.I. Arkhipkin 2001. A review of the biology of the jumbo squid Dosidicus
gigas (Cephalopoda: Ommastrephidae). Fisheries Research 54(1): 9–19. doi:10.1016/S0165-7836(01)00371-X
• Norman, M.D. 2000. Cephalopods: A World Guide. ConchBooks.
• Norman, M.D. & A. Reid 2000. A Guide to Squid, Cuttlefish and Octopuses of Australasia. CSIRO Publishing.
• Okutani, T. 1995. Cuttlefish and squids of the world in color. Publication for the 30th anniversary of the
foundation of the National Cooperative Association of Squid Processors. 185 pp.
• O'Shea, S. 2004. The giant octopus Haliphron atlanticus (Mollusca : Octopoda) in New Zealand waters. New
Zealand Journal of Zoology 31(1): 7-13.
• O'Shea, S. 2005a. Giant Squid and Colossal Squid Fact Sheet [10]. The Octopus News Magazine Online.
• O'Shea, S. 2005b. Kondakovia longimana [11]. In: Giant Squid and Colossal Squid Fact Sheet. The Octopus News
Magazine Online.
• Pisor, D.L. 2005. Registry of World Record Size Shells: Fourth Edition - 2005. Snail's Pace Productions and
ConchBooks.
• Reid, A., P. Jereb, & C.F.E. Roper 2005. Family Sepiidae. In: P. Jereb & C.F.E. Roper, eds. Cephalopods of the
world. An annotated and illustrated catalogue of species known to date. Volume 1. Chambered nautiluses and
sepioids (Nautilidae, Sepiidae, Sepiolidae, Sepiadariidae, Idiosepiidae and Spirulidae). FAO Species Catalogue
for Fishery Purposes. No. 4, Vol. 1. Rome, FAO. pp. 57–152.
• Roper C.F.E., M.J. Sweeney & C.E. Nauen 1984. Cephalopods of the world. Food and Agriculture Organization,
Rome, Italy.
• Teichert, C. & B. Kümmel 1960. Size of Endocerid Cephalopods. Breviora Mus. Comp. Zool. 128: 1–7.
• Tsuchiya, K. & T. Okutani 1993. Rare and interesting squids in Japan -X. Recent occurrences of big squids from
Okinawa. Venus 52: 299-311.
• Vecchione, M., R.E. Young, A. Guerra, D.J. Lindsay, D.A. Clague, J.M. Bernhard, W.W. Sager, A.F. Gonzalez,
F.J. Rocha & M. Segonzac 2001. Vecchione et al., 2001 [12]. Cephalopods in Action.
• Verrill, A.E. 1876. Notes on gigantic cephalopods, a correction. American Journal of Science and Arts 12(3):
236-237.
References
[1] http:/ / www. conchology. be/ ?t=68& u=118764& g=0d16facb6ebae4a31321e9f6e6298f46& q=a1d787a5674deb7b193549c6ccdff907
[2] http:/ / www. conchology. be/ ?t=68& u=126690& g=e8a21ed471661c24ca2aea9f5ea70400& q=4e4d53268a89f2aef5613d13d96327f0
[3] http:/ / news. yahoo. com/ s/ ap/ 20070322/ ap_on_sc/ colossal_squid;_ylt=ArrD9. iKLzOCbR7FKcFksAnMWM0F
[4] http:/ / www. beehive. govt. nz/ ViewDocument. aspx?DocumentID=28451
[5] http:/ / www. tonmo. com/ science/ public/ deepseacephs. php
[6] http:/ / news. bbc. co. uk/ 2/ hi/ science/ nature/ 813394. stm
[7] http:/ / www. tonmo. com/ science/ public/ belemnites. php
[8] http:/ / www. tonmo. com/ science/ fossils/ cretaceousGS. php
[9] http:/ / www. pubs. royalsoc. ac. uk/ media/ proceedings_b/ papers/ RSPB20060236. pdf
[10] http:/ / www. tonmo. com/ science/ public/ giantsquidfacts. php
[11] http:/ / web. archive. org/ web/ 20071227144307/ http:/ / www. tonmo. com/ science/ public/ giantsquidfacts2. php
[12] http:/ / www. mnh. si. edu/ cephs/ vetal01/ vetal01. html
Cephalopod ink 89
Cephalopod ink
Cephalopod ink is a dark pigment released into water by most species
of cephalopod, usually as an escape mechanism. All cephalopods, with
the exception of the Nautilidae and the species of octopus belonging to
the suborder Cirrina,[1] are able to release ink.
The ink is released from the ink sacs (located between the gills) and is
dispersed more widely by accompanying its release with a jet of water
Ventral view of the viscera of Chtenopteryx
from the funnel. Its dark color is caused by its main constituent, sicula, showing the location of the ink sac
melanin. Each species of cephalopod produces slightly differently
coloured inks; generally, octopuses produce black ink, squid ink is blue-black and cuttlefish ink is brown (see Use by
humans).
A number of other aquatic molluscs have evolved similar responses to attack, including sea hares. This is an example
of convergent evolution.
Inking behaviours
I was much interested, on several occasions, by watching the habits of an Octopus or cuttle-fish ... they darted tail first, with the
rapidity of an arrow, from one side of the pool to the other, at the same instant discolouring the water with a dark chestnut-brown
ink.
Charles Darwin, The Voyage of the Beagle
Two distinct behaviours have been observed in inking cephalopods. The first is the release of large amounts of ink
into the water by the cephalopod, in order to create a dark, diffuse cloud (much like a smokescreen) which can
obscure the predator’s view, allowing the cephalopod to make a rapid retreat by jetting away.
The second response to a predator is to release ‘pseudomorphs’ (‘false bodies’); smaller clouds of ink with a greater
mucus content, which allows them to hold their shape for longer. These are expelled slightly away from the
cephalopod in question, which will often release several pseudomorphs and change color (blanch) in conjunction
with these releases. The pseudomorphs are roughly the same volume and look similar to the cephalopod that released
them, and many predators have been observed attacking them mistakenly, allowing the cephalopod to escape (this
behavior is often referred to as the ‘Blanch-Ink-Jet Maneuver’). Furthermore, green turtle hatchlings (Chelonia
mydas) that have been observed mistakenly attacking pseudomorphs released by Octopus bocki have subsequently
ignored conspecific octopuses.[2] However, many cephalopod predators (for instance moray eels) have advanced
chemosensory systems, and some anecdotal evidence[3] suggests that compounds such as tyrosinase found in
cephalopod ink can irritate, numb or even deactivate such apparatus. Unfortunately, few controlled experiments have
been conducted to substantiate this. Cephalopod ink is nonetheless generally thought to be more sophisticated than a
simple ‘smokescreen;’ the ink of a number of squid and cuttlefish has been shown to function as a conspecific
chemical alarm.[4]
Octopuses have also been observed squirting ink at snails or crabs approaching their eggs.[4]
Cephalopod ink 90
Chemical composition
Cephalopod ink contains a number of chemicals in a variety of different concentrations, depending on the species.
However, its main constituents are melanin and mucus. It can also contain, among other things, tyrosinase, dopamine
and L-DOPA,[5] and small amounts of amino acids, including taurine, aspartic acid, glutamic acid, alanine and
lysine.[4]
Use by humans
Cephalopod ink has, as its name suggests, been used in the past as ink;
indeed, the Greek name for cuttlefish, and the taxonomic name of a
cuttlefish genus, Sepia, is associated with the brown colour of
cuttlefish ink (for more information, see Sepia (color)). Modern use of
cephalopod ink is generally limited to cooking, where it is used as a
food colouring, for example in pasta and sauces. For this purpose it is
generally obtainable from fishmongers or gourmet food suppliers. The
ink is extracted from the ink sacs during preparation of the dead Arròs negre owes its dark colour to squid ink
cephalopod, usually squid, and therefore contains no mucus. Recent
studies have shown that cephalopod ink is toxic to some cells, including tumor cells.[4]
References
[1] Roger T. Hanlon, John B. Messenger: Cephalopod Behaviour, page 2. Cambridge University Press, 1999, ISBN 0521645832
[2] Roy L. Caldwell (2005), "An Observation of Inking Behavior Protecting Adult Octopus bocki from Predation by Green Turtle (Chelonia
mydas) Hatchlings" http:/ / muse. jhu. edu/ journals/ pacific_science/ v059/ 59. 1caldwell. pdf
[3] G.E. MacGinitie, N. MacGinitie (1968) Natural History of Marine Animals, Pages 395-397, 2nd ed. McGraw-Hill, New York.
[4] Charles D. Derby (2007), "Escape by Inking and Secreting: Marine Molluscs Avoid Predators Through a Rich Array of Chemicals and
Mechanisms" http:/ / www. biolbull. org/ cgi/ reprint/ 213/ 3/ 274. pdf
[5] http:/ / nationalzoo. si. edu/ Animals/ Invertebrates/ Facts/ cephalopods/ inking. cfm
External links
• An article on harvesting squid ink (http://www.instructables.com/id/How-to-Harvest-Squid-Ink/)
Ink sac 91
Ink sac
With the exception of nocturnal and very deep water cephalopods, all
coeloids which dwell in light conditions have an ink sac, which can be
used to expel a cloud of dark ink to confuse predators.[1] This sac is a
muscular bag which originated as an extension of the hind gut. 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
Ventral view of the viscera of Chtenopteryx
funnel means that the ink can be distributed by ejected water as the
sicula
cephalopod uses its jet propulsion.[1] The ejected cloud of melanin is
bound by mucus particles, so it forms a lump approximately the size
and shape of the cephalopod, fixing the predator's attention while the mollusc itself makes a hasty escape.[1]
A general level of provocation is necessary to trigger an octopus to release its ink, as it is biologically costly to
produce. Some species can even use their ink to stun or numb their predators.
References
[1] Boyle, Peter; Rodhouse, Paul (2004). Cephalopods : ecology and fisheries (http:/ / books. google. com/ books?id=4UtCi2B4VnoC). Ames,
Iowa: Blackwell. doi:10.1002/9780470995310.ch2. ISBN 0632060484. .
Cephalopod arm
[[file:Illex illecebrosus Arm of Illex illecebrosus with two rows of suckers along its Tentacle of Illex illecebrosus with a
arm.jpg|thumb|right length]][[file:Illex illecebrosus tentacle.jpg|thumb|right distal tentacular club (right)]]
A cephalopod arm is distinct from a tentacle, though the terms are

often used interchangeably.
Generally, cephalopod arms have suckers along most of their length, as
opposed to tentacles, which have suckers only near their ends.[1]
Octopuses have eight arms and no tentacles, while squid and cuttlefish
have eight arms and two tentacles.[2] The limbs of nautiluses, which
number around 90 and lack suckers altogether, are called tentacles.[2]
[3] [4]
The tentacles of Decapodiformes are thought to be derived from the

fourth arm pair of the ancestral coleoid, but the term arms IV is used to
refer to the subsequent, ventral arm pair in modern animals (which is
evolutionarily the fifth arm pair).[1]
The males of most cephalopods develop a specialised arm for sperm
delivery, called a hectocotylus.
Octopus arm with two rows of suckers
Abnormalities
Cephalopod arm 92
Many octopus arm anomalies have been recorded,[5] [6] including a 6-armed octopus nicknamed Henry the Hexapus,
a 7-armed octopus,[7] a 10-armed Octopus briareus,[8] one with a forked arm tip,[9] an octopus with double
hectocotylization,[10] bilateral hectocotylization,[11] and specimens with up to 96 tentacle branches.[12] [13] [14]
Branched arms have also been recorded in cuttlefish.[15]
References
[1] Young, R.E., M. Vecchione & K.M. Mangold 1999. Cephalopoda Glossary (http:/ / tolweb. org/ accessory/
Cephalopoda_Glossary?acc_id=587). Tree of Life web project.
[2] Norman, M. 2000. Cephalopods: A World Guide. ConchBooks, Hackenheim. p. 15. "There is some confusion around the terms arms versus
tentacles. The numerous limbs of nautiluses are called tentacles. The ring of eight limbs around the mouth in cuttlefish, squids and octopuses
are called arms. Cuttlefish and squid also have a pair of specialised limbs attached between the bases of the third and fourth arm pairs [...].
These are known as feeding tentacles and are used to shoot out and grab prey."
[3] Fukuda, Y. 1987. Histology of the long digital tentacles. In: W.B. Saunders & N.H. Landman (eds.) Nautilus: The Biology and Paleobiology
of a Living Fossil. Springer Netherlands. pp. 249–256. doi:10.1007/978-90-481-3299-7_17
[4] Kier, W.M. 1987. The functional morphology of the tentacle musculature of Nautilus pompilius. (http:/ / biology. unc. edu/ faculty/ Kier/ lab/
pdf/ Kier_1987. pdf)PDF In: W.B. Saunders & N.H. Landman (eds.) Nautilus: The Biology and Paleobiology of a Living Fossil. Springer
Netherlands. pp. 257–269. doi:10.1007/978-90-481-3299-7_18
[5] Kumph, H.E. 1960. Arm abnormality in octopus. Nature 185(4709): 334-335. doi:10.1038/185334a0
[6] Toll, R.B. & L.C. Binger 1991. Arm anomalies: cases of supernumerary development and bilateral agenesis of arm pairs in Octopoda
(Mollusca, Cephalopoda) Zoomorphology 110(6): 313–316.doi:10.1007/BF01668021
[7] Gleadall, I.G. 1989. An octopus with only seven arms: anatomical details (http:/ / mollus. oxfordjournals. org/ cgi/ content/ abstract/ 55/ 4/
479). Journal of Molluscan Studies 55: 479–487.
[8] Minor birth defect resulting in 10-armed juvenile, all arms fully present and functional. (http:/ / cephbase. utmb. edu/ imgdb/ imgsrch3.
cfm?ID=66& PhotographerID=& CephID=510) CephBase.
[9] Minor birth defect showing bifurcated arm tip. Both tips were fully functional. (http:/ / cephbase. utmb. edu/ imgdb/ imgsrch3. cfm?ID=65&
PhotographerID=& CephID=510) CephBase.
[10] Palacio, F.J. 1973. On the double hectocotylization of octopods. (http:/ / ia311216. us. archive. org/ 2/ items/ nautilus87amer/
nautilus87amer. pdf)PDF The Nautilus 87: 99–102.
[11] Robson, G.C. 1929. On a case of bilateral hectocotylization in Octopus rugosus. Journal of Zoology 99(1): 95–97.
doi:10.1111/j.1469-7998.1929.tb07690.x
[12] Okada, Y.K. 1965. On Japanese octopuses with branched arms, with special reference to their captures from 1884 to 1964 (http:/ / www.
journalarchive. jst. go. jp/ english/ jnlabstract_en. php?cdjournal=pjab1945& cdvol=41& noissue=7& startpage=618). Proceedings of the
Japan Academy 41(7): 618–623.
[13] Okada, Y.K. 1965. Rules of arm-branching in Japanese octopuses with branched arms (http:/ / www. journalarchive. jst. go. jp/ english/
jnlabstract_en. php?cdjournal=pjab1945& cdvol=41& noissue=7& startpage=624). Proceedings of the Japan Academy 41(7): 624–629.
[14] Monster octopi with scores of extra tentacles (http:/ / pinktentacle. com/ 2008/ 07/ monster-octopi-with-scores-of-extra-tentacles/ ). Pink
Tentacle, July 18, 2008.
[15] Okada, Y.K. 1937. An occurrence of branched arms in the decapod cephalopod, Sepia esculenta Hoyle. Annotated Zoology of Japan 17:
93–94.
Hectocotylus 93
Hectocotylus
Lateral view of adult male Tremoctopus violaceus

with hectocotylus
[[file:Ocythoe tuberculata Detail of the hectocotylus of Ocythoe tuberculata]][[file:Haliphron Hectocotylus of a young

hectocotylus.jpg|thumb|right atlanticus hectocotylus.jpg|thumb|right Haliphron atlanticus]]
A hectocotylus is one of the arms of the male of most kinds of cephalopods that is modified in various ways to
effect the fertilization of the female's eggs. It is a specialized, extended muscular hydrostat used to store
spermatophores, the male gametophore. Males generally form a new hectocotylus in each new season.
Hectocotylus 94
The shape of the tip of the hectocotylus has been much used in octopod systematics. In many species it is
considerably elaborated. However, in the males of some species, such as the Seven-arm Octopus (Haliphron
atlanticus), the hectocotylus develops in an inconspicuous sac in front of the right eye that gives the male the
appearance of having only seven arms.
The term is also used to specifically refer to the greatly modified arm of Argonauta and allied genera. In argonauts,
the male transfers the spermatophores to the female by putting its hectocotylus into a cavity in the mantle of the
female. This mantle cavity is known as the pallial cavity. This is the only contact the male and female have with
each other during copulation, and it can be at a distance. During copulation, the hectocotylus breaks off from the
male. The funnel-mantle locking apparatus on the hectocotylus keeps it lodged in the pallial cavity of the female.
The name hectocotylus was devised by Georges Cuvier, who first found one embedded in the mantle of a female
argonaut; supposing it to be a parasitic worm, Cuvier gave it a generic name. The hectocotyl arm was first described
in the biological works of Aristotle, and it was widely disbelieved until its rediscovery in the nineteenth century.[1] [2]
Rare examples of double and bilateral hectocotylization have been recorded in incirrate octopuses.[3] [4]
References
[1] Thompson, Darcy Wentworth. 1913. On Aristotle as a biologist, with a prooemion on Herbert Spencer. Being the Herbert Spencer Lecture
before the University of Oxford, on February 14, 1913. Oxford University Press.
[2] Nixon M. & J.Z. Young J.Z. 2003. The brains and lives of Cephalopods. Oxford University Press.
[3] Robson, G.C. 1929. On a case of bilateral hectocotylization in Octopus rugosus. Journal of Zoology 99(1): 95–97.
doi:10.1111/j.1469-7998.1929.tb07690.x
[4] Palacio, F.J. 1973. On the double hectocotylization of octopods. (http:/ / ia311216. us. archive. org/ 2/ items/ nautilus87amer/ nautilus87amer.
pdf)PDF The Nautilus 87: 99–102.
Tentacle
A tentacle or bothrium (plural: bothria) is one of usually two or more
elongated flexible organs present in animals, especially invertebrates. The term
may also refer to the hairs of the leaves of some insectivorous plants. Usually,
tentacles are used for feeding, feeling and grasping. Anatomically, they work like
other muscular hydrostats.
Tentacles in animals
Invertebrates
The phylum Mollusca includes many species with muscular hydrostats in the
form of tentacles and arms. Octopuses do not have tentacles but rather have eight
arms. Tentacles are distinguished in this context as being longer than arms, with
suckers at their tips only. Squid and cuttlefish have eight arms like octopuses,
and two extra flexible tentacles. Cuttlefish with 2 tentacles and 8
arms
The tentacles of the Giant Squid and Colossal Squid have powerful suckers and
pointed teeth at the ends. The teeth of the Giant Squid resemble bottle caps, and
function like small, circular saws; while the tentacles of the Colossal Squid wield two long rows of swiveling,
tri-pointed hooks.
Tentacle 95
Snails are another class of Mollusca.

They have less elaborate tentacles than
the Cephalopods. Pulmonate land
snails usually have two sets of
tentacles on the head: the upper pair
have an eye at the end; the lower pair
are for olfaction. Both pairs are fully
retractable. Some marine snails such as
Front view of land snail showing upper and lower
sets of tentacles the abalone and the top snails,
Trochidae have numerous small
tentacles around the edge of the
mantle. These are known as pallial
tentacles.
Cnidarians, which include among

others the jellyfishes, are another
phylum with many tentacled
specimens. Cnidarians often have huge
numbers of cnidocytes on their
tentacles. Cnidocytes are cells
Abalone showing pallial tentacles
containing a coiled thread-like
structure called a nematocyst, fired at
potential prey.
Many species of the jellyfishlike ctenophores have two tentacles, while some have none. Their tentacles have
adhesive structures called colloblasts or lasso cells. These cells burst open when prey comes in contact with the
tentacle; sticky threads released from each of the colloblasts then captures food.
The tentacles of the Lion's mane jellyfish can reach 120 feet (37 meters).
Bryozoa (Moss animals) are tiny creatures with a ring of tentacles surrounding the mouth.
Tentacle 96
Amphibians
Some wormlike amphibians have tentacles. The caecilians have two tentacles at their heads, which are probably used
for smell.
Mammals
The star-nosed mole, Condylura cristata, possesses nasal tentacles which are mobile and extremely sensitive,
helping the animal to find its way about the burrow and detect prey.
Tentacles in plants
In carnivorous plants, tentacles refer to the stalked glands of the upper surface of
the leaves.
On a sundew plant, they are hairlike projections with a drop of nectar-like glue
which attract insects. When an insect is captured, the tentacles bend inward and
the leaf rolls together as shown in the picture. The tentacles then secrete enzymes
to dissolve and digest the insect.
External links
• Difference between arms and tentacles [1]
Leaf and tentacle movement on

Drosera capensis
References
[1] http:/ / www. cephbase. utmb. edu/ TCP/ faq/ TCPfaq2b. cfm?ID=50
Dactylus 97
Dactylus
The dactylus is the tip region of the tentacular club of cephalopods and of the leg of some crustaceans (see arthropod
leg). In cephalopods, the dactylus is narrow and often characterized by the asymmetrical placement of suckers (i.e.,
the ventral expansion of the club) and the absence of a dorsal protective membrane. In crustaceans, the dactylus is
the seventh and terminal segment of their thoracic appendages. In certain instances the dactylus, together with the
propodus, form the claw.
The term dactylus means "finger" in Greek.
References
Cephalopod eye
Vertebrate
Octopus
Vertebrates and octopuses developed the camera eye independently. In the vertebrate version the nerve fibers pass in front of the
retina, and there is a blind spot where the nerves pass through the retina. In the vertebrate example, 4 represents the blind spot,
which is notably absent from the octopus eye. In vertebrates, 1 represents the retina and 2 is the nerve fibers, including the optic
nerve (3), whereas in the octopus eye, 1 and 2 represent the nerve fibers and retina respectively.
Cephalopods, as active marine predators, possess sensory organs specialized for use in aquatic conditions.[1] They
have a camera-type eye, which consists of a lens projecting an image onto a retina. Unlike the vertebrate camera eye,
the cephalopods' form as invaginations of the body surface (rather than outgrowths of the brain), and consequently
they lack a cornea. Unlike the vertebrate eye, a cephalopod eye is focused through movement, much like the lens of a
camera or telescope, rather than changing shape as the lens in the human eye does. The eye is approximately
spherical, as is the lens, which is fully internal.[2]
Cephalopod eye 98
The crystalins used in the lens appear to have developed independently

from vertebrate crystalins, suggesting a homoplasious origin of the
lens.[3]
Most cephalopods possess complex extraocular muscle systems that
allow for very fine control over the gross positioning of the eyes.
Octopuses possess an autonomic response that maintains the
orientation of their pupils such that they are always horizontal.[1]
Eye of Bathyteuthis sp.

Polarized light
It has been documented that several types of cephalopods, most
notably squid and octopuses, and potentially cuttlefish, have eyes that can distinguish the orientation of polarized
light. This sensitivity is due to the orthogonal organization of neighboring photoreceptors. It is theorized that this
ability plays a role in communication among the color-changing cephalopods.[4]
Cephalopod eye 99
References
[1] Budelmann BU. "Cephalopod sense organs, nerves and the brain: Adaptations for high performance and life style." Marine and Freshwater
Behavior and Physiology. Vol 25, Issue 1-3, Page 13-33.
[2] Yamamoto, M. (Feb 1985). "Ontogeny of the visual system in the cuttlefish, Sepiella japonica. I. Morphological differentiation of the visual
cell". The Journal of Comparative Neurology 232 (3): 347–361. doi:10.1002/cne.902320307. ISSN 0021-9967. PMID 2857734.
[3] SAMIR K. BRAHMA1 (1978). "Ontogeny of lens crystallins in marine cephalopods" (http:/ / dev. biologists. org/ cgi/ reprint/ 46/ 1/ 111.
pdf). Embryol. Exp. Morph. Vol. 46,pp. 111-118, 46: 111–8. PMID 359745. .
[4] Shashar N, Rutledge P, and Cronin T. "Polarization vision in cuttlefish in a concealed communication channel?" Journal of Experimental
Biology, Vol 199, Issue 9 2077-2084
Chromatophore
Chromatophores are pigment-containing and light-reflecting cells
found in amphibians, fish, reptiles, crustaceans, and cephalopods. They
are largely responsible for generating skin and eye colour in
cold-blooded animals and are generated in the neural crest during
embryonic development. Mature chromatophores are grouped into
subclasses based on their colour (more properly "hue") under white
light: xanthophores (yellow), erythrophores (red), iridophores
(reflective / iridescent), leucophores (white), melanophores
(black/brown) and cyanophores (blue). The term can also refer to
coloured, membrane associated vesicles found in some forms of
photosynthetic bacteria.
Some species can rapidly change colour through mechanisms that

translocate pigment and reorient reflective plates within
chromatophores. This process, often used as a type of camouflage, is
called physiological colour change. Cephalopods such as octopus have
complex chromatophore organs controlled by muscles to achieve this,
while vertebrates such as chameleons generate a similar effect by cell
signaling. Such signals can be hormones or neurotransmitters and may
Zebrafish chromatophores mediate background
be initiated by changes in mood, temperature, stress or visible changes
adaptation on exposure to dark (top) and light
in local environment. environments (bottom).
Unlike cold-blooded animals, mammals and birds have only one class
of chromatophore-like cell type: the melanocyte. The cold-blooded equivalent, melanophores, are studied by
scientists to understand human disease and used as a tool in drug discovery.
Classification
Invertebrate pigment-bearing cells were first described as chromoforo in an Italian science journal in 1819.[1] The
term chromatophore was adopted later as the name for pigment bearing cells derived from the neural crest of
cold-blooded vertebrates and cephalopods. The word itself comes from the Greek words khrōma (χρωμα) meaning
"color," and phoros (φορος) meaning "bearing". In contrast, the word chromatocyte (cyte or κυτε being Greek for
"cell") was adopted for the cells responsible for colour found in birds and mammals. Only one such cell type, the
melanocyte, has been identified in these animals.
It wasn't until the 1960s that the structure and colouration of chromatophores were understood well enough to allow
the development of a system of sub-classification based on their appearance. This classification system persists to
this day even though more recent studies have revealed that certain biochemical aspects of the pigments may be
Chromatophore 100
more useful to a scientific understanding of how the cells function.[2]

Color-producing molecules fall into two distinct classes: biochromes and schemochromes.[3] The biochromes include
true pigments, such as carotenoids and pteridines. These pigments selectively absorb parts of the visible light
spectrum that makes up white light while permitting other wavelengths to reach the eye of the observer.
Schemochromes, also known as "structural colors", produce coloration by reflecting some wavelengths (colors) of
light and transmitting others, by causing light waves to interfere within the structure or by scattering light which falls
upon them.
While all chromatophores contain pigments or reflecting structures (except when there has been a genetic mutation
resulting in a disorder like albinism), not all pigment containing cells are chromatophores. Haem, for example, is a
biochrome responsible for the red appearance of blood. It is primarily found in red blood cells (erythrocytes), which
are generated in bone marrow throughout the life of an organism, rather than being formed during embryological
development. Therefore erythrocytes are not classified as chromatophores.
Xanthophores and erythrophores

Chromatophores that contain large amounts of yellow pteridine
pigments are named xanthophores; those with a preponderance of
red/orange carotenoids are termed erythrophores.[2] Pteridine and
carotenoid containing vesicles are sometimes found within the same
cell, in which case the overall colour depends on the ratio of red and
yellow pigments.[4] Therefore, the distinction between these
chromatophore types is not always clear.
The capacity to generate pteridines from guanosine triphosphate is a

feature common to most chromatophores, but xanthophores appear to
have supplemental biochemical pathways that result in an excess A veiled chameleon, Chamaeleo calyptratus.
Structural green and blue colors are generated by
accumulation of yellow pigment. In contrast, carotenoids are
overlaying chromatophore types to reflect filtered
metabolised from the diet and transported to erythrophores. This was light.
first demonstrated by rearing normally green frogs on a diet of
carotene-restricted crickets. The absence of carotene in the frogs' diet meant the red/orange carotenoid colour 'filter'
was not present in their erythrophores. This resulted in the frogs appearing blue in colour, instead of green.[5]
Iridophores and leucophores

Iridophores, sometimes also called guanophores, are pigment cells that reflect light using plates of crystalline
chemochromes made from guanine.[6] When illuminated they generate iridescent colors because of the diffraction of
light within the stacked plates. Orientation of the schemochrome determines the nature of the colour observed.[7] By
using biochromes as coloured filters, iridophores create an optical effect known as Tyndall or Rayleigh scattering,
producing bright blue or green colors.[8]
A related type of chromatophore, the leucophore, is found in some fish, particularly in the tapetum lucidum. Like
iridophores, they utilize crystalline purines (often guanine) to reflect light. Unlike iridophores, however, leucophores
have more organized crystals which reduce diffraction. Given a source of white light, they produce a white shine. As
with xanthophores and erythrophores, in fish the distinction between iridophores and leucophores is not always
obvious, but generally iridophores are considered to generate iridescent or metallic colors while leucophores produce
reflective white hues.[8]
Chromatophore 101
Melanophores
Melanophores contain eumelanin, a type of melanin, that appears black
or dark brown because of its light absorbing qualities. It is packaged in
vesicles called melanosomes and distributed throughout the cell.
Eumelanin is generated from tyrosine in a series of catalysed chemical
reactions. It is a complex chemical containing units of dihydroxyindole
and dihydroxyindole-2-carboxylic acid with some pyrrole rings.[9] The
key enzyme in melanin synthesis is tyrosinase. When this protein is
defective, no melanin can be generated resulting in certain types of
albinism. In some amphibian species there are other pigments
At the bottom a mutant zebrafish larva that fails
packaged alongside eumelanin. For example, a novel deep (wine) red
to synthesise melanin in its melanophores, at the
coloured pigment was identified in the melanophores of top a non-mutant, wildtype larva.
phyllomedusine frogs.[10] This was subsequently identified as
pterorhodin, a pteridine dimer that accumulates around eumelanin core, and it is also present in a variety of tree frog
species from Australia and Papua New Guinea. While it is likely that other lesser studied species have complex
melanophore pigments, it is nevertheless true that the majority of melanophores studied to date do contain eumelanin
exclusively.[11]
Humans have only one class of pigment cell, the mammalian equivalent of melanophores, to generate skin, hair and
eye colour. For this reason, and because the large number and contrasting colour of the cells usually make them very
easy to visualise, melanophores are by far the most widely studied chromatophore. However, there are differences
between the biology of melanophores and melanocytes. In addition to eumelanin, melanocytes can generate a
yellow/red pigment called phaeomelanin.
Cyanophores
In 1995 it was demonstrated that the vibrant blue colors in some types
of mandarinfish are not generated by schemochromes. Instead, a cyan
biochrome of unknown chemical nature is responsible.[8] This pigment,
found within vesicles in at least two species of callionymid fish, is
highly unusual in the animal kingdom, as all other blue colourings thus
far investigated are schemochromatic. Therefore a novel
chromatophore type, the cyanophore, was proposed. Although they
Dendrobates pumilio, a poison dart frog. Some
appear unusual in their taxonomic restriction, there may be
brightly coloured species have unusual
cyanophores (as well as further unusual chromatophore types) in other chromatophores of unknown pigment
fish and amphibians. For example, bright coloured chromatophores composition.
with undefined pigments have been observed in both poison dart frogs
and glass frogs.[12]
Pigment translocation
Many species have the ability to translocate the pigment inside chromatophores, resulting in an apparent change in
colour. This process, known as physiological colour change, is most widely studied in melanophores, since melanin
is the darkest and most visible pigment. In most species with a relatively thin dermis, the dermal melanophores tend
to be flat and cover a large surface area. However, in animals with thick dermal layers, such as adult reptiles, dermal
melanophores often form three-dimensional units with other chromatophores. These dermal chromatophore units
(DCU) consist of an uppermost xanthophore or erythrophore layer, then an iridophore layer, and finally a basket-like
melanophore layer with processes covering the iridophores.[13]
Chromatophore 102
Both types of dermal melanophores are important in physiological colour change. Flat dermal melanophores will
often overlay other chromatophores so when the pigment is dispersed throughout the cell the skin appears dark.
When the pigment is aggregated towards the centre of the cell, the pigments in other chromatophores are exposed to
light and the skin takes on their hue. Similarly, after melanin aggregation in DCUs, the skin appears green through
xanthophore (yellow) filtering of scattered light from the iridophore layer. On the dispersion of melanin, the light is
no longer scattered and the skin appears dark. As the other biochromatic chomatophores are also capable of pigment
translocation, animals with multiple chromatophore types can generate a spectacular array of skin colors by making
good use of the divisional effect.[14] ,[15]
The control and mechanics of rapid pigment translocation has been
well studied in a number of different species, particularly amphibians
and teleost fish.[16] ,[8] It has been demonstrated that the process can be
under hormonal, neuronal control or both. Neurochemicals that are
known to translocate pigment include noradrenaline, through its
receptor on the surface on melanophores.[17] The primary hormones
involved in regulating translocation appear to be the melanocortins,
melatonin and melanin concentrating hormone (MCH), that are
produced mainly in the pituitary, pineal gland and hypothalamus
respectively. These hormones may also be generated in a paracrine
A single zebrafish melanophore imaged by fashion by cells in the skin. At the surface of the melanophore the
time-lapse photography during pigment
hormones have been shown to activate specific G-protein coupled
aggregation
receptors that, in turn, transduce the signal into the cell. Melanocortins
result in the dispersion of pigment, while melatonin and MCH results
in aggregation.[18]
Numerous melanocortin, MCH and melatonin receptors have been identified in fish[19] and frogs,[20] including a
homologue of MC1R,[21] a melanocortin receptor known to regulate skin and hair colour in humans.[22] It has been
demonstrated that MC1R is required in zebrafish for dispersion of melanin.[23] Inside the cell, cyclic adenosine
monophosphate (cAMP) has been shown to be an important second messenger of pigment translocation. Through a
mechanism not yet fully understood, cAMP influences other proteins such as protein kinase A to drive molecular
motors carrying pigment containing vesicles along both microtubules and microfilaments.[24] ,[25] ,[26]
Background adaptation
Most fish, reptiles and amphibians undergo a limited physiological colour change in response to a change in
environment. This type of camouflage, known as background adaptation, most commonly appears as a slight
darkening or lightening of skin tone to approximately mimic the hue of the immediate environment. It has been
demonstrated that the background adaptation process is vision dependent (it appears the animal needs to be able to
see the environment to adapt to it),[27] and that melanin translocation in melanophores is the major factor in colour
change.[18] Some animals, such as chameleons and anoles, have a highly developed background adaptation response
capable of generating a number of different colors very rapidly. They have adapted the capability to change colour in
response to temperature, mood, stress levels and social cues, rather than to simply mimic their environment.
Chromatophore 103
Development
During vertebrate embryonic development,
chromatophores are one of a number of cell types
generated in the neural crest, a paired strip of cells
arising at the margins of the neural tube. These cells
have the ability to migrate long distances, allowing
chromatophores to populate many organs of the body,
including the skin, eye, ear and brain. Leaving the
neural crest in waves, chromatophores take either a
dorsolateral route through the dermis, entering the
ectoderm through small holes in the basal lamina, or a
ventromedial route between the somites and the neural
tube. The exception to this is the melanophores of the
retinal pigmented epithelium of the eye. These are not
derived from the neural crest, instead an outpouching
of the neural tube generates the optic cup which, in
turn, forms the retina.
When and how multipotent chromatophore precursor

Transverse section of a developing vertebrate trunk showing the
cells (called chromatoblasts) develop into their
dorsolateral (red) and ventromedial (blue) routes of chromatoblast
daughter subtypes is an area of ongoing research. It is migration
known in zebrafish embryos, for example, that by 3
days after fertilization each of the cell classes found in the adult fish — melanophores, xanthophores and iridophores
— are already present. Studies using mutant fish have demonstrated that transcription factors such as kit, sox10 and
mitf are important in controlling chromatophore differentiation.[28] If these proteins are defective, chromatophores
may be regionally or entirely absent, resulting in a leucistic disorder.
Practical applications
In addition to basic research into better understanding of chromatophores themselves, the cells are used for applied
research purposes. For example, zebrafish larvae are used to study how chromatophores organise and communicate
to accurately generate the regular horizontal striped pattern as seen in adult fish.[29] This is seen as a useful model
system for understanding patterning in the evolutionary developmental biology field. Chromatophore biology has
also been used to model human condition or disease, including melanoma and albinism. Recently the gene
responsible for the melanophore-specific golden zebrafish strain, Slc24a5, was shown to have a human equivalent
that strongly correlates with skin colour.[30]
Chromatophores are also used as a biomarker of blindness in cold-blooded species, as animals with certain visual
defects fail to background adapt to light environments.[27] Human homologues of receptors that mediate pigment
translocation in melanophores are thought to be involved in processes such as appetite suppression and tanning,
making them attractive targets for drugs.[21] Therefore pharmaceutical companies have developed a biological assay
for rapidly identifying potential bioactive compounds using melanophores from the African clawed frog.[31] Other
scientists have developed techniques for using melanophores as biosensors,[32] and for rapid disease detection (based
on the discovery that pertussis toxin blocks pigment aggregation in fish melanophores).[33] Potential military
applications of chromatophore mediated colour changes have been proposed, mainly as a type of active
camouflage.[34]
Chromatophore 104
Cephalopod chromatophores
Coleoid cephalopods have complex
multicellular organs which they use to
change colour rapidly. This is most notable
in brightly coloured squid, cuttlefish and
octopuses. Each chromatophore unit is
composed of a single chromatophore cell
and numerous muscle, nerve, glial and
sheath cells.[35] Inside the chromatophore
cell, pigment granules are enclosed in an
elastic sac, called the cytoelastic sacculus.
To change colour the animal distorts the
sacculus form or size by muscular
contraction, changing its translucency,
reflectivity or opacity. This differs from the
mechanism used in fish, amphibians and An infant cuttlefish, using background adaptation to mimic the local environment
reptiles, in that the shape of the sacculus is
being changed rather than a translocation of pigment vesicles within the cell. However a similar effect is achieved.
Octopuses can operate chromatophores in complex, wavelike chromatic displays, resulting in a variety of rapidly
changing colour schemes. The nerves that operate the chromatophores are thought to be positioned in the brain in a
pattern similar to that of the chromatophores they each control. This means the pattern of colour change matches the
pattern of neuronal activation. This may explain why, as the neurons are activated one after another, the colour
change occurs in waves.[36] Like chameleons, cephalopods use physiological colour change for social interaction.
They are also among the most skilled at background adaptation, having the ability to match both the colour and the
texture of their local environment with remarkable accuracy.
Bacteria
Chromatophores are also found in membranes of phototrophic bacteria. Used primarily for photosynthesis, they
contain bacteriochlorophyll pigments and carotenoids.[37] In purple bacteria, such as Rhodospirillum rubrum the
light-harvesting proteins are intrinsic to the chromatophore membranes. However, in green sulfur bacteria they are
arranged in specialised antenna complexes called chlorosomes.[38]
See also
• Chromophore
Chromatophore 105
Notes
[1] Sangiovanni G. Descrizione di un particolare sistema di organi cromoforo espansivo-dermoideo e dei fenomeni che esso produce, scoperto
nei molluschi cefaloso. G. Enciclopedico Napoli. 1819; 9:1–13.
[2] Bagnara JT. (1966). "Cytology and cytophysiology of non-melanophore pigment cells". Int Rev Cytol 20: 173–205.
doi:10.1016/S0074-7696(08)60801-3. PMID 5337298.
[3] Fox, DL. Animal Biochromes and Structural Colors: Physical, Chemical, Distributional & Physiological Features of Colored Bodies in the
Animal World. University of California Press, Berkeley, 1976. ISBN 0520023471
[4] Matsumoto J (1965). "Xiphophorus helleri" (http:/ / www. pubmedcentral. nih. gov/ articlerender. fcgi?tool=pmcentrez& artid=2106771). J
Cell Biol 27 (3): 493–504. PMID 5885426. PMC 2106771.
[5] Bagnara JT. Comparative Anatomy and Physiology of Pigment Cells in Nonmammalian Tissues. In: The Pigmentary System: Physiology and
Pathophysiology, Oxford University Press, 1998. ISBN 0-19-509861-7
[6] Taylor JD. (1969). "The effects of intermedin on the ultrastructure of amphibian iridophores". Gen Comp Endocrinol 12 (3): 405–16.
doi:10.1016/0016-6480(69)90157-9. PMID 5769930.
[7] Morrison RL. (1995). "A transmission electron microscopic (TEM) method for determining structural colors reflected by lizard iridophores".
Pigment Cell Res 8 (1): 28–36. doi:10.1111/j.1600-0749.1995.tb00771.x. PMID 7792252.
[8] Fujii R. (2000). "The regulation of motile activity in fish chromatophores". Pigment Cell Res 13 (5): 300–19.
doi:10.1034/j.1600-0749.2000.130502.x. PMID 11041206.
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review". Pigment Cell Res 16 (5): 523–31. doi:10.1034/j.1600-0749.2003.00072.x. PMID 12950732.
[10] Bagnara, J.T.; Taylor, JD; Prota, G (1973). "Color changes, unusual melanosomes, and a new pigment from leaf frogs" (http:/ / www.
sciencemag. org/ cgi/ content/ abstract/ 182/ 4116/ 1034). Science 182 (4): 1034–5. doi:10.1126/science.182.4116.1034. PMID 4748673. .
[11] Bagnara, J.T. (2003). "Enigmas of Pterorhodin, a Red Melanosomal Pigment of Tree Frogs" (http:/ / www. ingentaconnect. com/ content/
mksg/ pcr/ 2003/ 00000016/ 00000005/ art00014). Pigment Cell Research 16 (5): 510–516. doi:10.1034/j.1600-0749.2003.00075.x.
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[12] Schwalm PA, Starrett PH, McDiarmid RW (1977). "Infrared reflectance in leaf-sitting neotropical frogs". Science 196 (4295): 1225–7.
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[13] Bagnara JT, Taylor JD, Hadley ME (1968). "[http://www.jcb.org/cgi/reprint/38/1/67 "The dermal chromatophore unit" (http:/ / www.
pubmedcentral. nih. gov/ articlerender. fcgi?tool=pmcentrez& artid=2107474). J Cell Biol 38 (1): 67–79. PMID 5691979. PMC 2107474.
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dispersion in goldfish xanthophores". Cell Motil Cytoskeleton 13 (1): 9–20. doi:10.1002/cm.970130103. PMID 2543509.
[15] Porras MG, De Loof A, Breuer M, Aréchiga H (2003). "Procambarus clarkii". Peptides 24 (10): 1581–9. PMID 14706537.
[16] Deacon SW, Serpinskaya AS, Vaughan PS, Lopez Fanarraga M, Vernos I, Vaughan KT, Gelfand VI (2003). "Dynactin is required for
bidirectional organelle transport." (http:/ / www. jcb. org/ cgi/ content/ full/ 160/ 3/ 297). The Journal of cell biology 160 (3): 297–301.
doi:10.1083/jcb.200210066. PMID 12551954. PMC 2172679. .
[17] Aspengren S, Sköld HN, Quiroga G, Mårtensson L, Wallin M (2003). "Noradrenaline- and melatonin-mediated regulation of pigment
aggregation in fish melanophores". Pigment Cell Res 16 (1): 59–64. doi:10.1034/j.1600-0749.2003.00003.x. PMID 12519126.
[18] Logan DW, Burn SF, Jackson IJ (2006). "Regulation of pigmentation in zebrafish melanophores". Pigment Cell Res 19 (3): 206–13.
doi:10.1111/j.1600-0749.2006.00307.x. PMID 16704454.
[19] Logan DW, Bryson-Richardson RJ, Taylor MS, Currie P, Jackson IJ (2003). "Sequence characterization of teleost fish melanocortin
receptors". Ann N Y Acad Sci 994: 319–30. doi:10.1111/j.1749-6632.2003.tb03196.x. PMID 12851332.
[20] Sugden D, Davidson K, Hough KA, Teh MT (2004). "Melatonin, melatonin receptors and melanophores: a moving story". Pigment Cell Res
17 (5): 454–60. doi:10.1111/j.1600-0749.2004.00185.x. PMID 15357831.
[21] Logan DW, Bryson-Richardson RJ, Pagán KE, Taylor MS, Currie PD, Jackson IJ (2003). "The structure and evolution of the melanocortin
and MCH receptors in fish and mammals". Genomics 81 (2): 184–91. doi:10.1016/S0888-7543(02)00037-X. PMID 12620396.
[22] Valverde P, Healy E, Jackson I, Rees JL, Thody AJ (1995). "Variants of the melanocyte-stimulating hormone receptor gene are associated
with red hair and fair skin in humans". Nat Genet 11 (3): 328–30. doi:10.1038/ng1195-328. PMID 7581459.
[23] Richardson J, Lundegaard PR, Reynolds NL, Dorin JR, Porteous DJ, Jackson IJ, Patton EE (2008). "mc1r Pathway regulation of zebrafish
melanosome dispersion". Zebrafish 5 (4): 289–95. doi:10.1089/zeb.2008.0541. PMID 19133827.
[24] Snider J, Lin F, Zahedi N, Rodionov V, Yu CC, Gross SP (2004). "Intracellular actin-based transport: how far you go depends on how often
you switch." (http:/ / www. pnas. org/ cgi/ content/ full/ 101/ 36/ 13204). Proceedings of the National Academy of Sciences of the United
States of America 101 (36): 13204–9. doi:10.1073/pnas.0403092101. PMID 15331778. PMC 516548. .
[25] Rodionov VI, Hope AJ, Svitkina TM, Borisy GG (1998). "Functional coordination of microtubule-based and actin-based motility in
melanophores." (http:/ / www. sciencedirect. com/ science?_ob=ArticleURL& _udi=B6VRT-4D04MK2-4& _user=10& _coverDate=01/ 29/
1998& _alid=412398039& _rdoc=1& _fmt=full& _orig=search& _qd=1& _cdi=6243& _sort=d& _docanchor=& view=c&
_acct=C000050221& _version=1& _urlVersion=0& _userid=10& md5=53b4395208a20da195d9c1b3dbe0fe41& artImgPref=F). Current
biology : CB 8 (3): 165–8. doi:10.1016/S0960-9822(98)70064-8. PMID 9443917. .
[26] Kashina AS, Semenova IV, Ivanov PA, Potekhina ES, Zaliapin I, Rodionov VI (2004). "Protein kinase A, which regulates intracellular
transport, forms complexes with molecular motors on organelles." (http:/ / www. sciencedirect. com/ science?_ob=ArticleURL&
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md5=96f1a14140773f1e898c1ef410db1cb9& artImgPref=F). Current biology : CB 14 (20): 1877–81. doi:10.1016/j.cub.2004.10.003.
PMID 15498498. .
[27] Neuhauss SC. (2003). "Behavioral genetic approaches to visual system development and function in zebrafish" (http:/ / www3. interscience.
wiley. com/ cgi-bin/ fulltext/ 101526009/ PDFSTART) (PDF). J Neurobiol 54 (1): 148–60. doi:10.1002/neu.10165. PMID 12486702. .
[28] Kelsh RN, Schmid B, Eisen JS (2000). "Genetic analysis of melanophore development in zebrafish embryos". Dev Biol 225 (2): 277–93.
doi:10.1006/dbio.2000.9840. PMID 10985850.
[29] Kelsh RN (2004). "Genetics and evolution of pigment patterns in fish.". Pigment cell research / sponsored by the European Society for
Pigment Cell Research and the International Pigment Cell Society 17 (4): 326–36. doi:10.1111/j.1600-0749.2004.00174.x. PMID 15250934.
[30] Lamason RL, Mohideen MA, Mest JR, Wong AC, Norton HL, Aros MC, Jurynec MJ, Mao X et al. (2005). "SLC24A5, a putative cation
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[31] Jayawickreme CK, Sauls H, Bolio N, Ruan J, Moyer M, Burkhart W, Marron B, Rimele T et al. (1999). "Use of a cell-based, lawn format
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[33] Karlsson JO, Andersson RG, Askelöf P, Elwing H, Granström M, Grundström N, Lundström I, Ohman L (1991). "The melanophore
aggregating response of isolated fish scales: a very rapid and sensitive diagnosis of whooping cough". FEMS Microbiol Lett 66 (2): 169–75.
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[34] Lee I. Nanotubes for noisy signal processing: Adaptive Camouflage PhD Thesis. 2005; University of Southern California. Retrieved June
2006 (http:/ / biron. usc. edu/ ~ianlee/ index_files/ thesis_ch5. pdf)PDF (799 KiB).
[35] Cloney RA, Florey E (1968). "Ultrastructure of cephalopod chromatophore organs.". Zeitschrift fur Zellforschung und mikroskopische
Anatomie (Vienna, Austria : 1948) 89 (2): 250–80. doi:10.1007/BF00347297. PMID 5700268.
[36] Demski LS (1992). "Chromatophore systems in teleosts and cephalopods: a levels oriented analysis of convergent systems.". Brain,
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[37] Salton MR (1987). "Bacterial membrane proteins.". Microbiological sciences 4 (4): 100–5. PMID 3153178.
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External links
• Nature's Palette - how animals produce colour (http://www.bioscience-explained.org/EN1.2/pdf/paletteEN.
pdf)PDF (1.20 MiB)
• Video footage of octopus background adaptation (http://www.funny-games.biz/videos/78-octopus.html)
• Video footage of squid chromatophore patterning (http://www.gfai.de/~heinz/historic/biomodel/squids/
squids.htm)
• Tree of Life Web Project: Cephalopod Chromatophores (http://tolweb.org/tree/eukaryotes/animals/mollusca/
cephalopoda/glossary/glossaryLichen/chromatophoreLichen/Chromatophore.html)
Mantle 107
Mantle
The mantle (also known by the Latin word pallium meaning mantle,
robe or cloak, adjective pallial) is a significant part of the anatomy of
molluscs: it is the dorsal body wall which covers the visceral mass and
usually protrudes in the form of flaps well beyond the visceral mass
itself.
In many, but by no means all, species of molluscs, the epidermis of the
mantle secretes calcium carbonate and conchiolin, and creates a shell.
The words mantle and pallium both originally meant cloak or cape, see
mantle (vesture). This anatomical structure in molluscs often resembles European Squid (Loligo vulgaris). The mantle is
a cloak because in many groups the edges of the mantle extend far all that is visible behind the head: the outer body
wall and the fins are all part of the mantle.
beyond the main part of the body, forming flaps, double-layered
structures which have been adapted for many different uses, including
for example, the siphon.
The mantle cavity

The mantle cavity is a central feature of molluscan biology. This cavity
is formed by the mantle skirt, a double fold of mantle which encloses a
water space. This space contains the mollusc's gills, anus, osphradium,
nephridiopores, and gonopores.
The mantle cavity functions as a respiratory chamber in all molluscs.
The brightly coloured mantle of a Giant clam
In bivalves it is usually part of the feeding structure. In some mollusks
protects it from bright sunlight.
the mantle cavity is a brood chamber, and in cephalopods and some
bivalves such as scallops, it is a locomotory organ.
The mantle is highly muscular. In cephalopods the contraction of the mantle is used to force water through a tubular
siphon, the hyponome, and this propels the animal very rapidly through the water. In other mollusks, it is used as a
kind of "foot" for locomotion.
Mantle 108
Formation of mollusc shell

In shelled molluscs, the mantle is what forms the shell, and what adds to the shell to increase its size and strength as
the animal grows. Shell material is secreted by the ectodermic (epithelial) cells of the mantle tissue.[1]
Mantle of gastropods
The mantle of many gastropods is usually fully or partially hidden inside the gastropod shell.
The marine gastropod Cypraea chinensis, the Cypraea chinensis with its mantle fully The mantle of the land snail Indrella
Chinese cowry, showing partially extended extended. ampulla is off-white in color and it is partly
mantle. visible under the shell. The head and foot
are red, and the foot fringe is off-white with
narrow black lines.
In species where the shell is small compared to the size of the body, more of the mantle shows. Shell-less slugs have
the mantle fully visible. The dorsal surface of the mantle is called the notum, while the ventral surface of the mantle
is called the hyponotum. In the family Philomycidae, the mantle covers the whole back side of the body.[2]
The mantle and the head of this slug Bielzia Megapallifera mutabilis from Philomycidae shows enormously
coerulans is smooth, while the rest of the body is developed mantle
tubercled.
Mantle 109
This drawing shows that the mantle (in gray)

Photo of Haliotis asinina with the shell removed.
covers the majority of the dorsal surface of the
[3]
animal.
See also
• Mollusc shell, which is formed by the mantle
• Siphon (mollusc) which is a part of the mantle in some groups of mollusks
References
[1] "integument (mollusks)."Encyclopædia Britannica. 2009. Encyclopædia Britannica 2006 Ultimate Reference Suite DVD
[2] Tsai C.-L. & Wu S.-K. (2008). "A New Meghimatium Slug (Pulmonata: Philomycidae) from Taiwan". Zoological Studies 47(6): 759-766.
PDF (http:/ / zoolstud. sinica. edu. tw/ Journals/ 47. 6/ 759. pdf).
[3] Daniel J Jackson, Carmel McDougall, Kathryn Green, Fiona Simpson, Gert Wörheide & Bernard M Degnan. 2006. http:/ / www.
biomedcentral. com/ 1741-7007/ 4/ 40 A rapidly evolving secretome builds and patterns a sea shell]. BMC Biology 2006, 4:4.
0doi:10.1186/1741-7007-4-40.
Nidamental gland 110
Nidamental gland
The nidamental gland is an internal organ found in
some elasmobranchs and certain molluscs, including
cephalopods (specifically Decapodiformes and
nautiluses) and gastropods.[1] [2] [3]
In cephalopods, the nidamental gland is a large
glandular structure found in the mantle cavity. An
accessory nidamental gland may also be present.
Nidamental glands are composed of lamellae which
Ventral view of the viscera of Chtenopteryx sicula, showing the
secrete egg cases or the gelatinous substance presence of the nidamental gland and accessory nidamental gland.
comprising egg masses.[1]
References
[1] Young, R.E., M. Vecchione & K.M. Mangold 1999. Cephalopoda Glossary (http:/ / tolweb. org/ accessory/
Cephalopoda_Glossary?acc_id=587). Tree of Life web project.
[2] Prasad, R.R. (1948). "Observations on the Nidamental Glands of Hydrolagus colliei, Raja rhina and Platyrhinoidis triseriatus" (http:/ / www.
jstor. org/ pss/ 1438791). Copeia 1948 (1): 54–7. doi:10.2307/1438791. .
[3] Bloodgood RA (1977). "The squid accessory nidamental gland: ultrastructure and association with bacteria" (http:/ / linkinghub. elsevier.
com/ retrieve/ pii/ 0040-8166(77)90016-7). Tissue Cell 9 (2): 197–208. PMID 906013. .
Siphon
A siphon is an anatomical structure which is part of the soft parts of
aquatic molluscs in three classes: Gastropoda, Bivalvia and
Cephalopoda. In other words, a siphon is found in some saltwater and
freshwater snails, in some clams, and in octopus, squid and relatives.
Siphons are tube-like structures in which water flows (or more rarely in
which air flows). In molluscs, the water flow is used for one or more
purposes such as locomotion, feeding, respiration, and reproduction.
The siphon is part of the mantle of the mollusc, and the water flow is
directed to (or from) the mantle cavity.
A specimen of a venerid bivalve. The adductor
A single siphon occurs in some gastropods. In those bivalves which muscles have been cut, the valves are gaping. The
internal anatomy is visible, including the paired
have siphons, they are paired. In cephalopods, there is a single siphon
siphons to the right
or funnel which is known as a hyponome.
Siphon 111
The siphon of a large carnivorous marine volute, The sea snail Nassarius tiarula is a scavenger.
Cymbiola magnifica Siphon on the left
In some (but not all) sea snails, marine gastropod molluscs, the animal
has an anterior extension of the mantle called a siphon, or inhalant
siphon, through which water is drawn into the mantle cavity and over
the gill for respiration.[1]
This siphon is a soft fleshy tube-like structure equipped with
chemoreceptors which "smell" or "taste" the water, in order to hunt for
food.[2] [3] [4] Marine gastropods that have a siphon are either predators
or scavengers.[5]
Although in gastropods the siphon functions perfectly well as a tube, it Melo amphora moving across coral at low tide.
is not in fact a hollow organ, it is simply a flap of the mantle that is
rolled into the shape of a tube.[1]
In many marine gastropods where the siphon is particularly long, the structure of the shell has been modified in order
to house and protect the soft tissue of the siphon. This shell modification is known as the siphonal canal. For a
gastropod whose shell has an exceptionally long siphonal canal, see Venus comb murex.
In the case of some other marine gastropod shells, such as the volute and the Nassarius pictured above, the shell has
a simple "siphonal notch" at the anterior edge of the aperture instead of a long siphonal canal.
Aplysia gill and siphon withdrawal reflex is defensive reflex of Aplysia and it is valuable in neuroscience.
Siphon 112
Siphon as a snorkel
Freshwater apple snails in the genera Pomacea and Pila have an
extensible siphon made from a flap of the left mantle cavity. They use
this siphon in order to breathe air while they are submerged in water
which has a low oxygen content so they cannot effectively use their
gill.[6]
Apple snails use the siphon in a way that is reminiscent of a swimmer
using a snorkel, except that the apple snail's siphon can be retracted
completely, or extended to various lengths as needed.[6]
For these freshwater snails, the siphon is an anti-predator adaptation. It
reduces their vulnerability to being attacked and eaten by birds because
it enables the apple snails to breathe without having to come all the
way up to the surface, where they are easily visible to predators.[6]
Pomacea canaliculata, seen
through glass, has reached its The shells of these freshwater snails have simple round apertures; there
siphon up to the water surface to is no special notch for the siphon.
breathe air
Engraving of Florida freshwater applesnail

Pomacea paludosa; siphon on lower right
Siphon 113
The paired siphons of bivalves

Those bivalves that have siphons, have two of them. Not all bivalves
have siphons however: those that live on or above the substrate, as is
the case in scallops, oysters, etc, do not need them. Only those bivalves
that burrow in sediment, and live buried in the sediment, need to use
these tube-like structures. The function of these siphons is to reach up
to the surface of the sediment, so that the animal is able to respire,
feed, and excrete, and also to reproduce.[7] [8]
The deeper a bivalve species lives in the sediment, the longer its
siphons are. Bivalves which have extremely long siphons, like the
Geoducks pictured here, live very deeply buried, and are hard to dig up
when clamming[9] .
Three specimens of Panopea abrupta in a

seafood tank; the paired siphons (or "necks") of
this species can be one meter long
Veneridae with siphons out
Drawing of the venerid Venus verrucosa showing

paired siphons (upper inhalant and lower exhalant
siphon), shell and foot.
Siphon 114
Many bivalves that have siphons can withdraw them completely into
the shell when needed, but this is not true of all species. Bivalves that
can withdraw the siphons into the shell have a "pallial sinus", a sort of
pocket, into which the siphons can fit when they are withdrawn, so that
the two shell valves can close properly. The existence of this pocket
shows even in an empty shell, as a visible indentation in the pallial
line, a line which runs along parallel to the ventral margin of the
shell.[10]
Diagramatic drawing of the inside of one valve of
The bivalve's two siphons are situated at the posterior edge of the
a bivalve such as a venerid: pallial sinus on the
mantle cavity.[11] There is an inhalant or incurrent siphon, and an
lower left, at the posterior end of the clam
exhalant or excurrent siphon.[12] The water is circulated by the action
of the gills. Usually water enters the mantle cavity through the inhalant
siphon, moves over the gills, and leaves through the exhalant siphon. The water current is utilized for respiration, but
also for filter feeding, excretion, and reproduction.
Feeding
Depending on the species and family concerned, some bivalves utilize their inhalant siphon like the hose of a
vacuum cleaner, and actively suck up food particles from the marine substrate. Most other bivalves ingest
microscopic phytoplankton as food from the general water supply, which enters via the inhalant siphon and reaches
the mouth after passing over the gill.[13]
Please also see pseudofeces.
The hyponome of cephalopods

The hyponome or siphon is the organ used by cephalopods to expel
water, a function that produces a locomotive force. The hyponome
developed from the foot of the molluscan ancestor.[14]
Water enters the mantle cavity around the sides of the funnel, and
subsequent contraction of the hyponome expands and then contracts,
expelling a jet of water.
In most cephalopods, such as octopus, squid, and cuttlefish, the
hyponome is a muscular tube. The hyponome of the nautilus differs
however, in that it is a one-piece flap that is folded over. The presence Nautilus belauensis seen from the front, showing
or form of the ammonite hyponome is unknown.[15] the opening of the hyponome.
Siphon 115
References
[1] Örstan A. 13 April 2007. Melongena's siphon (http:/ / snailstales. blogspot. com/ 2007/ 04/ melongenas-siphon. html). Snail's Tales.
[2] Abbott, RT and Sandstrom, GF (2001) Seashells of North America (http:/ / books. google. co. nz/ books?id=i94pFP-Wb8YC& pg=PA142&
lpg=PA142& dq=Siphon+ taste+ smell+ sea|marine+ snail|gastropod& source=web& ots=3BE3ifHUL3&
sig=bwd0bZmDja2tDF-aS-m9Ug9wGY4& hl=en& sa=X& oi=book_result& resnum=9& ct=result) Macmillan. ISBN 9781582381251 Nassa
mud snails, p. 142.
[3] Cone snails (http:/ / www. mauioceancenter. com/ index. php?id=11& ss=0& page=marine& content=marine_detail& cat=2& CRid=21&
limitstart=0). Hawaiian Marine Life. Accessed 18 November 2008.
[4] Respiratory system (http:/ / www. applesnail. net/ content/ anatomy/ respiration. php). The apple snail website. Accessed 18 November 2008.
[5] Los Marineros Marine Life. Caption Mollusca. (http:/ / mrt. tripod. com/ marinelife. html#anchor135905) Accessed 21 November 2008.
[6] Respiratory system (http:/ / www. applesnail. net/ content/ anatomy/ respiration. php). The apple snail website, http:/ / www. applesnail. net,
accessed 26 February 2009.
[7] Bales, SL and Venable, S. 2007. Natural Histories: Stories from the Tennessee Valley (http:/ / books. google. co. nz/
books?id=aIZmfXFGydwC& pg=PA66& lpg=PA66& dq=siphon+ bivalve+ reproduction& source=web& ots=2HSA3zeFIs&
sig=WUMp4q52BuiZKw6Iz7Xlv1Z2XtI& hl=en& sa=X& oi=book_result& resnum=17& ct=result). University of Tennessee Press. ISBN
9781572335615. p. 66.
[8] Barnes, H. (Ed.) 2008. Oceanography and Marine Biology (http:/ / books. google. co. nz/ books?id=h2BZZcjAdnkC& pg=PA77&
lpg=PA77& dq=siphon+ bivalve+ reproduction& source=web& ots=0jd1EPDq19& sig=kFZiiEIDFAiYfXT9AONg9UD-9FQ& hl=en&
sa=X& oi=book_result& resnum=22& ct=result) CRC Press. ISBN 9781420065749. p. 77.
[9] Washington Department of Fish and Wildlife. 2000. WDFW - Shellfish: Geoduck clam (http:/ / www. wdfw. wa. gov/ fish/ shelfish/ beachreg/
2clam. htm). accessed 26 February 2009.
[10] M. Alan Kazlev. Palaeos Metazoa: Mollusca: Bivalvia: Bivalve Glossary (http:/ / www. palaeos. com/ Invertebrates/ Molluscs/ Bivalvia/
bivalgloss. html). Page uploaded 11 January 2003, last change 7 July 2007, accessed 26 February 2009.
[11] Anatomy of a Bivalve (http:/ / www. marietta. edu/ ~biol/ mussels/ anatomy. html). accessed 26 February 2009.
[12] Siphons (http:/ / www. marietta. edu/ ~biol/ mussels/ 11. html). accessed 26 February 2009.
[13] S. Peter Dance. 1977. The Encyclopedia of Shells. Blandford Press Limited, Poole, Dorset, ISBN 0-7137-0698-8, pp. 288, page 8.
[14] Class Cephalopoda: the Head-Feet (http:/ / userwww. sfsu. edu/ ~biol240/ labs/ lab_18molluscs/ pages/ cephalopods. html) Accessed 21
November 2008.
[15] Discussion (http:/ / palaeo-electronica. org/ 1998_1/ monks/ discus. htm). http:/ / palaeo-electronica. org/ Accessed 21 November 2008.
External links
• Glossary (http://www.manandmollusc.net/glossary.html)
• Bivalve anatomy (http://www.assateague.com/nt-bival.html)
• More bivalve anatomy (http://www.marietta.edu/~biol/mussels/anatomy.html)
Squid giant axon 116
Squid giant axon

The squid giant axon is the very large (up to 1 mm in diameter; typically around 0.5 mm) axon that controls part of
the water jet propulsion system in squid. It was discovered by English zoologist and neurophysiologist John Zachary
Young in 1936. Squid use this system primarily for making brief but very fast movements through the water.
Between the tentacles of a squid is a siphon through which water can be rapidly expelled by the fast contractions of
the body wall muscles of the animal. This contraction is initiated by action potentials in the giant axon. Action
potentials travel faster[1] in a larger axon than a smaller one, and squid have evolved the giant axon to improve the
speed of their escape response. The increased diameter of the squid axon decreases the internal resistance of the
axon, as resistivity is inversely proportional to the cross sectional area of the object. This increases the space
constant, λ=sqrt(rm/ri). The increased space constant propagates a given local depolarization further, which speeds up
the action potential, according to the equation E=Eoe(-x/λ).
In their Nobel Prize-winning work uncovering ionic mechanism of action potentials, Alan Hodgkin and Andrew
Huxley performed experiments on the squid giant axon. The prize was shared with John Eccles. The large diameter
of the axon provided a great experimental advantage for Hodgkin and Huxley as it allowed them to insert voltage
clamp electrodes inside the lumen of the axon.
While the squid axon is very large in diameter it is unmyelinated which decreases the conduction velocity potential
substantially. The conduction velocity of a typical 0.5 mm squid axon is about 25 m/s. During a typical action
potential in the cuttlefish Sepia giant axon, an influx of 3.7 pmol/cm2 (picomoles per centimeter2) of sodium is offset
by a subsequent efflux of 4.3 pmol/cm2 of potassium.[2]
References
[1] Gazzaniga, Ivry and Mangun 1998, Cognitive Neuroscience - The biology of mind
[2] Plonsey, Robert; Roger C. Barr (2007). Bioelectricity: A Quantitative Approach, 3rd. Edition. New York, NY: Springer. pp. 109.
ISBN 978-0-387-48864-6.
117
Cephalopod shells
Cuttlebone
Cuttlebone, also known as cuttlefish bone, is a hard, brittle internal
structure found in all members of the family Sepiidae, commonly
known as cuttlefish.
Cuttlebone is composed primarily of aragonite. It is a chambered,
gas-filled shell used for buoyancy control; its siphuncle is highly
modified and is on the ventral side of the shell.[1] The microscopic
structure of cuttlebone consists of narrow layers connected by Cuttlebone from Sepia sp.
numerous upright pillars.
Depending on the species, cuttlebones implode at a depth of 200 to 600
metres (660 to 2000 ft). Because of this limitation, most species of
cuttlefish live on the seafloor in shallow water, usually on the
continental shelf.[2]
Human uses
In the past, cuttlebones were used in making polishing powder. The
powder was added to toothpaste, and used as an antacid or as an
absorbent.
Today, cuttlebones are commonly used as calcium-rich dietary
Cuttlebone of Sepia officinalis
supplements for caged birds, chinchillas, hermit crabs, snails, and
turtles.[3]
Jewelry making
Because cuttlebone is able to withstand high temperatures and is easily carved, it serves as mold-making material for
small metal casting for the creation of jewelry and small sculptural objects.
Jewelers prepare cuttlebone for use as a mold by cutting it in half and rubbing the two sides together until they fit
flush against one another. Then the casting can be done by carving a design into the cuttlebone, adding the necessary
sprue, melting the metal in a separate pouring crucible, and pouring the molten metal into the mold through the
sprue. Finally, the sprue is sawed off and the finished piece is polished.[4]
Cuttlebone 118
References
[1] Rexfort, A.; Mutterlose, J. (2006). "Stable isotope records from Sepia officinalis—a key to understanding the ecology of belemnites?". Earth
and Planetary Science Letters 247: 212–212. doi:10.1016/j.epsl.2006.04.025.
[2] Norman, M.D. 2000. Cephalopods: A World Guide. ConchBooks.
[3] Norman, M.D. & A. Reid 2000. A Guide to Squid, Cuttlefish and Octopuses of Australasia. CSIRO Publishing.
[4] Casting Silver with Cuttlefish (http:/ / www. silverstall. com/ casting-silver-jewellery. html)
• Neige, P. 2003. Combining disparity with diversity to study the biogeographic pattern of Sepiidae. (http://
biogeosciences.u-bourgogne.fr/cv/neige/PDF/NeigeBerlin.pdf)PDF Berliner Paläobiologische Abhandlungen
3: 189–197.
Septum
Septa (singular septum) are thin walls or partitions between the
internal chambers (camerae) of the shell of a cephalopod, namely
nautiloids or ammonoids.
As the creature grows, its body moves forward in the shell to a new
living chamber, secreting septa behind it. This adds new chambers to
the shell, which can be clearly seen in cross-sections of the shell of the
living nautilus, or in ammonoid and nautiloid fossils. The septa are
attached to the inside wall of the shell, thus dividing the phragmocone
into camerae.
Cutaway of a nautilus shell showing the
Where the septum meets the shell a suture line forms; in some chambers
ammonoids these lines became extremely complex and elaborate,

providing strength without the necessity of added weight. Elaborate sutures allowed for thinner shells, and hence less
time needed for shell growth and less time spent in the vulnerable juvenile stage.
The nature and structure of the septa, as with the camerae, and siphuncle, and the presence or absence of deposits,
are important in classification of nautiloids. In some nautiloids, such as the Orthoceratidae, the septa tend to be
widely spaced, resulting in large, long camarae. In others such as the Ellesmerocerida, Oncocerida and Discosorida
the septa are crowded closely together. In some straight-shelled forms like Actinoceras, calcium carbonate deposits
extend from the camera (mural deposits) to the septa (episeptal deposits).
Septa can be used to identify extinct coral Order Rugosa and may be arranged as cardinal proseptum, counter
proseptum, alar proseptum orcounter lateral proseptum.
Aptychus 119
Aptychus
An aptychus is a type of marine fossil, a hard anatomical structure like
a curved shelly plate, which was part of the body of an ammonite.
Paired aptychi have, on rare occasions, been found at or within the
aperture of ammonite shells.
Aptychi are often found well-preserved as fossils, but are only very
rarely found connected to ammonite shells. This circumstance led to
them being initially classified as the valves of bivalves, which they
somewhat resemble. Aptychi are found in rocks from the Devonian
period through to the those of the Cretaceous period. The aptychus was
usually composed of calcite, whereas the ammonite shell was
aragonite.
A pair of fossil aptychi. The picture is 1 cm
There are many forms of aptychus, varying in shape and in the
across.
sculpture of the inner and outer surfaces. However, because they are so
rarely found in position within the ammonite shell, it is often unclear as
to which species of ammonite many aptychi belong.
When only a single plate is present, as is sometimes the case, the term
"anaptycus" is used.
Function
Aptychi seem to have most often existed as bilaterally-symmetrical
pairs, and were first described (incorrectly) as being the valves of
bivalve mollusks. Aptychi are now considered to be either: (1) a
two-valved closing hatch on the shells of extinct ammonites; or (2) a One of what would have been a pair of aptychi,
(at first given the name "Trigonellites latus" and
double-plate jaw-piece similar to that of some modern cephalopods.[1]
[2] [3] [4] described as a bivalve) from the Kimmeridge
Clay Formation in England
Set close to or against the shell's terminal opening (the living

chamber), the aptychi usually consisted of two identical but mirror image valves. Some authors consider the
aptychus to be a jaw apparatus (mandibles), while others believe them to be paired opercula. If the latter is the case,
then aptychi may have had a function similar to the head shield of modern nautiluses.
References
[1] Morton, N. 1981. Aptychi: the myth of the ammonite operculum. Lethaia 14(1): 57–61. doi:10.1111/j.1502-3931.1981.tb01074.x
[2] Morton, N. & M. Nixon 1987. Size and function of ammonite aptychi in comparison with buccal masses of modem cephalopods. Lethaia
20(3): 231–238. doi:10.1111/j.1502-3931.1987.tb02043.x
[3] Lehmann, U. & C. Kulicki 1990. Double function of aptychi (Ammonoidea) as jaw elements and opercula. Lethaia 23: 325–331.
doi:10.1111/j.1502-3931.1990.tb01365.x
[4] Seilacher, A. 1993. Ammonite aptychi; how to transform a jaw into an operculum? American Journal of Science 293: 20–32.
doi:10.2475/ajs.293.A.20
• Morphological Terminology: the Aptychus (http://www.calfrye.com/aptychi/morphological_terminology.

htm) from "North American Late Devonian Cephalopod Aptychi". Kirtlandia 46 (August 1991):49-71. By Calvin
J. Frye and Rodney M. Feldmann.
Orthocone 120
Orthocone
An orthocone is a usually long straight shell of a
nautiloid cephalopod. During the 18th and 19th
centuries, all shells of this type were named
Orthoceras, but it is now known that many groups of
nautiloids developed or retained this type of shell.
An orthocone can be thought of as like a Nautilus shell,
but with the shell straight and uncoiled. It was
previously believed that these represented the most
primitive form of nautiloid, but it is now known that
the earliest nautiloids had shells that were slightly Fossilised Orthoceras orthocones.
curved. An orthoconic form evolved several times

among cephalopods, and among nautiloid cephalopods is prevalent among the ellesmerocerids, endocerids,
actinocerids, orthocerids, and bactritids.
Orthocones existed from the Late Cambrian to the Late Triassic, but they were most common in the early Paleozoic.
Revivals of the orthocone design later occurred in other cephalopod groups, notably baculitid ammonites in the
Cretaceous Period. Orthocone nautiloids range in size from less than an inch to (in some giant endocerids of the
Ordovician) seventeen feet (or five meters) long.
Phragmocone
The phragmocone is the chambered portion of the shell of a
cephalopod. It is divided by septa into camerae.
In most nautiloids and ammonoids, the phragmocone is a long, straight,
curved, or coiled structure, in which the camarae are linked by a
siphuncle which determines buoyancy by means of gas exchange.
Despite this benefit, such a large shell adds to the mass of the animal,
and hence is not advantageous in catching fast-moving prey. Some
nautiloids, such as the Silurian Ascocerida, dropped the phragmocone
upon maturity, presumably to increase speed and maneuverability. Cutaway of a nautilus shell showing the
They thus became the early Paleozoic equivalent of coleoids. The early chambers
coleoids and belemnoids adopted a different approach. The
phragmocone was retained but became internal and reduced, and so like the shell in general it tends to be vestigial or
absent in most cephalopods.
Phragmocone 121
Fossil record
Being the only biomineralised part of most cephalopods, the phragmocone is typically the only part to enter the fossil
record. It is sometimes infilled with sediment, with sediment presumably getting in through the siphuncle.[1] There
are occasions where trilobites have been preserved within phragmocones, presumably where they crawled in for
refuge.[2]
References
[1] Henderson, ROBERT A.; McNamara, Kenneth J. (1985). "Taphonomy and ichnology of cephalopod shells in a Maastrichtian chalk from
Western Australia". Lethaia 18: 305. doi:10.1111/j.1502-3931.1985.tb00710.x.
[2] Arnold Davis, R. H. B. Fraaye, Char, Richard (2001). "Trilobites within nautiloid cephalopods". Lethaia 34: 37.
doi:10.1080/002411601300068251.
Siphuncle
The siphuncle is a strand of tissue passing longitudinally through the shell of a cephalopod mollusk. Only
cephalopods with chambered shells have siphuncles, such as the extinct ammonites and belemnites, and the living
nautiluses, cuttlefish, and Spirula. In the case of the cuttlefish, the siphuncle is indistinct and connects all the small
chambers of that animal's highly modified shell; in the other cephalopods it is thread-like and passes through small
openings in the walls dividing the chambers.
The siphuncle is used primarily in emptying water from new chambers as the shell grows.[1] Essentially what
happens is the cephalopod increases the saltiness of the blood in the siphuncle, and the water moves from the more
dilute chamber into the blood through osmosis. At the same time gas, mostly nitrogen, oxygen, and carbon dioxide,
diffuses from the blood in the siphuncle into the emptying chamber. Note that the cephalopod does not pump up the
shell; the gas moving into the chamber is a passive process, instead the energy is used in absorbing the water from
the chamber.
Removing water from the chambers of the shell reduces the overall
density of the shell, and thus the shell behaves as a flotation device
comparable to the swim bladder in bony fish. Typically, cephalopods
maintain a density close to that of sea water, allowing them to swim
with the minimum of effort. In the geologic past, many cephalopods
grew to an enormous size (over ten meters in length) thanks to this.
Generally, the siphuncle is unable to provide a way to change the

density of shell rapidly and thus cause the animal to rise or sink at will;
rather, the animal must swim up or down as required.
The siphuncle found in fossilised cephalopods is assumed to have
worked in the same general way. The siphuncle itself only rarely gets
An image showing the siphuncle, the tube which
preserved, but many fossils show the holes, called septal necks (or
connects the current living shell to the previous
siphuncle notches), through which the siphuncle passed. In most fossil ones.
nautiluses, the siphuncle runs more or less through the center of each
chamber, but in ammonites and belemnites it usually runs along the ventral surface. In some fossil straight shelled
nautiluses cylindrical calcareous growths ("siphuncular deposits") around the siphuncle can be seen towards the apex
of the shell. These were apparently counterweights for the soft body at the other end of the shell, and allowed the
nautilus to swim in a horizontal position. Without these deposits, the apex of the buoyant shell would have pointed
upwards and the heavier body downwards, making horizontal swimming difficult. The siphuncle of the Endocerida
also contained much of the organisms' body organs.[2]
Siphuncle 122
See also
• Phragmocone
• Orthoceras
• Baculites
References
[1] Mutvei, Harry; Zhang, Yun-bai; Dunca, Elena (2007). "Late Cambrian Plectronocerid Nautiloids and Their Role in Cephalopod Evolution".
Palaeontology 50 (6): 1327–1333. doi:10.1111/j.1475-4983.2007.00708.x
[2] Kroger, B; Yun-Bai, Zhang (2008). "Pulsed cephalopod diversification during the Ordovician". Palaeogeography Palaeoclimatology
Palaeoecology 273: 174. doi:10.1016/j.palaeo.2008.12.015.
Body chamber
The body chamber, also called the living chamber, is the outermost
or last chamber in the shell of a nautiloid or ammonoid cephalopod.
The body of the animal occupies the living chamber, apart from the
siphuncle which extends through the rest of septa (the phragmocone) to
provide buoyancy.
Nautilus pompilius
Article Sources and Contributors

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Frodesiak, Argo Navis, Arkuat, ArthurWeasley, Axl, AzaToth, Bender235, Bewareofdog, Bob the ducq, Bobo192, Brandon5485, Brothejr, CN0t3, CanadianLinuxUser, Cedders, Celithemis,
Ceph, Cephal-odd, Cs california, Da Joe, Dan D. Ric, Daniel.Cardenas, DanielCD, Dbratland, Debresser, Derek Ross, Dfred, Dinoguy2, Dlloyd, DocWatson42, Dominus, Dorathaexplora,
Dromiceiomimus, Dunning, Dureo, Dysmorodrepanis, Elassint, Eluchil404, Epolk, Eregli bob, Erianna, Estudiarme, Flutelover2008, Flyguy649, Fordmadoxfraud, FrancescaSF, Fubar Obfusco,
Funandtrvl, Gadfium, Gary2863, Gentgeen, Georgehenries, Gf2tw, Glenn, Gnomon, GoingBatty, Hadal, HaeB, Headbomb, Hephaestos, IRP, Imo is emo, Inrerumnatura, Ipatrol, IronChris,
Iustinus, J.H.McDonnell, J.delanoy, JNShutt, Jackhynes, Jamesofur, Jarry1250, Jerkov, Jimp, Jklin, John, John254, JohnCD, Jonadab, Jones8888, K-22-22, Katharineamy, Kazvorpal,
Kembangraps, Kevmin, Kialari, Kils, Kipoc, Kleopatra, Kmg90, Kungfuadam, Lando5, Luigifan, Macdonald-ross, Mani1, Mark K. Jensen, Marskell, Mav, Maxim Razin, Maxinuk, Mehrunes
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Nuvitauy07, Omnipedian, Orangesquid, Paul Ferber, Peter, Peter b, Pgan002, Philip Trueman, Physis, PierreAbbat, Pinkpedaller, Polarpanda, Postdlf, Quantumobserver, Quintote, Qworensk,
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Nautiloid Source: http://en.wikipedia.org/w/index.php?oldid=411746505 Contributors: Abyssal, Adrian.benko, Apokryltaros, Bactrite, Bayle Shanks, Bigfun, Bobo192, Brandizzi, Causantin,
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Hocevar, Schreibergasse, Skapur, Smith609, Spellmaster, StarManta, SteveSims, T callahan, The Rationalist, Tom harrison, UtherSRG, Wilson44691, Ziggy Sawdust, Zploskey, ‫ةيناريد دابع‬, 50
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Nautilus Source: http://en.wikipedia.org/w/index.php?oldid=412562056 Contributors: A. B., ARUNKUMAR P.R, Abigail-II, Adrian.benko, After Midnight, Alansohn, Alex.tan, Altenmann,
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Hadal, Headbomb, Hemoptysis, Hongooi, IanOsgood, Invertzoo, Island, Jackthesnake, Jakewaage, Jalwikip, JayEsJay, Jedi of redwall, Johnbod, Juliancolton, Junglecat, Kariteh, Kelson,
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harrison, Toytoy, Travis.barnett33, Trento54, Trusilver, UtherSRG, Valodzka, Vinne2, Visik, Ward3001, Weft, Wimt, Winhunter, Xneonvisionx, Zoney, 219 anonymous edits
Ammonite Source: http://en.wikipedia.org/w/index.php?oldid=412472112 Contributors: !!, Abigail-II, Abyssal, Adashiel, Adeliine, Adrian.benko, Amezcackle, Animum, AnonMoos,
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Spencer, J.H.McDonnell, Jacen Aratan, JamesPFisherIII, Jimp, Jiy, JoanneB, Johnelson, Jonathan350, Jonathunder, Josh Grosse, Jusdafax, Karmosin, Kevmin, Khfan93, Kleopatra, Kukini,
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Vsmith, Whatnwas, Whoelius, Wikipelli, Wilson2, Wilson44691, WolfmanSF, Xana500, Zzzzz, ‫ةيناريد دابع‬, 194 anonymous edits
Belemnoidea Source: http://en.wikipedia.org/w/index.php?oldid=412475067 Contributors: Abyssal, Adrian.benko, Aiko, Anthony Appleyard, Arpingstone, DanielCD, Dlloyd,
Dysmorodrepanis, Felagund, FunkMonk, Headbomb, Huttarl, Invertzoo, J. Spencer, Jashiin, Katharineamy, Kencasey98, Kevmin, Marcelo-Silva, Mgiganteus1, Moon&Nature, NatureA16, Neale
Monks, Obsidian Soul, Polarpanda, Rich Farmbrough, Richardleaky, Saintrain, Slawojarek, Smith609, UtherSRG, WOSlinker, Wetman, Wilson44691, WolfmanSF, ZeldaGamer31337, Zvar,
‫ةيناريد دابع‬, 32 anonymous edits
Argonaut Source: http://en.wikipedia.org/w/index.php?oldid=407759059 Contributors: Abigail-II, Abyssal, Altenmann, Anthony Appleyard, Cakehelmit, CatherineMunro, Cewvero, Citron,
D6, DanielCD, Dannajoy, Dysmorodrepanis, GrahamBould, Guidod, Headbomb, Iciac, InfernoXV, Invertzoo, Koven.rm, Kwamikagami, LilHelpa, Limonadis, M-le-mot-dit, Mgiganteus,
Mgiganteus1, Montyy0, Neale Monks, Orangemarlin, Palpalpalpal, Peter Horn, Phphello, PurpleHz, Qwertzy2, Rajah, Rama, Rolf Schmidt, SteveSims, Template namespace initialisation script,
TheAlphaWolf, Tonyrex, UtherSRG, Wetman, XQ fan, 18 anonymous edits
Cephalopod intelligence Source: http://en.wikipedia.org/w/index.php?oldid=406848079 Contributors: Acaeton, Alan Liefting, Andy Dingley, Animalplanetemployee, Ashesofoak, Atarr, Ceph,
ChildofMidnight, Coelacan, Coemgenus, Delicious carbuncle, Dentren, Eaefremov, Epastore, Epipelagic, Falcon982, Headbomb, HisSpaceResearch, Iam3dhomer, Invertzoo, Jefffire, Jonkerz,
Karlchwe, Kayau, LittleHow, Luigifan, Maproom, Marskell, MatthiasKabel, Mgiganteus1, Montyy0, Moshe Constantine Hassan Al-Silverburg, NEMT, Neurosojourn, Nibuod, Outriggr, Philip
Trueman, Proxima Centauri, Rjwilmsi, Serendipodous, Seric2, Simon80, Spotty11222, SteveSims, Steven Walling, Thumperward, Tktktk, Wood Thrush, Xanthine, 66 anonymous edits
Cephalopod size Source: http://en.wikipedia.org/w/index.php?oldid=408489088 Contributors: Archelon, Cadwaladr, Dysmorodrepanis, Headbomb, J Milburn, Lambyte, Mattg82, Mgiganteus1,
PenguinJockey, Rjwilmsi, Shaundakulbara, Taollan82, Twas Now, Uppland, Zuzster, 18 anonymous edits
Cephalopod ink Source: http://en.wikipedia.org/w/index.php?oldid=407130384 Contributors: Anxietycello, Cedders, Headbomb, Ignacio Bibcraft, Igodard, Intelligentsium, K-22-22,
Mgiganteus1, Scotsman240363, The Thing That Should Not Be, 9 anonymous edits
Ink sac Source: http://en.wikipedia.org/w/index.php?oldid=405070305 Contributors: Brz7, Cdcdoc, Crazycomputers, EncycloPetey, GL, Iciac, Jcvamp, Jessicapierce, Jni, Jwbarnes,
Mgiganteus1, Mukadderat, Skynoceanna, Smith609, Stevenharrower, UtherSRG, 17 anonymous edits
Cephalopod arm Source: http://en.wikipedia.org/w/index.php?oldid=407805213 Contributors: Berek, Combatking0, Headbomb, Henryhartley, Materialscientist, Mgiganteus1, Montecarlocars,
Stemonitis, Yobmod, ZayZayEM, 14 anonymous edits
Hectocotylus Source: http://en.wikipedia.org/w/index.php?oldid=387215087 Contributors: Altenmann, Bansp, DanielCD, DragonflySixtyseven, Ginkgo100, Ickle Ronnie, Imnowei, Jpgordon,
Kasyapa, MacRusgail, Macdonald-ross, Mgiganteus1, Ospalh, Robofish, UtherSRG, ZayZayEM, Zuzster, 5 anonymous edits
Tentacle Source: http://en.wikipedia.org/w/index.php?oldid=394274034 Contributors: Andycjp, Arrowned, Bioform 1234, Bobo192, Bogey97, Bongwarrior, Burmiester, Cflm001, Chris Roy,
Chrislk02, Clicketyclack, Delldot, Doulos Christos, Dragon's Light, E. Fokker, Ekko, Excirial, Ezzex, Facts707, Finn-Zoltan, Gadfium, Geary, Gene Nygaard, Gimme danger, Goodnightmush,
GrahamBould, Gurko, Invertzoo, Isnow, JenniferGan, Joaoantonio, Jobba, John, KnightRider, Kodemage, Lcarscad, Legoo15, Liu Bei, Lokicarbis, LorenzoB, Lugnuts, Lycaon, MOGoss,
Materialscientist, Mgiganteus1, Mmxx, Mnj21, Mohammedh15, Narvalo, NawlinWiki, Neutrality, No more bongos, Noloop, Okc, Omegatron, Owen, PhilHibbs, Piano non troppo, Pinkadelica,
Proxima Centauri, PurpleHz, Quistnix, Reywas92, Rkitko, Rlandmann, Rodhullandemu, Rogutaan, Rossj81, ScottishGuy, Shoeofdeath, Silly rabbit, Skittle, Tavilis, The High Fin Sperm Whale,
TheKMan, TheMadBaron, TheRanger, UnDeRsCoRe, Username381, Uyanga, Verbosemjp, Wombatcat, Woohookitty, Wyndclaw, Yair rand, Yobmod, ZayZayEM, 125 anonymous edits
Dactylus Source: http://en.wikipedia.org/w/index.php?oldid=404288285 Contributors: Agamemnon2, Athurber, Kaarel, Mgiganteus1, Nono64, Pgan002, 1 anonymous edits
Cephalopod eye Source: http://en.wikipedia.org/w/index.php?oldid=377389469 Contributors: Headbomb, Iciac, Komodo, Mgiganteus1, Smith609, Woohookitty, 2 anonymous edits
Chromatophore Source: http://en.wikipedia.org/w/index.php?oldid=407119547 Contributors: Adamharvey182, AdultSwim, Alphachimp, Anville, Axeman89, BD2412, BhaiSaab, Bobo192,
Boghog, BorgQueen, Brighterorange, Caesar Rodney, Ceyockey, Cinchjt, Dark Shikari, DeansFA, Deviator13, Dicklyon, Dispenser, Dj Capricorn, Djanvk, Dominus, DragonflySixtyseven,
Drummie06, EagleFan, Edgar181, Ejdzej, Element16, Emvee, Floyd Landis, Gadfium, Gamesmasterg9, GregorB, Gudeldar, Harmil, Hegar, ISpamThisSite, Icey, InvictaHOG, Jaibobs108,
Joyous!, Jpatokal, Kapuchinski, KimvdLinde, Kukini, LANA2007, Mav, Melchoir, MisfitToys, Mithril, Mr. Blackout, Nono64, NorwegianBlue, NuclearWarfare, P-Chan, PGWG, Pajast, Paul
venter, Peter Delmonte, Philip Trueman, Raul654, Reywas92, Rich Farmbrough, Rintrah, Rjwilmsi, Robert Brockway, RobertG, Rockpocket, RupertMillard, Ryulong, Samir, Sandip90, Saravask,
Savidan, Serephine, Stemonitis, Sucoyant, Super cyclist, Szquirrel, ToNToNi, Tpbradbury, Trevor MacInnis, Una Smith, Utcursch, UtherSRG, WoWFanatic, WolfmanSF, Zafiroblue05, 74
anonymous edits
Mantle Source: http://en.wikipedia.org/w/index.php?oldid=412714375 Contributors: Alexei Kouprianov, Altenmann, Barbara Shack, Brim, Daniel, DanielCD, Hadal, Hofoen, IlSoge,
Invertzoo, Jamoche, Jimmy Figsworth, JoJan, Jomegat, Kaarel, MacGyverMagic, Mgiganteus1, Mild Bill Hiccup, Mr. Billion, Nuvitauy07, Pharos, Potatoswatter, Rettetast, SF007, Secretlondon,
Serlin, Sidhekin, Snek01, Stfg, TheLimbicOne, TimBentley, UtherSRG, Versus22, Voyagerfan5761, Wickey-nl, 34 anonymous edits
Nidamental gland Source: http://en.wikipedia.org/w/index.php?oldid=404289268 Contributors: Mgiganteus1, RDBrown, Snek01, 2 anonymous edits
Siphon Source: http://en.wikipedia.org/w/index.php?oldid=404262442 Contributors: Amikake3, Arct, DanielCD, Epipelagic, Excirial, Invertzoo, Isthmus, J04n, JamesAM, Jeff G., Kaarel,
Kesal, Mgiganteus1, Mukadderat, Nono64, Rror, Smith609, Snek01, Tabletop, UtherSRG, 14 anonymous edits
Squid giant axon Source: http://en.wikipedia.org/w/index.php?oldid=407999385 Contributors: Alkaloids, Altenmann, Alvis, Diberri, Kalexander, MementoVivere, Mgiganteus1, Nrets,
Orlandoturner, Pmjboyle, Synaptidude, Welsh, Whosasking, Wiseoldman123, Woreno, 14 anonymous edits
Cuttlebone Source: http://en.wikipedia.org/w/index.php?oldid=397984794 Contributors: Darklilac, DavidFarmbrough, Dawynn, Dentren, Invertzoo, Lfstevens, Mervyn, Mgiganteus1,
Obsidianearth, Paxsimius, Reywas92, Smith609, Speciate, Thumperward, 21 anonymous edits
Septum Source: http://en.wikipedia.org/w/index.php?oldid=404262711 Contributors: Bobo192, DanielCD, Elkman, JHunterJ, Kaarel, M Alan Kazlev, Mgiganteus1, Nono64, Randomfrenchie,
1 anonymous edits
Aptychus Source: http://en.wikipedia.org/w/index.php?oldid=386681050 Contributors: DanielCD, Invertzoo, Kontos, Mgiganteus1, Snek01, UtherSRG
Orthocone Source: http://en.wikipedia.org/w/index.php?oldid=410456513 Contributors: Apokryltaros, Cephal-odd, D053, Everyking, J.H.McDonnell, JukoFF, M Alan Kazlev, Mgiganteus1,
UtherSRG, VanHelsing, 8 anonymous edits
Phragmocone Source: http://en.wikipedia.org/w/index.php?oldid=365143443 Contributors: Cephal-odd, M Alan Kazlev, Mgiganteus1, Michael Devore, Sam Hocevar, Schmiteye, Smith609,
UtherSRG, WolfmanSF, 3 anonymous edits
Siphuncle Source: http://en.wikipedia.org/w/index.php?oldid=376016720 Contributors: Bucephalus, DanielCD, Hephaestos, Iciac, Mgiganteus1, Neale Monks, P4en, Pazuzu413, Smith609,
Stewartadcock, UtherSRG, 7 anonymous edits
Body chamber Source: http://en.wikipedia.org/w/index.php?oldid=409758989 Contributors: Dawynn, Dina, M Alan Kazlev, Mgiganteus1, UtherSRG
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Cephalopoda

Anatomy and behaviour 78

Cephalopod shells 117

Bigfin reef squid (Sepioteuthis

Subclasses and orders

Nervous system and behaviour

[[file:Chtenopteryx Viscera of Chtenopteryx sicula]][[file:Ocythoe tuberculata Viscera of Ocythoe

Locomotion and buoyancy

Early cephalopods are thought to have produced jets by drawing their

Reproduction and life cycle

The ancestors of coleoids (including most modern

Basal Octopods (e.g. Argonauta)

Heteroteuthis (bobtail squid)

* Certain squid (e.g. Bathyteuthis)

Class Cephalopoda († indicates extinct groups)

Suprafamilial classification of the Treatise

• Order † Orthocerida (482.5 to 211.5 [99] Ma)

Monophyly of coeloids Pyritized fossil of Vampyronassa rhodanica, a vampyromorphid

The Common Octopus, Octopus

An octopus has a highly complex nervous system, only part of which is

In laboratory experiments, octopuses can be readily trained to

An octopus's camouflage is aided by certain specialized skin cells

Octopuses also have an excellent sense of touch. An octopus's suction

Octopuses swim headfirst, with arms trailing

Octopus at Tsukiji fish market

Bigfin Reef Squid, Sepioteuthis

†Plesioteuthididae (incertae sedis)

Modification from ancestral forms

Species Family Common name Catch Percent

Loligo gahi Loliginidae Patagonian squid 24,976 1.1

Loligo pealei Loliginidae Longfin squid 16,684 0.8

[13] Loliginidae 225,958 10.3

Ommastrephes bartramii Ommastrephidae Neon flying squid 22,483 1.0

Illex argentinus Ommastrephidae Argentine shortfin squid 511,087 23.3

Dosidicus gigas Ommastrephidae Jumbo flying squid 406,356 18.6

Todarodes pacificus Ommastrephidae Japanese flying squid 504,438 23.0

Nototoda russloani Ommastrephidae Wellington Flying Squid 62,234 2.8

[13] Various 414,990 18.6

Total squid 2,189,206 100.0

Sepia latimanus, East

Suborders and Families

This Broadclub Cuttlefish (Sepia

Range and habitat

Nautiluses are free swimming animals that possess a

Internally, the shell divides into camerae

The shell coloration also keeps the animal

Buoyancy and movement

In the wild, nautiluses usually inhabit depths of about 300 metres (980

Nautilus with extended tentacles and hyponome

Reproduction and lifespan

fertilized eggs to rocks in shallow waters, whereupon the eggs take

Range and habitat

Dubious or uncertain taxa

N. beccarii Linne, 1758 Non-cephalopod; Foraminifera [fide Frizzell and Keen

N. crispus Linne, 1758 Undetermined Mediterranean Sea Unresolved; Linnean Society of

N. crista Linne, 1758 Non-cephalopod; Turbo [fide Dodge (1953:14)]

N. fascia Linne, 1758 Undetermined Adriatic Sea Unresolved; Linnean Society of

N. granum Linne, 1758 Undetermined Mediterranean Sea Unresolved; Linnean Society of

N. lacustris Lightfoot, 1786 Non-cephalopod; Helix [fide Dillwyn (1817:339)]