Cephalopoda

An overview

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Contents
Articles
Cephalopods
Cephalopod Octopus Squid Cuttlefish Nautiloid Nautilus Ammonite Belemnoidea Argonaut 1 1 25 37 45 52 58 68 79 83 90 90 94 102 104 104 111 114 116 116 118 127 129 130 135 136 136 138 138 141 141

Anatomy and behaviour

Cephalopod intelligence Cephalopod size Cephalopod ink Ink sac Cephalopod arm Hectocotylus Tentacle Dactylus Cephalopod eye Chromatophore Mantle Nidamental gland Siphon Squid giant axon

Cephalopod shells
Cuttlebone Septum Aptychus Orthocone Phragmocone

Siphuncle Body chamber

142 143

References
Article Sources and Contributors Image Sources, Licenses and Contributors 144 147

Article Licenses
License 153

Cephalopods
Cephalopod
Cephalopods

Bigfin reef squid (Sepioteuthis lessoniana)

Scientific classification Kingdom: Phylum: Class: Animalia Mollusca Cephalopoda<br Subclasses and orders

Subclass Nautiloidea Plectronocerida Ellesmerocerida Actinocerida Pseudorthocerida Ascocerida Endocerida Tarphycerida Oncocerida Discosorida Nautilida Orthocerida Lituitida Dissidocerida Bactritida

Cephalopod

Subclass Ammonoidea Goniatitida Prolecanitida Ceratitida Ammonitida

Subclass Coleoidea Belemnoidea Aulacocerida Belemnitida Hematitida Phragmoteuthida Neocoleoidea (most living cephalopods) ?Boletzkyida Sepiida Sepiolida Spirulida Teuthida Octopoda Vampyromorphida

A cephalopod is any member of the molluscan class Cephalopoda (Greek plural (kephalpoda); "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).

Distribution

Cephalopod There are around 800 extant species of cephalopod,[1] although new species continue to be described. An estimated 11,000 extinct taxa have been described, although the soft-bodied nature of cephalopods means they are not easily fossilised.[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.[3] Cephalopods occupy most of the depth of the ocean, from the abyssal plain to the sea surface. Their diversity is greatest near the equator (~40 species retrieved in nets at 11N by a diversity study) and decreases towards the poles (~5 species captured at 60N).[4]

Nervous system and behavior

Cephalopods are widely regarded as the most intelligent of the invertebrates, and have well developed senses and large brains (larger than those 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 brain is protected in a cartilaginous cranium. The giant nerve fibers of the cephalopod mantle have been widely used as experimental material in neurophysiology for many years; their large diameter (due to lack of myelination) makes them relatively easy to study.[6] Cephalopods are social creatures; when isolated from their own kind, they will take to shoaling with fish.[7] Some cephalopods are able to fly through air for distances up to 50m. 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]

Cephalopod

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 construction differs, as cephalopods lack a cornea, and have an everted The primitive nautilus eye functions similarly to a pinhole camera. retina.[9] Cephalopods' eyes are also sensitive to the plane of polarization of light.[10] Surprisinglygiven their ability to change colorall octopuses[11] and most cephalopods[12] 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.[13] They also produce visual pigments throughout their body, and may sense light levels directly from their body.[14] Evidence of color vision has been found in the sparkling enope squid (Watasenia scintillans),[12][15] 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.[16] 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

Hearing
Cephalopods can use their statocysts to detect sound.[17]

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.[18] 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.[19]

Coloration
Cephalopods can change their colours and patterns in milliseconds, whether for signalling or active camouflage,[4] as their chromatophores are expanded or contracted.[19] Coloration is typically stronger in near-shore species than those living in the open ocean, whose functions tend to be restricted to disruptive camouflage.[4]:2

Evidence of original coloration has been detected in cephalopod fossils dating as far back as the Silurian; these orthoconic individuals bore concentric stripes, which are thought to have served as camouflage.[20] Devonian cephalopods bear more complex colour patterns, whose function may be more complex.[21]

This broadclub cuttlefish (Sepia latimanus) can go from camouflage tans and browns (top) to yellow with dark highlights (bottom) in less than a second.

Ink
With the exception of the Nautilidae and the species of octopus belonging to the suborder Cirrina,[22] all known cephalopods have an ink sac, which can be used to expel a cloud of dark ink to confuse predators.[11] 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 the ink can be distributed by ejected water as the cephalopod uses its jet propulsion.[11] 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.[11] 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.

Cephalopod

Viscera of Chtenopteryx sicula

Viscera of Ocythoe tuberculata

Circulatory system
Cephalopods are the only mollusks with a closed circulatory system. Coleoids have two gill hearts (also known as branchial hearts) that move blood through the capillaries of the gills. A single systemic heart then pumps the oxygenated blood through the rest of the body.[23] 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.[24]

Respiration
Cephalopods exchange gasses with the seawater by forcing water through their gills, which are attached to the roof of the organism.[25]:488[26] 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.[26] 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.[25] The water flow is controlled by contractions of the radial and circular mantle cavity muscles.[27] The gills of cephalopods are supported by a skeleton of robust fibrous proteins; the lack of mucopolysaccharides distinguishes this matrix from cartilage.[28][29] The gills are also thought to be involved in excretion, with NH4+ being swapped with K+ from the seawater.[26]

Cephalopod

Locomotion and buoyancy

While most cephalopods can move by jet propulsion, this is a very energy-consuming way to travel compared to the tail propulsion used by fish.[30] The efficiency of a propellor-driven waterjet (i.e. Froude efficiency) is a more efficient model than rocket efficiency.[31] The relative efficiency of jet propulsion decreases further as animal size increases; paralarvae are far more efficient than juvenile and adult individuals.[32] 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 Octopuses swim headfirst, with arms trailing sole mode of locomotion,[2]:208 the stop-start motion provided by the behind jets continues to be useful for providing bursts of high speed - not least when capturing prey or avoiding predators.[2] Indeed, it makes cephalopods the fastest marine invertebrates,[4]:Preface and they can out-accelerate most fish.[25] 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).[32] 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.[33] The velocity of the organism can be accurately predicted for a given mass and morhpology of animal.[34] 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.[35] Some cephalopods accompany this expulsion of water with a gunshot-like popping noise, thought to function to frighten away potential predators.[36] Cephalopods employ a similar method of propulsion despite their increasing size (as they grow) changing the dynamics of the water in which they find themselves. Thus their paralarvae do not extensively use their fins (which are less efficient at low Reynolds numbers) and primarily use their jets to propel themselves upwards, whereas large adult cephalopods tend to swim less efficiently and with more reliance on their fins.[32] Early cephalopods are thought to have produced jets by drawing their body into their shells, as Nautilus does today.[37] 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.[37] The jet velocity in Nautilus is much slower than in coleoids, but less musculature and energy is involved in its production.[38] Jet thrust in cephalopods is controlled primarily by the maximum diameter of the funnel orifice (or, perhaps, the average diameter of the funnel)[39]:440 and the diameter of the mantle cavity.[40] Changes in the size of the Nautilus belauensis seen from the front, showing orifice are used most at intermediate velocities.[39] 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.[39] Water refills the cavity by entering not only through the orifices, but also though the funnel.[39] Squid can expel up to 94% of the fluid within their cavity in a single jet thrust.[31] 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.[39]

Cephalopod 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.[30] 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;[30] others, like some fish, accumulate oils in the liver;[30] and some octopuses have a gelatinous body with lighter chlorine ions replacing sulfate in the body chemistry.[30]

Shell

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

Cuttlebone of Sepia officinalis

Gladius of Sepioteuthis lessoniana

Nautiluses are the only extant cephalopods with 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 (cuttlebone) that is internal in the adult.[41] The same is true of the chitinous gladius of squid[41] and octopus.[42] Cirrate octopuses have cartilaginous fin supports,[43] which are sometimes referred to as a "shell vestige" or "gladius".[44]: The Incirrina have no vestige of an internal shell, and some squid also lack a gladius.[45] Interestingly, the shelled coleoids do not form a clade or even a paraphyletic group.[46] The Spirula shell begins as an organic structure, and is then very rapidly mineralized.[47] Shells that are "lost" may be lost by resorption of the calcium carbonate component.[48] 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.[49]

Cephalopod

Head appendages
Cephalopods, as the name implies, have muscular appendages extending from their heads and surrounding their mouths. These are used in feeding, mobility, and even reproduction. In coleoids they number eight or ten. Decapods such as cuttlefish and squid have five pairs. The longer two, termed tentacles, are actively involved in capturing prey;[]:225 they can lengthen rapidly (in as little as 15 milliseconds[]:225). In giant squid they may reach a length of 8 metres. They may terminate in a broadened, sucker-coated club.[]:225 The shorter four pairs are termed arms, and are involved in holding and manipulating the captured organism.[]:225 They too have suckers, on the side closest to the mouth; these help to hold onto the prey.[]:226 Octopods only have four pairs of sucker-coated arms, as the name suggests, though developmental abnormalities can modify the number of arms expressed.[50] The tentacle consists of a thick central nerve cord (which must be thick to allow each sucker to be controlled independently)[51] 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%.[]:227 The shorter arms lack this capability. The size of the tentacle is related to the size of the buccal cavity; larger, stronger tentacles can hold prey as small bites are taken from it; with more numerous, smaller tentacles, prey is swallowed whole, so the mouth cavity must be larger.[52] Externally shelled nautilids (Nautilus and Allonautilus) have on the order of 90 finger-like appendages, termed tentacles, which lack suckers but are sticky instead, and are partly retractable.

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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[53]:110 They feed by capturing prey with their tentacles, drawing it in to their mouth and taking bites from it.[11] 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.[11] The salivary gland has a small tooth at its end which can be poked into an organism to digest it from within.[11]

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

The digestive gland itself is rather short.[11] It has four elements, with food passing through the crop, stomach and caecum before entering the intestine. Most digestion, as well as the absorption of nutrients, occurs in the digestive gland, sometimes called the liver. Nutrients and waste materials are exchanged between the gut and the digestive gland through a pair of connections linking the gland to the junction of the stomach and caecum.[11] 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.[11] Cephalopods tend to concentrate ingested heavy metals in their body tissue.[54]

Radula
The cephalopod radula consists of multiple symmetrical rows of up to nine teeth[55] thirteen in fossil classes.[56] The organ is reduced or even vestigial in certain octopus species and is absent in Spirula.[56] The teeth may be homodont (i.e. similar in form across a row), heterodont (otherwise), or ctenodont (comb-like).[56] Their height, width and number of cusps is variable between species.[56] 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.[56]:79 Cephalopod radulae are known from fossil deposits dating back to the Amphioctopus marginatus eating a crab Ordovician.[57] They are usually preserved within the cephalopod's body chamber, commonly in conjunction with the mandibles; but this need not always be the case;[58] many radulae are preserved in a range of settings in the Mason Creek.[59] 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,[60] and they are rarer still in non-ammonoid forms: only three pre-Mesozoic species possess one.[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.[61] Nautilus, unusually, possesses four nephridia, none of which are connected to the pericardial cavities.

Cephalopod A detailed discussion on cephalopod feeding strategies and behaviour is available from Rodhouse, P. G.; Nigmatullin, C. M. (1996). "Role as Consumers". Philosophical Transactions of the Royal Society B: Biological Sciences 351 (1343): 1003. doi:10.1098/rstb.1996.0090. and Hanlon, Roger T; Messenger, John B (1998-10-28). "Feeding and Foraging" [62]. Cephalopod Behaviour. pp.4765. ISBN978-0-521-64583-6., which discusses cephalotoxins.

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Ammonium
The handling of ammonia is thought to be important in shell formation in terrestrial molluscs, and in other nonmolluscan lineages.[63] 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.[64] 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 to produce buoyancy.[64] Other cephalopods use ammonium in a similar way, storing the ions (as ammonium chloride) themselves to reduce their overall density and thus become more buoyant.[64]

Reproduction and life cycle

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.[65] Penis elongation in Onykia ingens may result in a penis that is as long as the mantle, head and arms combined.[65][66] 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

Female Argonauta argo with eggcase and eggs

Detail of the hectocotylus of Ocythoe tuberculata

is

Cephalopod

12

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.[67] 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.[67]:227

A dissected male specimen of Onykia ingens, showing a non-erect penis (the white tubular structure located below most of the other organs)

Nidamental glands are involved in the secretion of egg cases or the gelatinous substance comprising egg masses.[68] The eggs may be brooded: female paper nautilus construct a shelter for the young, while Gonatiid squid carry a larva-laden membrane from the A specimen of the same species exhibiting elongation of the penis to hooks on their arms.[69] Other cephalopods deposit 67cm in length 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.[70] Fossilised egg clutches show that ammonites also laid clutches of eggs.[71] 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.[11] Most live for one to two years,[11] reproducing and then dying shortly thereafter.[72] 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.[73]

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.[61][74] The funnel of cephalopods develops on the top of their head, whereas the mouth develops on the opposite surface.[75]:86 The early embryological stages are reminiscent of ancestral gastropods and extant Monoplacophora.[74]
Egg cases laid by a female squid

Cephalopod The shells develop from the ectoderm as an organic framework which is subsequently mineralised.[41] In Sepia, which has an internal shell, the ectoderm forms an invagination whose pore is sealed off before this organic framework is deposited.[41] The gene engrailed is expressed first in the arms, funnel and optic vesicles, and is only later present in the tentacles and eyelids.[41] It is expressed in embryonic stages 1719 in all arm buds, and subsequently in the future-tentacles in stages 245, suggesting that it may serve a role in the differential development of tentacles. Sequential expression of Hox genes is also observed in cephalopod arms.[41]

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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 eggs span a large range of sizes, from 1 to 30mm in diameter.[76] 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.[76] 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.[76]

Cephalopod Young do not pass through a larval stage, strictly speaking, instead they are known as the paralarvae. They quickly learn how to hunt, using encounters with prey to refine their strategies.[76] Growth in juveniles is usually allometric, whilst adult growth is isometric.[77]

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Evolution
The traditional view of cephalopod evolution holds that they evolved in the Late Cambrian from a monoplacophoran-like ancestor[78] with a curved, tapering shell,[79] which was closely related to the gastropods (snails).[80] 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.[80] 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.[81] However, some morphological evidence is difficult to reconcile with this view, and the redescription of Nectocaris pteryx, which did not have a shell and appeared to possess jet propulsion in the manner of "derived" cephalopods, complicated the question of the order in which cephalopod features developed provided Nectocaris is a cephalopod at all.[82] 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.[11] They underwent pulses of diversification during the Ordovician period[83] to become diverse and dominant in the Paleozoic and Mesozoic seas.[84] In the Early Palaeozoic, their range was far more restricted than today; they were mainly constrained to sublittoral regions of shallow shelves of the low latitudes, and usually occur in association with thrombolites.[85] A more pelagic habit was gradually adopted as the Ordovician progressed.[85] Deep-water cephalopods, whilst rare, have been found in the Lower Ordovician - but only in high-latitude waters.[85] 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 100200m.[83] 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.[86] (Endogastric mean the shell is curved so as the ventral or lower side is longitudinally concave (belly in); exogastric means the shell is curved so as the ventral side is longitudinally convex (belly out) allowing the funnel to be pointed backwards beneath the shell.)[86] 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.[85][87] The Bactritida, a SilurianTriassic group of orthocones, are widely held to be paraphyletic to the coleoids and ammonoids that is, the latter groups arose from An ammonitic ammonoid with the body chamber missing, showing the septal surface (especially at right) with its undulating lobes and within the Bactritida.[88]: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.[88] Unlike most modern cephalopods, most ancient varieties had protective shells. These shells at first were conical but later developed into curved nautiloid shapes seen in modern nautilus species. Competitive pressure from fish is thought to have forced the shelled forms into deeper water, which provided an evolutionary pressure towards shell loss and gave rise to the modern coleoids, a change which led to greater metabolic costs associated with the loss of

Cephalopod buoyancy, but which allowed them to recolonise shallow waters.[80]: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.[]:289

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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)

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.

[46]

The internal phylogeny of the cephalopods is difficult to constrain; many molecular techniques have been adopted, but the results produced are conflicting.[46][89] 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.[46] 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.[90][91] Molecular estimates for clade divergence vary. One 'statistically robust' estimate has Nautilus diverging from Octopus at 415 [92]24 million years ago.[93]

Cephalopod

16

Taxonomy
The classification presented here, for recent cephalopods, follows largely from Current Classification of Recent Cephalopoda [94] (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.[95] Class Cephalopoda ( indicates extinct groups) Subclass Nautiloidea: Fundamental ectocochliate cephalopods that provided the source for the Ammonoidea and Coleoidea. Order Plectronocerida: the ancestral cephalopods from the Cambrian Period Order Ellesmerocerida (500to470 [96] Ma) Order Endocerida (485to430 [97] Ma) Order Actinocerida (480to312 [98] Ma) Order Discosorida (482to392 [99] Ma) Order Pseudorthocerida (432to272 [100] Ma) Order Tarphycerida (485to386 [101] Ma) Order Oncocerida (478.5to324 [102] Ma) Order Nautilida (extant; 410.5to0 [103] Ma) Order Orthocerida (482.5to211.5 [104] Ma) Order Ascocerida (478to412 [105] Ma) Order Bactritida (418.1to260.5 [106] Ma) Subclass Ammonoidea: Ammonites (479to65 Ma) Order Goniatitida (388.5to252 [108] Ma) Order Ceratitida (254to200 [109] Ma) Order Ammonitida (215to66 [110] Ma) Subclass Coleoidea (410.0 Ma-Rec) Cohort Belemnoidea: Belemnites and kin Genus Jeletzkya Order Aulacocerida (265to183 [111] Ma) Order Phragmoteuthida (189.6to183 [112] Ma) Order Hematitida (339.4to318.1 [113] Ma) Order Belemnitida (339.4to65.5 [114] Ma) Genus Belemnoteuthis (189.6to183 [112] Ma) Cohort Neocoleoidea Superorder Decapodiformes (also known as Decabrachia or Decembranchiata) ?Order Boletzkyida Order Spirulida: Ram's Horn Squid Order Sepiida: cuttlefish
Atlantic Bobtail (Sepiola atlantica) [107] Common Cuttlefish (Sepia officinalis)

Chambered Nautilus (Nautilus pompilius)

Cephalopod 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.

17

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 Order Intejocerida Subclass Actinoceratoidea Not used by Flower, ibid Order Actinocerida Subclass Nautiloidea Nautiloidea in the restricted sense. Order Ellesmerocerida Plectronocerida subsequently split off as separate order. Order Orthocerida Includes orthocerids and pseudorthocerids Order Ascocerida Order Oncocerida Order Discosorida Order Tarphycerida Order Barrandeocerida A polyphyletic group now included in the Tarphycerida Order Nautilida Subclass Bactritoidea Order Bactritida Paleozoic Ammonoidea ( Miller, Furnish, and Schindewolf, 1957) Suborder Anarcestina Suborder Clymeniina Suborder Goniatitina

Common Octopus (Octopus vulgaris)

Cephalopod 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.

18

Shevyrev classification
Shevyrev (2005) suggested a division into eight subclasses, mostly comprising the more diverse and numerous fossil forms,[115][116] although this classification has been criticized as arbitrary.[117] Class Cephalopoda Subclass Ellesmeroceratoidea Order Plectronocerida (501to490 [118] Ma) Order Protactinocerida Order Yanhecerida Order Ellesmerocerida (500to470 [96] Ma) Subclass Endoceratoidea (485to430 [97] Ma) Order Endocerida (485to430 [97] Ma) Order Intejocerida (485to480 [119] Ma) Subclass Actinoceratoidea Order Actinocerida (480to312 [98] Ma) Subclass Nautiloidea (490.0 Ma- Rec) Order Basslerocerida (490to480 [120] Ma) Order Tarphycerida (485to386 [101] Ma) Order Lituitida (485to480 [119] Ma) Order Discosorida (482to392 [99] Ma) Order Oncocerida (478.5to324 [102] Ma) Order Nautilida (410.5 Ma-Rec) Subclass Orthoceratoidea (482.5to211.5 [104] Ma) Order Orthocerida (482.5to211.5 [104] Ma) Order Ascocerida (478to412 [105] Ma) Order Dissidocerida (479to457.5 [121] Ma) Order Bajkalocerida Subclass Bactritoidea (422to252 [122] Ma) Subclass Ammonoidea (410to66 [123] Ma) Subclass Coleoidea (410.0 Ma-rec)[124]
A fossilised belemnite Holotype of Ostenoteuthis siroi from family Ostenoteuthidae. Various species of ammonites

Cephalopod

19

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.[126][127]

Monophyly of coeloids
The coeloids have been thought to possibly represent a polyphyletic group,[]:289 although this has not been supported by the rising body of molecular data.[128]

Pyritized fossil of Vampyronassa rhodanica, a vampyromorphid from the Lower Callovian (unknown operator: u'callovianround' [125] million years ago)

Post-mortem decay
After death, if undisturbed, cephalopods[129] 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.[73]

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, ISBN0-12-728702-7 [3] Bartol I.K., Mann R., Vecchione M. (2002). "Distribution of the euryhaline squid Lolliguncula brevis in Chesapeake Bay: effects of selected abiotic factors". Mar. Ecol. Prog. Ser. 226: 235247. doi:10.3354/meps226235. [4] Marion Nixon; J.Z. Young (2003). The brains and lives of cephalopods. New York: Oxford University Press. ISBN0-19-852761-6. [5] Kutsch, Wolfram (1995). The nervous systems of invertebrates: An evolutionary and comparative approach (http:/ / books. google. com/ ?id=dW5e6FHOH-4C& lpg=PA115& pg=PA115#v=onepage& f=false). ISBN978-3-7643-5076-5. . [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 (2): 241307. 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 Evolutions Trajectory: Using Molluscan Eye Diversity to Understand Parallel and Convergent Evolution". Evolution Education and Outreach 1 (4): 439447. doi:10.1007/s12052-008-0084-1. [10] Martin J. Wells (2011). [paleo.ku.edu/treatiseonline Part M, Chapter 4: Physiology of Coleoids]. Lawrence, Kansas, USA. paleo.ku.edu/treatiseonline. [11] Boyle, Peter; Rodhouse, Paul (2004). Cephalopods : ecology and fisheries (http:/ / books. google. com/ ?id=4UtCi2B4VnoC). Ames, Iowa: Blackwell. doi:10.1002/9780470995310.ch2. ISBN0-632-06048-4. . [12] Messenger, John B.; Roger T. Hanlon (1998). Cephalopod Behaviour. Cambridge: Cambridge University Press. pp.1721. ISBN0-521-64583-2. [13] Hanlon and Messenger, 68. [14] Mthger, L.; Roberts, S.; Hanlon, R. (2010). "Evidence for distributed light sensing in the skin of cuttlefish, Sepia officinalis". Biology Letters 6 (5): 600603. doi:10.1098/rsbl.2010.0223. PMC2936158. PMID20392722. [15] 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): 112. PMID9317205.

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[16] 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. [17] "The cephalopods can hear you" (http:/ / news. bbc. co. uk/ earth/ hi/ earth_news/ newsid_8095000/ 8095977. stm). BBC News. 2009-06-15. . Retrieved 2010-04-28. [18] Tong, D.; Rozas, S.; Oakley, H.; Mitchell, J.; Colley, J.; Mcfall-Ngai, J. (Jun 2009). "Evidence for light perception in a bioluminescent organ". Proceedings of the National Academy of Sciences of the United States of America 106 (24): 98369841. Bibcode2009PNAS..106.9836T. doi:10.1073/pnas.0904571106. ISSN0027-8424. PMC2700988. PMID19509343. [19] "integument (mollusks)."Encyclopdia Britannica. 2009. Encyclopdia Britannica 2006 Ultimate Reference Suite DVD [20] tpn Manda and Vojtch Turek Acta Palaeontologica Polonica 54 (3), 2009: 503-512. Minute Silurian oncocerid nautiloids with unusual colour patterns (http:/ / www. app. pan. pl/ article/ item/ app20080062. html). . [21] Colour patterns in Early Devonian cephalopods from the Barrandian Area: Taphonomy and taxonomy Vojtch Turek Acta Palaeontologica Polonica (http:/ / www. app. pan. pl/ article/ item/ app20070064. html). 54. 2009. pp.491502. . [22] Roger T. Hanlon, John B. Messenger: Cephalopod Behaviour, page 2. Cambridge University Press, 1999, ISBN 0-521-64583-2 [23] Wells, M.J. (1 April 1980). "Nervous control of the heartbeat in octopus" (http:/ / jeb. biologists. org/ cgi/ content/ abstract/ 85/ 1/ 111). Journal of Experimental Biology 85 (1): 11128. PMID7373208. . [24] Ghiretti-Magaldi, A. (October 1992). "The Pre-history of Hemocyanin. The Discovery of Copper in the Blood of Molluscs" (http:/ / www. springerlink. com/ content/ ph8v3ln7560v68pp/ ). Cellular and Molecular Life Sciences (Birkhuser Basel) 48 (10). . [25] Daniel L. Gilbert; Adelman, William J.; John M. Arnold (1990). Squid as Experimental Animals (illustrated ed.). Springer. ISBN978-0-306-43513-3. [26] Electron Microscopical and Histochemical Studies of Differentiation and Function of the Cephalopod Gill (Sepia officinalis L.) (http:/ / www. springerlink. com/ content/ m483628053742p34/ fulltext. pdf). 1979. . [27] . Journal of Experimental Biology. 1994. http:/ / jeb. biologists. org/ cgi/ reprint/ 194/ 1/ 153. pdf. [28] Cole, A. .; Hall, B. . (2009). "Cartilage differentiation in cephalopod molluscs". Zoology (Jena, Germany) 112 (1): 215. doi:10.1016/j.zool.2008.01.003. PMID18722759. [29] See also http:/ / tolweb. org/ articles/ ?article_id=4200 [30] Wilbur, Karl M.; Clarke, M.R.; Trueman, E.R., eds. (1985), "11: Evolution of Buoyancy and Locomotion in recent cephalopods", The Mollusca, 12. Paleontology and neontology of Cephalopods, New York: Academic Press, ISBN0-12-728702-7 [31] Anderson, E.; Demont, M. (2000). "The mechanics of locomotion in the squid Loligo pealei: Locomotory function and unsteady hydrodynamics of the jet and intramantle pressure". The Journal of experimental biology 203 (Pt 18): 28512863. PMID10952883. [32] Bartol, I. K.; Krueger, P. S.; Thompson, J. T.; Stewart, W. J. (2008). "Swimming dynamics and propulsive efficiency of squids throughout ontogeny". Integrative and Comparative Biology 48: 720. doi:10.1093/icb/icn043. [33] E., S.; M., V. (2002). "Quantification of ontogenetic discontinuities in three species of oegopsid squids using model II piecewise linear regression". Marine Biology 140: 971. doi:10.1007/s00227-001-0772-7. [34] W. Johnson; P. D. Soden; E. R. Trueman. "A Study in Jet Propulsion: An Analysis of the Motion of the Squid, Loligo Vulgaris". Journal of Experimental Biology 56 (1972): 155165. [35] Campbell, Reece, & Mitchell, p.612 [36] Guerra, A.; Martinell, X.; Gonzlez, A. F.; Vecchione, M.; Gracia, J.; Martinell, J. (2007). 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Lethaia 42 (2): 204217. doi:10.1111/j.1502-3931.2008.00133.x. - this reference also contains a good summary of the process of egg development. [72] Norman, M.D. (2000). Cephalopods: A World Guide. ConchBooks. [73] Kear, A.J.; Briggs, D.E.G.; Donovan, D.T. (1995). "Decay and fossilization of non-mineralized tissue in coleoid cephalopods" (http:/ / palaeontology. palass-pubs. org/ pdf/ Vol 38/ Pages 105-131. pdf). Palaeontology 38 (1): 105132. . Retrieved 2009-04-21. [74] Shigeno, S.; Sasaki, T.; Moritaki, T.; Kasugai, T.; Vecchione, M.; Agata, K. (Jan 2008). "Evolution of the cephalopod head complex by assembly of multiple molluscan body parts: Evidence from Nautilus embryonic development". Journal of Morphology 269 (1): 117. doi:10.1002/jmor.10564. PMID17654542. [75] Daniel L. Gilbert; William J. Adelman; John M. Arnold (1990). Squid as experimental animals. New York: Plenum Press. ISBN0-306-43513-6.

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You can jump the queue or expand by hand (http:/ / en. wikipedia. org/ wiki/ Template:cite_doi/ _10. 1002. 2f9780470995310. ch3?preload=Template:Cite_doi/ preload& editintro=Template:Cite_doi/ editintro& action=edit) [81] doi: 10.1111/j.1475-4983.2007.00644.x This citation will be automatically completed in the next few minutes. You can jump the queue or expand by hand (http:/ / en. wikipedia. org/ wiki/ Template:cite_doi/ _10. 1111. 2fj. 1475-4983. 2007. 00644. x?preload=Template:Cite_doi/ preload& editintro=Template:Cite_doi/ editintro& action=edit) [82] Smith, Martin R and Caron, Jean-Bernard (2010). "Primitive soft-bodied cephalopods from the Cambrian". Nature 465 (7297): 427428. Bibcode2010Natur.465..427B. doi:10.1038/465427a. PMID20505713 [83] doi: 10.1016/j.palaeo.2008.12.015 This citation will be automatically completed in the next few minutes. You can jump the queue or expand by hand (http:/ / en. wikipedia. org/ wiki/ Template:cite_doi/ _10. 1016. 2fj. palaeo. 2008. 12. 015?preload=Template:Cite_doi/ preload& editintro=Template:Cite_doi/ editintro& action=edit) [84] Dzik, J. (1981). "Origin of the cephalopoda" (http:/ / www. paleo. pan. pl/ people/ Dzik/ Publications/ Cephalopoda. pdf) (PDF). Acta Palaeontologica Polonica 26 (2): 161191. . [85] doi: 10.1371/journal.pone.0007262 This citation will be automatically completed in the next few minutes. You can jump the queue or expand by hand (http:/ / en. wikipedia. org/ wiki/ Template:cite_doi/ _10. 1371. 2fjournal. pone. 0007262_?preload=Template:Cite_doi/ preload& editintro=Template:Cite_doi/ editintro& action=edit) [86] Holland, C. H. (1987). "The nautiloid cephalopods: a strange success: President's anniversary address 1986". Journal of the Geological Society 144: 10. doi:10.1144/gsjgs.144.1.0001. [87] Krger, Bjrn (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): 129139. doi:10.1080/00241160600623749. . [88] 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.393420. [89] 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 & Evolution 37 (2): 426441. doi:10.1016/j.ympev.2005.03.020. PMID15935706. [90] Strugnell, J.; Jackson, J.; Drummond, A. J.; Cooper, A. (2006). "Divergence time estimates for major cephalopod groups: evidence from multiple genes". Cladistics 22: 8996. doi:10.1111/j.1096-0031.2006.00086.x. [91] 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): 135370. PMID10958852. [92] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. php?Ma=415 [93] 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): 362374. doi:10.1007/s00239-005-0160-x. PMID16501879. [94] http:/ / www. mnh. si. edu/ cephs/ newclass. pdf [95] Bather, F.A. (1888b). "Professor Blake and Shell-Growth in Cephalopoda" (http:/ / biodiversitylibrary. org/ page/ 26195416). Annals and Magazine of Natural History. 6 1: 421426. . [96] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. php?Ma=500470 [97] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. php?Ma=485430 [98] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. php?Ma=480312 [99] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. php?Ma=482392 [100] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. php?Ma=432272 [101] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. php?Ma=485386 [102] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. php?Ma=478. 5324. 0 [103] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. php?Ma=410. 50 [104] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. php?Ma=482. 5211. 5

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Cephalopod
[105] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. php?Ma=478412 [106] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. php?Ma=418. 1260. 5 [107] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. php?Ma=47965 [108] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. php?Ma=388. 5252. 0 [109] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. php?Ma=254200 [110] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. php?Ma=21566 [111] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. php?Ma=265183 [112] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. php?Ma=189. 6183. 0 [113] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. php?Ma=339. 4318. 1 [114] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. php?Ma=339. 465. 5 [115] 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): 606614. "Translated from Paleontologicheskii Zhurnal No. 6, 2005, 33-42." [116] 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): 4654. doi:10.1134/S0031030106010059. [117] Kroger, B.. "Peer review in the Russian "Paleontological Journal"" (http:/ / www. tiefes-leben. de/ 2009/ 03/ peer-review-in-the-russian-paleontological-journal). . [118] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. php?Ma=501490 [119] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. php?Ma=485480 [120] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. php?Ma=490480 [121] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. php?Ma=479. 0457. 5 [122] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. php?Ma=422252 [123] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. php?Ma=41066 [124] Bather, F.A. (1888a). "Shell-growth in Cephalopoda (Siphonopoda)" (http:/ / biodiversitylibrary. org/ page/ 26195293). Annals and Magazine of Natural History. 6 1: 298310. . [125] http:/ / toolserver. org/ ~verisimilus/ Timeline/ Timeline. php?Ma=Callovian [126] Berthold, Thomas, & Engeser, Theo (1987). "Phylogenetic analysis and systematization of the Cephalopoda (Mollusca)". Verhandlungen Naturwissenschaftlichen Vereins in Hamburg. (NF) 29: 187220. [127] 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. . [128] 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. [129] Experiments were performed on a variety of squid

23

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 (11): 11671284. 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. National Museum of Natural History: Department of Systematic Biology: Invertebrate Zoology: Cephalopods (http://www.mnh.si.edu/cephs) N. Joan Abbott, Roddy Williamson, Linda Maddock. Cephalopod Neurobiology. Oxford University Press, 1995. ISBN 0-19-854790-0 Marion Nixon & John Z. Young. The brains and lives of Cephalopods. Oxford University Press, 2003. ISBN 0-19-852761-6 Roger T. Hanlon & John B. Messenger. Cephalopod Behaviour. Cambridge University Press, 1996. ISBN 0-521-42083-0

Cephalopod Martin Stevens & Sami Merilaita. Animal camouflage: mechanisms and function. Cambridge University Press, 2011. ISBN 0-521-19911-5 Classification key to modern cephalopods: ftp://ftp.fao.org/docrep/fao/009/a0150e/a0150e03.pdf

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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) Rapid Adaptive Camouflage and Signaling in Cephalopods (http://www.ibioseminars.org/lectures/ ecology-a-behavior/roger-hanlon.html) Online seminar by Roger Hanlon (MBL)

Octopus

25

Octopus
Octopus Scientific classification Kingdom: Phylum: Class: Animalia Mollusca Cephalopoda

Superorder: Octopodiformes Order: Octopoda<br Suborders

Cirrina Incirrina Synonyms

Octopoida Leach, 1817[1]

The octopus ( /ktps/; plural: octopuses, octopi, or octopodes; see below) 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 mantles), 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.[2] Around 300 species are recognized, 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.

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.[3] 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. Schematic lateral aspect of octopod features 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 bodies. This enables them to squeeze through very narrow slits between underwater rocks, which is very helpful when they are fleeing from moray eels 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.

Octopus

26 Octopuses have a relatively short life expectancy, and some species live for as little as six months. Larger species, such as the giant pacific 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, eventually dying of starvation. In a scientific experiment, removal of both optic glands after spawning was found to result in cessation of broodiness, resumption of feeding, increased growth, and greatly extended lifespans.[4]

An octopus moving between tide pools during low tide

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. The octopus draws water into its mantle cavity, where it passes through its gills. As mollusks, their gills are finely divided and vascularized outgrowths of either the outer or the inner body surface.

Grimpoteuthis discoveryi, a finned octopus of the suborder Cirrina

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 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

27 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, using a nonsomatotopic system unique to large-brained invertebrates.[10] Some octopuses, such as the mimic octopus, will move their arms in ways that emulate the shape and 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 aquariums and then catching them.[12] Octopuses often break out of An octopus opening a container with a screw cap 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

28

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, the male uses 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).

Octopus

29

Sensation
Octopuses have keen eyesight. Like other cephalopods, they can distinguish the polarization of light. Color vision appears to vary from species to species, being present in O. aegina but absent in O. 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 the pupil slit is always horizontal. Octopuses also have an excellent sense of touch. An octopus's suction cups are equipped with chemoreceptors so the octopus can taste what it Eye of Octopus vulgaris is touching. The arms contain tension sensors so the octopus knows whether its arms are stretched out. However, it has a very poor proprioceptive sense. The tension receptors are not sufficient for the brain to determine the position of the octopus's body or arms. (It is not clear whether 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 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.

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30

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, some octopuses (Adopus aculeatus and Amphioctopus marginatus under current taxonomy) were found to 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 Video of an octopus in its natural habitat possibly not triggering that predator's search image for octopus (food).[24] A study of this behavior conducted by the Weymouth Sea Life Centre led to the suggestion that the two rearmost appendages may be more accurately termed 'legs' rather than 'arms'.[26] Some species of octopuses can crawl out of the water for a short period, which they may do between tide pools while hunting crustaceans or gastropods or to escape predators.[27][28] Octopuses swim by expelling a jet of water from a contractile mantle, and aiming it via a muscular siphon.

Prey
Bottom-dwelling octopuses eat mainly crabs, polychaete worms, and other molluscs such as whelks and clams. Open-ocean octopuses eat mainly prawns, fish and other cephalopods. They usually inject their prey with a paralysing saliva before dismembering it into small pieces with their beaks.[29] Octopuses feed on shelled molluscs either by using force, or by drilling a hole in the shell, injecting a secretion into the hole, and then extracting the soft body of the mollusc.[30]

Size
The giant Pacific octopus, Enteroctopus dofleini, is often cited as the largest octopus species. Adults usually weigh around 15kg (33lb), with an arm span of up to 4.3m (14ft).[31] The largest specimen of this species to be scientifically documented was an animal with a live mass of 71kg (156.5lb).[32] 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 75kg (165lb).[33][34] However, a number of questionable size records would suggest E. dofleini is the largest of all octopus species by a considerable margin;[35] one such record is of a specimen weighing 272kg (600lb) and having an arm span of 9m (30ft).[36]

An adult giant Pacific octopus, Enteroctopus dofleini

Etymology and pluralization

The term "octopus" is from Greek [37][38] (oktapous, "eight-footed"), with traditional plural forms "octopuses" (pronounced /ktpsz/) from English grammar and "octopodes" (pronounced /ktpdiz/) from the Greek. Currently, "octopuses" is the most common form in both the US and the UK. The term "octopod" (plural: "octopods" or "octopodes") is taken from the taxonomic order Octopoda, but has no classical equivalent. The collective plural "octopus" is usually reserved for animals consumed for food.

Octopus Some authorities consider "octopi" an objectionable[39] hypercorrection, feeling the form arose from the incorrect assumption that "octopus" is a Latin second-declension form. However, "octopus" is a scientific Latin third-declension noun with a plural of octopodes. Nevertheless, the Oxford English Dictionary (2008 Draft Revision)[40] lists "octopuses", "octopi", and "octopodes" (in that order), labelling "octopodes" 'rare' and noting that "octopi" derives from the misapprehension that octpus is a second-declension Latin noun. The book further maintains that if the word were native to Latin, it would be third declension octps (plural: octpedes) after the pattern of ps ("foot", plural peds).[41] The original Latin word for octopus and other similar species is polypus, from Greek polpous (, "many-footed"); again, usually the inappropriate plural polyp is used instead of polypods. Fowler's Modern English Usage states, 'the only acceptable plural in English is "octopuses"', that "octopi" is 'misconceived', and "octopodes" 'pedantic'. Chambers 21st Century Dictionary[42] and the Compact Oxford Dictionary[43] list only "octopuses", although the latter notes that "octopodes" is 'still occasionally used'. The descriptivist Merriam-Webster 11th Collegiate Dictionary lists "octopuses" and "octopi" in that order; likewise, Webster's New World College Dictionary lists in order "octopuses", "octopi", and "octopodes". In modern Greek, the word is (khtapdi; plural: , khtapdia), from Byzantine (oktapdion) derived from the Classical Greek variant (oktpous).

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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.[44] Octopuses were often depicted in the art of the Moche people of ancient Peru, who worshipped the sea and its animals.[45]

Octopus

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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.[46]

In literature
The octopus has a significant role in Victor Hugo's book Travailleurs de la mer (Toilers of the Sea).[47]

As a metaphor
Due to having numerous arms that emanate from a common center, the octopus is often used as a metaphor for a group or organization which is perceived as being powerful, manipulative or bent on domination. Use of this terminology is invariably negative and employed by the opponents of the groups or institutions so described.[48]

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. In Korea, some small species are sometimes eaten alive as a novelty food. A live octopus is usually sliced up, and it is eaten while still squirming.
Octopus at Tsukiji fish market

Octopuses are "tickled" out of their holes in the Hawaiian Islands with three-pronged polespears

Raw octopus arms

Lightly boiled octopus arm that turned a bright purple

Octopus 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 feira (market fair style octopus) is a local delicacy. Restaurants which specialize or serve this dish are known as pulperas. 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 them for octopuses sheltered there. A common scene in the Greek islands is octopuses hanging in the sunlight from a rope, just like laundry from a 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 about 139 kilocalories (Calories) per three-ounce portion, and is a source of vitamin B3, B12, potassium, phosphorus, and selenium.[49] Care must be taken to boil the octopus properly, to rid it of slime, smell, and residual ink.

33

As pets
Though octopuses can be difficult to keep in captivity, some people keep them as pets. They often escape even from supposedly secure tanks, due to their problem-solving skills, mobility and lack of rigid structure. The variation in size and lifespan 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 it is young with a full life ahead of it. Large octopuses have also been known to catch and kill some species of sharks.[50]

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 Family Opisthoteuthidae: umbrella octopus Family Cirroteuthidae Family Stauroteuthidae Suborder Incirrina
Cirrothauma murrayi

Octopus

34 Family Amphitretidae: telescope octopus Family Bolitaenidae: gelatinous octopus Family Octopodidae: benthic octopus Family Vitreledonellidae: glass octopus Superfamily Argonautoida
Amphitretus pelagicus

Family Alloposidae: seven-arm octopus Family Argonautidae: argonauts Family Ocythoidae: tuberculate pelagic octopus Family Tremoctopodidae: blanket octopus

References
[1] Helsinki.fi (http:/ / www. helsinki. fi/ ~mhaaramo/ metazoa/ protostoma/ mollusca/ Cephalopoda/ Coleoidea. htm), Mikko's Phylogeny Archive: Coleoidea Recent cephalopods [2] 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 [3] 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." [4] Wodinsky, Jerome (2 December 1977). "Hormonal Inhibition of Feeding and Death in Octopus: Control by Optic Gland Secretion" (http:/ / www. sciencemag. org/ content/ 198/ 4320/ 948. abstract). Science 198 (4320): 948951. doi:10.1126/science.198.4320.948. . Retrieved 27 November 2011. [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 OctopusMighty 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): 6972. doi:10.1353/psc.2005.0004. [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(312KB) Proceedings of the Royal Society 268: 17551758. [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: 5770. 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(453KB) . Nippon Suisan Gakkashi 67(1): 3539.

Octopus
[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] Octopuses have only six arms, the other two are actually legs! (http:/ / www. highbeam. com/ doc/ 1P3-1530507261. html) Hindustan Times, 8/3/2008 [27] Harmon, Katherine (24 November 2011). "Land-Walking Octopus Explained" (http:/ / blogs. scientificamerican. com/ octopus-chronicles/ 2011/ 11/ 24/ land-walking-octopus-explained-video/ ). Octopus Chronicles. Scientic American blogs. . Retrieved 24 November 2011. [28] Wood, James B.; Roland C. Anderson (2004). "Interspecific Evaluation of Octopus Escape Behavior" (http:/ / www. animalsandsociety. org/ assets/ library/ 186_s15327604jaws07022. pdf). Journal of Applied Animal Welfare Science (Lawrence Erlbaum Associates, Inc.) 7 (2): 95106. doi:10.1207/s15327604jaws0702_2. PMID15234886. . [29] Wassilieff, Maggy; and OShea, Steve (2009) "Octopus and squid - Feeding and predation" (http:/ / www. TeAra. govt. nz/ en/ octopus-and-squid/ 3) Te Ara - the Encyclopedia of New Zealand. Updated 2 March 2009. [30] Wodinsky, Jerome (1969) "Penetration of the Shell and Feeding on Gastropods by Octopus" (http:/ / icb. oxfordjournals. org/ content/ 9/ 3/ 997. short). Amer. Zool., 9(3): 9971010. doi:10.1093/icb/9.3.997 [31] Smithsonian National Zoological Park: Giant Pacific Octopus (http:/ / nationalzoo. si. edu/ Animals/ Invertebrates/ Facts/ cephalopods/ FactSheets/ Pacificoctopus. cfm) [32] 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. [33] O'Shea, S. 2004. The giant octopus Haliphron atlanticus (Mollusca : Octopoda) in New Zealand waters. New Zealand Journal of Zoology 31(1): 713. [34] O'Shea, S. 2002. Haliphron atlanticus a giant gelatinous octopus. Biodiversity Update 5: 1. [35] Norman, M. 2000. Cephalopods: A World Guide. ConchBooks, Hackenheim. p. 214. [36] High, W.L. (1976). "The giant Pacific octopus". U.S. National Marine Fisheries Service, Marine Fisheries Review 38 (9): 1722. [37] 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 [38] Scientific Latin from Greek -, (also - ) "eight-footed" > - or - [combination form of "eight"] and -, "foot". Cf. Modern Greek < < < . [39] Peters, Pam (2004). The Cambridge Guide to English Usage. Cambridge: Cambridge University Press. ISBN 0-521-62181-X, p. 388. [40] 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. [41] "centipede" (http:/ / oed. com/ search?searchType=dictionary& q=centipede). Oxford English Dictionary (3rd ed.). Oxford University Press. 2001. . [42] Chambersharrap.co.uk (http:/ / www. chambersharrap. co. uk/ chambers/ features/ chref/ chref. py/ main?title=21st& query=octopus), Retrieved October 19, 2007. [43] Askoxford.com (http:/ / www. askoxford. com/ concise_oed/ octopus), Retrieved October 19, 2007. [44] C. Michael Hogan. 2007 Knossos fieldnotes, The Modern Antiquarian (http:/ / www. themodernantiquarian. com/ site/ 10854/ knossos. html#fieldnotes) [45] Berrin, Katherine & Larco Museum. The Spirit of Ancient Peru:Treasures from the Museo Arqueolgico Rafael Larco Herrera. New York: Thames and Hudson, 199 7. [46] Dixon, Roland Burrage (1916). The Mythology of All Races. 9. Marshall Jones. p.15. [47] Chisholm, Hugh, ed. (1911). "Octopus". Encyclopdia Britannica (11th ed.). Cambridge University Press. [48] Nazi Tentacles: The octopus as visual metaphor (http:/ / scienceblogs. com/ bioephemera/ 2010/ 05/ nazi_tentacles_a_brief_history. php) [49] Octopus Calories And Nutrition (http:/ / www. annecollins. com/ calories/ calories-octopus-squid. htm) [50] Archived Google video of an octopus catching a shark (http:/ / video. google. com/ videoplay?docid=-7004909622962894202), from The Octopus Show by Mike deGruy

35

Octopus

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External links
The Cephalopod Page (http://www.thecephalopodpage.org/) 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 (25mm) 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(359KB) Video of walking octopuses (http://berkeley.edu/news/media/releases/2005/03/24_octopus.shtml)

Squid

37

Squid
Squid Temporal range: (at least)

European squid (Loligo vulgaris)

Scientific classification Kingdom: Phylum: Class: Superorder: Order: Animalia Mollusca Cephalopoda Decapodiformes Teuthida<br Suborders Plesioteuthididae (incertae sedis) Myopsina Oegopsina Squid are 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 material.

Squid

38

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 practically 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 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.

Composite diagram illustrating basic squid features (ventral aspect)

Nervous system
The giant axon, which may be up to 1mm (0.04in) 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

A dissected male specimen of Onykia ingens, showing a non-erect penis (the white tubular structure located below most of the other organs)

A specimen of the same species exhibiting elongation of the penis to 67cm 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

Squid 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]

39

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 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

Ventral view of the viscera of the female Chtenopteryx sicula

lower ventricle and two upper auricles.

Head
The head end bears eight arms and two 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 sexual 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). The eyes, on either side of the head, each contain a hard lens. The image is focused by changing the position of the lens, as in a camera or telescope, rather than changing the shape of the lens, as in the human eye. Squid appear to have limited hearing.[6]

Squid

40

Size
The majority are no more than 60cm (24in) long, although the giant squid may reach 13 metres (43ft).[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 largest squid known at the time.[8]

A frozen giant squid in Melbourne Aquarium

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 14m (46ft) in length, making it the largest invertebrate.[10] Squid have the largest eyes in the animal kingdom. 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 495kg (1,090lb) and measuring around 10m (33ft) 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 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 Nautiloidea: nautilus Subclass Coleoidea: squid, octopus, cuttlefish Superorder Octopodiformes Superorder Decapodiformes ?Order Boletzkyida Order Spirulida: Ram's Horn Squid Order Sepiida: cuttlefish Order Sepiolida: bobtail squid Order Teuthida: squid Family Plesioteuthididae (incertae sedis) Suborder Myopsina Family Australiteuthidae
Grimalditeuthis bonplandi

Bathyteuthis abyssicola

Squid 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 Family Chiroteuthidae Family Chtenopterygidae: comb-finned squid Family Cranchiidae: glass squid Family Cycloteuthidae Family Enoploteuthidae Family Gonatidae: armhook squid Family Histioteuthidae: jewel squid Family Joubiniteuthidae: Joubin's Squid Family Lepidoteuthidae: Grimaldi Scaled Squid Family Lycoteuthidae 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 Family Walvisteuthidae Parateuthis tunicata (incertae sedis)
Mastigoteuthis flammea Histioteuthis reversa

41

Commercial fishing
According to the FAO, the cephalopod catch for 2002 was 3,173,272 tonnes (6.995867109lb). Of this, 2,189,206 tonnes, or 75.8 percent, was squid.[12] The following table lists the squid species fishery catches which exceeded 10,000 tonnes (22,000,000lb) in 2002.
Onychoteuthis banksii

Squid

42

Pterygioteuthis giardi

World squid catch in 2002 Species Family

[12] Catch tonnes 24,976 16,684 225,958 22,483 Percent

Common name

Loligo gahi Loligo pealei Common squid nei [13]

Loliginidae Loliginidae Loliginidae

Patagonian squid Longfin squid

1.1 0.8 10.3 1.0 23.3 18.6 23.0 2.8 18.6

Ommastrephes bartramii Ommastrephidae Neon flying squid Illex argentinus Dosidicus gigas Todarodes pacificus Nototoda russloani Squid nei [13]

Ommastrephidae Argentine shortfin squid 511,087 Ommastrephidae Jumbo flying squid Ommastrephidae Japanese flying squid 406,356 504,438

Ommastrephidae Wellington Flying Squid 62,234 Various 414,990

Total squid

2,189,206 100.0

Squid

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As food
Many species are popular as food in cuisines as diverse as Chinese, Greek, Turkish, Japanese, Portuguese, Italian, Spanish, Korean, Vietnamese, 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 high in the recommended daily intake of copper,[14] selenium, vitamin B12, and riboflavin.[15]

Fried calamari: breaded, deep-fried squid

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. ISBN0-19-857603-X. [5] Miserez, A; Li, Y; Waite, H; Zok, F (2007). "Jumbo squid beaks: Inspiration for design of robust organic composites". Acta Biomaterialia 3 (1): 139149. doi:10.1016/j.actbio.2006.09.004. PMID17113369. [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/ ) [10] Anderton, Jim (22 February 2007). "World's largest squid landed in NZ" (http:/ / www. beehive. govt. nz/ release/ world039s-largest-squid-landed-nz). 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). ISBN955-1052-67-0. . [13] nei: not elsewhere included [14] Squid - Overview: Food Market Exchange - B2B e-marketplace for the food industry (http:/ / www. foodmarketexchange. com/ datacenter/ product/ seafood/ squid/ detail/ dc_pi_sf_squid_0204. htm) [15] FishWatch - Market Squid (http:/ / www. nmfs. noaa. gov/ fishwatch/ species/ market_squid. htm)

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External links
CephBase: Teuthida (http://www.cephbase.utmb.edu/spdb/squid.cfm) Colossal Squid at the Museum of New Zealand Te Papa Tongarewa (http://squid.tepapa.govt.nz/) Market squid mating, laying eggs (video) (http://diving.rogerbly.com/video/squid) Scientific American - Giant Squid (http://www.scientificamerican.com/article. cfm?id=live-giant-squid-photogra) The Cephalopod Page (http://www.thecephalopodpage.org/) The Octopus News Magazine Online (http://www.tonmo.com/)

Cuttlefish

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Cuttlefish
Cuttlefish

Sepia latimanus, East Timor

Scientific classification Kingdom: Phylum: Class: Superorder: Order: Animalia Mollusca Cephalopoda Decapodiformes Sepiida<br 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. 'Cuttle' is a reference to their unique internal shell, the cuttlebone; and despite their name, cuttlefish are true molluscs. Cuttlefish have large W-shaped pupils, eight arms and two tentacles furnished with denticulated (very finely toothed)[1] suckers, with which they secure their prey. They generally range in size from 15cm (5.9in) to 25cm (9.8in), with the largest species, Sepia apama, reaching 50cm (20in) in mantle length and over 10.5kg (23lb) in weight.[2] Cuttlefish eat small molluscs, crabs, shrimp, fish, octopuses, worms, and other cuttlefish. Their predators include dolphins, sharks, fish, seals, seabirds and other cuttlefish. Their life expectancy is about one to two years. Recent studies indicate that cuttlefish are among the most intelligent invertebrates.[3] Cuttlefish also have one of the largest brain-to-body size ratios of all invertebrates.[3] The 'cuttle' in 'cuttlefish' comes from the Old English word cudele, meaning 'cuttlefish', which may be cognate with the Old Norse koddi ('cushion') and the Middle Low German kdel ('pouch'). 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 both Greek and Latin, sepia, is now used to refer to a brown pigment in English.

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Physiology
Cuttlebone
Cuttlefish possess an internal structure called the cuttlebone, which is porous and is made of aragonite. This provides the cuttlefish with buoyancy, which it regulates by changing the gas-to-liquid ratio in the chambered cuttlebone via the ventral siphuncle.[4] Each species' cuttlebone has a distinct shape, size, and pattern of ridges or texture. The cuttlebone is unique to cuttlefish, and is one of the features that distinguish them from their squid relatives. Video of a cuttlefish in its natural habitat Jewellers and silversmiths traditionally use cuttlebones as moulds for casting small objects,[5] 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.

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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 polarisation 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 millimetre,[6] which would correspond to about 359 DPI if they could be An infant cuttlefish protects itself with camouflage individually controlled. The pigmented chromatophores have a sac of pigment and a large membrane that is folded when retracted. There are 6 to 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.[7]

Eyes
Cuttlefish, like other cephalopods, have sophisticated eyes. The organogenesis and final structure of the cephalopod eye differs fundamentally from that of vertebrates like humans.[8] Superficial similarities between cephalopod and vertebrate eyes are thought to be examples of convergent evolution. The cuttlefish pupil is a smoothly-curving W shape.[9][10] Although cuttlefish cannot see color,[11] 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 backward. The eye changes focus by shifting the position of the entire lens with respect to the retina, instead of reshaping the lens as in mammals. Unlike the vertebrate eye, there is no blind spot, because the optic nerve is positioned behind the retina.

Close up of a cuttlefish eye

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.[12]

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Pupil expansion in Sepia officinalis

Suckers
The suckers of cuttlefish extend most of the length of their arms and along the distal portion of their tentacles.

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 An illustration of the suckers of a cuttlefish's pair of gills (one heart for each), and the third pumps blood around cuttlefish, 1834 the rest of the body. Cuttlefish blood must flow more rapidly than that of most other animals because hemocyanin carries substantially less oxygen than hemoglobin.

Ink
Cuttlefish have ink, like squid and octopus species, which they use to help evade predators. This ink is stored inside an ink sac.

Toxicity
Like octopuses and some squid, all cuttlefish have bacterially-produced neurotoxins in their saliva.[13] The muscles of Pfeffer's Flamboyant Cuttlefish contain a highly toxic compound that is yet to be identified.[3] 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.[14]

Ecology
Diet
While the preferred diet of cuttlefish is crabs and fish, they feed on small shrimp shortly after hatching.[15]
Pfeffer's Flamboyant Cuttlefish from Sipadan, Malaysia

They use their camouflage to hunt and sneak up on their prey.[16] They swim at the bottom, where shrimp and crab are found and shoot out a jet of water to uncover the shrimp or crab that are buried in the sand. Then when the shrimp or crab are trying to get away the cuttlefish open their eight arms and shoot out two long feeding tentacles to grab them. On the end of each is a pad covered in suckers that grabs and pulls prey toward its beak, where it gets

Cuttlefish paralyzed by poison and then eaten.[15]

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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 (2,000ft).[17] 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.[18]

Mating
[19]

Males challenge one another for dominance and the best den during mating season. During this challenge no direct contact is usually made. They just threaten each other until one of them wimps out and gets called out on their bluff.

Taxonomy
Over 120 species of cuttlefish are currently recognised, grouped into 5 genera. Sepiadariidae contains seven species and 2 genera; all the rest are in Sepiidae. CLASS CEPHALOPODA Subclass Nautiloidea: nautilus Subclass Coleoidea: squid, octopus, cuttlefish Superorder Octopodiformes Superorder Decapodiformes ?Order Boletzkyida Order Spirulida: Ram's Horn Squid Order Sepiida: cuttlefish Suborder Vasseuriina Family Vasseuriidae Family Belosepiellidae Suborder Sepiina Family Belosaepiidae Family Sepiadariidae Family Sepiidae Order Sepiolida: bobtail squid Order Teuthida: squid
Sepia officinalis from Turkish waters

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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 are especially popular in Italy, where they are used in Risotto al Nero di Seppia (literally black cuttlefish rice). The Croatian Crni Riot 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 rice turns because of the cuttlefish ink. In Spain, breaded and deep-fried cuttlefish is a popular dish in Andalusia, where it is known as choco. Spanish cuisine, especially that of the coastal regions, uses cuttlefish and squid ink for the marine flavour and smoothness it provides; it is included in dishes such as rice, pasta and fish stews.

Linguine with cuttlefish and ink sauce served at a Venetian osteria

In Portugal, cuttlefish is present in many popular dishes, chocos com tinta (cuttlefish in black ink) being among the most popular. This dish is made with grilled cuttlefish served in a marinade of its own ink. Cuttlefish is also the regional dish of the city of Setbal and surrounding areas, where it is served as deep-fried strips or in a variant of feijoada, with red kidney beans. A somewhat popular food is black dry spaghetti (also present in other pasta) that one can find in any store right next to the normal dry variant. The black color comes from using the ink of the cuttlefish during industrial food processing and is known as esparguete de tinta de choco (cuttlefish ink spaghetti).

Sepia
Cuttlefish ink was formerly an important dye, called sepia. Today artificial dyes have mostly replaced natural sepia.

Research
Research into replicating biological color-changing has lead to engineering artificial chromatophores out of small devices known as dielectric elastomer actuators. Engineers at the University of Bristol[20] have engineered soft materials that mimic a cuttlefishs color-changing skin, paving the way for smart clothing and camouflage applications.[21]

References
[1] From Latin denticultus having small teeth "denticulate" (http:/ / www. thefreedictionary. com/ denticulate). TheFreeDictionary. . [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. 57152. [3] NOVA, 2007. Cuttlefish: Kings of Camouflage. (http:/ / www. pbs. org/ wgbh/ nova/ camo/ ) (television program) NOVA, PBS, April 3, 2007. [4] Rexfort, A.; Mutterlose, J. (2006). "Stable isotope records from Sepia officinalisa key to understanding the ecology of belemnites?". Earth and Planetary Science Letters 247 (34): 212212. Bibcode2006E&PSL.247..212R. doi:10.1016/j.epsl.2006.04.025. [5] Casting Silver with Cuttlefish (http:/ / www. silverstall. com/ casting-silver-jewellery. html). Silverstall.com. Retrieved on 2011-09-18. [6] http:/ / www. pbs. org/ wgbh/ nova/ camo/ anat-nf. html [7] Mthger, L. M., Shashar, N., and R.T. Hanlon. 2009. Do cephalopods communicate using polarized light reflections from their skin? Journal of Experimental Biology 212: 21332140. doi:10.1242/jeb.020800 [8] 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.

Cuttlefish
[9] Schaeffel, F.; Murphy, C. J.; Howland, H. C. (1999). "Accommodation in the cuttlefish (Sepia officinalis)". The Journal of experimental biology 202 (Pt 22): 31273134. PMID10539961. [10] Murphy, C. J.; Howland, H. C. (1990). "The functional significance of crescent-shaped pupils and multiple pupillary apertures". Journal of Experimental Zoology 256: 2228. doi:10.1002/jez.1402560505. [11] Mthger, Lydia M.. "Color blindness and contrast perception in cuttlefish (Sepia officinalis) determined by a visual sensorimotor assay" (http:/ / hermes. 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. [12] "Cuttlefish spot target prey early" (http:/ / news. bbc. co. uk/ 1/ hi/ sci/ tech/ 7435757. stm). BBC News. 2008-06-05. . Retrieved 2008-05-06. [13] All Octopuses Are Venomous, Study Says (http:/ / news. nationalgeographic. com/ news/ 2009/ 04/ 090417-octopus-venom. html). News.nationalgeographic.com (2010-10-28). Retrieved on 2011-09-18. [14] 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). [15] Cuttlefish Basics (http:/ / www. tonmo. com/ articles/ basiccuttlefish. php). Tonmo.com (2003-02-12). Retrieved on 2011-09-18. [16] Cousteau and Diol., Jacques and Philippe (1973). Octopus and Squid, the Soft Intelligence. Garden City, N.Y.. [17] 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'Universit de Caen: 192. [18] Young, R. E., M. Vecchione and D. Donovan (1998). "The evolution of coleoid cephalopods and their present biodiversity and ecology". South African Journal of Marine Science 20: 393. doi:10.2989/025776198784126287. [19] Bavendam, Fred. "The Giant Cuttlefish Chameleon of the Reef." National Geographic 1995: 94-107. Print. [20] "Biomimetic chromatophores for camouflage and soft active surfaces" (http:/ / iopscience. iop. org/ 1748-3190/ 7/ 3/ 036009?fromSearchPage=true). Bioinspiration & Biomimetics. 3 (IOPscience) 7 (036009). September 2012. doi:10.1088/1748-3182/7/3/036009. [21] Anthes, Emily. "Cuttlefish provide smart fashion tips" (http:/ / www. bbc. com/ future/ story/ 20120907-smart-fashion-tips-by-cuttlefish). BBC.com. . Retrieved 30 October 2012.

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External links
TONMO.com Community Forum Keeping cuttlefish in the home aquarium (http://www.tonmo.com/forums/ forumdisplay.php?f=18) University of Texas Medical Branch's Cephbase (http://web.archive.org/web/20060118140436/http://www. cephbase.utmb.edu/); photos and videos of cuttlefish and other cephalopods (archived version dated January 18, 2006) NO LONGER UPDATED, from the Wayback Machine. 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) Cuttlefish: Kings of Camouflage (http://www.pbs.org/wgbh/nova/camo/) NOVA episode on PBS.org Interactive: Anatomy of a Cuttlefish (http://www.pbs.org/wgbh/nova/nature/anatomy-cuttlefish.html) on NOVAbeta l

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Nautiloid
Nautiloids Temporal range: Late Cambrian Recent

Orthoceras

Scientific classification Kingdom: Phylum: Class: Subclass: Animalia Mollusca Cephalopoda Nautiloidea<br 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.

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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 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.

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Modern nautiloids
Much of what is known about the extinct nautiloids is based on what we know about modern nautiluses, such as the chambered nautilus, which is found in the southwest Pacific Ocean from Samoa to the Philippines, and in the Indian Ocean off 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 (2,300ft). Nautiluses are free swimming animals that possess a head with two simple lens-free eyes and arms (or tentacles). They have a smooth shell over a large body chamber, which is divided into subchambers filled with Nautilus belauensis an inert gas (similar to the composition of atmospheric air, but with more nitrogen and less oxygen) making the animal neutrally buoyant in the water. As many as 90 tentacles are arranged in two circles around the mouth. The animal is predatory, and has jaws which are horny and beak-like, allowing it to feed on crustaceans. 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, allowing the empty shell to be carried some distance from where the animal lived before finally sinking to the seafloor. 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. Unlike the belemnites and other cephalopods, modern nautiluses do not have an ink sac, and there is no evidence to suggest that the extinct forms possessed one either. Furthermore, unlike the extinct ammonoids, the modern nautilus lacks any sort of plate for closing its 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 nautilus is quite prominent, and, although somewhat rarely, the shell coloration has been known to be preserved in fossil nautiloids. They often show color patterns only on the dorsal side, suggesting that 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' shellsespecially in the late Paleozoic and early Mesozoicare ornamented with spines and ribs, but most have a smooth shell. 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

Fossil nautiloid Trilacinoceras from the Ordovician of China.

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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. Specimens of the Ordovician nautiloid Endoceras have been recorded measuring up to 3.5 meters (13ft) in length, and Cameroceras is (somewhat doubtfully) estimated to have reached 11 meters (36ft). These large nautiloids must have been formidable predators of other marine animals at the time they lived.

Fossil orthoconic nautiloid from the Ordovician of Kentucky; an internal mold showing siphuncle and half-filled camerae, both encrusted.

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.

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 this 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.

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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: 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 Endocerida, Actinocerida, and Discosorida. Three unite related orders that share a common ancestor and form a branch of the nautiloid taxonomic tree; Plectronoceratoidea, which consists mostly of small Cambrian forms that include the ancestors of subsequent stocks; Orthoceratoidea, which unites different primarily orthoconic orders of which one is the source for Bacritida and Ammonoidea; and Nautilitoidea, which includes the first coilded cephalopods, Tarphycerida, as well as Nautilida, which includes the recent Nautilus. Another order, Bactritida, which is derived from Orthocerida, is sometimes included with Nautiloidea, sometimes with Ammonoidea, and sometimes placed in a subclass of its own, Bactritoidea. Recently some workers in the field have come to recognize Dissidocerida as a distinct order, along with Pseudorthocerida, both previously included in Orthoceridaas subtaxa. A more recent interpretation (Engeser 1997-1998) suggests that nautiloids, and indeed cephalopods in general, fall into two main groups, Palcephalopoda (including all the nautiloids except Orthocerida and Ascocerida) and Neocephalopoda (the rest of the cephalopods).

References
[1] Stehli, F. G. (8 June 1956). "Shell Mineralogy in Paleozoic Invertebrates". Science 123 (3206): 10311032. Bibcode1956Sci...123.1031S. doi:10.1126/science.123.3206.1031. ISSN0036-8075. JSTOR1750042. PMID17800970. [2] Krger, 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 (04). doi:10.1017/S0016756808004585. [3] Krger, B.; Yun-bai, Y. B. (2009). "Pulsed cephalopod diversification during the Ordovician". Palaeogeography Palaeoclimatology Palaeoecology 273: 174201. 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., (19971998) 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,

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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)

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Nautilus
Nautilus

Nautilus belauensis

Scientific classification Kingdom: Phylum: Class: Subclass: Order: Superfamily: Family: Animalia Mollusca Cephalopoda Nautiloidea Nautilida Nautilaceae Nautilidae Blainville, 1825 Genera Carinonautilus Cenoceras Eutrephoceras Pseudocenoceras Strionautilus Allonautilus Nautilus Synonyms[1]

Eutrephoceratidae Miller, 1951

Nautilus (from Greek , 'sailor') is the common name of pelagic marine mollusks of the 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.[2] Having survived relatively unchanged for millions of years, nautiluses represent the only living members of the subclass Nautiloidea, and are often considered "living fossils."

Nautilus The name "nautilus" originally referred to the pelagic octopuses of the genus Argonauta, otherwise known as paper nautiluses, as the ancients believed these animals used their two expanded arms as sails.[3]

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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, and 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.[4]:56 Nautilus pompilius is the largest species in the genus. One form from northwestern Australia, once called Nautilus repertus, may reach 26.8 centimetres (10.6in) in diameter.[5] However, most nautilus species never exceed 20 centimetres (7.9in). Nautilus macromphalus is the smallest species, usually measuring only 16 centimetres (6.3in). A dwarf population from the Sulu Sea (Nautilus pompilius suluensis) is even smaller, with a mean shell diameter of 115.6mm.[6]

The anatomy of Nautilus. The top figure is dissected; the bottom just has the shell removed.

Shell

Nautilus hemishell showing the camerae in a logarithmic spiral

A nautilus shell viewed from above (left), and from underneath (right)

Nautilus Nautili are the sole living cephalopods whose bony body structure is externalized as a shell. The animal can withdraw completely into its shell and close the opening with a leathery hood formed from two specially folded tentacles. The shell is coiled, aragonitic,[7] nacreous and pressure resistant, imploding at a depth of about 800 metres (2,600ft). 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 osmea pearl, contrarily to its name, is not a pearl, but a jewellery 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 moment of hatching to thirty or more in adults. The shell colouration 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.

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Tentacles
Nautilus tentacles differ from those of other cephalopods. Lacking pads, the tentacles stick to prey by virtue of their ridged surface.[8] Nautiloids have a powerful grip. Attempts to take an object already seized by a nautilus may tear away the creature's tentacles, which remain firmly attached to the surface of the object.[8] 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.[8][9][10]

Physiology
Buoyancy and movement
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 only in long term density changes by osmosis, either removing liquid from its chambers or allowing water from the blood in the siphuncle to slowly refill the chambers. This is done in response to sudden changes in buoyancy that can occur with predatory attacks of fish, which can break off parts of the shell. This limits nautiluses in that they cannot operate under the extreme hydrostatic pressures found at depths greater than approximately 800 metres (2,600ft), and in fact implode at about that depth, causing instant death.[6] The maximum depth at which they can regulate buoyancy by osmotic removal of chamber liquid is not known.[11]
Nautilus with extended tentacles and hyponome visible

Nautilus

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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.[12]

Brain and intelligence

Head of Nautilus pompilius showing the eye Nautiluses are much closer to the first cephalopods that appeared about 500 million years ago than the early modern cephalopods that appeared maybe 100 million years later (ammonoids and coleoids). They have a seemingly simple brain, not the large complex brains of octopuses and squid, and had long been assumed to lack intelligence. But the cephalopod nervous system is quite different from that of other animals, and recent experiments have shown not only memory, but a changing response to the same event over time.[13][14][15]

In a study in 2008, a group of nautiluses (N. pompilius) were given food as a bright blue light flashed until they began to associate the light with food, extending their tentacles every time the blue light was flashed. The blue light was again flashed without the food 3 minutes, 30 minutes, 1 hour, 6 hours, 12 hours, and 24 hours later. The nautiluses continued to respond excitedly to the blue light for up to 30 minutes after the experiment. An hour later they showed no reaction to the blue light. However, between 6 and 12 hours after the training, they again responded to the blue light, but more tentatively. The researchers concluded that nautiluses had memory capabilities similar to the "short-term" and "long-term memories" of the more advanced cephalopods, despite having different brain structures.[13][14][15] However the long-term memory capability of nautiluses was much shorter than that of other cephalopods. The nautiluses completely forgot the earlier training 24 hours later, in contrast to octopuses, for example, which can remember conditioning for weeks afterwards. However, this may simply be the result of the conditioning procedure being suboptimal for sustaining long-term memories in nautiluses. Nevertheless the study showed that scientists had previously underestimated the memory capabilities of nautiluses.[15]

Reproduction and lifespan

Nautiluses reproduce by laying eggs. Gravid females attach the fertilized eggs to rocks in shallow waters, whereupon the eggs take eight to twelve months to develop until the 30 millimetres (1.2in) juveniles hatch. Females spawn once per year and regenerate their gonads, making nautiluses the only cephalopods to present iteroparity or polycyclic spawning.[16] 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. At sexual maturity, the male shell becomes slightly larger than the female's.[17] Males have been found to greatly outnumber females in practically all published studies, accounting for 60 to 94% of all recorded individuals at different sites (only one study from the Philippines found females to be more abundant, with males representing only 32% of the population).[6] The lifespan of nautiluses may exceed 20 years, which is exceptionally lengthy for a cephalopod.[18] However, nautiluses typically reach sexual maturity when they are about 15 years old.[6]

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Ecology
Range, habitat, and status
Nautiluses are only found in the Indo-Pacific, from 30 N to 30 S latitude and 90 to 175 E longitude. They inhabit the deep slopes of coral reefs. Nautiluses usually inhabit depths of several hundred metres. It has long been believed that nautiluses rise at night to feed, mate and lay eggs, but it appears that, in at least some populations, the vertical movement patterns of these animals are far more complex.[19] The greatest depth at which a nautilus has been sighted is 703m (N.pompilius).[19] Implosion depth for nautilus shells is thought to

Number of captured N.pompilius at various depths around the Osprey Reef Seamount, Coral Sea. The data was collated from 271 trapping events spread across all months of the year. Nautiluses were most common at 300350m. No specimens were recovered from a [6] depth of less than 150m during 18 trapping efforts.

Nautilus be around 800m.[6][19] Only in New Caledonia, the Loyalty Islands, and Vanuatu can nautiluses be observed in very shallow water (at a depth of as little as 5m).[11][19] This is due to the cooler surface waters found in these southern hemisphere habitats as compared to the many equatorial habitats of other nautilus populations (these usually being restricted to depths greater than 100m).[11][19] Nautiluses generally avoid water temperatures above 25C.[19] There is growing concern that nautiluses are being greatly overfished.[20] Their limited ecological range and the late onset of their sexual maturity combined with this overfishing has led to recent investigations into the need to protect them from possible endangerment or extinction.[21]

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Diet
A pair of N.pompilius feeding on two-spot red snapper (Lutjanus bohar) bait during daytime at 703m depth. This observation constitutes the deepest record of any nautilus species.

Nautiluses are scavengers and opportunistic predators.[22][23] They eat molts of lobsters, hermit crabs, and carrion of any kind.[11]

Evolution
Fossil records indicate that nautiloids 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 Late Cambrian period and became a significant group of sea predators during the Ordovician period. Certain species reached over 2.5 metres (8ft2in) in size. The other cephalopod subclass, Coleoidea, diverged from the nautiloids 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.

Shell characters of the genera Nautilus and Allonautilus

The Nautilidae has its origin in the Trigonocerataceae (Centroceratina), specifically in the Syringonautilidae of the Late Triassic[2] and continues to this day with Nautilus, the type genus, and its close relative, Allonautilus.

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 Eutrephoceras dorbignyanum 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.

Nautilus 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.

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Taxonomy
The family Nautilidae contains up to 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 Recent genetic data has pointed to there being only three extant species: A.scrobiculatus, N.macromphalus, and N.pompilius, with N.belauensis and N.stenomphalus both subsumed under N.pompilius, possibly as subspecies.[6]

Nautilus shells: N. macromphalus (left), A. scrobiculatus (centre), N. pompilius (right)

Dubious or uncertain taxa

The following taxa associated with the family Nautilidae are of uncertain taxonomic status:[24]
Binomial name and author citation N. alumnus Iredale, 1944 Current systematic status Type locality Type repository

Species dubium [fide Saunders (1987:49)]

Queensland, Australia Not designated

Not designated [fide Saunders (1987:49)] Unresolved

N. ambiguus Sowerby, 1848 N. beccarii Linne, 1758

Species dubium [fide Saunders (1987:48)] Non-cephalopod; Foraminifera [fide Frizzell and Keen (1949:106)] ?Non-cephalopod; Foraminifera Lenticulina

N. calcar Linne, 1758

Adriatic Sea

Unresolved; Linnean Society of London? Unresolved; Linnean Society of London?

N. crispus Linne, 1758

Undetermined

Mediterranean Sea

N. crista Linne, 1758

Non-cephalopod; Turbo [fide Dodge (1953:14)]

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Undetermined Adriatic Sea Unresolved; Linnean Society of London? Unresolved; Linnean Society of London?

N. fascia Linne, 1758

N. granum Linne, 1758

Undetermined

Mediterranean Sea

N. lacustris Lightfoot, 1786 N. legumen Linne, 1758

Non-cephalopod; Helix [fide Dillwyn (1817:339)] 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? Unresolved

N. pompilius marginalis Willey, Species dubium [fide Saunders (1987:50)] 1896 N. pompilius moretoni Willey, 1896 N. pompilius perforatus Willey, 1896 N. radicula Linne, 1758 Species dubium [fide Saunders (1987:49)]

New Guinea

New Guinea

Unresolved

Species dubium [fide Saunders (1987:49)]

New Guinea

Unresolved

?Non-cephalopod; Foraminifera Nodosaria

Adriatic Sea

Unresolved; Linnean Society of London? Unresolved; Linnean Society of London? Unresolved; Linnean Society of London? Unresolved; Linnean Society of London? Unresolved; Linnean Society of London?

N. raphanistrum Linne, 1758

Undetermined

Mediterranean Sea

N. raphanus Linne, 1758

Undetermined

Adriatic Sea

N. semi-lituus Linne, 1758

Undetermined

Liburni, Adriatic Sea "freto Siculo"

N. sipunculus Linne, 1758

Undetermined

N. texturatus Gould, 1857 Octopodia nautilus Schneider, 1784

Nomen nudum Rejected specific name [fide Opinion 233, ICZN (1954:278)]

In popular culture
The nautilus shell features prominently in the official emblem of New Caledonia.

References
Notes
[1] Cichowolski, M.; Ambrosio, A.; Concheyro, A. (2005). "Nautilids from the Upper Cretaceous of the James Ross Basin, Antarctic Peninsula". Antarctic Science 17 (2): 267. doi:10.1017/S0954102005002671. [2] Kmmel,B. 1964. Nautiloidae-Nautilida, in the Treatise on Invertebrate Paleontology, Geological Society of America and Univ of Kansas Press, Teichert and Moore eds. [3] cf. Aristotle Historia Animalium 622b [4] 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: 794. . [5] Nautilus repertus ID:118764 (http:/ / www. conchology. be/ ?t=68& u=118764& g=0d16facb6ebae4a31321e9f6e6298f46& q=a1d787a5674deb7b193549c6ccdff907). Shell Encyclopedia, Conchology, Inc. [6] Dunstan AJ; Ward PD; Marshall NJ (February 2011). Solan, Martin. ed. "Nautilus pompilius life history and demographics at the Osprey Reef Seamount, Coral Sea, Australia" (http:/ / www. plosone. org/ article/ info:doi/ 10. 1371/ journal. pone. 0016312). PLoS ONE 6 (2): e16312. doi:10.1371/journal.pone.0016312. PMC3037366. PMID21347356. .

Nautilus
[7] "Diagenesis of aragonite from Upper Cretaceous ammonites: a geochemical case-study". Sedimentology 28 (3): 423438. 1981. Bibcode1981Sedim..28..423B. doi:10.1111/j.1365-3091.1981.tb01691.x. [8] 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): 197201. . [9] 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. 249256. doi:10.1007/978-90-481-3299-7_17 [10] 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. 257269. doi:10.1007/978-90-481-3299-7_18 [11] Ward, P.D. (1987). The Natural History of Nautilus. Allen and Unwin, London. [12] Grasso, F.; Basil, J. (2009). "The evolution of flexible behavioral repertoires in cephalopod molluscs". Brain, Behavior and Evolution 74 (3): 231245. doi:10.1159/000258669. PMID20029186. [13] Ewen Callaway (2 June 2008). "Simple-Minded Nautilus Shows Flash of Memory" (http:/ / www. newscientist. com/ article/ dn14033-simpleminded-nautilus-reveals-flash-of-memory. html). New Scientist. . Retrieved 7 March 2012. [14] Kathryn Phillips (15 June 2008). "Living Fossil Memories" (http:/ / jeb. biologists. org/ content/ 211/ 12/ iii. full. pdf#page=1& view=FitH). Inside JEB 211 (12): iii. doi:10.1242/jeb.020370. . [15] Robyn Crook & Jennifer Basil (2008). "A biphasic memory curve in the chambered nautilus, Nautilus pompilius L. (Cephalopoda: Nautiloidea)" (http:/ / jeb. biologists. org/ content/ 211/ 12/ 1992. full. pdf#page=1& view=FitH). The Journal of Experimental Biology 211 (12): 19921998. doi:10.1242/jeb.018531. . [16] Rocha, F.; Guerra, .; Gonzlez, . F. (2001). "A review of reproductive strategies in cephalopods". Biological Reviews of the Cambridge Philosophical Society 76 (3): 291304. doi:10.1017/S1464793101005681. PMID11569786. [17] Bruce Saunders, W.; Spinosa, C. (1978). "Sexual Dimorphism in Nautilus from Palau". Paleobiology 4 (3): 349358. doi:10.2307/2400210. JSTOR2400210. [18] Saunders WB (June 1984). "Nautilus Growth and Longevity: Evidence from Marked and Recaptured Animals". Science 224 (4652): 990992. Bibcode1984Sci...224..990S. doi:10.1126/science.224.4652.990. PMID17731999. [19] Dunstan, A. J.; Ward, P. D.; Marshall, N. J. (2011). Solan, Martin. ed. "Vertical distribution and migration patterns of Nautilus pompilius". PLoS ONE 6 (2): e16311. doi:10.1371/journal.pone.0016311. PMC3043052. PMID21364981. [20] Broad WJ (24 October 2011). "Loving the Chambered Nautilus to Death" (http:/ / www. nytimes. com/ 2011/ 10/ 25/ science/ 25nautilus. html?_r=4). New York Times. . Retrieved 25 October 2011. [21] Dunstan AJ; Bradshaw CJA; Marshall NJ (February 2011). Solan, Martin. ed. "Nautilus at Risk Estimating Population Size and Demography of Nautilus pompilius" (http:/ / www. plosone. org/ article/ info:doi/ 10. 1371/ journal. pone. 0016716). PLoS ONE 6 (2): e16716. doi:10.1371/journal.pone.0016716. PMC3037370. PMID21347360. . [22] Saunders, W.B. (1984). "The role and status of Nautilus in its natural habitat: Evidence from deep-water remote camera photosequences". Paleobiology 10 (4): 469486. JSTOR2400618. [23] Wells, M. J.; Wells, J.; O'Dor, R. K. (2009). "Life at low oxygen tensions: The behaviour and physiology of Nautilus pompilius and the biology of extinct forms". Journal of the Marine Biological Association of the United Kingdom 72 (2): 313328. doi:10.1017/S0025315400037723. [24] 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.

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Bibliography Ward, P.D. 1988. In Search of Nautilus. Simon and Schuster. W. Bruce Saunders and Neil H. Landman (2010), "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), Topics in Geobiology (Dordrecht : Springer Science+Business Media B.V) 6, ISBN978-90-481-3299-7 CephBase: Nautilidae (http://www.cephbase.utmb.edu/spdb/familygroup.cfm?Family=Nautilidae)

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External links
Nautilidae discussion forum (http://www.tonmo.com/forums/forumdisplay.php?f=49), tonmo.com Waikk 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

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Ammonite
Ammonites Temporal range: Devonian - Cretaceous

Artist's reconstruction of Asteroceras

Scientific classification Kingdom: Phylum: Class: Subclass: Animalia Mollusca Cephalopoda < Orders and Suborders See text Ammonites /mnats/ 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 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 nonspiraled 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 () for "horn".

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Diagnostic characters
Ammonites (subclass Ammonoidea) can be distinguished by their septa, the dividing walls that separate the chambers in the phragmocone, by the nature of their sutures where the septa joint the outer shell wall, and in general by their siphuncles.

Septa
Ammonoid septa characteristically have bulges and indentations and are to varying degrees convex from the front, distinguishing them from nautiloid septa which are typically simple concave dish-shaped structures. The topology of the septa, especially around the rim, results in the various suture patterns found.

Suture patterns
Three major types of suture patterns are found in the Ammonoidea: 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.

Siphuncle
The siphuncle in most ammonoids by far is a narrow tubular structure that runs along the outer rim, known as the venter, connecting the chambers of the phragmocone to the body or living chamber. This distinguishes them from living nautiloides (Nautilus and Allonautilus) and typical Nautilida. However, the very earliest nautiloids from the Late Cambrian and Ordovician typically had ventral siphuncles, although often proportionally larger than those in ammonites and more internally structured. The word "siphuncle" comes from the New Latin siphunculus, meaning "little siphon".

Classification of the Ammonoidea

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 Mya) 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) is variably folded, forming saddles (or peaks) and lobes (or valleys).
An ammonite shell viewed in section, revealing the internal chambers and septa. Large polished examples are prized for their aesthetic, as well as scientific, value.

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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 Prolecanitida, Upper Devonian - Upper Triassic Ceratitida, Permian - Triassic Phylloceratida, Triassic - Cretaceous Lytoceratida, Jurassic - Cretaceous Ammonitida, Lower Jurassic - Upper Cretaceous In some classifications, these are left as suborders, included in only three orders: Goniatitida, Ceratitida, and Ammonitida.
An ammonitic ammonoid with the body chamber missing, showing the septal surface (especially at right) with its undulating lobes and saddles.

Taxonomy of the Treatise

The Treatise on Invertebrate Paleontology (Part L, 1957) divides the Ammonoidea, regarded simply as an order, into eight suborders, the Iridescent ancient ammonite fossil on display at Anarcestina, Clymeniina, Goniatitina, and Prolecanitina from the the American Museum of Natural History, New York City, around 2.5 feet in diameter Paleozoic; the Ceratitina from the Triassic; and the Ammonitina, Lytoceratina, and Phylloceratina from the Jurassic and Cretaceous. In subsequent taxonomies, these are sometimes regarded as orders within the subclass Ammonoidea.

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, because their fossils are often found in rocks 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 bottom-dwellers. Fossil shell of ammonite Placenticeras whitfieldi Synchrotron analysis of an aptychophoran ammonite revealed showing punctures caused by the bite of a mosasaur, Peabody Museum of Natural History, Yale remains of isopod and mollusc larvae in its buccal cavity, indicating at least this kind of ammonite fed on plankton.[2] 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

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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] Many ammonite shells have been found with round holes once interpreted as a result of limpets attaching themselves to the shells. The triangular formation of the holes, their size and shape, and their presence on both sides of the shells, corresponding to the upper and lower jaws, is evidence of the bite of a medium-sized mosasaur preying upon ammonites.

Shell anatomy and diversity

Jeletzkytes, a Cretaceous ammonite from the USA

Asteroceras, a Jurassic ammonite from England

Orthosphynctes , a Jurassic ammonite from Portugal

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Basic shell anatomy

The chambered part of the ammonite shell is called a phragmocone. It 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 the siphuncle of ammonites (excepting Clymeniina) runs along the ventral periphery of the septa and camerae (i.e., the inner surface of the outer axis of the shell), while the siphuncle of nautiloids runs more or less through the center of the septa and camerae.

A variety of ammonite forms, from Ernst Haeckel's 1904 Kunstformen der Natur (Art Forms of Nature)

Baculites ammonite from the Late Cretaceous of Wyoming: The original aragonite of the outer conch and inner septa has dissolved away, leaving this articulated internal mold.

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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 (the macroconch) being female, and the smaller shell (the 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. Only recently has sexual variation in the shells of ammonites been Discoscaphites iris, Owl Creek Formation (Upper recognized. The macroconch and microconch of one species were often Cretaceous), Ripley, Mississippi. previously mistaken for two closely related but different species occurring in the same rocks. However because these "pairs" were so consistently found together it became apparent they were, in fact, sexual forms of the same species.

Variations in shape
The majority of ammonite species feature planispiral, flat-coiled shells, but other species feature nearly straight (as in baculites) shells. 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. Upon closer inspection, though, 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.

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Aptychus
Some ammonites have been found in association with a single horny plate or a pair of calcitic plates. In the past, these plates were assumed to serve in closing the opening of the shell in much the same way as an operculum, but more recently, they are postulated to have been, 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. Anaptychi are relatively rare as fossils. They are found representing ammonites from the Devonian period through those of the Cretaceous period.

A drawing of an aptychus which was mistakenly described as a bivalve and given the name "Trigonellites latus", from the Kimmeridge Clay Formation in England

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.

Soft-part anatomy
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.[9] They likely bore a radula and beak, a marginal siphuncle, and ten arms.[10] They operated by direct development with sexual reproduction, were carnivorous, and had a crop for food storage. They are unlikely to have dwelt in fresh or brackish water.[11]

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Size
Few of the ammonites occurring in the lower and middle part of the Jurassic period reached a size exceeding 23 cm (9in) 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 cm (2 ft) in diameter, and Parapuzosia seppenradensis of the Cretaceous period of Germany, which is one of the largest known ammonites, sometimes reaching 2 m (6.5ft) in diameter. The largest documented North American ammonite is Parapuzosia bradyi from the Cretaceous, with specimens measuring 137 cm (4.5ft) in diameter, although a new 2.3-m (7.5-ft) British Columbian specimen, if authentic, would appear to trump even the European champion.[12]

2-metre (6.5-foot) Parapuzosia seppenradensis cast in Germany

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 as to prevent the establishment of animal life on the seafloor. When A specimen of Hoploscaphites from the Pierre upon death the ammonites fell to this seafloor and were gradually Shale of South Dakota: Much of the original 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, a concretion, 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.

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Other fossils, such as many found in Madagascar and Alberta, 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; the outer shell (composed of aragonite)[13] has been lost during the fossilization process. Only in these internal-mold specimens can the suture lines be observed; in life, the sutures would have been hidden by the outer shell. The ammonoids as a group continued through several major extinction events, although often only a few species survived. Each time, An iridescent ammonite from Madagascar 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 Mya in the CretaceousPaleogene extinction event. The reason why no ammonites survived the extinction event at the end of the Cretaceous, whereas some nautiloids 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 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 Ammonites in the permanent collection of The the Cretaceous, the order having gone through a more or less steady Childrens Museum of Indianapolis 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 the Cretaceous ammonites, being closely related to coleoids, had a similar reproductive strategy in which huge numbers of eggs were laid in a single batch at the end of the lifespan. 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 conversely thought to have had a reproductive strategy in which eggs were laid in smaller batches many times during the lifespan, and on the sea floor well away from any direct effects of such a bolide strike, and thus survived.

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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 Hilda of Whitby, a myth referenced in Sir Walter Scott's Marmion,[14] 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.[15] 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.[16]

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 This citation will be automatically completed in the next few minutes. You can jump the queue or expand by hand (http:/ / en. wikipedia. org/ wiki/ Template:cite_doi/ . 0910. 1007. 2f978-1-4020-6806-5_11?preload=Template:Cite_doi/ preload& editintro=Template:Cite_doi/ editintro& action=edit) [4] "Introduction to Ammonoidea" (http:/ / www. bbm. me. uk/ portsdown/ PH_232_Ammonites. htm). The Geology of Portsdown Hill. Archived (http:/ / web. archive. org/ web/ 20070502202153/ http:/ / www. bbm. me. uk/ portsdown/ PH_232_Ammonites. htm) from the original on 2 May 2007. . Retrieved 2007-04-26. [5] Morton, N. 1981. Aptychi: the myth of the ammonite operculum. Lethaia 14(1): 5761. 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): 231238. 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: 325331. 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: 2032. doi:10.2475/ajs.293.A.20 [9] 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 (5): 10931107. doi:10.1111/j.0031-0239.2004.00408.x. [10] Landman, Neil H; Tanabe, Kazushige; Davis, Richard Arnold (1996). Ammonoid paleobiology (http:/ / books. google. ca/ books?id=hKbkB4MzUIkC& pg=PA14). ISBN978-0-306-45222-2. . [11] Landman, Neil H; Tanabe, Kazushige; Davis, Richard Arnold (1996). Ammonoid paleobiology (http:/ / books. google. ca/ books?id=hKbkB4MzUIkC& pg=PA17). ISBN978-0-306-45222-2. . [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 (3): 423438. 1981. Bibcode1981Sedim..28..423B. doi:10.1111/j.1365-3091.1981.tb01691.x. [14] Lovett, Edward (September 1905). "The Whitby Snake-Ammonite Myth" (http:/ / en. wikisource. org/ wiki/ Folk-Lore. _Volume_16/ The_Whitby_Snake-Ammonite_Myth). Folk-Lore 16 (3): 3334. . [15] Cadbury, D. the Dinosaur Hunters. (Fourth Estate, 2000) (ISBN 1-85702-963-1), p.7 [16] "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.

Ammonite 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)

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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. William R. Wahl * MOSASAUR BITE MARKS ON AN AMMONITE. PRESERVATION OF AN ABORTED ATTACK? (http://www.wyodino.org/wp-content/uploads/2011/01/BittenAmmonite.pdf) Mosasaur diet (http://palaeoart.blogspot.it/2011/10/mosasaur-diets-everhart-2004.html)

Belemnoidea

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Belemnoidea
Belemnoids

Belemnite guards from the Jurassic of Wyoming, USA.

Scientific classification Orders Aulacocerida Phragmoteuthida Belemnitida Diplobelida 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 belemnoids 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] Belemnoids include belemnites (which belong to order Belemnitida proper), aulacocerids (order Aulacocerida), phragmoteuthids (order Phragmoteuthida), and diplobelids (order Diplobelida).

Occurrence
Belemnoids were numerous during the Jurassic and Cretaceous periods, and their fossils are abundant in Mesozoic marine rocks, often accompanying their cousins the ammonites. The belemnoids become Artist's reconstruction of belemnoids. extinct at the end of the Cretaceous period along with the ammonites. The belemnoids' origin lies within the bactritoid nautiloids, which date from the Devonian period; well-formed belemnoid 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.

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Anatomy
Belemnoids 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 (towards the head) 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, belemnoid arms carried a series of small hooks for grabbing prey. Belemnoids 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.

Ecology
Belemnoids 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 belemnoids only the back part of the shell (called the guard or rostrum) is found. The guard is usually elongated and bullet-shaped (though in some subgroups the rostrum may only exist as a thin layer coating the phragmocone). 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 preserved specimens. Projecting forwards from one side of the phragmocone is

A belemnoid fossil from the Franconian Jura.

the thin pro-ostracum. While belemnoid phragmocones are homologous with the shells of other cephalopods and are similarly composed of aragonite, belemnoid guards are evolutionarily novel and are composed of calcite or aragonite, 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 belemnoid Megateuthis gigantea, which is found in Europe and Asia, can measure up to 46 centimetres (18in) in length, giving the living animal an estimated length of 3 metres (9.8ft).

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Very exceptional belemnoid 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
The bulk geochemical signature contained within belemnoid guards of 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.

belemnoid fossils at Bristol City Museum, Bristol, England. Found in the Lower Lias strata, Gloucestershire, England.

Some belemnoids (such as Belemnites of Belemnitida) 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 Belemnotheutis Order Aulacocerida Family Aulacoceratidae Family Dictyoconitidae Family Hematitidae Family Palaeobelemnopseidae Family Xiphoteuthidae Order Belemnitida Suborder Belemnitina Family Cylindroteuthidae Family Hastitidae Family Oxyteuthidae Family Passaloteuthidae Family Salpingoteuthidae Suborder Belemnopseina Family Belemnitellidae
Fossilised belemnite belemnoids in the very top bedding plane of the Zohar Formation (Jurassic) near Neve Atif, the Golan. Note the central fold along the axis characteristic of some genera.

Family Belemnopseidae Family Dicoelitidae Family Dimitobelidae Family Duvaliidae Suborder Belemnotheutina

Belemnoidea Family Belemnotheutidae Family Chitinobelidae Family Sueviteuthidae Order Diplobelida Family Chondroteuthidae Family Diplobelidae Order Phragmoteuthida Family Phragmoteuthidae Family Rhiphaeoteuthidae

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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 (2): 267287. 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 (4): 983998. 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/ ). Palontologische Zeitschrift 70: 425431. 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

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Argonaut
For other uses, see Argonaut (disambiguation). Argonauts

Female Argonauta argo with eggs

Scientific classification Kingdom: Phylum: Class: Order: Superfamily: Family: Genus: Animalia Mollusca Cephalopoda Octopoda Argonautoida Argonautidae 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 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 funnelmantle 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 , 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.

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Physical description
Sexual dimorphism and reproduction
Argonauts exhibit extreme sexual dimorphism in size and lifespan. Females grow up to 10cm and make shells up to 30cm, while males rarely surpass 2cm. 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 detaches a special arm from his body, the hectocotylus, to transfer sperm to the female. For fertilization, the 'arm'(thingy) 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. nodosa

Juvenile female A. hians

Immature male A. hians

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

Argonaut eggcases are occasionally found floating on the sea, sometimes with the female argonaut clinging to it. It is not made 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 CretaceousPaleogene 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 300mm.[9][10] The smallest species is Argonauta bottgeri, with a maximum recorded size of 67mm.[9][11]

85

Female A. nodosa with its eggcase

The eggcase of A. argo

The eggcase of A. nodosa

The eggcase of A. hians

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.[11]

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.[11] Male argonauts have been observed residing inside salps, although little is known about this relationship.[12]

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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.[13]
Shells of various Argonauta species

Dubious or uncertain taxa

The following taxa associated with the family Argonautidae are of uncertain taxonomic status:[14]
Binomial name and author citation Argonauta arctica Fabricius, 1780 Argonauta bibula Rding, 1798 Argonauta compressa Blainville, 1826 Current systematic status Type locality Type repository

Undetermined

Unresolved; ?Tullukaurfak, Greenland Unresolved

Unresolved

Undetermined

Unresolved

Undetermined

Mer de Indes

Unresolved; [other Blainville types at MNHN] [not reported by Lu et al. (1995)] Unresolved

Argonauta conradi Parkinson, 1856 Argonauta cornu Gmelin, 1791 Argonauta cymbium Linn, 1758

Species of uncertain status [fide Robson (1932:200)] Undetermined

"New Nantucket, Pacific Ocean"

Unresolved

Unresolved; LS?

Non-cephalopod; foraminiferous shell [fide Von Martens (1867:103) Not designated Unresolved

Argonauta fragilis Parkinson, Species of uncertain status [fide 1856 Robson (1932:200)] Argonauta geniculata Gould, 1852 Argonauta maxima Dall, 1871 Species of uncertain status [fide Robson (1932:200)] Nomen nudum

Near Sugarloaf Mountain, Rio de Janeiro, Brazil

Type not extant [fide Johnson (1964:32)]

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87
Species dubium [fide Rehder (1967:11)] Non-cephalopod; Carcinaria sp. [fide Robson (1932:201)] Incertae sedis [fide Robson (1932:181)] Nomen nudum "Indian seas" ["South Pacific ocean" fide Owen (1842:114)] Unresolved; Museum of the Royal College of Surgeons? Holotype Not designated Unresolved

Argonauta navicula Lightfoot, 1786 Argonauta rotunda Perry, 1811 Argonauta rufa Owen, 1836

Argonauta sulcata Lamarck, 1801 Argonauta tuberculata f. aurita Von Martens, 1867 Argonauta tuberculata f. mutica Von Martens, 1867 Argonauta tuberculata f. obtusangula Von Martens, 1867 Argonauta vitreus Gmelin, 1791 Octopus (Ocythoe) raricyathus Blainville, 1826 Ocythoe punctata Say, 1819

Undetermined

Unresolved

ZMB

Undetermined

Coast of Brazil

ZMB Holotype

Undetermined

Not designated

ZMB Syntypes

Undetermined

Not designated

Unresolved; LS?

Undetermined [Argonauta?]

Not designated

MNHN Holotype; specimen not extant [fide Lu et al. (1995:323)] Unresolved; ANSP? Holotype [not traced by Spamer and Bogan (1992)]

Argonauta sp. [fide Robson (1929d:215)]

Atlantic Ocean near the North American coast (from stomach of dolphin) 442856N 464815W (Atlantic Ocean)

Tremoctopus hirondellei Joubin, 1895

Argonauta or Ocythoe [fide Thomas (1977:386)]

MOM Holotype [station 151] [fide Belloc (1950:3)]

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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.[15] A variation known as the double argonaut design was also found in Minoan jewelry.[16]

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.[17] 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: 1863. [2] (Italian) Delle Chiaje, S. (1825). Memorie sulla storia e notomia degli animali. Senza Vertebre del Regno di Napoli. I. [3] Nixon, M. & J.Z. Young (2003). The Brains and Lives of Cephalopods. Oxford University Press. Argonauts surrounding the Nautilus, in Jules Verne's novel Twenty Thousand Leagues Under the Sea

[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): 520531. doi:10.1666/0022-3360(2005)079<0520:FAMCOF>2.0.CO;2. ISSN0022-3360. . [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] Clarke, M. R. (1986). A Handbook for the Identification of Cephalopod Beaks. Oxford University Press. pp.273 pp. [12] 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: 648650. [13] 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. [14] 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. [15] Eleni M. Konstantinidi, Jewellery Revealed in the Burial Contexts of the Greek Bronze Age, 2001, Hadrian Books, 322 pages ISBN 1-84171-165-9 [16] C.Michael Hogan, Knossos Fieldnotes, The Modern Antiquarian (2007) (http:/ / themodernantiquarian. com/ site/ 10857/ phaistos. html#fieldnotes)

Argonaut
[17] Johann David Wyss and Jenny H. Stickney, The Swiss Family Robinson, Ginn & Co., 1898, 364 pages

89

External links
Tree of Life web project: Argonauta (http://tolweb.org/tree?group=Argonauta)

90

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 An octopus in a zoo creatures. Classical conditioning of cephalopods has been reported, and 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, an ability 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] An octopus at the Sea Star Aquarium in Coburg named Otto was known to juggle his fellow tankmates around, 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]

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91

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
The octopus is one of the prime examples of an invertebrate animal which has repeatedly been shown to exhibit flexibility in its use of 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.[11] It is surmised that the octopuses originally used bivalves for the same purpose, before humans made coconut shells widely available on the sea floor.[12][13] Most hermit crabs use discarded shells of other species for habitation A small octopus (4-5 cm diameter) individual and other crabs choose sea anemones to cultivate on their carapaces as using a nut shell and clam shell as shelter. 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.

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Nautiluses
Nautiluses are much closer to the first cephalopods that appeared about 500 million years ago than the early modern cephalopods that appeared maybe 150 million years later. They have a seemingly simple brain, not the large complex brains of octopuses and squid, and had long been assumed to lack intelligence. But the cephalopod nervous system is quite different from that of other animals, and recent experiments have shown not only memory, but a changing response to the same event over time.[14][15][16] In a study in 2008, a group of nautiluses (N. pompilius) were given food as a bright blue light flashed until they began to associate the light with food, extending their tentacles every time the blue light was flashed. The blue light was again flashed without the food 3 minutes, 30 minutes, 1 hour, 6 hours, 12 hours, and 24 hours later. The nautiluses continued to respond to the blue light for up to 30 minutes after the experiment. An hour later they showed no reaction to the blue light. However, between 6 and 12 hours after the training, they again responded to the blue light. The researchers concluded that nautiluses had memory capabilities similar to the "short-term" and "long-term memories" of the more advanced cephalopods, despite having different brain structures.[14][15][16] However the long-term memory capability of nautiluses was much shorter than that of other cephalopods. The nautiluses completely forgot the earlier training 24 hours later, in contrast to octopuses, for example, which can remember conditioning for weeks afterwards. However, this may simply be the result of the conditioning procedure being suboptimal for sustaining long-term memories in nautiluses. Nevertheless the study showed that scientists had previously underestimated the memory capabilities of nautiluses.[16]
Nautilus, of cephalopod family Nautilidae

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): 16326. doi:10.1016/j.cub.2009.07.067. PMID19765993. [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.; Florey, E. (1968). "Ultrastructure of cephalopod chromatophore organs". Z Zellforsch Mikrosk Anat 89 (2): 250280. doi:10.1007/BF00347297. PMID5700268. [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(3.86MB) Berliner Geowissenschaftliche Abhandlungen 3: 29-35. [11] Finn, J. K.; Tregenza, T., Tregenza, N. (2009). "Defensive tool use in a coconut-carrying octopus". Current Biology 19 (23): R1069-R1070. doi:10.1016/j.cub.2009.10.052. PMID20064403. [12] 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. [13] "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. [14] Ewen Callaway (2 June 2008). "Simple-Minded Nautilus Shows Flash of Memory" (http:/ / www. newscientist. com/ article/ dn14033-simpleminded-nautilus-reveals-flash-of-memory. html). New Scientist. . Retrieved 7 March 2012.

Cephalopod intelligence
[15] Kathryn Phillips (15 June 2008). "Living Fossil Memories" (http:/ / jeb. biologists. org/ content/ 211/ 12/ iii. full. pdf#page=1& view=FitH). Inside JEB 211 (12): iii. doi:10.1242/jeb.020370. . [16] Robyn Crook & Jennifer Basil (2008). "A biphasic memory curve in the chambered nautilus, Nautilus pompilius L. (Cephalopoda: Nautiloidea)" (http:/ / jeb. biologists. org/ content/ 211/ 12/ 1992. full. pdf#page=1& view=FitH). The Journal of Experimental Biology 211 (12): 19921998. doi:10.1242/jeb.018531. .

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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 National Zoo in Washington D.C. 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. ISBN0-521-42083-0. 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 (3): 308817. PMID16801504. 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.

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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

Giant Pacific Octopus, Enteroctopus dofleini

Octopoda (octopuses) Species Maximum mantle length 0.69 m 0.6 m References Notes

Haliphron atlanticus Enteroctopus dofleini

O'Shea (2004) Norman (2000:214) Sepiida (cuttlefish)

Species

Maximum mantle length 0.5 m 0.5 m <0.5 m 0.49 m 0.42 m

References

Notes

Sepia apama Sepia latimanus Sepia hierredda Sepia officinalis Sepia pharaonis

Reid et al. (2005:68) Reid et al. (2005:92) Reid et al. (2005:88) Reid et al. (2005:99) Reid et al. (2005:107)

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0.38 m 0.375 m Reid et al. (2005:96) Reid et al. (2005:114) Sepiolida (bobtail squid)

Sepia lycidas Sepia ramani

Species

Maximum mantle length

References

Notes

Austrorossia antillensis 0.09 m Rossia pacifica Rossia macrosoma Neorossia caroli 0.09 m (female) 0.085 m 0.083 m (female)

Reid & Jereb (2005:192) Reid & Jereb (2005:185) Reid & Jereb (2005:184) Reid & Jereb (2005:190) Male grows to 45mm in mantle length (Reid & Jereb, 2005). More typically the mantle length is 2060mm (Reid & Jereb, 2005). Male grows to 51mm in mantle length (Reid & Jereb, 2005).

Spirulida (Ram's Horn Squid) (only one extant species) Species Maximum mantle length References Notes

Spirula spirula

rarely exceeds 0.045 Reid (2005:211) m Teuthida (squid)

Species

Maximum mantle length 4 m (estimate)

References

Notes

Mesonychoteuthis hamiltoni Galiteuthis phyllura

O'Shea (2005a)

Estimate based on largest known beak (LRL: 49mm).

2.652.75 m (estimate) 2.25 m 2m

Nesis (1985)

Estimate based on 0.4 m long arm and 1.15 m tentacle.

Architeuthis sp. Onykia robusta

O'Shea (2005a) Norman (2000:174) Tsuchiya & Okutani (1993) Glaubrecht & Salcedo-Vargas (2004) Nesis (1982) O'Shea (2005b) Longest confirmed specimen measures 0.85 m (O'Shea, 2005b). Total length to at least 2.3 m (Carrington, 2000). Norman (2000:165) gives maximum of 1.5 m. Kubodera et al. (1998) give maximum of at least 1.615 m.

Megalocranchia fisheri 1.8 m Dosidicus gigas 1.75 m

Taningia danae

1.7 m

Kondakovia longimana probably 1.15+ m (estimate) Lepidoteuthis grimaldii 1 m Thysanoteuthis rhombus cf. Magnapinna 1m

Young & Vecchione (2009) Roper et al. (1984) Commonly grows to a mantle length of 0.6 m (Roper et al., 1984).

~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) Species Maximum mantle length 0.13 m References Notes

Vampyroteuthis infernalis

Nesis (1982)

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.

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Total length

A long-arm squid (cf. Magnapinna) filmed in the Gulf of Mexico.

Octopoda (octopuses) Not to be confused with armspan, which is approximately double the total length. Species Maximum total length >6.1 m 4 m (estimate) References Notes

Enteroctopus dofleini Haliphron atlanticus

Cosgrove (1987) O'Shea (2004) Estimate based on incomplete 2.90 m specimen.

Teuthida (squid) Total length including long feeding tentacles. Species Maximum total length 14 m (estimate) References Notes

Mesonychoteuthis hamiltoni Architeuthis sp.

O'Shea (2005a)

Estimate based on largest known beak (LRL: 49mm).

13 m (female)

O'Shea (2005a)

Measured post mortem and relaxed. Older records were exaggerated by stretching of the tentacles (O'Shea, 2005a). Estimate based on video evidence.

cf. Magnapinna

at least 8 m (estimate) 5.5 m

Bolstad (2003)

Asperoteuthis acanthoderma Onykia robusta Galiteuthis phyllura

Tsuchiya & Okutani (1993) Verrill (1876) Nesis (1985)

Length of immature specimen measuring 0.45 m ML. Largest known specimen (0.78 m ML) would presumably be longer if it were complete (Okutani, 1995).

over 4 m over 4 m (estimate)

Estimate based on 0.40 m long arm and 1.15 m tentacle.

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Weight

Sepia apama, the heaviest species of cuttlefish

Octopoda (octopuses) Species Maximum weight References Notes

Haliphron atlanticus Enteroctopus dofleini

75kg (estimate) O'Shea (2004) 71kg Cosgrove (1987)

Estimate based on incomplete 61.0kg specimen. Weight of live specimen. There exists a highly dubious record of a 272kg specimen (High, 1976). Sepiida (cuttlefish)

Species

Maximum weight >10.5kg 10kg >7.5kg 5kg 5kg

References

Notes

Sepia apama Sepia latimanus Sepia hierredda Sepia lycidas Sepia pharaonis

Reid et al. (2005:68) Reid et al. (2005:92) Reid et al. (2005:88) Reid et al. (2005:96) Reid et al. (2005:107) Reid et al. (2005:99) Teuthida (squid)

Sepia officinalis

4kg

Species

Maximum weight 495kg

References

Notes

Mesonychoteuthis hamiltoni Architeuthis sp. Taningia danae

[Anonymous] (2007)

Weight of mature specimen caught in early 2007. Originally estimated to weigh 450kg (Anderton, 2007).

275kg (female) 61.4kg

O'Shea (2005a) Kubodera et al. (2006) Roper et al. (1984) Nigmatullin et al. (2001) Miyahara et al. (2006)

Onykia robusta Dosidicus gigas

50kg 50kg

Thysanoteuthis rhombus

30kg

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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 diameter 268mm References Notes

Nautilus pompilius pompilius

[1]

Pisor (2005:93) lists maximum shell diameter of 254.0mm. Nautilus repertus is treated here in synonymy with N. pompilius pompilius. Pisor (2005:93) lists 230.0mm record for N. repertus.

Nautilus belauensis

226mm

Jereb (2005:54) Pisor (2005:93) Pisor (2005:93) Jereb (2005:55) Pisor (2005:93) [2] Pisor (2005:93) lists maximum shell diameter of 148.0mm.

Allonautilus scrobiculatus Nautilus stenomphalus Allonautilus perforatus Nautilus macromphalus Nautilus pompilius suluensis

215.0mm

201.0mm

around 180mm

180.0mm

160mm

Cephalopod size

99
Octopoda (octopuses) (all extant Argonauta species listed) Females of the genus Argonauta produce a calcareous eggcase in which they reside.

Species

Maximum shell diameter 300.0mm

References

Notes

Argonauta argo

Pisor (2005:12) Pisor (2005:12) Pisor (2005:12) Pisor (2005:12) Pisor (2005:12) Pisor (2005:12) Pisor (2005:12)

Argonauta nodosa

292.0mm

Argonauta pacifica 220.0mm

Argonauta hians

112.6mm

Argonauta cornuta

98.6mm

Argonauta nouryi

95.5mm

Argonauta bottgeri

67.0mm

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. Species Maximum shell diameter 28.8mm References Notes

Spirula spirula

[3]

Pisor (2005:108) lists maximum shell diameter of 26.9mm.

Species status questionable.

Extinct taxa

Cast of Parapuzosia seppenradensis

Fossilised guards of Megateuthis gigantea (top and centre)

Ammonoidea (ammonites) Species Maximum shell diameter 2.55 m (estimate) References Notes

Parapuzosia seppenradensis

Kennedy & Kaplan (1995)

Estimate based on 1.95 m diameter specimen with an incomplete living chamber.

Belemnoidea (belemnites)

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100
Maximum rostrum length 0.46 m References Notes

Species

Megateuthis gigantea

Eyden (2003)

The whole belemnite is estimated to have been 35 m long.

Nautiloidea (nautiloids) Species Cameroceras sp. Maximum shell length 11 m (estimate) References Teichert & Kmmel (1960) Notes Frey (1995) gives a maximum length of up to 6m.

Vampyromorphida (vampire squid) Species Maximum mantle length over 1.8 m (estimate) References Notes

Tusoteuthis longa

Eyden (2004)

References
[Anonymous] 2007. Colossal squid may be headed for oven [4]. Associated Press. Anderton, H.J. 2007. Amazing specimen of world's largest squid in NZ [5]. New Zealand Government website. Bolstad, K. 2003. Deep-Sea Cephalopods: An Introduction and Overview [6]. The Octopus News Magazine Online. Carrington, D. 2000. Big squid breaks record [7]. 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 [8]. The Octopus News Magazine Online. Eyden, P. 2004. Cretaceous Giant Squid [9]. The Octopus News Magazine Online. Frey, R.C. 1995. Middle and Upper Ordovician nautiloid cephalopods of the Cincinnati Arch region of Kentucky, Indiana, and Ohio. [10]PDF U.S. Geological Survey. 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): 5369. 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. FAO, Rome. pp. 5155. (German) Kennedy, W.J. & U. Kaplan 1995. Parapuzosia (Parapuzosia) seppenradensis (LANDOIS) und die Ammoniten fauna der Dlmener Schichten, Westfalen. Geol. Palont. 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. [11]PDF(295KB) Proceedings of the Royal Society B: Biological Sciences 274(1613): 10291034. 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): 199205. 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

Cephalopod size 1987. Cephalopods of the world. T.F.H. Publications, Neptune City, NJ. 351pp.] (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): 919. 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 [12]. The Octopus News Magazine Online. O'Shea, S. 2005b. Kondakovia longimana [13]. 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.

101

Reid, A. 2005. Family Spirulidae. 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. FAO, Rome. pp. 211212. Reid, A. & P. Jereb 2005. Family Sepiolidae. 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. FAO, Rome. pp. 153203. 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. FAO, Rome. pp. 57152. Roper C.F.E., M.J. Sweeney & C.E. Nauen 1984. Cephalopods of the world. Food and Agriculture Organization, Rome, Italy. Teichert, C. & B. Kmmel 1960. Size of Endocerid Cephalopods. Breviora Mus. Comp. Zool. 128: 17. 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 [14]. Cephalopods in Action. Verrill, A.E. 1876. Notes on gigantic cephalopods, a correction. American Journal of Science and Arts 12(3): 236-237. Young, R.E. & M. Vecchione 2009. Lepidoteuthidae Pfeffer 1912 [15]. The Tree of Life Web Project.

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References
[1] http:/ / www. conchology. be/ ?t=68& u=118764& g=0d16facb6ebae4a31321e9f6e6298f46& q=a1d787a5674deb7b193549c6ccdff907 [2] http:/ / www. conchology. be/ ?t=68& u=626793& g=1e1012d7a54ee03e9e40db969f89b3c8& q=a84eff41a380577a0fc6c2eada7d1fd4 [3] http:/ / www. conchology. be/ ?t=68& u=126690& g=e8a21ed471661c24ca2aea9f5ea70400& q=4e4d53268a89f2aef5613d13d96327f0 [4] http:/ / news. yahoo. com/ s/ ap/ 20070322/ ap_on_sc/ colossal_squid;_ylt=ArrD9. iKLzOCbR7FKcFksAnMWM0F [5] http:/ / www. beehive. govt. nz/ ViewDocument. aspx?DocumentID=28451 [6] http:/ / www. tonmo. com/ science/ public/ deepseacephs. php [7] http:/ / news. bbc. co. uk/ 2/ hi/ science/ nature/ 813394. stm [8] http:/ / www. tonmo. com/ science/ public/ belemnites. php [9] http:/ / www. tonmo. com/ science/ fossils/ cretaceousGS. php [10] http:/ / pubs. usgs. gov/ pp/ 1066p/ report. pdf [11] http:/ / www. pubs. royalsoc. ac. uk/ media/ proceedings_b/ papers/ RSPB20060236. pdf [12] http:/ / www. tonmo. com/ science/ public/ giantsquidfacts. php [13] http:/ / web. archive. org/ web/ 20071227144307/ http:/ / www. tonmo. com/ science/ public/ giantsquidfacts2. php [14] http:/ / www. mnh. si. edu/ cephs/ vetal01/ vetal01. html [15] http:/ / tolweb. org/ Lepidoteuthis_grimaldii/ 19833

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 the gastropod clade known as 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").

Cephalopod ink 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]

103

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). Modern use of cephalopod ink is generally limited to cooking, where it is used as a food colouring and flavouring, 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 Arrs negre owes its dark colour to squid ink preparation of the dead 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 0-521-64583-2 [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/)

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104

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. ISBN0-632-06048-4. .

Cephalopod arm

Arm of Illex illecebrosus with two rows of suckers along its length

Tentacle of Illex illecebrosus with a distal tentacular club (right)

Cephalopod arm

105

All cephalopods possess flexible limbs extending from their heads and surrounding their beaks. These appendages, which function as muscular hydrostats, have been variously termed arms or tentacles.

Description
In the scientific literature, a cephalopod arm is often treated as distinct from a tentacle, though the terms are sometimes used interchangeably. Generally, arms have suckers along most of their length, as opposed to tentacles, which have suckers only near their ends.[1] Barring a few exceptions, 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, the hectocotylus. Anatomically, cephalopod limbs function using a crosshatch of helical collagen fibres in opposition to internal hydrostatic pressure.[5]
Arms and buccal mass of the squid Taningia danae. As in other Octopoteuthidae, the tentacles are absent in adults.

Abnormalities
Many octopus arm anomalies have been recorded,[6][7] including a 6-armed octopus (nicknamed Henry the Hexapus), a 7-armed octopus,[8] a 10-armed Octopus briareus,[9] one with a forked arm tip,[10] an octopus with double hectocotylization,[11] bilateral hectocotylization,[12] and specimens with up to 96 arm branches.[13][14][15] Branched arms have also been recorded in cuttlefish.[16]

Oral view of the bobtail squid Semirossia tenera

Variability
Cephalopod limbs and the suckers they bear are shaped in many distinctive ways, and vary considerably between species. Some examples are shown below.

Arms
For hectocotylized arms see hectocotylus variability.

Head and limbs of the bobtail squid Rossia glaucopis

Cephalopod arm

106

Oral view of male Bathypolypus arcticus with hectocotylus on arm III (left)

Octopus arm with two rows of suckers

Cephalopod arm

107

Octopus suckers

A deformed octopus sucker cluster

Cephalopod arm

108

A common octopus with a forked arm that was found in south Crete, Greece

Shape of arm

Species

Family

Todarodes pacificus Ommastrephidae

Tentacular clubs
Shape of tentacular club Species Abraliopsis morisi Family Enoploteuthidae

Ancistroteuthis lichtensteini Onychoteuthidae

Architeuthis sp.

Architeuthidae

Austrorossia mastigophora

Sepiolidae

Berryteuthis magister

Gonatidae

Idioteuthis cordiformis

Mastigoteuthidae

Iridoteuthis iris

Sepiolidae

Mastigoteuthis glaukopis Onykia ingens Semirossia tenera

Mastigoteuthidae Onychoteuthidae Sepiolidae

Cephalopod arm

109
Spirula spirula Spirulidae

Todarodes pacificus

Ommastrephidae

Suckers
Shape of sucker Species Idioteuthis cordiformis Family Mastigoteuthidae

Idioteuthis latipinna

Mastigoteuthidae

Magnapinna talismani

Magnapinnidae

Mastigoteuthis agassizii Mastigoteuthidae

Mastigoteuthis atlantica Mastigoteuthidae

Mastigoteuthis dentata

Mastigoteuthidae

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110
Mastigoteuthis grimaldii Mastigoteuthidae

Mastigoteuthis magna

Mastigoteuthidae

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. 249256. 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. 257269. doi:10.1007/978-90-481-3299-7_18 [5] Inside natures giants, Giant squid episode. [6] Kumph, H.E. 1960. Arm abnormality in octopus. Nature 185(4709): 334-335. doi:10.1038/185334a0 [7] 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): 313316.doi:10.1007/BF01668021 [8] 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: 479487. [9] 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. [10] 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. [11] Palacio, F.J. 1973. On the double hectocotylization of octopods. (http:/ / ia311216. us. archive. org/ 2/ items/ nautilus87amer/ nautilus87amer. pdf)PDF The Nautilus 87: 99102. [12] Robson, G.C. 1929. On a case of bilateral hectocotylization in Octopus rugosus. Journal of Zoology 99(1): 9597. doi:10.1111/j.1469-7998.1929.tb07690.x [13] 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): 618623. [14] 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): 624629. [15] Monster octopi with scores of extra tentacles (http:/ / pinktentacle. com/ 2008/ 07/ monster-octopi-with-scores-of-extra-tentacles/ ). Pink Tentacle, July 18, 2008. [16] Okada, Y.K. 1937. An occurrence of branched arms in the decapod cephalopod, Sepia esculenta Hoyle. Annotated Zoology of Japan 17: 9394.

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111

Hectocotylus
A hectocotylus (plural: hectocotyli) is basically a cephalopod phallus. One of the arms of the male of most kinds of cephalopods is modified and specialized to transfer sperm to the female and fertilize her eggs. It is a muscular hydrostat that serves as a spermatophore. Depending on the species, the male may use it merely to store and transfer sperm to the female, or he may wrench it off and present it to the female. Males generally form a new hectocotylus in each new season. Among Decapodiformes (ten-limbed cephalopods), generally either one or both of arms IV are hectocotylized; in incirrate octopuses it is one of arm pair III.[1] Many coleoids lack hectocotyli altogether.[1] 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 funnelmantle locking apparatus on the hectocotylus keeps it lodged in the pallial cavity of the female.

Georges Cuvier's original illustration of an octopus hectocotylus, which he named "Hectocotyle octopodis"

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.[2][3] Rare examples of double and bilateral hectocotylization have been recorded in incirrate octopuses.[4][5]

Anatomy
Generalized anatomy of squid and octopod hectoctyli:

Hectocotylus

112

Squid

Octopod

Variability
Hectocotyli are shaped in many distinctive ways, and vary considerably between species. Some examples are shown below.
Shape of hectocotylus Species Abraliopsis morisi Family Enoploteuthidae

Argonauta bottgeri

Argonautidae

Bathypolypus arcticus

Octopodidae

Graneledone verrucosa Octopodidae

Haliphron atlanticus

Alloposidae

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113
Ocythoe tuberculata Ocythoidae

Scaeurgus patagiatus

Octopodidae

Tremoctopus violaceus Tremoctopodidae

Uroteuthis duvauceli

Loliginidae

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] 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. [3] Nixon M. & J.Z. Young J.Z. 2003. The brains and lives of Cephalopods. Oxford University Press. [4] Robson, G.C. 1929. On a case of bilateral hectocotylization in Octopus rugosus. Journal of Zoology 99(1): 9597. doi:10.1111/j.1469-7998.1929.tb07690.x [5] Palacio, F.J. 1973. On the double hectocotylization of octopods. (http:/ / ia311216. us. archive. org/ 2/ items/ nautilus87amer/ nautilus87amer. pdf)PDF The Nautilus 87: 99102.

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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. The word tentillum is a diminutive, but although it literally means "a little tentacle", it usually refers, irrespective of size, to a side branch of a larger tentacle. In some cases such tentilla are specialised for particular functions; for example, in the Cnidaria tentilla usually bear cnidocytes,[1] whereas in the Ctenophora they usually bear collocytes.[2][3]

Tentacles in animals
Invertebrates
Cuttlefish with 2 tentacles and 8 The phylum Mollusca includes many species with muscular hydrostats in the arms form of tentacles and arms. Octopuses do have tentacles but are rather known as 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.

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. 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 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 containing a coiled thread-like structure called a nematocyst, fired at

Front view of land snail showing upper and lower sets of tentacles

Abalone showing pallial tentacles

potential prey.

Tentacle Many species of the jellyfish-like 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.

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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.

References
[1] http:/ / species-identification. org/ species. php?species_group=zsao& selected=definitie& menuentry=woordenlijst& record=tentilla [2] Harmer, Sir Sidney Frederic; Shipley, Arthur Everett et alia: The Cambridge natural history Volume 1, Protozoa, Porifera, Coelenterata, Ctenophora, Echinodermata. Macmillan Company 1906 [3] Mackie G.O., Mills C.E., Singla C.L.: Structure and function of the prehensile tentilla of Euplokamis (Ctenophora, Cydippida). Zoomorphology (1988) 107:319-337 Leaf and tentacle movement on Drosera capensis

Dactylus

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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 cephalopods developed camera eyes independently. In vertebrate eyes, the nerve fibers route before the retina, blocking some light and creating a blind spot where the fibers pass through the retina and out of the eye. In cephalopod eyes, the nerve fibers route behind the retina, and do not block light or disrupt the retina. In the example, 4 denotes the vertebrate blind spot, which is notably absent in the cephalopod eye. In vertebrates, 1 denotes the retina and 2 the nerve fibers, including the optic nerve (3), whereas in cephalopods, such as octopuses, 1 and 2 denote 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

117 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. To illustrate, the vertebrate eye is normally insensitive to polarization differences because visual pigment in rods and cones is arrayed semi-randomly, and is thereby equally sensitive to any orientation of the e-vector axis of the light. Because of their orthogonal organization, the visual pigment molecules in cephalopod eyes have the highest light absorption when aligned properly with the light e-vector axis, allowing sensitivity to differences in polarization.[4] The precise function of this ability has not been proven, but is hypothesized to be for prey detection, navigation, and possibly communication among the color-changing cephalopods.[4][5]

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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): 347361. doi:10.1002/cne.902320307. ISSN0021-9967. PMID2857734. [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. 46 (1): 1118. PMID359745. . [4] Mathger, L.M.; Shashar, N.; Hanlon, R.T. (2009). "Do cephalopods communicate using polarized light reflections from their skin?". Journal of Experimental Biology 212 (Pt 14): 21332140. doi:10.1242/jeb.020800. PMID19561202. [5] 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 organelles in cells found in amphibians, fish, reptiles, crustaceans, cephalopods and bacteria. 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 or metachrosis.[1] 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 be initiated by changes in mood, temperature, stress or visible changes in local environment.

Zebrafish chromatophores mediate background adaptation on exposure to dark (top) and light 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.

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119

History
Aristotle mentioned the ability of the octopus to change colour for both camouflage and signalling in his Historia animalium (ca 400 BC):[2] The octopus ... seeks its prey by so changing its colour as to render it like the colour of the stones adjacent to it; it does so also when alarmed. G. Sangiovanni was the first to describe invertebrate pigment-bearing cells as chromoforo in an Italian science journal in 1819.[3] Charles Darwin described the colour-changing abilities of the cuttlefish in The Voyage of the Beagle (1860):[4] These animals also escape detection by a very extraordinary, chameleon-like power of changing their colour. They appear to vary their tints according to the nature of the ground over which they pass: when in deep water, their general shade was brownish purple, but when placed on the land, or in shallow water, this dark tint changed into one of a yellowish green. The colour, examined more carefully, was a French grey, with numerous minute spots of bright yellow: the former of these varied in intensity; the latter entirely disappeared and appeared again by turns. These changes were effected in such a manner that clouds, varying in tint between a hyacinth red and a chestnut-brown, were continually passing over the body. Any part, being subjected to a slight shock of galvanism, became almost black: a similar effect, but in a less degree, was produced by scratching the skin with a needle. These clouds, or blushes as they may be called, are said to be produced by the alternate expansion and contraction of minute vesicles containing variously coloured fluids.

Classification
The term chromatophore was adopted (following Sangiovanni's chromoforo) 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 khrma () meaning "colour," and phoros () meaning "bearing". In contrast, the word chromatocyte (cyte or 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 more useful to a scientific understanding of how the cells function.[5] Colour-producing molecules fall into two distinct classes: biochromes and schemochromes.[6] 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 colours", produce colouration by reflecting some wavelengths (colours) 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.

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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.[5] 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.[7] 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 A veiled chameleon, Chamaeleo calyptratus. have supplemental biochemical pathways that result in an excess Structural green and blue colours are generated accumulation of yellow pigment. In contrast, carotenoids are by overlaying chromatophore types to reflect metabolised from the diet and transported to erythrophores. This was filtered 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.[8]

Iridophores and leucophores

Iridophores, sometimes also called guanophores, are pigment cells that reflect light using plates of crystalline chemochromes made from guanine.[9] When illuminated they generate iridescent colours because of the diffraction of light within the stacked plates. Orientation of the schemochrome determines the nature of the colour observed.[10] By using biochromes as coloured filters, iridophores create an optical effect known as Tyndall or Rayleigh scattering, producing bright blue or green colours.[11] 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 colours while leucophores produce reflective white hues.[11]

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.[12] 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 packaged alongside eumelanin. For example, a novel deep (wine) red coloured pigment was identified in the melanophores of

At the bottom a mutant zebrafish larva that fails to synthesise melanin in its melanophores, at the top a non-mutant, wildtype larva.

Chromatophore phyllomedusine frogs.[13] 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.[14] 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.

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Cyanophores
In 1995 it was demonstrated that the vibrant blue colours in some types of mandarinfish are not generated by schemochromes. Instead, a cyan biochrome of unknown chemical nature is responsible.[11] 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 appear unusual in their taxonomic restriction, there may be The purple-striped dottyback, Pseudochromis diadema, generates its violet stripe with an cyanophores (as well as further unusual chromatophore types) in other unusual type of chromatophore fish and amphibians. For example, bright coloured chromatophores with undefined pigments have been observed in both poison dart frogs and glass frogs,[15] and atypical dichromatic chromatophores, named erythro-iridophores have been described in Pseudochromis diadema.[16]

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.[17] 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 colours by making good use of the divisional effect.[18],[19]

Chromatophore

122 The control and mechanics of rapid pigment translocation has been well studied in a number of different species, particularly amphibians and teleost fish.[20],[11] 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.[21] 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 fashion by cells in the skin. At the surface of the melanophore the hormones have been shown to activate specific G-protein coupled receptors that, in turn, transduce the signal into the cell. Melanocortins result in the dispersion of pigment, while melatonin and MCH results

A single zebrafish melanophore imaged by time-lapse photography during pigment aggregation

in aggregation.[22] Numerous melanocortin, MCH and melatonin receptors have been identified in fish[23] and frogs,[24] including a homologue of MC1R,[25] a melanocortin receptor known to regulate skin and hair colour in humans.[26] It has been demonstrated that MC1R is required in zebrafish for dispersion of melanin.[27] 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.[28],[29],[30]

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),[31] and that melanin translocation in melanophores is the major factor in colour change.[22] Some animals, such as chameleons and anoles, have a highly developed background adaptation response capable of generating a number of different colours 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.

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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.[32] 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.[33] 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.[34] 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.[31] 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.[25] Therefore pharmaceutical companies have developed a biological assay for rapidly identifying potential bioactive compounds using melanophores from the African clawed frog.[35] Other scientists have developed techniques for using melanophores as biosensors,[36] and for rapid disease detection (based on the discovery that pertussis toxin blocks pigment aggregation in fish melanophores).[37] Potential military applications of chromatophore mediated colour changes have been proposed, mainly as a type of active camouflage, along with invisibility.[38][39]

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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.[40] 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 An infant cuttlefish, using background adaptation to mimic the local environment mechanism used in fish, amphibians and 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.[41] 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.[42] 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.[43]

Notes
[1] Scott M. Boback & Lynn M. Siefferman (2010). "Variation in Color and Color Change in Island and Mainland Boas (Boa constrictor)". Journal of Herpetology 44 (4): 506515. doi:10.1670/09-026.1. [2] Aristotle. Historia Animalium. IX, 622a: 2-10. About 400 BC. Cited in Luciana Borrelli, Francesca Gherardi, Graziano Fiorito. A catalogue of body patterning in Cephalopoda. Firenze University Press, 2006. Abstract (http:/ / www. fupress. com/ scheda. asp?IDV=487) Google books (http:/ / books. google. co. uk/ books?id=CW5RKQEL7-EC& pg=PA3& lpg=PA3) [3] 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; 9:113. [4] Darwin, Charles (1860). Journal Of Researches Into The Natural History And Geology Of The Countries Visited During The Voyage Round The World Of H.M.S. 'Beagle' Under The Command Of Captain Fitz Roy, R.N.. John Murray, London. p.7. [5] Bagnara JT. (1966). "Cytology and cytophysiology of non-melanophore pigment cells". Int Rev Cytol. International Review of Cytology 20: 173205. doi:10.1016/S0074-7696(08)60801-3. ISBN978-0-12-364320-9. PMID5337298. [6] 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 0-520-02347-1 [7] Matsumoto J (1965). "STUDIES ON FINE STRUCTURE AND CYTOCHEMICAL PROPERTIES OF ERYTHROPHORES IN SWORDTAIL, XIPHOPHORUS HELLERI, WITH SPECIAL REFERENCE TO THEIR PIGMENT GRANULES (PTERINOSOMES)". J

Chromatophore
Cell Biol 27 (3): 493504. doi:10.1083/jcb.27.3.493. PMC2106771. PMID5885426. [8] 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 [9] Taylor JD. (1969). "The effects of intermedin on the ultrastructure of amphibian iridophores". Gen Comp Endocrinol 12 (3): 40516. doi:10.1016/0016-6480(69)90157-9. PMID5769930. [10] Morrison RL. (1995). "A transmission electron microscopic (TEM) method for determining structural colors reflected by lizard iridophores". Pigment Cell Res 8 (1): 2836. doi:10.1111/j.1600-0749.1995.tb00771.x. PMID7792252. [11] Fujii R. (2000). "The regulation of motile activity in fish chromatophores". Pigment Cell Res 13 (5): 30019. doi:10.1034/j.1600-0749.2000.130502.x. PMID11041206. [12] Ito, S; Wakamatsu, K. (2003). "Quantitative analysis of eumelanin and pheomelanin in humans, mice, and other animals: a comparative review". Pigment Cell Res 16 (5): 52331. doi:10.1034/j.1600-0749.2003.00072.x. PMID12950732. [13] 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 (4116): 10345. doi:10.1126/science.182.4116.1034. PMID4748673. . [14] 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): 510516. doi:10.1034/j.1600-0749.2003.00075.x. PMID12950730. . [15] Schwalm PA, Starrett PH, McDiarmid RW (1977). "Infrared reflectance in leaf-sitting neotropical frogs". Science 196 (4295): 12257. doi:10.1126/science.860137. PMID860137. [16] Goda M, Ohata M, Ikoma H, Fujiyoshi Y, Sugimoto M, Fujii R (2011). "Integumental reddish-violet coloration owing to novel dichromatic chromatophores in the teleost fish, Pseudochromis diadema". Pigment Cell Melanoma Res 24 (4): 6147. doi:10.1111/j.1755-148X.2011.00861.x. PMID21501419. [17] Bagnara JT, Taylor JD, Hadley ME (1968). "The dermal chromatophore unit" (http:/ / www. jcb. org/ cgi/ reprint/ 38/ 1/ 67). J Cell Biol 38 (1): 6779. doi:10.1083/jcb.38.1.67. PMC2107474. PMID5691979. . [18] Palazzo RE, Lynch TJ, Lo SJ, Taylor JD, Tchen TT (1989). "Rearrangements of pterinosomes and cytoskeleton accompanying pigment dispersion in goldfish xanthophores". Cell Motil Cytoskeleton 13 (1): 920. doi:10.1002/cm.970130103. PMID2543509. [19] Porras MG, De Loof A, Breuer M, Archiga H (2003). "Procambarus clarkii". Peptides 24 (10): 15819. doi:10.1016/j.peptides.2003.08.016. PMID14706537. [20] 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): 297301. doi:10.1083/jcb.200210066. PMC2172679. PMID12551954. . [21] Aspengren S, Skld HN, Quiroga G, Mrtensson L, Wallin M (2003). "Noradrenaline- and melatonin-mediated regulation of pigment aggregation in fish melanophores". Pigment Cell Res 16 (1): 5964. doi:10.1034/j.1600-0749.2003.00003.x. PMID12519126. [22] Logan DW, Burn SF, Jackson IJ (2006). "Regulation of pigmentation in zebrafish melanophores". Pigment Cell Res 19 (3): 20613. doi:10.1111/j.1600-0749.2006.00307.x. PMID16704454. [23] 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: 31930. doi:10.1111/j.1749-6632.2003.tb03196.x. PMID12851332. [24] Sugden D, Davidson K, Hough KA, Teh MT (2004). "Melatonin, melatonin receptors and melanophores: a moving story". Pigment Cell Res 17 (5): 45460. doi:10.1111/j.1600-0749.2004.00185.x. PMID15357831. [25] Logan DW, Bryson-Richardson RJ, Pagn 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): 18491. doi:10.1016/S0888-7543(02)00037-X. PMID12620396. [26] 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): 32830. doi:10.1038/ng1195-328. PMID7581459. [27] 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): 28995. doi:10.1089/zeb.2008.0541. PMID19133827. [28] 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): 132049. doi:10.1073/pnas.0403092101. PMC516548. PMID15331778. . [29] 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): 1658. doi:10.1016/S0960-9822(98)70064-8. PMID9443917. . [30] 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& _udi=B6VRT-4DMW0C4-11& _user=10& _coverDate=10/ 26/ 2004& _alid=412398862& _rdoc=1& _fmt=full& _orig=search& _qd=1& _cdi=6243& _sort=d& _docanchor=& view=c& _acct=C000050221& _version=1& _urlVersion=0& _userid=10& md5=96f1a14140773f1e898c1ef410db1cb9& artImgPref=F). Current biology : CB 14 (20): 187781. doi:10.1016/j.cub.2004.10.003. PMID15498498. .

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[31] 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): 14860. doi:10.1002/neu.10165. PMID12486702. . [32] Kelsh RN, Schmid B, Eisen JS (2000). "Genetic analysis of melanophore development in zebrafish embryos". Dev Biol 225 (2): 27793. doi:10.1006/dbio.2000.9840. PMID10985850. [33] 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): 32636. doi:10.1111/j.1600-0749.2004.00174.x. PMID15250934. [34] Lamason RL, Mohideen MA, Mest JR, Wong AC, Norton HL, Aros MC, Jurynec MJ, Mao X et al. (2005). "SLC24A5, a putative cation exchanger, affects pigmentation in zebrafish and humans". Science 310 (5755): 17826. doi:10.1126/science.1116238. PMID16357253. [35] 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 assay to rapidly screen a 442,368 bead-based peptide library". J Pharmacol Toxicol Methods 42 (4): 18997. doi:10.1016/S1056-8719(00)00083-6. PMID11033434. [36] Andersson TP, Filippini D, Suska A, Johansson TL, Svensson SP, Lundstrm I (2005). "Frog melanophores cultured on fluorescent microbeads: biomimic-based biosensing". Biosens Bioelectron 21 (1): 11120. doi:10.1016/j.bios.2004.08.043. PMID15967358. [37] Karlsson JO, Andersson RG, Askelf P, Elwing H, Granstrm M, Grundstrm N, Lundstrm 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): 16975. PMID1936946. [38] http:/ / news. nationalgeographic. com/ news/ 2008/ 08/ 080608-cuttlefish-camouflage-missions_2. html [39] Lee I. Nanotubes for noisy signal processing: Adaptive Camouflage PhD Thesis. 2005; University of Southern California. [40] Cloney RA, Florey E (1968). "Ultrastructure of cephalopod chromatophore organs". Zeitschrift fur Zellforschung und mikroskopische Anatomie (Vienna, Austria : 1948) 89 (2): 25080. doi:10.1007/BF00347297. PMID5700268. [41] Demski LS (1992). "Chromatophore systems in teleosts and cephalopods: a levels oriented analysis of convergent systems". Brain, behavior and evolution 40 (23): 14156. doi:10.1159/000113909. PMID1422807. [42] Salton MR (1987). "Bacterial membrane proteins". Microbiological sciences 4 (4): 1005. PMID3153178. [43] Frigaard NU, Bryant DA (2004). "Seeing green bacteria in a new light: genomics-enabled studies of the photosynthetic apparatus in green sulfur bacteria and filamentous anoxygenic phototrophic bacteria". Archives of Microbiology 182 (4): 26576. doi:10.1007/s00203-004-0718-9. PMID15340781.

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External links
Nature's Palette - how animals produce colour (http://www.bioscience-explained.org/ENvol1_2/pdf/ paletteEN.pdf)PDF(1.20MB) 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)

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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 a cloak because in many groups the edges of the mantle extend far 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.

European Squid (Loligo vulgaris). The mantle is all that is visible behind the head: the outer body wall and the fins are all part of the mantle.

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 most molluscs. In bivalves it is usually part of the feeding structure. In some molluscs the mantle cavity is a brood chamber, and in cephalopods and some bivalves such as scallops, it is a locomotory organ.
The brightly coloured mantle of a Giant clam protects it from bright sunlight.

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 gastropods it is used as a kind of "foot" for locomotion over the surface. In Patella the foot includes the entire ventral surface of the animal. The foot of the Bivalvia is a fleshy process adapted by its form to digging rather than to locomotion.

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Formation of mollusc shell

Further information: mollusc shell In shelled molluscs, the mantle is the organ that forms the shell, and 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 Chinese cowry, showing partially extended mantle.

Cypraea chinensis with its mantle fully extended.

The mantle of the land snail Indrella ampulla is off-white in color and it is partly 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 coerulans is smooth, while the rest of the body is tubercled.

Megapallifera mutabilis from Philomycidae shows enormously developed mantle

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Photo of Haliotis asinina with the shell removed.

This drawing shows that the mantle (in gray) covers the majority of the dorsal surface of the [3] animal.

References
[1] "integument (molluscs)." Encyclopdia Britannica. 2009. Encyclopdia 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 Wrheide & 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. 0 doi:10.1186/1741-7007-4-40.

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 secrete egg cases or the gelatinous substance comprising egg masses.[1]

Ventral view of the viscera of Chtenopteryx sicula, showing the presence of the nidamental gland and accessory nidamental gland.

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". Copeia 1948 (1): 547. doi:10.2307/1438791. JSTOR1438791. [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): 197208. doi:10.1016/0040-8166(77)90016-7. PMID906013. .

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Siphon
A siphon is an anatomical structure which is part of the body 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 in molluscs are tube-like structures in which water flows (or more rarely in which air flows). 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 single siphon occurs in some gastropods. In those bivalves which have siphons, the siphons are paired. In cephalopods, there is a single siphon or funnel which is known as a hyponome.

A specimen of a venerid bivalve. The adductor muscles have been cut, the valves are gaping. The internal anatomy is visible, including the paired siphons to the right

The siphon of a large carnivorous marine volute, Cymbiola magnifica

The sea snail Nassarius tiarula is a scavenger. Siphon on the left

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In gastropods
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 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]
Melo amphora moving across coral at low tide

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. The Aplysia gill and siphon withdrawal reflex is a defensive reflex which is found in sea hares of the genus Aplysia; this reflex has been much studied in neuroscience.

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 human 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] The shells of these freshwater snails have simple round apertures; there is no special notch for the siphon.
Pomacea canaliculata, seen through glass, has reached its siphon up to the water surface to breathe air

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Engraving of Florida freshwater applesnail Pomacea paludosa; siphon on lower right

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 [9] when clamming.

Four specimens of Panopea generosa in a seafood tank; the paired siphons (or "necks") of this species can be one meter long

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Veneridae with siphons out

Drawing of the venerid Venus verrucosa showing paired siphons (upper inhalant and lower exhalant siphon), shell and foot.

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.

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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.

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. Whether ammonites possessed a hyponome and if so what form it may have taken, is as yet not known.[15]

Nautilus belauensis seen from the front, showing the opening of the hyponome.

References
[1] rstan 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 978-1-58238-125-1 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 978-1-57233-561-5. 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 978-1-4200-6574-9. 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.

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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

The squid giant axon is the very large (up to 1mm in diameter; typically around 0.5mm) 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, =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/) 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 substantially. The conduction velocity of a typical 0.5mm squid axon is about 25m/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. ISBN978-0-387-48864-6.

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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 numerous upright pillars. Depending on the species, cuttlebones implode at a depth of 200 to 600 metres (660 to 2,000 ft). Because of this limitation, most species of cuttlefish live on the seafloor in shallow water, usually on the continental shelf.[2]

Cuttlebone from Sepia sp.

Cuttlebone of Sepia officinalis (left to right: ventral, dorsal, and lateral views)

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 supplements for caged birds, chinchillas, hermit crabs, and reptiles.[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.
Tortoise with cuttlebone

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]

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3D visualisation of a Sepia cuttlebone by industrial micro computed tomography

3D view of part of a cuttlebone at low resolution.

Overview of a part at high resolution, about 5m/voxel.

Higher magnification.

Detailed view at very high magnification. Wall thickness of the vertical structures is about 10m.

Flight through the corresponding tomographic image stacks

Flight through the corresonding CT image stack, section direction about 30, top view.

Flight through the aligned image stack, top view.

References
[1] Rexfort, A.; Mutterlose, J. (2006). "Stable isotope records from Sepia officinalisa key to understanding the ecology of belemnites?". Earth and Planetary Science Letters 247 (34): 212212. Bibcode2006E&PSL.247..212R. 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 Palobiologische Abhandlungen 3: 189197.

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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.

Where the septum meets the shell a suture line forms; in some 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).

Cutaway of a nautilus shell showing the chambers

Aptychus
An aptychus is a type of marine fossil. It is a hard anatomical structure, a sort of curved shelly plate, now understood to be part of the body of an ammonite. Paired aptychi have, on rare occasions, been found at or within the aperture of ammonite shells. The aptychus was usually composed of calcite, whereas the ammonite shell was aragonite. Aptychi can be found well-preserved as fossils, but usually quite separate from ammonite shells. This circumstance led to them being initially classified as valves of bivalves (clams), which they do somewhat resemble. Aptychi are found in rocks from the Devonian period through to the those of the Cretaceous period.

Some examples of aptychi (top right: Oppelia from Late Jurassic of Solnhofen, Germany; bottom left: aptychi (recto and versus) from Late Jurassic of Lombardy, Italy), and conceptual scheme of their function if indeed they were used to close the shell aperture, as opposed to being jaws.

Aptychus

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There are many forms of aptychus, varying in shape and in the sculpture of the inner and outer surfaces. However, because they are so rarely found in position within the ammonite shell, it is often unclear which kind of aptychus belonged to which species of ammonite. When only a single plate is present, as is sometimes the case, the term "anaptychus" is used.

Function
One of what would have been a pair of aptychi (at Aptychi seem to have most often existed as bilaterally-symmetrical first given the name Trigonellites latus and pairs, and were first described (incorrectly) as being the valves of described as a bivalve) from the Kimmeridge bivalve mollusks. Aptychi are now considered to be either: (1) a Clay Formation in England two-valved closing hatch on the shells of extinct ammonites; or (2) a double-plate jaw-piece similar to that of some modern cephalopods.[1][2][3][4]

Set near 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.

Pair of fossil aptychi (image is 1cm across)

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Fossil of Laevaptychus

Pair of Lingulaticeras solenoides with aptychi in place

Perisphictes with aptychi from the Solnhofen limestone

References
[1] Morton, N. 1981. Aptychi: the myth of the ammonite operculum. Lethaia 14(1): 5761. 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): 231238. 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: 325331. 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: 2032. 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.

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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 Fossilised Orthoceras orthocones. the earliest nautiloids had shells that were slightly 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 chambers They thus became the early Paleozoic equivalent of coleoids. The early 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.

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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 (4): 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 (1): 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]

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References
[1] Mutvei, Harry; Zhang, Yun-bai; Dunca, Elena (2007). "Late Cambrian Plectronocerid Nautiloids and Their Role in Cephalopod Evolution". Palaeontology 50 (6): 13271333. 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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Anderson, Paul H., Pedoleon, Piano non troppo, Postdlf, Psuanthguy, Ragesoss, Rbarreira, Rjwilmsi, Robert K S, Ronz, SWAdair, Santa Sangre, Saperaud, SarahStierch, Shanes, Shrewpelt, Silly rabbit, Sjc, Sk741, Skyskraper, Slakr, Sligocki, Smith609, Snowolf, Spotty11222, Steinsky, Sterwick, Syrthiss, Tarquin, Tbc2, Template namespace initialisation script, The Thing That Should Not Be, TheRingess, Theinsomniac4life, Thingg, Thryduulf, Tjunier, Toadfosky, TomGreen, Toyokuni3, UninvitedCompany, UtherSRG, Vassto, Vegaswikian, Velella, Vrenator, Vsmith, Whatnwas, Whoelius, Wikipelli, Wilson2, Wilson44691, WolfmanSF, Xana500, Zzzzz, 502 , anonymous edits Belemnoidea Source: http://en.wikipedia.org/w/index.php?oldid=510363068 Contributors: Abyssal, Adrian.benko, Aiko, Andy4789, Anthony Appleyard, Arpingstone, Dan Harkless, DanielCD, Dlloyd, Dysmorodrepanis, Felagund, FunkMonk, Headbomb, Huttarl, Invertzoo, J. Spencer, J.H.McDonnell, Jashiin, Katharineamy, Kencasey98, Kevmin, Marcelo-Silva, Mgiganteus1, Moon&Nature, Morning277, NatureA16, Neale Monks, Obsidian Soul, Polarpanda, Rich Farmbrough, Richardleaky, Saintrain, SchreiberBike, Slawojarek, Smith609, UtherSRG, WOSlinker, Wetman, Wilson44691, WolfmanSF, ZeldaGamer31337, Zvar, 53 , anonymous edits Argonaut Source: http://en.wikipedia.org/w/index.php?oldid=520035515 Contributors: Abigail-II, Abyssal, Altenmann, Anthony Appleyard, BD2412, Cakehelmit, CatherineMunro, Cewvero, Citron, D6, DanielCD, Dannajoy, Dysmorodrepanis, GrahamBould, Gribeco, Guidod, Hans Frrum, Headbomb, Iciac, InfernoXV, Invertzoo, Ka Faraq Gatri, Koavf, Koven.rm, Kwamikagami, LilHelpa, Limonadis, M-le-mot-dit, Mgiganteus, Mgiganteus1, Montyy0, Neale Monks, Orangemarlin, Palpalpalpal, Paul H., Peter Horn, Phphello, Pinethicket, PurpleHz, Qwertzy2, Rajah, Rama, Roberto Mura, Rolf Schmidt, SteveSims, Template namespace initialisation script, TheAlphaWolf, Tonyrex, UtherSRG, Wetman, XQ fan, 27 anonymous edits Cephalopod intelligence Source: http://en.wikipedia.org/w/index.php?oldid=518028412 Contributors: 2001:18E8:2:12BC:9C:D575:2894:303F, Acaeton, Alan Liefting, Andy Dingley, Animalplanetemployee, Ashesofoak, Atarr, BatteryIncluded, Ceph, ChildofMidnight, Coelacan, Coemgenus, Delicious carbuncle, Dentren, Donlammers, Eaefremov, Emijrp, Epastore, Epipelagic, Falcon982, Headbomb, HisSpaceResearch, Iam3dhomer, Invertzoo, Jasonz2z, Jefffire, Jonkerz, Karlchwe, Kayau, LittleHow, Luigifan, LukeSurl, Lusanaherandraton, Maproom, Marskell, Martarius, MatthiasKabel, Maunus, 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, 88 anonymous edits Cephalopod size Source: http://en.wikipedia.org/w/index.php?oldid=507472043 Contributors: Archelon, Cadwaladr, Dennis Bratland, Dysmorodrepanis, Headbomb, Hybridbus, J Milburn, Jarble, Jeancey, Lambyte, Materialscientist, Mattg82, Mgiganteus1, PenguinJockey, Rjwilmsi, Shaundakulbara, Taollan82, Twas Now, Uppland, Zuzster, 24 anonymous edits Cephalopod ink Source: http://en.wikipedia.org/w/index.php?oldid=513403979 Contributors: Angr, Anxietycello, Carnby, Cedders, CommonsDelinker, Erianna, Headbomb, Ignacio Bibcraft, Igodard, Intelligentsium, Invertzoo, K-22-22, Mgiganteus1, Nyar94, Panek, Scotsman240363, The Thing That Should Not Be, , 11 anonymous edits Ink sac Source: http://en.wikipedia.org/w/index.php?oldid=492029247 Contributors: Brz7, Cdcdoc, CommonsDelinker, 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=470161360 Contributors: Berek, Cephalopodstudies, Cesiumfrog, Combatking0, Dkontogiannnis, Headbomb, Henryhartley, Jesse V., Materialscientist, Mgiganteus1, Montecarlocars, Nono64, Plastikspork, Stemonitis, Sturm br, Tideflat, Yobmod, ZayZayEM, 19 anonymous edits Hectocotylus Source: http://en.wikipedia.org/w/index.php?oldid=521086060 Contributors: Altenmann, Bansp, Citron, CommonsDelinker, DanielCD, DragonflySixtyseven, Frietjes, Ginkgo100, Icarus source, Icarus source2, Ickle Ronnie, Imnowei, Jpgordon, Kasyapa, MacRusgail, Macdonald-ross, Mgiganteus1, Obsidian Soul, Ospalh, Robofish, UtherSRG, ZayZayEM, Zuzster, , 7 anonymous edits Tentacle Source: http://en.wikipedia.org/w/index.php?oldid=516003482 Contributors: Androstachys, Andycjp, Arrowned, Bioform 1234, Bobo192, Bogey97, Bongwarrior, Burmiester, Cephalopodstudies, Cflm001, Chris Roy, Chrislk02, Clicketyclack, Danger, Delldot, Doulos Christos, Dragon's Light, DrainBead, E. Fokker, Ekko, Epbr123, Excirial, Ezzex, Facts707, Finn-Zoltan, Gadfium, Geary, Gene Nygaard, Goodnightmush, GrahamBould, Gurko, Invertzoo, Isnow, Jarble, JenniferGan, Joaoantonio, Jobba, John, JonRichfield, Kinaro, KnightRider, Kodemage, Lcarscad, Legoo15, Liu Bei, Lokicarbis, LorenzoB, Lugnuts, Lycaon, MOGoss, Materialscientist, Maxima m, Mgiganteus1, Mmxx, Mnj21, Mohammedh15, Narvalo, NawlinWiki, Neutrality, No more bongos, Noloop, Okc, Omegatron, Owen, Oxycut, PhilHibbs, Piano non troppo, Pichpich, Pinkadelica, Proxima Centauri, PurpleHz, Quistnix, Reywas92, Rkitko, Rlandmann, Rodhullandemu, Rogutaan, Rossj81, ScottishGuy, Shoeofdeath, Silly rabbit, Skittle, Smalljim, Tavilis, The High Fin Sperm Whale, TheKMan, TheMadBaron, TheRanger, Thebestofall007, Ugncreative Usergname, UnDeRsCoRe, Username381, Uyanga, Verbosemjp, Wombatcat, Woohookitty, Wyndclaw, Yair rand, Yobmod, ZayZayEM, 140 anonymous edits Dactylus Source: http://en.wikipedia.org/w/index.php?oldid=473539618 Contributors: Agamemnon2, Athurber, Kaarel, Mgiganteus1, Nono64, Pgan002, 3 anonymous edits Cephalopod eye Source: http://en.wikipedia.org/w/index.php?oldid=505557376 Contributors: BD2412, Ccevo2011, Epipelagic, Headbomb, Iciac, Jerryobject, Khazar, Komodo, Mgiganteus1, Richard001, Smeagol 17, Smith609, Woohookitty, 4 anonymous edits Chromatophore Source: http://en.wikipedia.org/w/index.php?oldid=514369615 Contributors: 2607:F470:12:C:BCBA:6019:90CB:1D9C, Adamharvey182, AdultSwim, Alphachimp, Anville, Axeman89, BD2412, BhaiSaab, Bobo192, Boghog, BorgQueen, Brad101, Brighterorange, Caesar Rodney, Ceyockey, Chiswick Chap, Cinchjt, Dark Shikari, DeansFA, Deviator13, Dicklyon, Dispenser, Dj Capricorn, Djanvk, Dominus, DragonflySixtyseven, Drummie06, EagleFan, Edgar181, Ejdzej, Element16, Emvee, Epipelagic, Floyd Landis, Gadfium, Gamesmasterg9, GregorB, Gudeldar, Harmil, Hegar, ISpamThisSite, Icey, InvictaHOG, Jaibobs108, Joyous!, Jpatokal, Kapuchinski, Kilom691, KimvdLinde, Kukini, LANA2007, Mav, 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Sidhekin, Snek01, Stfg, Sun Creator, TheLimbicOne, TimBentley, UtherSRG, Versus22, Wickey-nl, 36 anonymous edits Nidamental gland Source: http://en.wikipedia.org/w/index.php?oldid=516722549 Contributors: CommonsDelinker, Invertzoo, Jarble, Magioladitis, Mgiganteus1, RDBrown, Rjwilmsi, Snek01, , 2 anonymous edits Siphon Source: http://en.wikipedia.org/w/index.php?oldid=451649701 Contributors: Amikake3, Andrewa, Anthony Appleyard, Arcadian, Arct, ChrisGualtieri, DanielCD, Epipelagic, Excirial, Invertzoo, Isthmus, J04n, JamesAM, Jeff G., Kaarel, Kesal, Lymantria, Magioladitis, Mgiganteus1, Mukadderat, Nono64, Rror, Smith609, Snek01, Tabletop, UtherSRG, 21 anonymous edits Squid giant axon Source: http://en.wikipedia.org/w/index.php?oldid=516331936 Contributors: Alkaloids, Altenmann, Alvis, Diberri, Jacopo Werther, Kalexander, MementoVivere, Mgiganteus1, Nrets, Orlandoturner, Pmjboyle, Robertgreer, Synaptidude, Welsh, Whosasking, Wiseoldman123, Woreno, 19 anonymous edits Cuttlebone Source: http://en.wikipedia.org/w/index.php?oldid=519089517 Contributors: Ameima, Darklilac, DavidFarmbrough, Dawynn, Dentren, Eadouth1231, Invertzoo, Lfstevens, Mervyn, Mgiganteus1, Morgankevinj huggle, Obsidian Soul, Obsidianearth, Paxsimius, Pluma, Reywas92, SecretDisc, Smith609, Speciate, Thumperward, 32 anonymous edits Septum Source: http://en.wikipedia.org/w/index.php?oldid=470184941 Contributors: Bobo192, DanielCD, Elkman, JHunterJ, Kaarel, M Alan Kazlev, Mgiganteus1, Nono64, Randomfrenchie, Snek01, 1 anonymous edits Aptychus Source: http://en.wikipedia.org/w/index.php?oldid=477461394 Contributors: DanielCD, Invertzoo, Kontos, Mgiganteus1, Smith609, Snek01, UtherSRG Orthocone Source: http://en.wikipedia.org/w/index.php?oldid=483275401 Contributors: Apokryltaros, Cephal-odd, Closedmouth, D053, Everyking, J.H.McDonnell, JukoFF, M Alan Kazlev, Mgiganteus1, TobbiM, UtherSRG, VanHelsing, 17 anonymous edits Phragmocone Source: http://en.wikipedia.org/w/index.php?oldid=470337434 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=470337496 Contributors: AvicAWB, 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=470337154 Contributors: Dawynn, Dina, M Alan Kazlev, Mgiganteus1, UtherSRG, 1 anonymous edits

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file:Squid komodo.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Squid_komodo.jpg License: Creative Commons Attribution-Sharealike 3.0 Contributors: Nhobgood File:Red Pencil Icon.png Source: http://en.wikipedia.org/w/index.php?title=File:Red_Pencil_Icon.png License: Creative Commons Zero Contributors: User:Peter coxhead Image:Sepia officinalis Linnaeus, 1758 .jpg Source: http://en.wikipedia.org/w/index.php?title=File:Sepia_officinalis_Linnaeus,_1758_.jpg License: Public Domain Contributors: Parent Gry Image:Benthoctopus sp.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Benthoctopus_sp.jpg License: Public Domain Contributors: NOAA/MBARI Image:Oktopus opening a container with screw cap 01.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Oktopus_opening_a_container_with_screw_cap_01.jpg License: Creative Commons Attribution-ShareAlike 3.0 Unported Contributors: User:MatthiasKabel Image:Euprymna scolopes (Bobtail squid) behavior .jpg Source: http://en.wikipedia.org/w/index.php?title=File:Euprymna_scolopes_(Bobtail_squid)_behavior_.jpg License: Creative Commons Attribution-Sharealike 3.0 Contributors: Nhobgood File:Nautilus pompilius (head).jpg Source: http://en.wikipedia.org/w/index.php?title=File:Nautilus_pompilius_(head).jpg License: Creative Commons Attribution-Sharealike 3.0 Contributors: Hans Hillewaert Image:Cuttlefish color.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Cuttlefish_color.jpg License: Creative Commons Attribution-Sharealike 3.0 Contributors: Nick Hobgood Image:Chtenopteryx sicula2 - from Commons.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Chtenopteryx_sicula2_-_from_Commons.jpg License: unknown Contributors: Image:Ocythoe tuberculata viscera - from Commons.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Ocythoe_tuberculata_viscera_-_from_Commons.jpg License: unknown Contributors: Bearcat Image:Octopus3.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Octopus3.jpg License: Creative Commons Attribution-Sharealike 3.0,2.5,2.0,1.0 Contributors: albert kok Image:Nautilus front.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Nautilus_front.jpg License: Creative Commons Attribution 2.5 Contributors: Original uploader was Profberger at en.wikipedia Image:Spirula spirula.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Spirula_spirula.jpg License: Public Domain Contributors: Ewald Rbsamen Image:Herklots 1859 I 2 Sepia officinalis - schelp.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Herklots_1859_I_2_Sepia_officinalis_-_schelp.jpg License: Public Domain Contributors: Herklots (scan by Tom Meijer, 26-6-2007) Image:Sepioteuthis lessoniana gladius.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Sepioteuthis_lessoniana_gladius.jpg License: Public Domain Contributors: L. 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Image:Ancient Greek pottery in the National Archaeological Museum in Athens 13.JPG Source: http://en.wikipedia.org/w/index.php?title=File:Ancient_Greek_pottery_in_the_National_Archaeological_Museum_in_Athens_13.JPG License: Creative Commons Attribution-ShareAlike 3.0 Unported Contributors: sailko Image:OctopusMocheLMC.jpg Source: http://en.wikipedia.org/w/index.php?title=File:OctopusMocheLMC.jpg License: GNU Free Documentation License Contributors: AxelBoldt, Badseed, Johnbod, Pieter Kuiper Image:Departure warrior Staatliche Antikensammlungen 1484 side A.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Departure_warrior_Staatliche_Antikensammlungen_1484_side_A.jpg License: Public Domain Contributors: User:Bibi Saint-Pol File:Octopuses in Tsukiji.JPG Source: http://en.wikipedia.org/w/index.php?title=File:Octopuses_in_Tsukiji.JPG License: Creative Commons Attribution-ShareAlike 3.0 Unported Contributors: Hu Totya File:Octopus 3-pronged tako.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Octopus_3-pronged_tako.jpg License: Creative Commons Attribution-Sharealike 3.0 Contributors: Rmirasol Image:Octoraw.JPG Source: http://en.wikipedia.org/w/index.php?title=File:Octoraw.JPG License: Creative Commons Attribution-Sharealike 3.0 Contributors: Beeblebrox Image:Octoboiled.JPG Source: http://en.wikipedia.org/w/index.php?title=File:Octoboiled.JPG License: Creative Commons Attribution-Sharealike 3.0 Contributors: Beeblebrox File:CirrothaumaMurDraw2.jpg Source: http://en.wikipedia.org/w/index.php?title=File:CirrothaumaMurDraw2.jpg License: Public Domain Contributors: Ewald Rbsamen File:Amphitretus.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Amphitretus.jpg License: Public Domain Contributors: Ewald Rbsamen file:Loligo vulgaris.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Loligo_vulgaris.jpg License: Creative Commons Attribution-Sharealike 3.0 Contributors: Hans Hillewaert File:Composite diagram illustrating basic squid features, ventral aspect.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Composite_diagram_illustrating_basic_squid_features,_ventral_aspect.jpg License: Public Domain Contributors: Obsidian Soul, 4 anonymous edits Image:Onykia ingens with non-erect penis.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Onykia_ingens_with_non-erect_penis.jpg License: Creative Commons Attribution-Sharealike 3.0 Contributors: Alexander Arkhipkin Image:Onykia ingens with erect penis.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Onykia_ingens_with_erect_penis.jpg License: Creative Commons Attribution-Sharealike 3.0 Contributors: Alexander Arkhipkin Image:Giant squid melb aquarium03.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Giant_squid_melb_aquarium03.jpg License: unknown Contributors: Fir0002, Lin1, Pristigaster, 2 anonymous edits File:Bathyteuthisabyssicola.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Bathyteuthisabyssicola.jpg License: Public Domain Contributors: Carl Chun File:Grimalditeuthis bonplandi.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Grimalditeuthis_bonplandi.jpg License: Public Domain Contributors: Jeanne Le Roux & L. Joubin File:Reversa1.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Reversa1.jpg License: Public Domain Contributors: Carl Chun File:Mastigoteuthis flammea.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Mastigoteuthis_flammea.jpg License: Public Domain Contributors: EugeneZelenko, Javaprog, Keith Edkins, Kilom691, Kristof vt, Lin1 File:Onychoteuthis banksii1.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Onychoteuthis_banksii1.jpg License: Public Domain Contributors: L. Joubin & Ch. Richard File:Cephalop.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Cephalop.jpg License: Public Domain Contributors: Carl Chun Image:Fried calamari.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Fried_calamari.jpg License: GNU Free Documentation License Contributors: en:User:Chensiyuan file:Sepia latimanus (Reef cuttlefish) dark coloration.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Sepia_latimanus_(Reef_cuttlefish)_dark_coloration.jpg License: Creative Commons Attribution-Sharealike 3.0 Contributors: Nhobgood Nick Hobgood File:Cuttlefish.ogv Source: http://en.wikipedia.org/w/index.php?title=File:Cuttlefish.ogv License: Creative Commons Attribution 2.0 Contributors: prilfish (Silke Baron) Image:Camouflage.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Camouflage.jpg License: GNU Free Documentation License Contributors: Raul654 Image:Cuttlefishhead.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Cuttlefishhead.jpg License: Public Domain Contributors: FireFly5 File:Sepia eyelid shape.theora.ogv Source: http://en.wikipedia.org/w/index.php?title=File:Sepia_eyelid_shape.theora.ogv License: Creative Commons Attribution-Sharealike 3.0 Contributors: User:Tliglesias File:SuckersOfACuttle-Fish,1834.PNG Source: http://en.wikipedia.org/w/index.php?title=File:SuckersOfACuttle-Fish,1834.PNG License: Public Domain Contributors: Google Books Image:Metasepia pfefferi 1.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Metasepia_pfefferi_1.jpg License: Creative Commons Attribution 2.0 Contributors: Bricktop, Dodo, Lin1 Image:kalamar.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Kalamar.jpg License: Public Domain Contributors: Original uploader was Borazont at en.wikipedia. Later version(s) were uploaded by Mgiganteus1 at en.wikipedia. Image:linguine with cuttlefish.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Linguine_with_cuttlefish.jpg License: Public Domain Contributors: Original uploader was Schellack at en.wikipedia file:Orthoceras BW.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Orthoceras_BW.jpg License: Creative Commons Attribution 3.0 Contributors: Nobu Tamura (http://spinops.blogspot.com) Image:Nautilus profile.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Nautilus_profile.jpg License: GNU Free Documentation License Contributors: Lee R Berger Image:Nautiloid trilacinoceras.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Nautiloid_trilacinoceras.jpg License: GNU Free Documentation License Contributors: User:Dlloyd Image:OrdNautiloidInternalMold.jpg Source: http://en.wikipedia.org/w/index.php?title=File:OrdNautiloidInternalMold.jpg License: Public Domain Contributors: Wilson44691 file:Nautilus Palau.JPG Source: http://en.wikipedia.org/w/index.php?title=File:Nautilus_Palau.JPG License: Creative Commons Attribution-Sharealike 3.0 Contributors: Manuae File:Nautilus anatomy.png Source: http://en.wikipedia.org/w/index.php?title=File:Nautilus_anatomy.png License: Public Domain Contributors: John Denis Macdonald Image:NautilusCutawayLogarithmicSpiral.jpg Source: http://en.wikipedia.org/w/index.php?title=File:NautilusCutawayLogarithmicSpiral.jpg License: GNU Free Documentation License Contributors: User:Chris 73 Image:NautilusTop.jpg Source: http://en.wikipedia.org/w/index.php?title=File:NautilusTop.jpg License: GNU Free Documentation License Contributors: DanielCD, Haplochromis, Javaprog, Petwoe Image:NautilusBottom.jpg Source: http://en.wikipedia.org/w/index.php?title=File:NautilusBottom.jpg License: GNU Free Documentation License Contributors: DanielCD, Haplochromis, Javaprog, Petwoe Image:Nautilus oceanworld thailand.png Source: http://en.wikipedia.org/w/index.php?title=File:Nautilus_oceanworld_thailand.png License: Public Domain Contributors: Keskival Image:Size frequency distribution for Nautilus pompilius at Osprey Reef.png Source: http://en.wikipedia.org/w/index.php?title=File:Size_frequency_distribution_for_Nautilus_pompilius_at_Osprey_Reef.png License: Creative Commons Attribution 2.5 Contributors: Dunstan AJ, Ward PD, Marshall NJ Image:Sexual dimorphism in mature male and female Nautilus pompilius.png Source: http://en.wikipedia.org/w/index.php?title=File:Sexual_dimorphism_in_mature_male_and_female_Nautilus_pompilius.png License: Creative Commons Attribution 2.5 Contributors: Dunstan AJ, Ward PD, Marshall NJ File:Comparison of Nautilus capture rates with trapping depth.png Source: http://en.wikipedia.org/w/index.php?title=File:Comparison_of_Nautilus_capture_rates_with_trapping_depth.png License: Creative Commons Attribution 2.5 Contributors: Dunstan AJ, Ward PD, Marshall NJ File:Deepest record of Nautilus 703 meters.png Source: http://en.wikipedia.org/w/index.php?title=File:Deepest_record_of_Nautilus__703_meters.png License: Creative Commons Attribution 2.5 Contributors: Dunstan AJ, Ward PD, Marshall NJ File:Allonautilus vs Nautilus.png Source: http://en.wikipedia.org/w/index.php?title=File:Allonautilus_vs_Nautilus.png License: Public Domain Contributors: Antonov File:Eutrephoceras dorbignyanum (Forbes in Darwin, 1846) - Santiago specimen.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Eutrephoceras_dorbignyanum_(Forbes_in_Darwin,_1846)_-_Santiago_specimen.jpg License: Public Domain Contributors: Obsidian Soul Image:Nautilus species shells.png Source: http://en.wikipedia.org/w/index.php?title=File:Nautilus_species_shells.png License: GNU Free Documentation License Contributors: User:Mgiganteus1

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file:Asteroceras BW.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Asteroceras_BW.jpg License: Creative Commons Attribution 3.0 Contributors: Nobu Tamura (http://spinops.blogspot.com) Image:Ammoniteplit.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Ammoniteplit.jpg License: Creative Commons Attribution-Share Alike Contributors: John Alan Elson Image:Iridescent Ammonite Fossil.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Iridescent_Ammonite_Fossil.jpg License: Creative Commons Attribution 3.0 Contributors: JamesPFisherIII File:Placenticeratidae - Placenticeras whitfieldi.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Placenticeratidae_-_Placenticeras_whitfieldi.jpg License: Creative Commons Attribution-Sharealike 3.0 Contributors: Hectonichus Image:Ammonite Jeletzkytes.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Ammonite_Jeletzkytes.jpg License: GNU Free Documentation License Contributors: User:Dlloyd Image:Ammonite Asteroceras.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Ammonite_Asteroceras.jpg License: GNU Free Documentation License Contributors: Dlloyd Image:Amonite-portugal.JPG Source: http://en.wikipedia.org/w/index.php?title=File:Amonite-portugal.JPG License: Creative Commons Attribution-Sharealike 3.0 Contributors: User:LMCoelho Image:Haeckel Ammonitida.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Haeckel_Ammonitida.jpg License: Public Domain Contributors: ComputerHotline, Dysmorodrepanis, Kevmin, M0tty, Pengo, Ragesoss, Ulrichstill, 1 anonymous edits Image:BaculitidArticulated.jpg Source: http://en.wikipedia.org/w/index.php?title=File:BaculitidArticulated.jpg License: Creative Commons Attribution-Sharealike 3.0 Contributors: User:Wilson44691 Image:DiscoscaphitesirisCretaceous.jpg Source: http://en.wikipedia.org/w/index.php?title=File:DiscoscaphitesirisCretaceous.jpg License: Public Domain Contributors: Wilson44691 File:Trigonellites latus.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Trigonellites_latus.jpg License: Public Domain Contributors: Cooke, A. H., Shipley, A. E. & Reed, F. R. C. File:Parapuzosia seppenradensis 5.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Parapuzosia_seppenradensis_5.jpg License: GNU Free Documentation License Contributors: Markus Schweiss Image:Hoploscaphites ammonite.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Hoploscaphites_ammonite.jpg License: GNU Free Documentation License Contributors: User:DanielCD, User:Deadstar Image:IridescentAmmonite.jpg Source: http://en.wikipedia.org/w/index.php?title=File:IridescentAmmonite.jpg License: GNU Free Documentation License Contributors: DanielCD, Kevmin, Saperaud, Tonton Bernardo, Ulrichstill, 1 anonymous edits File:The Childrens Museum of Indianapolis - Ammonites.jpg Source: http://en.wikipedia.org/w/index.php?title=File:The_Childrens_Museum_of_Indianapolis_-_Ammonites.jpg License: unknown Contributors: HstryQT, Kevmin, Missvain file:BelemnitesJurassicWyoming.jpg Source: http://en.wikipedia.org/w/index.php?title=File:BelemnitesJurassicWyoming.jpg License: Public Domain Contributors: Wilson44691 Image:BelemniteDB2.jpg Source: http://en.wikipedia.org/w/index.php?title=File:BelemniteDB2.jpg License: Creative Commons Attribution-Sharealike 3.0,2.5,2.0,1.0 Contributors: DiBgd, Kevmin, Putnik, Ra'ike Image:Belemnit.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Belemnit.jpg License: GNU Free Documentation License Contributors: AnRo0002, DanielCD, Glenn, Kevmin, Lin1, Muriel Gottrop, 1 anonymous edits Image:belemnite at bristol museum arp.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Belemnite_at_bristol_museum_arp.jpg License: Public Domain Contributors: AnRo0002, Arpingstone, Kevmin, PurpleHz Image:ZoharBelemnite.JPG Source: http://en.wikipedia.org/w/index.php?title=File:ZoharBelemnite.JPG License: Public Domain Contributors: Wilson44691 file:Papierboot Argonauta 200705181139.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Papierboot_Argonauta_200705181139.jpg License: Creative Commons Attribution-Sharealike 2.0 Contributors: Original uploader was Bernd Hofmann at de.wikipedia Image:Argonauta nodosa lithograph.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Argonauta_nodosa_lithograph.jpg License: Public Domain Contributors: Citron, Lin1, Mgiganteus Image:Argonauta sp.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Argonauta_sp.jpg License: Public Domain Contributors: Ewald Rbsamen Image:Argonauta hians male.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Argonauta_hians_male.jpg License: Public Domain Contributors: Ewald Rbsamen Image:Argonauta nodosa with eggcase lithograph.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Argonauta_nodosa_with_eggcase_lithograph.jpg License: Public Domain Contributors: Citron, Lin1, Mgiganteus Image:Argonauta argo shell.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Argonauta_argo_shell.jpg License: Public Domain Contributors: Verrill, A. E. 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N. Lea Image:LongArmSquid.jpg Source: http://en.wikipedia.org/w/index.php?title=File:LongArmSquid.jpg License: Creative Commons Zero Contributors: NOAA File:GiantCuttlefish6.jpg Source: http://en.wikipedia.org/w/index.php?title=File:GiantCuttlefish6.jpg License: Creative Commons Attribution-Sharealike 3.0 Contributors: Jacob Bridgeman Image:Argonauta hians.JPG Source: http://en.wikipedia.org/w/index.php?title=File:Argonauta_hians.JPG License: Public Domain Contributors: Original uploader was XQ fan at en.wikipedia Image:Spirula fg1.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Spirula_fg1.jpg License: Creative Commons Attribution-Sharealike 2.5 Contributors: Fritz Geller-Grimm Image:Parapuzosia seppenradensis cast.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Parapuzosia_seppenradensis_cast.jpg License: GNU Free Documentation License Contributors: Markus Schweiss File:Megateuthis 1.JPG Source: http://en.wikipedia.org/w/index.php?title=File:Megateuthis_1.JPG License: Creative Commons Attribution-Sharealike 3.0 Contributors: Ghedoghedo File:Chtenopteryx sicula2 - from Commons.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Chtenopteryx_sicula2_-_from_Commons.jpg License: unknown Contributors: File:Arrosnegre.png Source: http://en.wikipedia.org/w/index.php?title=File:Arrosnegre.png License: Public Domain Contributors: Gveret Tered, Lobo, Toniher Image:Illex illecebrosus arm.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Illex_illecebrosus_arm.jpg License: Public Domain Contributors: James H. Emerton Image:Illex illecebrosus tentacle.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Illex_illecebrosus_tentacle.jpg License: Public Domain Contributors: James H. Emerton File:Taningia danae7.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Taningia_danae7.jpg License: Public Domain Contributors: L. Joubin File:Semirossia tenera2.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Semirossia_tenera2.jpg License: Public Domain Contributors: Lin1, Lycaon, Mgiganteus File:Rossia glaucopis mouth.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Rossia_glaucopis_mouth.jpg License: Public Domain Contributors: James H. Emerton File:Bathypolypus arcticus.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Bathypolypus_arcticus.jpg License: Public Domain Contributors: James H. Emerton

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File:Img octopus arm and suckers 057513.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Img_octopus_arm_and_suckers_057513.jpg License: Creative Commons Attribution 3.0 Contributors: Henryhartley File:Suckers of octopus by steve lodefink.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Suckers_of_octopus_by_steve_lodefink.jpg License: Creative Commons Attribution 2.0 Contributors: Citron, FlickreviewR, Opponent File:Octopus-sucker-deformity.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Octopus-sucker-deformity.jpg License: Creative Commons Attribution-Sharealike 3.0 Contributors: sehsuan (talk) File:Octapus forked arm.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Octapus_forked_arm.jpg License: Creative Commons Attribution-Sharealike 3.0 Contributors: User:Dkontogiannnis File:Todarodes pacificus arm.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Todarodes_pacificus_arm.jpg License: Public Domain Contributors: A. Pollock File:Tentacule Abraliopsis morisi-2.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Tentacule_Abraliopsis_morisi-2.jpg License: Public Domain Contributors: Tentacule_Abraliopsis_morisi.jpg: Carl Chun derivative work: Mgiganteus (talk) File:AncistroteuClub.jpg Source: http://en.wikipedia.org/w/index.php?title=File:AncistroteuClub.jpg License: Public Domain Contributors: Javaprog, Kersti Nebelsiek, Lin1, Mgiganteus File:Giant squid tentacle club.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Giant_squid_tentacle_club.jpg License: Public Domain Contributors: Original uploader was Mgiganteus1 at en.wikipedia File:Austrorossia mastigophora2.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Austrorossia_mastigophora2.jpg License: Public Domain Contributors: Kersti Nebelsiek, Lin1, Lycaon, Mgiganteus File:Berryteuthis magister5.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Berryteuthis_magister5.jpg License: Public Domain Contributors: Kersti Nebelsiek, Lin1, Mgiganteus File:Idioteuthis cordiformis4.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Idioteuthis_cordiformis4.jpg License: Public Domain Contributors: Javaprog, Kersti Nebelsiek, Lin1, Mgiganteus File:Iridoteuthis iris tentacle.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Iridoteuthis_iris_tentacle.jpg License: Public Domain Contributors: H. V. Poor File:Mastigoteuthis glaukopis2.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Mastigoteuthis_glaukopis2.jpg License: Public Domain Contributors: Javaprog, Kersti Nebelsiek, Lin1, Mgiganteus File:Morotuthis ingens2.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Morotuthis_ingens2.jpg License: Public Domain Contributors: Kersti Nebelsiek, Mgiganteus File:Semirossia tenera3.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Semirossia_tenera3.jpg License: Public Domain Contributors: Kersti Nebelsiek, Lin1, Lycaon, Mgiganteus File:Spirula tentacle club.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Spirula_tentacle_club.jpg License: Public Domain Contributors: Ewald Rbsamen File:Todarodes pacificus tentacle.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Todarodes_pacificus_tentacle.jpg License: Public Domain Contributors: A. Pollock File:Idioteuthis cordiformis2.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Idioteuthis_cordiformis2.jpg License: Public Domain Contributors: Javaprog, Kersti Nebelsiek, Lin1, Mgiganteus File:Idioteuthis cordiformis3.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Idioteuthis_cordiformis3.jpg License: Public Domain Contributors: Javaprog, Kersti Nebelsiek, Lin1, Mgiganteus File:Idioteuthis cordiformis5.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Idioteuthis_cordiformis5.jpg License: Public Domain Contributors: Javaprog, Kersti Nebelsiek, Lin1, Mgiganteus File:Idioteuthis latipinna2.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Idioteuthis_latipinna2.jpg License: Public Domain Contributors: Javaprog, Kersti Nebelsiek, Lin1, Mgiganteus File:Idioteuthis latipinna3.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Idioteuthis_latipinna3.jpg License: Public Domain Contributors: Javaprog, Kersti Nebelsiek, Lin1, Mgiganteus File:Magnapinna talismani2.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Magnapinna_talismani2.jpg License: Public Domain Contributors: Javaprog, Kersti Nebelsiek, Lin1, Mgiganteus File:Magnapinna talismani3.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Magnapinna_talismani3.jpg License: Public Domain Contributors: Javaprog, Kersti Nebelsiek, Lin1, Mgiganteus File:Mastigoteuthis agassizii.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Mastigoteuthis_agassizii.jpg License: Public Domain Contributors: Javaprog, Kersti Nebelsiek, Lin1, Mgiganteus File:Mastigoteuthis agassizii3.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Mastigoteuthis_agassizii3.jpg License: Public Domain Contributors: Javaprog, Kersti Nebelsiek, Lin1, Mgiganteus File:Mastigoteuthis atlantica2.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Mastigoteuthis_atlantica2.jpg License: Public Domain Contributors: Javaprog, Kersti Nebelsiek, Lin1, Mgiganteus File:Mastigoteuthis atlantica3.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Mastigoteuthis_atlantica3.jpg License: Public Domain Contributors: Javaprog, Kersti Nebelsiek, Lin1, Mgiganteus File:Mastigoteuthis dentata.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Mastigoteuthis_dentata.jpg License: Public Domain Contributors: Kersti Nebelsiek, Lin1, Mgiganteus File:Mastigoteuthis grimaldii2.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Mastigoteuthis_grimaldii2.jpg License: Public Domain Contributors: Javaprog, Kersti Nebelsiek, Lin1, Mgiganteus File:Mastigoteuthis magna2.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Mastigoteuthis_magna2.jpg License: Public Domain Contributors: Javaprog, Kersti Nebelsiek, Lin1, Mgiganteus File:Hectocotyle1.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Hectocotyle1.jpg License: Public Domain Contributors: Georges Cuvier Image:Hectocotylized arm of a squid.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Hectocotylized_arm_of_a_squid.jpg License: Public Domain Contributors: Obsidian Soul Image:Hectocotylized arm of an octopod.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Hectocotylized_arm_of_an_octopod.jpg License: Public Domain Contributors: Obsidian Soul File:Abraliopsis morisi hectocotylus-English.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Abraliopsis_morisi_hectocotylus-English.jpg License: Public Domain Contributors: Abraliopsis_morisi_hectocotylus.jpg: Carl Chun derivative work: Mgiganteus (talk) File:Argonauta bottgeri hectocotylus-2.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Argonauta_bottgeri_hectocotylus-2.jpg License: Public Domain Contributors: Argonauta_bottgeri_hectocotylus.jpg: Berry, S. Stillman derivative work: Mgiganteus (talk) File:Bathypolypus arcticus hectocotylus-2.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Bathypolypus_arcticus_hectocotylus-2.jpg License: Public Domain Contributors: Bathypolypus_arcticus_hectocotylus.jpg: James H. Emerton derivative work: Mgiganteus (talk) File:Graneledone verrucosa hectocotylus.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Graneledone_verrucosa_hectocotylus.jpg License: Public Domain Contributors: Joubin File:Haliphron atlanticus hectocotylus.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Haliphron_atlanticus_hectocotylus.jpg License: Public Domain Contributors: A.E Verrill (Addison Emery), 1839-1926; File:Scaeurgus patagiatus hectocotylus-2.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Scaeurgus_patagiatus_hectocotylus-2.jpg License: Public Domain Contributors: Scaeurgus_patagiatus_hectocotylus.jpg: R. L. Hudson derivative work: Mgiganteus (talk) File:Tremoctopus violaceus5.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Tremoctopus_violaceus5.jpg License: Public Domain Contributors: R. L. Hudson File:Uroteuthis duvauceli hectocotylus.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Uroteuthis_duvauceli_hectocotylus.jpg License: Public Domain Contributors: A. Pollock Image:cuttlefish.png Source: http://en.wikipedia.org/w/index.php?title=File:Cuttlefish.png License: Public Domain Contributors: unknown. Image:Snail-front-0A.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Snail-front-0A.jpg License: Creative Commons Attribution-ShareAlike 3.0 Unported Contributors: Adamantios Image:White abalone Haliotis sorenseni.jpg Source: http://en.wikipedia.org/w/index.php?title=File:White_abalone_Haliotis_sorenseni.jpg License: Creative Commons Attribution-Sharealike 3.0 Contributors: Original uploader was Geographer at en.wikipedia Image:Drosera capensis bend.JPG Source: http://en.wikipedia.org/w/index.php?title=File:Drosera_capensis_bend.JPG License: Creative Commons Attribution-ShareAlike 3.0 Unported Contributors: Infrogmation, NoahElhardt, Wim b, Yarl, 3 anonymous edits Image:Evolution eye.svg Source: http://en.wikipedia.org/w/index.php?title=File:Evolution_eye.svg License: GNU Free Documentation License Contributors: Caerbannog

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File:Eye squid.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Eye_squid.jpg License: Public Domain Contributors: Carl Chun Image:Octopusv cropped.JPG Source: http://en.wikipedia.org/w/index.php?title=File:Octopusv_cropped.JPG License: Creative Commons Attribution-Sharealike 3.0,2.5,2.0,1.0 Contributors: Octopusv.JPG: Gronk derivative work: Mgiganteus (talk) Image:Squid eye.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Squid_eye.jpg License: Creative Commons Attribution 2.0 Contributors: wildxplorer Image:Sepia officinalis Linnaeus, 1758 cropped.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Sepia_officinalis_Linnaeus,_1758_cropped.jpg License: Public Domain Contributors: Sepia_officinalis_Linnaeus,_1758.jpg: Parent Gry derivative work: Mgiganteus (talk) Image:Nautilus pompilius (head).jpg Source: http://en.wikipedia.org/w/index.php?title=File:Nautilus_pompilius_(head).jpg License: Creative Commons Attribution-Sharealike 3.0 Contributors: Hans Hillewaert Image:Zfishchroma.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Zfishchroma.jpg License: Public Domain Contributors: Original uploader was Rockpocket at en.wikipedia Image:C Calyptratus female.jpg Source: http://en.wikipedia.org/w/index.php?title=File:C_Calyptratus_female.jpg License: GNU Free Documentation License Contributors: Original uploader was Geoff at en.wikipedia Image:Zebrafish embryos.png Source: http://en.wikipedia.org/w/index.php?title=File:Zebrafish_embryos.png License: Creative Commons Attribution 2.5 Contributors: Adam Amsterdam, Massachusetts Institute of Technology, Boston, Massachusetts, United States. File:Pseudochromis Diadema Larry.JPG Source: http://en.wikipedia.org/w/index.php?title=File:Pseudochromis_Diadema_Larry.JPG License: Creative Commons Attribution 3.0 Contributors: Mrbrefast Image:melanophore.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Melanophore.jpg License: Public Domain Contributors: Original uploader was Rockpocket at en.wikipedia Image:Neural.crest.cells.migration.svg Source: http://en.wikipedia.org/w/index.php?title=File:Neural.crest.cells.migration.svg License: Public Domain Contributors: Neuralcrestroute.jpg: Original uploader was Rockpocket at en.wikipedia derivative work: Mithril (talk) Image:Giant clam komodo.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Giant_clam_komodo.jpg License: Creative Commons Attribution-Sharealike 3.0 Contributors: Nhobgood Nick Hobgood File:Cypraea chinensis with partially extended mantle.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Cypraea_chinensis_with_partially_extended_mantle.jpg License: Public Domain Contributors: Bricktop, GrahamBould, JoJan, JorisvS, Kilom691 File:Cypraea chinensis with fully extended mantle.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Cypraea_chinensis_with_fully_extended_mantle.jpg License: Public Domain Contributors: Bricktop, GrahamBould, JoJan File:SnailWynaad.jpg Source: http://en.wikipedia.org/w/index.php?title=File:SnailWynaad.jpg License: Creative Commons Attribution-Sharealike 3.0 Contributors: L. Shyamal File:Bielzia coerulans-3.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Bielzia_coerulans-3.jpg License: Creative Commons Attribution-Sharealike 2.5 Contributors: User:Snek01 File:Megapallifera mutabilis.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Megapallifera_mutabilis.jpg License: Creative Commons Attribution-Sharealike 2.0 Contributors: Paul J. Morris File:Haliotis asinina anatomy.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Haliotis_asinina_anatomy.jpg License: Creative Commons Attribution 2.0 Contributors: Daniel J Jackson, Carmel McDougall, Kathryn Green, Fiona Simpson, Gert Wrheide & Bernard M Degnan File:Haliotis asinina anatomy 2.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Haliotis_asinina_anatomy_2.jpg License: Creative Commons Attribution 2.0 Contributors: Daniel J Jackson, Carmel McDougall, Kathryn Green, Fiona Simpson, Gert Wrheide & Bernard M Degnan File:A clam.jpg Source: http://en.wikipedia.org/w/index.php?title=File:A_clam.jpg License: Creative Commons Attribution 2.0 Contributors: FlickrLickr, FlickreviewR, Kersti Nebelsiek, 3 anonymous edits File:Cymbiola magnifica.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Cymbiola_magnifica.jpg License: Creative Commons Attribution-Sharealike 2.0 Contributors: Richard Ling File:Nassarius tiarula.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Nassarius_tiarula.jpg License: Public Domain Contributors: Steve Lonhart (SIMoN / MBNMS) File:Heron Island Giant balor S01.OGG Source: http://en.wikipedia.org/w/index.php?title=File:Heron_Island_Giant_balor_S01.OGG License: Creative Commons Attribution-Sharealike 3.0 Contributors: Sirrob01 File:Pomacea canaliculata siphonout.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Pomacea_canaliculata_siphonout.jpg License: GNU Free Documentation License Contributors: Kristjan, Mattes, Pristigaster, Schimmelreiter File:Pomacea paludosa drawing.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Pomacea_paludosa_drawing.jpg License: Public Domain Contributors: drawn by Helen Lawson ( 1854), engraved by the Alexander Lawson firm, colored by Helen Lawson. File:GooeyduckSeafood.jpg Source: http://en.wikipedia.org/w/index.php?title=File:GooeyduckSeafood.jpg License: Creative Commons Attribution-Sharealike 3.0,2.5,2.0,1.0 Contributors: Original uploader was Bachcell at en.wikipedia File:Muscheln mit Sipho Nahaufnahme.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Muscheln_mit_Sipho_Nahaufnahme.jpg License: Creative Commons Attribution-Sharealike 3.0,2.5,2.0,1.0 Contributors: Stefan Didam - Schmallenberg File:Venus verrucosa 3.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Venus_verrucosa_3.jpg License: Public Domain Contributors: Kurt Floericke (1869-1934) File:Valve-InternalView.png Source: http://en.wikipedia.org/w/index.php?title=File:Valve-InternalView.png License: Creative Commons Attribution-ShareAlike 1.0 Generic Contributors: Muriel Gottrop Image:Cuttlebone.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Cuttlebone.jpg License: Creative Commons Attribution-ShareAlike 3.0 Unported Contributors: Dysmorodrepanis, Freaky Fries, Mgiganteus, Moumou82, Salix, Snek01, TomCatX, Totodu74, 1 anonymous edits File:Herklots 1859 I 2 Sepia officinalis - schelp.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Herklots_1859_I_2_Sepia_officinalis_-_schelp.jpg License: Public Domain Contributors: Herklots (scan by Tom Meijer, 26-6-2007) File:Turtle and Sepia.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Turtle_and_Sepia.jpg License: GNU Free Documentation License Contributors: Foto von Richard Mayer. Original uploader was Richard Mayer at de.wikipedia File:3D visualisation of CT-data of a cuttlebone 01.jpg Source: http://en.wikipedia.org/w/index.php?title=File:3D_visualisation_of_CT-data_of_a_cuttlebone_01.jpg License: Creative Commons Attribution-Sharealike 3.0 Contributors: User:SecretDisc File:3D visualisation of CT-data of a cuttlebone 03.jpg Source: http://en.wikipedia.org/w/index.php?title=File:3D_visualisation_of_CT-data_of_a_cuttlebone_03.jpg License: Creative Commons Attribution-Sharealike 3.0 Contributors: User:SecretDisc File:3D visualisation of CT-data of a cuttlebone 04.jpg Source: http://en.wikipedia.org/w/index.php?title=File:3D_visualisation_of_CT-data_of_a_cuttlebone_04.jpg License: Creative Commons Attribution-Sharealike 3.0 Contributors: User:SecretDisc File:3D visualisation of CT-data of a cuttlebone 05.jpg Source: http://en.wikipedia.org/w/index.php?title=File:3D_visualisation_of_CT-data_of_a_cuttlebone_05.jpg License: Creative Commons Attribution-Sharealike 3.0 Contributors: User:SecretDisc File:Flight through image stack of CT-data of a cuttlebone, top view.ogg Source: http://en.wikipedia.org/w/index.php?title=File:Flight_through_image_stack_of_CT-data_of_a_cuttlebone,_top_view.ogg License: Creative Commons Attribution-Sharealike 3.0 Contributors: User:SecretDisc File:Aligned flight through image stack of CT-data of a cuttlebone, top view.ogg Source: http://en.wikipedia.org/w/index.php?title=File:Aligned_flight_through_image_stack_of_CT-data_of_a_cuttlebone,_top_view.ogg License: Creative Commons Attribution-Sharealike 3.0 Contributors: User:SecretDisc File:Aptychi examples function.PNG Source: http://en.wikipedia.org/w/index.php?title=File:Aptychi_examples_function.PNG License: Public Domain Contributors: Antonov File:Aptychus.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Aptychus.jpg License: GNU Free Documentation License Contributors: DanielCD, Kevmin, Saperaud, Snek01, 1 anonymous edits File:Duoaptico.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Duoaptico.jpg License: Creative Commons Attribution-Sharealike 3.0,2.5,2.0,1.0 Contributors: Bramfab File:Lingulaticeras solenoides.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Lingulaticeras_solenoides.jpg License: Creative Commons Attribution-Sharealike 3.0 Contributors: Ghedoghedo File:Perisphinctes with aptychi.JPG Source: http://en.wikipedia.org/w/index.php?title=File:Perisphinctes_with_aptychi.JPG License: Public Domain Contributors: Antonov

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File:Ooland baculite Image0022 lo res 3 x1 7.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Ooland_baculite_Image0022_lo_res_3_x1_7.jpg License: GNU Free Documentation License Contributors: Original uploader was Dcwade at en.wikipedia Image:Siphuncle.gif Source: http://en.wikipedia.org/w/index.php?title=File:Siphuncle.gif License: Attribution Contributors: Dysmorodrepanis, Kevmin, P4en, Smith609 Image:Nautilus pompilius.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Nautilus_pompilius.jpg License: Public Domain Contributors: Citron, Daggerstab, Lin1

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