Cephalopod
A variety of cephalopod forms from Ernst Haeckel's 1904 Kunstformen der Natur | |||||||
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A cephalopod (Greek plural Κεφαλόποδα (kephalópoda); "head-feet") is any member of the mollusc class Cephalopoda, characterized by bilateral body symmetry, a prominent head, and a modification of the mollusk foot, a muscular hydrostat, into the form of arms or tentacles. Teuthology, a branch of malacology, is the study of cephalopods. These creatures became dominant around the Ordovician period. The fishing industry name for this class is inkfish, referring to many cephalopods' ability to squirt ink.
The class contains two extant subclasses. In the Coleoidea, the mollusk shell has been internalized or is absent; this subclass includes the octopus, squid, and cuttlefish. In the Nautiloidea, the shell remains; this subclass includes the nautilus. About 800 distinct living species of cephalopods have been identified. Two important extinct taxa are Ammonoidea, the ammonites, and Belemnoidea, the belemnites.
Distribution
There are around 800 extant species of cephalopod,[1] although new species continue to be described. It is estimated that around 11,000 extinct taxa have been described,[2] although the soft-bodied nature of cephalopods means that they are not easily fossilised.[3]
Cephalopods are found in all the oceans of Earth. None of them can tolerate freshwater, but the brief squid, Lolliguncula brevis, found in Chesapeake Bay may be a notable exception in that it tolerates brackish water which has a low salinity.[4]
Cephalopods occupy most of the depth of the ocean, from hydrothermal vents to the sea surface. Their diversity is greatest near the equator (~40 species retrieved in nets at 11°N by a diversity study) and decreases towards the poles (~5 species captured at 60°N).[5]:11
An octopus opening a container with a screw cap
Nervous system and behaviour
See also: Cephalopod intelligence and squid giant axon
Cephalopods are widely regarded as the most intelligent of the invertebrates and have well developed senses and large brains; larger than the brains of gastropods. The nervous system of cephalopods is the most complex of the invertebrates,[citation needed] and their brain to body mass ratio falls between that of warm and cold blooded vertebrates.[5]:14 The giant nerve fibers of the cephalopod mantle have been a favorite[vague] experimental material of neurophysiologists for many years; their large diameter (due to lack of myelination) makes them easier to study.[citation needed]
Cephalopods are social creatures; when isolated from their own kind, they will take to shoaling with fish.[6]
Some cephalopods are able to fly distances up to 50 m. While the organisms are not particularly aerodynamic, they achieve these rather impressive ranges by use of jet-propulsion; water continues to be expelled from the funnel while the organism is in flight.[citation needed]
Senses
Cephalopods have advanced vision, can detect gravity with statocysts, and have a variety of chemical sense organs.[5]:34 Octopuses use their tentacles to explore their environment and can use them for depth perception.[5]
Vision
Main articles: Cephalopod eye and mollusk eye
Most cephalopods rely on vision to detect predators and prey, and to communicate with one another.[7] Consequently, cephalopod vision is acute: training experiments have shown that the Common Octopus can distinguish the brightness, size, shape, and horizontal or vertical orientation of objects. The morphological construction gives cephalopod eyes the same performance as sharks'; however, their construction differs as cephalopods lack a cornea, and have an everted retina.[7] Cephalopods' eyes are also sensitive to the plane of polarization of light. Surprisingly—given their ability to change color—all octopuses[8] and most cephalopods[9] 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.[10] Evidence of color vision has been found in only one species, the Sparkling Enope Squid.[9]
The primitive nautilus eye functions similarly to a pinhole camera.
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.
Hearing
Cephalopods can use their statocyst to detect sound.[11]
Use of light
This Broadclub Cuttlefish (Sepia latimanus) can go from camouflage tans and browns (top) to yellow with dark highlights (bottom) in less than a second.
Most cephalopods possess chromatophores - that is, coloured pigments - which they can use in a startling array of fashions.[5] 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.[5] The bioluminescence is produced by bacterial symbionts; the host cephalopod is able to detect the light produced by these organisms.[12] 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.[5] It is not certain whether bioluminescence is actually of epithelial origin or if it is a bacterial production.[13]
Colouration
Colouration can be changed in milliseconds as they adapt to their environment,[5] and the pigment cells are expandable by muscular contraction.[13] Colouration is typically more pronounced in near-shore species than those living in the open ocean, whose functions tend to be restricted to camouflage by breaking their outline.[5]:2
Evidence of original colouration 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.[14] Devonian cephalopods bear more complex colour patterns, whose function may be more complex.[15]
Ink
Main articles: Cephalopod ink and ink sac
With the exception of the Nautilidae and the species of octopus belonging to the suborder Cirrina[16], all known cephalopods have an ink sac, which can be used to expel a cloud of dark ink to confuse predators.[8] This sac is a muscular bag which originated as an extension of the hind gut. It lies beneath the gut and opens into the anus, into which its contents – almost pure melanin – can be squirted; its proximity to the base of the funnel means that the ink can be distributed by ejected water as the cephalopod uses its jet propulsion.[8] 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.[8] For more information, see Inking behaviors.
Single Oxygenated Functional Unit from the hemocyanin of an octopus
Circulatory system
Cephalopods are the only mollusks with a closed circulatory system. They 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.[17]
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.[18]
Gills
The gills of cephalopods are supported by a skeleton of robust fibrous proteins; the lack of mucopolysaccharides distinguishes this matrix from cartilage.[19] [20]
Respiration
Cephalopods exchange gasses with the seawater by forcing water through their gills, which are attached to the roof of the organism.[21]:488[22] 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.[22] 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.[21] The water flow is controlled by contractions of the radial and circular mantle cavity muscles.[23]
Their gill lamellae are supported by a cartilage framework.[22] The gills are also thought to be involved in excretion, with NH4+ being swapped with K+ from the seawater.[22]
Locomotion and buoyancy
Octopuses swim headfirst, with arms trailing behind
While all cephalopods can move by jet propulsion, this is a very energy-consuming way to travel compared to the tail propulsion used by fish.[24] The relative efficiency of jet propulsion decreases further as animal size increases. Since the Paleozoic era, as competition with fish produced an environment where efficient motion was crucial to survival, jet propulsion has taken a back role, with fins and tentacles used to maintain a steady velocity.[3] Whilst jet propulsion is never the sole mode of locomotion,[3]:208 the stop-start motion provided by the jets continues to be useful for providing bursts of high speed - not least when capturing prey or avoiding predators.[3] Indeed, it makes cephalopods the fastest marine invertebrates,[5]:Preface and they can out-accelerate most fish.[21] 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.[25] The velocity of the organism can be accurately predicted for a given mass and morhpology of animal.[26] 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.[27] Some cephalopods accompany this expulsion of water with a gunshot-like popping noise, thought to function to frighten away potential predators.[28]
Early cephalopods are thought to have produced jets by drawing their body into their shells, as Nautilus does today.[29] 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.[29] The jet velocity in Nautilus is much slower than in coleoids, but less musculature and energy is involved in its production.[30] Jet thrust in cephalopods is controlled primarily by the maximum diameter of the funnel orifice (or, perhaps, the average diameter of the funnel)[31]:440 and the diameter of the mantle cavity.[32] Changes in the size of the orifice are used most at intermediate velocities.[31] The absolute velocity achieved is limited by the cephalopod's requirement to inhale water for expulsion; this intake limits the maximum velocity to eight body-lengths per second, a speed which most cephalopods can attain after two funnel-blows.[31] Water refills the cavity by entering not only through the orifices, but also though the funnel.[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.[31]
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; in fact most cephalopods are about 2-3% denser than seawater[6]), they achieve this in different ways.[24] 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;[24] others, like some fish, accumulate oils in the liver;[24] and some octopuses have a gelatinous body with lighter chlorine ions replacing sulfate in the body chemistry.[24]
Shell
See also: Cuttlebone
Cuttlebone of Sepia officinalis
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, for example, an invagination of the ectoderm forms during the embryonic period, resulting in a shell that is internal in the adult.[33] The same is true of the chitinous gladius of squid[33] and octopus.[34] Spirula, and cirrate octopuses also have small internal "shells", although the most common octopus suborder, the Incirrina, have no vestige of an internal shell,[citation needed] and some squid also lack a gladius.[35]
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.
A giant squid found in Logy Bay, Newfoundland, in 1873. The two long feeding tentacles are visible on the extreme left and right.
Head appendages
Main articles: Cephalopod arm and tentacle
Cuttlefish and squid have five pairs of muscular appendages surrounding their mouths. The longer two, termed tentacles, are actively involved in capturing prey;[36]:225 they can lengthen rapidly (in as little as 15 milliseconds[36]:225). In giant squid they may reach a length of 8 metres. They may terminate by broadening into a sucker-coated club.[36]:225 The shorter four pairs are termed arms, and are involved in holding and manipulating the captured organism.[36]:225 They too have suckers, on the side closest to the mouth; these help to hold onto the prey.[36]:226
The tentacle consists of a thick central nerve cord (which must be thick to allow each sucker to be controlled independently)[37] 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%.[36]:227
The size of the tentacle is related to the size of the buccal cavity; larger, stronger tentacles can hold prey as small bites are taken from it; with more numerous, smaller tentacles, prey is swallowed whole, so the mouth cavity must be larger.[38]
Feeding
All cephalopods have a two-part beak;[5]:7 most have a radula, although it is reduced in most octopus and absent altogether in Spirula.[5]:7[39]:110 They feed by capturing prey with their tentacles, drawing it in to their mouth and taking bites from it.[8] 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.[8] The salivary gland has a small tooth at its end which can be poked into an organism to digest it from within.[8]
The digestive gland itself is rather short.[8] 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.[8] 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.[8]
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.[40]
Nautilus, unusually, possesses four nephridia, none of which are connected to the pericardial cavities.
Reproduction and life cycle
Female Argonauta argo with eggcase and eggs
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. They tend towards a semelparous reproduction strategy; they lay many small eggs in one batch and die afterwards. The Nautiloidea, on the other hand, stick to iteroparity; they produce a few large eggs in each batch and live for a long time.
External sexual characteristics are lacking in cephalopods. So cephalopods use colour communication. A courting male will approach a likely looking opposite number flashing his brightest colours, often in rippling displays. If the other cephalopod is female and receptive, her skin will change colour to become pale, and mating will occur. If the other cephalopod remains brightly coloured, it is taken as a warning[41].
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[41]:227.
The eggs may be brooded: female paper nautilus construct a shelter for the young, while Gonatiid squid carry a larva-laden membrane from the hooks on their arms.[42] Other cephalopods deposit their young under rocks and aerate them with their tentacles hatching. Often, though, the eggs are left to their own devices; many squid lay sausage-like bunches of eggs in crevices or occasionally on the sea floor. Cuttlefish lay their eggs separately in cases and attach them to coral or algal fronds.[43] Fossilised egg clutches show that ammonites also laid clutches of eggs.[44]
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.[8] Most live for one to two years,[8] reproducing and then dying shortly thereafter.[45]
In order to free up resources for reproduction, many squid are known to resorb the muscle tissue of their mantle and tentacles, breaking down the tissue and using the energy contained therein to produce more gametes.[46]
Egg cases laid by a female squid
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.[40][47]
The funnel of cephalopods develops on the top of their head, whereas the mouth develops on the opposite surface.[48]:86 The early embryological stages are reminiscent of ancestral gastropods and extant Monoplacophora.[47]
The shells develop from the ectoderm as an organic framework which is subsequently mineralised.[33] In Sepia, which has an internal shell, the ectoderm forms an invagination whose pore is sealed off before this organic framework is deposited.[33]
engrailed is expressed first in the arms, funnel and optic vesicles, and is only later present in the tentacles and eyelids.[33] It is expressed in embryonic stages 17–19 in all arm buds, and subsequently in the future-tentacles in stages 24–5, suggesting that it may serve a role in the differential development of tentacles. Sequential expression of Hox genes is also observed in cephalopod arms.[33]
Development
Left: Immature specimens of Chiroteuthis veranyi. In this paralarval form, known as the doratopsis stage, the pen is longer than the mantle and 'neck' combined
Right: A mature Chiroteuthis veranyi. This species has some of the longest tentacles in proportion to its size of any known cephalopod.
Cephalopod eggs span a large range of sizes, from 1 to 30 mm in diameter.[49] 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.[49]
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.[49]
Young do not pass through a larval stage sensu stricto. They quickly learn how to hunt, using encounters with prey to refine their strategies.[49]
Growth in juveniles is usually allometric, whilst adult growth is isometric.[50]
Evolution
The class developed during the Late Cambrian, and underwent pulses of diversification during the Ordovician period[51] to become diverse and dominant in the Paleozoic and Mesozoic seas. Small shelly fossils such as Tommotia were once interpreted as early cephalopods, but today these tiny fossils are recognized as sclerites of larger animals,[52] and the earliest accepted cephalopods date to the Late Cambrian Period.[note 1][53] During the Cambrian, cephalopods are most common in shallow near-shore environments, but they have been found in deeper waters too.[54] Cephalopods were thought to have "undoubtedly" arisen from within the tryblidiid monoplacophoran clade.[55] However genetic studies suggest that they are more basal, forming a sister group to the scaphopoda but otherwise basal to all other major mollusc classes.[56] The internal phylogeny of Mollusca, however, is wide open to interpretation - see Mollusca#Phylogeny.
Fossil orthoconic nautiloid from the Ordovician of Kentucky; an internal mold showing siphuncle and half-filled camerae, both encrusted.
The cephalopods are thought to have evolved from a monoplacophoran-like ancestor[57] with a curved, tapering shell,[58] and to be closely related to the gastropods (snails).[59] The development of a siphuncle allowed their shells to become gas-filled (thus buoyant) in order to support them and keep the shells upright while the animal crawled along the floor, and separates the true cephalopods from putative ancestors such as Knightoconus, which lacked a siphuncle.[59] Negative buoyancy (i.e. the ability to float) came later, followed by swimming in the Plectronocerida and eventually jet propulsion in more derived cephalopods.[60] However, because chambered shells are found in a range of molluscs - monoplacophora and gastropods as well as cephalopods - a siphuncle is essential to ally a fossil shell conclusively to the cephalopoda.[58]:57 The earliest such shells do not have the muscle scars which would be expected if they truly had a monoplacophoran affinity.[58]:57
The earliest cephalopod order to emerge was the Ellesmerocerida, which were quite small organisms; their shells were slightly curved, and the internal chambers were closely spaced. The siphuncle penetrated the septa with meniscus-like holes.[51] Early cephalopods had fine shells which could not cope with the pressures of deep water.[51] In the mid Tremadoc, these were supplemented by larger shells around 20 cm in length; these larger forms included straight and coiled shells, and fall into the orders Endocerida (with wide siphuncles) and Tarphycerida (with narrow siphuncles).[51] By the mid Ordovician these orders are joined by the Orthocerids, whose first chambers are small and spherical, and Lituitids, whose siphuncles are thin. The Oncocerids also appear during this time; they are restricted to shallow water and have short conchs which surround the stomach.[51] The mid Ordovician saw the first cephalopods with septa strong enough to cope with the pressures associated with deeper water, and could inhabit depths greater than 100–200 m.[51] The wide-siphuncled Actinocerida and the Discocerida both emerged during the Darriwilian.[51] The direction of 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.[61] (Endogastric mean the shell is curved so as the ventral or lower side is longitudinally concave (belly in); exogastric means the shell is curve so as the ventral side is longitudinally convex (belly out) allowing the funnel to be pointed backwards beneath the shell.)[61]
Curved shells brought a number of benefits. Firstly, minerals are not required in as large quantities, as each successive whorl builds on the one before. Also, the organism is more stable (its centre of mass coincides with its centre of buoyancy) and more manoeuvrable.[29]
Early cephalopods were likely predators near the top of the food chain.[8] In the Early Palaeozoic, their range was far more restricted than today; they were mainly constrained to sub-littoral regions of shallow shelves of the low latitudes, and usually occur in association with thrombolites.[62] A more pelagic habit was gradually adopted as the Ordovician progressed.[62] Deep-water cephalopods, whilst rare, have been found in the Lower Ordovician - but only in high-latitude waters.[62]
An ammonitic ammonoid with the body chamber missing, showing the septal surface (especially at right) with its undulating lobes and saddles.
The ancestors of coleoids (including most modern cephalopods) and the ancestors of the modern nautilus, had diverged by the Floian Age of the Early Ordovician Period, over 470 million years ago. We know this because the orthocerids were the first known representatives of the neocephalopoda,[63] were ultimately the ancestors of ammonoids and coleoids, and had appeared by the Floian.[62] It is widely held that the Bactritida, an Silurian–Triassic group of orthocones, are paraphyletic to the coleoids and ammonoids – that is, the latter groups arose from within the Bactritida.[64]:393 An increase in the diveristy 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.[64]
Unlike most modern cephalopods, most ancient varieties had protective shells. These shells at first were conical but later developed into curved nautiloid shapes seen in modern nautilus species. It is thought that competitive pressure from fish forced the shelled forms into deeper water, which provided an evolutionary pressure towards shell loss and gave rise to the modern coleoids, a change which led to greater metabolic costs associated with the loss of buoyancy, but which allowed them to recolonise shallow waters.[59]:36 However, some of the straight-shelled nautiloids evolved into belemnites, out of which some evolved into squid and cuttlefish.[verification needed] The loss of the shell may also have resulted from evolutionary pressure to increase manoeuvrability, resulting in a more fish-like habit.[36]:289 This pressure may have increased as a result of the increased complexity of fish in the late Palaeozoic, increasing the competitive pressure.[36]:289 Internal shells still exist in many non-shelled living cephalopod groups but most truly shelled cephalopods, such as the ammonites, became extinct at the end of the Cretaceous.
The tentacles of the ancestral cephalopod developed from the mollusc's foot;[47] the ancestral state is thought to have had five pairs of tentacles which surround the mouth.[47] Smell-detecting organs evolved very early in the cephalopod lineage.[47]
The earliest cephalopods[note 2], like Nautilus and some coeloids, appeared to be able to propel themselves forwards by directing their jet backwards.[36]:289 Because they had an external shell, they would not have been able to generate their jets by contracting their mantle, so must have used alternate methods: such as by contracting their funnels or moving the head in and out of the chamber.[36]:289
The preservation of cephalopod soft parts is not entirely unusual; soft-bodied fossils, especially of coeloids (squid), are relatively widespread in the Jurassic,[46] but phosphatized remains are unknown before this period.[65] On the other hand, soft parts – including a possible ink sac — are known from the Paleozoic Hunsruck shale and Francis Creek shale.[66]
Phylogeny
The internal phylogeny of the cephalopods is difficult to constrain; many molecular techniques have been adopted, but the results produced are conflicting.[67] 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.[67]
Molecular estimates for clade divergence vary. One 'statistically robust' estimate has Nautilus diverging from Octopus at 415 ± 24 million years ago.[68]
Taxonomy
Chambered Nautilus (Nautilus pompilius)
Common Cuttlefish (Sepia officinalis)
Atlantic Bobtail (Sepiola atlantica)
European Squid (Loligo vulgaris)
Common Octopus (Octopus vulgaris)
The classification presented here, for recent cephalopods, follows largely from Current Classification of Recent Cephalopoda (May 2001), for fossil cephalopods takes from Arkell et al. 1957, Teichert and Moore 1964, Teichert 1988, and others. The three subclasses are traditional, corresponding to the three orders of cephalopods recognized by Bather.[69]
Class Cephalopoda († indicates extinct groups)
· Subclass Nautiloidea: Fundamental ectocochliate cephalopods that provided the source for the Ammonoidea and Coleoidea.
o Order † Plectronocerida: the ancestral cephalopods from the Cambrian Period
o Order † Ellesmerocerida (500.0—470.0 Ma)
o Order † Endocerida (485.0—430.0 Ma)
o Order † Actinocerida (480.0—312.0 Ma)
o Order † Discosorida (482.0-392.0 Ma)
o Order † Pseudorthocerida (432.0-272.0 Ma)
o Order † Tarphycerida (485.0-386.0 Ma)
o Order † Oncocerida (478.5-324.0 Ma)
o Order Nautilida (extant;410.5 Ma-Rec)
o Order † Orthocerida (482.5-211.5 Ma)
o Order † Ascocerida (478.0-412.0 Ma)
o Order † Bactritida (418.1—260.5 Ma)
· Subclass † Ammonoidea: Ammonites (479.0—65.0 Ma)
o Order † Goniatitida (388.5- 252.0 Ma)
o Order † Ceratitida (254.0-200.0 Ma)
o Order † Ammonitida (215.0-66.0 Ma)
· Subclass Coleoidea (410.0 Ma-Rec)
o Cohort † Belemnoidea: Belemnites and kin
§ Genus † Jeletzkya
§ Order † Aulacocerida (265.0—183.0 Ma)
§ Order † Phragmoteuthida (189.6—183.0 Ma)
§ Order † Hematitida (339.4—318.1 Ma)
§ Order † Belemnitida (339.4—65.5 Ma)
§ Genus † Belemnoteuthis (189.6—183.0 Ma)
o Cohort Neocoleoidea
§ Superorder Decapodiformes (also known as Decabrachia or Decembranchiata)
§ (?Order Boletzkyida)
§ Order Spirulida: Ram's Horn Squid
§ Order Sepiida: cuttlefish
§ Order Sepiolida: pygmy, bobtail and bottletail squid
§ Order Teuthida: squid
§ Superorder Octopodiformes (also known as Vampyropoda)
§ 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.
Suprafamilial Classifiction of the Treatise
This is the older classification that combines those found in parts K and L of the Treatise on Invertebrate Paleontology, which forms the basis for and is retained in large part by classifications that have come later.
Nautiloids in general, (Teichert and Moore 1964) Sequence as given.
Subclass † Endoceratoidea. Not used by Flower, e.g. Flower and Kummel 1950, interjocerids included in the Endocerida.
Order † Endocerida
Order † Intejocerida
Subclass † Actinoceratoidea Not used by Flower, ibid
Order † Actinocerida
Subclass † Nautiloidea Nautiloidea in the restricted sense.
Order † Ellesmerocerida Plectronocerida subsequently split off as separate order.
Order † Orthocerida Includes orthocerids and pseudorthocerids
Order † Ascocerida
Order † Oncocerida
Order † Discosorida
Order † Tarphycerida
Order † Barrandeocerida A polyphyletic group now included in the Tarphycerida
Order Nautilida
Subclass † Bactritoidea
Order † Bactritida
Paleozoic Ammonoidea ( Miller, Furnish, and Schindewolf, 1957)
Suborder † Anarcestina
Suborder † Clymeniina
Suborder † Goniatitina
Suborder † Prolecanitina
Mesozoic Ammonoidea (Arkel et al., 1957)
Suborder † Ceratitina
Suborder † Phylloceratina
Suborder † Lytoceratina
Suborder † Ammonitina
Subsequent revisions include the establishment of three Upper Cambrian orders, the Plectronocerida, Protactinocerida and Yanhecerida; separation of the pseudorthocerids as the Pseudorthocerida, and elevating orthoceritoids as the Subclass Orthoceratoidea.
Shevyrev classification
Shevyrev (2005) suggested a division into eight subclasses, mostly comprising the more diverse and numerous fossil forms.[70] [71]
Holotype of Ostenoteuthis siroi from family Ostenoteuthidae.
Class Cephalopoda
· Subclass † Ellesmeroceratoidea
o Order † Plectronocerida (501.0-490.0 Ma)
o Order † Protactinocerida
o Order † Yanhecerida
o Order † Ellesmerocerida (500.0-470.0 Ma)
· Subclass † Endoceratoidea (485.0-430.0 Ma)
o Order † Endocerida (485.0-430.0 Ma)
o Order † Intejocerida (485.0-480.0 Ma)
· Subclass † Actinoceratoidea
o Order † Actinocerida (480.0—312.0 Ma)
· Subclass Nautiloidea (490.0 Ma- Rec)
o Order † Basslerocerida (490.0-480.0 Ma)
o Order † Tarphycerida (485.0-386.0 Ma)
o Order † Lituitida (485.0—480.0 Ma)
o Order † Discosorida (482.0-392.0 Ma)
o Order † Oncocerida (478.5-324.0 Ma)
o Order Nautilida (410.5 Ma-Rec)
· Subclass † Orthoceratoidea (482.5-211.5 Ma)
o Order † Orthocerida (482.5-211.5 Ma)
o Order † Ascocerida (478.0-412.0 Ma)
o Order † Dissidocerida (479.0—457.5 Ma)
o Order † Bajkalocerida
· Subclass † Bactritoidea (422.0-252.0 Ma)
· Subclass † Ammonoidea (410.0-66.0 Ma)
· Subclass Coleoidea (410.0 Ma-rec)[72]
Cladistic classification
Pyritized fossil of Vampyronassa rhodanica, a vampyromorphid from the Lower Callovian (164.7 million years ago)
Another recent system divides all cephalopods into two clades. One includes nautilus and most fossil nautiloids. The other clade (Neocephalopoda or Angusteradulata) is closer to modern coleoids, and includes belemnoids, ammonoids, and many orthocerid families. There are also stem group cephalopods of the traditional Ellesmerocerida that belong to neither clade.[73][74]
Monophyly of coeloids
The coeloids have been thought to possibly represent a polyphyletic group,[36]:289 although this has not been supported by the rising body of molecular data.[75]
Post-mortem decay
After death, if undisturbed, cephalopods[note 3] decay relatively quickly.[46] Their muscle softens within a couple of days, and may swell; egg sacs can swell so much that they rip through the mantle.[46] 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.[46] After around a week, the carcass collapses in on itself and begins to disintegrate. The ink sac solidifies around this point.[46] After a fortnight little is left but a blob with eyes, arms and ink sac visible.[46] 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.[46]
See also
Books are collections of articles which can be downloaded or ordered in print. | |
· Kraken
Notes
1. ^ The genus Plectronoceras
2. ^ Ordovician orthocone nautiloids are the first for which trace fossil evidence is available
3. ^ Experiments were performed on a variety of squid
References
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Further reading
· Barskov, I.S., M.S. Boiko, V.A. Konovalova, T.B. Leonova & S.V. Nikolaeva (2008). "Cephalopods in the marine ecosystems of the Paleozoic". Paleontological Journal 42: 1167–1284. doi:10.1134/S0031030108110014. A comprehensive overview of Paleozoic cephalopods.
· Campbell, Neil A., Reece, Jane B., and Mitchell, Lawrence G.: Biology, fifth edition. Addison Wesley Longman, Inc. Menlo Park, California. 1999. ISBN 0-8053-6566-4.
· Felley, J., Vecchione, M., Roper, C. F. E., Sweeney, M. & Christensen, T., 2001-2003: Current Classification of Recent Cephalopoda. Internet: National Museum of Natural History: Department of Systematic Biology: Invertebrate Zoology: http://www.mnh.si.edu/cephs/
External links
· CephBase - cephalopod database
· TONMO.COM - The Octopus News Magazine Online - cephalopod articles and discussion
· Tree of Life Web Project - Cephalopoda
· Will Fast Growing Squid Replace Slow Growing Fish?
· Biomineralisation in modern and fossil cephalopods
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