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Cephalopods (Class Cephalopoda)are exclusively marine mollusks (Phyllum ) with well developed prehensile grasping appediges the encircle the mouth and protrude from a well defined head, and which have a well developed nervous system. All are carnivorous. Other mollusks include , snails, , clams and other bivalves, and , tusk shells.

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The name Cephalopoda literally means "head feet" which refers to the cluster of and/or that project forward from the head, surrounding the mouth. The group includes living (squid, octopods, and cuttlefish) and , and a large number of ancient (mostly and ) forms. All are active marine predators (although some early types were drifters), able to swim swiftly, and easily competing with fish in the marine habitat. There are 650 living species, but more than 7,500 fossil forms are known (and as in all cases like this this number is obviously a gross underestimate of the real number of cephalopod species that have ever lived through the time). Like fish they are equipped with highly developed eyes and other sense organs, include both active swimmers and bottom-dwellers, and in many cases have a streamlined body for more efficient locomotion. Swimming is by rapidly expelling water from the . The water is forced out through a funnel or - the - actually a tube-like flap of modified foot, thus driving the animal in the opposite direction. This is the key to the so-called "jet-propulsion" of these animals The funnel is highly maneuverable and can be directed in any direction, allowing motion backwards or forwards. However, the fastest movement is backward escape swimming, with powerful contractions of the mantle ejecting water through the forward facing funnel. A cloud of "ink" can also be ejected as a sort of underwater smoke screen to hide the fleeing animal.

All cephalopods are carnivorous, feeding primarily on fish, other , , and worms. The head projects into a crown of prehensile tentacles - ranging from 8 in the octopus to 80 or 90 in the living nautilus. These tentacles are actually a specialized form of the standard molluscan foot, and used for grasping prey. Once the prey is snared it is bitten with strong beak-like jaws and pulled into the mouth by the .

Cephalopods are quite large by molluscan standards (most species being between 6 and 70 cm including tentacles), with the giants of the group - such as the modern day , the giant squid, with a body length (including tentacles) of up to 20 meters, the Ordovician , with a straight shell up to 10 metres in length, and the Cretaceous , with a coiled shell 3 metres in diameter - the largest invertebrates ever to live, with weights of one to two tons. Such giant cephalopods play or played a similar ecological role of top predator to that of , and .

Cephalopods have a highly developed nervous system, unequalled among the invertebrates, and correlated with locomotor dexterity and carnivorous lifestyle (predators generally always have larger brains than prey animals). There is a high level of cephalization (development and concentration of sensory and neural centers in the head). The nerve ganglia are concentrated and more or less fused to form a brain that encircles the esophagus. A bundle of giant nerve fibres tied to the give them very rapid reflexes. They are visual creatures, changing colour to express mood. The eyes of the are very elaborate, with a retinal structure remarkably like that found in vertebrates. The eye of the giant squid is the largest of any animal - 40 cm across. Nautiloids have smaller and more primitive eyes.

As with vertebrates, the brain is partitioned into different areas that control particular functions. For example, the brain centres for both forward swimming and closing of the suckers are located in the cerebral ganglia. These animals are most intelligent and highly evolved of the mollusks, indeed they are the most intelligent of all the invertebrates, exhibiting complex patterns of behavior. Octopods can easily be trained to distinguish between classes of objects. See

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Basic Divisions

  • Phyllum Mollusca
    • Class Cephalopoda
      • Subclass
      • Subclass
      • Subclass

Most classification divide the Cephalopoda in the three subclasses as shown. Nautiloids are also known as Palcephalopoda, a term which is applied to most but not all members of the subclass. Ammonoids and Coleoids and a few nautiloides come under the term Neocephalopoda. Both terms are based on the morphology and biology of living cephalopods, Palcephalopoda for the presumed primitive Nautilus and Allonautilus, representing the nautilids and Neocephalopoda for the derived and presumed advanced coleoids.

Systematics

The class Cephalopoda has been traditionally divided into three subclasses on the basis of shell structure, or two subclasses on the basis of gills and other soft parts. Both shown here:

>Shell based classification, used exclusively in paleontology, with three basic subclasses.

Nautiloidea -- includes primitive forms, all have an external shell and a retrosiphonate siphuncle; represented by modern, tetrabranchiate Nautilus and Allonautilus.

Ammonoidea -- evolved forms with an external shell and typically a prosiphonate siphuncle; derived from the Nautilioidea but closely related biologically to the dibranchiate coleoids. Became extinct at the end of the Cretaceous.

Coleoidea -- shell-less forms and those with an internal vestigial shell represented by living octopods, cuttlefish, squid,and Spirula, all dibranchite. Includes belemnites and other extinct forms.

>>Anatomical (gill) based classification, used in cephalopod biology.

Tetrabranchiata-- forms with four gills (2 pairs), represented by Nautilus, thought to be primitive. The Palcephalopoda.

Dibranchiata-- forms with two gills, considered advanced. Equivalent to the Coleoidea, possibly the Ammonoidea and may be even some Nautilodea

The trouble with the anatomical scheme is that it is impossible to know from shell structure alone alone whether extinct forms had two or four gills, or anything really much about their soft body anatomy. From this it is apparent that the two-fold anatomical classification is inadequate except in modern forms.

The Subclass Nautiloidea forms a large and diverse, although paraphyletic, group, divisible into phyllogenetically defined superorders. All nautiloids are united by the fact they have simple, adorally concave septa, retrosiphonate septal necks, sutures that are never elaborate, and are descended from a common ancestor, Plectronoceras They are more than simply cephalopods that are neither ammonoids or coleoids.

While there is general agreement regarding classification of recent cephalopods, mostly coleoids, based on soft part anatomical features, classification of extinct fossil forms based on such criteria is impossible other than for a few inferences that night be drawn from preserved muscle attachment scars, preserved radula, tentacle imprints, and isolated body impressions. Shell based features on the other hand provide the basis for a reliable, phylogenetically plausible and generally accepted classification of extinct forms.

Palcephalopoda-Neocephalopoda

The Palcephalopoda-Neocephalopoda distinction is based on essential differences between the living nautiloid genera, Nautilus and Allonautilus, and the huge variety of living coleoids. Living nautiloides are tetrabranchiate (have 4 gills in 2 pairs), are lateradulate (13 elements per radular row), and produce multiple batches of well develovoped offspring from large yolk-rich eggs. The have an external shell, retained from their ancestors but have mutliple finger-like protractable tentacles, probably a derived character. Also their eyes have no lens but operate like a pin-hole camera, again apparently primitive, Coleoids are dibranchiate (have 2 gills), are angusteradulate (9 elements per radular row) and in general produce a single batch of numerous planctonic offspring before dying. They lack an external shell, a obvious derived character but have 8 or 10 muscular suckered and/or hooked tentacles which seems to be a retained primitive trait. On the other hand coleoids have well developed eyes with lenses, sometimes covered by a corneal membrane.

The extent to which these distinctions can be extrapolated into fossil forms is questionable and subject to individual interpretation. Ammonoids for example are thought to be more closely related to coleoids than to nautiloids on the basis of being angueradulate (with 9 radular elements like coleoids) from the few radula found, in spite of having an external shell, and are therefor considered neocephalopods. Paleozoic orthocerids are considered neocephalopods because their protoconch is like that of ammonoids in spite of having an obviously nautiloid phragmocone.

Alternative Taxonomies

An Infraclass based taxonomy

The following basically follows Teichert (1988) except that Teichert's two Subclasses Endoceratoidea and Actinoceratoidea have been discarded, since the two orders and are not so distinct from their contemporaries as to justify such a high taxonomic ranking.

Infraclass unnamed ("Ellesmeroceroidea"? or several infraclasses?)

Order Flower, 1964.
Order Chen & Qi, 1979.(or included in Plectronocerida?)
Order Chen & Qi, 1979.(or included in Plectronocerida?)
Order Flower, 1950.
Order Teichert, 1933.
Order Balashov, 1960. (or included in Endocerida?)
Order Flower, 1950
Order Teichert, 1933
Order

Infraclass Nautiloidea Agassiz, 1947
(Superorder Nautilitoidea, Wade)

Order Flower, 1950.
Suborder Flower, 1950.
Suborder Flower, 1950.
Order Flower, 1950.
Order Agassiz, 1847.

Infraclass Orthoceratoidea Kuhn, 1940 (Orthoceratoidea Wade, in part)

Order Kuhn, 1940.
Order Kuhn, 1949.

Infraclass Ammonoidea Agassiz, 1947
(=Subclass )

Order Shimanskiy
Order Miller & Furnish 1954
Order Hyatt 1884
Order Wedekind 1927
Order Miller & Furnish 1954
Order Hyatt 1884
Order Kuhn, 1940
Order Hyatt 1889
Order Agassiz 1847
Order Wedman 1966

Infraclass Coleoidea Bather, 1888
(=Subclass )

Order Wedman 1966
Order Bandel, Reitner & Stürmer 1983
Order Zittel 1885
Order Stolley 1919
Order Jeletzky 1964
Order Naef 1916
Order Naef 1916
Order Leach 1818
Order Grimpe 1917 (or in Octopodida)


Superordinal Classification from Wade, 1988.

>Group (Subclass) Nautiloidea

Superorder

Order:
Order: (or in Plectronocerida)
Order: (or in Plectronocerida)
Order: .

Superorder Endoceratoidea, = Order:

Superorder Actinoceratoidea, = Order:

Superorder Discosoroidea, = Order:

Superorder

Order:
Order:
Order:

Superorder

Order:
Order:
Order:

>Group (subclass Dibranchia)

Superorder

Order:
Order:
Order:

Superorder (= subclass Ammonoidea, see for content)

Superorder (= subclass Coleoidea)

Order:
Order:
Order:
Order:
Order:


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Teichert,Curt 1988. "Main Features of Cephalopod Evolution", pp.19-20, 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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© M. Alan Kazlev 1998-2002 with revisions by John M 2010-2011.

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