class of mollusks

The Cephalopoda (Greek meaning "head-foot") is an important mollusc class. They have bilateral symmetry, a head, and arms or tentacles.[1] Teuthology, a branch of malacology, is the study of cephalopods.

Juvenile squid from plankton
Scientific classification


Subclass Nautiloidea
Subclass †Ammonoidea
Subclass Coleoidea

The class has two living subclasses. In the Coleoidea, the mollusc shell has become smaller, or is not there at all; this subclass has the octopus, squid, and cuttlefish. The Nautiloidea have a shell; Nautilus is its only living genus.

There are at least 800 different living species of Cephalopods. Two important extinct taxa are the ammonites, and the belemnites (order Belemnoidea, of class Coleoidea). Cephalopods are found in all the oceans of the world and at all pelagic levels. None of them can live in freshwater (water with no salt in it), but a few species live in brackish (partly salty) water.

Cephalopod groups


Number of species


There are still discoveries of new species of cephalopods:

  • 1998: 703 recent species [2]
  • 2001: 786 recent species [3]
  • 2004: estimate: from 1000 to 1200 species [4]

There are many more fossil species. It is thought there are around 11,000 extinct taxa.[5]

Nervous system and behaviour


Cephalopods are the most intelligent invertebrates and have good senses and large brains. The nervous system of cephalopods is the most complex of the invertebrates, and their brain to body mass ratio falls between that of warm and cold blooded vertebrates.[6] The giant nerve fibers of the cephalopod mantle have been a favourite experimental material for many years; their large diameter makes them easier to study.

Colour and light

This Broadclub Cuttlefish (Sepia latimanus) can go from blending tans and browns (top) to yellow with dark bits (bottom) in less than a second.

Most cephalopods have chromatophores – that is, cells with different colours – which they can use in a number of surprising ways.[6] As well as blending with their background, some cephalopods bioluminesce, shining light downwards to hide their shadows from any attackers.[6] The bioluminescence is made by bacterial symbionts; the host cephalopod is able to find the light made by these animals.[7] Bioluminescence may also be used to attract prey, and some species use colourful shows to get mates, amaze predators, or even signal to one another.[6]



Colouration can be changed in milliseconds as they adjust to their environment,[6] and the pigment cells can expand or contract.[8] Rapid colour change is usually more common in near-shore species than those living in the open ocean. Those in open ocean mostly use camouflage to make their body outline less easy to see.[6]

Evidence of original colouration has been found in cephalopod fossils as far back as the Silurian; certain straight-shelled species had lines round their shell, which are thought to have used as camouflage of their body outline.[9] Devonian cephalopods bear more complex colour patterns, whose function may be more complex.[10]

Moving around


Cephalopods usually move by jet propulsion (squirting water). This uses a lot of energy to travel compared to the tail propulsion used by fish. They use jet propulsion because they do not have fins or flippers. The efficiency of jet propulsion goes down with larger animals. This is probably the reason why many species use their fins or arms for moving if possible.

Oxygenated water is taken into the mantle cavity to the gills. By contracting the mantle's muscles, the water is pushed out through the siphon, made by a fold in the mantle. Motion of the cephalopods is usually backward as water is forced out forwards, but the siphon can be pointed in different directions. Some cephalopods can adjust their body shape to move through the water more easily.

Some octopus species are also able to walk along the sea bed. Squids and cuttlefish can move short distances in any direction by moving a flap of muscle around the mantle.

With the exception of the Nautilidae and the species of octopus of the suborder Cirrina,[11] all known cephalopods have an ink sac, which can be used to push out a cloud of dark ink to confuse predators.[12]

Like most molluscs, cephalopods use haemocyanin, a copper-containing protein, rather than haemoglobin to transport oxygen. As a result, their blood is colourless when deoxygenated and turns blue when placed in air.[13]

Reproduction and life cycle


With a few exceptions, Coleoideans live short lives with fast growth. Most of the energy from their food is used for growing. The penis in most male Coleoidea is a long and muscular end of the vas deferens (tube for sperms) used to transfer spermatophores (sperm packages) to a modified arm called a hectocotylus.[14] That in turn is used to transport 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. The females lay many small eggs in one batch, and die afterwards. The Nautiloidea, on the other hand make a few large eggs in each batch, and live for a long time.



The class developed during the late Cambrian and were the most common and varied marine life forms during the Palaeozoic and Mesozoic eras. Tommotia, an early cephalopod, had squid-like tentacles but also a snail-like foot it used to move across the sea floor. Early cephalopods were at the top of the food chain.

The old (cohort Belemnoidea) and modern (cohort Neocoleoidea) coleoids, as well as the ammonoids, all diverged (evolved away) from the external shelled nautiloids during the middle Paleozoic Era, between 450 and 300 million years ago. Most ancient varieties had protective shells. These shells at first were conical but later developed into the curved shapes seen in modern nautilus species. Shells inside the body still exist in many living cephalopod groups, such as cuttlefish. The most famous group with external shells, the ammonites, became extinct at the end of the Cretaceous period.


  1. These are adaptations of the basic mollusc foot.
  2. [updated 28-Nov-2000] [cit. 12-Dec-2003] http://www.cephbase.dal.ca/spdb/allsp.cfm Archived 2003-12-11 at the Wayback Machine
  3. [updated 13-Jun-2003] [cit. 27-Feb-2005] http://www.cephbase.utmb.edu/spdb/allsp.cfm Archived 2005-12-18 at the Wayback Machine
  4. Brune R.H. 2004. Encyklopedie ulit a lastur. Rebo Productions, Dobřejovice. p16 (in Czech)
  5. Ivanov M., Hrdličková S. & Gregorová R. 2001. Encyklopedie zkamenělin. Rebo Productions, Dobřejovice. p139. (in Czech)
  6. 6.0 6.1 6.2 6.3 6.4 6.5 Marion Nixon & J.Z. Young 2003. The brains and lives of cephalopods. New York: Oxford University Press. ISBN 0-19-852761-6.
  7. Tong D. Rozas S.; Oakley H.; Mitchell J.; Colley J.; Mcfall-Ngai J. 2009. Evidence for light perception in a bioluminescent organ. PNAS 106: 9836–9841.
  8. Encyclopædia Britannica 2006 Ultimate Reference Suite DVD
  9. Štěpán Manda and Vojtěch Turek 2009. Acta Palaeontologica Polonica 54: 503-512. "Minute Silurian oncocerid nautiloids with unusual colour patterns". {{cite journal}}: Cite journal requires |journal= (help)CS1 maint: numeric names: authors list (link)
  10. "Colour patterns in Early Devonian cephalopods from the Barrandian Area: Taphonomy and taxonomy. Vojtěch Turek 2009. Acta Palaeontologica Polonica 54: 491-502". {{cite journal}}: Cite journal requires |journal= (help)
  11. Hanlon, Roger T. & Messenger John B. 1999. Cephalopod behaviour. Cambridge. ISBN 0521645832
  12. Boyle, Peter; Rodhouse, Paul (2004). Cephalopods: ecology and fisheries. Ames, Iowa: Blackwell. doi:10.1002/9780470995310.ch2. ISBN 0632060484.
  13. Ghiretti-Magaldi A.; Ghiretti F. (1992). "The pre-history of hemocyanin: the discovery of copper in the blood of molluscs". Cellular and Molecular Life Sciences. 48 (10). Birkhäuser Basel: 971–972. doi:10.1007/BF01919143. S2CID 33290596.[permanent dead link]
  14. Gilbert L. Voss (27 September 2013). "cephalopod". Encyclopædia Britannica Online. Encyclopædia Britannica, Inc.

Other sources

  • Berthold, Thomas, & Engeser, Theo. 1987. Phylogenetic analysis and systematization of the Cephalopoda (Mollusca). Verhandlungen Naturwissenschaftlichen Vereins in Hamburg. (NF) 29: 187-220.
  • Engeser, Theo. 1997. Fossil Nautiloidea page. <http://userpage.fu-berlin.de/~palaeont/fossilnautiloidea/fossnautcontent.htm>
  • 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/
  • 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 molluscs. Paleontological Journal. 39: 606-614. Translated from Paleontologicheskii Zhurnal #6, 2005, 33-42.

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