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Incirrata Grimpe, 1916

Common octopods, octopuses or devilfishes

Katharina M. Mangold (1922-2003) and Richard E. Young
The suborder contains four families and one family-group with four families.
Containing group: Octopoda


Incirrate octopods are about 85% of all octopodan species. They are very small to large in size. The members of the Octopodidae, the family with the most species and individuals, are benthic. The other seven incirrate families have pelagic species. The benthic incirrate octopods live from the intertidal zone to at least 4000 m, and bathypelagic forms may be found down to 2000 m. The octopodids exhibit sophisticated behavior. The brain of benthic octopuses, especially that of Octopus vulgaris, has become a model for relating brain structure to function (e.g., J. Z. Young, 1971). Incirrate octopods are commercially fished throughout the world.

Fins are absent and the arms bear one or two rows of suckers and lack cirri. The shape of the body is saccular and rather broad. The arms are often of about equal length, but sometimes the dorsal or lateral or ventral arms are distinctly longer than the others.


  1. Arms
    1. One member of Arms III hectocotylized.
    2. Cirri absent.

  2. Eyes
    1. Cornea present (greatly reduced in some pelagic species).

  3. Fins
    1. Absent.

  4. Gills
    1. Branchial canal present
    2. Asymmetrical in cross-section.

  5. Female reproductive system
    1. Paired oviducts present.
    2. Egg chorion with stalk.

  6. Internal shell
    1. Pair of stylets or absent.


Naef (1923) placed the argonautoid families within a single family, the Argonautidae. Robson (1932) placed these families in the Tribe Argonautida and the remaining families in the Tribes Heteroglossa and Ctenoglossa based on the structure of the radula. The latter tribe contained the families Bolitaenidae and Amphitretidae. However the placement of these families in their own clade has been questioned (e.g., Thore, 1949; Voight, 1997).

Idioctopus gracilipes described by Taki (1962, 1964) and placed in its own family is probably a synonym of Amphitretus pelagicus (see Hochberg, et al, 1992).

Discussion of Phylogenetic Relationships

Voight (1997) attempted to unravel the relationships among incirrate genera using cladistic methods based on morphology. Her family-level phylogeny is as follows:

    ----------- CIRRATA
/ ---------- Bolitaenidae
/ / --------- Amphitretidae
\/ / / \----- Vitreledonellidae
\/ / ------ Alloposidae
\/ \ / ----- Argonautidae
\ \/ /
\ \/ Ocythoidae
\ \
\ \----- Tremoctopodidae
\------- Octopodidae

Thore (1949) considered the Amphitretidae and Vitreledonellidae to be closely related due to the shared position of the stomach and caecum relative to the digestive gland. Young et al. (1999) caution that the unusual position of the stomach and caecum serves to reduce the silhouette in upper mesopelagic waters and that the similarity could be convergent. In addition, the very different structure of their radulas has complicated the understanding of their relationships. Nevertheless, in a first look at relationships via a cladistic analysis based on morphology, Voight (1997) found these families grouping together, and placed the Vitreledonellidae in the synonomy of the Amphitretidae. This study, however, requires confirmation.

Molecular data from the mitochondrial Cytochrome Oxidase I gene and the nuclear Actin II gene, while without strong bootstrap support, suggest that our present classification of the Incirrata may be in error (Carlini, 1998; Carlini and Graves, 1999). Indeed, the possibility exists that all incirrate families are derived from the Octopodidae.

Life history

Where known, incirrate octopods have a life span of 6 months to 4 years; one year is typical. Those living to 4 years are large and/or cold water species. Most species appear to be terminal spawners, i. e. they lay all eggs toward the end of their life-span and die shortly after the last of the brooded embryos hatch. Some species, however, lay several egg batches and feed between spawning periods. Spawning in the epipelagic incirrates is in this latter mode but egg batches overlap (i.e. a second batch is deposited before the first one hatches).

All incirrate octopods, benthic and pelagic, brood their eggs until the young hatch. Egg strings are mostly fixed to a substrate although some species carry their eggs on the arms. Specialized brooding habits are present in pelagic incirrates. In Argonauta the dorsal arms secrete a "shell" that functions as a brood chamber and in Tremoctopus and the bolitaenids the arms form brood-chambers. In Ocythoe, the eggs develop in the oviducts and in Vitreledonella, apparently, in the mantle cavity.


Carlini, D. B. 1998. The phylogeny of coleoid cephalopods inferred from molecular evolutionary analyses of the cytochrome oxidase I, muscle actin, and cytoplasmic actin genes. Ph.D. diss. Coll. William and Mary, 273 pp.

Carlini, D. B. and J. E. Graves. 1999. Phylogenetic analysis of cytochrome c oxidase I sequences to determine higher-level relationships within the coleoid cephalopods. Bull. Mar. Sci., 64: 57-76.

Hochberg, F. G., M. Nixon and R. B. Toll. 1992. Order Octopoda Leach, 1818. In: Sweeney, M. J., C. F. E. Roper, K. M. Mangold, M. R. Clarke and S. v. Boletzky (eds.) "Larval" and juvenile cephalopods: A manual for their identification. Smithson. Contr. Zool., 513:1-282.

Mangold, K. 1989. Cephalopodes. Traité de Zoologie. Tome V. Masson, Paris. 804pp.

Naef, A. 1921/23. Cephalopoda. Fauna und Flora des Golfes von Neapel. Monograph, no. 35.

Robson, G. C. (1932). A monograph of the Recent Cephalopoda. Part II. The Octopoda (excluding the Octopodinae). Brit. Mus. (Nat. Hist.), London.

Taki, I. 1962. On species newly added to the fauna of Japanese Cephalopoda. Zool. Mag., Tokyo, 71: 397-398. [In Japanese.]

Taki, I. 1964. On eleven new species of the Cephalopoda from Japan, including two new genera of Octopodidinae. Jour. Fac. Fish. Animal Husb., Hiroshima Univ., 5: 277-343.

Thore, S. 1949. Investigations on the "Dana" Octopoda. Dana-Report No. 33, 85pp.

Voight, J. R. 1997 -- Cladistic analysis of the octopods based on anatomical characters. J. Moll. Stud. 63: 311-325.

Young, J. Z. 1971. The Anatomy of the Nervous System of Octopus vulgaris. Claredon Press, Oxford.

Young, R. E., M. Vecchione and D. Donovan. 1999. The evolution of coleoid cephalopods and their present biodiversity and ecology. South African Jour. Mar. Sci. .

Title Illustrations
Scientific Name Octopus ornatus
Location Great Barrier Reef
Specimen Condition Live Specimen
Copyright © 1996
About This Page

Katharina M. Mangold (1922-2003)

Richard E. Young

Dept of Oceanography
University of Hawaii
Honolulu, Hawaii 96822

Citing this page:

Mangold (1922-2003), Katharina M. and Young, Richard E. 1995. Incirrata Grimpe, 1916. Common octopods, octopuses or devilfishes. Version 01 January 1995 (under construction). http://tolweb.org/Incirrata/20087/1995.01.01 in The Tree of Life Web Project, http://tolweb.org/

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