Under Construction
This is an archived version of a Tree of Life page. For up-to-date information, please refer to the current version of this page.

Gonatus Gray, 1849

Tsunemi Kubodera, F. G. Hochberg, Richard E. Young, and Michael Vecchione
This genus contains the following 12 species:
taxon links [down<--]Gonatidae [up-->]Gonatus antarcticus [up-->]Gonatus berryi [up-->]Gonatus californiensis [up-->]Gonatus fabricii [up-->]Gonatus madokai [up-->]Gonatus middendorffi [up-->]Gonatus onyx [up-->]Gonatus oregonensis [up-->]Gonatus pyros [up-->]Gonatus steenstrupi [up-->]Gonatus ursabrunae Interpreting the tree
close box

This tree diagram shows the relationships between several groups of organisms.

The root of the current tree connects the organisms featured in this tree to their containing group and the rest of the Tree of Life. The basal branching point in the tree represents the ancestor of the other groups in the tree. This ancestor diversified over time into several descendent subgroups, which are represented as internal nodes and terminal taxa to the right.

example of a tree diagram

You can click on the root to travel down the Tree of Life all the way to the root of all Life, and you can click on the names of descendent subgroups to travel up the Tree of Life all the way to individual species.

For more information on ToL tree formatting, please see Interpreting the Tree or Classification. To learn more about phylogenetic trees, please visit our Phylogenetic Biology pages.

close box
Containing group: Gonatidae


Gonatus, the most speciose genus in the family, has its highest diversity in the high North Pacific. These squids reach a maximum length of 39 cm ML.


A gonatid ...


  1. Arms
    1. Hooks in medial two armature series on arms I-III.

  2. Tentacles
    1. Tentacles present in subadults.
    2. Tentacular club with one or more hooks in median line; one greatly enlarged.
    3. Proximary locking-apparatus of club with suckers and knobs medial to 4-6 large ridges and grooves, and usually several smaller ones.
       image info

      Figure. Oral view of tentacular club of G. californiensis, 315 mm GL, immature female, preserved. Photograph of preserved club by R. Young.

  3. Head
    1. Radula with 5 teeth in transverse row.

  4. Mantle
    1. Mantle tissue muscular or flabby depending on species.

  5. Fins
    1. Fins sagittate, drawn out posteriorly into a short tail.

  6. Photophores
    1. Ocular photophores present only in G. pyros.


Many of the species characteristics are found on the tentacles. The tentacles are complex and for descriptive purposes the club and stalk are divided into various zones, regions and series as follows:

 image info

Figure. Oral view of the club and distal stalk of G. californiensis showing the terminology (much of it color coded) used in describing the tentacle. Drawing modified from Young (1972).

The following table compares some characteristics of subadult species of Gonatus. The most useful character states are indicated in bold-red. The table doesn't separate all species.

Species / Character  Habitat  Ocular light organs  Proximal club hooks  Distal club hook  Medial suckers of tentacular stalk  Suckers on club  Club suckers of dorsal- and ventral marginal zones merge proximally. Arm II< III length  Club length 
G. antarcticus Antarctic waters  No  Yes Yes
 120-140 250-315   No 40-50% ML
16-17%  ML
G. berryi  North Pacific   No  Yes  Yes  0-2 159-181   No 60-70% GL  30-37% GL
G. californiensis  North Pacific
 No  Yes  Yes  40-80 215-270   No 46-53% GL  17-24% GL 
G. fabricii  North Atlantic   No  Yes  Yes  38-109 155-229  No 53-59% GL  12-20% GL 
G. madokai  North Pacific
 No  Yes  Yes  Many  ?  Yes 90% ML  20% ML 
G. middendorffi  North Pacific
 No  No  Yes  Few  ?  No 50% ML
10% ML 
G. onyx  North Pacific
 No  No  No  0-27  160-200   No 48-54% GL
20-25% GL 
G. oregonensis  North Pacific
 No  Yes  Yes  70 295-370   No 59-63% ML
21-30% ML 
G. pyros  North Pacific
 Yes  Yes  Yes  50-125 151-184   No 60-70% GL  20-25% GL 
G. steenstrupi  North Atlantic
 No  Yes  Yes  75-165 190-225   No 50-70% GL
20-36% GL 
G. ursabrunae North Pacific
 No  ?  ?  ?  ?  ? 42-56% ML  13-25% ML 

 *Known only from juveniles but with distinctive relative sucker sizes on arms and club dactylus.


Life History

Brooding of egg masses in G. onyx (Seibel et al., 2000) [suspected in G. fabricii (Bjorke et al., 1997)] occurs in deep water. Such brooding behavior, rare in oceanic squids, may prove to be characteristic of all members of the genus and perhaps the entire family. The long brooding period demanded by cold temperatures in deep water and its resulting slow population turnover rate, presumably is offset, in evolutionary time, by low egg mortality in the vast, dark, lowly populated bathypelagic environment (see Seibel, et al., 2000). For more information on brooding behavior, go to the G. onyx page.

 image info

Figure. Side view of Gonatus onyx brooding an egg mass at 2522 m depth off California in Monterey Canyon. ROV photograph from Seibel et al. (2005). © 2002 MBARI


Nesis, K. N. (1982). Abridged dey to the cephalopod mollusks of the world's ocean. 385+ii pp. Light and Food Industry Publishing House, Moscow. (In Russian.). Translated into English by B. S. Levitov, ed. by L. A. Burgess (1987), Cephalopods of the world. T. F. H. Publications, Neptune City, NJ, 351pp.

Okutani, T. and M. R. Clarke (1992). Family Gonatidae Hoyle, 1886. P. 139-156. 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. Smiths. Contr. Zool., No. 513.

Seibel, B. A., F. G. Hochberg, and D. B. Carlini. 2000. Life history of Gonatus onyx (Cephalopoda: Teuthoidea): deep-sea spawning and post-spawning egg care. Marine Biology 137 (3): 519-526.

Seibel, B. A., B. H. Robison and S. H. D. Haddock. 2005. Post-spawning egg care by a squid. Nature 438: 929.

Title Illustrations
Scientific Name Gonatus steenstrupi
Location Central North Atlantic
Specimen Condition Live Specimen
View Dorsal
Copyright ©
About This Page

Tsunemi Kubodera
National Science Museum, Tokyo, Japan

F. G. Hochberg
Santa Barbara Museum of Natural History, Santa Barbara, California, USA

Richard E. Young
University of Hawaii, Honolulu, HI, USA

National Museum of Natural History, Washington, D. C. , USA

Citing this page:

Kubodera, Tsunemi, Hochberg, F. G., Young, Richard E., and Vecchione, Michael. 2006. Gonatus Gray, 1849. Version 31 May 2006 (under construction). http://tolweb.org/Gonatus/19767/2006.05.31 in The Tree of Life Web Project, http://tolweb.org

close box

This page is a Tree of Life Branch Page.

Each ToL branch page provides a synopsis of the characteristics of a group of organisms representing a branch of the Tree of Life. The major distinction between a branch and a leaf of the Tree of Life is that each branch can be further subdivided into descendent branches, that is, subgroups representing distinct genetic lineages.

For a more detailed explanation of the different ToL page types, have a look at the Structure of the Tree of Life page.

close box


Page Content

articles & notes



Explore Other Groups

random page

  go to the Tree of Life home page