Brief Summary


Gonatids are mostly muscular squids of moderate size but are unusual in having the armature on the arms in four, rather than two, series (occasionally more than four series near the arm tips). The two medial series are usually hooks. Often the tentacular club has one very large centrally located hook. Species of the genus Gonatopsis are unusual in that they lack tentacles which are lost in the early juvenile stage.

Brief diagnosis:

An oegopsid ...

  • with tetraserial armature on arms.
  • usually with hooks on arms and often on tentacular clubs.


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Comprehensive Description


A list of all nominal genera and species in the Gonatidae can be found here. The list includes the current status and type species of all genera, and the current status, type repository and type locality of all species and all pertinent references.


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  1. Arms
    1. Arms with quadraserial armature except at arm tips in some species where sucker series increases.
    2. Arms I-III with hooks in two medial series except in Berryteuthis anonychus; the latter with hooks only in females and only at bases of arms I-III.

      Figure. Oral view of two arms of Gonatus steenstrupi with large hooks in the medial two series (white arrows) and a series of small suckers on each arm margin (black arrows). Transmitted-light photograph by M. Vecchione aboard the R/V G. O. SARS, MarEco cruise to the central North Atlantic.

  2. Tentacles
    1. Tentacular clubs with numerous irregular series of suckers; additional hooks in some genera.
    2. Tentacular clubs with unique locking-apparatus in Gonatus and Eogonatus consisting of a series of elongated ridges with medial suckers and knobs.

      Figure. Oral view of the proximal region of the club of G. steenstrupi. Arrow points to one ridge of the locking-apparatus. Transmitted-light photograph by M. Vecchione aboard the R/V G. O.SARS, MarEco cruise to the central North Atlantic.

  3. Buccal crown
    1. Buccal-crown connectives attach to ventral margins of arms IV.

  4. Funnel
    1. Funnel locking-apparatus with straight groove.

  5. Photophores
    1. Photophores absent except in G. pyros (ocular photophores).

  6. Gladius
    1. Gladius with primary conus.


The following table compares characteristics of subadults for the genera of Gonatidae.

Genus / Character Tentacles present Hooks present on clubs Club locking-apparatus
Berryteuthis Yes No Single series of suckers and knobs along entire dorsal margin of manus.
Eogonatus Yes No Short series of suckers, knobs and elongate ridges at base of manus.
Gonatopsis No NA NA
Gonatus Yes Yes Short series of suckers, knobs and elongate ridges at base of manus.


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Habitat and distribution

Gonatids are pelagic squids that occur from the surface to over 1000 m depth. Some species undergo extensive diurnal (diel) vertical migrations, ascending at night and descending during the day. A few species are associated with the ocean floor over the continental slope. Gonatids occur in high latitudes of both hemispheres. One species lives in antarctic waters, two in the North Atlantic and 16 in the high North Pacific where they are among the most abundant oceanic squids.


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Life History and Behavior

Life Cycle

Life History

Historically the paralarvae of gonatid squids have been virtually impossible to identify. Recent studies, however, found that the dorsal-head chromatophores can allow specific identification of the smallest paralarvae of at least six species in the North Pacific (Jorgensen, 2006). Jorgensen (2006) recognizes two basic patterns, Type 1 and Type 2, with species-specific variations in each.

Figure. Dorsal view of the two classes of head chromatophores presently recognized in gonatids. Type I has three tear-shaped chromatophores on each side that come to a point over each eye. Type II has three transverse rows of chromatophores with one chromatophore in the anterior row, two in the middle row and three in the posterior row. Drawings from Jorgensen (2007).


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Evolution and Systematics


Discussion of Phylogenetic Relationships

View Gonatidae Tree

There have been several recent attempts to unravel the phylogeny of the Gonatidae. Nesis (1997) relied on similarity of morphology and, apparently, results of protein electrophoresis by Katugin (1993 and 1995) to derive the following relationships:

Figure. Phylogeny of the Gonatidae. Chart modified from Nesis (1997). Subgenera are in red. Nesis included only species that were well established as valid species. Nesis considered Eogonatus to be a subgenus of Gonatus.

Clearly if this phylogeny is correct then Gonatopsis is polyphyletic. More recently Katugin (2004) has reassessed the phylogeny of the gonatidae using electrophoretic analysis of allozymes. His results are seen below.

Figure. Phylogenetic tree of the Gonatidae. Chart modified from Katugin (2004). Numbers represent genetic distance based on protein electrophoresis. Branch lengths are not drawn proportional to genetic distance.

In the paper Katugin proposes a new classification of the Gonatidae with two subfamilies (Berryteuthinae and Gonatinae) based on the radula. Within the Berryteuthinae he includes Berryteuthis (B. magister), Boreoteuthis (for Gonatopsis borealis; he elevates the subgenus Boreoteuthis to generic level) and a new genus Okutania for Berryteuthis anonychus. In the Gonatinae he includes Gonatus (including Eogonatus) and the remaining species of Gonatopsis. Katugin's phylogeny based on allozymes is similar to that of Nesis (1997) however the allozyme data shows the Gonatinae nested within the Berryteuthinae.

Recent molecular data analyzed using parsimony from 12S rRNA, 16S rRNA, and COI genes (Lindgren et al. 2005) offer a somewhat different phylogeny of the family but without good bootstrap support for most nodes. This data suggests that the Gonatus is paraphyletic. Gonatopsis borealis groups more closely with Berryteuthis than other species of Gonatopsis as suggested by both Nesis (1997) and Katugin (2004).

Figure. Phylogenetic tree of the Gonatidae. Chart simplified from Lindgren et al. (2005) by collapsing nodes with bootstrap support of 50 or less, and eliminating uncertain identifications. Numbers represent bootstrap support for relationships derived from combined molecular analyses. Numberered G. middendorffi represent different morphological types.

Because of the uncertainty in gonatid phylogeny, we retain here the standard classification for the family.


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Molecular Biology and Genetics

Molecular Biology

Statistics of barcoding coverage

Barcode of Life Data Systems (BOLD) Stats
Specimen Records:166
Specimens with Sequences:125
Specimens with Barcodes:125
Species With Barcodes:18
Public Records:66
Public Species:17
Public BINs:17
Creative Commons Attribution 3.0 (CC BY 3.0)

© Barcode of Life Data Systems

Source: Barcode of Life Data Systems (BOLD)


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Barcode data

Creative Commons Attribution 3.0 (CC BY 3.0)

© Barcode of Life Data Systems

Source: Barcode of Life Data Systems (BOLD)


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The Gonatidae, also known as armhook squid, are a family of moderately sized squid. The family contains about 19 species in three genera, widely distributed and plentiful in cold boreal waters of the Pacific Ocean. At least one species is known from Antarctic waters, and two from the North Atlantic.

Physical description[edit]

Oral view of Berryteuthis magister, showing the arrangement of suckers

Morphologically, armhook squid are fairly uniform: all species are characterised by the suckers of their arms, which are arranged in four rows (series) rather than the typical count of two. In most species, the arm's two mesial rows of suckers have been modified into hooks and the tentacular clubs—which are covered with many irregular rows of tiny suckers—may possess an enlarged central hook, with or without several smaller hooks. In the magister armhook squid (Berryteuthis magister), only the females possess hooks. Species of the genus Gonatus differ from the rest of the family (and from most squid) by their lack of tentacles as adults.

Only one species, the fiery armhook squid (Gonatus pyros), possesses photophores; these are located on the ventral periphery of the eyes.

Gonatids typically have muscular, cylindrical bodies with very soft, reddish to purplish-brown skin. The arms are thick and capable; the fins vary in shape and size, from sagittate and about 50% of the mantle length, to reniform and about 30% of the mantle length. Of moderate size, these squid range in size from 11 to 40 cm—most species are 25 cm or less. Females are somewhat larger than males.

Life history[edit]

Gonatus onyx on the Davidson Seamount at a depth of 1,328 m
Gonatus fabricii, the boreoatlantic armhook squid
Gonatus fabricii swims by the Pisces V submersible during a dive off New Zealand

These squid are pelagic, associated with the continental shelf and may roam as deep as 4,500 m or more, depending on the species. Their habits are poorly studied, but the squid are thought to undertake diel migration; by day, the squid remain in the blackness of the depths in midwater. By night, they ascend to the upper layers of the water column to feed by starlight. One species, however, Gonatopsis octopedatus, has curiously recurved arms, suggesting a benthic existence.

Little is known about the reproductive cycle of armhook squid. Most squid species whose reproduction has been obeserved have been seen to deposit eggs on the sea floor, then leave the eggs to hatch on their own. Five female Gonatus onyx squids have been observed in Monterey Canyon dragging a membrane sack containing 2,000 to 3,000 developing eggs.[1] It is uncertain if this behavior extends to other members of the Gonatidae family or if it is particular to this species.

Prey items include both benthic and pelagic species, including smaller fish, such as sculpins and juvenile pollock, crustaceans, including euphausiids and amphipods, and other squid. Cannibalism is also known to occur among the Gonatidae.

Cetaceans are important predators of gonatids; Baird's beaked whale, the narwhal, the short-finned pilot whale, Dall's porpoise, and sperm whales are all known to feed upon them. Other predators include large seabirds, northern fur seals, elephant seals, and large fish, such as grenadiers, halibut and several species of salmon. In far southern waters, Weddell seals and southern fur seals, as well as several species of albatross and penguin, feed upon Gonatus antarcticus.


The species listed above with an asterisk (*) is questionable and needs further study to determine if it is a valid species or a synonym.


  1. ^ Seibel, BA; Robison BH; Haddock SH (December 15, 2005). "Post-spawning egg care by a squid". Nature 438 (7070): 929. Bibcode:2005Natur.438..929S. doi:10.1038/438929a. PMID 16355206. 
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