Taxon profile


merluccid hakes
Merlucciidae Rafinesque, 1815

kingdom Animalia - animals »  phylum Chordata - chordates »  class Actinopterygii - ray-finned fishes »  order Gadiformes


Inada in Cohen et al. (1990) following Inada (1989) criterion, included Merluccius Rafinesque, 1810; Macruronus Günther, 1873; Lyconus Günther,1887 and Steindachneria Goode and Bean, 1896 in the family Merlucciidae: the first three genera in the subfamily Merlucciinae and the fourth in Steindachneriinae. However, it must be pointed out that there is no existing consensus on the extent of the family Merlucciidae or whether it is a family as such, or simply a subfamily of the Gadidae. Even though all authors include in it the genus Merluccius, there are different opinions as to where Macruronus, Lyconus, Lyconodes, and Steindachneria belong.

Additional problems are whether to accept genus Lyconodes, as well as recent dissension on the validity of Lyconus.

Adams (1864) separated Merluccius from Gadidae and created for it the family Merlucciidae, which characters Gill (1884: 172-173) specified in the following way: “Gadoidea with a moderate caudal region coniform behind and with the caudal rays procurrent forwards, the anus submedian, moderate suborbital bones, terminal mouth, subjugular ventral fins, dorsal double, a short anterior and long posterior one, a long anal corresponding to the second dorsal; ribs wide, approximated, and channelled before or with inflected sides, and paired excavated frontal bones with divergent crests continuous from the forked occipital crest.”

Günther (1887) included Merluccius in the Gadidae, Macruronus in Macrouridae, and created the family Lyconidae for Lyconus. Goode and Bean (1896) accepted the families Lyconidae and Merlucciidae, but included Macruronus and Steindachneria in Macrouridae.

Gilchrist (1922) described Lyconodes, a genus of controversial affinities but usually associated with Lyconus, and included it in the subfamily Lyconinae under the Coryphaenoididae. Barnard (1925) acted in the same manner.

Svetovidov (1948), in his revision of Gadiformes, considers Merlucciinae as a subfamily of Gadidae together with Lotinae and Gadinae.

Norman (1966) was the first to consider Macruronus and Lyconus as Merlucciidae, but in a subfamily of its own (Macruroninae). Marshall (1966) and Cohen (1984) included them all in the family Merlucciidae made up of Merlucciinae (Merluccius), Macruroninae (Macruronus, Lyconus and Lyconodes) and Steindachneriinae (Steindachneria). According to Marshall (1966) the three subfamilies have the following in common: terminal mouth, front vomerine teeth biserial, 7 branchiostegal rays, the upper of which rests on the epihyal, pectoral fins with narrow base and 12 to 16 rays, ventral fins with 7 to 9 rays, and no barbel. The Merlucciinae have, among other characters, a separate caudal fin, second dorsal and anal fin of similar length and height, less than 30 vertebrae in the caudal region of the vertebral column, and prominent lower jaw; the Macruroninae and Steindachneriinae have dorsal and anal fins confluent caudally, dorsal-fin rays much longer than those of the anal, more than 30 caudal vertebrae, lower and upper jaw of equal length, and both with a few very long teeth; Steindachneria (Steindachneriinae) is singular in having the anus located between the pelvic fins and clearly separated from the urogenital orifice, which is located just before the anal fin, and also for possessing a complex bioluminescent system.

Marshall and Cohen (1973), Fahay and Markle (1984), and Okamura (1989) separate Steindachneria into a family (Steindachneriidae) and put the rest of the aforementioned genera in Merlucciidae. Nolf and Steurbaut (1989) came to a similar conclusion by analysing otoliths, but included them, respectively, in Steindachneriinae and Merlucciinae, within the Gadidae.

Using osteological and ontogenetic characters, Fahay (1989) separated Steindachneria from Merlucciidae and related them to the Macruroidei. Markle (1989) spoke of three families: Steindachneriidae, as monotypic and under Macruroidei, whereas Merlucciidae, also monotypic, and Macruronidae (Macruronus and Lyconus) are grouped under Gadoidei.

Howes (1991) states “Presumed synapomorphies relating Macruronus and Merluccius are shown to be homoplastic. Macruronus, Lyconus and possibly Lyconodes form a monophyletic group recognized as family Macruronidae; Merluccius is the sole member of Merlucciidae.” Steindachneriidae would also be monotypic.

As can be seen, at present there is no consensus on the extent of the family Merlucciidae. An in-depth revision of all genera that could be part of it is necessary.


Lloris, D.; Matallanas, J.; Oliver P., Hakes of the world (Family Merlucciidae). An annotated and illustrated catalogue of hake species known to date., FAO Species Catalogue for Fishery Purposes. No. 2. Rome, FAO. 2005
Author: Petr Čech

Included taxa

Number of records: 5

genus Lyconodes Gilchrist, 1922
genus Lyconus Günther, 1887
genus Macruronus Günther, 1873
genus Merluccius Rafinesque, 1810
genus Steindachneria Goode & Bean, 1888

Links and literature

CZ AQUATAB. World Wide Web electronic publication [family/276]

Plíštil J. (Ed.) (2009): AQUATAB. World Wide Web electronic publication [] [as Merlucciidae]
Data retrieved on: 17 January 2010
EN Australian Faunal Directory [5689d1e3-244b-4a22-b5df-2576894e8ac9]

ABRS (2009-2019): Australian Faunal Directory [], Australian Biological Resources Study, Canberra [as Merlucciidae]
Data retrieved on: 19 January 2015
CZ Hanel L., Novák J. (2007): České názvy živočichů V. Ryby a rybovití obratlovci (Pisces), 5. Paprskoploutví (Actinopterygii), kostnatí (Neopterygii) - stříbrnicotvární (Argentiniformes), ďasové (Lophiiformes), Národní muzeum (zoologické oddělení), Praha, 91 str. [as Merlucciidae]
Data retrieved on: 16 November 2013
EN Sheiko B.A. (2013): On family-group names of extant fishes and fish-like vertebrates of the world, Explorations of the fauna of the seas. St. Petersburg: Zoological Institute RAS. Vol. 74 (82). P. 1–204. [as Merlucciidae Rafinesque, 1815]
Data retrieved on: 23 November 2019

Data source info

Froese R., Pauly D. (eds.): FishBase
[Data retrieved on: 22 March 2007]

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