''Heterosteus'' (also known as ''Heterostius'') is an

extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...

genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...

of heterosteid

placoderm Placoderms (from Ancient Greek πλάξ 'plax'', ''plakos'''Plate (animal anatomy), plate' and δέρμα 'derma'''skin') are vertebrate animals of the class (biology), class Placodermi, an extinct group of prehistoric fish known from Pal ...

of the

Middle Devonian In the geological timescale, the Middle Devonian epoch (from 393.3 ± 1.2 million years ago to 382.7 ± 1.6 million years ago) occurred during the Devonian period, after the end of the Emsian age. The Middle Devonian epoch is subdivided into two ...

known from remains discovered in

Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...

and

Greenland Greenland is an autonomous territory in the Danish Realm, Kingdom of Denmark. It is by far the largest geographically of three constituent parts of the kingdom; the other two are metropolitan Denmark and the Faroe Islands. Citizens of Greenlan ...

. According to Denison, 1978, Heterosteus might have been

planktivorous A planktivore is an aquatic organism that feeds on planktonic food, including zooplankton and phytoplankton. Planktivorous organisms encompass a range of some of the planet's smallest to largest multicellular animals in both the present day and i ...

, along with ''

Homosteus ''Homosteus'' (also commonly spelled ''Homostius'') is a genus of flattened arthrodire placoderm from the Middle Devonian. Fossils are found primarily in Eifelian-epoch aged strata of Europe, Canada, Greenland, and Estonia. All of the species h ...

'', and '' Titanichthys''.

Name

''Heterosteus'' was originally described in 1837 as species of ''

Trionyx ''Trionyx'' is a genus of softshell turtles belonging to the family Trionychidae. In the past many species in the family were classified in this genus, but today '' T. triunguis'', the African or Nile softshell turtle, is the only extant softshel ...

'', a softshelled turtle. Also in later studies it was often misidentified and given names like ''Ichthyosauroides'', '' Asterolepis asmussi'' and ''Chelonichthys asmusii''. Even in recent studies, it is controversial as to whether to use genus name ''Heterosteus'' or ''Heterostius''. According to

International Code of Zoological Nomenclature The International Code of Zoological Nomenclature (ICZN) is a widely accepted Convention (norm), convention in zoology that rules the formal scientific name, scientific naming of organisms treated as animals. It is also informally known as the I ...

, suffix ‘-ostius’ in

scientific name In Taxonomy (biology), taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin gramm ...

should not be corrected as ‘-osteus’, so some studies use the genus name ''Heterostius''.

Description

This genus includes the largest species in the family, and are among the largest

arthrodires Arthrodira (Greek for "jointed neck") is an order of extinct armored, jawed fishes of the class Placodermi that flourished in the Devonian period before their sudden extinction, surviving for about 50 million years and penetrating most marine eco ...

, as well, with the type species, ''H. asmussi'', having an estimated body length of up to . The genus differs from ''Herasmius'' by having the orbits on slightly longer eyestalk-like projections. The various species are found in

Givetian The Givetian is one of two faunal stages in the Middle Devonian Period. It lasted from million years ago to million years ago. It was preceded by the Eifelian Stage and followed by the Frasnian Stage. It is named after the town of Givet in ...

-aged deposits in Europe and Greenland. With the except of the German ''H. rhenanus'', all species are known from freshwater deposits: ''H. rhenanus'' is based on fragments found in a marine deposit.

Phylogeny

''Heterosteus'' is the

type genus In biological taxonomy, the type genus (''genus typica'') is the genus which defines a biological family and the root of the family name. Zoological nomenclature According to the International Code of Zoological Nomenclature, "The name-bearin ...

for the

family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...

Heterosteidae, which belongs to the

superfamily SUPERFAMILY is a database and search platform of structural and functional annotation for all proteins and genomes. It classifies amino acid sequences into known structural domains, especially into SCOP superfamilies. Domains are functional, str ...

Dunkleosteoidea Dunkleosteoidea is an extinct superfamily of arthrodire placoderms that lived during the Devonian period. The gigantic apex predator ''Dunkleosteus terrelli'' is the best known member of this group. Phylogeny Eubrachythoraci is divided into the ...

. It is a relative of the giant ''

Dunkleosteus ''Dunkleosteus'' is an extinct genus of large arthrodira, arthrodire ("jointed-neck") fish that existed during the Late Devonian period, about 382–358 million years ago. It was a pelagic fish inhabiting open waters, and one of the first verteb ...

'', as shown in the

cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...

below:

References

{{Taxonbar, from=Q3785387, from2=Q112325922 Arthrodires Placoderm families Middle Devonian first appearances Middle Devonian extinctions Fossil taxa described in 1844 Fossil taxa described in 1856