Stegocephali
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Stegocephali (often spelled Stegocephalia) is a group containing all four-limbed
vertebrates Vertebrates () comprise all animal taxa within the subphylum Vertebrata () (chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with c ...
. It is equivalent to a broad definition of Tetrapoda: under this broad definition, the term "tetrapod" applies to any animal descended from the first vertebrate with limbs and toes, rather than fins. This includes both the modern lineage of limbed vertebrates (the crown group, including modern amphibians,
mammals Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur o ...
, reptiles and birds) as well as a portion of the
stem group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...
, limbed vertebrates that evolved prior to the origin of the crown group. Members of the tetrapod stem group include the earliest limbed
tetrapodomorphs The Tetrapodomorpha (also known as Choanata) are a clade of vertebrates consisting of tetrapods (four-limbed vertebrates) and their closest sarcopterygian relatives that are more closely related to living tetrapods than to living lungfish. Advance ...
such as '' Ichthyostega'' and ''
Acanthostega ''Acanthostega'' (meaning "spiny roof") is an extinct genus of stem-tetrapod, among the first vertebrate animals to have recognizable limbs. It appeared in the late Devonian period (Famennian age) about 365 million years ago, and was anatomic ...
,'' which evolved in the Devonian Period long before any modern form of tetrapod. Many paleontologists prefer a stricter definition of Tetrapoda which applies solely to the crown group, excluding earlier types of limbed tetrapodomorphs. Stegocephali was re-established to replace the broad definition of Tetrapoda, resolving the usage of two conflicting definitions in discussions of tetrapod evolution. Stegocephali (from
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
'roofed head') was coined in 1868 by the American
paleontologist Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
Edward Drinker Cope, who used it as a general category of prehistoric amphibians. This name was in reference to the skull form of many early tetrapods, with a low, solid shape combining numerous strongly-textured dermal bones. In its original usage, the term quickly became obsolete. In 1998, Canadian paleontologist
Michel Laurin Michel Laurin is a Canadian-born French vertebrate paleontologist whose specialities include the emergence of a land-based lifestyle among vertebrates, the evolution of body size and the origin and phylogeny of lissamphibians. He has also made impo ...
repopularized the term and provided a formal phylogenetic definition as a monophyletic clade containing both crown-group and stem-group tetrapods. Laurin's Stegocephali is roughly defined as including all vertebrates closer to modern tetrapods than to ''
Panderichthys ''Panderichthys'' is a genus of extinction, extinct Sarcopterygii, sarcopterygian (lobe-finned fish) from the late Devonian period, about 380 Myr, Mya. ''Panderichthys'', which was recovered from Frasnian (early Late Devonian) deposits in Latvia, ...
''. This definition was intended to include taxa with digits rather than fins, except where secondarily lost. Another definition, published in
Phylonyms The ''International Code of Phylogenetic Nomenclature'', known as the ''PhyloCode'' for short, is a formal set of rules governing phylogenetic nomenclature. Its current version is specifically designed to regulate the naming of clades, leaving the ...
, defines the group as including all taxa closer to ''
Eryops ''Eryops'' (; from Greek , , 'drawn-out' + , , 'face', because most of its skull was in front of its eyes) is a genus of extinct, amphibious temnospondyls. It contains the single species , the fossils of which are found mainly in early Permian (a ...
'' than to ''
Tiktaalik ''Tiktaalik'' (; Inuktitut ) is a monospecific genus of extinct sarcopterygian (lobe-finned fish) from the Late Devonian Period, about 375 Mya (million years ago), having many features akin to those of tetrapods (four-legged animals). It may ha ...
'', ''Panderichthys'', or ''
Eusthenopteron ''Eusthenopteron'' (from el, εὖ , 'good', el, σθένος , 'strength', and el, πτερόν 'wing' or 'fin') is a genus of prehistoric sarcopterygian (often called lobe-finned fishes) which has attained an iconic status from its close ...
''. The discovery of the
Zachelmie trackways The Zachelmie trackways are a series of Middle Devonian-age Trace fossil, trace fossils in Poland, purportedly the oldest evidence of terrestrial Vertebrate, vertebrates (Tetrapod, tetrapods) in the fossil record. These trackways were discovered in ...
in 2010 suggests that stegocephalians possibly emerged 395 Ma or earlier.


History of classification

Originally, the term was used as a systematic unit at the rank of order. The term rose to prominence in American and British science in the 19th century, though the largely equivalent term
Labyrinthodontia "Labyrinthodontia" (Greek, 'maze-toothed') is an informal grouping of extinct predatory amphibians which were major components of ecosystems in the late Paleozoic and early Mesozoic eras (about 390 to 150 million years ago). Traditionally consi ...
had been coined 18 years earlier by
Hermann Burmeister Karl Hermann Konrad Burmeister (also known as Carlos Germán Conrado Burmeister) (15 January 1807 – 2 May 1892) was a German Argentine zoologist, entomologist, herpetologist, botanist, and coleopterologist. He served as a professor at the Uni ...
in reference to the tooth structure. The terms were used interchangeably during the early 20th century, usually divided into three orders. Cope originally spelled the term as "Stegocephali", though A.S. Woodward introduced a popular alternative spelling, "Stegocephalia", in 1898. In their original usage, Stegocephali (and the Labyrinthodontia) are paraphyletic, the name is now used in an informal way to denote the early non-piscine
vertebrates Vertebrates () comprise all animal taxa within the subphylum Vertebrata () (chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with c ...
, excluding amniotes (the first reptiles and their descendants) and modern lissamphibians. Paleontologist
Michel Laurin Michel Laurin is a Canadian-born French vertebrate paleontologist whose specialities include the emergence of a land-based lifestyle among vertebrates, the evolution of body size and the origin and phylogeny of lissamphibians. He has also made impo ...
took up the older term and defined stegocephalians phylogenetically as all
vertebrates Vertebrates () comprise all animal taxa within the subphylum Vertebrata () (chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with c ...
more closely related to
Temnospondyli Temnospondyli (from Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') is a diverse order of small to giant tetrapods—often considered primitive amphibians—that flourished worldwide during the Carb ...
than to ''
Panderichthys ''Panderichthys'' is a genus of extinction, extinct Sarcopterygii, sarcopterygian (lobe-finned fish) from the late Devonian period, about 380 Myr, Mya. ''Panderichthys'', which was recovered from Frasnian (early Late Devonian) deposits in Latvia, ...
'' (the closest relative of
tetrapods Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids (reptiles, including dinosaurs and therefore birds) and synapsids ( pelycosaurs, extinct therapsi ...
known to have retained paired fins, see below). Laurin M. (1998): The importance of global parsimony and historical bias in understanding tetrapod evolution. Part I-systematics, middle ear evolution, and jaw suspension. ''Annales des Sciences Naturelles'', Zoologie, Paris, 13e Series 19: pp 1–42. Therefore, Stegocephali includes all vertebrate groups that have toes rather than fins, and a few (''
Elginerpeton ''Elginerpeton'' is a genus of stegocephalian (stem-tetrapod), the fossils of which were recovered from Scat Craig, Morayshire in the UK, from rocks dating to the late Devonian Period (Early Famennian stage, 368 million years ago). The only know ...
'', ''
Metaxygnathus ''Metaxygnathus'' is an extinct genus of ichthyostegalian found in Late Devonian deposits of New South Wales, Australia . It is known only from a lower jawbone. Previously thought to be a lobe-finned fish, it has now been reassigned to the ...
'', '' Ventastega'' and possibly ''
Hynerpeton ''Hynerpeton'' ( ) is an extinct genus of early four-limbed vertebrate that lived in the rivers and ponds of Pennsylvania during the Late Devonian period, around 365 to 363 million years ago. The only known species of ''Hynerpeton'' is ''H. basse ...
'') that may have retained paired fins. Contrary to the old usage of this term, the Stegocephali refers to a clade in this scheme. This concept of the clade Stegocephali was chosen to substitute for the name Tetrapoda by those who sought to restrict Tetrapoda to the crown group.
Laurin Laurin is both a surname and a given name. Notable people with the name include: Surname * Anna-Lena Laurin (born 1962), Swedish composer * Camille Laurin (1922–1999), psychiatrist and politician in Quebec, Canada * Dan Laurin (born 1960), Swe ...
& al
Terrestrial Vertebrates. Stegocephalians: Tetrapods and other digit-bearing vertebrates
Tree of life
As such, it encompasses all presently living land vertebrates as well as their early amphibious ancestors.


Phylogeny

Below is an evolutionary tree of stegocephalians, as recovered from an analysis by Swartz in 2012. As recovered by Clack ''et al.'' 2016:


Subgroups

*
Elginerpetontidae Elginerpetontidae is an extinct family of basal stegocephalian tetrapodomorphs which lived in Europe during the Frasnian stage of the Late Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 m ...
* Aistopoda *
Whatcheeriidae Whatcheeriidae is an extinct family of tetrapods which lived in the Mississippian sub-period, a subdivision of the Carboniferous period. It contains the genera ''Pederpes'', '' Whatcheeria'', and possibly ''Ossinodus''. Fossils of a possible w ...
*
Colosteidae Colosteidae is a family of stegocephalians (tetrapod-like vertebrates) that lived in the Carboniferous period. They possessed a variety of characteristics from different tetrapod or stem-tetrapod groups, which made them historically difficult to ...
* Adelospondyli * Baphetoidea *
crown A crown is a traditional form of head adornment, or hat, worn by monarchs as a symbol of their power and dignity. A crown is often, by extension, a symbol of the monarch's government or items endorsed by it. The word itself is used, partic ...
- Tetrapoda *Stegocephali ''
incertae sedis ' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertain ...
'' ** ''
Acanthostega ''Acanthostega'' (meaning "spiny roof") is an extinct genus of stem-tetrapod, among the first vertebrate animals to have recognizable limbs. It appeared in the late Devonian period (Famennian age) about 365 million years ago, and was anatomic ...
'' ** ''
Antlerpeton ''Antlerpeton'' is an extinct genus of early tetrapod from the Early Carboniferous of Nevada. It is known from a single poorly preserved skeleton from the Diamond Peak Formation of Eureka County. A mix of features in its compound vertebrae sugg ...
'' ** ''
Aytonerpeton ''Aytonerpeton'' is an extinct genus of Crown group#Stem groups, stem-tetrapod from the Ballagan Formation of Scotland. It was one of five new genera of early limbed vertebrates from the Ballagan Formation described by Clack ''et al.'' in 2016. T ...
'' ** ''
Brittagnathus ''Brittagnathus'' is an extinct genus of four-limbed vertebrate ("tetrapod") from the Late Devonian of Greenland. It contains a single species, ''Brittagnathus minutus'', which is based on a complete lower jaw recovered from an ''Acanthostega'' ...
'' ** '' Crassigyrinus'' ** ''
Diploradus ''Diploradus'' is an extinct genus of four-limbed stem-tetrapod from the Mississippian (mid- Tournaisian) of Scotland. It contains a single species, ''Diploradus austiumensis'', based on an incomplete skull and jaw fragments from the Ballagan ...
'' ** ''
Densignathus ''Densignathus'' is an extinct genus of early stem-tetrapod from the Late Devonian of Pennsylvania in the United States. A lower jaw has been found from the Red Hill fossil site, which is known for a diversity of lobe-finned fishes and other ear ...
'' ** ''
Doragnathus ''Doragnathus'' is an extinct genus of tetrapod from the Early Carboniferous of Scotland. References

Carboniferous tetrapods of Europe Prehistoric tetrapod genera Mississippian animals Basal tetrapods of Europe Enigmatic vertebrate ta ...
'' ** ''
Elpistostege ''Elpistostege'' is an extinct genus of finned tetrapodomorphs that lived during the late Givetian to early Frasnian ages of the Late Devonian epoch. Its only known species, ''E. watsoni'', was first described in 1938 by the British palaeontolog ...
?'' ** ''
Hynerpeton ''Hynerpeton'' ( ) is an extinct genus of early four-limbed vertebrate that lived in the rivers and ponds of Pennsylvania during the Late Devonian period, around 365 to 363 million years ago. The only known species of ''Hynerpeton'' is ''H. basse ...
'' ** ''
Ichthyerpeton ''Ichthyerpeton'' is an Extinction, extinct genus of Stegocephalia, stegocephalian ("tetrapod") from the Pennsylvanian (geology), Pennsylvanian of Ireland. It was originally described by Edward Percival WE.P. Wright and Thomas Henry Huxley in 1866 ...
'' ** '' Ichthyostega'' ** ''
Jakubsonia ''Jakubsonia'' is an extinct genus of early tetrapod from the Late Devonian of Russia. The type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is consid ...
'' ** ''
Koilops ''Koilops'' is an Extinction, extinct genus of four-limbed Stem tetrapoda, stem-tetrapod from the Mississippian (geology), Mississippian (mid-Tournaisian) of Scotland. It contains a single species, ''Koilops herma'', based on a Fossil#Casts and ...
'' ** '' Livoniana''? ** ''
Mesanerpeton ''Mesanerpeton'' is an extinct genus of four-limbed stem-tetrapod from the Mississippian ( Tournaisian) of Scotland. It contains a single species, ''Mesanerpeton woodi'', who based on a disarticulated specimen including a clavicle, vertebrae, ...
'' ** ''
Metaxygnathus ''Metaxygnathus'' is an extinct genus of ichthyostegalian found in Late Devonian deposits of New South Wales, Australia . It is known only from a lower jawbone. Previously thought to be a lobe-finned fish, it has now been reassigned to the ...
'' ** '' Occidens'' ** ''
Ossirarus ''Ossirarus'' is an extinct genus of four-limbed stem-tetrapod from the Mississippian (mid- Tournaisian) of Scotland. It contains a single species, ''Ossirarus kierani'', based on disarticulated skull and postcranial bones from the Ballagan For ...
'' ** ''
Parmastega ''Parmastega'' is an extinct genus of tetrapod from the Devonian, dated to the earliest Famennian age (about 372 million years ago), in contrast to later fossils known from late Famennian (365–359 million years ago). These later fossils are c ...
'' ** ''
Perittodus ''Perittodus'' is an extinct genus of four-limbed stem-tetrapod from the Mississippian (mid- Tournaisian) of Scotland. It contains a single species, ''Perittodus apsconditus'', based on disarticulated skull and postcranial bones from the Ballag ...
'' ** '' Sigournea'' ** '' Sinostega'' ** '' Tantallognathus'' ** ''
Tiktaalik ''Tiktaalik'' (; Inuktitut ) is a monospecific genus of extinct sarcopterygian (lobe-finned fish) from the Late Devonian Period, about 375 Mya (million years ago), having many features akin to those of tetrapods (four-legged animals). It may ha ...
?'' ** '' Tulerpeton'' ** '' Tutusius'' ** '' Umzantsia'' ** '' Ventastega'' ** ''
Ymeria ''Ymeria'' is an extinct genus of early stem tetrapod from the Devonian of Greenland. Of the two other genera of stem tetrapods from Greenland, ''Acanthostega'' and ''Ichthyostega'', ''Ymeria'' is most closely related to ''Ichthyostega'', thoug ...
''


References


External links


Tree of Life: Terrestrial Vertebrates
{{Taxonbar, from=Q7460384 Extant Late Devonian first appearances no:Panserpadder