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Stegocephali (often spelled Stegocephalia, from
Greek Greek may refer to: 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 of all kno ...
, lit. "roofed head") is a
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
of
vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
animal Animals are multicellular, eukaryotic organisms in the Biology, biological Kingdom (biology), kingdom Animalia (). With few exceptions, animals heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, ...
s containing all fully limbed
tetrapodomorph Tetrapodomorpha (also known as Choanata) is 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. Advanced for ...
s. It is equivalent to a broad definition of the superclass Tetrapoda: under this broad definition, the term "tetrapod" applies to any animal descended from the first vertebrate with four limbs each with digits in the extremity (
pentadactyly In biology, dactyly is the arrangement of wikt:digit, digits (fingers and toes) on the hands, feet, or sometimes wings of a tetrapod animal. The term is derived from the Greek (language), Greek word () meaning "finger." Sometimes the suffix "- ...
), rather than fins of their
sarcopterygian Sarcopterygii (; )—sometimes considered synonymous with Crossopterygii ()—is a clade (traditionally a class or subclass) of vertebrate animals which includes a group of bony fish commonly referred to as lobe-finned fish. These vertebrates ar ...
relatives. Stegocephalians include both the modern lineage of limbed vertebrates (the
crown 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 ...
tetrapods, including modern
amphibian Amphibians are ectothermic, anamniote, anamniotic, tetrapod, four-limbed vertebrate animals that constitute the class (biology), class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all Tetrapod, tetrapods, but excl ...
s,
reptile Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology), orders: Testudines, Crocodilia, Squamata, and Rhynchocepha ...
s,
bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
s and
mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
s) as well as a portion of the stem group, the earliest limbed tetrapodomorphs such as ''
Ichthyostega ''Ichthyostega'', from Ancient Greek ἰχθύς (''ikthús''), meaning "fish", and στέγη (''stégē''), meaning "roof", is an Extinction, extinct genus of limbed tetrapodomorpha, tetrapodomorphs from the Devonian, Late Devonian of what is ...
'' and ''
Acanthostega ''Acanthostega'', from Ancient Greek ἄκανθα (''ákantha''), meaning "spine", and στέγη (''stégē''), meaning "roof", is an extinct genus of stem tetrapoda, stem-tetrapod, among the first vertebrates, vertebrate animals to have recogn ...
'', which evolved in the Devonian period long before the origin of the crown group. Many
paleontologist Paleontology, also spelled as palaeontology or palæontology, is the scientific study of the life of the past, mainly but not exclusively through the study of fossils. Paleontologists use fossils as a means to classify organisms, measure geolo ...
s 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 was coined in 1868 by the American paleontologist
Edward Drinker Cope Edward Drinker Cope (July 28, 1840 – April 12, 1897) was an American zoologist, paleontology, paleontologist, comparative anatomy, comparative anatomist, herpetology, herpetologist, and ichthyology, ichthyologist. Born to a wealthy Quaker fam ...
, 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 In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
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, defines the group as including all taxa closer to '' Eryops'' than to ''
Tiktaalik ''Tiktaalik'' (; ) 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). ''Tiktaalik'' is est ...
'', ''Panderichthys'', or '' Eusthenopteron''. The discovery of the Zachelmie trackways 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 Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...
. The term rose to prominence in American and British science in the 19th century, though the largely equivalent term Labyrinthodontia 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, botany, botanist, and coleopterologist. He served as a professor at ...
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 Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
, the name is now used in an informal way to denote the early non-piscine
vertebrates Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
, excluding
amniotes Amniotes are tetrapod vertebrate animals belonging to the clade Amniota, a large group that comprises the vast majority of living terrestrial and semiaquatic vertebrates. Amniotes evolved from amphibious stem tetrapod ancestors during the ...
(the first
reptiles Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology), orders: Testudines, Crocodilia, Squamata, and Rhynchocepha ...
and their descendants) and modern
lissamphibia The Lissamphibia (from Greek λισσός (lissós, "smooth") + ἀμφίβια (amphíbia), meaning "smooth amphibians") is a group of tetrapods that includes all modern amphibians. Lissamphibians consist of three living groups: the Salientia ( ...
ns. 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 In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical data ...
as all
vertebrates Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
more closely related to
Temnospondyli Temnospondyli (from Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') or temnospondyls is a diverse ancient order of small to giant tetrapods—often considered primitive amphibians—that flourished ...
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 A tetrapod (; from Ancient Greek τετρα- ''(tetra-)'' 'four' and πούς ''(poús)'' 'foot') is any four- limbed vertebrate animal of the clade Tetrapoda (). Tetrapods include all extant and extinct amphibians and amniotes, with the lat ...
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 (Late Frasnian stage, 375 million years ago). The type spe ...
'', ''
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 Sarcopterygii (; )—sometimes conside ...
'', '' 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 biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
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 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 ...
. Laurin & 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 * Aistopoda * Whatcheeriidae *
Colosteidae Colosteidae is a Family (biology), family of stegocephalians (stem-group tetrapod, tetrapods) that lived in the Carboniferous period. They possessed a variety of characteristics from different tetrapod or stem-tetrapod groups, which made them his ...
* 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, parti ...
- Tetrapoda **
Batrachomorpha The Batrachomorpha ("frog forms") are a clade containing extant and extinct amphibians that are more closely related to modern amphibians than they are to mammals and reptiles (including birds). According to many analyses they include the extinct ...
(
stem Stem or STEM most commonly refers to: * Plant stem, a structural axis of a vascular plant * Stem group * Science, technology, engineering, and mathematics Stem or STEM can also refer to: Language and writing * Word stem, part of a word respon ...
-
Lissamphibia The Lissamphibia (from Greek λισσός (lissós, "smooth") + ἀμφίβια (amphíbia), meaning "smooth amphibians") is a group of tetrapods that includes all modern amphibians. Lissamphibians consist of three living groups: the Salientia ( ...
, likely equivalent to
Temnospondyli Temnospondyli (from Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') or temnospondyls is a diverse ancient order of small to giant tetrapods—often considered primitive amphibians—that flourished ...
) **
Reptiliomorpha Reptiliomorpha (meaning reptile-shaped; in PhyloCode known as ''Pan-Amniota'') is a clade containing the amniotes and those tetrapods that share a more recent common ancestor with amniotes than with living amphibians (lissamphibians). It was defi ...
(
stem Stem or STEM most commonly refers to: * Plant stem, a structural axis of a vascular plant * Stem group * Science, technology, engineering, and mathematics Stem or STEM can also refer to: Language and writing * Word stem, part of a word respon ...
-
Amniota Amniotes are tetrapod vertebrate animals belonging to the clade Amniota, a large group that comprises the vast majority of living terrestrial and semiaquatic vertebrates. Amniotes evolved from amphibious stem tetrapod ancestors during the C ...
, a.k.a. Anthracosauria ''sensu lato'') *Stegocephali ''
incertae sedis or is a term used for a taxonomy (biology), 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, uncertainty ...
'' ** ''
Acanthostega ''Acanthostega'', from Ancient Greek ἄκανθα (''ákantha''), meaning "spine", and στέγη (''stégē''), meaning "roof", is an extinct genus of stem tetrapoda, stem-tetrapod, among the first vertebrates, vertebrate animals to have recogn ...
'' ** '' Antlerpeton'' ** '' Aytonerpeton'' ** '' Brittagnathus'' ** '' Crassigyrinus'' ** '' Diploradus'' ** '' Densignathus'' ** '' Doragnathus'' ** '' Elpistostege?'' ** '' Gaiasia'' ** ''
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'' ** ''
Ichthyostega ''Ichthyostega'', from Ancient Greek ἰχθύς (''ikthús''), meaning "fish", and στέγη (''stégē''), meaning "roof", is an Extinction, extinct genus of limbed tetrapodomorpha, tetrapodomorphs from the Devonian, Late Devonian of what is ...
'' ** ''
Jakubsonia ''Jakubsonia'' is an extinct genus of early tetrapod from the Late Devonian of Russia. The type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with w ...
'' ** '' Koilops'' ** '' Livoniana''? ** '' Mesanerpeton'' ** ''
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 Sarcopterygii (; )—sometimes conside ...
'' ** '' Occidens'' ** '' Ossirarus'' ** ''
Parmastega ''Parmastega'' is an extinct genus of stem 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 ...
'' ** '' Perittodus'' ** '' Sigournea'' ** '' Sinostega'' ** '' Tantallognathus'' ** ''
Tiktaalik ''Tiktaalik'' (; ) 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). ''Tiktaalik'' is est ...
?'' ** '' Tulerpeton'' ** '' Tutusius'' ** '' Umzantsia'' ** '' Ventastega'' ** '' Ymeria''


References


External links


Tree of Life: Terrestrial Vertebrates
{{Taxonbar, from=Q7460384 Extant Late Devonian first appearances Taxa named by Edward Drinker Cope no:Panserpadder