Stegocephali (often spelled Stegocephalia, from
Greek , 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
tetrapodomorphs. 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), rather than
fins of their
sarcopterygian relatives.
Stegocephalians include both the modern lineage of limbed vertebrates (the
crown group 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'' and ''
Acanthostega'', which evolved in the
Devonian period long before the origin of the crown group. 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 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''. 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'', ''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. 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 (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 () 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 than to ''
Panderichthys'' (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'', ''
Metaxygnathus'', ''
Ventastega'' and possibly ''
Hynerpeton'') 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.
[ 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-
Tetrapoda
**
Batrachomorpha (
stem-
Lissamphibia, likely equivalent to
Temnospondyli)
**
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-
Amniota, 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''
** ''
Antlerpeton''
** ''
Aytonerpeton''
** ''
Brittagnathus''
** ''
Crassigyrinus''
** ''
Diploradus''
** ''
Densignathus''
** ''
Doragnathus''
** ''
Elpistostege?''
** ''
Gaiasia''
** ''
Hynerpeton''
** ''
Ichthyerpeton''
** ''
Ichthyostega''
** ''
Jakubsonia''
** ''
Koilops''
** ''
Livoniana''?
** ''
Mesanerpeton''
** ''
Metaxygnathus''
** ''
Occidens''
** ''
Ossirarus''
** ''
Parmastega''
** ''
Perittodus''
** ''
Sigournea''
** ''
Sinostega''
** ''
Tantallognathus''
** ''
Tiktaalik?''
** ''
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