Stegocephalia Temporal range: Late Devonian – Present, | |
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Examples of stegocephalians (clockwise from top left): Litoria phyllochroa , Tyto alba , Tiktaalik roseae and Vulpes vulpes . | |
Scientific classification | |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Elpistostegalia |
Clade: | Stegocephalia Cope, 1868 |
Subgroups | |
Stegocephalia is a group containing all four-limbed vertebrates. 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, reptiles and birds) as well as a portion of the stem group, limbed vertebrates that evolved prior to the origin of the crown group. Members of the tetrapod stem group include the earliest limbed tetrapodomorphs such as Ichthyostega and Acanthostega, 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. Stegocephalia was re-established to replace the broad definition of Tetrapoda, resolving the usage of two conflicting definitions in discussions of tetrapod evolution.
Stegocephalia (from Greek στεγοκεφαλια 'roofed head') was coined in 1868 by the American paleontologist 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. [1] In its original usage, the term quickly became obsolete. In 1998, Canadian paleontologist Michel Laurin repopularized the term and provided a formal phylogenetic definition as a monophyletic clade containing both crown-group and stem-group tetrapods. Laurin's Stegocephalia is roughly defined as including all vertebrates with digits rather than fins, except where secondarily lost. [2]
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 in reference to the tooth structure. [3] The terms were used interchangeably during the early 20th century, usually divided into three orders. [4] However, the Stegocephalia (and the Labyrinthodontia) being paraphyletic, the name is now used in an informal way to denote the early non-piscine vertebrates, excluding amniotes (the first reptiles and their descendants) and modern lissamphibians.
Paleontologist Michel Laurin took up the older term and defined stegocephalians phylogenetically as all vertebrates more closely related to Temnospondyli than to Panderichthys (the closest relative of tetrapods known to have retained paired fins, see below). [5] Therefore, Stegocephalia 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 this scheme. This concept of the clade Stegocephalia was chosen to substitute for the name Tetrapoda by those who sought to restrict Tetrapoda to the crown group. [6] As such, it encompasses all presently living land vertebrates as well as their early amphibious ancestors.
Below is an evolutionary tree of stegocephalians, as recovered from an analysis by Swartz in 2012. [7]
Elpistostegalia |
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As recovered by Clack et al. 2016: [8]
Elpistostegalia |
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Tetrapoda | |||||||||||||||||||||||||||||||||||||||||||||||||
Tetrapods are four-limbed vertebrate animals constituting the superclass Tetrapoda. It includes extant and extinct amphibians, reptiles, and synapsids. Tetrapods evolved from a group of animals known as the Tetrapodomorpha which, in turn, evolved from ancient sarcopterygian fish around 390 million years ago in the middle Devonian period; their forms were transitional between lobe-finned fishes and the four-limbed tetrapods. The first crown-tetrapods appeared by the very early Carboniferous, 350 million years ago. The specific aquatic ancestors of the tetrapods and the process by which they colonized Earth's land after emerging from water remains unclear. The change from a body plan for breathing and navigating in water to a body plan enabling the animal to move on land is one of the most profound evolutionary changes known. Tetrapods have numerous anatomical and physiological features that are distinct from their aquatic ancestors. These include the structure of the jaw and teeth for feeding on land, limb girdles and extremities for land locomotion, lungs for respiration in air, and eyes and ears for seeing and hearing in air.
Sarcopterygii —sometimes considered synonymous with Crossopterygii —is a taxon of the bony fishes whose members are known as lobe-finned fishes. The group Tetrapoda, a superclass including amphibians, reptiles, and mammals, evolved from certain sarcopterygians; under a cladistic view, tetrapods are themselves considered a group within Sarcopterygii.
Ichthyostega is an early genus of tetrapodomorphs that lived at the end of the Late Devonian Period. It was one of the first four-limbed vertebrates in the fossil record. Ichthyostega possessed lungs and limbs that helped it navigate through shallow water in swamps. Although Ichthyostega is often labelled a "tetrapod" due to the possession of limbs and fingers, it was more basal ("primitive") than true crown-tetrapods, and could more accurately be referred to as a stegocephalian or stem tetrapod. Likewise, while undoubtedly of amphibian build and habit, it is not considered a true member of the group in the narrow sense, as the first modern amphibians appeared in the Triassic Period. Until finds of other early stegocephalians and closely related fishes in the late 20th century, Ichthyostega stood alone as a transitional fossil between fish and tetrapods, combining fish- and tetrapod-like features. Newer research has shown that it had an unusual anatomy, functioning more akin to a seal than a salamander, as previously assumed.
Jennifer Alice Clack, was an English palaeontologist and evolutionary biologist. She specialised in the early evolution of tetrapods, specifically studying the "fish to tetrapod" transition: the origin, evolutionary development and radiation of early tetrapods and their relatives among the lobe-finned fishes. She is best known for her book Gaining Ground: the Origin and Early Evolution of Tetrapods, published in 2002 and written with the layperson in mind.
"Labyrinthodontia" is an informal grouping of extinct predatory amphibians which were major components of ecosystems in the late Paleozoic and early Mesozoic eras. Traditionally considered a subclass of the class Amphibia, modern classification systems recognize that labyrinthodonts are not a formal natural group (clade) exclusive of other tetrapods. Instead, they consistute an evolutionary grade, ancestral to living tetrapods such as lissamphibians and amniotes. "Labyrinthodont"-grade vertebrates evolved from lobe-finned fishes in the Devonian, though a formal boundary between fish and amphibian is difficult to define at this point in time.
Acanthostega is an extinct genus of stem-tetrapod, among the first vertebrate animals to have recognizable limbs. It appeared in the late Devonian period about 365 million years ago, and was anatomically intermediate between lobe-finned fishes and those that were fully capable of coming onto land.
The Batrachomorpha are a clade containing recent and extinct amphibians that are more closely related to modern amphibians than they are to mammals and reptiles. According to many analyses they include the extinct Temnospondyli; some show that they include the Lepospondyli instead. The name traditionally indicated a more limited group.
Lepospondyli is a diverse taxon of early tetrapods. With the exception of one late-surviving lepospondyl from the Late Permian of Morocco, lepospondyls lived from the Early Carboniferous (Mississippian) to the Early Permian and were geographically restricted to what is now Europe and North America. Five major groups of lepospondyls are known: Adelospondyli; Aïstopoda; Lysorophia; Microsauria; and Nectridea. Lepospondyls have a diverse range of body forms and include species with newt-like, eel- or snake-like, and lizard-like forms. Various species were aquatic, semiaquatic, or terrestrial. None were large, and they are assumed to have lived in specialized ecological niches not taken by the more numerous temnospondyl amphibians that coexisted with them in the Paleozoic. Lepospondyli was named in 1888 by Karl Alfred von Zittel, who coined the name to include some tetrapods from the Paleozoic, that shared some specific characteristics in the notochord and teeth. Lepospondyls have sometimes been considered to be either related or ancestral to modern amphibians or to Amniota. It has been suggested that the grouping is polyphyletic, with aïstopods being primitive stem-tetrapods, while recumbirostran microsaurs are primitive reptiles.
Anthracosauria is an order of extinct reptile-like amphibians that flourished during the Carboniferous and early Permian periods, although precisely which species are included depends on one's definition of the taxon. "Anthracosauria" is sometimes used to refer to all tetrapods more closely related to amniotes such as reptiles, mammals, and birds, than to lissamphibians such as frogs and salamanders. An equivalent term to this definition would be Reptiliomorpha. Anthracosauria has also been used to refer to a smaller group of large, crocodilian-like aquatic tetrapods also known as embolomeres.
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. bassetti, named after the describer's grandfather, city planner Edward Bassett. Hynerpeton is known for being the first Devonian four-limbed vertebrate discovered in the United States, as well as possibly being one of the first to have lost internal (fish-like) gills.
The Tetrapodomorpha are a clade of vertebrates consisting of tetrapods and their closest sarcopterygian relatives that are more closely related to living tetrapods than to living lungfish. Advanced forms transitional between fish and the early labyrinthodonts, such as Tiktaalik, have been referred to as "fishapods" by their discoverers, being half-fish, half-tetrapods, in appearance and limb morphology. The Tetrapodomorpha contains the crown group tetrapods and several groups of early stem tetrapods, which includes several groups of related lobe-finned fishes, collectively known as the osteolepiforms. The Tetrapodamorpha minus the crown group Tetrapoda are the Stem Tetrapoda, a paraphyletic unit encompassing the fish to tetrapod transition.
Crassigyrinus is an extinct genus of carnivorous stem tetrapod from the Early Carboniferous Limestone Coal Group of Scotland and possibly Greer, West Virginia. The type specimen was originally described as Macromerium scoticum and lacked a complete skull. With subsequent discoveries, Crassigyrinus is now known from three skulls, one of which is in articulation with a fairly complete skeleton, and two incomplete lower jaws. Crassigyrinus grew up to 2 meters in length, coupled with tiny limbs and unusually large jaws. Crassigyrinus is taxonomically enigmatic, having confused paleontologists for decades with its apparent fish-like and tetrapod features. It was traditionally placed within the group Labyrinthodontia along with many other early tetrapods. Some paleontologists have even considered it as the most basal crown group tetrapod, while others hesitate to even place it within the Tetrapoda superclass. Crassigyrinus had unusually large jaws, enabling it to eat other animals it could catch and swallow. It had two rows of sharp teeth in its jaws, the second row having a pair of fangs. Crassigyrinus had large eyes, suggesting that it was either nocturnal, or lived in very murky water.
Ventastega is an extinct genus of stem tetrapod that lived during the Upper Fammenian of the Late Devonian, approximately 372.2 to 358.9 million years ago. Only one species is known that belongs in the genus, Ventastega curonica, which was described in 1996 after fossils were discovered in 1933 and mistakenly associated with a fish called Polyplocodus wenjukovi. ‘Curonica’ in the species name refers to Curonia, the Latin name for Kurzeme, a region in western Latvia. Ventastega curonica was discovered in two localities in Latvia, and was the first stem tetrapod described in Latvia along with being only the 4th Devonian tetrapodomorph known at the time of description. Based on the morphology of both cranial and post-cranial elements discovered, Ventastega is more primitive than other Devonian tetrapodomorphs including Acanthostega and Ichthyostega, and helps further understanding of the fish-tetrapod transition.
Tulerpeton is an extinct genus of Devonian four-limbed vertebrate, known from a fossil that was found in the Tula Region of Russia at a site named Andreyevka. This genus and the closely related Acanthostega and Ichthyostega represent the earliest tetrapods.
Embolomeri is an order of tetrapods or stem-tetrapods, possibly members of Reptiliomorpha. Embolomeres first evolved in the Early Carboniferous (Mississippian) Period and were the largest and most successful predatory tetrapods of the Late Carboniferous (Pennsylvanian) Period. They were specialized semiaquatic predators with long bodies for eel-like undulatory swimming. Embolomeres are characterized by their vertebral centra, which are formed by two cylindrical segments, the pleurocentrum at the rear and intercentrum at the front. These segments are equal in size. Most other tetrapods have pleurocentra and intercentra which are drastically different in size and shape.
Polydactyly in stem-tetrapods should here be understood as having more than five digits to the finger or foot, a condition that was the natural state of affairs in the very first stegocephalians during the evolution of terrestriality. The polydactyly in these largely aquatic animals is not to be confused with polydactyly in the medical sense, i.e. it was not an anomaly in the sense it was not a congenital condition of having more than the typical number of digits for a given taxon. Rather, it appears to be a result of the early evolution from a limb with a fin rather than digits.
Ichthyostegalia is an order of extinct amphibians, representing the earliest landliving vertebrates. The group is thus an evolutionary grade rather than a clade. While the group are recognized as having feet rather than fins, most, if not all, had internal gills in adulthood and lived primarily as shallow water fish and spent minimal time on land.
The Stem Tetrapoda are a cladistically defined group, consisting of all animals more closely related to extant four-legged vertebrates than to their closest extant relatives, but excluding the crown group Tetrapoda. They are thus paraphyletic, though acceptable in phylogenetic nomenclature as the group is defined by strict reference to phylogeny rather than to traits as in traditional systematics. Thus, some finned sarcopterygians are considered to be stem tetrapods.
Tinirau is an extinct genus of sarcopterygian fish from the Middle Devonian of Nevada. Although it spent its entire life in the ocean, Tinirau is a stem tetrapod close to the ancestry of land-living vertebrates in the crown group Tetrapoda. Relative to more well-known stem tetrapods, Tinirau is more closely related to Tetrapoda than is Eusthenopteron, but farther from Tetrapoda than is Panderichthys. The type and only species of Tinirau is T. clackae, named in 2012.
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 bonebed in the Britta Dal Formation. It is the fourth named genus of "tetrapod" from the Late Devonian of Greenland, after Ichthyostega, Acanthostega, and Ymeria.