Tetrapodomorpha | |
---|---|
Life restoration of Panderichthys | |
Modern tetrapods | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Sarcopterygii |
Clade: | Rhipidistia |
Clade: | Tetrapodomorpha Ahlberg, 1991 |
Subgroups | |
See also below. |
Tetrapodomorpha (also known as Choanata [3] ) 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 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 (the last common ancestor of living tetrapods and all of its descendants) and several groups of early stem tetrapods, which includes several groups of related lobe-finned fishes, collectively known as the osteolepiforms. The Tetrapodomorpha minus the crown group Tetrapoda are the stem Tetrapoda, a paraphyletic unit encompassing the fish to tetrapod transition.
Among the characteristics defining tetrapodomorphs are modifications to the fins, notably a humerus with convex head articulating with the glenoid fossa (the socket of the shoulder joint). Another key trait is the internal nostril or choana. Most fish have two pairs of nostrils, one on either side of the head for incoming water (incurrent nostrils) and another pair for outgoing water (excurrent nostrils). In early tetrapodomorphs like Kenichthys , the excurrent nostrils had shifted towards the mouth's perimeter. In later tetrapodomorphs, including tetrapods, the excurrent nostril is positioned inside the mouth, where it is known as the choana. [4] The nearly-equivalent clade Choanata often refers to these later forms specifically. [2]
Tetrapodomorph fossils are known from the early Devonian onwards, and include Osteolepis , Panderichthys , Kenichthys and Tungsenia . [1] Tetrapodomorpha evolved from ancient lobe-finned fish (sarcopterygians) around 390 million years ago in the Middle Devonian period. [5]
After Benton, 2004; [7] and Swartz, 2012. [8]
Other clades include the Eotetrapodiformes ( Tinirau , Platycephalichthys , the Tristichopteridae and Elpistostegalia). [8] Older taxa which include late stem tetrapods and early tetrapods are the Labyrinthodontia and Ichthyostegalia.
The cladogram is based on a phylogenetic analysis of 46 taxa using 204 characters by B. Swartz in 2012. [8]
Tetrapodomorpha | |
The following cladogram follows the results found by Clement et al. (2021). [9]
Tetrapodomorpha | |
A tetrapod is any four-limbed vertebrate animal of the superclass Tetrapoda. Tetrapods include all extant and extinct amphibians and amniotes, with the latter in turn evolving into two major clades, the sauropsids and synapsids. Some tetrapods, such as snakes, legless lizards, and caecilians, have evolved to become limbless via mutations of the Hox gene. Nevertheless, these limbless groups still qualify as tetrapods through their ancestry, and some retain a pair of vestigial spurs that are remnants of the hindlimbs.
Sarcopterygii — sometimes considered synonymous with Crossopterygii — is a clade of vertebrate animals which includes a group of bony fish commonly referred to as lobe-finned fish. These vertebrates are characterised by prominent muscular limb buds (lobes) within their fins, which are supported by articulated appendicular skeletons. This is in contrast to the other clade of bony fish, the Actinopterygii, which have only skin-covered bony spines supporting the fins.
Rhipidistia, also known as Dipnotetrapodomorpha, is a clade of lobe-finned fishes which includes the tetrapods and lungfishes. Rhipidistia formerly referred to a subgroup of Sarcopterygii consisting of the Porolepiformes and Osteolepiformes, a definition that is now obsolete. However, as cladistic understanding of the vertebrates has improved over the last few decades, a monophyletic Rhipidistia is now understood to include the whole of Tetrapoda and the lungfishes.
Eusthenopteron is a genus of prehistoric sarcopterygian fish known from several species that lived during the Late Devonian period, about 385 million years ago. It has attained an iconic status from its close relationship to tetrapods. Early depictions of animals of this genus show them emerging onto land, but paleontologists now think that eusthenopteron species were strictly aquatic animals, though this is not completely known.
Panderichthys is a genus of extinct sarcopterygian from the late Devonian period, about 380 Mya. Panderichthys, which was recovered from Frasnian deposits in Latvia, is represented by two species. P. stolbovi is known only from some snout fragments and an incomplete lower jaw. P. rhombolepis is known from several more complete specimens. Although it probably belongs to a sister group of the earliest tetrapods, Panderichthys exhibits a range of features transitional between tristichopterid lobe-fin fishes and early tetrapods. It is named after the German-Baltic paleontologist Christian Heinrich Pander. Possible tetrapod tracks dating back to before the appearance of Panderichthys in the fossil record were reported in 2010, which suggests that Panderichthys is not a direct ancestor of tetrapods, but nonetheless shows the traits that evolved during the fish-tetrapod evolution
The choanae, posterior nasal apertures or internal nostrils are two openings found at the back of the nasal passage between the nasal cavity and the pharynx, in humans and other mammals. They are considered one of the most important synapomorphies of tetrapodomorphs, that allowed the passage from water to land.
Tristichopterids (Tristichopteridae) were a diverse and successful group of fish-like tetrapodomorphs living throughout the Middle and Late Devonian. They first appeared in the Eifelian stage of the Middle Devonian. Within the group sizes ranged from a few tens of centimeters (Tristichopterus) to several meters.
Kenichthys is a genus of sarcopterygian fish from the Devonian period, and a member of the clade Tetrapodomorpha. The only known species of the genus is Kenichthys campbelli, the first remains of which were found in China in 1993. The genus is important to the study of the evolution of tetrapods due to the unique nature of its nostrils, which provide vital evidence regarding the evolutionary transition of fish-like nostrils to the tetrapod choanae.
Rhizodontida is an extinct group of predatory tetrapodomorphs known from many areas of the world from the Givetian through to the Pennsylvanian - the earliest known species is about 377 million years ago (Mya), the latest around 310 Mya. Rhizodonts lived in tropical rivers and freshwater lakes and were the dominant predators of their age. They reached huge sizes - the largest known species, Rhizodus hibberti from Europe and North America, was an estimated 7 m in length, making it the largest freshwater fish known.
Eusthenodon is an extinct genus of tristichopterid tetrapodomorphs from the Late Devonian period, ranging between 383 and 359 million years ago. They are well known for being a cosmopolitan genus with remains being recovered from East Greenland, Australia, Central Russia, South Africa, Pennsylvania, and Belgium. Compared to the other closely related genera of the Tristichopteridae clade, Eusthenodon was one of the largest lobe-finned fishes and among the most derived tristichopterids alongside its close relatives Cabonnichthys and Mandageria.
Osteolepiformes, also known as Osteolepidida, is a group of prehistoric lobe-finned fishes which first appeared during the Devonian period. The order contains the families Canowindridae, Megalichthyidae, Osteolepididae and Tristichopteridae, in addition to several monotypic families. The order is generally considered to be paraphyletic because the characters that define it are mainly attributes of stem tetrapodomorphs.
Stegocephali is a clade of vertebrate animals 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.
Elpistostegalia is a clade containing Panderichthys and all more derived tetrapodomorph taxa. The earliest elpistostegalians, combining fishlike and tetrapod-like characters, such as Tiktaalik, are sometimes called fishapods. Although historically Elpistostegalia was considered an order of prehistoric lobe-finned fishes, it was cladistically redefined to include tetrapods.
Megalichthyidae is an extinct family of tetrapodomorphs which lived from the Middle–Late Devonian to the Early Permian. They are known primarily from freshwater deposits, mostly in the Northern Hemisphere, but one genus (Cladarosymblema) is known from Australia, and the possible megalichthyid Mahalalepis is from Antarctica.
Marsdenichthys is an extinct genus of Devonian tetrapodomorph. Fossils have been found from Mount Howitt in Victoria, Australia from strata that are Givetian-Frasnian in age. Mount Howitt is an important site that has been the source of many tetrapodomorph fossils, including Beelarongia and Howittichthys, both of which were first described from the locality.
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.
Eotetrapodiformes is a clade of tetrapodomorphs including the four-limbed vertebrates and their closest finned relatives, two groups of stem tetrapods called tristichopterids and elpistostegalids.
The evolution of tetrapods began about 400 million years ago in the Devonian Period with the earliest tetrapods evolved from lobe-finned fishes. Tetrapods are categorized as animals in the biological superclass Tetrapoda, which includes all living and extinct amphibians, reptiles, birds, and mammals. While most species today are terrestrial, little evidence supports the idea that any of the earliest tetrapods could move about on land, as their limbs could not have held their midsections off the ground and the known trackways do not indicate they dragged their bellies around. Presumably, the tracks were made by animals walking along the bottoms of shallow bodies of water. The specific aquatic ancestors of the tetrapods, and the process by which land colonization occurred, remain unclear. They are areas of active research and debate among palaeontologists at present.
Cladistic classification of Sarcopterygii is the classication of Sarcopterygii as a clade containing not only the lobe-finned fishes but also the tetrapods, which are closely related to lungfish. The taxon Sarcopterygii was traditionally classified as a paraphyletic group considered either a class or a subclass of Osteichthyes. Identification of the group is based on several characteristics, such as the presence of fleshy, lobed, paired fins, which are joined to the body by a single bone.