A tetrapod (; from Ancient Greek τετρα (tetra) 'four' and πούς (poús) 'foot') is any vertebrate animal of the clade Tetrapoda (). The main diagnostic and defining trait (synapomorphy) of tetrapods is having four limbs, connected to the shoulder- and pelvic girdles, for which the group is named. As a clade, Tetrapoda is defined such that it includes every organism which descended from four-limbed ancestors, including those which have lost some or all of these limbs, e.g. snakes. Tetrapods include all extant and extinct amphibians and amniotes, with the latter in turn evolving into two major clades, the sauropsids (reptiles, including dinosaurs and therefore birds) and synapsids (extinct "pelycosaurs", therapsids and all extant mammals, including humans). Hox gene mutations have resulted in some tetrapods becoming limbless (snakes, legless lizards, and caecilians) or two-limbed (cetaceans, sirenians, some lizards, kiwis, and the extinct moa and elephant birds). Nevertheless, they still qualify as tetrapods through their ancestry, and some retain a pair of vestigial spurs that are remnants of the hindlimbs. Tetrapods evolved from a group of semiaquatic animals within the tetrapodomorphs which, in turn, evolved from ancient lobe-finned fish (sarcopterygians) around 390 million years ago in the Middle Devonian period. Early tetrapodomorphs were transitional between lobe-finned fishes and true four-limbed tetrapods, though most still fit the body plan expected of other lobe-finned fishes. The oldest fossils of four-limbed vertebrates (tetrapods in the broad sense of the word) are trackways from the Middle Devonian, and body fossils became common near the end of the Late Devonian, around 370–360 million years ago. These Devonian species all belonged to the tetrapod stem group, meaning that they did not belong to any modern tetrapod group. Limbs evolved prior to terrestrial locomotion, but by the start of the Carboniferous Period, 360 million years ago, a few stem-tetrapods were experimenting with a semiaquatic lifestyle to exploit food and shelter on land. The first crown-tetrapods (those descended from the last common ancestors of extant tetrapods) appeared by the Tournaisian age of the Early Carboniferous. The specific aquatic ancestors of the tetrapods and the process by which they colonized Earth's land after emerging from water remains unclear. The transition from a body plan for gill-based aquatic respiration and tail-propelled aquatic locomotion to one that enables the animal to survive out of water and move around on land is one of the most profound evolutionary changes known. Tetrapods have numerous anatomical and physiological features that are distinct from their aquatic fish ancestors. These include distinct head and neck structures for feeding and movements, appendicular skeletons (shoulder and pelvic girdles in particular) for weight bearing and locomotion, more versatile eyes for seeing, middle ears for hearing, and more efficient heart and lungs for oxygen circulation and exchange outside water. Stem-tetrapods and "fish-a-pods" were primarily aquatic. Modern amphibians are generally semiaquatic; the first stages of their lives are as waterborne eggs and fish-like larvae with gills known as tadpoles, and later undergo metamorphosis to grow limbs and lungs and become partly terrestrial and partly aquatic. However, most tetrapod species today are amniotes, a group of mostly terrestrial tetrapods that evolved early in the Late Carboniferous. The key amniote innovation is the amnion, which enables the eggs to retain their aqueous contents on land. Some tetrapods, such as snakes and caecilians, have lost some or all of their limbs through further speciation and evolution; some only have concealed vestigial bones as a remnant of the limbs of their distant ancestors. Others returned to being amphibious or otherwise living partially or fully aquatic lives, the first during the Carboniferous period, while others, such as whales, as recently as the Cenozoic. Amniotes diverged into two branches: one, sauropsids, includes the reptiles: lepidosaurs (lizards, snakes, and the tuatara), archosaurs (crocodilians and dinosaurs, of which birds are a subset), turtles, and various other extinct forms. The other branch, synapsids, include mammals and their extinct relatives. Amniotes include the only animals apart from insects that further evolved powered flight: the extinct pterosaurs and extant birds and bats. In addition, tetrapods are the only terrestrial lineage among animals, excluding the various terrestrial lineages of Protostomia.
Definitions The precise definition of "tetrapod" is a subject of strong debate among paleontologists who work with the earliest members of the group.
Apomorphy-based definitions
A majority of paleontologists use the term "tetrapod" to refer to all vertebrates with four limbs and distinct digits (fingers and toes), as well as legless vertebrates with limbed ancestors. Limbs and digits are major apomorphies (newly evolved traits) which define tetrapods, though they are far from the only skeletal or biological innovations inherent to the group. The first vertebrates with limbs and digits evolved in the Devonian, including the Late Devonian-age Ichthyostega and Acanthostega, as well as the trackmakers of the Middle Devonian-age Zachelmie trackways. Defining tetrapods based on one or two apomorphies can present a problem if these apomorphies were acquired by more than one lineage through convergent evolution. To resolve this potential concern, the apomorphy-based definition is often supported by an equivalent cladistic definition. Cladistics is a modern branch of taxonomy which classifies organisms through evolutionary relationships, as reconstructed by phylogenetic analyses. A cladistic definition would define a group based on how closely related its constituents are. Tetrapoda is widely considered a monophyletic clade, a group with all of its component taxa sharing a single common ancestor. In this sense, Tetrapoda can also be defined as the "clade of limbed vertebrates", including all vertebrates descended from the first limbed vertebrates.
Crown group tetrapods
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