This article may be too technical for most readers to understand.(February 2018) |
Eusthenopteron Temporal range: Late Devonian, | |
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Life restoration of Eusthenopteron foordi | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Sarcopterygii |
Clade: | Tetrapodomorpha |
Clade: | Eotetrapodiformes |
Family: | † Tristichopteridae |
Genus: | † Eusthenopteron Whiteaves, 1881 |
Species [1] | |
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Eusthenopteron (from Greek : εὖeû, 'good', Greek : σθένοςsthénos, 'strength', and Greek : πτερόνpteron 'wing' or 'fin') [2] is a genus of prehistoric sarcopterygian (often called "lobe-finned") 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. [3]
The genus was first described by J. F. Whiteaves in 1881, as part of a large collection of fishes from Miguasha, Quebec, Canada. [4] Some 2,000 Eusthenopteron specimens have been collected from Miguasha, one of which was the object of intensely detailed study and several papers by paleoichthyologist Erik Jarvik between the 1940s and the 1990s. [5]
Eusthenopteron is a medium- to large-sized tristichopterid. The species E. foordi is estimated to have exceeded 1.5 m (4 ft 11 in) in length, while the species E. jenkinsi probably reached 2.1 m (6 ft 11 in). [6] [1] Eusthenopteron may have weighed around 50 kilograms. [7]
The earliest known fossilized evidence of bone marrow has been found in Eusthenopteron, which may be the origin of bone marrow in tetrapods. [8]
Eusthenopteron shares many unique features among fishes but in common with the earliest-known tetrapods. It shares a similar pattern of skull roofing bones with stem tetrapoda forms such as Ichthyostega and Acanthostega . Eusthenopteron, like other tetrapodomorph fishes, had internal nostrils (or a choana), one of the defining traits of tetrapodomorphs, including tetrapods. It also had labyrinthodont teeth, characterized by infolded enamel, which characterizes all of the earliest known tetrapods as well.
Unlike the early tetrapods, Eusthenopteron did not have larval gills. [9]
Like other fish-like sarcopterygians, Eusthenopteron possessed a two-part cranium, which hinged at mid-length along an intracranial joint. Eusthenopteron's notoriety comes from the pattern of its fin endoskeleton, which bears a distinct humerus, ulna, and radius in the fore-fin and femur, tibia, and fibula in the pelvic fin. These appendicular long bones had epiphyseal growth plates that allowed substantial longitudinal growth through endochondral ossification, as in tetrapod long bones. [10] These six appendicular bones also occur in tetrapods and are a synapomorphy of a large clade of sarcopterygians, possibly Tetrapodomorpha (the humerus and femur are present in all sarcopterygians). Similarly, its elasmoid scales lack superficial odontodes composed of dentine and enamel; this loss appears to be a synapomorphy with more crownward tetrapodomorphs. [11] Eusthenopteron differs significantly from some later Carboniferous tetrapods in the apparent absence of a recognized larval stage and a definitive metamorphosis. [6] In even the smallest known specimen of Eusthenopteron foordi, with a length of 29 millimetres (1.1 in), the lepidotrichia cover all of the fins, which does not happen until after metamorphosis in genera like Polydon (the American paddlefish). This might indicate that Eusthenopteron developed directly, with the hatchling already attaining the adult's general body form (Cote et al., 2002).
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 including both a group of bony fish commonly referred to as lobe-finned fish, and tetrapods. They 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.
Anders Erik Vilhelm Jarvik was a Swedish paleontologist who worked extensively on the sarcopterygian fish Eusthenopteron. In a career that spanned some 60 years, Jarvik produced some of the most detailed anatomical work on this fish, making it arguably the best known fossil vertebrate.
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.
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
Tetrapodomorpha is 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 Tetrapodomorpha minus the crown group Tetrapoda are the stem Tetrapoda, a paraphyletic unit encompassing the fish to tetrapod transition.
Cheirolepis is an extinct genus of marine and freshwater ray-finned fish that lived in the Devonian period of Europe and North America. It is the only genus yet known within the family Cheirolepididae and the order Cheirolepidiformes. It was among the most basal of the Devonian actinopterygians and is considered the first to possess the "standard" dermal cranial bones seen in later actinopterygians.
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.
The Fram Formation is an Upper Devonian (Frasnian) sequence of rock strata on Ellesmere Island that came into prominence in 2006 with the discovery in its rocks of examples of the transitional fossil, Tiktaalik, a sarcopterygian or lobe-finned fish showing many tetrapod characteristics. Fossils of Laccognathus embryi, a porolepiform lobe-finned fish, and Qikiqtania, a close relative of Tiktaalik, were also found in the formation. The Fram Formation is a Middle to Upper Devonian clastic wedge forming an extensive continental facies consisting of sediments derived from deposits laid down in braided stream systems that formed some 375 million years ago, at a time when the North American craton ("Laurentia") was straddling the equator.
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.
Tristichopterus, with a maximum length of sixty centimetres, is the smallest genus in the family of prehistoric lobe-finned fish, Tristichopteridae that was believed to have originated in the north and dispersed throughout the course of the Upper Devonian into Gondwana. Tristichopterus currently has only one named species, first described by Egerton in 1861. The Tristichopterus node is thought to have originated during the Givetian part of the Devonian. Tristichopterus was thought by Egerton to be unique for its time period as a fish with ossified vertebral centers, breaking the persistent notochord rule of most Devonian fish but this was later reinspected and shown to be only partial ossification by Dr. R. H. Traquair. Tristichopterus alatus closely resembles Eusthenopteron and this sparked some debate after its discovery as to whether it was a separate taxon.
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.
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.
Devonosteus is an extinct genus of prehistoric marine lobe-finned fish known from the Late Devonian. It contains a single species, D. proteus from the late Frasnian of Wildungen, Germany. It has sometimes been considered a lungfish of the family Holodontidae, but this remains uncertain as the original specimen may be lost. Alternatively, it may be a tristichopterid, a type of basal tetrapodomorph.
Laccognathus is an extinct genus of amphibious lobe-finned fish from Europe and North America. They existed from the Middle Devonian to the Late Devonian. The name comes from Greek for 'pitted jaw'.
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.
The Escuminac Formation is a geologic formation in Quebec. It preserves fossils dating back to the Frasnian, in the Devonian period.
Innovations conventionally associated with terrestrially first appeared in aquatic elpistostegalians such as Panderichthys rhombolepis, Elpistostege watsoni, and Tiktaalik roseae. Phylogenetic analyses distribute the features that developed along the tetrapod stem and display a stepwise process of character acquisition, rather than abrupt. The complete transition occurred over a period of 30 million years beginning with the tetrapodomorph diversification in the Middle Devonian.