Lobe-finned fishes | |
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From top to bottom and left to right, examples of sarcopterygians: Guiyu oneiros , West Indian Ocean coelacanth, Australian lungfish and the tetrapodomorph Panderichthys rhombolepis . | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Superclass: | Osteichthyes |
Clade: | Sarcopterygii Romer, 1955 |
Subgroups | |
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Sarcopterygii ( /ˌsɑːrkɒptəˈrɪdʒi.aɪ/ ; from Ancient Greek σάρξ (sárx) 'flesh'and πτέρυξ (ptérux) 'wing, fin') — sometimes considered synonymous with Crossopterygii (from Ancient Greek κροσσός (krossós) 'fringe') — is a clade (traditionally a class or subclass) 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.
The tetrapods, a mostly terrestrial superclass of vertebrates, are now recognized as having evolved from sarcopterygian ancestors and are most closely related to lungfishes. Their paired pectoral and pelvic fins evolved into limbs, and their foregut diverticulum eventually evolved into air-breathing lungs. Cladistically, this would make the tetrapods a subgroup within Sarcopterygii and thus sarcopterygians themselves. As a result, the phrase "lobe-finned fish" normally refers to not the entire clade but only aquatic members that are not tetrapods, i.e. a paraphyletic group.
Non-tetrapod sarcopterygians were once the dominant predators of freshwater ecosystems during the Carboniferous and Permian periods, but suffered significant decline after the Great Dying. The only known extant non-tetrapod sarcopterygians are the two species of coelacanths and six species of lungfishes.
Early lobe-finned fishes are bony fish with fleshy, lobed, paired fins, which are joined to the body by a single bone. [5] The fins of lobe-finned fishes differ from those of all other fish in that each is borne on a fleshy, lobelike, scaly stalk extending from the body that resembles a limb bud. The scales of sarcopterygians are true scaloids, consisting of lamellar bone surrounded by layers of vascular bone, cosmine (similar to dentin), and external keratin. [6] The physical structure of tetrapodomorphs, fish bearing resemblance to tetrapods, provides valuable insights into the evolutionary shift from aquatic to terrestrial existence. [7] Pectoral and pelvic fins have articulations resembling those of tetrapod limbs. The first tetrapod land vertebrates, basal amphibian organisms, possessed legs derived from these fins. Sarcopterygians also possess two dorsal fins with separate bases, as opposed to the single dorsal fin in ray-finned fish. The braincase of sarcopterygians primitively has a hinge line, but this is lost in tetrapods and lungfish. Early sarcopterygians commonly exhibit a symmetrical tail, while all sarcopterygians possess teeth that are coated with genuine enamel.
Most species of lobe-finned fishes are extinct. The largest known lobe-finned fish was Rhizodus hibberti from the Carboniferous period of Scotland which may have exceeded 7 meters in length. Among the two groups of living species, the coelacanths and the lungfishes, the largest species is the West Indian Ocean coelacanth, reaching 2 m (6 ft 7 in) in length and weighing up 110 kg (240 lb). The largest lungfish is the marbled lungfish which can reach 2 m (6.6 ft) in length and weigh up to 50 kg (110 lb). [8] [9]
Taxonomists who adhere to the cladistic approach include Tetrapoda within this classification, encompassing all species of vertebrates with four limbs. [10] The fin-limbs found in lobe-finned fishes like the coelacanths display a strong resemblance to the presumed ancestral form of tetrapod limbs. Lobe-finned fishes seemingly underwent two distinct evolutionary paths, leading to their classification into two subclasses: the Rhipidistia (comprising the Dipnoi, or lungfish, and the Tetrapodomorpha, which includes the Tetrapoda) and the Actinistia (represented by coelacanths).
The classification below follows Benton (2004), [11] and uses a synthesis of rank-based Linnaean taxonomy and also reflects evolutionary relationships. Benton included the superclass Tetrapoda in the subclass Sarcopterygii in order to reflect the direct descent of tetrapods from lobe-finned fish, despite the former being assigned a higher taxonomic rank. [11]
Actinistia | Actinistia, coelacanths, are a subclass of lobe-finned fishes, all but two of which are species only known through fossils. The subclass Actinistia contains the coelacanths, including the two living coelacanths: the West Indian Ocean coelacanth and the Indonesian coelacanth. | |
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Dipnoi | Dipnoi, commonly referred to as lungfish, but also known as salamanderfish, [12] are a subclass of freshwater fish. Lungfish are best known for retaining characteristics primitive within the bony fishes, including the ability to breathe air, and structures primitive within the lobe-finned fishes, including the presence of lobed fins with a well-developed internal skeleton. Today, lungfish live only in Africa, South America, and Australia. While vicariance would suggest this represents an ancient distribution limited to the Mesozoic supercontinent Gondwana, the fossil record suggests advanced lungfish had a widespread freshwater distribution and the current distribution of modern lungfish species reflects extinction of many lineages following the breakup of Pangaea, Gondwana, and Laurasia. | |
Tetrapodomorpha | Advanced tetrapodomorph Tiktaalik | Tetrapodomorpha, tetrapods and their extinct relatives, are 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. [13] Advanced forms transitional between fish and the early labyrinthodonts, like Tiktaalik , have been referred to as "fishapods" by their discoverers, being half-fish, half-tetrapods, in appearance and limb morphology. The Tetrapodomorpha contain the crown group tetrapods (the last common ancestor of living tetrapods and all of its descendants) and several groups of early stem tetrapods, and 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. Among the characters defining tetrapodomorphs are modifications to the fins, notably a humerus with convex head articulating with the glenoid fossa (the socket of the shoulder joint). Tetrapodomorph fossils are known from the early Devonian onwards, and include Osteolepis , Panderichthys , Kenichthys , and Tungsenia . [14] |
Lobe-finned fishes and their sister group, the ray-finned fishes, make up the superclass Osteichthyes, characterized by the presence of swim bladders (which share ancestry with lungs) as well as the evolution of ossified endoskeleton instead of cartilages like the skeletons of acanthodians, chondrichthyians and most placoderms. There are otherwise vast differences in fin, respiratory and circulatory structures between the Sarcopterygii and the Actinopterygii, such as the presence of cosmoid layers in the scales of sarcopterygians. The earliest sarcopterygian fossils were found in the uppermost Silurian, about 418 Ma. They closely resembled the acanthodians (the "spiny fish", a taxon that became extinct at the end of the Paleozoic). In the early–middle Devonian (416–385 Ma), while the predatory placoderms dominated the seas, some sarcopterygians came into freshwater habitats.
In the Early Devonian (416–397 Ma), the sarcopterygians, or lobe-finned fishes, split into two main lineages: the coelacanths and the rhipidistians. Coelacanths never left the oceans and their heyday was the late Devonian and Carboniferous, from 385 to 299 Ma, as they were more common during those periods than in any other period in the Phanerozoic. Actinistians, a group within the lobe-finned fish, have been around for almost 380 million years. Over time, researchers have identified 121 species spread across 47 genera. Some species are well-documented in their evolutionary placement, while others are harder to track. The greatest boom in actinistian diversity happened during the Early Triassic, just after the Great Dying. [16] Coelacanths of the genus Latimeria still live today in the open oceans and retained many primordial features of ancient sarcopterygians, earning them a reputation as living fossils.
The rhipidistians, whose ancestors probably lived in the oceans near river mouths and estuaries, left the marine world and migrated into freshwater habitats. They then split into two major groups: the lungfish and the tetrapodomorphs, and both of them evolved their swim bladders into air-breathing lungs. Lungfish radiated into their greatest diversity during the Triassic period; today, fewer than a dozen genera remain, having evolved the first proto-lungs and proto-limbs, adapting to living outside a submerged water environment by the middle Devonian (397–385 Ma). The tetrapodomorphs, on the other hand, evolved into the fully-limbed stegocephalians and later the fully terrestrial tetrapods during the Late Devonian, when the Late Devonian Extinction bottlenecked and selected against the more aquatically adapted groups among stem-tetrapods. [17] [18] The surviving tetrapods then underwent adaptive radiation on dry land and become the dominant terrestrial animals during the Carboniferous and the Permian periods.
There are three major hypotheses as to how lungfish evolved their stubby fins (proto-limbs).
The first tetrapodomorphs, which included the gigantic rhizodonts, had the same general anatomy as the lungfish, who were their closest kin, but they appear not to have left their water habitat until the late Devonian epoch (385–359 Ma), with the appearance of tetrapods (four-legged vertebrates). Tetrapods and megalichthyids are the only tetrapodomorphs which survived after the Devonian, with the latter group disappearing during the Permian. [31]
Non-tetrapod sarcopterygians continued until towards the end of Paleozoic era, suffering heavy losses during the Permian–Triassic extinction event (251 Ma).
The cladogram presented below is based on studies compiled by Janvier et al. (1997) for the Tree of Life Web Project, [32] Mikko's Phylogeny Archive [33] and Swartz (2012). [34]
Sarcopterygii |
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[37] ==References==
A popular exposition of the doctrine of evolution in general, and of that of Darwin, Goethe, and Lamarck in particular.
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. Hox gene mutations have resulted in some tetrapods becoming limbless or two-limbed. 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.
Euteleostomi is a successful clade that includes more than 90% of the living species of vertebrates. Both its major subgroups are successful today: Actinopterygii includes most extant bony fish species, and Sarcopterygii includes the tetrapods.
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.
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
Tiktaalik is a monospecific genus of extinct sarcopterygian from the Late Devonian Period, about 375 Mya, having many features akin to those of tetrapods. Tiktaalik is estimated to have had a total length of 1.25–2.75 metres (4.1–9.0 ft) on the basis of various specimens.
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.
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.
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.
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 earliest 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.
Guiyu oneiros is one of the earliest articulated bony fish discovered. Fossils of Guiyu have been found in what is now Qujing, Yunnan, China, in late Silurian marine strata, about 425 million years old.
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'.
Fins are moving appendages protruding from the body of fish that interact with water to generate thrust and help the fish swim. Apart from the tail or caudal fin, fish fins have no direct connection with the back bone and are supported only by muscles.
The evolution of fish began about 530 million years ago during the Cambrian explosion. It was during this time that the early chordates developed the skull and the vertebral column, leading to the first craniates and vertebrates. The first fish lineages belong to the Agnatha, or jawless fish. Early examples include Haikouichthys. During the late Cambrian, eel-like jawless fish called the conodonts, and small mostly armoured fish known as ostracoderms, first appeared. Most jawless fish are now extinct; but the extant lampreys may approximate ancient pre-jawed fish. Lampreys belong to the Cyclostomata, which includes the extant hagfish, and this group may have split early on from other agnathans.
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.
Cosmine is a spongy, bony material that makes up the dentine-like layers in the scales of the lobe-finned fishes of the class Sarcopterygii. Fish scales that include layers of cosmine are known as cosmoid scales.
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.
The evolution of fishes took place over a timeline which spans the Cambrian to the Cenozoic, including during that time in particular the Devonian, which has been dubbed the "age of fishes" for the many changes during that period.