Lobe-finned fishes Temporal range: Late Silurian–Recent, 418–0Ma PreЄ Є O S D C P T J K Pg N
Coelacanth
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Subphylum:	Vertebrata
Class:	Sarcopterygii Romer, 1955
Subclasses
Actinistia — Coelacanths Dipnoi — Lungfish Tetrapodomorpha — Tetrapods and their extinct relatives.

Guiyu oneiros, the earliest known bony fish, lived during the Late Silurian, 419 million years ago).^[1][2] It has the combination of both ray-finned and lobe-finned features, although analysis of the totality of its features place it closer to lobe-finned fish.^[3][4][5][6]

The Sarcopterygii /ˌsɑrkɒptəˈrɪdʒi.aɪ/ or lobe-finned fish (from Greek σαρξ sarx, flesh, and πτερυξ pteryx, fin) – sometimes considered synonymous with Crossopterygii ("fringe-finned fish", from Greek κροσσός krossos, fringe) – constitute a clade (traditionally a class or subclass) of the bony fish, though a strict cladistic view includes the terrestrial vertebrates. The living sarcopterygians are the coelacanths, lungfish, and the tetrapods.

Characteristics [edit]

Early lobe-finned fishes are bony fish with fleshy, lobed, paired fins, which are joined to the body by a single bone.^[7] 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. The scales of sarcopterygians are true scaloids, consisting of lamellar bone surronded by layers of vascular bone, dentine-like cosmine, and external keratin.^[8] Pectoral and pelvic fins have articulations resembling those of tetrapod limbs. These fins evolved into legs of the first tetrapod land vertebrates, amphibians. They also possess two dorsal fins with separate bases, as opposed to the single dorsal fin of actinopterygians (ray-finned fish). The braincase of sarcopterygians primitively has a hinge line, but this is lost in tetrapods and lungfish. Many early sarcopterygians have a symmetrical tail. All sarcopterygians possess teeth covered with true enamel.

Most species of lobe-finned fishes are extinct. The largest known lobe-finned fish is the prehistoric predatory Hyneria that lived during the Devonian period around 360 million years ago. Hyneria grew up to 5 m (16 ft) in length^[9] and may have weighed as much as two tons.^[10] Among the two groups of extant (living) species, the coelacanths and the lungfishes, the largest species is the West Indian Ocean coelacanth, reaching 2 m (6.5 ft) in length and weighing up 110 kg (240 lb). The largest lungfish is the African lungfish which can reach 2 m (6.6 ft) in length and weigh up to 50 kg (110 lb).^[11][12]

Classification [edit]

Taxonomists who subscribe to the cladistic approach include the grouping Tetrapoda within this group, which in turn consists of all species of four-limbed vertebrates.^[13] The fin-limbs of lobe-finned fishes such as the coelacanths show a strong similarity to the expected ancestral form of tetrapod limbs. The lobe-finned fishes apparently followed two different lines of development and are accordingly separated into two subclasses, the Rhipidistia (including the Dipnoi, the lungfish, and the Tetrapodomorpha which include the Tetrapoda) and the Actinistia (coelacanths).

Taxonomy [edit]

The classification below follows Benton 2004, 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.^[14]

Actinistia	West Indian Ocean coelacanth	Actinistia, coelacanths, is a subclass of mostly fossil lobe-finned fishes. This subclass contains the coelacanths, including the two living coelacanths, the West Indian Ocean coelacanth and the king of the sea.
Dipnoi	Queensland lungfish	Dipnoi, lungfish, also known as salamanderfish,[15] is 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, 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, like Tiktaalik, have been referred to as "fishapods" by their discoverers, being half-fish half-tetrapods, in appearance and limb morphology. 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. Tetrapodomorpha contains several groups of related lobe-finned fishes, collectively known as the osteolepiforms. The Tetrapodamorpha minus the crown group Tetrapoda is 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.[16]

A modern coelacanth, Latimeria chalumnae

Queensland lungfish

• Subclass Sarcopterygii
  • Infraclass Dipnomorpha
    • Subclass Dipnoi
      • Order Ceratodontiformes
      • Order Lepidosireniformes
  • Infraclass Crossopterygii
    • • †Order Porolepiformes
      • †Order Onychodontida
      • Order Actinistia
  • Infraclass Tetrapodomorpha
    • †Order Rhizodontida
    • Superorder Osteolepidida
      • †Order Osteolepiformes
        • †Family Tristichopteridae
      • †Order Panderichthyida
      • Superclass Tetrapoda

Phylogeny [edit]

The cladogram presented below is based on studies compiled by Philippe Janvier and others for the Tree of Life Web Project.^[17]

Evolution [edit]

Evolution of lobe-finned fishes

Spindle diagram for the evolution of lobe-finned fishes, tetrapods and other vertebrate classes.^[18]

In Late Devonian vertebrate speciation, descendants of pelagic lobe-finned fish — like Eusthenopteron — exhibited a sequence of adaptations:

• Panderichthys, suited to muddy shallows;
• Tiktaalik with limb-like fins that could take it onto land;
• Early tetrapods in weed-filled swamps, such as:
  • Acanthostega, which had feet with eight digits,
  • Ichthyostega with limbs.

Descendants also included pelagic lobe-finned fish such as coelacanth species.

Lobe-finned fishes (sarcopterygians) and their relatives the ray-finned fishes (actinopterygians) comprise the superclass of bony fishes (Osteichthyes) characterized by their bony skeleton rather than cartilage. 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 fossils of sarcopterygians, found in the uppermost Silurian (ca 418 Ma), closely resembled the acanthodians (the "spiny fish", a taxon that went 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 split into two main lineages — the coelacanths and the rhipidistians. The former 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; coelacanths still live today in the oceans (genus Latimeria).

The Rhipidistians, whose ancestors probably lived in the oceans near the river mouths (estuaries), left the ocean world and migrated into freshwater habitats. They in turn split into two major groups: the lungfish and the tetrapodomorphs. The lungfish's greatest diversity was in the Triassic period; today there are fewer than a dozen genera left. The lungfish evolved the first proto-lungs and proto-limbs; developing the ability to live outside a water environment in the middle Devonian (397 - 385 Ma).

There are three major hypotheses as to how they evolved their stubby fins (proto-limbs). The traditional explanation is the "shrinking waterhole hypothesis" or "desert hypothesis" posited by the American paleontologist Alfred Romer. He believed that limbs and lungs may have evolved from the necessity of having to find new bodies of water as old waterholes dried up.^[19]

The second hypothesis is the "inter-tidal hypothesis" put forward in 2010 by a team of Polish paleontologists (Grzegorz Niedźwiedzki, Piotr Szrek, Katarzyna Narkiewicz, Marek Narkiewicz, & Per Ahlberg). They argued that sarcopterygians may have first emerged unto land from intertidal zones rather than inland bodies of water. Their hypothesis is based on the discovery of the 395 million-year-old Zachełmie tracks in Zachełmie, Świętokrzyskie Voivodeship, Poland, the oldest ever discovered fossil evidence of tetrapods.^[20][21]

The third hypothesis is dubbed the "woodland hypothesis" and was proposed by the American paleontologist Gregory J. Retallack in 2011. He argues that limbs may have developed in shallow bodies of water in woodlands as a means of navigating in environments filled with roots and vegetation. He based his conclusions on the evidence that transitional tetrapod fossils are consistently found in habitats that were formerly humid and wooded floodplains.^[19][22]

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 are the only tetrapodomorphs which survived after the Devonian.

Non-tetrapod sarcopterygians continued until towards the end of Paleozoic era, suffering heavy losses during the Permian-Triassic extinction event (251 Ma).

See also [edit]

• List of Sarcopterygii

References [edit]

Citations [edit]

Bibliography [edit]

• Nelson, Joseph S. (2006), Fishes of the World, John Wiley & Sons, ISBN 0-471-25031-7 

v t e Evolution of fish Forerunners Chordata Lancelet Haikouichthys Jawless fish Conodonts †Prioniodontida †Promissum Ostracoderms †Pteraspida †Thelodonti †Anaspida †Galeaspida †Pituriaspida †Osteostraci Jawed fish Spiny sharks †Climatiiformes †Ischnacanthiformes Placoderms †Antiarchi †Arthrodira †Brindabellaspida †Petalichthyida †Phyllolepida †Ptyctodontida †Rhenanida †Acanthothoraci † †Pseudopetalichthyida? †Stensioellida? Cartilaginous Elasmobranchii Holocephali Bony Lobe-finned Actinistia Coelacanth Lungfish Tetrapodomorpha Ray-finned Chondrostei Neopterygii Holostei Teleostei Lists Lists of prehistoric fish spiny sharks placoderms cartilaginous bony lobe-finned List of transitional fossils Related Prehistoric life Transitional fossils Vertebrate paleontology

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