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Sarcopterygii (/ˌsɑːrkɒptəˈrɪi./; 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) 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.

Lobe-finned fishes
Temporal range:
Late SilurianPresent, 425–0 Ma[1]
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 Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Superclass: Osteichthyes
Clade: Sarcopterygii
Romer, 1955

The tetrapods, a mostly terrestrial superclass of vertebrates, are now recognized as having evolved from sarcopterygian ancestors and are most closely related to lungfishes. Cladistically, this would make the tetrapods a subgroup of Sarcopterygii and thus sarcopterygians themselves. As a result, the phrase "lobe-finned fish" normally refers to not the entire clade but only its members which are not tetrapods.

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.

Characteristics edit

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

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]

Classification edit

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).

Taxonomy edit

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.
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.
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]

Evolution edit

Evolution of lobe-finned fishes
Spindle diagram for the evolution of lobe-finned fishes, tetrapods and other vertebrate classes[15]
In Late Devonian vertebrate speciation, descendants of pelagic lobe-finned fish—like Eusthenopteron — exhibited a sequence of adaptations: Descendants also included pelagic lobe-finned fish such as coelacanth species.
Tooth from the sarcopterygian Onychodus from the Devonian of Wisconsin

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. 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.[16][17] The surviving tetrapods then underwent adaptive radiation on dry land and become the dominant terrestrial animals during the Carboniferous and the Permian periods.

Hypotheses for means of pre-adaptation edit

There are three major hypotheses as to how lungfish evolved their stubby fins (proto-limbs).

Shrinking waterhole
The first, traditional explanation is the "shrinking waterhole hypothesis", or "desert hypothesis", posited by the American paleontologist Alfred Romer, who believed that limbs and lungs may have evolved from the necessity of having to find new bodies of water as old waterholes dried up.[18]
Inter-tidal adaptation
Niedźwiedzki, Szrek, Narkiewicz, et al. (2010)[19] proposed a second, the "inter-tidal hypothesis": That sarcopterygians may have first emerged unto land from intertidal zones rather than inland bodies of water, based on the discovery of the 395 million-year-old Zachełmie tracks, the oldest discovered fossil evidence of tetrapods.[19][20]
Woodland swamp adaptation
Retallack (2011)[21] proposed a third hypothesis is dubbed the "woodland hypothesis": Retallack 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.[18][21]
Habitual escape onto land
A fourth, minority hypothesis posits that advancing onto land achieved more safety from predators, less competition for prey, and certain environmental advantages not found in water—such as oxygen concentration,[24] and temperature control[26]—implying that organisms developing limbs were also adapting to spending some of their time out of water. However, studies have found that sarcopterygians developed tetrapod-like limbs suitable for walking well before venturing onto land.[29] This suggests they adapted to walking on the ground-bed under water before they advanced onto dry land.

History through to the end-Permian extinction edit

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.[30]

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

Phylogeny edit

The cladogram presented below is based on studies compiled by Janvier et al. (1997) for the Tree of Life Web Project,[31] Mikko's Phylogeny Archive[32] and Swartz (2012).[33]

Life restoration of Sparalepis tingi and other fauna from the Silurian of Yunnan

See also edit

Footnotes edit

  1. ^ The Osteolepida taxa were not addressed by Ahlberg & Johanson (1998).[citation needed]

References edit

  1. ^ a b Zhao, W.; Zhang, X.; Jia, G.; Shen, Y.; Zhu, M. (2021). "The Silurian-Devonian boundary in East Yunnan (South China) and the minimum constraint for the lungfish-tetrapod split". Science China Earth Sciences. 64 (10): 1784–1797. Bibcode:2021ScChD..64.1784Z. doi:10.1007/s11430-020-9794-8. S2CID 236438229.
  2. ^ Zhu, M.; Zhao, W.; Jia, L.; Lu, J.; Qiao, T.; Qu, Q. (2009). "The oldest articulated osteichthyan reveals mosaic gnathostome characters". Nature. 458 (7237): 469–474. Bibcode:2009Natur.458..469Z. doi:10.1038/nature07855. PMID 19325627. S2CID 669711.
  3. ^ Coates, M.I. (2009). "Palaeontology: Beyond the age of fishes". Nature. 458 (7237): 413–414. Bibcode:2009Natur.458..413C. doi:10.1038/458413a. PMID 19325614. S2CID 4384525.
  4. ^ "Pharyngula – Guiyu oneiros". Science Blogs (blog). 1 April 2009. Archived from the original on 9 March 2012.
  5. ^ Clack, J.A. (2002). Gaining Ground. Indiana University.
  6. ^ Kardong, Kenneth V. (1998). Vertebrates: Comparative anatomy, function, evolution (second ed.). USA: McGraw-Hill. ISBN 0-07-115356-X. ISBN 0-697-28654-1
  7. ^ Clack, J.A. (2009). "The fin to limb transition: New data, interpretations, and hypotheses from paleontology and developmental biology". Annual Review of Earth and Planetary Sciences. 37 (1): 163–179. Bibcode:2009AREPS..37..163C. doi:10.1146/
  8. ^ Froese, Rainer, and Daniel Pauly, eds. (2009). "Lepidosirenidae" in FishBase. January 2009 version.
  9. ^ "Protopterus aethiopicus". Lung fishes. Archived from the original on 3 August 2011.
  10. ^ Nelson, Joseph S. (2006). Fishes of the World. John Wiley & Sons. ISBN 978-0-471-25031-9.
  11. ^ a b Benton, M.J. (2004). Vertebrate Paleontology (3rd ed.). Blackwell Science.
  12. ^ Haeckel, Ernst Heinrich Philipp August (1892). Lankester, Edwin Ray; Schmitz, L. Dora (eds.). The History of Creation, or, the Development of the Earth and Its Inhabitants by the Action of Natural Causes (8th, German ed.). D. Appleton. p. 289. A popular exposition of the doctrine of evolution in general, and of that of Darwin, Goethe, and Lamarck in particular.
  13. ^ Amemiya, C.T.; Alfoldi, J.; Lee, A.P.; Fan, S.H.; Philippe, H.; MacCallum, I.; Braasch, I.; et al. (2013). "The African coelacanth genome provides insights into tetrapod evolution". Nature. 496 (7445): 311–316. Bibcode:2013Natur.496..311A. doi:10.1038/nature12027. hdl:1912/5869. PMC 3633110. PMID 23598338.
  14. ^ Lu, Jing; Zhu, Min; Long, John A.; Zhao, Wenjin; Senden, Tim J.; Jia, Liantao; Qiao, Tuo (2012). "The earliest known stem-tetrapod from the lower Devonian of China". Nature Communications. 3: 1160. Bibcode:2012NatCo...3.1160L. doi:10.1038/ncomms2170. hdl:1885/69314. PMID 23093197.
  15. ^ Benton, M. J. (2005) Vertebrate Palaeontology, Blackwell, 3rd edition, Fig 2.10 on page 35 and Fig 3.25 on page 73
  16. ^ George r. Mcghee, Jr (12 November 2013). When the Invasion of Land Failed: The Legacy of the Devonian Extinctions. Columbia University Press. ISBN 9780231160575. Archived from the original on 27 December 2019. Retrieved 1 March 2016.
  17. ^ "Research project: The Mid-Palaeozoic biotic crisis: Setting the trajectory of Tetrapod evolution". Archived from the original on 12 December 2013. Retrieved 31 May 2014.
  18. ^ a b "Fish-tetrapod transition got a new hypothesis in 2011". Science 2.0. 27 December 2011. Retrieved 2 January 2012.
  19. ^ a b Niedźwiedzki, Grzegorz; Szrek, Piotr; Narkiewicz, Katarzyna; Narkiewicz, Marek; Ahlberg, Per E. (2010). "Tetrapod trackways from the early Middle Devonian period of Poland". Nature. 463 (7277): 43–48. Bibcode:2010Natur.463...43N. doi:10.1038/nature08623. PMID 20054388. S2CID 4428903.
  20. ^ Barley, Shanta (6 January 2010). "Oldest footprints of a four-legged vertebrate discovered". New Scientist. Retrieved 3 January 2010.
  21. ^ a b Retallack, Gregory (May 2011). "Woodland hypothesis for Devonian tetrapod evolution". Journal of Geology. 119 (3). University of Chicago Press: 235–258. Bibcode:2011JG....119..235R. doi:10.1086/659144. S2CID 128827936.
  22. ^ Carroll, R.L.; Irwin, J.; Green, D.M. (2005). "Thermal physiology and the origin of terrestriality in vertebrates". Zoological Journal of the Linnean Society. 143 (3): 345–358. doi:10.1111/j.1096-3642.2005.00151.x.
  23. ^ a b Hohn-Schulte, B.; Preuschoft, H.; Witzel, U.; Distler-Hoffmann, C. (2013). "Biomechanics and functional preconditions for terrestrial lifestyle in basal tetrapods, with special consideration of Tiktaalik roseae". Historical Biology. 25 (2): 167–181. Bibcode:2013HBio...25..167H. doi:10.1080/08912963.2012.755677. S2CID 85407197.
  24. ^ Carroll, Irwin, & Green (2005),[22] cited in[23]
  25. ^ Clack, J.A. (2007). "Devonian climate change, breathing, and the origin of the tetrapod stem group" (PDF). Integrative and Comparative Biology. 47 (4): 1–14. doi:10.1093/icb/icm055. PMID 21672860.[full citation needed]
  26. ^ Clack (2007),[25] cited in[23]
  27. ^ King, H.M.; Shubin, N.H.; Coates, M.I.; Hale, M.E. (2011). "Behavioural evidence for the evolution of walking and bounding before terrestriality in sarcopterygian fishes". Proceedings of the National Academy of Sciences USA. 108 (52): 21146–21151. Bibcode:2011PNAS..10821146K. doi:10.1073/pnas.1118669109. PMC 3248479. PMID 22160688.
  28. ^ Pierce, S.E.; Clack, J.A.; Hutchinson, J.R. (2012). "Three-dimensional limb joint mobility in the early tetrapod Ichthyostega". Nature. 486 (7404): 523–526. Bibcode:2012Natur.486..523P. doi:10.1038/nature11124. PMID 22722854. S2CID 3127857.
  29. ^ King (2011),[27] cited in[28]
  30. ^ Witzmann, F.; Schoch, R. R. (2012). "A megalichthyid sarcopterygian fish from the Lower Permian (Autunian) of the Saar-Nahe Basin, Germany". Geobios. 45 (2): 241–248. doi:10.1016/j.geobios.2011.03.002.
  31. ^ Janvier, Philippe (1 January 1997). "Vertebrata: Animals with backbones". (Version 01 January 1997 (under construction) ed.). The Tree of Life Web Project.
  32. ^ Haaramo, Mikko (2003). "Sarcopterygii". Mikko's Phylogeny Archive. University of Helsinki. Retrieved 4 November 2013.
  33. ^ Swartz, B. (2012). "A marine stem-tetrapod from the Devonian of western North America". PLOS ONE. 7 (3): e33683. Bibcode:2012PLoSO...733683S. doi:10.1371/journal.pone.0033683. PMC 3308997. PMID 22448265.
  34. ^ Choo, Brian; Zhu, Min; Qu, Qingming; Yu, Xiaobo; Jia, Liantao; Zhao, Wenjin (8 March 2017). "A new osteichthyan from the late Silurian of Yunnan, China". PLOS ONE. 12 (3): e0170929. Bibcode:2017PLoSO..1270929C. doi:10.1371/journal.pone.0170929. ISSN 1932-6203. PMC 5342173. PMID 28273081.
  35. ^ "Ancient southern China fish may have evolved prior to the 'Age of Fish'". (Press release). PLoS. March 2017. Archived from the original on 8 March 2017. Retrieved 11 March 2017.