Sarcopterygii

Sarcopterygii
Temporal range: Late Silurian–Recent, 418–0 Ma
Queensland Lungfish
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Subphylum: Vertebrata
Class: Sarcopterygii
Romer, 1955
Subclasses

The Sarcopterygii /ˌsɑrkɒptəˈrɪdʒi.aɪ/ or lobe-finned fishes (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 fishes, though a strict classification would include the terrestrial vertebrates. The living sarcopterygians are the coelacanths, lungfishes, and the tetrapods.

Contents

Characteristics

Early sarcopterygians or crossopterygians are bony fish with fleshy, lobed, paired fins, which are joined to the body by a single bone.[1] The fins of crossopterygians differ from those of all other fish in that each is borne on a fleshy, lobelike, scaly stalk extending from the body. 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 sarcoptergygians 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.

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.[2] The fin-limbs of sarcopterygians such as the coelacanths show a strong similarity to the expected ancestral form of tetrapod limbs. The crossopterygians apparently followed two different lines of development and are accordingly separated into two subclasses, the Rhipidistia (including the Dipnoi, the lungfishes, and the Tetrapodomorpha which include the Tetrapoda) and the Actinistia (coelacanths).

Evolution

Sarcopterygians and their relatives the actinopterygians ("ray-finned fish") comprise the superclass Osteichthyes, the "bony fish", 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. The first 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.[3]

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.[4][5]

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.[6][3]

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).

Classification

Taxonomy

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.[7]

Phylogeny

The cladogram presented below is based on studies compiled by Philippe Janvier and others for the Tree of Life Web Project.[8]

Sarcopterygii

?†Onychodontiformes



Coelacanthimorpha (coelacanths)


unnamed

Porolepimorpha



Dipnoi (lungfish)



unnamed

Rhizodontimorpha


unnamed

Osteolepimorpha



Terrestrial vertebrates





See also

References

  1. ^ Clack, J. A. (2002) Gaining Ground. Indiana University
  2. ^ Nelson, Joseph S. (2006). Fishes of the World. John Wiley & Sons, Inc. ISBN 0-471-25031-7. 
  3. ^ a b "Fish-Tetrapod Transition Got A New Hypothesis In 2011". Science 2.0. December 27, 2011. http://www.science20.com/news_articles/fishtetrapod_transition_got_new_hypothesis_2011-85782. Retrieved January 2, 2012. 
  4. ^ Grzegorz Niedźwiedzki, Piotr Szrek, Katarzyna Narkiewicz, Marek Narkiewicz & Per E. Ahlberg (2010). "Tetrapod trackways from the early Middle Devonian period of Poland". Nature (Nature Publishing Group) 463: 43–48. doi:10.1038/nature08623. http://www.nature.com/nature/journal/v463/n7277/full/nature08623.html. Retrieved January 3, 2012. 
  5. ^ Shanta Barley (January 6, 2010). "Oldest footprints of a four-legged vertebrate discovered". New Scientist. http://www.newscientist.com/article/dn18346-oldest-footprints-of-a-fourlegged-vertebrate-discovered.html. Retrieved January 3, 2010. 
  6. ^ Retallack, Gregory (May 2011). "Woodland Hypothesis for Devonian Tetrapod Evolution". Journal of geology (University of Chicago Press) 119 (3): 235-258. doi:10.1086/659144. 
  7. ^ Benton, M.J. (2004). Vertebrate Paleontology. 3rd ed. Blackwell Science Ltd
  8. ^ Janvier, Philippe. 1997. Vertebrata. Animals with backbones. Version 01 January 1997 (under construction). http://tolweb.org/Vertebrata/14829/1997.01.01 in The Tree of Life Web Project, http://tolweb.org/