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for|the band|Lungfish (band)Taxobox| name = Lungfishes| fossil_range = Fossil range|Early Devonian|Recent| image = Australian-Lungfish.jpg| image_width = 250px| image_caption = Queensland lungfish | regnum = Animal ia| phylum = Chordate|Chordata | subphylum = Vertebrate|Vertebrata | classis = Sarcopterygii | subclassis = Dipnoi | subclassis_authority = Johannes Peter Müller|J. P. Müller , 1844| subdivision_ranks = Orders| subdivision =
Ceratodontiformes
Lepidosireniformes
Lungfish ( common name|also known as salamanderfish cite book |title=The History of Creation, Or, The Development of the Earth and Its Inhabitants by the Action of Natural Causes: A Popular Exposition of the Doctrine of Evolution in General, and of that of Darwin, Goethe, and Lamarck in Particular : from the 8. German Ed. of Ernst Haeckel |author= Ernst Haeckel|Ernst Heinrich Philipp August Haeckel , Edwin Ray Lankester, L. Dora Schmitz |year=1892 |publisher=D. Appleton |pages=422 |url= http://books.google.com/books? id=ltUj8vk3auEC page 289) are freshwater fish belonging to the Subclass (biology)|subclass Dipnoi . Lungfish are best known for retaining characteristics primitive within the Osteichthyes , including the ability to breathe air, and structures primitive within Sarcopterygii , 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 .
Anatomy and morphology
All lungfish demonstrate an uninterrupted cartilaginous notochord and an extensively developed palatal dentition. Lungfish are Omnivore|omnivorous , feeding on fish, insects, crustaceans, worms, molluscs, amphibians and plant matter. They have a spiral valve rather than a true stomach . http://onlinelibrary.wiley.com/doi/10.1002/ar.1091820109/abstract Electron microscopy of the intestine of the african lungfish, Protopterus aethiopicus Basal lungfish groups may retain marginal teeth and an ossified braincase, but derived lungfish groups, including all modern species, show a significant reduction in the marginal bones and a cartilaginous braincase. The bones of the skull roof in primitive lungfish are covered in a mineralized tissues|mineralized tissue called cosmine , but in post- Devonian lungfishes, the skull roof lies beneath the skin and the cosmine covering is lost. All modern lungfish show significant reductions and fusions of the bones of the skull roof, and the specific bones of the skull roof show no Homology (biology)|homology to the skull roof bones of ray-finned fish es or tetrapods . During the breeding season, the South American lungfish develops a pair of feathery appendages that are actually highly modified pelvic fins. These fins are thought to improve gas exchange around the fish's eggs in its nest. Ross Piper|Piper, Ross (2007), Extraordinary Animals: An Encyclopedia of Curious and Unusual Animals , Greenwood Press (publisher)|Greenwood Press .
Through convergent evolution , lungfishes have evolved internal nostrils similar to the tetrapods' choana . http://scienceweek.com/2004/sa041224-3.htm Evolution: On the evolution of internal nostrils (choanae)
The dentition of lungfish is different from that of any other vertebrate group. " Odontode s" on the palate and lower jaws develop in a series of rows to form a fan-shaped Occlusion (dentistry)|occlusion surface. These odontodes then wear to form a uniform crushing surface. In several groups, including the modern lepidosireniformes , these ridges have been modified to form occluding blades.
The modern lungfishes have a number of larval features, which suggest paedomorphosis . They also demonstrate the largest genome among the vertebrates.
Modern lungfish all have an elongate body with fleshy, paired pectoral fin|pectoral and pelvic fin s and a single unpaired caudal fin replacing the dorsal, caudal and anal fins of most fishes.
Lungs
All lungfish have two lungs, with the exception of the Australian lungfish, which only has one. The lung(s) connect to the pharynx . The lungs of lungfish are homology (biology)|homologous to the lungs of tetrapod s. As in tetrapods and bichir s, the lungs extend from the ventral surface of the esophagus and gut. http://cwx.prenhall.com/bookbind/pubbooks/martini10/chapter24/custom2/deluxe-content.html Chapter 24: The Respiratory System Evolution Atlas http://people.biology.ufl.edu/sahilber/VertZooLab2007/Lab2.htm LAB 2 - GNATHOSTOME FORM & FUNCTION
While other species of fish can breathe air via modified, vascularized gas bladder s,citation|title=Did lungs and the intracardiac shunt evolve to oxygenate the heart in vertebrates|author=Colleen Farmer|journal=Paleobiology|year=1997|url= http://www.biology.utah.edu/farmer/publications%20pdf/1997%20Paleobiology23.pdf these bladders are usually simple sacs, devoid of complex internal structure. In contrast, the lungs of lungfish are subdivided into numerous smaller air sacs, maximizing the surface area available for gas exchange.
Perfusion of water
Of extant lungfish, only the Australian lungfish can respire through its gills. In other species, the gills are too atrophied to allow for adequate gas exchange . When a lungfish is obtaining oxygen from its gills, its circulatory system is configured similarly to the common fish. The spiral valve of the conus arteriosus is open, the bypass arterioles of the third and fourth gill arches (which do not actually have gills) are shut, the second, fifth and sixth gill arch arterioles are open, the ductus arteriosus branching off the sixth arteriole is open, and the pulmonary arteries are closed. As the water passes through the gills, the lungfish uses a buccal pump. Flow through the mouth and gills is unidirectional. Blood flow through the secondary lamellae is countercurrent to the water, maintaining a more constant concentration gradient.
Perfusion of air
When breathing air, the spiral valve of the conus arteriosus closes (minimizing the mixing of oxygenated and deoxygenated blood), the third and fourth gill arches open, the second and fifth gill arches close (minimizing the possible loss of the oxygen obtained in the lungs through the gills), the sixth arteriole's ductus arteriosus is closed, and the pulmonary arteries open. Importantly, during air breathing, the sixth gill is still used in respiration; deoxygenated blood loses some of its carbon dioxide as it passes though the gill before reaching the lung. This is because carbon dioxide is more soluble in water. Air flow through the mouth is tidal, and through the lungs it is bidirectional and observes "uniform pool" diffusion of oxygen.
Ecology and life history
African and South American lungfish are capable of surviving seasonal drying out of their habitats by burrowing into mud and estivation|estivating throughout the dry season. Changes in physiology allow it to slow its metabolism to as little as 1/60th of the normal metabolic rate, and protein waste is converted from ammonia to less-toxic urea (normally, lungfish excrete nitrogenous waste as ammonia directly into the water).
Burrowing is seen in at least one group of fossil lungfish, the Gnathorhizidae . It has been proposedBy whom|date=March 2009 both that burrowing is plesiomorphy|plesiomorphic for lungfish, and that gnathorhizids are directly ancestral to modern Lepidosireniformes, but the similarity possibly is simply due to convergent evolution|convergent or parallel evolution .
Lungfish can be extremely long-lived. The Queensland lungfish at the Shedd Aquarium in Chicago has been part of the permanent live collection since 1933. http://www.sheddaquarium.org/granddad.html
Taxonomy
The relationship of lungfishes to the rest of the Osteichthyes|bony fish is well understood:
Lungfishes are most closely related to Powichthyes , and then to the Porolepiformes .
Together, these taxa form the Dipnomorpha , the sister group to the Tetrapodomorpha .
Together, these form the Rhipidistia , the sister group to the Coelacanth s.
The relationships among lungfishes are significantly more difficult to resolve. While Devonian lungfish had enough bone in the skull to determine relationships, post-Devonian lungfish are represented entirely by skull roofs and teeth, as the rest of the skull is cartilage|cartilaginous . Additionally, many of the taxa already identified may not be monophyletic . Current phylogenetic studies support the following relationships of major lungfish taxa:
Ahlberg, PE, Smith, MM and Johanson, Z, (2006). Developmental plasticity and disparity in early dipnoan (lungfish) dentitions. Evolution and Development 8(4):331-349.
Palmer, Douglas, Ed. The Simon & Schuster Encyclopedia of Dinosaurs & Prehistoric Creatures. A Visual Who's Who of Prehistoric Life. Pg. 45. Great Britain: Marshall Editions Developments Limited. 1999.
Schultze, HP, and Chorn, J., (1997). The Permo-Herbivorus genus Sagenodus and the beginning of modern lungfish. Contributions to Zoology 61(7):9-70.
cite journal | last = Sepkoski | first = Jack | title = A compendium of fossil marine animal genera | journal = Bulletins of American Paleontology | volume = 364 | pages = p.560 | year = 2002 | url = http://strata.ummp.lsa.umich.edu/jack/showgenera.php? taxon=611& rank=class | accessdate = 2011-05-17
