lungfish

(Lungfishes)

Class: Sarcopterygii

Order: Lepidosireniformes

Number of families: 2

Evolution and systematics

The South American lungfish (Lepidosiren paradoxa) was the first lungfish species to be described (in 1837 by Leopold J. F. J. Fitzinger; 1802–1884). It was originally discovered and collected from the Amazon River by Johann Natterer (1787–1843), a dedicated and gifted Austrian naturalist who collected extensively in Brazil from 1817–1835. The first African lungfish was described shortly thereafter in 1839 by the British anatomist Richard Owen (1810–1890), who firmly believed that lungfishes were true "fishes" and not amphibians, mostly on the basis of his erroneous conviction that lungfishes did not have a second (internal) nostril (known as a choana) or a divided auricle. Zoologists of the mid-nineteenth century were divided about the ancestry of lungfishes, wondering whether they were more closely related to amphibians or to other bony fishes. It is now well established that lungfishes belong to a higher group, the Sarcopterygii, which also includes the tetrapods, coelacanths, and many other lobe-finned fossil fishes, and are therefore unequivocally more closely related to land vertebrates than to ray-finned bony fishes (Actinopterygii).

The genus Lepidosiren is monotypic, but detailed comparative studies of specimens from most of its extended range are needed (two other nominal species exist, but they are considered synonyms of L. paradoxa). Four species of African lung-fishes (Protopterus) are recognized: P. annectens, P. aethiopicus, P. dolloi, and P. amphibius. Some of these species are difficult to distinguish, and are in need of critical taxonomic evaluation, including the validity of certain subspecies (a total of 10 nominal species have been described); their evolutionary relationships to each other have yet to be fully investigated. Lepidosiren and Protopterus are placed in the same order, but are classified in distinct families. They are closely related to the Australian lungfish (Neoceratodus forsteri) and its immediate fossil relatives, while the vast majority of remaining fossil lungfishes are more distantly related (some 60 genera and 280 species and of lungfishes are known). Extant lungfishes are "living fossils," and belong to an ancient lineage, the Dipnoi, that was much more diverse in the Devonian (ca. 417–354 million years ago) and Triassic (ca. 248–205 million years ago) periods. Fossil relatives of Lepidosiren and Protopterus are known from the late Cretaceous of South America and Africa, respectively, and these genera, along with Neoceratodus, are among the oldest vertebrates living today.

Physical characteristics

South American and African lungfishes are morphologically similar, presenting elongated, eel-like bodies, with relatively small heads, and filamentous pectoral and pelvic fins. The pelvic fins are stouter than the pectorals in both genera; the pectorals are slightly more robust in Protopterus, and may resemble simple filaments in Lepidosiren. The caudal fin is confluent with the dorsal and anal fins (as in the Australian lung-fish), tapering distally. The body is compressed laterally, especially at the anus, but not as much as in Neoceratodus. The eyes are minute and the mouth is terminal, with a lateral groove extending to the sides of the head. Sensory canals on the head and cheek appear as sinuous, deep lines that extend posteriorly at midbody height towards the tail; sensory pores are also present on the head. The anus is asymmetrical, situated laterally just posterior to the pelvic fins, and not directly in the middle as in Neoceratodus (the side may vary among individuals of both Protopterus and Lepidosiren). The scales are mostly embedded in the skin and are very thin, but clearly visible. The nostrils are on the internal lip margin, and the teeth are fused into sharp tooth plates.

Both Lepidosiren and Protopterus have two highly vascularized and separated lungs (modified swim bladders), positioned on each side of the gut and connected to the esophagus ventrally, as in tetrapods and bichirs (Polypteriformes). The lungs have many alveoli, similar to the lungs of tetrapods. The gill openings are small, not nearly as large as in Neoceratodus. Newly hatched individuals have flared external gill filaments, absent in Neoceratodus, and these may persist vestigially above the pectoral fins in subadults and adults of Protopterus. The skeleton is mostly cartilaginous. Both Protopterus and Lepidosiren vary slightly in color, from dark brown to deep gray dorsally and laterally, with many varied blotches and spots; usually dark ventrally, although Protopterus may be lighter ventrally. Lepidosiren may reach 4.1 ft (1.25 m) in length, while Protopterus varies from between 17.7 in (45 cm) (P. amphibius) to 6.5 ft (2 m) in length (P. aethiopicus, which can weigh some 37.5 lb [17 kg]).

Distribution

Lepidosiren has the greatest distribution of any extant lung-fish, occurring in many tributaries of the Amazon and Paraná-Paraguay River systems, as well as in French Guiana. Species of Protopterus are slightly more restricted. P. annectens is present in central and West Africa; P. aethiopicus occurs in central and East Africa; P. dolloi is restricted to the Congo basin; and P. amphibius occurs in coastal East Africa.

Habitat

The South American and African lungfishes are generally found in lentic (slow-moving) rivers, with plenty of associated vegetation and swampy, stagnant conditions (especially L. paradoxa). They can also be found in open lakes (e.g., P. aethiopicus in Lake Victoria); floodplains (e.g., P. dolloi in the Congo River basin, and P. annectens in the Senegal, Gambia, Niger, and Volta Rivers in West Africa); near river deltas (P. aethiopicus in Lake Tanganyika, P. amphibius in the Zambesi River delta); and in small pools.

Behavior

Lepidosiren and Protopterus species are sluggish, swimming through sinuous movements or by "crawling" on their pectoral and pelvic fins, especially to scavenge the bottom. Both genera are obligate air breathers, unlike Neoceratodus, which breathes primarily through the gills. Lepidosiren and Protopterus individuals will drown if forced to stay underwater, as the gill surfaces of these fishes are not large enough to satisfy their oxygen needs. Both genera also employ estivation, being capable of remaining inside a resting chamber for protracted periods during dry seasons and emerging when wet conditions return (estivation has been documented for Permian lungfishes, in the form of fossilized burrows). The degree of estivation varies among the species, but has been particularly well documented for P. annectens. The burrows are excavated by biting the soil and expelling mud through the gill openings. The fish will then turn around and remain with its head facing the burrow opening, from where it obtains oxygen. The individual suffers metabolic changes during this period to endure the lack of moisture (detailed below for P. annectens). One individual of P. aethiopicus remained in its cocoon for four years in captivity. Lungfishes do not feed during estivation. To sustain themselves, they initially metabolize fat reserves and then muscle mass.

Feeding ecology and diet

Lungfishes are mostly carnivorous, feeding mainly on invertebrates (insects, insect larvae, mollusks, crustaceans) but also on fishes and amphibians. Both genera may occasionally feed on aquatic plants. Lungfishes approach potential prey items through ambush or stalking, capturing them by quickly opening their mouths to create negative pressure that pulls them in. Little is known concerning their natural predators, but presumably larger carnivorous fishes and other vertebrates prey on lungfishes, especially when they are juveniles.

Reproductive biology

Spawning is usually seasonal, taking place during the wet season. Fertilization is external. In both genera the adult male guards and aerates the hatchlings and young temporarily. Female Protopterus usually lay eggs in burrows excavated by the males. The eggs are small (from 0.16–0.27 in/4–7 mm in diameter), and take one to two weeks to hatch, at which time they resemble tadpoles with slender, featherlike external gills. Only after a period from one month to 55 days do the larvae breathe air. At this stage, they range from 1 in (2.5 cm) to 1.6 in (4 cm) in length, and still have external gills. The larvae remain relatively inactive and are attached to the nest through their cement glands until their yolk reserves have been depleted, at which time they begin to forage for insect larvae and crustaceans and inhale air.

Conservation status

No species of Lepidosireniformes are listed by the IUCN.

Significance to humans

Lungfishes are common in both public and private aquaria. Although they are consumed as food in some parts of Africa, they are not important food fishes. They are harmless, but if provoked can inflict painful bites because of their strong jaws and sharp teeth.

Species accounts

Resources

Bemis, W. E., W. W. Burggren, and N. E. Kemp. The Biology and Evolution of Lungfishes. New York: A. R. Liss, 1987.

Berra, T. M. Freshwater Fish Distribution. San Diego: Academic Press, 2001.

Britski, H. A., K. Z. S. de Silimon, and B. S. Lopes. Peixes do Pantanal, Manual de Identificação. Brasília: Embrapa, 1999.

Cloutier, R, and P. E. Ahlberg. "Morphology, Characters, and the Interrelationships of Basal Sarcopterygians." In Interrelationships of Fishes, edited by M. L. J. Stiassny, L. Parenti, and G. D. Johnson. San Diego: Academic Press, 1996.

Conant, E. B. "Bibliography of Lungfishes, 1811–1985." In The Biology and Evolution of Lungfishes, edited by W. E. Bemis, W. W. Burggren, and N. E. Kemp. New York: A. R. Liss, 1987.

Gosse, J. P. "Protopteridae." In Check-List of the Freshwater Fishes of Africa (CLOFFA), edited by J. Daget, J. P. Gosse, and D. F. E. Thys van den Audenaerde. Paris: ORSTOM; Tervuren: MRAC, 1984.

Graham, J. B. Air-Breathing Fishes. San Diego: Academic Press, 1997.

Greenwood, P. H. "The Natural History of African Lungfishes." In The Biology and Evolution of Lungfishes, edited by W. E. Bemis, W. W. Burggren, and N. E. Kemp. New York: A. R. Liss, 1987.

Janvier, P. Early Vertebrates. Oxford: Oxford University Press, 1996.

Lévêque, C. "Protopteridae." In Faune des poissons d'eaux douces et saumâtres d'Afrique de l'Ouest. Tome 1, edited by C. Lévêque, D. Paugy, and G. G. Teugels. Paris: ORSTOM, 1990.

Merrick, J. R., and G. E. Schmida. Australian Freshwater Fishes, Biology and Management. North Ryde, Australia: Macquarie University, 1984.

Nelson, J. S. Fishes of the World, 3rd edition. New York: John Wiley & Sons, 1994.

Planquette, P., P. Keith, and P. Y. LeBail. Atlas des Poissons d'Eau Douce de Guyane, Tome 1. Paris: Museum National d'Histoire Naturelle, 1996.

Skelton, P. Freshwater Fishes of Southern Africa, 2nd edition. Cape Town: Struik, 2001.

Atz, J. W. Narial. "Breathing in Fishes and the Evolution of Internal Nares." Quarterly Review of Biology 27, no. 4 (1952): 366–377.

Bemis, W. E. "Morphology and Growth of Lepidosirenid Lungfish Tooth Plates (Pisces: Dipnoi)." Journal of Morphology 179 (1984): 73–93. ——. "Paedomorphosis and the Evolution of the Dipnoi." Paleobiology 10, no. 3 (1984): 293–307.

Bertmar, G. "The Olfactory Organ and the Upper Lips in Dipnoi, a Comparative Embryological Study." Acta Zoologica 46 (1965): 1–40.

Carter, G. S., and L. C. Beadle. "Notes on the Habitat and Development of Lepidosiren paradoxa." Journal of the Linnaean Society, Zoology 37 (1930): 197–203.

Coates, C. W. "Slowly the Lungfish Gives Up Its Secrets." Bulletin of the New York Zoological Society 40 (1937): 25–34.

Cunningham, J. T., and D. M. Reid. "Pelvic Filaments of Lepidosiren." Nature 131 (1933): 913.

Dollo, L. "Sur la phylogénie des dipneustes." Bull. Soc. Belge Geol., Paleont. Hydrologie. 9, no. 2 (1896): 79–128.

Johnels, A. G., and G. S. O. Svensson. "On the Biology of Protopterus annectens (Owen)." Ark. Zool. Stockholm 7, no. 7(1954): 131–164.

Littrell, L. "African Lungfishes." Tropical Fish Hobbyist 19, no. 8 (1971): 40–57.

McMahon, B. R. "A Functional Analysis of the Aquatic and Aerial Respiratory Movements of an African Lungfish, Protopterus aethiopicus, with Reference to the Evolution of the Lung-ventilation Mechanism in Vertebrates." Journal of Experimental Biology 51, no. 2 (1969): 407–430.

Miles, R. S. "Dipnoan (Lungfish) Skulls and the Relationships of the Group: A Study Based on New Specimens from the Devonian of Australia." Zoological Journal of the Linnaean Society 61 (1977): 1–328.

Poll, M. "Revision systématique et raciation géographique des Protopteridae de lÁfrique centrale." Ann. Mus. R. Afr. Cent., Zool., 8, no. 103 (1961): 1–50, pls. 1–6.

Rosen, D. E., P. L. Forey, B. G. Gardiner, and C. Patterson. "Lungfishes, Tetrapods, Paleontology, and Plesiomorphy." Bulletin of the American Museum of Natural History 167, no. 4(1981): 163–275.

"FishBase" [cited January 15, 2003]. "Catalog of Fishes On-Line" [cited January 15, 2003].

"Palæos: Vertebrates" [cited January 15, 2003].

[Article by: Marcelo Carvalho, PhD]

The noun has one meaning:

Meaning #1: air-breathing fish having an elongated body and fleshy paired fins; certain species construct mucus-lined mud coverings in which to survive drought

For the band, see Lungfish (band).

Lungfishes Fossil range: Early Devonian–Recent PreЄ Є O S D C P T J K Pg N
Queensland Lungfish
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Subphylum:	Vertebrata
Class:	Sarcopterygii
Subclass:	Dipnoi Müller, 1844
Orders
See text.

Lungfish (also known as salamanderfish^[1]) are freshwater fish belonging to the 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, they 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 that advanced lungfish had a widespread freshwater distribution and that the current distribution of modern lungfish species reflects extinction of many lineages following the breakup of Pangaea, Gondwana, and Laurasia.

Contents 1 Anatomy and morphology 2 Lungs 3 Ecology and life history 3.1 Behavior 4 Taxonomy 5 See also 6 References 7 External links

Anatomy and morphology

All lungfish demonstrate an uninterrupted cartilaginous notochord and an extensively developed palatal dentition. The lungfish is a true carnivore. 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 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 to the skull roof bones of ray-finned fishes or tetrapods. During the breeding season, the South American lungfish develops a pair of feathery appendages that are actually highly modified pelvic fins. It is thought these fins improve gas exchange around the fish's eggs in its nest.^[2]

The dentition of lungfish is different from that of any other vertebrate group. "Odontodes" on the palate and lower jaws develop in a series of rows to form a fan-shaped 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 and pelvic fins and a single unpaired caudal fin replacing the dorsal, caudal, and anal fin of most fishes.

Lungs

Lateral view of lungs of a dissected Protopterus dolloi

All lungfish have two lungs (except for the Australian, which has one) that connect to the pharynx. While other species of fish can breathe air via modified, vascularized gas bladders, these bladders are usually simple sacs, devoid of complex internal structure. In contrast, the lungs of lungfish are subdivided into numerous smaller air sacs, greatly increasing surface area for improved gas exchange. Furthermore, while the aforementioned vascularized swim bladders have arisen independently in several lineages of fish, only in the lungfish are they homologous to the lungs of tetrapods.

Ecology and life history

Behavior

African and South American lungfish are capable of surviving seasonal drying out of their habitats by burrowing into mud and estivating throughout the dry season. Changes in physiology allow the lungfish 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 proposed^{[by whom?]} both that burrowing is plesiomorphic for lungfish as well as that gnathorhizids are directly ancestral to modern Lepidosireniformes, but it is possible that the similarity is simply due to 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.[1]

Taxonomy

The relationship of lungfishes to the rest of the 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 coelacanths.

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 cartilaginous. Additionally, many of the taxa that have been identified may not be monophyletic. Current phylogenetic studies support the following relationships of major lungfish taxa:

Class Osteichthyes
Subclass Sarcopterygii
Order Dipnoi

,--†Family Diabolichthyidae
| ,--†Family Uranolophidae
| |  __,--†Family Speonesydrionidae
'-|-|  '--†Family Dipnorhynchidae
    |     ,--†Family Stomiahykidae
    '----|___ ,--†Family Chirodipteridae
          |      '-|--†Family Holodontidae
          |------†Family Dipteridae
          |  __,--†Family Fleurantiidae
          '-|  '--†Family Rhynchodipteridae
              '--†Family Phaneropleuridae
                     | ,--†Family Ctenodontidae
                       '-| ,--†Family Sagenodontidae
                          '-|--†Family Gnathorhizidae
                             '--Order Ceratodontiformes
                                  |--†family Asiatoceratodontidae
                                  |--†Family Ptychoceratodontidae
                                  |--Family Ceratodontidae
                                  |  '--†Genus Ceratodus
                                  |  '--†Genus Metaceratodus
                                   '--Family Neoceratodontidae
                                        | '--†Genus Mioceratodus
                                        | '--Genus Neoceratodus - Queensland lungfish
                                        '--Order Lepidosireniformes
                                               '--Family Lepidosirenidae - South American lungfish 
                                               '--Family Protopteridae - African lungfish

See also

• Lepidogalaxias salamandroides

References

This article includes a list of references, related reading or external links, but its sources remain unclear because it lacks inline citations. Please improve this article by introducing more precise citations where appropriate. (January 2008)

1. ^ Ernst Heinrich Philipp August Haeckel, Edwin Ray Lankester, L. Dora Schmitz (1892). 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. D. Appleton. pp. 422. http://books.google.com/books?id=ltUj8vk3auEC.  page 289
2. ^ Piper, Ross (2007), Extraordinary Animals: An Encyclopedia of Curious and Unusual Animals, Greenwood Press.

• 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 Cretures. 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.

External links

Wikispecies has information related to: Dipnoi

The Wikibook Dichotomous Key has a page on the topic of Dipnoi

• Dr Anne Kemps - Lungfish Information site
• Lungfish information site
• Dipnoiformes at Palaeos.com
• Dipnoi at the University of California Museum of Paleontology
• Tree of life illustration showing lungfish's relation to other organisms

v • d • e Extant sarcopterygii orders by subclass Kingdom Animalia · Phylum Chordata · Subphylum Vertebrata · Infraphylum Gnathostomata · Superclass Osteichthyes Coelacanthimorpha Coelacanthiformes Dipnoi Ceratodontiformes · Lepidosireniformes

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