Lungfish are freshwater vertebrates belonging to the

order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...

Dipnoi. Lungfish are best known for retaining ancestral characteristics within the

Osteichthyes Osteichthyes (), popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartilage ...

, including the ability to breathe air, and ancestral structures within Sarcopterygii, including the presence of lobed fins with a well-developed internal skeleton. Lungfish represent the closest living relatives of the tetrapods. Today there are only six known species of lungfish, living in Africa, South America, and

Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...

. The fossil record shows that lungfish were abundant since the Triassic. While

vicariance Allopatric speciation () – also referred to as geographic speciation, vicariant speciation, or its earlier name the dumbbell model – is a mode of speciation that occurs when biological populations become geographically isolated from ...

would suggest this represents an ancient distribution limited to the Mesozoic supercontinent

Gondwana Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final stages ...

, the fossil record suggests advanced lungfish had a widespread freshwater distribution and the current distribution of modern lungfish species reflects extinction of many lineages subsequent to the breakup of Pangaea,

and

Laurasia Laurasia () was the more northern of two large landmasses that formed part of the Pangaea supercontinent from around ( Mya), the other being Gondwana. It separated from Gondwana (beginning in the late Triassic period) during the breakup of Pan ...

. Lungfish have historically been referred to as salamanderfish, but this term more often refers to '' Lepidogalaxias salamandroides''.

Anatomy and morphology

All lungfish demonstrate an uninterrupted cartilaginous

notochord In anatomy, the notochord is a flexible rod which is similar in structure to the stiffer cartilage. If a species has a notochord at any stage of its life cycle (along with 4 other features), it is, by definition, a chordate. The notochord consis ...

and an extensively developed palatal dentition.

Basal Basal or basilar is a term meaning ''base'', ''bottom'', or ''minimum''. Science * Basal (anatomy), an anatomical term of location for features associated with the base of an organism or structure * Basal (medicine), a minimal level that is nec ...

primitive Primitive may refer to: Mathematics * Primitive element (field theory) * Primitive element (finite field) * Primitive cell (crystallography) * Primitive notion, axiomatic systems * Primitive polynomial (disambiguation), one of two concepts * Pr ...

") 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 The skull roof, or the roofing bones of the skull, are a set of bones covering the brain, eyes and nostrils in bony fishes and all land-living vertebrates. The bones are derived from dermal bone and are part of the dermatocranium. In comparati ...

in primitive lungfish are covered in a mineralized tissue called cosmine, but in post-

Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...

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 Homology may refer to: Sciences Biology *Homology (biology), any characteristic of biological organisms that is derived from a common ancestor * Sequence homology, biological homology between DNA, RNA, or protein sequences *Homologous chrom ...

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. These fins are thought to improve gas exchange around the fish's eggs in its nest. Through convergent evolution, lungfishes have evolved internal nostrils similar to the tetrapods' choana, and a brain with certain similarities to the

Lissamphibia The Lissamphibia is a group of tetrapods that includes all modern amphibians. Lissamphibians consist of three living groups: the Salientia (frogs, toads, and their extinct relatives), the Caudata (salamanders, newts, and their extinct relatives), ...

n brain (except for the Queensland lungfish, which branched off in its own direction about 277 million years ago and has a brain resembling that of the '' latimeria''). 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 Lev Berg Lev Semyonovich Berg, also known as Leo S. Berg (russian: Лев Семёнович Берг; 14 March 1876 – 24 December 1950) was a leading Russian geographer, biologist and ichthyologist who served as President of the Soviet Geogra ...

, 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 fins of most fishes.

Lungs

Lungfish have a highly specialized respiratory system. They have a distinct feature in that their lungs are connected to the larynx and pharynx without a trachea. While other species of fish can breathe air using modified, vascularized

gas bladder The swim bladder, gas bladder, fish maw, or air bladder is an internal gas-filled organ that contributes to the ability of many bony fish (but not cartilaginous fish) to control their buoyancy, and thus to stay at their current water depth w ...

s, 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. Most extant lungfish species have two lungs, with the exception of the Australian lungfish, which has only one. The lungs of lungfish are

homologous Homology may refer to: Sciences Biology *Homology (biology), any characteristic of biological organisms that is derived from a common ancestor *Sequence homology, biological homology between DNA, RNA, or protein sequences * Homologous chrom ...

to the lungs of tetrapods. As in tetrapods and bichirs, the lungs extend from the ventral surface of the esophagus and gut.

Perfusion of water

Of extant lungfish, only the Australian lungfish can breathe through its gills without needing air from its lungs. 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 through 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

Lungfish are omnivorous, feeding on fish, insects, crustaceans, worms, mollusks,

amphibian Amphibians are tetrapod, four-limbed and ectothermic vertebrates of the Class (biology), class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terres ...

s and plant matter. They have an

intestinal spiral valve A spiral valve or scroll valve is the corkscrew-shaped lower portion of the intestine of some sharks, Acipenseriformes ( sturgeon and paddlefish), rays, skates, bichirs, Lepisosteiformes (gars), and lungfishes. A modification of the ileum, th ...

rather than a true stomach. 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 it to slow its metabolism to as little as one sixtieth 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. Lungfish can be extremely long-lived. A Queensland lungfish at the Shedd Aquarium in Chicago was part of the permanent live collection from 1933 to 2017 after a previous residence at the Sydney Aquarium; at perhaps 90 years old, it was euthanized following a decline in health consistent with old age. A primitive Australian fish living in a San Francisco museum is believed to be the oldest living aquarium fish in the world. Methuselah is a four-foot-long (1.2-meter) Australian lungfish, weighing around 40 lb (18.1 kg).

Evolution

About 420 million years ago, during the

, the last common ancestor of both lungfish and the tetrapods split into two separate evolutionary linages, with the ancestor of the extant coelacanths diverging a little earlier from a sarcopterygian progenitor. The fossil of Youngolepis from about 415 million years ago already show significant modification of their jaws and jaw muscles towards the specialized lungfish feeding mechanism. '' Diabolepis'', a 400 million year old lungfish, is the oldest known unmistakingly durophagous vertebrate, and shows that the lungfishes had adapted to a diet including hard-shelled prey very early in their evolution. The transition could have taken as little as 7 million years. It is worth noting that, despite the name "lungfish", fish evolved lungs before lungfish, or even lobe-finned fish. The common ancestor of lobe-finned and ray-finned fish had lungs, but in most surviving branches of ray-finned fish these evolved into

swim bladder The swim bladder, gas bladder, fish maw, or air bladder is an internal gas-filled Organ (anatomy), organ that contributes to the ability of many bony fish (but not cartilaginous fish) to control their buoyancy, and thus to stay at their curren ...

s used for flotation, instead of breathing. Some, like the bichirs, do retain their lungs, and several other traits that appear to have been common to lobe-finned and ray-finned fish. While the coelacanth shares some traits with the tetrapods (reptiles, amphibians, birds and mammals), the lungfish have other tetrapod traits which are absent in the coelacanth. Evidence suggests that the tetrapods are related more closely to lungfish than to coelacanths.

Extant lungfish

The Queensland lungfish, ''Neoceratodus forsteri'', is endemic to Australia. Fossil records of this group date back 380 million years, around the time when the higher vertebrate classes were beginning to evolve. Fossils of lungfish belonging to the genus ''Neoceratodus'' have been uncovered in northern New South Wales, indicating that the Queensland lungfish has existed in Australia for at least 100 million years, making it a living fossil and one of the oldest living vertebrate genera on the planet. It is the most primitive surviving member of the ancient air-breathing lungfish (Dipnoi) lineages. The five other freshwater lungfish species, four in Africa and

one 1 (one, unit, unity) is a number representing a single or the only entity. 1 is also a numerical digit and represents a single unit of counting or measurement. For example, a line segment of ''unit length'' is a line segment of length 1. I ...

in South America, are very different morphologically to ''N. forsteri''. The Queensland lungfish can live for several days out of the water if it is kept moist, but will not survive total water depletion, unlike its African counterparts. F de Castelnau-poissonsPl50

The South American lungfish, ''Lepidosiren paradoxa'', is the single species of lungfish found in

swamp A swamp is a forested wetland.Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. 497 p. Swamps are considered to be transition zones because both land and water play a role in ...

s and slow-moving waters of the Amazon, Paraguay, and lower

Paraná River The Paraná River ( es, Río Paraná, links=no , pt, Rio Paraná, gn, Ysyry Parana) is a river in south-central South America, running through Brazil, Paraguay, and Argentina for some ."Parana River". Encyclopædia Britannica. Encyclopædia Br ...

basins in South America. Notable as an

obligate {{wiktionary, obligate As an adjective, obligate means "by necessity" (antonym ''facultative'') and is used mainly in biology in phrases such as: * Obligate aerobe, an organism that cannot survive without oxygen * Obligate anaerobe, an organism that ...

air-breather, it is the sole member of its family native to the Americas. Relatively little is known about the South American lungfish, or ''scaly salamander-fish''. When immature it is spotted with gold on a black background. In the adult this fades to a brown or gray color. Its tooth-bearing

premaxillary The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has ...

and maxillary bones are fused like other lungfish. South American lungfishes also share an autostylic jaw suspension (where the palatoquadrate is fused to the

cranium The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...

) and powerful adductor jaw muscles with the extant lungfish (Dipnoi). Like the African lungfishes, this species has an elongate, almost eel-like body. It may reach a length of . The pectoral fins are thin and threadlike, while the pelvic fins are somewhat larger, and set far back. The fins are connected to the shoulder by a single bone, which is a marked difference from most fish, whose fins usually have at least four bones at their base; and a marked similarity with nearly all land-dwelling vertebrates. The gills are greatly reduced and essentially non-functional in the adults. Marbled lungfish 1

The marbled lungfish, ''Protopterus aethiopicus'', is found in Africa. The marbled lungfish is smooth, elongated, and cylindrical with deeply embedded scales. The tail is very long and tapers at the end. They are the largest of the African lungfish species as they can reach a length of up to 200 cm.Fishbase.org
/ref> The pectoral and pelvic fins are also very long and thin, almost spaghetti-like. The newly hatched young have branched external gills much like those of newts. After 2 to 3 months the young transform (called

metamorphosis Metamorphosis is a biological process by which an animal physically develops including birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation. Some inse ...

) into the adult form, losing the external gills for gill openings. These fish have a yellowish gray or pinkish toned ground color with dark slate-gray splotches, creating a marbling or leopard effect over the body and fins. The color pattern is darker along the top and lighter below. The marbled lungfish's genome contains 133 billion

base pairs A base pair (bp) is a fundamental unit of double-stranded nucleic acids consisting of two nucleobases bound to each other by hydrogen bonds. They form the building blocks of the DNA double helix and contribute to the folded structure of both DNA ...

, making it the largest known genome of any vertebrate. The only organisms known to have more base pairs are the protist '' Polychaos dubium'' and the flowering plant ''

Paris japonica is a Japanese species of plants in the genus ''Paris'' in the family Melanthiaceae.Ohwi, J. (1984). Flora of Japan (in English): 1-1067. Smithsonian Institution, Washington, D.C.. It is native to sub-alpine regions of Japan. A slow growing per ...

'' at 670 billion and 150 billion, respectively.

The gilled lungfish, ''Protopterus amphibius'' is a species of lungfish found in

East Africa East Africa, Eastern Africa, or East of Africa, is the eastern subregion of the African continent. In the United Nations Statistics Division scheme of geographic regions, 10-11-(16*) territories make up Eastern Africa: Due to the historical ...

. It generally reaches only long, making it the smallest

extant Extant is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Extant taxon, a taxon which is not extinct, ...

lungfish in the world. This lungfish is uniform blue, or slate grey in colour. It has small or inconspicuous black spots, and a pale grey belly. LepidosirenFord

The west African lungfish ''Protopterus annectens'' is a species of lungfish found in West Africa. It has a prominent

snout A snout is the protruding portion of an animal's face, consisting of its nose, mouth, and jaw. In many animals, the structure is called a muzzle, rostrum, or proboscis. The wet furless surface around the nostrils of the nose of many mammals is c ...

and small

eye Eyes are organs of the visual system. They provide living organisms with vision, the ability to receive and process visual detail, as well as enabling several photo response functions that are independent of vision. Eyes detect light and conv ...

s. Its body is long and eel-like, some 9-15 times the length of the head. It has two pairs of long, filamentous fins. The pectoral fins have a basal fringe and are about three times the head length, while its

pelvic fin Pelvic fins or ventral fins are paired fins located on the ventral surface of fish. The paired pelvic fins are homologous to the hindlimbs of tetrapods. Structure and function Structure In actinopterygians, the pelvic fin consists of two en ...

s are about twice the head length. In general, three external gills are inserted posterior to the gill slits and above the pectoral fins. It has cycloid scales embedded in the skin. There are 40-50 scales between the operculum and the

anus The anus (Latin, 'ring' or 'circle') is an opening at the opposite end of an animal's digestive tract from the mouth. Its function is to control the expulsion of feces, the residual semi-solid waste that remains after food digestion, which, d ...

and 36-40 around the body before the origin of the dorsal fin. It has 34-37 pairs of

rib In vertebrate anatomy, ribs ( la, costae) are the long curved bones which form the rib cage, part of the axial skeleton. In most tetrapods, ribs surround the chest, enabling the lungs to expand and thus facilitate breathing by expanding the ches ...

s. The dorsal side is olive or brown in color and the ventral side is lighter, with great blackish or brownish spots on the body and fins except on its belly. They reach a length of about 100 cm in the wild . Protopterus dolloi Boulenger2

The spotted lungfish, ''Protopterus dolloi'', is a species of lungfish found in Africa. Specifically, it is found in the Kouilou-Niari Basin of the

Republic of the Congo The Republic of the Congo (french: République du Congo, ln, Republíki ya Kongó), also known as Congo-Brazzaville, the Congo Republic or simply either Congo or the Congo, is a country located in the western coast of Central Africa to the w ...

and Ogowe River basin in Gabon. It is also found in the lower and Middle Congo River Basins.Fishbase.org
/ref> ''Protopterus dolloi'' can

aestivate Aestivation ( la, aestas (summer); also spelled estivation in American English) is a state of animal dormancy, similar to hibernation, although taking place in the summer rather than the winter. Aestivation is characterized by inactivity and a ...

on land by surrounding itself in a layer of dried mucus. It can reach a length of up to 130 cm.

Taxonomy

The relationship of lungfishes to the rest of the

bony fish Osteichthyes (), popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartilag ...

is well understood: * Lungfishes are most closely related to ''

Powichthys ''Powichthys'' is a genus of prehistoric lobe-finned fish which lived during the Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ...

'', and then to the

Porolepiformes Porolepiformes is an order of prehistoric lobe-finned fish which lived during the Devonian period (about 416 to 359 million years ago). The group contains two families: Holoptychiidae and Porolepididae. Porolepiformes was established by the ...

. * Together, these taxa form the Dipnomorpha, the sister group to the Tetrapodomorpha. * Together, these form the Rhipidistia, the sister group to the coelacanths. Recent molecular genetic analyses strongly support a sister relationship of lungfishes and tetrapods ( Rhipidistia), with coelacanths branching slightly earlier. 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 already identified may not be

monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...

. Current phylogenetic studies support the following relationships of major lungfish taxa: Class Osteichthyes, subclass Sarcopterygii, order Dipnoi. In 2017, Anne Kemp, Lionel Calvin, and Guillaume Guinot found a different classification for more derived lungfish. In this new analysis, all extant lungfish belong in the suborder Ceratodontoidei, and despite their similar appearance there is no distinct clade containing only ''

Ceratodus ''Ceratodus'' (from el, κέρας , 'horn' and el, ὀδούς 'tooth') was a wide-ranging genus of extinct lungfish. Fossil evidence dates back to the Early Triassic. A wide range of fossil species from different time periods have been found ...

'', ''

Neoceratodus ''Neoceratodus'' is a genus of lungfish in the family Neoceratodontidae. The extant Australian lungfish (''Neoceratodus forsteri'') is the only surviving member of this genus, but it was formerly much more widespread, being distributed throughout ...

'', and their closest allies. New analyses of the relationships of the recently discovered ''Persephonichthys chthonica'' and modern lungfishes robustly places both taxa within dipterid-grade dipnoans rather than within a clade containing Late Devonian ‘phaneropleurids’ and common Late Paleozoic lungfishes such as ''

Sagenodus ''Sagenodus'' is an extinct genus of prehistoric lungfish. It is a lungfish from the Permo-Carboniferous period found in Europe and North America. See also * Sarcopterygii * List of sarcopterygians * List of prehistoric bony fish A ''list'' ...

''. Monophyly of post-Devonian lungfishes is not supported and the Carboniferous-Permian taxon ''Sagenodus'' is found to be incidental to the origins of modern lungfishes, suggesting widespread convergence in Late Paleozoic lungfishes.Pardo, Jason D et al. “An exceptionally preserved transitional lungfish from the lower permian of Nebraska, USA, and the origin of modern lungfishes.” PLOS ONE vol. 9,9 e108542. 29 Sep. 2014, doi:10.1371/journal.pone.0108542

Timeline of lungfish genera of the Devonian

ImageSize = width:1000px height:auto barincrement:15px PlotArea = left:10px bottom:50px top:10px right:10px Period = from:-416 till:-349.2 TimeAxis = orientation:horizontal ScaleMajor = unit:year increment:5 start:-416 ScaleMinor = unit:year increment:1 start:-416 TimeAxis = orientation:hor AlignBars = justify Colors = #legends id:CAR value:claret id:ANK value:rgb(0.4,0.3,0.196) id:HER value:teal id:HAD value:green id:OMN value:blue id:black value:black id:white value:white id:paleozoic value:rgb(0.6,0.75,0.55) id:cambrian value:rgb(0.49,0.63,0.33) id:ordovician value:rgb(0,0.57,0.44) id:silurian value:rgb(0.70,0.88,0.71) id:devonian value:rgb(0.8,0.55,0.22) id:earlydevonian value:rgb(0.90,0.71,0.43) id:middledevonian value:rgb(0.96,0.81,0.51) id:latedevonian value:rgb(0.96,0.89,0.71) id:carboniferous value:rgb(0.4,0.65,0.6) id:mississippian value:rgb(0.4,0.56,0.4) id:pennsylvanian value:rgb(0.8,0.77,0.53) id:permian value:rgb(0.94,0.25,0.24) id:mesozoic value:rgb(0.38,0.77,0.79) id:triassic value:rgb(0.51,0.17,0.57) id:jurassic value:rgb(0.2,0.7,0.79) id:cretaceous value:rgb(0.5,0.78,0.31) id:cenozoic value:rgb(0.95,0.98,0.11) id:paleogene value:rgb(0.99,0.6,0.32) id:neogene value:rgb(0.999999,0.9,0.1) id:quaternary value:rgb(0.98,0.98,0.50) BarData= bar:eratop bar:space bar:periodtop bar:space bar:NAM1 bar:NAM2 bar:NAM3 bar:NAM4 bar:NAM5 bar:NAM6 bar:NAM7 bar:NAM8 bar:NAM9 bar:NAM10 bar:NAM11 bar:NAM12 bar:NAM13 bar:NAM14 bar:NAM15 bar:NAM16 bar:NAM17 bar:space bar:period bar:space bar:era PlotData= align:center textcolor:black fontsize:M mark:(line,black) width:25 shift:(7,-4) bar:periodtop from: -416 till: -411.2 color:earlydevonian text: Lochkovian from: -411.2 till: -407 color:earlydevonian text:

Pragian The Pragian is one of three faunal stages in the Early Devonian Epoch. It lasted from 410.8 ± 2.8 million years ago to 407.6 ± 2.8 million years ago. It was preceded by the Lochkovian Stage and followed by the Emsian Stage. The most important La ...

from: -407 till: -397.5 color:earlydevonian text: Emsian from: -397.5 till: -391.8 color:middledevonian text: Eifelian from: -391.8 till: -385.3 color:middledevonian text: Givetian from: -385.3 till: -374.5 color:latedevonian text: Frasnian from: -374.5 till: -359.2 color:latedevonian text: Famennian bar:eratop from: -416 till: -359.2 color:devonian text:

PlotData= align:left fontsize:M mark:(line,white) width:5 anchor:till align:left color:earlydevonian bar:NAM1 from:-416 till:-411.2 text: Diabolepis color:earlydevonian bar:NAM2 from:-416 till:-359.2 text: Devonesteus color:earlydevonian bar:NAM3 from:-416 till:-359.2 text: Grossipterus color:earlydevonian bar:NAM4 from:-416 till:-359.2 text:

Palaedaphus ''Palaedaphus'' is an extinct genus of prehistoric bony fish. See also * Prehistoric fish * List of prehistoric bony fish A ''list'' is any set of items in a row. List or lists may also refer to: People * List (surname) Organizations ...

color:earlydevonian bar:NAM5 from:-411.2 till:-407 text:

Uranolophus ''Uranolophus'' is a genus of prehistoric lungfish which lived during the Late Devonian period. It is the type genus of the family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or ...

color:earlydevonian bar:NAM6 from:-411.2 till:-359.2 text:

Dipterus ''Dipterus'' (from el, δίς , 'two' and el, πτερόν 'wing') is an extinct genus of lungfish from the middle Devonian period of Europe and North America. In most respects, ''Dipterus'', which was about long, closely resembled modern ...

color:earlydevonian bar:NAM7 from:-407 till:-403.83 text:

Sponysedrion ''Sponysedrion'' is an extinct genus of prehistoric bony fish. See also * Prehistoric fish * List of prehistoric bony fish A ''list'' is any set of items in a row. List or lists may also refer to: People * List (surname) Organizations ...

color:earlydevonian bar:NAM8 from:-407 till:-391.8 text:

Dipnorhynchus ''Dipnorhynchus'' is an extinct genus of lungfish from the middle Devonian period of Australia and Europe. ''Dipnorhynchus'' was a primitive lungfish, but still it had features that set it apart from other sarcopterygians. Its skull lacked the j ...

color:middledevonian bar:NAM9 from:-397.5 till:-385.3 text:

Melanognathus ''Melanognathus'' (Greek for "black jaw") is a genus of prehistoric lungfish which lived during the Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silur ...

color:middledevonian bar:NAM10 from:-397.5 till:-385.3 text:

Stomiahykus ''Stomiahykus'' is an extinct genus of prehistoric lobe-finned fish. See also * Prehistoric fish * List of prehistoric bony fish A ''list'' is any set of items in a row. List or lists may also refer to: People * List (surname) Organiz ...

color:middledevonian bar:NAM11 from:-397.5 till:-359.2 text:

Rhinodipterus ''Rhinodipterus'' is an extinct genus of prehistoric dipnoan sarcopterygians or lobe-finned fish, that lived in the Devonian Period, between 416 and 359 million years ago. It is believed to have inhabited shallow, salt-water reefs, and is one of ...

color:middledevonian bar:NAM12 from:-391.8 till:-359.2 text:

Ganorhynchus ''Ganorhynchus'' is an extinct genus of prehistoric lungfish from the Devonian period. It is only found in Mansfield, Pennsylvania Mansfield is a Borough (Pennsylvania), borough located in east-central Tioga County, Pennsylvania, United States ...

color:middledevonian bar:NAM13 from:-387.5 till:-359.2 text:

Conchodus ''Conchodus'' is an extinct genus of lungfish which lived during the Devonian period. References Prehistoric lungfish genera Devonian fish of Europe {{paleo-lobefinned-fish-stub ...

color:latedevonian bar:NAM14 from:-385.3 till:-381.7 text: Chirodipterus color:latedevonian bar:NAM15 from:-385.3 till:-381.7 text:

Pillararhynchus ''Pillararhynchus'' is an extinct genus of prehistoric sarcopterygians or lobe-finned fish. See also * Sarcopterygii * List of sarcopterygians * List of prehistoric bony fish A ''list'' is any set of items in a row. List or lists may also re ...

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Griphognathus ''Griphognathus'' (from el, γρίφος 'riddle') and el, γνάθος 'jaw') is an extinction, extinct genus of lungfish from the late Devonian period of Europe and Australia. ''Griphognathus'' was a specialized lungfish, about long, with ...

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Holodipterus ''Holodipterus'' is an extinct genus of prehistoric sarcopterygian or lobe-finned fish.Campbell, KSW T., and R. E. Barwick. "Paleozoic lungfishes—a review."Journal of Morphology 190.S1 (1986): 93-131. Species * Holodipterus elderae * Holod ...

PlotData= align:center textcolor:black fontsize:M mark:(line,black) width:25 bar:period from: -416 till: -411.2 color:earlydevonian text: Lochkovian from: -411.2 till: -407 color:earlydevonian text:

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* * * * * {{Authority control Articles which contain graphical timelines Extant Early Devonian first appearances