The Gnathorhizidae are an extinct family of

lungfish Lungfish are freshwater vertebrates belonging to the order Dipnoi. Lungfish are best known for retaining ancestral characteristics within the Osteichthyes, including the ability to breathe air, and ancestral structures within Sarcopterygii, i ...

that lived from the late

Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonifero ...

until the middle

Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...

. Gnathorhizid

fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...

s have been found in

North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...

Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Africa ...

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 ...

, and possibly

Eastern Europe Eastern Europe is a subregion of the Europe, European continent. As a largely ambiguous term, it has a wide range of geopolitical, geographical, ethnic, cultural, and socio-economic connotations. The vast majority of the region is covered by Russ ...

and

South Africa South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring countri ...

. They are characterized by high-ridged toothplates that form cutting blades and a reduction in cranial bones.

Taxonomy

Phylogenetic evidence supports Gnathorhizidae being the

sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...

to the

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, ...

family

Lepidosirenidae Lepidosirenidae is a family of lungfish containing the genera ''Lepidosiren'' (the South American lungfish) and ''Protopterus'' (the African lungfish). Both genera were formerly thought to represent the distinct families Lepidosirenidae and Proto ...

, which still lives in

South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...

and

Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...

. The clade containing Gnathorhizidae and Lepidosirenidae is the

crown group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...

of the suborder

Ceratodontoidei Ceratodontoidei is a suborder of lungfish that is defined as "the clade including all taxa more closely related to ''Lepidosiren'', '' Neoceratodus'' and '' Gnathorhiza'' than to '' Uronemus'', '' Conchopoma'' and ''Sagenodus''". Members of this ...

, which also contains the extant family

Neoceratodontidae Neoceratodontidae is a family of lungfish containing the extant Australian lungfish and several extinct genera. It and Lepidosirenidae represent the only lungfish families still extant. Fossils from this family are first known from Triassic-aged ...

and many other extinct families.

Distribution

Gnathorhizids are found in North America, Eastern Europe, Australia, and Africa. Gnathorhizids from North America range from the

Gzhelian The Gzhelian ( ) is an age in the ICS geologic timescale or a stage in the stratigraphic column. It is the youngest stage of the Pennsylvanian, the youngest subsystem of the Carboniferous. The Gzhelian lasted from to Ma. It follows the Kasim ...

through the

Roadian In the geologic timescale, the Roadian is an age or stage of the Permian. It is the earliest or lower of three subdivisions of the Guadalupian Epoch or Series. The Roadian lasted between and million years ago (Ma). It was preceded by the Kung ...

. In Africa, gnathorhizids are found in

Olenekian In the geologic timescale, the Olenekian is an age in the Early Triassic epoch; in chronostratigraphy, it is a stage in the Lower Triassic series. It spans the time between Ma and Ma (million years ago). The Olenekian is sometimes divided into ...

of Madagascar and possibly South Africa. Lungfish

teeth A tooth ( : teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, tear ...

attributed to gnathorhizids have been reported from the

Lopingian The Lopingian is the uppermost series/last epoch of the Permian. It is the last epoch of the Paleozoic. The Lopingian was preceded by the Guadalupian and followed by the Early Triassic. The Lopingian is often synonymous with the informal terms l ...

to the Olenekian in Poland and Western Russia. It is likely, then, that gnathorhizids had a

Pangea Pangaea or Pangea () was a supercontinent that existed during the late Paleozoic and early Mesozoic eras. It assembled from the earlier continental units of Gondwana, Euramerica and Siberia during the Carboniferous approximately 335 million y ...

n distribution throughout the late

Paleozoic The Paleozoic (or Palaeozoic) Era is the earliest of three geologic eras of the Phanerozoic Eon. The name ''Paleozoic'' ( ;) was coined by the British geologist Adam Sedgwick in 1838 by combining the Greek words ''palaiós'' (, "old") and ' ...

and early

Mesozoic The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceo ...

Paleoecology and behavior

Gnathorhizids are found primarily in

paleosol In the geosciences, paleosol (''palaeosol'' in Great Britain and Australia) is an ancient soil that formed in the past. The precise definition of the term in geology and paleontology is slightly different from its use in soil science. In geolo ...

s representing

ephemeral Ephemerality (from the Greek word , meaning 'lasting only one day') is the concept of things being transitory, existing only briefly. Academically, the term ephemeral constitutionally describes a diverse assortment of things and experiences, fr ...

wetlands A wetland is a distinct ecosystem that is flooded or saturated by water, either permanently (for years or decades) or seasonally (for weeks or months). Flooding results in oxygen-free (anoxic) processes prevailing, especially in the soils. The ...

. Additionally, gnathorhizids, unlike most groups of fossil lungfish, are often found in association with regular

burrow An Eastern chipmunk at the entrance of its burrow A burrow is a hole or tunnel excavated into the ground by an animal to construct a space suitable for habitation or temporary refuge, or as a byproduct of locomotion. Burrows provide a form of sh ...

structures, suggesting this group of lungfish may have estivated during the dry season, much like modern

African African or Africans may refer to: * Anything from or pertaining to the continent of Africa: ** People who are native to Africa, descendants of natives of Africa, or individuals who trace their ancestry to indigenous inhabitants of Africa *** Ethn ...

and

South American South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...

lungfish. Unlike most fossil lungfish, but again, like modern South American and African lungfish, gnathorhizids have bladelike toothplates. This suggests gnathorhizids were active predators unlike most lungfish, which feed primarily on

benthic The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from ancient Greek, βένθος (bénthos), meaning "t ...

invertebrates Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...

References

Sources

* Berman, D. S., 1976, ''Cranial morphology of the Lower Permian lungfish Gnathorhiza (Osteichthyes: Dipnoi): Journal of Paleontology'', v. 50(6), p. 1020-1033. * Cunningham, C. R. and Dickson, E. D III, 1996, ''Distributions of Kansas Permo-Carboniferous vertebrate assemblages as a function of wet and dry seasons: Transactions of the Kansas Academy of Science'', v. 99(1-2), p. 16-28. * Huttenlocker, A.K. ''et al.'', 2005, ''An earliest Permian nonmarine vertebrate assemblage from the Eskridge Formation, Nebraska: * Lucas, S.G. and Zeigler, K.E., eds., 2005, ''The Nonmarine Permian, New Mexico Museum of Natural History and Science Bulletin No. 30''., pp. 133–143. {{Taxonbar, from=Q16982395 Prehistoric lungfish Prehistoric lobe-finned fish families Middle Triassic extinctions Pennsylvanian first appearances