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Ceratodontiformes is the only extant order of
lungfish Lungfish are freshwater vertebrates belonging to the class Dipnoi. Lungfish are best known for retaining ancestral characteristics within the Osteichthyes, including the ability to breathe air, and ancestral structures within Sarcopterygii, inc ...
, containing the families
Neoceratodontidae Neoceratodontidae is a Family (biology), family of lungfish containing ''Neoceratodus'' (represented by the extant Australian lungfish) and the extinct ''Mioceratodus''. It, South American lungfish, Lepidosirenidae, and Protopterus, Protopteridae ...
, Lepidosirenidae, and
Protopteridae ''Protopterus'' is the genus of four species of lungfish found in Africa. ''Protopterus'' is considered the monotypic, sole genus in the family Protopteridae, which is grouped with ''South American lungfish, Lepidosiren'' in the order Lepidosir ...
as well as many other extinct groups. Members of this group are the only lungfish known to have survived the Permian-Triassic extinction event. Although lungfish originated in marine environments, the Ceratodontiformes have been an exclusively freshwater group since the
Carboniferous The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic era (geology), era that spans 60 million years, from the end of the Devonian Period Ma (million years ago) to the beginning of the ...
. This order was formerly considered the suborder Ceratodontoidei. All lungfish of the order can and often do estivate (except the spotted African lungfish, which can but rarely does so). All members of the order are obligatory air-breathers; only the
Australian lungfish The Australian lungfish (''Neoceratodus forsteri''), also known as the Queensland lungfish, Burnett salmon and barramunda, is the only surviving member of the family Neoceratodontidae. It is one of only six extant lungfish species in the world. ...
has functioning
gills A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they are ...
when adult; members of the Lepidosirenidae have gills only when they are larvae. The South American and African lungfish also all have generally small scales and two lungs as opposed to the Australian lungfish's single lung.


Taxonomy

The suborder was formerly defined as being within the order Ceratodontiformes and including the families Neoceratodontidae and Ceratodontidae, as they were formerly thought to be closely related to one another. However, phylogenetic analyses indicate that this classification is
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
, as Ceratodontidae was found to be a
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 ...
to a clade containing Lepidosirenidae, which was formerly classified as Lepidosireniformes, a distinct order from Ceratodontiformes. Due to this, Lepidosireniformes and Ceratodontiformes were redefined as families within the order Ceratodontiformes, redefined as including all lungfish more closely allied with Neoceratodontidae and Lepidosirenidae. This order contains the following subtaxa: * Order Ceratodontiformes ** Genus †'' Ameghinoceratodus'' ** Genus †'' Chaoceratodus'' ** Genus †''
Ferganoceratodus ''Ferganoceratodus'' (from Fergana + '' Ceratodus'') is a genus of prehistoric freshwater lungfish known from worldwide during the Mesozoic. Based on morphological evidence, it has either been recovered as a basal member of the Ceratodontiformes ...
'' ** Genus †''
Gosfordia ''Gosfordia'' is an extinct genus of prehistoric lungfish known from the Triassic of Australia. See also * Sarcopterygii * List of sarcopterygians * List of prehistoric bony fish This list of prehistoric bony fish is an attempt to create a c ...
'' ** Genus †''
Paraceratodus ''Paraceratodus'' is an extinct genus of prehistoric lungfish. Only one species, ''P. germaini'', is known from the latest Permian or earliest Triassic period of Madagascar. Phylogenetic evidence supports it being the most basal member of the su ...
'' ** Genus †'' Permoceratodus'' ** Genus †''
Potamoceratodus ''Potamoceratodus'' is an extinct genus of lungfish belonging to the family Ceratodontidae known from the Late Jurassic Morrison Formation of Colorado, USA. It was first named by Jason D. Pardo, Adam K. Huttenlocker, Bryan J. Small and Mark A. ...
'' ** Family † Arganodontidae (sometimes synonymized with Asiatoceratodontidae or placed within Ceratodontidae) ** Family † Asiatoceratodontidae ** Family † Ceratodontidae ** Family †
Gnathorhizidae The Gnathorhizidae are an extinct family of lungfish that lived from the late Carboniferous until the middle Triassic. Gnathorhizid fossils have been found in North America, Madagascar, Australia, and possibly Eastern Europe and South Africa. ...
** Suborder Lepidosirenoidei *** Family
Protopteridae ''Protopterus'' is the genus of four species of lungfish found in Africa. ''Protopterus'' is considered the monotypic, sole genus in the family Protopteridae, which is grouped with ''South American lungfish, Lepidosiren'' in the order Lepidosir ...
*** Family ?† Lavocatodidae *** Family Lepidosirenidae ** Family
Neoceratodontidae Neoceratodontidae is a Family (biology), family of lungfish containing ''Neoceratodus'' (represented by the extant Australian lungfish) and the extinct ''Mioceratodus''. It, South American lungfish, Lepidosirenidae, and Protopterus, Protopteridae ...
** Family † Ptychoceratodontidae Some of these groups, such as Ceratodontidae, are thought to be paraphyletic due to being based entirely on morphology. Other groups are often synonymized with one another or subsumed within others, due to the taxonomic confusion surrounding this group.


Phylogenies

Kemp ''et al'' (2017) found the following taxonomy based on morphological evidence: Based on this treatment, the oldest fossils of the Ceratodontiformes are of
Gnathorhizidae The Gnathorhizidae are an extinct family of lungfish that lived from the late Carboniferous until the middle Triassic. Gnathorhizid fossils have been found in North America, Madagascar, Australia, and possibly Eastern Europe and South Africa. ...
from the
Late Carboniferous Late or LATE may refer to: Everyday usage * Tardy, or late, not being on time * Late (or the late) may refer to a person who is dead Music * Late (The 77s album), ''Late'' (The 77s album), 2000 * Late (Alvin Batiste album), 1993 * Late!, a pseudo ...
, which are thought to be closely related to modern Lepidosirenidae and
Protopteridae ''Protopterus'' is the genus of four species of lungfish found in Africa. ''Protopterus'' is considered the monotypic, sole genus in the family Protopteridae, which is grouped with ''South American lungfish, Lepidosiren'' in the order Lepidosir ...
. This would indicate that the order itself originated slightly earlier in the late Carboniferous and rapidly diversified into the multiple families between then and the start of the
Permian The Permian ( ) is a geologic period and System (stratigraphy), stratigraphic system which spans 47 million years, from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.902 Mya. It is the s ...
, leading to a very deep split between the
Neoceratodontidae Neoceratodontidae is a Family (biology), family of lungfish containing ''Neoceratodus'' (represented by the extant Australian lungfish) and the extinct ''Mioceratodus''. It, South American lungfish, Lepidosirenidae, and Protopterus, Protopteridae ...
and the Lepidosirenidae + Protopteridae. Brownstein, Harrington & Near (2023) found a different taxonomy, based on both phylogenetic and morphological evidence. In this treatment, the former Lepidosirenformes is kept. This analysis found a more recent divergence, with the Neoceratodontidae and the "Lepidosireniformes" diverging during the
Late Jurassic The Late Jurassic is the third Epoch (geology), epoch of the Jurassic Period, and it spans the geologic time scale, geologic time from 161.5 ± 1.0 to 143.1 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic stratum, strata.Owen ...
, in contrast to the
Paleozoic The Paleozoic ( , , ; or Palaeozoic) Era is the first of three Era (geology), geological eras of the Phanerozoic Eon. Beginning 538.8 million years ago (Ma), it succeeds the Neoproterozoic (the last era of the Proterozoic Eon) and ends 251.9 Ma a ...
divergences estimated by Kemp ''et al'''s morphological study. This divergence was found to be tied to the breakup of
Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...
during the same time.


References

{{Taxonbar, from1=Q1328838, from2=Q21230686 Lungfish Extant Pennsylvanian first appearances Fish orders Taxa named by Lev Berg