Amia (fish)
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''Amia'', commonly called bowfin, is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
ray-finned fish Actinopterygii (; ), members of which are known as ray-finned fish or actinopterygians, is a class of bony fish that comprise over 50% of living vertebrate species. They are so called because of their lightly built fins made of webbings of sk ...
related to
gar Gars are an ancient group of ray-finned fish in the family Lepisosteidae. They comprise seven living species of fish in two genera that inhabit fresh, brackish, and occasionally marine waters of eastern North America, Central America and Cuba ...
s in the infraclass
Holostei Holostei is a group of ray-finned bony fish. It is divided into two major clades, the Halecomorphi, represented by the single living genus, '' Amia'' with two species, the bowfins (''Amia calva'' and '' Amia ocellicauda''), as well as the Gin ...
. They are regarded as
taxonomic 280px, Generalized scheme of taxonomy Taxonomy is a practice and science concerned with classification or categorization. Typically, there are two parts to it: the development of an underlying scheme of classes (a taxonomy) and the allocation ...
relicts A relict is a surviving remnant of a natural phenomenon. Biology A relict (or relic) is an organism that at an earlier time was abundant in a large area but now occurs at only one or a few small areas. Geology and geomorphology In geology, a r ...
, being the sole surviving species of the
order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...
Amiiformes The Amiiformes order (biology), order of fish has only two extant taxa, extant species, the bowfins: ''Amia calva'' and ''Amia ocellicauda'', the latter recognized as a separate species in 2022. These Amiiformes are found in the freshwater syste ...
and clade
Halecomorphi Halecomorphi is a taxon of Actinopterygii, ray-finned bony fish in the clade Neopterygii. The only extant Halecomorph species are the bowfin (''Amia calva'') and eyespot bowfin (''Amia ocellicauda''), but the group contains many extinct species i ...
, which dates from the
Triassic The Triassic ( ; sometimes symbolized 🝈) is a geologic period and system which spans 50.5 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.4 Mya. The Triassic is t ...
to the
Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
, persisting to the present. There are two living species in ''Amia'', ''
Amia calva The ruddy bowfin (''Amia calva'') is a ray-finned fish native to North America. Common names include mudfish, mud pike, dogfish, grindle, grinnel, swamp trout, and choupique. It is regarded as a relict species, relict, being one of only two sur ...
'' and ''
Amia ocellicauda ''Amia ocellicauda'', the eyespot bowfin or emerald bowfin, is a species of Amia (fish), bowfin native to North America. Originally described by John Richardson (naturalist), John Richardson from Lake Huron in 1836, it was synonymized with ''Amia ...
'', and a number of
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
species which have been described from the fossil record.


Etymology

The genus name ''Amia'' derives from the Ancient Greek and Latin name for an unknown fish (most likely the
Atlantic bonito The Atlantic bonito (''Sarda sarda'') is a large mackerel-like fish of the family Scombridae. It is common in shallow waters of the Atlantic Ocean, the Mediterranean Sea, and the Black Sea, where it is an important commercial and game fish. Desc ...
) referred to by ancient and medieval authors including
Pliny the Elder Gaius Plinius Secundus (AD 23/24 79), known in English as Pliny the Elder ( ), was a Roman Empire, Roman author, Natural history, naturalist, and naval and army commander of the early Roman Empire, and a friend of the Roman emperor, emperor Vesp ...
,
Isidore of Seville Isidore of Seville (; 4 April 636) was a Spania, Hispano-Roman scholar, theologian and Roman Catholic Archdiocese of Seville, archbishop of Seville. He is widely regarded, in the words of the 19th-century historian Charles Forbes René de Montal ...
, and
Thomas of CantimprĂ© Thomas of CantimprĂ© (Latin: Thomas Cantimpratensis or Thomas Cantipratensis) (Sint-Pieters-Leeuw, 1201 – Louvain, 15 May 1272) was a Flemish Region, Flemish Catholic medieval writer, preacher, theologian and a friar belonging to the Dominican ...
.


Evolution and phylogeny

''Amia'' is thought to have diverged from its
sister genus 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 ...
, ''
Cyclurus ''Cyclurus'' (Ancient Greek for "rounded tail") is an extinct genus of freshwater Amiidae, amiid Actinopterygii, ray-finned fish known from the Late Cretaceous to the Rupelian, Early Oligocene across much of the Northern Hemisphere. It is though ...
'', during the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''cre ...
. These two genera, their sister amiine '' Pseudoamiatus'', and the marine vidalamiine ''
Maliamia ''Maliamia'' ("Malian bowfin") is an extinct genus of Amiidae, amiid ray-finned fish from the Early Eocene, known from fragmentary remains found in the Tamaguélelt Formation of Mali. It was described in 1989, based on fossils recovered by three ...
'' were the only amiids, and halecomorphs as a whole, to survive the Cretaceous-Paleogene extinction event. Following the extinction event, ''Amia'' and ''Cyclurus'' survived in a refugium in western North America and experienced a rapid evolutionary radiation. The earliest fossil remains of ''Amia'' date to the
Middle Paleocene The Selandian is a stage in the Paleocene. It spans the time between . It is preceded by the Danian and followed by the Thanetian. Sometimes the Paleocene is subdivided in subepochs, in which the Selandian forms the "middle Paleocene". Stratigr ...
, several million years after the K-Pg extinction. During this time period, some species of ''Amia'' evolved very large body sizes, most notably '' A. basiloides'', which is one of the largest
holosteans Holostei is a group of ray-finned bony fish. It is divided into two major clades, the Halecomorphi, represented by the single living genus, '' Amia'' with two species, the bowfins (''Amia calva'' and ''Amia ocellicauda''), as well as the Ginglymo ...
known to have existed. In contrast to the modern distribution of ''Amia'', most of these fossils are from western North America. Although ''Cyclurus'' dispersed to Europe and Asia shortly after the K-Pg extinction event, ''Amia'' largely remained in western North America until the very end of the
Paleogene The Paleogene Period ( ; also spelled Palaeogene or PalĂŠogene) is a geologic period and system that spans 43 million years from the end of the Cretaceous Period Ma (million years ago) to the beginning of the Neogene Period Ma. It is the fir ...
, when it dispersed west to Asia and east to eastern North America. ''Cyclurus'' went extinct during the
Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...
, leaving ''Amia'' as the only surviving amiid. At some point afterwards, ''Amia'' went extinct in western North America, and also went extinct in Asia during the
Neogene The Neogene ( ,) is a geologic period and system that spans 20.45 million years from the end of the Paleogene Period million years ago ( Mya) to the beginning of the present Quaternary Period million years ago. It is the second period of th ...
, leaving only the eastern American populations, which have since undergone a small level of diversification.


Species

List of species. * ''
Amia calva The ruddy bowfin (''Amia calva'') is a ray-finned fish native to North America. Common names include mudfish, mud pike, dogfish, grindle, grinnel, swamp trout, and choupique. It is regarded as a relict species, relict, being one of only two sur ...
'' Linnaeus 1766 (bowfin) * ''
Amia ocellicauda ''Amia ocellicauda'', the eyespot bowfin or emerald bowfin, is a species of Amia (fish), bowfin native to North America. Originally described by John Richardson (naturalist), John Richardson from Lake Huron in 1836, it was synonymized with ''Amia ...
'' (eyespot bowfin) * ''
Amia basiloides ''Amia basiloides'' is an extinct species of giant bowfin that inhabited western North America during the Middle or Late Paleocene, about 5-10 million years after the Cretaceous-Paleogene extinction event. The species name originates from the A ...
'' Brownstein & Near, 2024 (middle
Paleocene The Paleocene ( ), or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), ...
of Montana, USA) * '' Amia godai'' Yabumoto & Grande, 2013 (
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
of Japan) * '' Amia hesperia'' Wilson, 1982 (
Eocene Okanagan Highlands The Eocene Okanagan Highlands or Eocene Okanogan Highlands are a series of Early Eocene geological formations which span a transect of British Columbia, Canada, and Washington state, United States. Known for a highly diverse and detailed plant ...
) * ''
Amia pattersoni Amia, AMIA, or AMiA may refer to: * ''Amia'' (fish), a genus of fish *American Medical Informatics Association *Anglican Mission in the Americas *AsociaciĂłn Mutual Israelita Argentina, a Jewish community center in Buenos Aires, Argentina **AMIA ...
'' Grande & Bemis, 1998 (late Paleocene of Alberta, Canada) * '' Amia scutata'' Cope 1875 (late Eocene of Colorado, USA) The species '''Amia uintaensis''' Leidy, 1873, which formerly contained several ''Amia'' specimens from western North America dating from the Late Cretaceous to the Eocene (including the holotype of ''A. basiloides''), is considered a ''
nomen vanum This is a list of terms and symbols used in scientific names for organisms, and in describing the names. For proper parts of the names themselves, see List of Latin and Greek words commonly used in systematic names. Many of the abbreviations are ...
''. Another large Paleocene amiid, ''Amia'' '''robusta''' Smith & Radcliffe, 1911 from France, is known from non-diagnostic remains, and is thus considered a ''
nomen dubium In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application. Zoology In case of a ''nomen dubium,'' it may be impossible to determine whether a ...
''.


Phylogeny

Based on Brownstein & Near (2024):


Genome evolution

The bowfin
genome A genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding genes, other functional regions of the genome such as ...
contains an intact
ParaHox The ParaHox gene cluster is an array of homeobox genes (involved in morphogenesis, the regulation of patterns of anatomical development) from the Gsx, Xlox ( Pdx) and Cdx gene families. Regulatory gene cluster These genes were first shown to ...
gene cluster A gene cluster is a group of two or more genes found within an organism's DNA that encode similar peptide, polypeptides or proteins which collectively share a generalized function and are often located within a few thousand base pairs of each othe ...
, similar to the
bichir Bichirs and the reedfish comprise Polypteridae , a family (biology), family of archaic Actinopterygii, ray-finned fishes and the only family in the order (biology), order Polypteriformes .Helfman GS, Collette BB, Facey DE, Bowen BW. 2009. The D ...
and to most other
vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
s. This is in contrast, however, with
teleost Teleostei (; Ancient Greek, Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts (), is, by far, the largest group of ray-finned fishes (class Actinopterygii), with 96% of all neontology, extant species of f ...
fish, which have a fragmented ParaHox cluster, probably because of a
whole genome duplication Paleopolyploidy is the result of genome duplications which occurred at least several million years ago (MYA). Such an event could either double the genome of a single species ( autopolyploidy) or combine those of two species ( allopolyploidy). ...
event in their lineage. The presence of an intact ParaHox gene cluster suggests that bowfin ancestors separated from other fish before the
last common ancestor A most recent common ancestor (MRCA), also known as a last common ancestor (LCA), is the most recent individual from which all organisms of a set are inferred to have descended. The most recent common ancestor of a higher taxon is generally assu ...
of all
teleost Teleostei (; Ancient Greek, Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts (), is, by far, the largest group of ray-finned fishes (class Actinopterygii), with 96% of all neontology, extant species of f ...
s appeared. Bowfin are thus possibly a better model to study vertebrate genome organization than common teleost
model organism A model organism is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the model organism will provide insight into the workings of other organisms. Mo ...
s such as
zebrafish The zebrafish (''Danio rerio'') is a species of freshwater ray-finned fish belonging to the family Danionidae of the order Cypriniformes. Native to South Asia, it is a popular aquarium fish, frequently sold under the trade name zebra danio (an ...
.


References

{{Authority control Ray-finned fish genera Extant Selandian first appearances Taxa named by Carl Linnaeus