Arapaiminae
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Arapaimidae is a
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
of freshwater
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 ...
belonging to the order
Osteoglossiformes Osteoglossiformes , meaning "bony tongues" in Ancient Greek, is a relatively primitive order of ray-finned fish that contains two sub-orders, the Osteoglossoidei and the Notopteroidei. All of at least 245 living species inhabit freshwater. They ...
. It includes the South American
arapaima The arapaima, pirarucu, or paiche is any large species of bonytongue in the genus ''Arapaima'' native to the Amazon Basin, Amazon and Essequibo River, Essequibo basins of South America. ''Arapaima'' is the type genus of the subfamily Arapaiminae ...
s of the
Amazon Amazon most often refers to: * Amazon River, in South America * Amazon rainforest, a rainforest covering most of the Amazon basin * Amazon (company), an American multinational technology company * Amazons, a tribe of female warriors in Greek myth ...
and Essequibo basins and the African arowana (''Heterotis niloticus'') from the watersheds of the Sahelo-Sudanese region,
Senegal Senegal, officially the Republic of Senegal, is the westernmost country in West Africa, situated on the Atlantic Ocean coastline. It borders Mauritania to Mauritania–Senegal border, the north, Mali to Mali–Senegal border, the east, Guinea t ...
,
Gambia The Gambia, officially the Republic of The Gambia, is a country in West Africa. Geographically, The Gambia is the List of African countries by area, smallest country in continental Africa; it is surrounded by Senegal on all sides except for ...
, and parts of Eastern Africa. This family is sometimes treated as the subfamily Arapaiminae. A commonly used synonym is Heterotidinae, but according to the
ICZN The International Code of Zoological Nomenclature (ICZN) is a widely accepted convention in zoology that rules the formal scientific naming of organisms treated as animals. It is also informally known as the ICZN Code, for its formal author, t ...
, Arapaimidae has priority. Arapaimides, along with other osteoglossomorphs, are of
phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
and evolutionary interest due to their trans-oceanic distribution, excellent fossil record, and position as one of the oldest living
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 ...
lineages. The type-species of the group, ''
Arapaima gigas ''Arapaima gigas'', also known simply as Arapaima, pirarucu, or paiche,Miranda-Chumacero, G., Wallace, R., Calderón, H., Calderón, G., Willink, P., Guerrero, M., ... & Chuqui, D. (2012). Distribution of arapaima (Arapaima gigas)(Pisces: Arapaim ...
'', is an important South American food source and charismatic representative of the region. Both ''Arapaima'' and ''Heterotis'' are cultured for food in their respective countries due to their heartiness and meat, and the arapaima is a prized sport-fish, being the largest truly freshwater fish.


Phylogeny and systematics

The internal placement of Osteoglossomorpha within
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. ...
teleosts is contested, with competing morphological and
molecular A molecule is a group of two or more atoms that are held together by attractive forces known as chemical bonds; depending on context, the term may or may not include ions that satisfy this criterion. In quantum physics, organic chemistry, ...
analyses placing them either as sister to all other
extant Extant or Least-concern species, least concern 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 * Exta ...
teleosts, or internal to Elopimorpha and sister to the clade consisting of
Otocephala Otocephala is a clade of ray-finned fishes within the infraclass Teleostei that evolved some 230 million years ago. It is named for the presence of a hearing (otophysic) link from the swimbladder to the inner ear. Other names proposed for the gro ...
and
Euteleostei Euteleostei, whose members are known as euteleosts, is a clade of bony fishes within Teleostei that evolved some 240 million years ago, although the oldest known fossil remains are only from the Early Cretaceous. It is divided into Protacanthopt ...
. The placement and name of the clade containing ''Arapaima'' and ''Heterotis'' is also uncertain. Some include this clade in the family Osteoglossidae with the South American and Asian arowana. Others place ''Arapaima'' and ''Heterotis'' together in their own family, Arapaimidae.


Taxonomy

''Arapaima'' taxonomy was recently revised to revalidate old names and describe a new species, proposing 6 existing species (see below) and invalidating current museum specimens. However, these four proposed or reestablished species are known only from singular
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
specimens, and only that of ''A. mapae'' and ''A. leptostoma'' still currently exist. Typically, all species of ''Arapaima'' described by
Valenciennes Valenciennes (, also , , ; ; or ; ) is a communes of France, commune in the Nord (French department), Nord Departments of France, department, Hauts-de-France, France. It lies on the Scheldt () river. Although the city and region experienced ...
,
Spix Johann Baptist Ritter von Spix (9 February 1781 – 13 March 1826) was a German natural history, biologist. From his expedition to Brazil, he brought to Germany a large variety of specimens of plants, insects, mammals, birds, amphibians and fish. ...
, and
Agassiz Jean Louis Rodolphe Agassiz ( ; ) FRS (For) FRSE (May 28, 1807 – December 14, 1873) was a Swiss-born American biologist and geologist who is recognized as a scholar of Earth's natural history. Spending his early life in Switzerland, he recei ...
are referred to as ''A. gigas'', though current taxonomy could be revised with more thorough evidence. Currently, population genetic evidence supports a singular ''Arapaima'' species with two distinct genetic populations: an Amazonas population (exhibiting a pattern of isolation by distance), and an Araguaia-Tocantins basin population. There is little debate that ''Heterotis'' is a monotypic genus represented by only ''H. niloticus''. * Arapaimidae Bonaparte, 1846 ** '' Arapaimidarum'' tolith** '' Heterotidinarum'' Nolf, Rana & Prasad 2008 tolith** '' Thrissopterus'' Heckel 1856 ** ''
Joffrichthys ''Joffrichthys'' is a genus of prehistoric bony fish. This North American genus includes three species, ''J. symmetropterus'', ''J. tanyourus'' and ''J. triangulpterus''. The last species is known from the Paleocene of the Sentinel Butte Format ...
''? Li & Wilson 1996 ** '' Sinoglossus''? Su 1986 ** '' Heterotis'' Rüppell 1829 ex Ehrenberg 1836 (African arowana) *** '' H. niloticus'' (Cuvier, 1829) ** ''
Arapaima The arapaima, pirarucu, or paiche is any large species of bonytongue in the genus ''Arapaima'' native to the Amazon Basin, Amazon and Essequibo River, Essequibo basins of South America. ''Arapaima'' is the type genus of the subfamily Arapaiminae ...
'' Müller 1843 (arapaima) *** '' A. agassizii'' (Valenciennes, 1847) *** '' A. gigas'' (Schinz, 1822) *** '' A. leptosoma'' Stewart, 2013 *** '' A. mapae'' (Valenciennes, 1847) *** ''A. sp.'' ''
incertae sedis or is a term used for a taxonomy (biology), taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...
''


Description and biology


Description

Arapaimides are characterized by elongate, slender bodies with large scales and long dorsal and anal fins positioned close to a short caudal peduncle. The pelvic fins are small and abdominal if present. They lack chin barbels, have a glossolaryngeal (tongue) bone with teeth present, and the
premaxilla 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 mammals h ...
e are fixed to the skull. Branched caudal fin rays are less than sixteen, branchiostegal rays between three and seven, and hypurals less than 6. ''Heterotis'' possesses a specialized suprabranchial organ for concentrating and filtering small food particles.


Biology

Both genera make use of similar freshwater habitats in the respective region, with ''Arapaima'' found in the floodplains of the
Amazon Amazon most often refers to: * Amazon River, in South America * Amazon rainforest, a rainforest covering most of the Amazon basin * Amazon (company), an American multinational technology company * Amazons, a tribe of female warriors in Greek myth ...
and Esequibo river basins of South America and ''Heterotis'' found in
littoral zone The littoral zone, also called litoral or nearshore, is the part of a sea, lake, or river that is close to the shore. In coastal ecology, the littoral zone includes the intertidal zone extending from the high water mark (which is rarely flood ...
s of large, open rivers in all Sahelo-Sudanese basins of Africa. ''Arapaima'' is typically a
top Top most commonly refers to: * Top, a basic term of orientation, distinguished from bottom, front, back, and sides * Spinning top, a ubiquitous traditional toy * Top (clothing), clothing designed to be worn over the torso * Mountain top, a moun ...
-water fish predator, while ''Heterotis'' is a
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 the Ancient Greek word (), meaning "the depths". ...
mud- filterer primarily feeding on
phytoplankton Phytoplankton () are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater Aquatic ecosystem, ecosystems. The name comes from the Greek language, Greek words (), meaning 'plant', and (), mea ...
and small crustaceans with their suprabranchial organ. Both groups are obligate air-breathers and nest-builders, with males guarding eggs and young.


Evolution

A genetic study shows that Arapaimidae diverged from
Osteoglossinae Arowanas are freshwater bony fish of the subfamily Osteoglossinae, also known as bony tongues (the latter name is now often reserved for Arapaiminae). In this family of fish, the head is bony and the elongated body is covered by large, heavy sca ...
about 220 million years ago, during the
Late Triassic The Late Triassic is the third and final epoch (geology), epoch of the Triassic geologic time scale, Period in the geologic time scale, spanning the time between annum, Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch a ...
. Within Osteoglossinae, the lineage leading to the South American ''
Osteoglossum ''Osteoglossum'' is a genus of fish in the family Osteoglossidae. They reach about in length and are restricted to freshwater habitats in tropical South America. These predators mostly feed on arthropods like insects and spiders, but may also t ...
'' arowanas diverged about 170 million years ago, during the
Middle Jurassic The Middle Jurassic is the second Epoch (geology), epoch of the Jurassic Period (geology), Period. It lasted from about 174.1 to 161.5 million years ago. Fossils of land-dwelling animals, such as dinosaurs, from the Middle Jurassic are relativel ...
. The Asian and Australian arowanas in the genus ''
Scleropages ''Scleropages'' is a genus of fish in the family (biology), family Osteoglossidae found in Asia and Australia. All of these species are carnivorous and have great jumping ability. These species are highly valued as aquarium fish, particularly by ...
'' separated about 140 million years ago, during the
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
. Originally, it was thought that 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 ...
150 – 30 million years ago was the evolutionary cause of the trans-continental distribution of the osteoglossomorphs. However, minimum ages of intercontinental clades and presence of marine forms in the fossil records imply that ancestral trans-oceanic dispersal is possible. Tests of these hypotheses are currently inconclusive as they are dependent on an ''
a priori ('from the earlier') and ('from the later') are Latin phrases used in philosophy to distinguish types of knowledge, Justification (epistemology), justification, or argument by their reliance on experience. knowledge is independent from any ...
'' calibrated age of crown-group
Teleostei 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 ...
, about which fossil and molecular evidence disagree. I.e., hypotheses do not fail only if Teleostei are of
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 ...
origin, but molecular inferences push crown ages further back.


Use by humans

Both ''Arapaima'' and ''Heterotis'' are farmed in their respective regions as relatively large and hardy food-fish.


Etymology

The family is named after the
monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unisp ...
genus ''Arapaima'', whose name derives from the Tupi-Guyarana indigenous name for ''Arapaima gigas''. File:Arapaima gigas in aqua park.jpg, ''A. gigas'' in aquarium File:Arapaima gigas at Beijing aquarium.JPG, ''A. gigas'' in aquarium File:Heterotis niloticus.png, ''H. niloticus'' in aquarium File:Heterotis niloticus MHNT ICHT 1995 69.jpg, Museum specimen of ''H. niloticus''


References

{{Taxonbar, from1=Q107552925, from2=Q525579, from3=Q18910351 Osteoglossidae Ray-finned fish subfamilies