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Catfish

Catfish (or catfishes; order Siluriformes /sɪˈljʊərɪfɔːrmz/ or Nematognathi) are a diverse group of ray-finned fish. Named for their prominent barbels, which resemble a cat's whiskers, catfish range in size and behavior from the three largest species alive, the Mekong giant catfish from Southeast Asia, the wels catfish of Eurasia, and the piraíba of South America, to detritivores (species that eat dead material on the bottom), and even to a tiny parasitic species commonly called the candiru, Vandellia cirrhosa. Neither the armour-plated types nor the naked types have scales. Despite their name, not all catfish have prominent barbels or "whiskers". Members of the Siluriformes order are defined by features of the skull and swimbladder. Catfish are of considerable commercial importance; many of the larger species are farmed or fished for food. Many of the smaller species, particularly the genus Corydoras, are important in the aquarium hobby. Many catfish are nocturnal,[6][7] but others (many Auchenipteridae) are crepuscular or diurnal (most Loricariidae or Callichthyidae, for example).

Taxonomy

Molecular evidence suggests that in spite of the great morphological diversity in the order, all catfish form a monophyletic group.[8] Catfish belong to a superorder called the Ostariophysi, which also includes the Cypriniformes (carps and minnows), Characiformes (characins and tetras), Gonorynchiformes (milkfish and beaked salmons) and Gymnotiformes (South American knifefish), a superorder characterized by the Weberian apparatus. Some place Gymnotiformes as a sub-order of Siluriformes; however, this is not as widely accepted. Currently, the Siluriformes are said to be the sister group to the Gymnotiformes, though this has been debated due to more recent molecular evidence.[9] As of 2007 there were about thirty-six extant catfish families, and about 3,093 extant species have been described.[10] This makes the catfish order the second or third most diverse vertebrate order; in fact, one out of every twenty vertebrate species is a catfish.[11]

Catfish are believed to have a Gondwanan origin primarily centered around South America, as the most basal living catfish groups are known from there. The earliest known definitive members lived in the Americas from the Campanian to Maastrichtian stages of the Late Cretaceous, including the Andinichthyidae, Vorhisia vulpes and possibly Arius.[1][12][13] A potential fossil record is known from the earlier Coniacian-Santonian stages in Niger of West Africa,[2] though this has been considered unreliable,[13] and the putative earliest armored catfish known from the fossil record, Afrocascudo, lived during the Cenomanian age of the Late Cretaceous in Morocco of North Africa (Kem Kem Group).[3] The describers of Afrocascudo claimed that the presence of a derived loricariid so early on would indicate the extensive diversification of catfish, or at least loricarioids, prior to the beginning of the Late Cretaceous. As extant loricariids are only known from South America, much of this diversification must have occurred on the supercontinent of West Gondwana prior to its fragmentation into South America and Africa.[3] Britz and colleagues suggested that Afrocascudo instead represents a juvenile obaichthyid lepisosteiform, possibly a junior synonym of Obaichthys.[14] The authors of the original study still stood by their original conclusion based on the absence of important holostean characters, and noted that it could not be a juvenile, since the bones were completely ossified.[15]

Blue catfish (Ictalurus furcatus) skeleton on display at the Museum of Osteology.

The taxonomy of catfish is quickly changing. In a 2007 and 2008 paper, Horabagrus, Phreatobius, and Conorhynchos were not classified under any current catfish families.[10] There is disagreement on the family status of certain groups; for example, Nelson (2006) lists Auchenoglanididae and Heteropneustidae as separate families, while the All Catfish Species Inventory (ACSI) includes them under other families. FishBase and the Integrated Taxonomic Information System lists Parakysidae as a separate family, while this group is included under Akysidae by both Nelson (2006) and ACSI.[9][16][17][18] Many sources do not list the recently revised family Anchariidae.[19] The family Horabagridae, including Horabagrus, Pseudeutropius, and Platytropius, is not shown by some authors but presented by others as a true group.[8] Thus, the actual number of families differs between authors. The species count is in constant flux due to taxonomic work as well as description of new species.[9] Between 2003 and 2005, over one hundred species were named, a rate three times faster than that of the past century.[20] In June 2005, researchers named the newest family of catfish, Lacantuniidae, only the third new family of fish distinguished in the last seventy years, the others being the coelacanth in 1938 and the megamouth shark in 1983. The new species in Lacantuniidae, Lacantunia enigmatica, was found in the Lacantun river in the Mexican state of Chiapas.[21]

The higher-level phylogeny of Siluriformes has gone through several recent changes, mainly due to molecular phylogenetic studies. While most studies, both morphological and molecular, agree that catfishes are arranged into three main lineages, the relationship among these lineages has been a contentious point in which these studies, performed for example by Rui Diogo, differ.[22][23][24][25][26] The three main lineages in Siluriformes are the family Diplomystidae, the denticulate catfish suborder Loricarioidei (containing the Neotropical "suckermouth" catfishes), and the suborder Siluroidei, which contains the remaining families of the order. According to morphological data, Diplomystidae is usually considered to be the earliest branching catfish lineage and the sister group to the other two lineages, Loricarioidei and Siluroidei.[25][26][27] Molecular evidence usually contrasts with this hypothesis, and shows the suborder Loricarioidei as the earliest branching catfish lineage, and sister to a clade that includes the Diplomystidae and Siluroidei; this phylogeny has been obtained in numerous studies based on genetic data.[8][22][23][28] However, it has been suggested that these molecular results are errors as a result of long branch attraction, incorrectly placing Loricarioidei as the earliest-branching catfish lineage.[24] When a data filtering method[29] was used to reduce lineage rate heterogeneity (the potential source of bias) on their dataset, a final phylogeny was recovered which showed the Diplomystidae are the earliest-branching catfish, followed by Loricarioidei and Siluroidei as sister lineages, providing both morphological and molecular support for Diplomystidae being the earliest branching catfish.[30]

Below is a list of family relationships by different authors. Lacantuniidae is included in the Sullivan scheme based on recent evidence that places it sister to Claroteidae.[31]

Phylogeny

Phylogeny of living Siluriformes based on 2017[32] and extinct families based on Nelson, Grande & Wilson 2016.[33]