Catfish

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Catfish (order Siluriformes) are a very diverse group of bony fish. Named for their prominent barbels, which give the image of cat-like whiskers, catfish range in size and behavior from the heaviest, the Mekong giant catfish in Southeast Asia and the longest, the wels catfish of Eurasia, to detritivores (species that eat dead material on the bottom), and even to a tiny parasitic species commonly called the candiru, Vandellia cirrhosa. There are armour-plated types and also naked types, neither having scales. Despite their common name, not all catfish have prominent barbels; what defines a fish as being in the order Siluriformes are in fact certain features of the skull and swimbladder. Catfish are of considerable commercial importance; many of the larger species are farmed or fished for food, and some are exploited for sport fishing, including a kind known as noodling. Many of the smaller species, particularly the genus Corydoras, are important in the aquarium hobby.

[edit] Taxonomy

The catfishes are a monophyletic group. This is supported by molecular evidence.^[1]

Catfish belong to a superorder called the Ostariophysi, which also includes the Cypriniformes, Characiformes, Gonorynchiformes and Gymnotiformes, 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.^[2] As of 2007 there are about 36 extant catfish families, and about 3,023 extant species have been described.^[3] This makes the catfish order the second or third most diverse vertebrate order; in fact, 1 out of every 20 vertebrate species is a catfish.^[4]

The taxonomy of catfishes is quickly changing. In a 2007 paper, Horabagrus, Phreatobius, and Conorhynchos were not classified under any current catfish families.^[3] 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. Also, 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.^[2][5][6][7] Many sources do not list the recently revised family Anchariidae.^[8] The family Horabagridae, including Horabagrus, Pseudeutropius, and Platytropius, is also not shown by some authors but presented by others as a true group.^[1] 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. On the other hand, our understanding of catfishes should increase in the next few years due to work by the ACSI.^[2]

The rate of description of new catfishes is at an all-time high. Between 2003 and 2005, over 100 species have been named, a rate three times faster than that of the past century.^[9] In June, 2005, researchers named the newest family of catfish, Lacantuniidae, only the third new family of fish distinguished in the last 70 years (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 Chiapas, Mexico.^[10]

[edit] Relationships between families

The relationship between the families is relatively unknown.^[11] Classifications of superfamilies varies. Many catfish families are classified into their own superfamilies.^[2]

Based on morphological data, Diplomystidae is usually considered to be the most primitive of catfishes and the sister group to the remaining catfishes, grouped in a clade called Siluroidei. Recent molecular evidence contrasts the prevailing hypothesis, where the suborder Loricarioidei are the sister group to all catfishes, including Diplomystidae (Diplomystoidei) and Siluroidei; though they were not able to reject the past hypothesis, the new hypothesis is not unsupported. Siluroidei was found to be monophyletic without Loricarioid families or Diplomystidae with molecular evidence; morphological evidence is unknown that supports Siluroidei without Loricarioidea.^[1]

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.^[12]

[edit] Evolution

A number of catfish fossils are known. Catfishes often have large, heavy bones that lend themselves to fossilization and, comparatively large otoliths. As such, a large number of species of catfishes have been named from complete or partial skeletal fossils or even from only otoliths.^[3] 19 valid genera and 72 species are based exclusively on fossil remains.^[3] There are two fossil families, Andinichthyidae, from the Lower Maastrichtian to Paleocene, as well as Hypsidoridae, from the Middle Eocene.^[2]

The earliest known catfish are known from the late Campanian-early Maastrichtian of Argentina.^[13] Catfish fossils are known from every continent except Australia.^[2] Fossils of the Eocene period have been found from Seymour Island in Antarctica.^[4] The order dispersed early throughout the continents primarily through land bridges.^[13] Australian species of catfish are all species from families that can enter saltwater; these fish traveled to Australia through saltwater, and then reverted to a freshwater lifestyle.^{[citation needed]}

The catfish must have spread through Africa to Asia during the late Jurassic if they were to reach Asia.^[13] During the Cretaceous period, the rift between South America and Africa would be forming; this may explain the contrast in families between the two continents. Most of the freshwater catfish of the two continents appear to be completely unrelated. Their relatively low diversity in Africa may explain why some primitive fish families coexist with them while they are absent in South America, where the more primitive fish may have been driven extinct.^[13] The earliest they could have spread into Central America was the late Miocene.^[13]

[edit] Distribution and habitat

Extant catfish species live in inland or coastal waters of every continent except Antarctica. Catfish have inhabited all continents at one time or another.^[2] Catfish are most diverse in tropical South America, Africa, and Asia.^[4] More than half of all catfish species live in the Americas. They are the only ostariophysans that have entered freshwater habitats in Madagascar, Australia, and New Guinea.^[14]

They are found primarily in freshwater environments of all kinds, though most inhabit shallow, running water habitats.^[14] Representatives of a at least eight families are hypogean (live underground) with three familes that are also troglobitic (inhabiting caves). Thus, catfishes are some of the most successful cave colonizers among fishes.^[15][16] One such species is Phreatobius cisternarum, known to live underground in phreatic habitats.^[17] Numerous species from the families Ariidae and Plotosidae, and a few species from among the Aspredinidae and Bagridae, are also found in marine environments.^[18][19]

[edit] Ecology

Most catfish are benthic in nature, meaning they normally associate with the bottom of the water column.^[14] However, variety of other lifestyles are also represented among the catfishes. A few species are pelagic in nature.^[14]

A wide range of feeding behaviors and diets are represented by the catfishes. In the family Trichomycteridae alone, there are species that feed on algae, fish scales, mucus, carrion, or even blood in the infamous candirú.^[20] Panaque and some species of Hypostomus are unique among catfishes in that are the only fishes able to eat and digest wood.^[21] Members of the aspredinid genus Amaralia are known to specialize in feeding on loricariid eggs.^[22]

Representatives of several catfish families utilize their pectoral spines to produce stridulatory sounds by rubbing a ridged process of the pectoral spine within the cleithral groove, including members of Aspredinidae, Mochokidae, Doradidae, Pimelodidae, and Ictaluridae. Catfishes make a "creaking" sound during defense or appeasement behavior when being attacked by conspecifics. They also vocalize when they are captured or prodded.^[23]

In catfishes, fertilization of eggs can be internal, external, or even include sperm passage through female digestive tracts, the so called sperm drinking type of fertilization.^[24] Internal insemination is probable in all species of Auchenipteridae.^[2] Catfishes express varying levels of care reproductive strategies. In loricariids, parental care is usually well-developed and the male guards the eggs and sometimes the larvae, either carrying eggs or having the eggs attached to the underside of rocks or in cavities.^[25] In most of Ariidae, if not all species, the male is a mouthbrooder; he carries the relatively large eggs in his mouth until the young hatch.^[2]

[edit] Physical characteristics

[edit] External anatomy

Most catfish are adapted for a benthic lifestyle. In general, they are negatively buoyant, which means that they will usually sink rather than float due to a reduced gas bladder and a heavy, bony head.^[14] Catfish have a variety of body shapes, though most have a cylindrical body with a flattened ventrum to allow for benthic feeding.^[14]

A flattened head allows for digging through the substrate as well as perhaps serving as a hydrofoil. Most have a mouth that can expand to a large size and contains no incisiform teeth; catfish generally feed through suction or gulping rather than biting and cutting prey.^[14] However, some families, notably Loricariidae and Astroblepidae, have a suckermouth that allows them to fasten themselves to objects in fast-moving water. Catfish also have a maxilla reduced to a support for barbels; this means that they are unable to protrude their mouths as other fish such as carp.^[14]

Catfish may have up to four pairs of barbels: nasal, maxillary (on each side of mouth), and two pairs of chin barbels, although pairs of barbels may be absent, depending on the species. Because their barbels are more important in detecting food, the eyes on catfish are generally small. Like other ostariophysans, they are characterized by the presence of a Weberian apparatus.^[2] Their well-developed Weberian apparatus and reduced gas bladder allow for improved hearing as well as sound production.^[14]

Catfish have no scales; their bodies are often naked. In some species, the mucus-covered skin is used in cutaneous respiration, where the fish breathes through its skin.^[14] In some catfishes, the skin is covered in bony plates called scutes; some form of body armor has evolved a number of times within the order. In loricarioids and in the Asian genus Sisor, the armor is primarily made up of one or more rows of free dermal plates. Similar plates are found in large specimens of Lithodoras. These plates may be supported by vertebral processes, as in scoloplacids and in Sisor, but the processes never fuse to the plates or form any external armor. By contrast, in the subfamily Doumeinae (family Amphiliidae) and in hoplomyzontines (Aspredinidae), the armor is formed solely by expanded vertebral processes that form plates. Finally, the lateral armor of doradids, Sisor, and hoplomyzontines consists of hypertrophied lateral line ossicles with dorsal and ventral lamina.^[26]

All catfish, except members of Malapteruridae (electric catfish), possess a strong, hollow, bonified leading spine-like ray on their dorsal and pectoral fins. As a defense, these spines may be locked into place so that they stick outwards, which can inflict severe wounds.^[4] In several species catfish can use these fin rays to deliver a stinging protein if the fish is irritated.^[27] This venom is produced by glandular cells in the epidermal tissue covering the spines.^[2] In members of the family Plotosidae, and of the genus Heteropneustes, this protein is so strong it may hospitalize humans unfortunate enough to receive a sting; in Plotosus lineatus, the stings may result in death.^[2]

Juvenile catfishes, like most fishes, have relatively large heads, eyes and posterior median fins in comparison to larger, more mature individuals. These juveniles can be readily placed in their families, particularly those with highly derived fin or body shapes; in some cases identification of the genus is possible. As far as known for most catfish, features that are often characteristic of species such as mouth and fin positions, fin shapes, and barbel lengths show little difference between juveniles and adults. For many species, pigmentation pattern is also similar in juveniles and adults. Thus, juvenile catfishes generally resemble and develop smoothly into their adult form without distinct juvenile specializations. Exceptions to this are the ariid catfishes, where the young retain yolk sacs late into juvenile stages, and many pimelodids, which may have elongated barbels and fin filaments or coloration patterns.^[28]

Sexual dimorphism is reported in about half of all families of catfish.^[29] The modification of the anal fin into an intromittent organ (in internal fertilizers) as well as accessory structures of the reproductive apparatus (in both internal and external fertilizers) have been described in species belonging to 11 different families.^[24]

[edit] Size

Catfish have one of the greatest range in size within a single order of bony fish.^[14] Many catfish have a maximum length of under 12 cm.^[2] Some of the smallest species of Aspredinidae and Trichomycteridae reach sexual maturity at only 10 millimetres (.4 in).^[4]

The wels catfish, Silurus glanis, is the only native catfish species of Europe, besides the much smaller related Aristotle's catfish found in Greece. Mythology and literature record wels catfish of astounding proportions, yet to be proven scientifically. The average size of the species is about 1.2–1.6 m (3.9–5.2 ft), and fish more than 2 m (6.6 ft) are very rare. The largest specimens on record measure more than 2.5 m (8.2 ft) in length and sometimes exceeded 100 kg (220 lb).

The largest Ictalurus furcatus, caught in the Mississippi River on May 22, 2005, weighed 56.25 kg (124 lb). The largest flathead catfish, Pylodictis olivaris, ever caught was in Independence, Kansas, weighing 56 kg (123 lb 9 oz). However, these records pale in comparison to a giant Mekong catfish caught in northern Thailand in May 1, 2005 and reported to the press almost 2 months later that weighed 293 kg (646 lb). This is the largest giant Mekong catfish caught, but only since Thai officials started keeping records in 1981.^[30] The giant Mekong catfish are not well studied since they live in developing countries and it is quite possible that they can grow even larger.

[edit] Internal anatomy

In many catfishes, the humeral process is a bony process extending backward from the pectoral girdle immediately above the base of the pectoral fin. It lies beneath the skin where its outline may be determined by dissecting the skin or probing with a needle.^[31]

The retina of catfish are composed of single cones and large rods. Many catfish have a tapetum lucidum which may help enhance photon capture and increase low-light sensitivity. Double cones, though present in most teleosts are absent from catfish.^[32]

The anatomical organization of the testis in catfish is variable among the families of catfish, but the majority of them present fringed testis: Ictaluridae, Claridae, Auchenipteridae, Doradidae, Pimelodidae, and Pseudopimelodidae.^[33] In the testes of some species of Siluriformes, organs and structures such as a spermatogenic cranial region and a secretory caudal region are observed, in addition to the presence of seminal vesicles in the caudal region.^[34] The total number of fringes and their length are different in the caudal and cranial portions between species.^[33] Fringes of the caudal region may present tubules, in which the lumen is filled by secretion and spermatozoa.^[33] Spermatocysts are formed from cytoplasmic extensions of Sertoli cells; the release of spermatozoa is allowed by breaking of the cyst walls.^[33]

The occurrence of seminal vesicles, in spite of their interspecific variability in size, gross morphology and function, has not been related to the mode of fertilization. They are typically paired, multi-chambered, and connected with the sperm duct, and have been reported to play a glandular and a storage function. Seminal vesicle secretion may include steroids and steroid glucuronides, with hormonal and pheromonal functions, but it appears to be primarily constituted of mucoproteins, acid mucopolysaccharides, and phospolipids.^[24]

Fish ovaries may be of two types: gymnovarian or cystovarian. In the first type, the oocytes are released directly into the coelomic cavity and then eliminated. In the second type, the oocytes are conveyed to the exterior through the oviduct.^[34] Many catfish are cystovarian in type, including Pseudoplatystoma corruscans, P. fasciatum, Lophiosilurus alexandri, and Loricaria lentiginosa.^[33][34]

[edit] Catfish as food

Catfish have been widely caught and farmed for food for hundreds of years in Africa, Asia, and Europe. Judgements as to the quality and flavor vary, with some food critics considering catfish as being excellent food, others dismiss them as watery and lacking in flavour.^[35] In Central Europe, catfish were often viewed as a delicacy to be enjoyed on feast days and holidays. Migrants from Europe and Africa to the United States brought along this tradition, and in the southern United States catfish is an extremely popular food. The most commonly eaten species in the United States are the channel catfish and blue catfish, both of which are common in the wild and increasingly widely farmed. Catfish is eaten in a variety of ways; in Europe it is often cooked in similar ways to carp, but in the United States it is typically breaded with cornmeal and fried.^[35] In Indonesia catfish are very popular food. They are usually served grilled in street stalls called warung and eaten with vegetables, the dish is called Pecel Lele (Lele is the Indonesian word for catfish). The iridescent shark is a common food fish in parts of Asia.^{[citation needed]}

Catfish is also high in Vitamin D.^[36]

[edit] Aquaculture

Catfish are easy to farm in warm climates, leading to inexpensive and safe food in local grocers. Ictalurids are cultivated in North America (especially in the Deep South, with Mississippi being the largest domestic catfish producer).^[37] Channel catfish (Ictalurus punctatus) supports a $450 million/yr aquaculture industry.^[4]

In Asia, many catfish species are important as food. Several walking catfish (Clariidae) and shark catfish (Pangasiidae) species are heavily cultured in Africa and Asia. Exports of one particular shark catfish species from Vietnam, Pangasius bocourti, has met with pressures from the U.S. catfish industry. In 2003, The United States Congress passed a law preventing the imported fish from being labeled as catfish.^[38] As a result, the Vietnamese exporters of this fish now label their products sold in the U.S. as "basa fish."

There is a large and growing ornamental fish trade, with hundreds of species of catfish, such as Corydoras and armored suckermouth catfish (often called plecos), being a popular component of many aquaria. Other catfish commonly found in the aquarium trade are banjo catfish, talking catfish, and long-whiskered catfish.

[edit] Catfish as invasive species

Representatives of the genus Ictalurus have been misguidedly introduced into European waters in the hope of obtaining a sporting and a food resource. However, the European stock of American catfishes has not achieved the dimensions of these fishes in their native waters, and have only increased the ecological pressure on native European fauna. Walking catfish has also been introduced in the freshwaters of Florida, with the voracious catfish becoming a major alien pest there. Flathead catfish, Pylodictis olivaris, is also a North American pest on Atlantic slope drainages.^[4] Pterygoplichthys species, released by aquarium fishkeepers, has also established feral populations in many warm waters around the world.^{[39][40][41][42][43]}