any member of what is widely believed to be the most primitive group of bony fishes (bony fish). This reputation stems from their rudimentary caudal skeleton and the lack of a set of intermuscular bones throughout the abdominal and anterior caudal regions of the body. Osteoglossomorphs are unique among fishes (fish) in that they use their tongue as the opposing surface for the teeth when they bite into food. They are divided into two orders, Osteoglossiformes and Hiodontiformes. Osteoglossiforms are a group of morphologically and biologically diverse forms primarily found in freshwater environments in Africa, Asia, South America, Australia, and islands of the tropical Pacific Ocean; however, a few osteoglossiform species enter slightly brackish water. In contrast, hiodontiforms occur only in North America and are probably the most primitive group of living osteoglossomorphs.

General features
 Osteoglossomorphs include five extant families and more than 200 species. Although the group is of little importance to humans, certain osteoglossomorph species are sometimes sought commercially as food fishes (fish) in parts of Africa, Asia, and South America.

      Osteoglossiforms are tropical fishes. The families Mormyridae (mormyrid) (elephant-snout fishes, mormyrs) and Gymnarchidae are confined to Africa; the Notopteridae (notopterid) (featherbacks) occur in Africa, Southeast Asia, and India. The distribution of the Osteoglossidae (such as the pirarucu [Arapaima], the arowana [Scleropages], and the butterfly fish [Pantodon]) in Africa, South America, and Australasia (believed by many authorities to have once been joined as a single landmass called Gondwana) is of particular zoogeographical interest.

      The pirarucu of the Amazon (A. gigas), one of the world's largest freshwater fishes, attains a length of 3 metres (about 10 feet); other osteoglossomorphs—for example, certain mormyrids—are only a few centimetres long.

      In contrast, the hiodontiforms are limited to North America and are represented by only two species—the goldeye (Hiodon alosides) and the mooneye (H. tergisus). Mooneyes may grow to lengths of 42.5 cm (17 inches).

Natural history

Life cycle and reproduction
      A variety of breeding habits have evolved among theosteoglossomorphs. In some species there is considerable care of the young by the parents. Although no species is known to undertake extensive breeding migrations, many leave the usual habitat and move into floodplains or streams at breeding time.

      The breeding biology of the mormyrids (mormyrid) has been little studied; it does not seem likely, however, that they prepare spawning nests or exercise much parental care. In contrast, Gymnarchus niloticus (Gymnarchidae) prepares a large floating nest from the matted stems of swamp grasses, biting off the stems and fashioning them into a trough-shaped structure with an internal length of about 50 cm (20 inches). Spawning takes place in the nest, and one or both parents guard the developing young for approximately 18 days.

      Nests are not made by notopterids (notopterid), but they do establish a breeding territory. Both Hiodon species ( goldeye and mooneye) spawn in the spring. Eggs are laid on gravel or rocks in shallow, quiet water, which the adults reached after a short migration. The young of the goldeye remain there until late summer before migrating downstream. In Chitala chitala (Notopteridae), one parent, probably the male, clears an area of the bottom near some submerged object (such as a rock, plant stem, or piling), on which the eggs are later laid in circular bands; the male guards the developing embryos.

      Members of the Osteoglossidae care for their young in a variety of ways. The South American arawana (Osteoglossum bicirrhosum) and its Indo-Australian relatives, the spotted bonytongue (Scleropages leichardti) and the arowana (S. formosus), carry the eggs and young in the mouth of one parent; little else is known of their breeding habits. The African arowana (Heterotis niloticus) prepares a crude nest from grasses in newly flooded swamp plains. The male guards the young and leads them from the nest on feeding excursions. Both sexes of Arapaima gigas (pirarucu) of South America dig a spawning pit and guard the developing embryos, which hatch and leave the pit after about seven days; care is provided by the male, around whose head the young congregate. The dark colour of the male's head seems to provide good camouflage. This colour is the chief stimulus for forming shoals—that is, groups or schools. Apparently, there is also a gustatory (taste) or olfactory (smell) stimulus in a secretion from glands on the male's head. Parent-young groups persist for two to three months; if the male dies, the young join other shoals.

      Spawnings in aquaria suggest that the African butterfly fish (Pantodon) of family Pantodontidae produces floating eggs and provides minimal parental care. A complicated courtship in this species has also been observed.

Behaviour and ecology
      Mormyridae and Gymnarchidae are of particular interest because they have electrical (bioelectricity) organs. Electrical discharges from these organs are in the form of pulses, the frequency and nature of which are different in each species. In nature, electrical discharges ranging from an output of about 120 to 300 pulses per second have been recorded.

      The flow pattern of the electric field around the fish is distorted by the different conductivity of objects that pass into it. These variations are detected by modified nerve cells (mormyromasts) in the skin. The greatly varying conductivity differences among animals and inorganic objects make it possible for mormyrids to use their electrical organs to distinguish between prey, predators, and obstacles in the turbid water they often inhabit. Discharges from the organs also serve as signals to other mormyrids.

      Some mormyrs tend to swim with little body movement, using instead the dorsal fins for propulsion. This unusual swimming method is probably associated with the use of electric organs in navigation and detection; Gymnarchus, for example, swims with its body held straight, propulsion being provided by undulations of the dorsal, or back, fin. Since electrical organs lie near the tail, side-to-side movements of the tail end (as in normal swimming movement) would constantly change their position relative to that of the receptor organs, which are in the head area.

      Rigid-bodied swimming like that of the mormyrs also occurs in the featherbacks (Notopteridae), which use the long anal fin for propulsion. There is no electrical organ in the notopterids, however; the rigid body of these fishes may be correlated with the long gas-filled swim bladder that extends into the tail end of the body.

      All other osteoglossomorphs swim by using the body musculature and caudal (tail) fin in the usual manner. Pantodon has greatly expanded winglike pectoral fins (behind the gills (gill)), which are used for short flights in the air, either to escape predators or to catch insects. It habitually swims or drifts just below the water surface and leaps from the water by means of a powerful thrust of the pectorals, sending the fish 30 cm (about 12 inches) or more vertically out of the water. Short horizontal flights of about 1 metre (about 3 feet) are also executed.

      Some species of Osteoglossidae (Heterotis, Arapaima, and Pantodon), Notopteridae (Chitala, Notopterus, Papyrocranus, Xenomystus), and the Gymnarchidae are able to breathe air at the surface; thus, they can live in areas where the water is deoxygenated.

      The notopterid Xenomystus produces sounds that are used as warnings and in courtship. Swim bladder structure in other Notopteridae suggests that they are also capable of emitting sounds. In all Notopteroidei and the Hiodontiformes, the swim bladder is closely connected with the inner ear, a condition that may be an aid to hearing. Except for the osteoglossid Heterotis, all osteoglossomorphs are carnivorous—the smaller species (and young of all species) feeding on insects and other invertebrates, the larger species on fish. Heterotis feeds on microscopic plants and animals filtered from the water.

      Osteoglossomorph fishes occupy a diversity of habitats in rivers and lakes, often in turbid waters or in regions with dense aquatic vegetation. A few species seem to require open waters, and some notopterids can even tolerate slightly brackish water.

Form and function
      All living osteoglossomorphs have strongly toothed jaws. The jaws are not protrusible, but in piscivorous (that is, fish-eating) species the gape is sometimes considerable. The Mormyridae (mormyrid) show a great variety in head shape and mouth form. Many insectivorous (insect-eating) species have a small mouth at the tip of a long, curved, tubular snout; they feed by probing among rocks or in soft mud. Piscivores have larger mouths and short snouts.

      The mouth is large in all Osteoglossidae, which, like the Notopteridae (notopterid) and the Hiodontiformes, have a more typical head shape than do the Mormyridae. Heterotis has complex spiraled structures lying above the gill arches on each side of the head and opening into the pharynx. Food particles, drawn into the pharynx with the respiratory current, are filtered out and concentrated in these organs.

      In all Mormyridae and in the Gymnarchidae, a short length of body musculature toward the tail is modified to form the electric organ. These muscles have lost the ability to contract and have undergone considerable cellular reorganization, from slender fibres of ordinary muscle into thin, flat, electroplates, the structures in which electricity is produced. There are about 300 to 400 electroplates in mormyrid electric organs and about 600 to 800 in those of Gymnarchus. The brain, particularly the cerebellum, of mormyrs is the largest known among fish; the cerebellum is associated with the electroreceptor organs (mormyromasts) in the skin (see mechanoreception: Ampullary lateral-line organs (mechanoreception)).

      A well-developed swim bladder is present in all osteoglossomorphs. In the air-breathing osteoglossids, pantodontids, and gymnarchids, it is highly vascularized (that is, has many blood vessels), and its inner surface is honeycombed with tiny pits.

      Paired extensions in the Notopteridae and Hiodontidae connect the swim bladder with the auditory region of the skull. In the African genus Papyrocranus, diverticula (outpocketings) of the swim bladder actually penetrate the skull, a condition that probably improves hearing. Posteriorly, the swim bladder in notopterids extends beyond the abdominal cavity and runs back on either side of the vertebral column. In the early embryos of Mormyridae and Gymnarchidae, a pair of thin tubes extends forward from the swim bladder into the skull; later the tubes atrophy, leaving an isolated vesicle—a blister, or balloonlike structure—within the skull surrounded by the semicircular canals of each ear.

      The inner ear shows various modifications; in notopterids, gymnarchids, and mormyrids, the upper portion (for balance) is completely separated from the lower part (for hearing).

      In the Mormyridae and the genus Pantodon (Osteoglossidae), the anal fin shows considerable sexual dimorphism in shape—that is, the anal fin of males differs from that of females. These differences may be related to spawning activity.


Annotated classification
      Classification in the superorder Osteoglossomorpha is based largely on skeletal characters, in particular the caudal fin skeleton, the bones around the eye, and the gill arches and their associated dentition. Details of the inner ear and swim bladder anatomy are also of importance. The classification used here is based on that of J.S. Nelson (2006). Groups indicated by a dagger (†) are extinct and known only from fossils.

Superorder Osteoglossomorpha
      Order Osteoglossiformes
 Primitive; well-developed teeth on tongue, skull base, and bones of the mouth cavity; caudal fin skeleton of characteristic form. Early Cretaceous to present.

      Suborder Osteoglossoidei
 Swim bladder not connected with skull; semicircular canals and lower part of inner ear connected; electric organs absent.

      Family Osteoglossidae (bonytongues (bony tongue) and arapaimas)
 Fishes of diverse body form; pectoral fins not greatly enlarged, pelvic fins abdominal in position. 6 genera, about 8 species. Living genera include Arapaima (1 species) and Osteoglossum (2 species), South America; Scleropages (2 species), Australia, New Guinea, Borneo, Sumatra, Malaysia, Thailand; Heterotis (1 species), Africa; and Pantodon (1 species), Africa.

      †Family Singididae
 Apparently toothless; monotypic genus (Singida) from Paleocene of Tanzania (East Africa).

      Suborder Notopteroidei
 Swim bladder connected with the skull; semicircular canals separate from lower part of ear or, if connected, utriculus greatly enlarged; no electric organs.

      †Family Lycopteridae
 Small freshwater fishes resembling hiodontiforms. 4 genera, 6 species. Early Cretaceous of northeastern Asia.

      Family Notopteridae (knifefishes (notopterid) or featherbacks).
 Long anal fin confluent with reduced caudal; dorsal fin small or absent. 8 species in 4 genera: Papyrocranus (2 species) and Xenomystus (1 species) in Africa, Chitala (4 species) and Notopterus (1 species) in Asia. Fossil Notopterus from Miocene and Late Paleocene of Sumatra.

      Suborder Mormyroidei
 With electrical organs; very diverse head shape and mouth form. 2 families, about 200 species. Confined to Africa; fossils from Pliocene of Egypt.

      Family Mormyridae (mormyrs (mormyrid) and elephant-snout fishes)
 Anal, caudal, and dorsal fins present; about 18 genera, approximately 200 species.

      Family Gymnarchidae
 No anal fin; long dorsal confluent with reduced caudal fin. 1 genus, 1 species (Gymnarchus niloticus).

      Order Hiodontiformes

      Family Hiodontidae (goldeyes and mooneyes (mooneye))
 Probably the most primitive living osteoglossomorphs. 1 genus (Hiodon), 2 species (H. alosoides and H. tergisus), confined to North America. Fossil from Eocene of British Columbia (Eohiodon rosei).

Critical appraisal
      Taxonomic problems of the osteoglossomorphs primarily concern intragroup relationships. The superorder as a whole presents problems of relationship with the other teleostean lineages. American ichthyologist Gloria Arratia, who claims that the caudal skeleton of the elopomorph, Elops, is more primitive, has challenged the basal position of osteoglossomorphs. Certain fossil groups—such as the Eocene genus Brychaetus and the Mesozoic families Cladocyclidae and Ichthyodectidae—are osteoglossomorphs, but their relationship with the living groups is still obscure. In addition, within the superorder, some classifications demote the hiodontiforms to a family (Hiodontidae) contained within the order Osteoglossiformes.

Peter Humphry Greenwood

Additional Reading
P.H. Greenwood, “On the Genus Lycoptera and Its Relationship with the Family Hiodontidae (Pisces, Osteoglossomorpha),” Bull. Br. Mus. Nat. Hist. (Zool.), 19:259–285 (1970); and et al., “Phyletic Studies of Teleostean Fishes, with a Provisional Classification of Living Forms,” Bull. Am. Mus. Nat. Hist., 131:339–455 (1966), includes a classification of the Osteoglossomorpha and a discussion of the reasons for the arrangement; E.S. Herald, Living Fishes of the World (1961, reprinted 1972), a popular, well-illustrated account of the various families and of some species, particularly their biology; K.H. Luling, “Arapaima, Giant Fish of Amazonas,” Animals, 11:222–225 (1968), a popular account; G.J. Nelson, “Infraorbital Bones and Their Bearing on the Phylogeny and Geography of Osteoglossomorph Fishes,” Am. Mus. Novit., no. 2394 (1969); “Gill Arches of Teleostean Fishes of the Division Osteoglossomorpha,” J. Linn. Soc. (Zool.), 47:261–277 (1968); J.R. Norman, A History of Fishes, 3rd rev. ed. by P.H. Greenwood (1977), includes a general account of osteoglossomorph biology, distribution, and anatomy (high school and college level); G. Sterba, Freshwater Fishes of the World, rev. ed., 2 vol. (1989; originally published in German, 1959), particularly for the aquarist.

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Universalium. 2010.

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