any member of a group of primitive bony fishes (bony fish) that make up one of the three major subdivisions of the superclass Actinopterygii (ray-finned fishes). Holosteans are represented today by the bowfins (bowfin) (order Amiiformes) of North America and the gars (gar) (order Semionotiformes) of North and Central America and Cuba. Holosteans diverged from their chondrostean ancestors in the order Palaeonisciformes during the Permian Period and were particularly abundant in the Mesozoic Era (251–65.5 million years ago); however, only three living genera remain. The genus Amia contains the single remaining species of bowfin, and the genera Lepisosteus and Atractosteus contain the seven living species of gars (gar). The extinct order Pycnodontiformes is often associated with the holosteans. There is disagreement as to whether infraclass Holosei should be recognized as a natural taxonomic entity, since many authorities believe that bowfins, gars, and their fossil relatives did not descend from a common ancestor.

General features
      Bowfins (bowfin) (also known as grindles, mudfishes, and dogfishes) are found in slow-moving waters from the Great Lakes to the Gulf of Mexico. Female bowfins reach a length of about 75 cm (30 inches) and weigh up to 3.5 kg (about 8 pounds); males are smaller. Bowfins eat all kinds of fish and invertebrates and are sometimes destructive to game fish populations. Bowfins are seldom caught as food fish. Relatives once common in Europe and Asia are extinct.

      Gars (gar) occur only in North America, Central America, and Cuba—from southeastern Canada to Panama—but are not found west of the Rocky Mountains. The longnose gar (Lepisosteus osseus) is the most widely distributed species. Gars are primarily freshwater fish that sometimes venture into salt water or brackish water. The alligator gar (Atractosteus spatula), one of the largest freshwater fishes, is particularly abundant in the Everglades region of southern Florida, where it is caught locally as a food fish. It sometimes grows to a length of nearly 3 metres (10 feet) and may attain a weight of 136 kg (300 pounds).

      The names gar, garfish, and garpike are sometimes also applied, especially in Europe, to the needlefishes (needlefish) (Belonidae). Needlefishes are coastal fishes of warm seas and have very long and slender jaws; however, they are not closely related to the Holostei.

Natural history

      Bowfins (bowfin) spawn in weedy areas along the edges of streams and lakes. The male constructs the nest and guards the eggs as well as the newly hatched young. The young bowfin has an adhesive organ at the tip of its snout that enables it to cling to weeds. The fish grows rapidly, and it may be as long as 23 cm (9 inches) by the end of its first year.

      In the spring, female gars (gar) lay large yolk-filled eggs in shallow water. The gar hatchlings grow rapidly, feeding on minnows. The long rows of needle-sharp teeth are effective in capturing fast-swimming prey.

      Gars and bowfins are voracious predators that feed on invertebrates and other fishes. All amiiform fishes (that is, the bowfin's extinct relatives) were probably predaceous. For the most part, the amiiforms were a marine group. The modern bowfin, however, is confined to fresh water. In addition, the bowfin is not limited to its gills (gill) for respiration. Its highly developed swim bladder can also function as a lung in times when the water temperature is high and oxygen concentration is low. In contrast, gars are known to occasionally venture into salt water, but apparently they do not attempt to feed there. Gars, which also possess a versatile swim bladder, often float quietly at the surface of slow-moving waters, breathing air.

Form and function

Skeletal features
      In the Holostei the dermal bone of the upper jaw (maxilla) is freed from the cheek elements, and its attachment to the skull only occurs in the ethmoid region or near the nasal chambers. The palate is separated from the cheek elements, and the adductor mandibulae muscle, which closes the jaws, is larger and more subdivided than it is in the Polypteriformes (bichirs (bichir)) and Acipenseriformes (sturgeons (sturgeon) and paddlefish). Primitively, the centrum (that is, the central and circular part of a vertebra) surrounding the notochord (a flexible rod that passes through the vertebral column) is absent, but this structure apparently developed independently in most of the holostean orders. The scales (scale) are primitively rhomboidal, or diamond shaped, and covered with an enamel-like substance; however, they have become thin and cycloidal (that is, rounded and overlapping) in several groups. The fin rays of the unpaired fins are always equal in number to their basal supports, and the fins themselves may or may not be bordered at the anterior end by fulcra, which are modified scales or spines. The caudal, or tail, fin is typically hemiheterocercal (that is, the body lobe turns up slightly) and externally symmetrical. The braincase is always composed of separate ossifications (centres of bone formation) that resemble, in number and placement, those found in the teleosts (teleost).

      The infraclass Holostei includes the orders Semionotiformes, Pycnodontiformes, and Amiiformes. In these orders the preoperculum (an L-shaped bone anterior to the operculum, or gill cover) is tied to the palatal elements and provides part of the originating area for the adductor mandibulae muscle.

Extant groups
      The order Seminonotiformes includes two families. The oldest known holostean, Acentrophorus (a form dating to the Late Permian, about 260–251 million years ago), belongs to the Semionotidae. Members of this family have small mouths and strong teeth, heavily ossified (that is, composed of true bone rather than cartilage) dermal bones, and hemiheterocercal tails. The body may be fusiform (tapered at both ends), as in Semionotus, or flat and disk-shaped, as in Dapedium.

      The other semionotiform family, the Lepisosteidae, includes the living gars (gar). The characteristic snout of the gar is greatly lengthened by multiplication of the small tooth-bearing bones anterior to the eye. The premaxilla bone is situated at the anterior end of the series; the maxilla bone itself elongated seems to be reduced to a small bony sliver at the angle of the mouth.

      The body of the gar is encased in an armour of thick, diamond-shaped, enamelled scales. The jaw ends in a beak that in the alligator gar (Atractosteus spatula) is broad and relatively short; in the longnose gar (Lepisosteus osseus) the beak is long and forcepslike. The dorsal and anal fins, both located far back on the body, are without spines and have fewer than 12 rays each.

      The Amiiformes, represented today by one species of bowfin (Amia calva), include about six families that showed considerable diversity in the length of the jaw, the development of the teeth and fins, and details of the dermal skull pattern. In general, the earlier amiiforms had well-developed rhombic scales and a persistent notochord. In later forms, including the extant Amia, the scales usually became thinner and cycloidal. Ossified centra developed around the notochord, either restricting it or eliminating it. The caudal fin was either forked or lobed. The amiiform body was generally fusiform, similar to that of the living bowfin.

      The bowfin has a long spineless dorsal fin with about 58 rays. This extends over most of the back to near the tail. The males have an orange- or yellow-encircled dark spot near the tail. In females either the outer circle or the entire marking is absent. Bony plates cover the head; the rest of the body has cycloid scales that are roughly circular and lack dentine and enamel layers.

Extinct groups
      The Pycnodontiformes, which may be related to the Semionotiformes, are unique among the holosteans in having their upper and lower dentitions modified to form an open pavement of crushing teeth. In many cases, however, the anterior teeth of the premaxilla and the dentary are incisiform and thus must have been used for grasping (as such teeth are in the living porgies (porgy)). In addition to skull modifications related to feeding, the pycnodonts are characterized by deep, almost disk-shaped bodies, elongated anal and dorsal fins, and an externally symmetrical caudal fin. In a number of genera scales are absent on the posterior part of the body, a condition that apparently increased flexibility. Scales were usually present but modified on the anterior half. The body and fin form of pycnodonts suggest that they were slow but highly maneuverable swimmers. The affinities of this order remain problematic as the ossification pattern of the braincase and the caudal-fin skeleton do not closely resemble those of other holosteans or halecostomes (a group that gave rise to the teleosts).

      The gars (gar) probably arose during the Cretaceous Period (145.5–65 million years ago) from some semionotid stock. They are known from freshwater deposits in India, Africa, North America, and Europe, dating from the Paleocene Epoch (65.5–56 million years ago). The bowfins (bowfin) also made their first appearance in Cretaceous times. Pycnodont fossils range from the Late Triassic to the Eocene (about 228–34 million years ago).

      Groups marked with a dagger (†) are extinct and known only from fossils.

Distinguishing taxonomic features
      The principal features on which classification of the Holostei is based include general body shape, scale structure, and the number and placement of head bones.

Annotated classification
Infraclass Holostei
 Tail hemiheterocercal; maxilla free of preopercle; rays of median fins approximately equal in number to basal elements; trend toward thinning of scales and loss of ganoid (enamel) layer.
      Order Amiiformes ( bowfin and fossil relatives)
 Body generally fusiform; carnivorous fishes, early forms with well-developed diamond-shaped scales covered with enameloid; scales thinner and cycloid in later forms; early forms with persistent notochord, centra developed later. 4 families, 1 living species. Middle Jurassic (about 176–161 million years ago) to present.

      Family Amiidae
 Dorsal fin long extending over most of back; tail rounded; scales thin and circular. 1 genus, 1 living species (Amia calva).

      Order Semionotiformes (gars (gar) and fossil relatives)
 Body of variable shape, some fusiform, others deep-bodied; cheek shows many tiny bones behind eye; scales well-developed and diamond-shaped covered with an enamel-like layer; 3 families, one of which (Lepisosteidae) is recent. Late Permian (about 260–251 million years ago) to present.

      Family Lepisosteidae
 Body elongate; jaws essentially a long snout and equipped with needlelike teeth; dorsal and anal fins located posteriorly on the body close to the tail. 2 genera (Lepisosteus and Atractosteus), 7 species.

      †Order Pycnodontiformes
 Deep-bodied fishes with dorsal and anal fins elongated; scales often absent from part of body; upper and lower teeth modified to form crushing pavementlike structures; front teeth often incisiform. Late Triassic to Eocene (about 228–34 million years ago).

Critical appraisal
      Long regarded as a sister group to the Chondrostei and Teleostei, the infraclass Holostei is not recognized by many authorities as a legitimate taxon, since the orders Amiformes, Semionotiformes, and their fossil relatives do not appear to descend collectively from a single ancestor. Some authorities state that amiiforms are more closely related to the Teleostei than they are to the Semionotiformes, as evidenced by the structure of the braincase. DNA analysis continues to investigate the genetic connections between the Amiiformes and the Semionotiformes. They are retained together in this classification as the Holostei chiefly because emerging molecular data suggest that there is a close relationship between them.

Peter L. Forey

Additional Reading
S.M. Andrews et al., “Pisces,” in W.B. Harland et al. (eds.), The Fossil Record: A Symposium with Documentation, ch. 26 (1967), a classification of fish, with first and last occurrences for each family; B.G. Gardiner, “A Revision of Certain Actinopterygian and Coelachanth Fishes, Chiefly from the Lower Lias,” Bull. Br. Mus. Nat. Hist. (Geol.), 4:239–384 (1960), important revised descriptions of early Jurassic fish from Great Britain; E.S. Goodrich, “Vertebrata craniata,” fasc. 1, “Cyclostomes and Fishes,” in E.R. Lankester (ed.), A Treatise on Zoology (1909, reprinted 1964), a classic work on the anatomy of fish that is still useful; J.P. Lehman, “Actinopterygii,” in J. Piveteau (ed.), Traité de paléontologie, vol. 4 (1966), a summary of important characteristics of the higher bony fishes, along with their geologic and geographic distributions; D.V. Obruchev (ed.), Fundamentals of Paleontology, vol. 11, Agnatha, Pisces (1967), a summary treatment of all fishes, living and fossil; D.H. Rayner, “The Structure and Evolution of the Holostean Fishes,” Biol. Rev., 16:218–237 (1941), an attempt to relate the various families of holostean fishes mainly on the basis of braincase design; A.C. Weed, The Alligator Gar (1923).

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Universalium. 2010.

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