chondrostean

▪ fish

Introduction

      any member of a group of primitive ray-finned bony fishes (bony fish) that make up one of the three major subdivisions of the superclass Actinopterygii, the other two being the holosteans (holostean) and the teleosts (teleost). The only living representatives are the sturgeons (sturgeon) and paddlefishes (paddlefish) (order Acipenseriformes) and the bichirs (bichir) and reedfish of Africa (order Polypteriformes). Fossil chondrostean species are known as palaeonisciforms (order Palaeonisciformes) and first appear in rocks near the end of the Silurian Period (about 419 million years ago).

      Subclass Chondrostei is not a natural group, since the acipenseriforms are genealogically more closely related to the Holostei than they are to the polypteriforms; however, acipenseriforms and polypteriforms are kept together for convenience. Although the relationships of the living forms with fossil forms are poorly known, different authorities usually retain the name chondrostean.

General features

      The chondrosteans were most numerous and possessed the greatest diversity during the last part of the Paleozoic Era and the beginning of the Mesozoic Era (some 251 million years ago). Extinct chondrosteans are known as palaeonisciforms, a label derived from a Greek word meaning “ancient scale.” Like the living members of Chondrostei, order Palaeonisciformes is not a natural group but rather a series of families connected by interrelationships that are poorly understood. With the rise of the holosteans (holostean) and teleosts (teleost) (the other two major subdivisions of the Actinopterygii) during the Mesozoic, the chondrosteans declined. By the end of the Cretaceous Period (some 65 million years ago), they had been reduced to a few genera, which survive today. The few living chondrosteans are highly specialized and have differentiated markedly from their Paleozoic ancestors. Consequently, comparisons between living forms and Paleozoic ones are difficult, and problems have arisen in classification as well as understanding the interrelationships between past and present forms.

      Living chondrosteans are contained within order Acipenseriformes and order Polypteriformes. The acipenseriforms include the living sturgeons (sturgeon) and paddlefishes (paddlefish) and are inhabitants of the Northern Hemisphere. Most acipenseriforms live in fresh water and immediate coastal waters.

      There are six genera of acipenseriforms. The genus Huso contains the kaluga (H. dauricus), which inhabits the Amur River basin in Asia, and the beluga (Huso huso), which is found in the Caspian and Black Sea basins. The beluga is one of the world's largest freshwater fish; specimens have been documented at 6 metres (about 20 feet) long, although an unconfirmed report places the size of the fish at 8 meters (26 feet) long and weighing up to 1,300 kg (about 2,900 pounds). The source of the world's prime caviar, the beluga is listed by the International Union for Conservation of Nature (IUCN) as an endangered species.

      Sturgeons are spread throughout the genera Acipenser, Scaphirhynchus, and Pseudoscaphirhynchus. The genus Acipenser contains approximately 17 species. Most of these species are Eurasian; however, there are five North American species. The common sturgeon (Acipenser sturio) is found along the European coast from Norway to the Mediterranean Sea. A closely related form, probably of the same species, is found along the east coast of North America from the St. Lawrence River to the Gulf of Mexico. The Baikal sturgeon (A. baerii) occurs in Lake Baikal and in nearby regions of Russia (Siberia), China, and Kazakhstan. A smaller species, the sterlet (A. ruthenus), inhabits the Black and Caspian seas. The starry sturgeon (A. stellatus) occurs in rivers leading to the Black Sea, the Sea of Azov, and the Caspian Sea. The lake sturgeon of North America (A. fulvescens) occurs in the Mississippi valley, the Great Lakes, and northward into Canada. The white, Oregon, or Sacramento sturgeon (A. transmontanus) inhabits the waters of the Pacific coast of North America from California to Alaska. The shovelnose sturgeons (genus Scaphirhynchus) occur in the Mississippi drainage system of North America, and the Aral Sea shovelnose sturgeons (Pseudoscaphirhynchus) are found in rivers that drain into the Aral Sea in Asia.

      Paddlefishes include two living species: Polyodon spathula in the Mississippi drainage basin and Psephurus gladius of the Yangtze River (Chang Jiang) basin in China. Both of these are highly distinctive with long paddle-shaped snouts. The Mississippi paddlefish grows to about 2.2 metres (7.2 feet); however, the Chinese paddlefish sometimes reaches 6.3 metres (about 21 feet) in length.

      The polypteriforms, which include the bichirs (bichir) (Polypterus) and the closely related reedfish (Erpetoichthys calabaricus), live in freshwater lakes and streams of western and central Africa. Polypteriforms are eel-shaped fishes covered with thick rhomboid scales. The largest species of bichir grows to about 70 cm (28 inches), and the reedfish reaches a length of 90 cm (35 inches).

Natural history

Reproduction and life cycle

      Sturgeons (sturgeon) ascend rivers in spring or summer to deposit their spawn. They are abundant in the rivers leading to the Black and Caspian seas and to the Sea of Azov during the two weeks of the upstream migration. Early in summer the fish migrate into the rivers or toward the shores of freshwater lakes in large shoals for breeding purposes. The eggs are small and numerous, and the growth of the young is rapid. After the sturgeon attains maturity, growth continues at a slow rate for several years. Some attain great age: observations made in Russia indicate that the beluga (Huso huso) may attain an age of 200 to 300 years. In addition, some sturgeons, such as those of the genus Scaphirhynchus, are entirely freshwater and make migrations to habitual spawning grounds.

      Paddlefishes (paddlefish) breed when seven or eight years old and spawn during spring floods. The larvae hatch in about two weeks and feed on their large yolk sac. The paddle, a long broad extension of the snout, is absent at birth but begins to appear after two or three weeks. Bichirs (bichir) initiate courtship by leaping from the water. However, little is known of their spawning habits. Young fish have external branching gills and are newtlike in appearance.

Ecology

      Sturgeons occur in both salt water and fresh water, although some species are restricted to fresh water. They are bottom feeders and spend much time foraging, dragging their tactile whiskerlike barbels over the bottom in search of small invertebrates and fishes. The American paddlefish (Polyodon) feeds by straining plankton (mostly tiny, drifting aquatic organisms) through its gill system and has been described as a living plankton net. The Chinese paddlefish (Psephurus) is more carnivorous and has shorter gill rakers. Bichirs and reedfish mainly inhabit the edges of streams and floodplains. They remain concealed by day and forage at night for worms, insect larvae, crustaceans, and small fishes.

Form and function

Extinct forms

      Most palaeonisciforms had fusiform (that is, tapered at both ends) bodies with blunt snouts, eyes situated far forward, pelvic fins located at about the middle of the body, dorsal (back) and anal (on the lower side) fins nearly opposite one another on the posterior part of the body, and heterocercal (that is, with the top lobe longer than the lower lobe) caudal fins. With few exceptions, their bodies were covered with rhomboidal (diamond-shaped) scales (scale) with a shiny enameloid layer. These scales, called ganoid scales, articulated with one another by a peg-and-socket joint; in some groups, the scales tended to become thin and cycloidal, or rounded, as in the coccolepids. The rays of the unpaired fins were usually more numerous than their basal supports, and all the fins were usually bordered by scales that were generally larger and stronger than other scales (fulcral scales). A few families, such as the Late Paleozoic platysomids and amphicentrids, evolved deep, compressed bodies with elongated anal and dorsal fins. A few, such as the Tarrasiids, had eel-like bodies.

      In all palaeonisciforms, the upper jaw was tied to the cheekbones, which completely covered the area between the eyes and the gill covers. The jaw suspension may have had an oblique orientation (associated with a wide mouth gape) or a nearly vertical orientation (associated with a relatively smaller gape). The teeth, where present, were usually small and needlelike. Some of the deep-bodied palaeonsiciforms showed grinding or cutting teeth. On structural grounds, there is reason to believe that the biting mechanism in palaeonisciforms was less powerful than that of the holosteans. In addition, the arrangement of the fins and the structure of the tail in paleonisciforms suggest that maneuverability in swimming was not as great as in either the holosteans (holostean) or the teleosts (teleost).

Extant forms

      Much of the internal skeleton of modern sturgeons (sturgeon) is made of cartilage, and it is for this reason that the group to which they belong is called chondrostei, which means “cartilage bone.” The modern sturgeon has thick bony plates on the head and five rows of enlarged scales (scutes) along the body: one along the back, one on each side above the pectoral fins, and one on each side near the belly. The tail fin is heterocercal. The mouth is subterminal (that is, behind and below the snout tip), and this and other specializations are clearly related to bottom feeding. The mouth is toothless and is preceded by four fleshy barbels; the protractile lips have taste buds surrounding them. The form of the snout becomes more blunt and abbreviated with age.

      The skeleton of the paddlefishes (paddlefish), like that of the sturgeons, has lost much of its ossification. The body is fusiform, the fins are well-developed, and the tail is heterocercal. The elongated paddle-shaped snout, which is composed entirely of cartilage, is one-third to one-half the total body length. The snout is covered with electroreception organs and thus is highly sensitive. The skin is smooth except for a few scattered vestigial scales at the base of the tail. The mouth is subterminal but large. The gills are equipped with comblike rakers to strain food particles out of the water. In addition, the Chinese paddlefish (Psephurus gladius) has tiny deeply embedded scales all over the body.

      The bichir is rather elongated in form, and the reedfish is eel-like; both have hard diamond-shaped scales. The dorsal fin is made up of a few to several separate finlets, and the tail is rounded. The upper body is brown, grayish, or greenish, the lower side often white or yellowish.

Evolution

      The long history of the chondrosteans, which extends over a period of 375 million years, is marked by several important evolutionary events. The first is related to the appearance of the earliest ray-finned fishes, the palaeonisciforms. These fishes possess essentially the same feeding mechanism design and the same pattern, including a fully heterocercal tail, as in later forms.

      The main groups of holosteans (holostean) and halecostomes (which gave rise to the teleosts) apparently arose from palaeoniscid-like ancestors during the Permian and Triassic periods. These advanced palaeoniscids are sometimes called subholosteans, a reference to the fact that they had some of the holostean features, such as upright jaw suspensions. Fishes that were referred to this unnatural group were characteristic of the Triassic Period (251–200 million years ago), although a few families continued into the Jurassic Period (200–145.5 million years ago). In general, the subholosteans can be said to show a diversity in the structure of the skeleton that was never attained by the more primitive palaeonisciforms. This diversity suggests the kind of evolutionary “experiments” that must have occurred during the rise of the various families of more-advanced actinopterygians.

      The origin of the order Acipenseriformes (which includes the sturgeon) is not known for certain, although they were clearly derived from some palaeonisciform groups. Fossils that are without doubt related to the sturgeons and paddlefishes (paddlefish) date back to the Middle Jurassic (about 176–160 million years ago); the earlier history of this order is poorly documented and confused. Both the sturgeons and the paddlefishes became specialized early in their history and have shown only minor diversification since then.

      The Polypteriformes show a confusing array of palaeonisciform, holostean, and specialized characters. Some skull and scale features indicate derivation from palaeonisciform ancestors. The palate and jaws, on the other hand, suggest attainment of a nearly holostean-like pattern. However, the specialized fins, including the diphycercal tail, indicate that the polypteriforms have had a long independent history. Fossil occurrences, which extend back to the Late Cretaceous Period (beginning some 100 million years ago), offer no clues to their affinity.

Classification

Distinguishing taxonomic features

      The approximately 37 families of the Chondrostei, which include extinct forms, are grouped into about 12 orders and separated from one another, for the most part, on the basis of differences in dermal bone pattern, body shape, and fin form and position. Only two orders exist at present. Groups of living forms are presented in the classification below.

Annotated classification

      Order Acipenseriformes

 Endoskeleton formed largely of cartilage, scale covering of the body greatly reduced, fin rays outnumber the internal supports, tail heterocercal, notochord persistent, intestine retains a spiral valve. 3 families, 2 of which contain living species. Middle Jurassic to present.

      Family Acipenseridae (sturgeons (sturgeon))

 Body covered with 5 rows of enlarged scales (a median row along back, paired rows along flank and belly), head covered with bony plates, mouth small and subterminal, pectoral fin with stout anterior spine. 4 genera and about 25 species, coastal marine and fresh waters of Eurasia and North America. Late Cretaceous to present.

      Family Polyodontidae (paddlefishes (paddlefish))

 Snout greatly elongated and paddle-shaped and supported by stellate ossicles. In Polyodon the skin is smooth except for a few scales at the base of the tail. Psephurus additionally has tiny deeply embedded scales all over the body. The mouth is subterminal but large. The gills are equipped with comblike rakers to strain food particles out of the water, less developed in Psephurus. 4 genera, 2 of which are living and each containing 1 living species, fresh waters of North America and China. Late Cretaceous to present.

      Order Polypteriformes

 

      Family Polypteridae (bichirs (bichir) and reedfish)

 Elongate fishes; scales rhomboid, interlocking with peg and socket joint, covered with enameloid, dorsal fin long and consisting of 5–18 finlets each supported by a spine, notochord replaced by bony centra, tail rounded. 2 living genera with about 10 species; fresh waters of central and west Africa. Late Cretaceous to present.

Critical appraisal

      Because they are a fairly uniform group, the classification of the Chondrostei is difficult and unsettled. About 37 families are now recognized; however, only three families are extant. The relationships of bichirs and the reedfish are especially controversial. Some authorities place them in a separate subclass; others conclude that they are related to the crossopterygians (crossopterygian).

Bobb Schaeffer 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; E.S. Goodrich, “Vertebrata craniata,” fasc. 1, “Cyclostomes and Fishes,” in E.R. Lankester (ed.), A Treatise on Zoology, 9 vol. (1909, reprinted 1964), a classic work on the anatomy of fish that is still useful; D. Heyler, Vertébrés de l'autunien de France (1969), a detailed account of some palaeonisciform fish from a classic locality in France; J.P. Lehman, “Etude complementaire des poissons de l'Eotrias de Madagascar,” K. Svenska Vetensk-Akad. Handl., 2:1–201 (1952), a detailed study of early Triassic fish from Madagascar; “Super-ordre des Chondrostéens (Chondrostei): Formes fossiles,” in P.P. Grassé (ed.), Traité de zoologie, vol. 13 (1958), a comprehensive reference work on the organization and classification of the chondrosteans, and “Actinopterygii,” in J. Piveteau (ed.), Traité de paléontologie, vol. 4 (1966); D.V. Obruchev (ed.), Fundamentals of Paleontology, vol. 11, Agnatha, Pisces (1967); and A.S. Romer, Vertebrate Paleontology, 3rd ed. (1966), two comprehensive reference works.

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