agnathan

/ag"neuh theuhn/, n.

1. any member of the vertebrate class Agnatha.

adj.

2. agnathous.

[AGNATH(A) + -AN]

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▪ fish

Introduction

  any member of the group of primitive, jawless fishes (fish) that includes the lampreys (lamprey) (order Petromyzoniformes), hagfishes (hagfish) (order Myxiniformes), and several extinct groups.

      Hagfishes are minor pests of commercial food fisheries of the North Atlantic, but lampreys, because of their parasitic habit, have been a serious pest of food fisheries in the Great Lakes in North America, where they have reduced the numbers of lake trout and other species. Agnathans are otherwise of little economic importance. The group is of great evolutionary interest, however, because it includes the oldest known craniate fossils and because the living agnathans have many primitive characteristics.

General features

      The body of the hagfish is soft-skinned, scaleless, and nearly cylindrical, with a single nostril at the anterior end, overlying the mouth, and a low caudal fin around the tail. In some species the gills open to the surface through separate pores; however, in others the gills open to a common duct, which in turn opens to the surface through a single pore. The eyes are vestigial and covered by skin. All of the approximately 70 known species are restricted to cold, marine bottom waters at depths ranging from 10 metres (about 33 feet) in high latitudes to 1,300 metres (about 4,300 feet) in equatorial oceans. Adults are 40 to 80 cm (about 15 to 30 inches) long. All species are superficially similar except in the number and position of the gill apertures.

      Lampreys (lamprey), which number about 43 species, are found in cool, fresh, and coastal waters of all continents except Africa. All species are rather similar. The body is smooth, scaleless, and eel-shaped, with well-developed dorsal and caudal fins; the mouth is surrounded by a suctorial oral disk bearing horny teeth. The eyes are well developed, and the single nostril is on the top of the head. Lampreys possess seven pairs of external gills. Adults range from 15 to 100 cm (6 to 39 inches) long.

      Although the gonad of a hagfish usually includes both ovary and testis, there is no evidence of either hermaphroditism (the reproductive organs of both sexes functioning in the same individual) or self-fertilization. A female produces a small number of tough-skinned yolk-filled eggs about 2 cm (0.8 inch) long that hatch into miniature adults.

      Hagfishes locate their food by scent. Although some are known to eat fishes (fish) immobilized in nets, the best-studied species, Myxine glutinosa, normally feeds on soft-bodied invertebrates and larger dead animals. Myxine burrows into soft marine sediments and rests with only the tip of the head protruding. During respiration, water enters through the nostril and passes by a nasopharyngeal duct to the pharynx and gills. When stimulated by the scent of a dead fish, Myxine leaves its burrow and swims against the current. On making contact with the fish, Myxine coils around it and bites into it by protruding and retracting the comblike horny tooth plates on the floor of the mouth. Along each side of Myxine's body is a row of prominent glands that produce a gelatinous slime when it is disturbed.

      Lampreys mate in a nestlike depression excavated by the male in the gravel bed of a stream; the numerous eggs, about 1 mm (0.04 inch) in diameter, lodge in the gravel around the nest. The egg hatches into an ammocoete larva—a blind, wormlike animal that burrows in silt. The larva's mouth is overhung by a hoodlike upper lip that protrudes above the surface of the silt. A continuous stream of water passes in through the mouth and out through the seven pairs of gills. Microscopic plants, the food of the ammocoete, are filtered from this respiratory current by strands of mucus produced by the endostyle, which is a gland in the floor of the pharynx.

      After about three years, when the ammocoete has grown to about 10 cm (about 4 inches), it undergoes a radical metamorphosis. The eyes complete their development; the upper lip becomes transformed into a suctorial oral disk; the endostyle changes into a thyroid gland; and the fins along the back increase in height. On completion of metamorphosis, a typical lamprey such as Petromyzon marinus migrates to the sea, where it feeds by attaching itself with its sucker to bony fishes (bony fish). It rasps into the flesh with a toothed, tonguelike structure on the floor of the mouth. Saliva containing an anticoagulant facilitates the ingestion of blood and muscle tissue. On attaining full adult size, the lamprey ceases to feed, migrates upstream to a spawning ground, mates, and dies.

      Members of several lamprey species do not migrate to sea but feed in fresh water. Petromyzon marinus dorsatus once seriously affected commercial fishing in the Great Lakes until measures were undertaken to control it. The brook lampreys do not feed after metamorphosis but mature sexually, reproduce, and die.

Form and function

      The elongated bodies of hagfishes (hagfish) and lampreys (lamprey) are adaptations to a burrowing habit, throughout life in the hagfish but only during the larval period in the lamprey. Considerable variation in body form occurs among extinct agnathans. Some Osteostraci, for example, possessed a flattened head shield and, although possessing a powerful swimming tail, appear to have been bottom dwellers. The size and shape of the mouth suggest that they were filter feeders. The laterally compressed, fishlike form of the anaspids (such as Pharyngolepis) indicates a free-swimming habit. The extinct heterostracans include obvious bottom dwellers (Drepanaspis) and others ( Pteraspis) that were apparently adapted to mid-water, or nektonic, life. Some heterostracans, for example, had movable enamel plates inside the lower lip, probably to provide a biting or grazing mechanism. Osteostracans, anaspids, hagfishes, and lampreys have one median nostril, but heterostracans appear to have had two, one at each corner of the mouth.

Evolution and classification

Evolution

      If evidence from fossil and living forms is combined, the Agnatha are distinguishable from the other craniates (Gnathostomata) by what they lack: jaws, lateral fins supported by fin rays, vertebrae, a horizontal semicircular canal in the ear, and genital ducts. The nervous, sensory, endocrine, circulatory, excretory, and muscular systems have the same basic structure as those of gnathostomes but are generally simpler. Unlike the slitlike gills of gnathostomes, the gills of agnathans are pouchlike and open through pores. The presence in the ammocoete larva of an endostyle, a gland that otherwise is found only in protochordates (protochordate) (such as the amphioxi (amphioxus) and tunicates (tunicate)), suggests that the Agnatha represent an evolutionary level intermediate between the protochordates and gnathostomes; however, the degree of specialization of known agnathans (particularly the single nostril of most forms) rules out the possibility that they are ancestral to the gnathostomes.

Annotated classification

      In the classification below, the groups indicated by a dagger (†) are extinct and known only from fossils.

Superclass Agnatha

 Craniate chordates, with pouchlike gills and lacking jaws. Ordovician Period to present.

      Order Petromyzontiformes (lampreys (lamprey))

 7 pairs of gills opening through pores, laterally placed eyes, single nostril dorsal, horny teeth on an oral sucker, horny teeth on tongue. 1 or 2 dorsal fins. 10 genera, 43 species. Pennsylvanian Subperiod to present.

      Family Petromyzontidae

 Single horny tooth plate above mouth carrying pointed or rounded teeth. 8 genera and approximately 39 species; Eurasia and North America.

      Family Mordaciidae

 Tooth plate above mouth paired, each being tricuspid. 1 genus, 3 species; eastern Australia and western South America.

      Family Geotriidae

 Single tooth plate above mouth carrying 4 teeth. 1 genus, 1 species; southern Australia, New Zealand, South America.

      Order Myxiniformes (hagfishes (hagfish))

 Nostril at tip of snout, eyes vestigial, tongue with 2 rows of horny teeth, mouth surrounded by barbels, slime glands along body, 1–16 pairs of gills, single fin extending around posterior end of body. Pennsylvanian Subperiod to present.

      Family Myxinidae

 Approximately 7 genera, about 70 species.

      Subfamily Eptatretinae

 5–15 pairs of gills opening separately to outside. 2 genera, about 43 species; Atlantic, Indian and Pacific oceans.

      Subfamily Myxininae

 5–7 pairs of gills open inside the body into collecting ducts, which in turn open to the outside through a single pore. 4–5 genera, about 26 species; all oceans.

      †Order Heterostraci

 Head covered with 2 large bony plates, 1 above and 1 below, separated by a variable number of smaller plates. Single gill opening on each side, body covered with large scales, possibly paired nostrils. Bone lacks enclosed bone cells. 6 or 7 minor taxa with questionable affinities. Early Silurian to Late Devonian (about 444–359 million years ago).

      †Order Arandaspida

 Eyes placed frontally, head covered in 2 large bony plates separated by small plates which each surround separate gill openings. Bone lacks enclosed bone cells. About 4 genera, 4 species. Middle to Late Ordovician (about 472–444 million years ago).

      †Order Astraspida

 Head covered with small mushroom-shaped plates, gill openings separate. 3 genera, 3 species. Late Ordovician to Early Silurian (461–428 million years ago).

      †Order Osteostraci

 Head covered in broad bony shield, undersurface of head covered with tiny scales, gill openings on undersurface, eyes dorsal, nostril median and placed on top of head, areas of bony shield covered with tiny tesserae. About 7 families. Early Silurian to Late Devonian (444–359 million years ago).

      †Order Galeaspida

 Head covered in broad semicircular head shield that is sometimes drawn out to a pointed snout, eyes dorsal, medium nostril very large, gills on the undersurface. 2 major clades (Polybranchiaspidida and Eugaleaspidiformes) and numerous species. Early Silurian to Late Devonian (444–359 million years ago).

      †Order Anaspida

 Small streamlined fishes, body and head covered with elongate scales, nostril between eyes, gill openings lateral and arranged in slanting line, hypocercal tail bent downward. 3 families. Early Silurian to Late Devonian (444–359 million years ago).

      †Order Thelodonti

 A little-known group of unknown affinities. Body covered in tiny scales; in some the body is flattened from top to bottom, in others from side to side. About 7 families. Early Silurian to Late Devonian (444–359 million years ago).

Critical appraisal

      Superclass Agnatha is made up of living and extinct forms. In older classifications, extant forms—that is, the lampreys and hagfishes— were grouped together as Cyclostomata (round mouth). It is now thought that the similarities between lampreys and hagfishes are primitive vertebrate features not necessarily indicative of a close relationship. Lampreys share many similarities with jawed vertebrates (Gnathostomes)—such as the ability to regulate the ion concentration in the body, the nervous regulation of heartbeat, and well-developed lateral eyes. The extinct agnathans appear to be more advanced than either hagfishes or lampreys and possess features indicative of a relatively close relationship with gnathosomes.

Ronald Strahan Peter L. Forey

Additional Reading

V.C. Applegate, Natural History of the Sea Lamprey, Petromyzon marinus, in Michigan, Spec. Scient. Rep. U.S. Fish Wildl. Serv. 55 (1950), an account of lamprey life history and the pest problem; A. Brodal and R. Fange (eds.), The Biology of Myxine (1963), an authoritative work by 24 authors; M. Fontaine, “Formes actuelles de cyclostomes,” in P.P. Grassé (ed.), Traité de zoologie, vol. 13, pp. 1–172 (1958), the best general account of cyclostomes (well illustrated); M.W. Hardisty and I.C. Potter (eds.), The Biology of Lampreys (1972), a comprehensive reference; A.J. Marshall, “Agnatha,” in T.J. Parker and W.A. Haswell, Textbook of Zoology, 7th ed., vol. 1 (1962), a good account of comparative anatomy, written for university students; E. Stensio, “Les Cyclostomes fossiles ou ostracodermes,” in J. Piveteau (ed.), Traité de paléontologie, vol. 4, pp. 96–382 (1964); R. Strahan, “The Behaviour of Myxinoids,” Acta Zool. Stockh., 44:73–102 (1963).

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