Fishes

Agnathans: Jawless Fishes

The superclass Agnatha describes fish that lack jaws and includes the extant species of hagfish and lampreys.

Learning Objectives

Differentiate between the taxa of jawless fishes

Key Takeaways

Key Points

  • Most agnathans are now extinct, but two branches exist today: hagfishes (not true vertebrates) and lampreys (true vertebrates).
  • The earliest jawless fishes were the ostracoderms, which had bony scales as body armor.
  • Hagfish are eel-like marine scavengers in the clade Myxini that produce slime and can tie themselves into knots.
  • Lampreys are in the clade Petromyzontidae and appear morphologically similar to hagfish, but contain cartilaginous vertebral elements as an adult; thus, they are considered true vertebrates.

Key Terms

  • hagfish: any of several primitive eellike creatures, of the family Myxinidae, having a sucking mouth with rasping teeth; considered edible in Japan, their skin is used to make a form of leather
  • lamprey: any long slender primitive eel-like freshwater and saltwater fish of the Petromyzontidae family, having a sucking mouth with rasping teeth, but no jaw
  • agnathan: a member of the superclass Agnatha of jawless vertebrates

Agnathans: Jawless Fishes

Jawless fishes or agnathans are craniates that represent an ancient vertebrate lineage that arose over one half-billion years ago. “Gnathos” is Greek for “jaw” and the prefix “a” means “without,” so agnathans are “without jaws. ” Most agnathans are now extinct, but two branches still exist today: hagfishes and lampreys. Hagfishes and lampreys are recognized as separate clades, primarily because lampreys are true vertebrates, whereas hagfishes are not. A defining feature of agnathans is the lack of paired lateral appendages or fins.

Some of the earliest jawless fishes were the ostracoderms (Greek for “bone-skin”). Ostracoderms were vertebrate fishes encased in bony armor, unlike present-day jawless fishes, which lack bone in their scales.

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Ostracoderm: Ostracoderms were some of the earliest jawless fishes and were covered in bony armor. Present-day jawless fishes lack bone in their scales.

Myxini: Hagfishes

The clade Myxini includes at least 20 species of hagfishes. Hagfishes are eel-like scavengers that live on the ocean floor and feed on dead invertebrates, other fishes, and marine mammals. Hagfishes are entirely marine and are found in oceans around the world, except for the polar regions. Hagfish have slime glands beneath the skin that constantly release mucus, allowing them to escape from the grip of predators. Hagfish can also twist their bodies into a knot to gain a mechanical advantage while feeding and are notorious for eating carcasses from the inside out.

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Hagfishes: Pacific hagfish are scavengers that live on the ocean floor. These agnathans are classified as Myxini and do not have a vertebral column.

The skeleton of a hagfish is composed of cartilage, which includes a cartilaginous notochord that runs the length of the body. This notochord provides support to the hagfish’s body. Unlike true vertebrates, hagfishes do not replace the notochord with a vertebral column during development. Since they have a cartilaginous skull, they are classified in the clade Craniata.

Petromyzontidae: Lampreys

The clade Petromyzontidae includes approximately 35–40 or more species of lampreys. Lampreys are morphologically similar to hagfishes and also lack paired appendages. However, lampreys develop some vertebral elements as an adult. Their notochord is surrounded by a cartilaginous structure called an arcualia, which may resemble an evolutionarily-early form of the vertebral column.

As adults, lampreys are characterized by a toothed, funnel-like sucking mouth. Many species have a parasitic stage of their life cycle during which they are ectoparasites of fishes. Lampreys live primarily in coastal and fresh waters. They are distributed worldwide, except for the tropics and polar regions. Some species are marine, but all species spawn in fresh water; eggs are fertilized externally. The larvae differ distinctly from the adult form, spending 3 to 15 years as suspension feeders. Once they reach sexual maturity, the adults die within days of reproduction.

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Parasitic lampreys: These parasitic sea lampreys attach to their lake trout host by suction and use their rough tongues to rasp away flesh in order to feed on the trout’s blood.

Gnathostomes: Jawed Fishes

Gnathostomes, jawed vertebrates, can be divided into two types of fish: Chondrichthyes (cartilaginous fish) or Osteichthyes (bony fish).

Learning Objectives

Differentiate among the types of jawed fishes

Key Takeaways

Key Points

  • Early jawed fish (gnathostomes) were able to exploit new nutrient sources because of their jaws and paired fins.
  • Chondrichthyes includes all jawed fish with cartilagenous skeletons, such as sharks, rays, skates, and chimaeras.
  • Osteichthyes includes all jawed fish with ossified (bony) skeletons; this includes the majority of modern fish.
  • Osteichthyes can be further separated into Actinopterygii (the ray-finned fishes) and Sarcopterygii (lobe-finned fishes).
  • The majority of modern fish species are actinopterygii, from trout to clownfish.
  • Early Sarcopterygii (lobe-finned fishes) evolved into modern tetrapods, including reptiles, amphibians, birds, and mammals.

Key Terms

  • ossified: composed of bone, which is a calcium phosphate matrix created by special cells called osteoblasts
  • operculum: a covering flap or lidlike structure in plants and animals, such as a gill cover
  • Chondrichthyes: a taxonomic class within the subphylum Vertebrata: the cartilaginous fish
  • Osteichthyes: a taxonomic class within the subphylum vertebrata: the bony fish

Gnathostomes: Jawed Fishes

Gnathostomes or “jaw-mouths” are vertebrates that possess jaws. One of the most significant developments in early vertebrate evolution was the development of the jaw, which is a hinged structure attached to the cranium that allows an animal to grasp and tear its food. The evolution of jaws allowed early gnathostomes to exploit food resources that were unavailable to the jawless animals. In early evolutionary history, there were gnathostomes (jawed fishes) and agnathans (jawless fishes). Gnathostomes later evolved into all tetrapods (animals with four limbs) including amphibians, birds, and mammals.

Early gnathostomes were jawed fishes that possessed two sets of paired fins, which increased their ability to maneuver accurately. These paired fins were pectoral fins, located on the anterior body, and pelvic fins, on the posterior. The evolution of the jaw combined with paired fins permitted gnathostomes to expand from the sedentary suspension feeding of jawless fishes and become mobile predators. The gnathostomes’ ability to exploit new nutrient sources led to their evolutionary success during the Devonian period. Two early groups of gnathostomes were the acanthodians and placoderms, which arose in the late Silurian period and are now extinct. Most modern gnathostomes belong to the clades Chondrichthyes and Osteichthyes.

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Placoderms: Dunkleosteous was an enormous placoderm from the Devonian period, 380–360 million years ago. It measured up to 10 meters in length and weighed up to 3.6 tons. As gnathostomes, they were more mobile and could exploit more food resources than the agnathostomes.

Chondrichthyes: Cartilaginous Fishes

The clade Chondrichthyes consists of sharks, rays, and skates, together with sawfishes and a few dozen species of fishes called chimaeras, or “ghost,” sharks. Chondrichthyes are jawed fishes that possess paired fins and a skeleton made of cartilage. This clade arose approximately 370 million years ago in the early or middle Devonian.

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Hammerhead shark: Hammerhead sharks tend to school during the day and hunt prey at night. As members of Chondrichthyes, their skeletons are composed of cartilage.

Most cartilaginous fishes live in marine habitats, although a few species live in fresh water for part or all of their lives. Most sharks are carnivores that feed on live prey, either swallowing it whole or using their jaws and teeth to tear it into smaller pieces. Shark teeth probably evolved from the jagged scales that cover their skin called placoid scales. Some species of sharks and rays are suspension feeders that feed on plankton.

Sharks have well-developed sense organs that aid them in locating prey, including a keen sense of smell and electroreception. Organs called ampullae of Lorenzini enable sharks to detect the electromagnetic fields that are produced by all living things, including their prey. Only aquatic or amphibious animals possess electroreception. Sharks, together with most fishes and aquatic and larval amphibians, also have a sense organ called the lateral line, which is used to detect movement and vibration in the surrounding water. It is often considered homologous to “hearing” in terrestrial vertebrates. The lateral line is visible as a darker stripe that runs along the length of a fish’s body.

Rays and skates comprise more than 500 species and are closely related to sharks. They can be distinguished from sharks by their flattened bodies, pectoral fins that are enlarged and fused to the head, and gill slits on their ventral surface. Like sharks, rays and skates have a cartilaginous skeleton. Most species are marine and live on the sea floor, with nearly a worldwide distribution.

Osteichthyes: Bony Fishes

Members of the clade Osteichthyes, also called bony fish, are characterized by a bony skeleton. The vast majority of present-day fish belong to this group, which consists of approximately 30,000 species, making it the largest class of vertebrates in existence today.

Nearly all bony fish have an ossified skeleton with specialized bone cells (osteocytes) that produce and maintain a calcium phosphate matrix. A few groups of Osteichthyes, such as sturgeons and paddlefish, have primarily cartilaginous skeletons, but retain some bony elements. The skin of bony fish is often covered by overlapping scales. Skin glands secrete mucus that reduces drag when swimming and aids the fish in osmoregulation. Like sharks, bony fish have a lateral line system that detects vibrations in water. All bony fish use gills for gas exchange. Water is drawn over gills that are located in chambers covered and ventilated by a protective, muscular flap called the operculum. Many bony fish also have a swim bladder, a gas-filled organ that helps to control the buoyancy of the fish.

Bony fish are further divided into two extant clades: Actinopterygii (ray-finned fish) and Sarcopterygii (lobe-finned fish). Actinopterygii, the ray-finned fish include many familiar fish, such as tuna, bass, trout, and salmon, among others. Ray-finned fish are named for their fins that are webs of skin supported by bony spines called rays. In contrast, the fins of Sarcopterygii are fleshy and lobed, supported by bone. Although most members of this clade are extinct, living members include the less-familiar lungfishes and coelacanths. Early Sarcopterygii evolved into modern tetrapods, including reptiles, amphibians, birds, and mammals.

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Actinopterygii and Sarcopterygii: The (a) sockeye salmon (Actinopterygii) and (b) coelacanth (Sarcopterygii) are both bony fishes of the Osteichthyes clade. The coelacanth, sometimes called a lobe-finned fish, was thought to have gone extinct in the Late Cretaceous period, 100 million years ago, until one was discovered in 1938 near the Comoros Islands between Africa and Madagascar.