Characteristics of Amphibians

As tetrapods, most amphibians are characterized by four well-developed limbs. Some species of salamanders and all caecilians are functionally limbless; their limbs are vestigial. An important characteristic of extant amphibians is a moist, permeable skin that is achieved via mucus glands that keep the skin damp; thus, exchange of oxygen and carbon dioxide with the environment can take place through it (cutaneous respiration). Additional characteristics of amphibians include pedicellate teeth—teeth in which the root and crown are calcified, separated by a zone of noncalcified tissue—and a papilla amphibiorum and papilla basilaris, structures of the inner ear that are sensitive to frequencies below and above 10,00 hertz, respectively. Amphibians also have an auricular operculum, which is an extra bone in the ear that transmits sounds to the inner ear.

The fossil record provides evidence of amphibian species, now extinct, that arose over 400 million years ago as the first tetrapods. Amphibia can be divided into three clades: salamanders (Urodela), frogs (Anura), and caecilians (Apoda). The life cycle of frogs, like the majority of amphibians, consists of two distinct stages: the larval stage that will metamorphosis to an adult stage. Some species in all orders bypass a free-living larval stage. All extant adult amphibians are carnivorous, and some terrestrial amphibians have a sticky tongue that is used to capture prey.

Evolution of Amphibians

The fossil record provides evidence of the first tetrapods: now-extinct amphibian species dating to nearly 400 million years ago. Evolution of tetrapods from fishes represented a significant change in body plan from one suited to organisms that respired and swam in water, to organisms that breathed air and moved onto land; these changes occurred over a span of 50 million years during the Devonian period.

Figure 1. The recent fossil discovery of Tiktaalik roseae suggests evidence for an animal intermediate to finned fish and legged tetrapods. (credit: Zina Deretsky, National Science Foundation)

One of the earliest known tetrapods is from the genus Acanthostega. Acanthostega was aquatic; fossils show that it had gills similar to fishes. However, it also had four limbs, with the skeletal structure of limbs found in present-day tetrapods, including amphibians. Therefore, it is thought that Acanthostega lived in shallow waters and was an intermediate form between lobe-finned fishes and early, fully terrestrial tetrapods. What preceded Acanthostega?

In 2006, researchers published news of their discovery of a fossil of a “tetrapod-like fish,” Tiktaalik roseae, which seems to be an intermediate form between fishes having fins and tetrapods having limbs (Figure 1). Tiktaalik likely lived in a shallow water environment about 375 million years ago.

The early tetrapods that moved onto land had access to new nutrient sources and relatively few predators. This led to the widespread distribution of tetrapods during the early Carboniferous period.

The Paleozoic Era and the Evolution of Vertebrates

The climate and geography of Earth was vastly different during the Paleozoic Era, when vertebrates arose, as compared to today. The Paleozoic spanned from approximately 542 to 251 million years ago. The landmasses on Earth were very different from those of today. Laurentia and Gondwana were continents located near the equator that subsumed much of the current day landmasses in a different configuration (Figure 2). At this time, sea levels were very high, probably at a level that hasn’t been reached since. As the Paleozoic progressed, glaciations created a cool global climate, but conditions warmed near the end of the first half of the Paleozoic. During the latter half of the Paleozoic, the landmasses began moving together, with the initial formation of a large northern block called Laurasia. This contained parts of what is now North America, along with Greenland, parts of Europe, and Siberia. Eventually, a single supercontinent, called Pangaea, was formed, starting in the latter third of the Paleozoic. Glaciations then began to affect Pangaea’s climate, affecting the distribution of vertebrate life.

Figure 2. During the Paleozoic Era, around 550 million years ago, the continent Gondwana formed. Both Gondwana and the continent Laurentia were located near the equator.

During the early Paleozoic, the amount of carbon dioxide in the atmosphere was much greater than it is today. This may have begun to change later, as land plants became more common. As the roots of land plants began to infiltrate rock and soil began to form, carbon dioxide was drawn out of the atmosphere and became trapped in the rock. This reduced the levels of carbon dioxide and increased the levels of oxygen in the atmosphere, so that by the end of the Paleozoic, atmospheric conditions were similar to those of today.

As plants became more common through the latter half of the Paleozoic, microclimates began to emerge and ecosystems began to change. As plants and ecosystems continued to grow and become more complex, vertebrates moved from the water to land. The presence of shoreline vegetation may have contributed to the movement of vertebrates onto land. One hypothesis suggests that the fins of aquatic vertebrates were used to maneuver through this vegetation, providing a precursor to the movement of fins on land and the development of limbs. The late Paleozoic was a time of diversification of vertebrates, as amniotes emerged and became two different lines that gave rise, on one hand, to mammals, and, on the other hand, to reptiles and birds. Many marine vertebrates became extinct near the end of the Devonian period, which ended about 360 million years ago, and both marine and terrestrial vertebrates were decimated by a mass extinction in the early Permian period about 250 million years ago.