Amphibians usually require moist environments

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Most modern amphibians are confined to moist environments because they lose water rapidly through the skin when exposed to dry air. In addition, their eggs are enclosed within delicate gelatinous envelopes that cannot prevent water loss in dry conditions. In some amphibian species, adults live mostly on land but return to fresh water to lay and fertilize their eggs (Figure 32.17). The fertilized eggs give rise to larvae that live in water until they undergo metamorphosis to become terrestrial adults. However, many amphibians (especially those in tropical and subtropical areas) have evolved a wide variety of additional reproductive modes and types of parental care. Internal fertilization, for example, evolved several times among amphibian species. Many species develop directly into adultlike forms from fertilized eggs laid on land or carried by the parents. Other species of amphibians are entirely aquatic, never leaving the water at any stage of their lives, and many of these species retain a larval-like morphology.

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Figure 32.17 In and Out of the Water Many amphibian species have life cycles like the one diagrammed here, in which the early stages take place in water and the aquatic tadpole transforms into a terrestrial adult through metamorphosis. Some species of amphibians, however, have direct development (with no aquatic larval stage), and others are aquatic throughout life.

Question

Q: What is a potential advantage of maintaining an aquatic tadpole stage, even though the adults are terrestrial?

Most adult frogs are insectivorous, but insects can be very limited during early spring or the beginning of wet seasons. By reproducing in the water, frogs can take advantage of an abundant seasonal resource (algal blooms in temporary water) as a food source for their offspring. This allows the developing tadpoles to specialize on an abundant resource until they are large enough to feed on insects.

Animation 32.2 Life Cycle of a Frog

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The more than 7,000 known species of amphibians living today belong to three major groups: the wormlike, limbless, tropical, burrowing, or aquatic caecilians (Figure 32.18A), the tailless frogs and toads (collectively called anurans) (Figure 32.18B), and the tailed salamanders (Figure 32.18C and D).

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Figure 32.18 Diversity among the Amphibians (A) Burrowing caecilians superficially look more like worms than like amphibians. They have internal fertilization and many species are viviparous. (B) Male golden toads in the cloud forest of Monteverde, Costa Rica. This is one of many amphibian species lost to extinction in the past few decades. (C) An adult spring salamander. This species has aquatic larvae but terrestrial adults. (D) This Austin blind salamander’s life cycle is completely aquatic; it has no terrestrial stage. The eyes of this cave dweller are greatly reduced.

Anurans are most diverse in wet tropical and warm temperate regions, although a few are found at very high latitudes. There are far more anurans than any other amphibians, with well over 6,000 described species and more being discovered every year. Some anurans have tough skins and other adaptations that enable them to live for long periods in deserts, whereas others live in moist terrestrial and arboreal environments. Some species are completely aquatic as adults. All anurans have a short vertebral column and a pelvic region that is modified for leaping, hopping, or propelling the body through water by kicking the hind legs.

The more than 600 described species of salamanders are most diverse in temperate regions of the Northern Hemisphere and in cool, moist environments in the mountains of Central America, although a few species penetrate into tropical regions. Many salamanders live in rotting logs or moist soil. One major group has lost the lungs, and these species exchange gases entirely through the skin and mouth lining—body parts that all amphibians use, in addition to their lungs, for gas exchange. A completely aquatic lifestyle has evolved several times among the salamanders (see Figure 32.18D). These aquatic species have arisen through a developmental process known as neoteny, or the retention of juvenile traits (in this case, gills) by delayed somatic development. Most species of salamanders have internal fertilization, which is usually achieved through the transfer of a small, jellylike, sperm-embedded capsule called a spermatophore.

Many amphibians have complex social behaviors. Most male anurans utter loud, species-specific calls to attract females of their own species (and sometimes to defend breeding territories), and they compete for access to females that arrive at the breeding sites. Many amphibians lay large numbers of eggs, which they abandon once they are deposited and fertilized. Other amphibians lay only a few eggs, which are fertilized and then cared for. A few species of frogs, salamanders, and caecilians are viviparous, meaning that they give birth to well-developed young that have received nutrition from the female during gestation.

Media Clip 32.4 Answering a Mating Call

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Amphibians are the focus of much attention today because populations of many species are declining rapidly, especially in mountainous regions of western North America, Central and South America, and northeastern Australia. Worldwide, about one-third of amphibian species are now threatened with extinction or have disappeared completely in the last few decades. Scientists are investigating several hypotheses to account for these *amphibian declines.

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*connect the concepts One of the major factors associated with amphibian declines is the global spread of a pathogenic chytrid fungus, as explored in Key Concept 29.2 and Figure 29.6.