第28章無脊椎動物

The annelids and the mollusks are the most familiar of the lophotrochozoan protostomes. They are also more “typical” lophotrochozoans, since both groups include aquatic species with trochophore larvae, which unite both taxa in common ancestry. These phyla show how a flexible body plan can lead to biological success in terms of abundance and species diversity. The phylum Mollusca has the second greatest number of species of all animal phyla with nearly 100,000 described extant species, and about 80,000 described extinct species. In fact, it is estimated that about 25 percent of all known marine species are mollusks! The annelids and mollusks are both bilaterally symmetrical, cephalized, triploblastic, schizocoelous eucoelomates They include animals you are likely to see in your backyard or on your dinner plate!

The name “Mollusca” means “soft” body, since the earliest descriptions of mollusks came from observations of “squishy,” unshelled cuttlefish. Molluscs are predominantly a marine group of animals; however, they are also known to inhabit freshwater as well as terrestrial habitats. This enormous phylum includes chitons, tusk shells, snails, slugs, nudibranchs, sea butterflies, clams, mussels, oysters, squids, octopuses, and nautiluses. Mollusks display a wide range of morphologies in each class and subclass, but share a few key characteristics (Figure 28.21). The chief locomotor structure is usually a muscular foot. Most internal organs are contained in a region called the visceral mass. Overlying the visceral mass is a fold of tissue called the mantle; within the cavity formed by the mantle are respiratory structures called gills, that typically fold over the visceral mass. The mouths of most mollusks, except bivalves (e.g., clams) contain a specialized feeding organ called a radula, an abrasive tonguelike structure. Finally, the mantle secretes a calcium-carbonate-hardened shell in most mollusks, although this is greatly reduced in the class Cephalopoda, which contains the octopuses and squids.

The muscular foot is the ventral-most organ, whereas the mantle is the limiting dorsal organ that folds over the visceral mass. The foot, which is used for locomotion and anchorage, varies in shape and function, depending on the type of mollusk under study. In shelled mollusks, the foot is usually the same size as the opening of the shell. The foot is both retractable and extendable. In the class Cephalopoda (“head-foot”), the foot takes the form of a funnel for expelling water at high velocity from the mantle cavity; and the anterior margin of the foot has been modified into a circle of arms and tentacles.

The visceral mass is present above the foot, in the visceral hump. This mass contains digestive, nervous, excretory, reproductive, and respiratory systems. Molluscan species that are exclusively aquatic have gills that extend into the mantle cavity, whereas some terrestrial species have "lungs" formed from the lining of the mantle cavity. Mollusks are schizocoelous eucoelomates, but the coelomic cavity in adult animals has been largely reduced to a cavity around the heart. However, a reduced coelom sometimes surrounds the gonads, part of the kidneys, and intestine as well. This overall coelomic reduction makes the mantle cavity the major internal body chamber.

Most mollusks have a special rasp-like organ, the radula, which bears chitinous filelike teeth. The radula is present in all groups except the bivalves, and serves to shred or scrape food before it enters the digestive tract. The mantle (also known as the pallium) is the dorsal epidermis in mollusks; all mollusks except some cephalopods are specialized to secrete a calcareous shell that protects the animal's soft body.

Most mollusks are dioecious animals and fertilization occurs externally, although this is not the case in terrestrial mollusks, such as snails and slugs, or in cephalopods. In most aquatic mollusks, the zygote hatches and produces a trochophore larva, with several bands of cilia around a toplike body, and an additional apical tuft of cilia. In some species, the trochophore may be followed by additional larval stages, such as a veliger larvae, before the final metamorphosis to the adult form. Most cephalopods develop directly into small versions of their adult form.