45 Mollusks and Annelids Concept Outline 45.1 Mollusks were among the first coelomates. Coelomates. Next to the arthropods, mollusks comprise the second most diverse phylum and include snails, clams, and octopuses. There are more terrestrial mollusk species than terrestrial vertebrates! Body Plan of the Mollusks. The mollusk body plan is characterized by three distinct sections, a unique rasping tongue, and distinctive free-swimming larvae also found in annelid worms. The Classes of Mollusks. The three major classes of mollusks are the gastropods (snails and slugs), the bivalves (oysters and clams), and the cephalopods (octopuses and squids). While they seem very different at first glance, on closer inspection, they all have the same basic mollusk body plan. 45.2 Annelids were the first segmented animals. Segmented Animals. Annelids are segmented coelomate worms, most of which live in the sea. The annelid body is composed of numerous similar segments. FIGURE 45.1 Classes of Annelids. The three major classes of annelids An annelid, the Christmas-tree worm, Spirobranchus are the polychaetes (marine worms), the oligochaetes giganteus. Mollusks and annelids inhabit both terrestrial and (earthworms and related freshwater worms), and the aquatic habitats. They are large and successful groups, with some hirudines (leeches). of their most spectacular members represented in marine environments. 45.3 Lophophorates appear to be a transitional group. Lophophorates. The three phyla of lophophorates share lthough acoelomates and pseudocoelomates have proven a unique ciliated feeding structure, but differ in many other very successful, a third way of organizing the animal ways. A body has also evolved, one that occurs in the bulk of the ani- mal kingdom. We will begin our discussion of the coelomate animals with mollusks, which include such animals as clams, snails, slugs, and octopuses. Annelids (figure 45.1), such as earthworms, leeches, and seaworms, are also coelomates, but in addition, were the earliest group of animals to evolve seg- mented bodies. The lophophorates, a group of marine ani- mals united by a distinctive feeding structure called the lophophore, have features intermediate between those of pro- tostomes and deuterostomes and will also be discussed in this chapter. The remaining groups of coelomate animals will be discussed in chapters 46, 47, and 48. 899 45.1 Mollusks were among the first coelomates. Coelomates As a group, mollusks are an impor- tant source of food for humans. Oys- The evolution of the coelom was a sig- ters, clams, scallops, mussels, octopuses, nificant advance in the structure of the and squids are among the culinary deli- animal body. Coelomates have a new cacies that belong to this large phylum. body design that repositions the fluid Mollusks are also of economic signifi- Annelids Sponges and allows the development of com- Mollusks cance to us in many other ways. For ex- Flatworms Chordates Cnidarians Arthropods plex tissues and organs. This new body Nematodes ample, pearls are produced in oysters, Echinoderms plan also made it possible for animals and the material called mother-of- to evolve a wide variety of different pearl, often used in jewelry and other body architectures and to grow to decorative objects, is produced in the much larger sizes than acoelomate ani- shells of a number of different mol- mals. Among the earliest groups of lusks, but most notably in the snail coelomates were the mollusks and the called abalone. Mollusks are not wholly annelids. beneficial to humans, however. Bivalve mollusks called shipworms burrow through wood submerged in the sea, Mollusks damaging boats, docks, and pilings. Mollusks (phylum Mollusca) are an ex- The zebra mussel has recently invaded tremely diverse animal phylum, second North American ecosystems from Eu- only to the arthropods, with over rope via the ballast water of cargo ships 110,000 described species. Mollusks from Europe, wreaking havoc in many include snails, slugs, clams, scallops, aquatic ecosystems. Slugs and terres- oysters, cuttlefish, octopuses, and trial snails often cause extensive dam- many other familiar animals (figure age to garden flowers, vegetables, and 45.2). The durable shells of some mol- crops. Other mollusks serve as hosts to lusks are often beautiful and elegant; the intermediate stages of many serious they have long been favorite objects parasites, including several nematodes for professional scientists and amateurs and flatworms, which we discussed in alike to collect, preserve, and study. chapter 44. Chitons and nudibranchs are less fa- Mollusks range in size from almost miliar marine mollusks. Mollusks are microscopic to huge, although most characterized by a coelom, and while measure a few centimeters in their there is extraordinary diversity in this largest dimension. Some, however, are phylum, many of the basic compo- FIGURE 45.2 minute, while others reach formidable nents of the mollusk body plan can be A mollusk. The blue-ringed octopus is one sizes. The giant squid, which is occa- of the few mollusks dangerous to humans. seen in figure 45.3. sionally cast ashore but has rarely been Strikingly beautiful, it is equipped with a Mollusks evolved in the oceans, and sharp beak and poison glands—divers give observed in its natural environment, most groups have remained there. Ma- it a wide berth! may grow up to 21 meters long! rine mollusks are widespread and Weighing up to 250 kilograms, the often abundant. Some groups of mol- giant squid is the largest invertebrate lusks have invaded freshwater and terrestrial habitats, in- and, along with the giant clam (figure 45.4), the heaviest. cluding the snails and slugs that live in your garden. Ter- Millions of giant squid probably inhabit the deep regions of restrial mollusks are often abundant in places that are at the ocean, even though they are seldom caught. Another least seasonally moist. Some of these places, such as the large mollusk is the bivalve Tridacna maxima, the giant crevices of desert rocks, may appear very dry, but even clam, which may be as long as 1.5 meters and may weigh as these habitats have at least a temporary supply of water at much as 270 kilograms. certain times. There are so many terrestrial mollusks that only the arthropods have more species adapted to a ter- restrial way of life. The 35,000 species of terrestrial mol- Mollusks are the second-largest phylum of animals in lusks far outnumber the roughly 20,000 species of terres- terms of named species; mollusks exhibit a variety of body forms and live in many different environments. trial vertebrates. 900 Part XII Animal Diversity PHYLUM MOLLUSCA: Coelom The mantle is a heavy fold of tissue Snails have a three-chambered heart wrapped around the mollusk body like a and an open circulation system. The cape. The cavity between the mantle and coelom is confined to a small cavity the body contains gills, which capture around the heart. oxygen from the water passing through the mantle cavity. In some mollusks, like snails, the mantle secretes a hard Mollusks were among the first outer shell. animals to develop an efficient excretory system. Tubular structures Gonad called nephridia gather wastes Heart from the coelom and discharge Intestine Coelom them into the mantle cavity. Stomach Nephridium Digestive gland Shell Mantle Gill Mantle cavity Anus Mouth Radula Nerve collar Foot Retractor muscles Many mollusks are carnivores. They locate Snails creep along the ground on a prey using chemosensory structures. Within muscular foot. Squid can shoot through the mouth of a snail are horny jaws and a the water by squeezing water out of the unique rasping tongue called a radula. mantle cavity, in a kind of jet propulsion. FIGURE 45.3 Evolution of the coelom. A generalized mollusk body plan is shown above. The body cavity of a mollusk is a coelom, which is completely enclosed within the mesoderm. This allows physical contact between the mesoderm and the endoderm, permitting interactions that lead to development of highly specialized organs such as a stomach. FIGURE 45.4 Giant clam. Second only to the arthropods in number of described species, members of the phylum Mollusca occupy almost every habitat on earth. This giant clam, Tridacna maxima, has a green color that is caused by the presence of symbiotic dinoflagellates (zooxanthellae). Through photosynthesis, the dinoflagellates probably contribute most of the food supply of the clam, although it remains a filter feeder like most bivalves. Some individual giant clams may be nearly 1.5 meters long and weigh up to 270 kilograms. Chapter 45 Mollusks and Annelids 901 Body Plan of the Mollusks In aquatic mollusks, a continuous stream of water passes into and out of the mantle cavity, drawn by the cilia on the In their basic body plan (figure 45.5), mollusks have dis- gills. This water brings in oxygen and, in the case of the bi- tinct bilateral symmetry. Their digestive, excretory, and valves, also brings in food; it also carries out waste materi- reproductive organs are concentrated in a visceral mass, als. When the gametes are being produced, they are fre- and a muscular foot is their primary mechanism of loco- quently carried out in the same stream. motion. They may also have a differentiated head at the The foot of a mollusk is muscular and may be adapted anterior end of the body. Folds (often two) arise from the for locomotion, attachment, food capture (in squids and oc- dorsal body wall and enclose a cavity between themselves topuses), or various combinations of these functions. Some and the visceral mass; these folds constitute the mantle. In mollusks secrete mucus, forming a path that they glide some mollusks the mantle cavity acts as a lung; in others it along on their foot. In cephalopods—squids and octo- contains gills. Gills are specialized portions of the mantle puses—the foot is divided into arms, also called tentacles.