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hic09617_ch16.qxd 5/30/00 12:43 PM Page 325 CHAPTER 16 Molluscs Phylum Mollusca Fluted giant clam, Tridacna squamosa. A Significant Space space within the mesoderm, the coelom. This meant that the Long ago in the Precambrian era, the most complex animals space was lined with mesoderm and the organs were sus- populating the seas were acoelomate. They must have been pended by mesodermal membranes, the mesenteries. Not inefficient burrowers, and they were unable to exploit the only could the coelom serve as an efficient hydrostatic rich subsurface ooze. Any that developed fluid-filled spaces skeleton, with circular and longitudinal body-wall muscles within the body would have had a substantial selective acting as antagonists, but a more stable arrangement of advantage because these spaces could serve as a hydro- organs with less crowding resulted. Mesenteries provided an static skeleton and improve burrowing efficiency. ideal location for networks of blood vessels, and the ali- The simplest, and probably the first, mode of achieving mentary canal could become more muscular, more highly a fluid-filled space within the body was retention of the specialized, and more diversified without interfering with embryonic blastocoel, as in pseudocoelomates. This was not other organs. the best evolutionary solution because, for example, the Development of a coelom was a major step in the evo- organs lay loose in the body cavity. lution of larger and more complex forms. All the major Some descendants of Precambrian acoelomate organ- groups in chapters to follow are coelomates. I isms evolved a more elegant arrangement: a fluid-filled 325 hic09617_ch16.qxd 5/30/00 12:45 PM Page 326 326 PART 3 The Diversity of Animal Life Position in Animal common ancestor with annelids 3. The efficiency of the respiratory Kingdom before the advent of metamerism. and circulatory systems in the 5. All organ systems are present and cephalopods has made greater body 1. Molluscs are one of the major groups well developed. size possible. Invertebrates reach their of true coelomate animals. largest size in some cephalopods. 2. They belong to the protostome Biological Contributions 4. They have a fleshy mantle that in branch, or schizocoelous coelomates, most cases secretes a shell and is and have spiral cleavage and 1. In molluscs gaseous exchange occurs variously modified for a number of determinate (mosaic) development. not only through the body surface as functions. 3. Many molluscs have a trochophore in phyla discussed previously, but 5. Features unique to the phylum are the larva similar to trochophore larvae of also in specialized respiratory radula and the muscular foot. marine annelids and other marine organs in the form of gills or lungs. 6. The highly developed direct eye of protostomes. Developmental evidence 2. Most classes have an open cephalopods is similar to the indirect thus indicates that molluscs and circulatory system with pumping eye of vertebrates but arises as a skin annelids share a common ancestor. heart, vessels, and blood sinuses. In derivative in contrast to the brain eye 4. Because molluscs are not metameric, most cephalopods the circulatory of vertebrates. they must have diverged from their system is closed. The Molluscs ,Mollusca (mol-luska) (L. molluscus soft) is one of the largest animal phyla after Arthropoda. There are nearly 50,000 living species and some 35,000 fossil species. The name Mollusca indicates one of their distinctive char- acteristics, a soft body. This very diverse group (Figure 16-1) includes chitons, tooth shells, snails, slugs, A D nudibranchs, sea butterflies, clams, mussels, oysters, squids, octopuses, and nautiluses. The group ranges from fairly simple organisms to some of the most complex of invertebrates, and in size from almost microscopic to the giant squid Architeuthis. These huge molluscs may grow to 18 m long, including their tentacles. They may weigh 450 kg (1,000 pounds). The shells of some giant clams, Tridacna gigas, which inhabit Indo-Pacific coral B E reefs, reach 1.5 m in length and weigh more than 225 kg. These are extremes, Figure 16-1 however, for probably 80% of all mol- Molluscs: a diversity of life forms. The basic luscs are less than 5 cm in maximum body plan of this ancient group has become shell size. The phylum includes some variously adapted for different habitats. A, A of the most sluggish and some of the chiton (Tonicella lineata), Class Polyplacophora. B, A marine snail (Calliostoma annulata), Class swiftest and most active invertebrates. Gastropoda. C, A nudibranch (Chromodoris sp.) It includes herbivorous grazers, preda- Class Gastropoda. D, Pacific giant clam ceous carnivores, filter feeders, detritus (Panope abrupta), with siphons to the left, Class feeders, and parasites. Bivalvia. E, An octopus (Octopus briareus), Molluscs are found in a great range Class Cephalopoda, forages at night on a Caribbean coral reef. of habitats, from the tropics to polar C hic09617_ch16.qxd 5/30/00 12:45 PM Page 327 CHAPTER 16 Molluscs 327 seas, at altitudes exceeding 7000 m, in ponds, lakes, and streams, on mud Characteristics of Phylum (usually three chambered), blood flats, in pounding surf, and in open Mollusca vessels, and sinuses; respiratory pig- ocean from the surface to abyssal ments in blood 1. Body bilaterally symmetrical (bilater- depths. Most of them live in the sea, 8. Gaseous exchange by gills, lungs, al asymmetry in some); unsegment- and they represent a variety of life- mantle, or body surface ed; often with definite head styles, including bottom feeders, bur- 9. One or two kidneys (metaneph- 2. Ventral body wall specialized as a rowers, borers, and pelagic forms. ridia) opening into the pericardial muscular foot, variously modified cavity and usually emptying into the According to fossil evidence, mol- but used chiefly for locomotion luscs originated in the sea, and most of mantle cavity 3. Dorsal body wall forms pair of folds 10. Nervous system of paired cerebral, them have remained there. Much of called the mantle, which encloses their evolution occurred along the pleural, pedal, and visceral ganglia, the mantle cavity, is modified into with nerve cords and subepidermal shores, where food was abundant and gills or lungs, and secretes the plexus; ganglia centralized in nerve habitats were varied. Only bivalves shell (shell absent in some) ring in gastropods and cephalopods and gastropods moved into brackish 4. Surface epithelium usually ciliated 11. Sensory organs of touch, smell, taste, and freshwater habitats. As filter feed- and bearing mucous glands and sen- equilibrium, and vision (in some); ers, bivalves were unable to leave sory nerve endings eyes highly developed in aquatic surroundings. Only snails (gas- 5. Coelom limited mainly to area cephalopods tropods) actually invaded the land. around heart, and perhaps lumen of 12. Internal and external ciliary tracts Terrestrial snails are limited in their gonads and part of kidneys often of great functional importance range by their need for humidity, shel- 6. Complex digestive system; rasping 13. Both monoecious and dioecious ter, and presence of calcium in the soil. organ (radula) usually present; anus forms; spiral cleaveage; larva primi- Many kinds of molluscs are used usually emptying into mantle cavity tively a trochophore, many with a as food. Pearl buttons are obtained 7. Open circulatory system (mostly veliger larva, some with direct from shells of bivalves. The Mississippi closed in cephalopods) of heart development and Missouri river basins have fur- nished material for most of this indus- try in the United States; however, supplies are becoming so depleted that attempts are being made to propagate bers of class Polyplacophora (chitons) Form and Function bivalves artificially. Pearls, both natural are common to abundant marine ani- and cultured, are produced in the mals, especially in the intertidal zone. The enormous variety, great beauty, shells of clams and oysters, most of Bivalves (class Bivalvia) have evolved and easy availability of shells of mol- them in a marine oyster, Meleagrina, many species, both marine and fresh- luscs have made shell collecting a pop- found around eastern Asia. water. Largest and most intelligent of ular pastime. However, many amateur Some molluscs are destructive. Bur- all invertebrates are in the class shell collectors, even though able to rowing shipworms, which are bivalves Cephalopoda (squids, octopuses, and name hundreds of the shells that grace of several species (see Figure 16-27), others). Most abundant and wide- our beaches, know very little about the do great damage to wooden ships and spread of molluscs, however, are snails living animals that created those shells wharves. To prevent the ravages of and their relatives (class Gastropoda). and once lived in them. Reduced to its shipworms, wharves must be either Although enormously diverse, mol- simplest dimensions, the mollusc body creosoted or built of concrete (unfortu- luscs have in common a basic body plan may be said to consist of a head- nately, some ignore the creosote, and plan, which is described later in the foot portion and a visceral mass por- some bivalves bore into concrete). chapter. It seems peculiar, though, that tion (Figure 16-2). The head-foot is the Snails and slugs frequently damage molluscs have failed to exploit the more active area, containing the feed- garden and other vegetation. In addi- coelom. The coelom in molluscs is lim- ing, cephalic sensory, and locomotor tion, snails often serve as intermediate ited to a space around the heart, and organs. It depends primarily on mus- hosts for serious parasites. The boring perhaps around the gonads and part of cular action for its function. The vis- snail Urosalpinx rivals sea stars in the kidneys. Although it develops ceral mass is the portion containing destroying oysters. embryonically in a manner similar to digestive, circulatory, respiratory, and In this chapter we explore the the coelom of annelids, the functional reproductive organs, and it depends various major groups of molluscs, consequences of the space are quite primarily on ciliary tracts for its func- including those that apparently met different.