Definition Molluscs are astonishingly diverse. This group includes worm-like animals and giant squids, as well as animals with a single or multiple shells, or no shell at all. They are one of the largest animal phyla after Anthropods. Molluscs are coelomate lophotrochozoan protostomes, and as such they develop via spiral mosaic cleavage and make a coelom by schizocoely. The name Mollusca indicates one of their distinctive characteristics: a soft body. This diverse group ranges from fairly simple organisms to some of the most complex of invertebrates; sizes range from microscopic to the giant squid Architeuthis. Origin According to fossil evidence, molluscs originated in the sea, and most of them have remained there. The evolution occurred along the shores: Bivalves and gastropods moved into brackish and freshwater habitats Slugs and snails invaded the land Importance Food source for people around the world In 2006, 450,687,000 tons of molluscs were harvested commercially on the eastern seaboard of US, with the western seaboard and the Gulf of Mexico adding 3,687,000 and 44,537,000 tons, respectively. Forms and Functions Characteristics of Phylum Mollusca 1. Dorsal body wall forms pair of folds (mantle), which encloses the mantle cavity, and secretes the shell; ventral body wall specialized as a muscular foot; radula in mouth 2. Live in marine, freshwater, and terrestrial habitats Characteristics of Phylum Mollusca 3. Free-living or occasionally parasitic 4. Body bilaterally symmetrical (bilateral asymmetry in some); unsegmented; often with definite head 5. Triploblastic body 6. Coelom limited mainly to area around heart, and perhaps lumen of gonads, part of kidneys, and occasionally part of the intestine Characteristics of Phylum Mollusca 7. Surface epithelium usually ciliated and bearing mucous glands and sensory nerve endings 8. Complex digestive system; rasping organ (radula) usually present; anus usually emptying into mantle cavity; internal and external ciliary tracts often of great functional importance 9. Circular, diagonal, and longitudinal muscles in the body wall; mantle and foot highly muscular in some classes (for example, cephalopods and gastropods) Characteristics of Phylum Mollusca 10. Nervous system of paired cerebral, pleural, pedal, and visceral ganglia, with nerve cords and subepidermal plexus; ganglia centralized in nerve ring in gastropods and cephalopods 11. Sensory organs of touch, smell, taste, equilibrium, and vision (in some); the highly developed direct eye of cephalopods is similar to the indirect eye of vertebrates but arises as a skin derivative in contrast to the brain eye of vertebrates Characteristics of Phylum Mollusca 12. No asexual reproduction 13. Both monoecious and dioecious forms; spiral cleavage; ancestral larva a trochophore, many with a veliger larva, some with direct development 14. One or two kidneys (metanephridia) opening into the pericardial cavity and usually emptying into the mantle cavity Characteristics of Phylum Mollusca 15. Gaseous exchange by gills, lungs, mantle, or body surface 16. Open circulatory system (secondarily closed in cephalopods) of heart (usually three chambered), blood vessels, and sinuses; respiratory pigments in blood Head-Foot Head-Foot Most mollusks have well-developed heads, which bear their mouth and some specialized structures Within the mouth is a structure called radula, and usually posterior to the mouth is the chief locomotor organ, or foot Head-Foot Radula The radula is a rasping, protrusible, tonguelike organ found in all mollusks except bivalves and most solenogasters. Ribbonlike membrane bearing a surface of tiny, backward-pointing teeth used for feeding. Foot Head-Foot Foot Adapted for locomotion, for attachment to a substratum, or for a combination of functions. Usually ventral, sole-like structure in which waves of muscular contraction effect a creeping locomotion. Visceral Mass Mantle & Mantle Cavity The mantle is a sheath of skin, extending from the visceral mass that hangs down on each side of the body Mantle cavity houses respiratory organs, which develop from the mantle, and the mantle’s own exposed surface serves also for gaseous exchange. Visceral Mass Shell The shell of a mollusk is secreted by the mantle and is lined by it. Three layers: periostracum, primastic layer, nacreous layer Periostracum: the outer organic layer, composed of an organic substance called conchiolin, which is consists of quinine-tanned protein. The middle prismatic layer is composed of densely packed prisms of calcium carbonate laid down in a protein matrix. The inner nacreous layer of the shell lies next to the mantle and is secreted continuously by the mantle surface Visceral Mass Internal Structure and Function There is an open circulatory system with a pumping heart, blood vessels, and blood sinuses. In a closed circulatory system, blood moves to and from tissues within blood vessels. The digestive tract is complex and highly specialized, according to feeding habits of the various mollusks. The nervous system contains neurosecretory cells that, atleast in certain air-breathing snails, produce a growth hormone and function in osmoregulation. Reproduction and Life History The free- swimming trochophore larva that emerges from the egg in many mollusks is remarkably similar to that seen in annelids. Reproduction and Life History In many molluscan groups, the trochophore stage gives rise to a uniquely molluscan larval stage called a veliger. Classes of Molluscs Classes of Molluscs For more than 50 years taxonomists recognized five classes of living molluscs: Amphineura, Gastropoda, Scaphopoda, Bivalvia (also called Pelecypoda), and Cephalopoda. Discovery of Neopilina in the 1950s added another class (Monoplacophora), and Hyman contended that solenogasters and chitons were separate classes (Aplacophora and Polyplacophora), lapsing the name Amphineura. Class Caudofoveata Members of class Caudofoveata comprise about 120 species of wormlike, marine organisms ranging from 2 to 140 mm in length. They are mostly burrowers and orient themselves vertically, with the terminal mantle cavity and gills at the entrance of the burrow. This class is sometimes called Chaetodermomorpha. Class Solenogasters Solenogasters are a small group of about 250 species of marine animals similar to caudofoveates. Solenogasters, however, usually have no radula and no gills (although secondary respiratory structures may be present). Their foot is represented by a midventral, narrow furrow, the pedal groove. This class is sometimes called Neomeniomorpha. Class Polyplacophora: Chitons Chitons represent a somewhat more diverse molluscan group with about 1000 currently described species. They are rather flattened dorsoventrally and have a convex dorsal surface that bears seven or eight articulating limy plates, or valves; hence their name Polyplacophora (“many plate bearers”). They prefer rocky surfaces in intertidal regions, although some live at great depths. The mantle cavity of polyplacophorans is extended along the side of the foot, and the gills are more numerous. Class Polyplacophora: Chitons Class Monoplacophora Monoplacophora were long thought to be extinct; they were known only from Paleozoic shells. These molluscs are small and have a low, rounded shell and a creeping foot. The mouth bears a characteristic radula. They superficially resemble limpets, but unlike most other molluscs, have some serially repeated organs. These animals have three to six pairs of gills, two pairs of heart atria, three to seven pairs of metanephridia, one or two pairs of gonads, and a ladderlike nervous system with 10 pairs of pedal nerves. Class Monoplacophora Class Monoplacophora Class Gastropoda Among molluscs, class Gastropoda is by far the largest and most diverse, containing over 70,000 living and more than 15,000 fossil species. It contains snails, limpets, slugs, whelks, conchs, periwinkles, sea slugs, sea hares, and sea butterflies. Class Gastropoda Gastropods are usually sluggish, sedentary animals because most of them have heavy shells and slow locomotion. Some are specialized for climbing, swimming, or burrowing. Shells are their chief defense. The range of gastropod habitats is large. Marine gastropods are common both in littoral zones and at great depths, and some are even pelagic. Some are adapted to brackish water and others to freshwater. Class Gastropoda Forms and Functions Torsion is a gastropod synapomorphy which occurs in all gastropods during larval development. Torsion is the rotation of the visceral mass, mantle, and shell 180˚ with respect to the head and foot of the gastropod. This rotation brings the mantle cavity and the anus to an anterior position above the head. Class Gastropoda: Torsion Class Gastropoda Forms and Functions Coiling, or spiral winding, of the shell and visceral mass is not the same as torsion. Coiling may occur in the larval stage at the same time as torsion, but the fossil record shows that coiling was a separate evolutionary event and originated in gastropods earlier than did torsion. Nevertheless, all living gastropods have descended from coiled, torted ancestors, although some have lost these characteristics. Class Gastropoda: Coiling Class Gastropoda Forms and Functions Feeding habits of gastropods are as varied as their shapes and habitats, but all include use of some adaptation of the radula. Most gastropods are herbivorous, rasping particles of algae from hard surfaces. Some herbivores are grazers, some are browsers, and some are planktonic feeders. Some are also scavengers that feeds on decaying animals and some are also carnivores. Class Gastropoda Forms and Functions Internal Forms and Functions: Respiration in most gastropods is performed by a ctenidium (two gills is the primitive condition, found in some prosobranchs) located in the mantle cavity, though some aquatic forms lack gills and instead depend on the mantle and skin. Class Gastropoda Forms and Functions Internal Forms and Functions: Most gastropods have a single nephridium (kidney). The circulatory and nervous systems are well developed Many gastropods perform courtship ceremonies. During copulation in monoecious species there is an exchange of spermatozoa or spermatophores (bundles of sperms). Class Gastropoda Forms and Functions Internal Forms and Functions: The latter incorporates three pairs of ganglia connected by nerves. Sense organs include eyes or simple photoreceptors, statocysts, tactile organs, and chemoreceptors. Class Gastropoda Major of Groups of Gastropod Prosobranchs: This group contains most marine snails and some freshwater and terrestrial gastropods. Class Gastropoda Major of Groups of Gastropod Prosobranchs: Prosobranchs have one pair of tentacles. Sexes are usually separate. An operculum is often present. They range in size from periwinkles and small limpets horse conchs (Pleuroploca), which grow shells up to 60 cm in length, making them the largest gastropods in the Atlantic Ocean. Class Gastropoda Major of Groups of Gastropod Opisthobranchs: Opisthobranchs are an odd assemblage of molluscs that include sea slugs, sea hares, sea butterflies, and bubble shells. They are nearly all marine; most are shallow-water forms, hiding under stones and seaweed; a few are pelagic. Currently nine or more orders of opisthobranchs are recognized. Class Gastropoda Major of Groups of Gastropod Pulmonates: Pulmonates include land and most freshwater snails and slugs (and a few brackish and saltwater forms). They have lost their ancestral ctenidia, but their vascularized mantle wall has become a lung, which fills with air by contraction of the mantle floor (some aquatic species have developed secondary gills in the mantle cavity). Class Bivalvia (Pelycypoda) Bivalvia are also called Pelecypoda (pel-e-sip′o-da), or “hatchet-footed” animals, as their name implies (Gr. pelekys, hatchet, + pous, podos, foot). They are bivalved molluscs that include mussels, clams, scallops, oysters, and shipworms. Most bivalves are sedentary filter feeders that depend on currents produced by cilia on their gills to gather food materials. Unlike gastropods, they have no head, no radula, and very little cephalization. Most bivalves are marine, but many live in brackish water and in streams, ponds, and lakes. Class Gastropoda Forms and Functions Shell: Bivalves are laterally compressed, and their two shells (valves) are held together dorsally by a hinge ligament that causes the valves to gape ventrally. The valves are drawn together by adductor muscles that work in opposition to the hinge ligament. The umbo is the oldest part of the shell, and growth occurs in concentric lines around it. Class Gastropoda Forms and Functions Body and Mantle: The visceral mass is suspended from the dorsal midline, and the muscular foot is attached to the visceral mass anteroventrally. The ctenidia hang down on each side, each covered by a fold of the mantle. The posterior edges of the mantle folds are modified to form dorsal excurrent and ventral incurrent openings. In some marine bivalves the mantle is drawn out into long muscular siphons that allow the clam to burrow into the mud or sand and to extend the siphons to the water above. Class Gastropoda Forms and Functions Locomotion: Bivalves initiate movement by extending a slender muscular foot between the valves. They pump blood into their foot, causing it to swell and to act as an anchor in the mud or sand; then longitudinal muscles contract to shorten the foot and pull the animal forward. Class Gastropoda Forms and Functions Gills: Gaseous exchange occurs through both mantle and gills. Gills of most bivalves are highly modified for filter-feeding; they are derived from primitive ctenidia by a great lengthening of filaments on each side of the central axis. Class Gastropoda Forms and Functions Feeding: Most bivalves are filter feeders. Respiratory currents bring both oxygen and organic materials to the gills, where ciliary tracts direct currents to the tiny pores of the gills. Class Gastropoda Forms and Functions Internal Structures: The floor of the stomach of filter-feeding bivalves is folded into ciliary tracts for sorting a continuous stream of particles. The three-chambered heart, which lies in the pericardial cavity, has two atria and a ventricle and beats slowly, ranging from 0.2 to 30 times per minute. A pair of U-shaped kidneys (nephridial tubules) lies just ventral and posterior to the heart Class Gastropoda Forms and Functions Reproduction and Development: Sexes are usually separate. Gametes are discharged into the suprabranchial chamber to be carried out with the excurrent flow. An oyster may produce 50 million eggs in a single season. In most bivalves fertilization is external. The embryo develops into trochophore, veliger, and spat (juvenile shelled form) stages. Class Scaphopoda Phylogeny and Adaptive Diversification Molluscs Arose during Precambrian times A Cambrian fossil radula was recently found in Alberta, Canada Considered to be as protostomes, allied with the annelids in the lophotrochozoan subgroup Trochozoa Molluscs and Annelids The body is metameric; composed of There are no true serially repeated segments segments found
Coelom is greatly reduced
Well-developed coelom Opinions Molluscs were derived from a wormlike ancestor independent of annelids Share an ancestor with annelids after the advent of the coelom Share a segmented common ancestor with annelids Hypothesis: Annelids and Molluscs share a segmented ancestor is strengthened if the repeated body parts present in Neopilina, and in some chitons, can be considered evidence of metamerism. However, morphological and developmental studies indicate that these parts are not remnants of an ancestral metameric body. Perspectives on the Evolution of Repeated Body Parts: Comes from analysis of molecular characters from a wide range of molluscs, including a monoplacophoran Monoplacophora Chitons Clade Serialia
Further, clade Serialia is not the sister taxon to other
molluscs, Instead, clade Serialia is phylogenetically nested within a clade of unsegmented molluscs, indicating that the repeated structures are derived molluscan features, not ancestral features. Diversity of Molluscs Adaptation to different habitats Modes of life Wide variety of feeding methods (ranges from sedentary filter feeding to active predation) Glandular Mantle Shown adaptive capacity than any other molluscan structure: Secreting the shell and forming mantle cavity Modified into gills, lungs, siphons, and apertures Functions as locomotion Feeding processes Sensory capacity Shell Undergone a variety of evolutionary adaptations: Supports and protects the soft parts of molluscs