wo mhbe camshickman: 16.1 MOLLUSCS ‘Mollusca (mol-us’ka) (L. molluseus, sot) is one ofthe largest ani- ‘mal phyla after Arthropoda, There are over 90,000 living species and some 70,000 fossil species. Molluscs are coelomate lophotro- chozoan protosiomes, and as such they develop via spiral mosaic cleavage and make a cocloi by schizocoely, The ancestral larval stage is a tochophore, but development is variously modified within the classes “The name Mollusca indicates one of ther distinctive charactris- tics, a soft body. This very diverse group (Figure 16.1) includes chi- tons, tusk shells, snails, slugs, nudibranchs, sea butterflies, clams, nusies, aysters, squids, octopuses, and nautilses, The group ranges {com fail simple organisms lo some of the most complex of inverte- brates; sizes range from almost microscopic to the giant squid Architeuthi. These buge molluses may grow to nearly 20 m long, including their tentacles. They may weigh up (0 900 kg (1980 pounds), ‘The shels of some giant clams, Tridacna gigas, which inhabit Indo~ Pacific coral refs, each 1.5 min length and weigh more than 250 kg, ‘These are extremes, however, for prebably 80% of all moliuses are less than 10 em in maximam shell size. The phylum includes some of ‘the most sluggish and some of the swiftest and most active inverte- brates. I includes herbivorous grazers, predaceous carnivores, filter Teeders, detritus feoders, and putas Molluses occupy & great range of habitats, from the tropics to polar seas, at altitudes exceeding 7000 m, in ponds, lakes, and streams, on mud fats, in pounding suf, and in open avean from the surface to abyssal depths, They represent a variety of lifestyles, including botiom feeders, buxawers, borers, and pelagic forms, A 8 Figure 16.1. Motuses:a dversty ofttetoms Become variously adopted for eerent hats. A A ctiton Toicete neat), less Potyplacophora B.A marine sel(Caostomo), less Gesvopocs €, A nulbranch(Chromodans sp), cass Gasvopod.D, ‘Geodick clams fram Puget Sound, Washington, ‘ctopus (Octopus bares), css Cephalpode, forages stnighton Carabesncorelree. According to fossil evidence, molluscs originated in the sea, and most of them ave remained there. Mach of thet evolution courred along the shores, wheze food was abundant and habitats \were varied. Only bivalves and gastropods moved into brackish and freshwater habitats, As fle fooders, bivalves were usable Wo leave aquatic surroundings. Only slugs and snails (gastropods) actually isvaded the land. Teresa snail ae lined in het ange by thet need for humidity, shelter, and presence of calcium in the soi Molluscs are an extemely important food source for people sound the word: in 2006, 450,687,000 toanes of malluses were ar vested commercially on the eastern seaboard of the US, with te wwestem seaboard and the Gull of Mexico adding 3,870,000 and 4,537,000 tonnes, eespctvely. The value ofthe easier seaboard catch alone was neatly $6.28 hilion dolar! so the economie impor tance of molluscs cannot be overstated. A healthy mollse “Lshery depends on healthy aceans, bt recent treats to moluss populations come from an unexpected dtetion: ocean acidification. Increased amounis of CO; inthe atmosphere inte « set of chemical reactions in the oceans that lower pH, As the ocean becomes more acidic, levels of biologically available calciu decline, making it more dificult fr marin oxgeisms to form ale ciuim skeletons. Cale is essential for the foundation beneath Lv ig coral tissue (sep, 280), ad is ciel to healthy mollusc shells, Clams, oysters, mussel, and scallops prodice thinner and weaker shells when reared at low pH, Similarly, larval abalone fil to develop normally, and many cannot create normal sbels. Larval survivorship fr oysters seduced when ocean acidity increases, 0 the ayster industry in the Peific Norwest is concerned abo is3a PARTTAREE Overs of imal Lie 5278 million dolar revense (2009 data), One Oregon oyster farm adds calcium chloride and sodium carbonate to seawater to increase the amount of biologically available calcium for young animals. ‘The one bright spot in the future of molluse harvesting lies with cephalopods such as squid and octopus. They appear to be much less affected by ocean acidification than their kin, presumably because they do not have shells. Research has shown dizect adverse effects of ocean acidifica- lion on coral settlement and growth, and on the neurological evelopment of larval tropical ref fishes, bul indirect damage to the coral reef ecosystem is equally worrying. The complex food webs (see p. 280) present on the reef will change with the distrib tion and abundance of corals and coralline algae, If acidification Jeads to increased abundance of algal turfs and sea grass beds, com> ‘munity changes may act on those molluscs and other (axa not brecly impacted by the acidification of our oceans. ‘In this chapter we explore the various major groups of molluscs (Figure 162), including those groups having limited diversity (classes Mollusca ——_— oS Joe@@ad. Causofoveata Solenogastres PolyplacophoraMonoplacophora ‘Shetlath 7-8 tes Merle cout extendes long side of ot Caleateous Less of gis Seles) pasigewe ‘Maine as formseales —epulatoy spicules stpestenor nsvalve, espe shel Shell reduced lorie body ‘wth reduce foot 7 dea shel plates Aragont spicules Mane Figure 16.2_cindogron showing hypothetealielntonehne song eases of Wolics Synapomerphies hatter "troughs suber ofthese have Seen mocfes or sin some descencans Fr example the save shel os wet she cling has et Sevalrepettion off pate Gastropoda ——-Bvalvia_-—“Scaphopoda_Cephalogo¢a I . . Headlost — Heacrecuces Tossion Byssu Captacul Concentration Larsofraduhh Loss af ie efvseet ss Biya spel Tuskeshaped, ——Sphuncle opevended sone Laer compression Tet ae ‘Amstentacles and sigh Septte shel Closed exeubon syste Tek. mublayere shell Single, welldefnes shel lane Perosracu,prsmati ne nacreousbyers Shell unwalve Preoraltentacles Posterior mente ety with tr mote paisa ge Chamberes heart wn aia and ventrele Muscular fot (or fot precutson Coesreous spicules arocuces by mantle shel gone Recucton of coelom ane development ethemacoe! vatlous cages se chow, eaueed or lestin many gastropods and cephalopods, and ary getropods have undergone detersin. The bivalve shell of te Bivalvia wes derived from an lancertalunivahe shel The byssus f not present in most aGuk Dvalves But functions In larval attachment i man, therefore the byssus i consideree & ‘symapomerphy of Bvabiawom mbhe comshickmanipat 7 Caudofoveata, Solenogastes, Monoplacophors, and Seaphopods). ‘Mennbers of class Polyplacophora(chitons) are common to abundant ‘azine animals, especially in the intertidal zone. Bivalves (class Bivalvia) have evolved many species, both marine and freshwater. Class Cephalopoda (squids, cutlefish, octopuses, and their kin) ‘conttins the largest an most intelligent ofall invertebrates. Most Abundant and widespread of molluscs. however, ate snails and theit relatives (class Gastropoda). Although enormously diverse, molluses have in common a basic body plan (pp. 335-337). The coelom in ‘molluscs is limited to a space around the hear, and perhaps around the gonads aud part of the kidaeys, Although it develops embryoni- cally ina manner similar tothe coetom of annelids (see Chapter 17), the functional consequences ofthis space aze quite different because iis not used in locomotion 16.2, FORM AND FUNCTION ‘The enormous variety, great beauty, and easy availability of shells of ‘molluscs have made shell collecting a popular pastime. However, ‘many amateur shell colleciors, even though able to name hundreds of the shells that grace cur beaches, know very litle about the living animals that created thas shells and once oecupied them. Reduced to its simples dimensions, the molluse body plan has a head-foot por- tion and a visceral mass portion (Figure 16:3). The head-foot isthe ‘more alive area, containing the feeding, cephalic sensory, and loco- ‘motor organs 1t depends primarily on muscular action far is function, ‘The visceral mass is the portion containing digestive, circulatory, respiratory, and reproductive organs, and it depends primarily on cili- ay tracts for is functioning, Two folds of skin, omgrowths of the dorsal body wall, fem a protective mantle, which encloses a space ‘between the manile and body wall called the mantle cavity. The man~ le cavity houses gills (ctenidia) or hung, and in some molluscs the ‘mantle seeretesa protective shell over the visceral mass. Modifications ‘ofthe structures that form the head-foot and the visceral mass peeduce the gzeat diversity of patezns observed in Mollusca. Greate emphasis ‘on either the headfoot postion or the visceral mass portion can be ‘observed in various classes of molluscs Head-Foot ‘Most molluscs have well-developed heads, which bear their moxth ‘and some specialized suctues such as sensory tentacles ad photo- receptors, Photoreceptors may be simple, but some, such as the eyes ‘of cephalopods, ae quite complex. Within the mouth is a structure ‘unique to molluss, the radula, and usualy posterior tothe mouth is the chief locomotor organ, er foo Radula ‘The radula is a rasping, protrsible,tonguelike organ Fouad im all molluses except bivalves and. most Figure 16.3 cenersized motuse. athough tis ‘constuctis often presented os» "hypateteal ances Inaluse (HAN) most experts now reject this Interpretation Such cingrim ute howe © CHRPTER 16 Mobures 335 Characteristics of Phylum Mollusca 1, Doral body wal forms pair of fol called the mantle hich encloses the mantle eavity, i modified nto gil or lungs, nd ‘cree the shell (hel absent som); vental body wall specialized a a muscular foot, variously modified bur used ety for locomotion; radulan outs 2, Live n marine, eshwatr, and teresa habitats 5, Free-lving or oecasionaly parasitic 4, Body bilatealysymmetical (lateral asymmetry in some) tuscgmented; often with definite head 5, Triploblanie body {6 Coelom limited mainly wo area around heat, and perhaps lumen of gonads, part of kidney, and oceasionaly pat ofthe intestine 17, Sutfce epithelium usually ciated and bearing mucous glands sd sensory nerve endings 1. Complex digestive system; rasping organ (radu) usually pesent anus usually emptying into mate eavty; internal and external eliary tracts often of great functional importance 9, Cueular, diagonst, and longitudinal muscles in the body wall: ‘mantle and foot highly muscularis seme lasses or example, cephalopods and gartopods) 10, Nervous system of pazedcerebra, pleut, pedal, and visceral ‘angla, with nerve cords and subepidermal plexus; ganglia. eltalied in nerve rng in gastopods and cephalopods 11, Sensory organs of touch, sll, taste, equilibrium, and vision in ome); the highly developed direct eye (photosensive retina faze light source) of cephalopods is simular tothe ye (photosensitive cells face away from light source) of ‘vertchates tate ae skin desivative in contrast the bin oye of vertebrates 12, No asexual reproduction 13, Both monoecious snd dioecious frm; spiral cleavage: socestral ava a trochophore, many wi a veliger larva, come with diet development 14, One or to kidneys (metanephridia) opening into the pericardial cavity and usually emptying into the mantle cavity 15, Gaseous exchange by gills, lungs, mantle or body surface 16. Open circulatory system (secondarily closed in cephalopods) cof heart (usualy three chambered), blood vessels, and sinuses; respiratory pigments in blood ond Digesive lene ARM Moun Newe col coelom Foot Retractor muscles Mantle cavity