Classification of The Animal Kingdom (Metazoa)

olluscsinclude someof the best-knowninvertebrates;almost everyone
is familiar with snails,clams,slugs,squids,and octopuses.Molluscan

shells have been popular since ancient times, and some cultures still
use them as tools, containers,musical devices,money, fetishes,reli-
gious symbols, ornaments, and decorations and art objects. Evidence of histori-
cal use and knowledge of molluscs is seen in ancient texts and hierogllphics,
on coins, in tribal customs, and in archaeological sites and aboriginal kitchen
middens or shell mounds. Royal or Tyrian purple of ancient Greece and Rome,
and even Biblical blue (Num. 15:38), were molluscan pigments extracted from
certain marine snails.1 Many aboriginal groups have for millenia relied on mol-
luscs for a substantial portion of their diet and for
use as tools. Today, coastal nations annually har-
vest millions of tons of molluscs commercially for
Classification of The Animal food.
Kingdom (Metazoa) There are approximately 80,000 described, Iiv-
Non-Bilateria* Lophophorata
ing mollusc species and about the same number of
(a.k.a.the diploblasts) PHYLUM PHORONIDA described fossil species. Flowever, many species
PHYLUM PORIFERA PHYLUM BRYOZOA still await names and descriptions, especially those
PHYLUM PTACOZOA PHYLUM BBACHIOPODA from poorly studied regions and time periods, and
PHYLUM CNIDARIA EcpYSozoA it has been estimated that only about half of the liv-
PHYLUM CTENOPHOBA Nematoida ing molluscs have so far been described. In addi-
PHYLUM NEMATODA
Bilateria PHYLUM NEMATOMORPHA
tion to three familiar molluscan classescomprising
(a.k.a.the triploblasts) the clams (Bivalvia), snails and slugs (Gastropoda),
Scalidophora
PHYLUM XENACOELOMORPHA
PHYLUM KINORHYNCHA and squids and octopuses (Cephalopoda), five
Protostomia PHYLUM PRIAPULA other extant classesexisl chitons (Polyplacophora),
PHYLUM CHAETOGNATHA PHYLUM LORICIFERA
tusk shells (Scaphopoda), Neopilina and its kin
SprRarn Panarthropoda
PHYLUM PLAWHELMINTHES (Monoplacophora), and the vermiform sclerite-
PHYLUM TARDIGRADA
PHYLUM GASTROTRICHA PHYLUM ONYCHOPHORA
bearing aplacophoran classes-Caudofoveata
PHYLUM RHOMBOZOA PHYLUM ARTHROPODA (or Chaetodermomorpha) and Solenogastres (or
PHYLUM ORTHONECTIDA cnustncee*
sUBPHYLUM Neomeniomorpha). Although members of these
PHYLUM NEMERTEA SUBPHYLUM HEXAPODA
PHYLUM MOLLUSCA
eight classesdiffer enormously in superficial ap-
SUBPHYLUM MYRIAPODA
PHYLUM ANNELIDA pearance, there is a suite of characters that diag-
SUBPHYLUM CHELICERATA
PHYLUM ENTOPROCTA nose their fundamentalbody plan (Box 13A).
Deuterostomia
PHYLUM CYCLIOPHORA
PHYLUM ECHINODERMATA
Gnathifera PHYLUM HEMICHORDATA
PHYLUM GNATHOSTOMULIDA
This chapterhas been revised by Richard C. Brusca,David R.
PHYLUM CHORDATA
Lindberg, and Winston F. Ponder
PHYLUM MICROGNATHOZOA
*Paraphyletic group lArchaeologicalsitesin Israel reveal the probable use of two
PHYLUM ROTIFERA
muricid snails(Murex brandarisand Trunculariopsis trunculus)
as sourcesof the Royal purple dye.
454 Chaoter Thirteen
BOX 13A Characteristics
of the
Taxonomic History and PhylumMollusca
Classification 1. Bilaterallysymmetrical (or secondarily asymmetri-
cal), unsegmented, coelomate protostomes
Molluscs carry the burden of a very long and convolut- 2. Coelom limited to small spaces in nephridia, heart
ed taxonomic history, in which hundreds of names for and gonads
various taxa have eome and gone. Aristotle recognized 3. Principal body cavity is a hemocoel (open circula-
molluscs, dividing thern into Malachia (the cephalo- tory system)
pods) and Ostrachodermata (the shelled forms), the lat- 4. Viscera concentrated dorsally as a "visceral mass"
ter being divided into univalves and bivalves. joannes 5. Body covered by a cuticle-covered epidermal sheet
jonston (or ]onstonus) created the name Mollusca2 in of skin, the mantle
1650 for the cephalopods and barnacles, but this name 6. Mantle with shell glands that secrete calcareous
was not accepted rmtil it was resurrected and redefined epidermal sclerites, shell plates, or shells
by Linnaeus nearly a hundred years later. Linnaeus's 7. Mantle overhangs and forms a cavity (the mantle
Mollusca included cephalopods, slugs, and pteropods, cavity) in which are housed the ctenidia, osphradia,
as well as tunicates, anemones, medusae, echinoderms, nephridiopores, gonopores, and anus
and polychaetes-but included chitons, bivalves, uni- B. Heart situated in a pericardial chamber and com-
valves, nautiloids, barnacles, and the serpulid poly- posed of a single ventricle and one or more sepa-
chaetes (which secrete calcareous tubes) in another rate atria
group/ Testacea.In 7795 Ceorges Cuvier published a 9. Typically with large, well-defined muscular foot,
revised classification of the Mollusca that was the first often with a flattened creeping sole
to approximate modern views. Henri de Blainville 10. Buccal region provided with a radula and muscular
(1825) altered the name Mollusca to Malacozoa, which odontophore
won little favor but survives in the terms malacology, 1 1. Complete (through) gut, with marked regional spe-
malacologist, etc. cialization,including large digestive glands
Much of the nineteenth century passed before the 12. With large, complex metanephridial "kidneys"
phylum was purged of all extraneous groups. In the 13. Cleavage spiral and embryogeny protostomous
1830s,j. Thompson and C. Brumeister identified the 14. With trochophore larva, and a veliger larva in two
larval stages ofbarnacles and revealed them to be crus- major groups
taceans, and in 1866 Alexander Kowalevsky removed
the tunicates from Mollusca. Separation of the brachio-
pods from the molluscs was long and controversial
and not resolved until near the end of the nineteenth possess calcareous sclerites by placing Pollplacophora
century. as the sister taxon of the aplacophorans (Caudofoveata
The first sclerite-covered wormlike aplacopho- + Solengastres).Aculifera was sometimes also called
rans, members of what today we recognize as the class Amphineura, although this latter term has also been
Caudofoveata, were discovered in 1841by the Swedish used by some workers to refer only to chitons. Sclerites
naturalist Sven Lov6n. He classified them with ho- are spicules, scales/ and so on that cover or are embed-
lothuroid echinoderms because of their vermiform ded in the epidermis of molluscs and are often calcified.
bodies and the presence of calcareous sclerites in the The history of classification of species in the class
body walls of both groups. In 1886, another Swede, Gastropoda has been volatile, undergoing constant
Tycho Tullberg, described the first representative of change since Cuvier's time. Most modern malacolo-
the other aplacophoran group-the Solenogastres. gists adhere more or less to the basic schemes of Henri
Ludwig von Graff (7875) recognized both groups as Milne-Edwards (1848)andJ. W. Spengel (1881).The for-
molluscs and they were united in the Aplacophora mer, basing his classification on the respiratory organs,
in1876 by Hermann von Ihering. The Aculifera hy- recognized the groups Pulmonata, Opisthobranchia,
pothesis of Am6lie Scheltema unites molluscs that and Prosobranchia. Spengel based his scheme on the
nervous system and divided the gastropods into the
2The name of the phylum derives from the Latin
molluscus, mean- Streptoneura and Euthyneura. hr subsequent classifica-
ing "soft," in allusion to the similarity of clams and snails to tions, Streptoneura was equivalent to Prosobranchia;
the mollusca, a kind of OId World soft nut with a thin but hard
Euthlmeura included Opisthobranchia and Pulmonata.
shell. The vernacular for Mollusca is often spelled mollusks in
the United States, whereas in most of the reit of the world it is The bivalves have been called Bivalvia, Pelecypoda,
typically spelled molluscs. In biology, a vernacular or diminutive and Lamellibranchiata. More recently, anatomical,
name is generally derived from the proper Latin name; thus the
ultrastructural, and molecular studies have brought
custom of altering the spelling of Mollusca by changing the c to k
seems to be an aberration (although it may have its historic roots about considerable changes to molluscan classification,
in the German language, which does not have the free-standing c; as outlined below. Manv taxa have multiple names and
e.g., Molluskenkunde). We prefer the more widely used spelling
the more commonly eniountered ones arênoted below.
"molluscs," which seems to be the proper vernacularization and is
in line with other accepted terms, suchas molluscan, molluscoid, Molluscan classification at the generic and species
molluscivore, etc. levels is also troublesome. Many species of gastropods
PHYLUM MoLLUSCA 455
and bivalves are also burdened with numerous names C OH OR TH Y P S OGA S TR OP OD A

H igher caeno-
(synonyms) that have been proposed for the same spe- gastropods
cies. This tangle is partly the result of a long history of S U P E R OR D E LITTOR
R IN IMOR P HA
Per iwinKI es,
amateur shell collecting beginning with the natural his- cowries,tritonshells,etc.
tory cabinets of seventeenth century Europe, which re-
quired documentation and promoted multiple taxono- S U P E R OR D E R N E OGA S TR OP O DA WheI Ks.
volutes,rock shellsetc.
mies and names ba3ed only on shell characters. Today,
species are recognized based on a combination of shell, SUBCLASSHETEROBRANCHIAMarine, freshwater
anatomical, and, most recently molecular characters. and landsnails,most sea slugs,all landslugs,and some
However, because of the tremendous diversity of gas- "falselimpets"
tropods and bivalves many species still remain known
"LOWERHETEROBRANCHIA"A few orimitivehetero-
only from their shells.
branchgroupsincludingsundialshells,valvatids,etc.
Only taxa with extant members are included in the
following classification and not all families are listed INFRACLASS EUTHYNEURA ..Ooisthobranchs,' and
in the taxonomic synopses. The classification is mostly "pulmonates"
ranked, but in a few cases unranked group names are COHORTNUDIPLEURA Side-gilledsea slugs and
used.3 Examples of the major molluscan taxa appear in nudibranchs
Figure 13.1.
COHORTEUOPISTHOBRANCHIABubbleshells,sea
hares,pteropods,etc.
ABBREVIATED CLASSIFICATION OF
COHORTPANPULMONATA "Pulmonates,"pyra-
THE PHYLUM MOLLUSCA midellids,sacoglossansea slugs,most land snails,all
CLASSCAUDOFOVEATA Caudofoveatanaolacoohorans landslugs
(spicule"worms")
CLASSBIVALVIA Clams and their kin (bivalves)
CLASSSOLENOGASTRES
Solenogasteraplacophorans
(spicule"worms") SUBCLASSPROTOBRANCHlA "Primitive" deoosit-
feedingbivalves
CLASSMONOPLACOPHORA Monoplacophorans.Deep
ti,..^^+ t;t'^ SUBCLASSAUTOBRANCHlA "Lamellibranch"
susoen-
^^^
DVd. iltItuut-ilnu
sion-feedingbivalves
CLASS POLYPLACOPHORAChitons, with eight shell
COHORTPTERIOMORPHIAMussels,oysters,scal-
VAIVES
lops,and their kin
CLASSGASTROPODA Snails,slugs and limpel
COHORTHETEROCONCHIAMarineand freshwater
SUBCLASSPATELLOGASTROPODA
The true limpets CIAMS
SUBCLASSVETIGASTROPODA "Pdmitive"marinetop- MEGAORDER PALAEOHETERODONTAFTeshwateT

shellsnails,abalonesand "limpets" clams (mussels),broch shells
SUBCLASS NERITIMORPHA Marine,landand freshwa- MEGAORDER
HETERODONTAMost marineclams
ter neritesnailsand "limoets"
SUPERORDER ARCHIHETERODONTA
A few fami-
SUBCLASSCAENOGASTROPODAMarine,freshwater liesof primitivemarineclams
and land snails (creepers,periwinkles,conchs, whelks,
SUPERORDER EUHETERODONTAThe maioritvof
cowriesetc.) and some "limpets"
marineand some freshwaterclams
"ARCHITAENIOGLOSSA"Nonmarinebasal caeno-
CLASSSCAPHOPODA TusKshells
gastropods(paraphyletic)
CLASSCEPHALOPODANautilus,squids,octopuses
I NF R AC L AS SSOR BE O C O N C H A AII remai ni no
caenogastropods SUBCLASSPALCEPHALOPODA
SUPERORDERCERITHIOMORPHA
Creeoers.turret COHORTNAUTILIDIAChamberednautilus
shells,etc.
SUBCLASSNEOCEPHALOPODA
rMultitudes of extinct molluscs have been described. Perhaps the

COHORTCOLEOIDEAOctopuses,squids,cuttlefish
most well-known are some of the groups of cephalopods that had
hard external shells, similar to those of living Nautilus. One of
S U P E R OR D EOC
R TOP OD IFOR MES
O ct oouses,
these groups was the ammonites. They diffeied from nautiloids vampiresquid
in having shell septa that were highly fluted on the periphery,
forming complex mazelike septal sutures. Ammonites also had S U P E R OR D E RD E C A P OD IFOR M ESCut t lef ish,
the siphuncle lying against the outer wall of the shell, as opposed S OU Id
to the condition seen in manv nautiloids where the siphuncle runs
t h r o u e h t h e c e n te r o f th e sh e ll.
I
I
i
I
Figure 13.1 Morphological diversity among the (Scaphopoda). (M) Scallops (Bivalvia: Pteriomorphia:
mof fuscs. (A) Laevipilina hyalina (Monoplacophora). Pectinidae), with a hermit crab in the foreground. (N) The
(B) Mopalia muscosa, the mossy chiton (Polyplacophora). giant clam Tridacna maxima (note zooxanthellate mantle)
(C) Epimenia australis (Solenogastres). (D) Haliotis rufe- from the Marshall lslands, Northwest Pacific (Bivalvia:
scens, the red abalone, (Gastropoda); note the exhalant Heterodonta: Cardiida). (O) The European cockle Acantho-
holes in the shell. (E) Conus, a predatory neogastropod; cardia tuberculafa (Bivalvia: Heterodonta: Cardiida).
note anterior siphon extending beyond shell. (F) The Note the partly extended foot. (P\ Lima, a tropical clbm
common garden snail, Cornu aspersum (Gastropoda). that swims by clapping the valves together (Bivalvia).
(G) Aplysia, the sea hare (Gastropoda: Euopisthobranchia). (Q) The highly modified bivalve Brechites. (Heterodonta:
(H) The chambered Nautilus (Cephalopoda). (l) Octopus Poromyata). Brechites are known as watering pot shells.
bimacu loides (Cephalopoda). (J) Sepiofeuthis lessoniana, They begin their life as a typical small bivalve, but then
the bigfin reef squid (Cephalopoda). (K) Histioteuthis, a secrete a large calcareous tube around themselves
pelagic squid (Cephalopoda). (L) Fustiaria, a tusk shell through which water is pumped for suspension feeding.
PHYLUMMOLLUSCA 457
(L)
(A)
(c) Radular Dorsal Gonopericardial

Esophagus aorta Intestine duct
1"t stomach Ventricle
Salivary
gland Pericardium
Cerebral ganglion
Suprarectal
ganglion
Mouth/
Oral shield Mantle cavity
Ventral 1= pedal) Lateral

nerve cord Odontophore Vertical Ventral
Digestive nerve Vertical
sePtum 1= pedal) cecum cord sePtum
sinus
Vestibular
Pedal pit opening
Mouth
Pedal groove
Pedal pit
Pedal groove
Gill folds
SYNOPSES OF MOLLUSCAN GROUPS CLASSSOLENOGASTRES (= NEOMENIOMORPHA) (Figure

CLASSCAUDOFOVEATA (= CHAETODERMOMORPHA) 13.2D-14. Spicule"worms." Marine, benthic;body vermi-
(Figure13,2A-C). "worms."
Spicule Marine,benthic,bur- form and nearly vestibulum
cylindrical; (= atrium)with sen-
rowing;bodyvermiform, sory papillaeanterior
to the mouth; smallposteriormantle
cylindrical,
lackinganytraceof
a shell;bodywallwitha chitinous cuticleand imbricating cavitylackingctenidiabut oftenwithrespiratory folds;body
scale-likearagonitic
calcareoussclerites;
mouthshieldan- wallwith a chitinous cuticleand imbuedwith calcareous
(as
the mouth;smdl posterior
teriorto or surrounding mantle sclerites spinesor scales);with or withoutradula;her-
cavitywith a pairof bipectinate maphroditic; pedalglandsopeningintoa pre-pedal ciliary
ctenidia;radulapresent;
gonochoristic. pit, foot weakly muscular, narrow, and can be retracted
Withoutfoot, eyes,tentacles,statocysts,
"pedalgroove."Withouteyes,ten-
crystalline
style,osphradia, About120species; intoa ventralfurrowor
or nephridia.
burrowin muddysedimentsand consumemicroorgan- tacles, statocysts,crystalline
style,osphradiaor nephridia.
ismssuchas foraminiferans. (e.9.,Chaetoderma, Chevro- About 260 describedspecies, but manyundescribed spe-
derma,Falcidens,Limifossor,Prochaetoderma, Psilodens, cies arethoughtto exist;epibenthiccarnivores,oftenfound
Scufopus) on (andconsuming) cnidarians and a few othertypesof
PHYLUMMOLLUSCA 459
50 pm
50 pm
(L) Cerebral Buccal Lateral

nerve cord Radula
ganglion nerve Dorsal cecum
Vestibule
Salivary gland
)l Esophagus
Ventral
nerve cord I
Pedal Pit
Anterior
uoontopnore
raouralrpocKet
Figure 13.2 General anatomy of aplacophorans. tricarinata, ventral view (X-ray micro-CT). (J) Macellomenia
(A-C) Caudof oveata. (A) C h aetod e rm a p ro d u ctu m. morseae. SEM of ventral surface showing two types
(B) Chaetoderma loveni. (C) Internal anatomy ot Limi- of scale-like sclerites surrounding the foot and spiny
fossor (highly stylized sagittal section drawing). (D-L) sclerites covering the rest of the surface of the body.
Solenogastres. (D) Kruppomenia minima. (E) Pruvotina (K) Macellomenia schanderl. SEM of ventral surface of
impexa, ventral view. (F) Proneomenia antarctica. anterior end showing densely ciliated pedal pit and
(G) Epimenia verrucosa. The body is covered with warts. mouth. (L) Anterior region of Spengelomenia bathybia
(H) Neomenia carinata, ventral view. (l) Entonomenia (highly stylized sagittal section drawing).
invertebrates. are proba-

and Caudofoveata
Solenogastres aroundfoot encloses3-6 pairsctenidia;2 pairsgonads;
bly sistergroupsand aresometimesregardedas subclass- 3-7 pairsnephridia;2 pairsheartatria;a pairof statocysts;
es withinthe classAplacophora.(e.g.,Alexandromenia, withradulaanddistinctbutsmallheadregion;withouteyes;
Dondersia,Epimenia,Kruppomenia,Neomenia,Proneo- shortoraltentaclespresentaroundmouth;with posterior
menia,Pruvotina,Rhopalomenia, Spengelomenia,Wirenia) anus;withouta crystalline style;gonochoristic or, rarely,
hermaphroditic(Figures13.'1A and13.3).Untilthefirstliving
CLASSMONoPLACOPHORA Witha
Monoplacophorans. galatheae)was discoveredby the Dan-
species(,,leopilina
single,cap-likeshell;foot formsweaklymuscularventral ishGalatheaExpedition in 1952,monoplacophorans were
disc,with8 pairsof retractor shallowmantlecavity
muscles;
460 ChaoterThirteen
Anterior lip
Mantle groove
Minute preoral tentacle
Velum
Statocyst
ephridium
Lateral nerve cord
Nephridia serving
gonoducts
Pedal nerve cord

Figure 13.3 General anatomy of a mono-
placophoran (Neopilina). (A) Dorsal view
Ctenidium
(shell). (B) Ventral view. (C) Photograph of
the ventral surface of a oreserved specimen
of Neopilina. (D) Ventral view, foot removed.
Heart atria (E) One of the gills.
Lateropedal sometimes haveshelleyes(Figure13.43C,D). Marine,inter-

comm$Srlle tidalto deepsea.Chitonsareuniquein theirpossession
Anus of
8 separateshellplates,calledvalves,and a thickmarginal
girdle;about850 described speciesin onelivingorder.4
knownonlyfromlowerPaleozoic fossils.Sincethentheir
unusualanatomyhas beena sourceof muchevolutionary ORDER NEOLORICATA Shellswith uniquearticula-
speculation.Monoplacophorans are limpet-like
in appear- platesthat inter-
mentumlayer,whichformsinsertion
ance,livingspeciesare lessthan3 cm in length,and most lockthevalves.
liveat considerable
depths.About30 described species,in
8 genera(AdenopiIina, LaevipiIina, MonopIacophorus,/Veo- SUBORDER LEPIDOPLEURIDA Chitonswithouter
pilina,Rokopella,Veleropilina,Vema,Micropilina). edgeof shellplateslackingattachment teeth;gir-
dle not extendingoverplates;ctenidialimitedto a
CLASSPOLYPLACOPHORA Chitons(Figures 13.1Band few posteriorpairs.(e.9.,Choriplax,
Lepidochiton,
13.4).Flattened, elongated molluscs witha broadventral Lepidopleurus,Oldroydia)
foot and B dorsalshellplates(composedof aragonite);
mantleformsthickgirdlethatbordersandmaypartlyor en- SUBORDER CHITONIDA Outer edges of shell
tirelycovershellplates;epidermis plateswith attachmentteeth;girdlenot extending
of girdleusuallywithcal-
careousspines,scales,or bristles;mantlecavityencircles overplates,or extending
partlyoverplates;ctenidia
foot and bearsfrom6 to morethan80 pairsof bipectinate occupying mostof mantlegroove,exceptnearanus.
ctenidia;1 pairnephridia;headwithouteyesor tentacles; (e.9.,Callistochiton,
Chaetopleura, lschnochiton,
crystalline
style,statocystsand osphradia absen| nervous
systemlackingdiscreteganglia,exceptin buccalregion; alJncommon, aberrant individuals have been found with only 7
well-developed radulapresent.Shellcanals(aesthetes) valves.
PHYLUMMOLLUSCA 461
Figure 13.4 Generalized anatomy (A) Head (anterior) valve (B) Head
of chitons (Polyplacophora). (A,B)
A typical chiton (dorsal and ventral Mouth
views). (C) The Pacific lined chiton, Palps
Intermediate
Tonicella lineata. (D) Dorsal view of a valve
Mantle groove
chiton, shell plates (valves) removed.
(E) Dorsal view of a chiton, dorsal Girdle
musculature removerd to reveal internal
Ctenidia
organs. (F) Dorsal view of a chiton,
showing extensive nephridia. (G) The Exhalant ch
arrangement of internal organs in a
Inhalant charnber
chiton (lateral view).
Girdle
Tail (posterior)valve
Anterior insertions of
longitudinal and (D)
lateral muscles
Buccal Buccal
Oblique muscle
rnuscles
Esophagus
Longitudinal Dorsal
muscle artery
Nephridium
Position of
dorsal artery
Digestive 'Intestine
gland
Transverse
muscle
Gonad
Oviduct
Ventricle
Pericardium Atrium
Posterior insertion of
longitudinal and
lateral muscles
Groove
between
head and foot
Foot
Main nephridial (G)

canal Dorsal
Rieht atrium .. . Inteshne
Gonad valves / artery
Gonoduct Ventricle Mantle
Nephrostome Rericardium
in pericardium
Gonopore
Nephridiopore Salivary
gland
Pericardial
Girdle
chamber
Renopericardial Anus
canal \\H ead
Right Right Digestive Foot -Mouth
Pedal Nerve
nephridiopore gonoPore nephridium gland nerves ring Esophagus
462 ChaoterThirteen
(A) Mantle edge papillae protruding (c) Right nephridium

out of shell foramen Pericardial Digestive gland
Slit in Stomach
mantle skirt
Style sac region
of stomach
Epipodial
with papillae sense organ
(B)
o
E
Ctenidium Epipodial
(in mantle cavity)
Figure 13.5 General anatomy of limpet-like gastropods.

(A) The vetigastropod limpet F/ssure//a(Fissurellidae)
(lateral view). (B) The patellogastropod limpet Loffra
(Lottiidae) (ventral view). The arrows indicate the direction
of water currents. (C) The vetigastropod limpet Puncturella
(Fissurellidae),removed from shell and seen from the left.
The arrows indicate the water currents. Certain structures
are visualizedthrough the mantle skirt: ctenidium, eye,
anus, and epipodial sense organs.
Shell muscle
glands;ctenidiasometimes
and hypobranchial lostand re-
placedwithsecondary
gasexchange structures
Gastropods compriseapproximately 70,000describedliv-
ingspeciesof marine,terrestrial,
andfreshwater snailsand
slugs.The classwas traditionally dividedintothreesub-
classes:prosobranchs (largely
shelledmarinesnails),opis-
Katharina, Lepidozona, Mopalia, Nuttallina, thobranchs (marine
slugs),andpulmonates (terrestrial
snails
Placiphorella, Schizoplax, Tonicella) andslugs).However, recentanatomical andmolecular stud-
ieshaveshownthatclassification to be incorrect.
as reflect-
SUBORDER ACANTHOCHITONIDA. Outeredgeof ed in theclassification
below.
shellplateswith well-developedattachment teeth;
shellvalvespartiallyor completelycoveredby gir- SUBCLASSPATELLOGASTROpODA Cap-shaped
(limpets) with porcelaneous, nonnacreous shell;oper-
dle;ctenidia
do not extendfulllengthof foot.(e.9.,
Acanthochitona,Cryptochiton, culumabsentin adult;cephalictentacleswith eyesat
Cryptoplax)
outerbases;raduladocoglossate, with ironimpregnat-
cLAsSGASTRoPoDA Snails,limpetsand slugs(Figures ed teeth,restof gut with largeesophageal glandsand
13.1D-G,13.5,13.6,and 13.7).Asymmetrical molluscs simplestomachlackinga crystalline style;intestine long
withsingle,usually spirally
coiledshellintowhichbodycan and looped;gillconfiguration variable,singlebipecti-
be withdrawn; shelllostor reducedin manygroups;dur- natectenidium sometimes present(Figure 13.58),and/
ingdevelopment, visceralmassand mantlerotate90-180' or with mantlegroovesecondary gills,or gillslacking;
on foot (a processknownas torsion),so mantlecavitylies shellmuscledividedintodiscretebundles;mantlecavity
anteriorlyor on rightside(ratherthanposteriorly as in other withoutsiphonor hypobranchial glands;2 rudimentary
molluscs), and gut and nervoussystemaretwisted;some osphradia; singleatria;2 nephridia; usually gonochoris-
taxahavepartlyor totallyreversed the rotation(detorsion); tic;nervoussystemweaklyconcentrated, pleuralganglia
with muscularcreepingfoot (modified in swimmingand nearpedalganglia,pedaland lateralcordspresent.Pri-
burrowingtaxa);foot with operculumin larvaand oftenin marilymarinewitha few estuarine species;herbivorous.
adult;headwitheyes(oftenreducedor lost),and 1-2 pairs Thepatellogastropods include6 families:Patellidae(e.9.,
of tentacles,and a snout;mostwith radulaand somewith PateIIa, ScuteIIastra),NacelIidae(e.9.,CelIana), Lottiidae
style,the latterbeingabsentin mostprimitive
crystalline and (e.9.,Lottia),
Acmaeidae (e.9.,Acmaea),Lepetidae (e.g.,
in manyadvanced groups;1-2 nephridia; mantle(=pallium) Lepeta),and Neolepetopsidae (e.9.,lVeo/epefopsis).
usuallyformsanteriorcavityhousingctenidia, osphradia, Theseareoftenregardedas the "true"limpets.
PHYLUM MOLLUSCA 463
Posterior aorta Figure 13.6 Generalanatomy of coiled gastropods.

rd (A)A generalizedcoiled-shellgastropod(female),indi-
cating positionsof internalorgans.(B)The periwinkle,
Littorina,removedfrom its shell (anteriorview).
Nephridium
-
(B)
Visceral mass
Ctenidium
Anus
Ospfuadium
Cephalic Mantle edge

tentacle
Eye
lip Buccal mass
aPPararus
SUBCLASS VETIGASTROPODA SheIIsboth Oorcela- ORDER NEOMPHALIDA Comprises manyof the hot
neousand nacreous;cephalictentaclesusuallywith ventsnailsand limpetsNeomphalidae (e.9.,Neom-
eyes on short processeson outer bases;operculum phalus), Peltospi pira),Lepetodri
ridae(e.9.,Peltas Iidae
usuallycircular,with a centralnucleusand oftenmany (e.9.,Lepetodilus).
spirals,hornyor calcareous; radulausuallyrhipidoglos-
ORDERCOCCULINIDA (= COCCULINIFORMES lN
sate(withnumeroustransverse rowsof teeth),restof gut
PART) Thesmall,deep-seawood and bonelimpets
with esophagushavinglargeglands,complexstomach
(e.9.,Cocculina).
Cocculinidae
with stylesac but no crystallinestyle,loopedintestine;
1-2 bipectinatectenidia;shellmusclespairedor single; SUBCLASS NERITIMORPHA Shellcoiled,limpet-like,
mantlecavitywith 2 hypobranchial glands,2 aIria,and or losi fl-itiscaniidae). Shellporcelaneous, with interior
2 nephridia;usuallygonochoristic; malegenerally with- whorlsreabsorbed in manycoiledgroups;operculum
out penis;nervoussystemweaklyconcentrated, gan- typicallypresent,of few spiralsandwithnon-central nu-
gliapoorlyformed,pedalcordspresent;1-2 osphradia, cleus,hornyor calcified, usually
withinternal peg;shell
small,inconspicuous. Allmarine andbenthic. Manyspe- muscledividedintodiscretebundles;onlyleftctenidium
ciesare microdetritivoresor feedon filmsof bacteriaor present;hypobranchial glandsoftenlost on left side;
otherorganisms, or aremicroherbivores; somearemac- stomachhighlymodified;rightnephridium incorporated
roherbivores,somegrazingcarnivores, and a few sus- intocomplexreproductive systemwithmultiple openings
pensionfeeders.Mostgastropodsfoundat hydrother- into mantlecavity;radularhipidoglossate; most spe-
malvents,coldseeps,andon deepseahardsubstrates ciesgonochoristic, wiih copulatorystructures;nervous
arevetigastropods. Vetigastropods compriseabout30 systemwith gangliaconcentrated, pleuralganglianear
familiesand whilethe internalclassification
remainsun- pedalganglia,pedalcordspresent.Globallydistributed
settled,threemaingroupsare oftenrecognized, which in marine,estuarine, freshwater,andterrestrialhabitats.
we treathereas orders. Thereare 9 familiesof neritimorphans, four of which,
Helicinidae (e.9.,Alcadia,Helicinia),Hydrocenidae (Hy-
ORDER TROCHIDAMostof the vetigastropods, in-
drocena,Georissa), (e.9.,Proserpinella),
Proserpinellidae
cludingthe slit-shelled snailsPleurotomaridae (e.9.,
and Proserpinidae (Proserpina)areexclusively terrestrial;
Perotrochus,Pleurotomaria), Scissurellidae(e.9.,
alsoNeritopsidae (neritopsids,
lVerltopsr.s),
Titiscaniidae
Scrssure/k)and Anatomidae(e,9.,Anatoma),Ihe
(titiscaniid,
Titiscan (nerites,
ia),Neritidae e.9.,Nerita,The-
abalonesHaliotidae (e.9.,Haliotis),
the keyholeand
odoxus),Neritiliidae (cavenerites,e.9.,Pisulina,Neritilia)
slitlimpetsFissurellidae (e.9.,Diodora,Fissurella,
Lu-
and Phenacolepadidae (Phenacolepas).
capinella,Puncturella), deep sea limpetscomprising
the Lepetellidae and relatedfamilies(e.9.,Lepetella, SUBCLASS CAENOGASTROPODA Shellmainlypor-
Pseu dococculina),trochidsTrochidae (e.9.,Trochus, celaneous; operculumusuallypresentand corneous,
Monodonta)andrelatedfamiliessuchas Calliostomat- rarelycalcified,
withfew spiralsand usuallywith a non-
idae(e.9.,CalIio stoma),Margaritidae(e.9.,Margarites), centralnucleus,mostlynon-nacreous, rarelywithinternal
Tegulidae (eg , Tegula), andturbansTurbinidae (e.9., peS(s);headwithpairof cephalictentacles,witheyesat
Turbo,Astrea). outerbases;mantlecavityasymmetrical, with incurrent
464 Chapter Thirteen
Figure 13.7 More gastropod anatomy; some caenogastropods

(A-C) and heterobranchs (D-J). (A) The petagic sheiled heteropod
Carinaria (Caenogastropoda). (B) Anatomy of Carinaria. (C) The
shell-less heteropod Pterotrachea (Caenogastropoda). (D) The
pelagic shelled pteropod C/lo (Heterobranchia: Euopisthobranchia).
The arrows indicate the direction of water flow; water enters all
around the narrow neck and is forcibly expelled together with
fecal, urinary, and genital products by contraction of the sheath.
(E) A swimming pteropod, Corolla (Heterobranchia: Euopistho-
branchia). (F-l) Various nudibranchs (Heterobranchia: Nudipleura).
(F) A dorid nudibranch, Diaulula. (G) An aeolid nudibranch,
Phidiana. (H) A dorid nudibranch, Chromodoris geminus (from
the Red Sea). (l) The eastern Pacific "Spanish shawl" aeolid nudi-
branch, Flabellina. (J) The lettuce sea slug, Tridachia crispata
(Heterobranchia: Panpulmonata), from the Cariboean.
(B) Esophagus (D) Foot

(parapodium)
:----_l
Intestine
Longitudinal
Salivary Protoconch
gland
Ctenidium
Median keel on shell
Visceral mass
Stomach Ctenidia
(F) Branchial plume
Rhinophore
Cerata
Rhinophore
Cephalic
tentacle
PHYLUMMOLLUSCA 465
dividedinto two main groups,Cerithiomorpha

and
Hypsogastropoda.
SUPERORDER CERITHIOMORPHA Usuallywithout
a penis;eggsusuallylaidin jelly,oftenin strings,or
arebrooded.Theanterioraperturemayor maynot
havea notch,whichhousesa shortsiphon.Include
marine,brackish,and freshwaterspecies.About
19 familiesare recognized, includingthe marine
Campanilidae (e.9.,Campanile), Cerithiidae (horn
shells,e.9., Cerithidea,Cerithium,Liocerithium),
Siliquariidae(slitworm shells,e.9.,Siliquaria),
and
(toweror turretshells,e.9., Turritella);
Turritellidae
andthefreshwater Melanopsidae (e.9.,Melanopsls),
Thiaridae(e.9.,Thiara),and Pleuroceridae (e.9.,
openingon anteriorleft,sometimeselaborated intoan Pleurocera).
inhalantsiphon;rightctenidiumlost;leftctenidium mono-
pectinate;lefthypobranchialglandlost;rightnephddium COHORT HYPSOGASTROPODA Comorisesihe re-
lost exceptfor remnantincorporated into reproductive mainingcaenogastropods. The anteriormantlemay
system;heartwithonlyleftatrium,Radulataenioglossate be simpleor can be enrolled formingan anterior si-
(7 rows of teeth),ptenoglossate (manyrowsof similar phonwhichemerges froman anteriornotchintheap-
teeth),rachiglossate (1-3 rowsof teeth),or toxoglos- ertureor. in some.is containedwithinan extension of
sate(teethmodifiedas harpoons), or occasionally lost. theshell,thesiphonal canal.Malewithcephalicpenis;
Higherformswith concentrated ganglia,pleuralganglia eggsusuallylaidin capsulesor sometimes brooded.
usuallynearcerebralganglia,pedalcordsusuallyab- Nervoussystemconcentrated; operculum, if present,
sent;osphradium conspicuous, oftenlarge,sometimes chitinous, Thislargegroupis divided
rarelycalcareous.
surfacesubdivided Mostcaenogastropods
intolamellae. intothe LittorinimorphaandNeogastropoda.
aregonochoristic. Thecaenogastropods, comprisethe SUPERORDER LITTORINIMORPHA Classification
former"mesogastropods" andneogastropods, andthey unsettled;includesthe marinegrazingsnails
areoftendividedintotwo groups,as follows: (periwinkles,
Littorinidae e.9.,Littorina),
a numberof
"ARoHITAENIoGLoSSA" Althoughthis is not a mono- small-sized marinefamilies includingthe Rissoidae
phyleticgroup,we retainit informally.Architaenioglos- (e.9.,Rlssoa,Alvania),and largersnailssuchas the
sansdifferfromothercaenogastropods in detailsof their Strombidae (conchsor strombids,e.9.,Strombus),
nervoussystemand in the ultrastructure of theirsperm andthecarrier shellsXenophoridae (e.9.,Xenophora).
andosphradia. Theyaredividedamong10 families, in- Also includesthe uncoiledsuspension-feeding
(applesnails,e.9., "worm"gastropodsVermetidae (e.9.,Serpu/orbis,
cludingthefreshwater Ampullariidae
Ampullaria,Pomacea,Pila)andViviparidae(riversnails, Dendropoma)andthelimpet-like Hipponicidae(e.9.,
e.9.,Viviparus),
and the terrestrial
Cyclophoridae (e.9., Hipponix)whicharedepositfeeders, whileCapulidae
Cyclophorus) andseveralrelatedfamiliessuchas Diplom- (e.9.,Capulus) attachto othermolluscsand mostly
matinidae(e.9.,DipIommatina andOpi stho stoma). feedon theirfeces.TheslippershellsCalyptraeidae
(e.9.,Calyptraea,Crepidula,Crucibulum)are sus-
INFRACLASS SORBEOCONCHA This groupingcon- pension-feeders. The Carinariidae (one of sev-
tainsall the rest of the caenoqastrooods.
Theseare eralfamiliesof pelagicmolluscscollectively called
heteropods, e.9.,Carinaria)
alsohavea cap-shaped (e.9.,Turris)
andseveral
otheralliedfamilies
including
shell.5Cypraeidae (cowries, e.g.,Cypraea) are her- theaugershellsTerebridae
(e.9.,Terebra).
bivoresor grazingcarnivores, whileseveralother
littorinimorph snail-like
familiesare strictlycarnivo- SUBCLASS HETEROBRANCHIA The heterobranchs
rous,including Naticidae (moonsnails, werepreviously organized as two subclasses-Opis-
e.9.,Natica,
Polinices) thatfeedmostlyon bivalves, thobranchia(seaslugsand theirkin)and Pulmonata
theascidian-
feedingEraloidae(coffeebeanshelts,e.9.,Erato, (airbreathingsnails).
Althoughthis divisionwas long
Trivia),andthesoftOoralfeeding Ovulidae (ovulids accepted,recentmorphological and molecularstudies
or
egg shells,e.9.,Jenneria, Ovula,Simnia). nowdividethesubclassintotwo maingroups-an infor-
Tonnidae
(tunshells,e.9.,Malea)and relatedfamilies mal paraphyletic
groupoftenreferredto as the "Lower
suchas
Cassididae (helmetshells,e.9.,CassrS) Heterobranchia"1=4ll6nu.tropoda,Heterostropha)and
mainlyfeed
on echinoderms, whereasFicidae(figshells,e.9., the Euthyneura,
whichincludesbothpulmonates andthe
Ficus)are primarily polychaete feeders.Epitoniidae opisthobranchs.
(wentletraps or epitoniids,
e.9.,Epitonium)feed on ThesubclassHeterobranchia characterized by lackinga
cnidarians, whilethefloatingvioletsnailsJanthinidae truectenidium and,usually,a smallto absentosphradi-
(e.9.,Janthina) feedon siphonophores that drifton um,a simplegutwiththeesophagus lackingglands,the
thesurfaceoftheocean.Eulimidae areectoparasites stomachlackinga crystalline stylein all but one group,
on echinoderms, andthesponge{eeding Triphoridae andthe intestine usuallybeingshort.Theradulais highly
(e.9., Triphora)and Cerithiopsidae(e.9., Cerifhropsrs) variablerangingfrom rhipidoglossate to a singlerow of
arehighlydiverse. Therearesomediverse familiesof teethor lostaltogether.
Theshellmaybewell-developed,
small-sized freshwater snailssuchastheHydrobiidae reduced,or absent;the operculum, if present,is horny;
(e.9.,Hydrobia) andseveralrelatedfamilies including the larvalshellis heterostrophic(i.e.,coilsin a different
the Pomatiopsidae (e.9.,Pomatiopsis, Tricula),
and planeto theadultshell).Theheadbearsoneor two pairs
thereare alsoa few terrestrial taxain familiessuch of tentacles,withthe eyesvariously placed;all are her-
as Pomatiasidae (e.9.,Pomatias) andthe otheruirise maphroditic. The nervoussystemis streptoneurous or
mainlysupralittoral Assimineidae. euthyneurous with variousdegreesof concentration of
SUPERORDER NEOGASTROPODA the ganglia;pleuralganglianearpedalor cerebralgan-
The most
derivedhypsogastropod glia,pedalcordsabsent.Mostlybenthic;with marine,
clade.Radularachiglossate
or toxoglossate,with 1-5 teethin eachrow; ante- freshwater andterrestrial
species.
riorsiphonpresent;operculum, if present,chitinous; "LOWER HETEROBRANCHS" Thisinformalgroupin-
osphradium largeand pectinate, lyingnearbaseof cludessomesnailslongthoughtto be "Mesogastrop-
siphon.Thishighlydiversegroupcomprisesmosfly oda,"suchas staircase or sundialshellsArchitectonici-
carnivoroustaxa. dae (e.9.,Architectonica,Philippia)and somegroupsof
The neogastropods comprisemore than 30 llv- small-sizedmarinesnailsincluding Rissoellidae(e.9.,,?is-
ingfamilies of almostentirely marinesnalls,includ- soella),
Omalogyridae (e.9.,Omalogyra), and the fresh-
ing;whelkssuchas Buccinidae (e.9.,Buccinum, waterValvatidae (e.9.,Valvata)andmarinerelatives such
Cantharus, Macron,andtheAsianfreshwatergenus as Cornirostridae.Thesesnailsare superlicially similar
C/ea);Fasciolariidae (tulipshellsand spindteshells, to caenogastropods, but oftenpossesssecondarygills
e.9.,Fasciolaria, Fusinus, Leucozonia, Troschelia); and longcephalictentacles withcephaliceyesset rnthe
Melongenidae (e.9.,Melongena); Nassariidae (dog middleof theirbasesor on theirinnersides.Another
whelksand basketshells,e.9., lVassanr,rs); groupincludedherearetinyinterstitial slugsof thefamily
dove
shellsColumbellidae (e.9.,Anachis,Columbeila, Fhodooidae.
Mitrella,Pyrene,Strombina); harpshellsHarpidae INFRACLASS EUTHYNEURA Includemostof theformer
(e.9.,Harpa);marginshellsMarginetlidae (e.9., opisthobranchsandpulmonates. Theeuthyneuran body
Marginella,Granula); mitersheltsMitridae(e.9.,Mitra, is characterized
by: the shellbeingexternalor internal,
SubcanciI la)andCostellariidae (e.9., Vexil
Ium, Pusia); or lostaltogether;
a heterostrophiclarvalshell;opercu-
rock shellsand thaidsMuricidae(e.9.,Hexaplex, lumhorny,oftenabsentinadult;bodyvariously detorled;
Murex, Phyllonotus, Fterynotus,Acanthina,Morula, headusuallywithoneor two pairsof tentacies, eyeson
Neorapana,Nucella,Purpura,Ihals)andthe related innersidesor on separatestalks;ctenidiaand manfle
coralassociated Coralliophilidae (e.9.,Coralliophila, cavityusuallyreducedor lost;hermaphroditic; euthy-
Latiaxis);
Olividae (oliveshells,e.9.,Agaronia, Otiva); neurouswith variousdegreesof nervoussystemcon-
Olivellidae (e.9.,Olivella);the votutesVolutidae centration.Mostlybenthic;withmarine,freshwater, and
(e.9.,Cymbium,Lyria,Voluta)and nutmegshells terrestrial
soecies.
Cancellariidae (e.9.,Admete, Cancellaria); cone
shellsConidae(e.9.,Conus)andtherelatedTurridae TheEuthyneura is dividedintothreemajorgroupswhich
we treathereas cohorls.
sThe term "heteropod" is an COHORT NUDIPLEURA Includesboththe internal-
old taxonomic name now used infor-
maily for a groupbf planktonic, predatory caenogastropods that shelledPleurobranchidae(e.9.,Berthella,Pleuro-
have a reduced shell or no shell at all.
branchus)and the Nudibranchia
(shell-less
or "true"
PHYLUMMOLLUSCA 467
nudibranchs) whichincludesmanyfamilies, someex- freshwater species)andthe ectoparasitic hTramidelli-

amplesbeingthedoridoidnudibranchs suchasOnchi- dae(e.9.,Odostomia, Pyramidella, Turbonilla,
Amathi-
dorididae(e.9.,Acanthodoris, Corambe), Polyceridae na),allof whichwerepreviously includedin the Opis-
(e.9.,Polycera,Tambja),Aegiretidae (e.9.,Aegr,res), thobranchia. The remaining panpulmonaies include
Chromodorididae (e.9.,Chromodoris), Phyllidiidae all the membersof the group previouslyknownas
(e.9.,Phyll
idia),Dendrodorididae (e.9.,Dendrodoris), Pulmonata, namelythe mainlyintertidal Siphonariidae
Discodorididae (e.9.,Discodoris, Diaulula,Rostanga), (falselimpets:e.9.,Sphonaria,Williamia); two oper-
Dorididae(e.9.,Dorb),Platydorid idae(e.9.,PIatydoris), culatefamilies); the freshwater Glacidorbidae (e.9.,
Hexibranchidae (e.9.,Hexabranch us),Goniodorididae Glacidorbis);the estuarine Amphibolidae (Amphibola,
(e.9.,Okenia), andthe cladobranch nudibranchs in- Salinator);the Hygrophila, including the mainlyfresh-
cludingthe Arminidae (frmina),Proctonotidae (e.9., waterSouthAmericanChilinidae (e.9.,Chilina),and
Janolus), Embletoni idae(e.9., Embletonia), Scyllaei- freshwater Physidae(e.9.,Physa),Planorbidae (e.9.,
dae (e.9.,Scyllaea), and Dendronotidae (e.9.,Den- Bulinus,Planorbis, Ancylus)and Lymnaeidae (e.9.,
dronotus).Alsoincludedare the cladobranch group Lymnaea,Lanx),these latterfamiliesbeing mostly
knownas aeolididoids-including
collectively theAe- snails,but somesuchas LanxandAncylusare lim-
(e.9.,Aeolidia),Flabellinidae
olidiidae (e.g.,Coryphella), pets.Theremaining "pulmonates" arecontained with-
Fionidae (e.9.,Fiona),Facelinidae (e.9.,Hermissenda, in a superorder, Eupulmonata. The best knownand
Phidiana),Tergipedidae (e.9.,Trinchesn),Tethydidae largestgroupof eupulmonates istheorderStylomma-
(e.9.,Melibe),andGlaucidae (e.9.,Glaucus). tophora,comprising thelandsnailsandslugs.Insome
of the shelledforms,the shellis partlyor completely
COHORT EUOPISTHOBRANCHIA Includes six main
enveloped by dorsalmantle.Theireyesareon thetips
groupsthatcouldbetreatedat theordinallevel:(1)the
of longsensorystalksandthereis an anteriorpairof
basalacteonoideans includingActeonidae (banelor
tentacles. Eupulmonates areallterrestrialand arean
bubblesnails,e.9.,Acteon, Pupa, (2)
Rictaxis); several
enormousgroupwithover26,000describedspecies
familiesgroupedas Cephalaspidea, for examplethe
in 104families. Someof thoseincluded arethe land
slugsAglajidae(e.9.,Aglaja,Chelidonura,Navanax), (e.9.,Cornul= Helixl,Cepaea),
snailfamilies Helicidae
Bullidae(bubbleshells,e.9.,BuIIa),Haminoeidae (e.9.,
Achatinidae (e.9.,
Achatina),Bulimulidae (e.9.,BuIimu-
Haminoea), Retusidae (e.9.,Fetusa), andScaphandri-
/us),Haplotrematidae (e.9.,Haplotrema), Orthalicidae
dae(e.9.,Scaphander); (3)theRuncinoidea, contain- (e.9.,Lrguus),Cerionidae (e.9.,Cerion),Oreohelicidae
rngtwo familiesof tinyslugs,llbiidae(e.9.,llbia)and (e.9.,Oreohelrx), (e.9.,Pupilla),Cerastidae
Pupillidae
Runcinidae (e.9.,Runcina); (4)theAplysiomorpha (=
(e.9.,,?hachis),
Succineidae (e.9.,Succinea),andVer-
Anaspidea) or sea hares,including Aplysidae (e.9.,
tiginidae (e.9.,Vertigo), slugfami-
as wellas terrestrial
Aplysia,Dolabella,Stylocheilus); (5)the pelagicptero-
liessuchasArionidae (e.9.,Arion)andLimacidae (e.9.,
pods,comprising two distantgroups,theThecosoma-
Limax).
ta or shelledpteropods, whichincludethefamilies Ca-
voliniidae(e.9.,Clio, Cavolin ia)and Limacinidae, (e.9., Theremaining Eupulmonata includetheordersSystel-
Limacina), andGymnosomata or nakedpteropods in- lommatophora and Ellobiacea. Theformerare slug-
cludingClionidae (e.9.,Clione); and(6)theUmbrachu- like,withoutinternalor external shell;dorsalmantle
lida,composedof the umbrella slugs,Umbraculidae integument formsa keeledor roundednotum;head
(e.9.,Umbraculum) andTylodinidae (e.9.,Tylodina). usuallywith2 pairstentacles,upperonesformingcon-
tractilestalksbeanngeyes,Includedare the mainly
COHORT PANPULMONATA Thishighlydiversegroup
marinefamilyOnchidiidae (e.9.,Onchidella, Onchidi-
is characterized by: variableshellshapeor loss of
um) andtheterrestrialVeronicelIidae(e.9.,VeronicelIa).
shell,minuteor in moderatesize;generally spirally
The Ellobiacea includes the threesuperfamilies; the
coiled,planispiral,
or limpet-shaped; withoutan
usually
mainlysupralittoralhollow-shelled ear snailsEllobioi-
operculum as adults;eyesat basesof sensorystalks;
dea(, e.9.,Ellobium,Melampus,Carychium, Ovatella),
secondary gillspresentin somemembers(e.9.,Pyra-
smallintertidalsnailsor slugsof the Otinoidea (e.9.,
midella,Siphonaria); body detorted;nervoussystem
Otina),and the limpet-like intertidal
Trimusculoidea
highlyconcentrated(euthyneurous); mantlecavity-
(e.9.,Trimusculus).
derivedlungin the derivedgroups,witha contractile
apertureintheEupulmonata; marine(intertidal),
brack- CLASSBIVALVIA (= PELECYPODA, = LAMELLIBRANCHI-
ish,freshwater and amphibious; includesfreshwater ATA) Clams,oysters,mussels,scallops,etc. (Figures
limpetsand the minute,meiofaunal Acochlidioidea 13.1M-Qand13.8).Laterally compressed;shelltypicallyof
(e.9.,Acochlidium,Unela). two valveshingedtogetherdorsallyby elasticligamentand
shellsclosedby adductormuscles
usuallyby shell-teeth;
Otherpanpulmonate groupsinclude: thesap-sucking
derivedfrom mantlemuscles;headrudimentary, without
sea slugsSacoglossa (e.9.,Berthelinia,
Elysia,Oxy-
eyes,tentaclesor radula,but eyesmayoccurelsewhere on
noe,Tridachia,andthe "bivalvegastropods"Juliidae),
body;pairof largelabialpalpspresentcomposedof inner
whichareshelledor shell-less;
thesmallshell-less
and
and outerpartsthat lie againstone another;pairof stato-
sometimes spiculate
acochlidioidslugsthatareoften
cystspresent,associated with pedalganglia,foot typically
interstitial marine(although
andusually therearesome
laterallycompressed, oftenwithouta sole;1 pairof large
(B) ligament
Intestine Pericardial
chamber
Nephridium
Ventricle
Nerve Right atrium
Efferent Suprabranchial
branchial canal
vessel
Mantle
Afferent vessel
branchial
vessel demibranch
of ctenidium
(c) DORSAL
Inner
Hinge teeth demibranch
Ventricle of ctenidium
Ctenidium
Posterior aorta Food groove
Anterior foot Rectum Gonad
retlactor
rnuscle Posterior foot
retractor muscle Mantle cavity
Mouth Exhalant
siphon
Labial palp
POSTERIOR
ANTERIOR (E) Cut end of right
lnhalant siphon labial palps Mouth
Cut edge of
Anterior adductor Posterior right ctenidium Cerebral ganglia
muscle adductor muscle
Plicate Cerebrovisceral
membtanes connective
VENTRAL
Labial palp
(D) Digestive
Foot
Byssalthreads
Right urogenital pore
Ventricle
Cerebrovisceralconnective
Posterior
foot retractor Right visceral ganglion
Recfum Left ctenidium
Anus
Inhalant area
Posterior adductor
Exhalant siphon
Inhalant siphon
Mouth
ganglion
Anterior aorta Esophagus
Pedal ganglion I7 Cerebral ganglion
Stomach
Right ahium Nephridium
Cerebropedalconnective
Right pedal ganglion
Crystalline style sac retractor muscle
Posterior adductor muscle Posterior end of kidneY
Cerebrovisceralconnective
Right visceral ganglion

PHYLUMMOLLUSCA 469
{ Figure 13.8 General anatomy of bivalves. (A) Iresus, a mainlyfor housingsymbiotic Gutreducedor
bacteria.
deep-burrowing eulamellibranch(Mactridae),with a dig- absent.(e.9.,Solemyidae,
So/emya).
ging foot and long, fused siphons. (B) A typical eulamel-
libranch (cross section). (C) The eulamellibranch Mercenaria SUPERORDER NUCULANIFORMII Mantlefusedoos-
(Veneridae), with the left shell valve and mantle removed. teriorly,
withsiphons,inhalantwaterentersposteriorly;
(D) lnternal anatomy of Mercenaria. The visceral mass is shellswithoutnacreand gillfilamentsalongctenidial
opened up, the foot is dissected, and most of the gills are cut axisalternate.Severalmainlydeep-seafamiliesrn-
away. (E) The commbn mussel, Mytilus (Mytilidae), seen from cluding (e.9.,
Nuculanidae (e.9.,
NucuIana),Malletiidae
the right side after removal of the right shell valve and mantle. Malletia)
and Sareptidae(e.9.,Yoldia).
(F) Mytilus, with the visceral mass opened up, the foot dis-
sected, and most of the gills cut away. SUBCLASS AUTOBRANCHIA (= AUTOLAMELLIBRAN-
CHIATA)Pairedctenidia, withverylongfilaments folded
backon themselves so thateachrow of filaments forms
bipeciinatectenidia;largemantlecavitysurroundsanimal; two lamellae;adjacentfilaments usuallyattachedto one
mantlemay be variously fused,sometimesformingexten- anotherby ciliarytufts(filibranch
condition),or by tissue
sions(siphons);1 pairnephridia; nervoussystemsimple, bridges(eulamellibranch Thegreatlyenlarged
condition).
typicallycomposedof cerebropleural, pedaland visceral ctenidiaare usedin combination with the two pairsof
ganglia labialpalpsin ciliaryfeeding;ctenidialsurfacescapture
Bivalves are marineor freshwater molluscs, primarily
mi- waterborneoarticlesand transferthem to the labial
palpswherethe capturedebrisis sortedand potential
crophagousor suspension feeders.The classincludes
about9,200livingspeciesrepresented at alldepthsand in foodparticlesroutedto the mouth.
all marineenvironments. Bivalveclassificationhas beenin COHORT PTERIOMORPHIA (=FILIBMNGHIA) Ctenid-
a stateof turmoiloverthe past50 yearsand continuesto ia with outerfold not connecteddorsallyto visceral
be unsettled. Highertaxahavebeendelimited on the basis mass,with freefilamentsor with adjacentfilaments
of shellcharacters (e.9.,hingeanatomy,positionof muscle attachedby ciliarytufts(filibranchcondition);
shellara-
scars),or, in otherclassifications,internalorgananatomy goniticor calcitic,sometimes nacreous;mantlemar-
(e.9.,ctenidia,stomach)havebeenused.However, begin- gin unfused,withweaklydifferentiated incurrentand
ningwith the work of Giribetand Wheeler(2002),bivalve excurrent apertures or siphons; footwelldeveloped or
taxonomyhasbecomemorestableas bothmolecules and extremely reduced;usually attachedby byssalthreads
morphologyhavebeencombinedwiththe fossilrecordto or cementedto substratum(orsecondarily free).These
understand the relationshipsof theclass. primitivelamellibranchs includeseveraldivergentan-
SUBCLASS PRoTOBRANCHIA IncludestheformerPal- cientlineages separated as orders.
aeotaxodonta in part.Ctenidiaare2 pairsof simple,un- ORDER MYTILIDAThetruemussels, Mytilidae(e.9.,
folded,bipectinate,plate-likeleafletssuspendedin the Adula,Brachidontes,Lithophaga,Modiolus,Mytilus).
mantlecavity.Thectenidiaare mainlyrespiratory struc-
tures,whilethe labialpalpsarethe primaryfoodcollect- ORDER ARCIDAThearkshellsArcidae(e.9.,Anada-
ing organs.Thesearethe mostprimitivelivingbivalves, ra, Arca,Barbatia),
and dog cocklesGlycymerididae
comprising two superorders. (e.9.,Glycymeris).
SUPERORDER NUCULIFORMII (= OPPONO-BRAN- ORDER OSTREIDAThetrueoystersOstreidae(true

CHIA) Mantleopen,withinhalant waterentering
ante- oysters,e.9.,Crassostrea,
Ostrea).
riorly;shellswithnacreandgillfrlaments
alongctenidial ORDERMALLDIDA (= PTERIOIDA, PTERIIDA)Pearl
axisarranged oppositeoneanother; footlongitudinally oystersand their relativesPteriidae(e.9.,Pinctada,
groovedandwitha plantarsole,withoutbyssalgland; Pteria),
hammeroystersMalleidae (e.9.,Mal/eus),
and
nervoussystemprimitive, oftenwithincomplete union penshellsPinnidae (e.9.,Atina, Pinna).
of cerebralandpleuralganglia. Twoorders.
(e.9.,Lima),
LIMOIDAFileshellsLimidae
ORDER
oRDERNUcULIDAShellaragonitic, interior
nacreous
or porcelaneous; periostracumsmooth;shellvalves oRDERPECTINIDINA Scallops.Pectinidae (e.9.,
equalandtaxodont(i.e.,thevalveshavea rowof simi- Chlamys,Lyropecten,Pecten),thornyoystersSpon-
jingleshellsAnomiidae
dylidae(e.9.,Spondylus), (e.9.,
shortteethalongthe hingemargins);
lar interlocking
adductormusclesequalin size;withlargelabialpalps Anomia,Pododesmus).
extendedas proboscidesusedfor food collection; COHoRT HETERocoNcHIA Thiscladeencomoass-
marine(particularly
ctenidiasmall,for respiration; in esthe Paleoheterodonta previously
and Heterodonta,
the deepsea),mainlyinfaunal (e.9.,Nu-
detritivores. treatedas separatehighergroupsbut shownto be
culidae,Nucula). phylogenies.
sistergroupsin recentmolecular
ORDER SOLEMYIDA (=CRYPTODONTA) Shellvalves MEGAORDER PALAEOHETERODONTA SheIIaTa-
thin,elongate,and equalin size;uncalcified
along gonitic,pearlyinternally;
periostracum
usuallywell
outeredges,withouthingeteeth;anterioradductor developed; valvesusuallyequal,with few hinge
musclelargerthanposterior
one;ctenidialarge,used teeth;elongatelateralteeth(whenpresent)are not
470 Chapter Thirteen Water movement
Sediment surface
separatedfromthelargecardinalteeth;
usually
dimy-
arian;mantleopensbroadlyventrally,
mostlyunfused
posteriorly
but withexhalant
and inhalant
apertures.
About 1,200 speciesof marineand freshwater Mantle cavity
clams.Includestwoverydistinctgroupsclassified
as Gonad
orders. Nephridium
Mantle (ciliated)
ORDERTRtcO\iltDA "The relictualmarinebroach Digestive gland
shellsfl-rigoniidae),
with onlya few livingspeciesof Retractormuscle
Neotrigonia in Australia.
Stomach
ORDER UNIONtDAEntirely freshwater, including the
freshwater Shell
clams(ormussels), e.9.,Unionidae (e.9.,
Anodonta, Unio), Margaritiferi
dae (e.9.,Margaritifera), Pleural ganglion
(e.9.,Hyridella).
and Hyriidae
Cerebral ganglion
MEGAORDER HETERODONTA Two main groups,
rankedas superorders, ganglion
arerecognized-theArchiheter-
odonta(witha singlelivingorder)andthe Euheterodonta
(withfourlivingorders).
Pedal ganglion
SUPERORDER
ARCHIHETERODONTA
Foot
ORDER CARD|T|DA Thisgroupof primitive heter- Adhesive knob
odontsis represented by the familiesCrassatellidae
(e.9., Crassafe//a),
Carditidae(e.9.,Cardita)andAstart-
;A^^ I^ n ^+^4^\ Captacula
luoE ^
\v.v,r nJLd.t LYJ,
Figure 13.9 General anatomy of a scaphopod.

SUPERORDER
EUHETERODONTA
ORDER LUCINIDAIncludes the families
Luncinidae
(e.9.,Lucina,Codakia), a groupwith symbioticbac- welldeveloped; mantleusuallyfusedventrally, with
teriain theirgillsand an anterior
watercurrent,and pedalgape,and posteriorly
anteroventral withventral
Thysiridae. incurrentand dorsalexcurrentaperturesor siphons;
ctenidiaeulamellibranchiate
or septibranchiate(modi-
ORDER CARDIIDA (=VENERIDA) Usuallythick-valved, fiedas a muscularpumpinghorizontal septum).This
equi-valved, and isomyarian, with posteriorsiphons.
ancientand verydiversegroupof marinebivalves
lncludes:cocklesandtheirkin,Cardiidae (e.9.,Cardj-
includesabout20 livingfamilies, includingthe rare
um, CIinocardium, Laevicardium, Trachycardium, and
deepwaterPholadomyidae (e.9.,Pholadomya) and
the giantclamse.9.,Tridacna); surfclams,Mactridae
the aberrantwateringpot shellsClavagellidae (e.g.,
(e.9.,Mactrd; solens,Solenidae (e.9,,Ensis,So/en);
Brechites),
as wellas Pandoridae,Poromyidae, Cus-
tellins, (e.9.,Flori
Tellinidae metis, Macoma,Tellina);se- pidariidae,
Laternulidae,
Thraciidae, Cleidothaeridae,
melids,Semelidae (e.9.,Leptomya,Semele); wedge
Myochamidae, andPeriplomatidae.
shells,Donacidae (e.9.,Donax);Venusclams,Veneri-
dae (e.9.,Chione,Dosinia,Pitar,Protothaca,Tivela); CLASSSCAPHOPODA Tusk shells(Figures 13.1Land
freshwater peaclams,Sphaeriidae (e.9.,Sphaerium, 13.9).Shellof onepiece,tubular,usuatly tapering, openat
Pisidium); theestuarine-to-freshwaterCyrenidae bothends;headrudimentary,
(e.g., projecting from largeraper-
Corbicula,Bafissa);and zebramussels,Dreissenidae ture;mantlecavitylarge,extending alongentireventralsur-
(e.9.,Drerbsena).Thelattertwofamilies containimpor- face;withoutctenidia or eyes;withradula,Iong,snout-like
tantinvasive soecies. "proboscis," andpairedclustersof long,thincontractile ten-
tacleswithclubbedends(captacula) that serve to capture
ORDERPHOLADIDA (= MYIDA)Thin-shelled bur- and manipulate minuteprey;hearlabsent;foot somewhat
rowingformswithwell-developed siphons.Includes:
cylindrical,with epipodium-likefringe.Over500 species
soft-shell
clams,Myidae(e.9.,Mya);rockborers or pld- of marine,benthicmolluscsin fourteenfamiliesand two
docks,Pholadidae (e.9.,Barnea,Martesia,Pholas);
orders.
shipworms,Teredinidae (e.9.,Bankia,Teredo);and
basketclams,Corbulidae (e.9.,Corbula).Themono- ORDER DENTALilDAShellregularly tapering,paired
phylyof thisorderis uncertain. digestive glandwith a muscular foot terminating in
epipodiallobesand a cone-shaped process.Several
OHDER POROMYATA (=ANOMALODESMATA) SheIIs familiesincludingDentaliidae (e.9.,Dentalium,Fusti-
equivalved, aragonitic,of 2-3 layers,innermostcon- aria) andLaevidentaliidae(e.9., Laevidentalium).
sistingof sheetnacre;periostracum oftenincorpo-
ratesgranulations; with O-l hingeteeth;generally ORDER GADILIDAShellregularly taperingor bulbous
isomyarian, rarelyamyarian; posteriorsiphonsusually with maximumshelldiameterapproximately nearer
Siphuncle PHYLUMMOLLUSCA 471
SUBCLASS NEOCEPHALOPODA Includesone fossil

groupin additionto the coleoids.Theshellis reduced
in most(andin alllivingtaxa).
andinternal
COHORT COLEOIDIA (= DIBRANCHIATA) Octopuses,
squids,andtheir kin,Shell reduced and internalor ab-
sent;headandfootunitedintoa commonanteriorstruc-
ture bearingB or 10 prehensile appendages (armsand
teniacles)bearingsuckersand,often,cirri,onearmusu-
allymodifiedin malefor copulation;7 toothrowsin radu-
la;withchitinousbeak;funnela singleclosedtube;I pair
1 pairnephridia;
ctenidia("dibranchiate"); eyescomplex,
withlensandoftenwithcornea;neryoussysiemwellde-
velopedand concentrated intoa brain;with a complex
statocyst;withchromatophores and inksac.
SUPERORDER OCTOPODIFORMES MEMbETS Of this
group,whichincludes theoctopuses andvampiresquid,
do not havethe headdistinctlyseparatedfromthe rest
Figure 13.1 0 The anatomy ot Nautilus (sagittal
of the body;have8 arms,with2 additional fila-
retractile
mentsin thevampiresquids;lateralfinson the bodyare
nrocont nr ahqont
glandandfootwitha
centerof shell.Singledigestive
disk
terminal surrounded by papilla.
epipodial Families ORDER OCTOPODA Octopuses. Bodyshort,round,
includePulsellidae(e.9.,Pulsellum,
Annulipulsellum) usuallywithout fins; shell
internal vestigialor absent;8
andGadilidae(e.9.,Cadulus, Gadila). similararmsjoinedby web of skin(interbrachial web);
Nautilus, squids,cuttlefish, and suckerswith narrowstalks,mostare benthic,About
CLASSCEPHALOPODA
(Figures13,1H-K, 13.10, 13.1 1 13.12, 13.17 200species, intwo groups:thelncirrata, includingthe
octopuses , ,
shell,usuallyreduced benthicoctopusesand somepelagictaxathat lack
and 13.22).Withlinearly chambered
(e.9.,
finsandciniwithexamples beingOctopodidae
or lostin livingtaxa;if externalshellpresent(Nautlus), ani-
(f,rgonaufa),
Octopus)and Argonaulidae the paper
malinhabitslast(youngest) chamber, witha filament of living
througholderchambers; nautilus;and Cirrata,whichare mainlypelagicdeep
tissue(thesiphuncle) extending
seacephalopods withfinsandcirri,suchas Cirroteu-
circulatorysystemlargelyclosed;headwith large,com- (e.9.,Opis-
(e.
thidae 9., Cirroteuthis),Opisthoteuthidae
plexeyesandcircleof prehensile armsor tentacles around
thoteuthis),and Stauroteuthidae(e.9., S/aurofeufhr.s).
mouth;with radulaand beak;1-2 pairsctenidia,and 1-2
pairscomplexnephridia; mantleformslargeventralmantle ORDERVAMPYROMORPHA The vampiresquid.
cavitycontaining 1-2 pairsof ctenidia;with muscularfun- Bodyplump,with 1 pairfins;shellreducedto thin,
nel(thesiphon)throughwhichwateris forced,providing jet leaf-shaped, transparent
uncalcified, vestige;4 pairs
propulsion; sometentacles of malemodified for copulation; eoual-sizedarms,eachwith one row of unstalked
benthicor pelagic,marine;about700 livingspecies. distalsuckersand two rowsof cirri;armsjoinedby
Includesmanyfossil extensiveweb of skin(interbrachialmembrane); fifth
SUBCLASS PALGEPHALOPoDA
pairof armsrepresented by 2 tendril-like,retractable
taxa,all with externalshells,as wellas the livingpearly
lacking;
hectocotylus
filaments; radulawelldeveloped;
nautilus.
ink sac degenerate;mostlydeep water.One living
COHORTNAUTILIDIA (= TETRABRANCHIATA) The species,Vampyroteuthis infernalis,that lives in the
pearlynautilus.Shellexternal,many-chambered, coiled oxygenminimumzoneof the deepsea.
in one plane,exteriorporcelaneous, interiornacreous
(= DECAPODA)
tentacles(4 SUPERORDER DECAPODIFORMES
(pearly);headwithmany(80-90)suckerless
and protected Membersof thisgroup,whichincludesthe squidand
modifiedas spadixin mde for copulation
cuttlefish,havethe headdistinctlyseparatedfrom the
by a fleshyhood);13 tooth rowsin radula;beakof chi-
funnelof 2 separatefolds;2 restof the body;with 8 armsand 2 retractile(intopits)
tin and calciumcarbonate;
2 pairsnephridia;
pairsctenidia("tetrabranchiale"): eyes tentacleswith suckersonlyon expandedtips; suckers
withwidebases,sometimes withspinesor hooks;lateral
likea pinholecamera,withoutcorneaor lens;nervous
concentrated intoa brain, finson body.Theinternalshellis large(asinthecuttlefish
systemwithanteriorelements gladius,or lost.
simple; withoutchromato- reducedto an uncalcified
or in Spirula),
optic lobes large;statocyst
phoresor ink sac. Fossilrecordrich,but represented oRDERSPIRULIDA The only livingspeciesis the
todayby a singleorder(Nautilida)and singlegenus,the ram'shorn,Spirulaspirula(Spirulidae),
a smalldeep
chamberedor pearlynautilus(Nautilus), withfiveor six chambered
seasquidwitha coiled,internal, shell.
Indo-Pacific species(although a second,controversial
genus,Allonautilus,hasalsobeenproposed). ORDER SEPIIDACuttlefish.Bodyshort,dorsoventral-
calcareous,
withlateralfins;shellinternal,
lv flattened,
straightor slightlycurved,chambered; or shellhorny, structures, most notably eyes and tentacles; statocysts
or absent;8 shortarms,and 2 longtentacles; suck- may be located in the foot region and chemosensory
erslackhooks.lncludes theshell-lessSepiolidae(e.9,, structures can also be present.
,?ossra, Sepiola),
the Sepiidae (e.9.,Sepa)withan in- The visceral mass is covered by a thick epidermal
ternalcalcareous shell,the cuttlebone,and ldiosepj- sheet of skin called the rnantle (also known ai the pal-
idae(e.9.,/drr:seprS),
beingtinysquidsthatlivein sea- lium), which is sometimes covered in cuticle and plavs
grassto whichtheyaltachwitha specialsucker.Their
a critical role in the organization of the body. It secret-es
shellis reducedto a hornygladius.
the hard calcareous skeleton, either as minute sclerites,
ORDER MYOPSIDASquidswiththeeyecoveredwith or plates, that are embedded in the body wall or as a
a corneaand havinga welldeveloped gladius.Body solid internal or external shell. Ventrally the body usu-
elongate, tubular, withlateralfins.
Loliginidae (e.9.,
Lo- ally bears a large, muscular foot, which typically has a
ligo, Doryteuthis). creeping sole.
ORDER OEGOPSIDA Includes the majorityof squids Surrounding or posterior to the visceral mass is a
(andtheformerTeuthoida in part);theeyelacksa cor- cavity-a space between the visceral mass and folds
neaandtheshellis a gladius.Bodyelongate, tubular, of the mantle itself. This mantle cavity (also known
with lateralfins;suckersoftenwith hooks.Someof as the pallial cavity) often houses the gills (the origi-
the manyfamiliesin this groupincludeArchiteuthi- nal molluscan gills are known as ctenidia), along with
dae (frchiteufhls),Bathyteuthidae (e.9.,Bathyteu - the openings of the gut, nephridial, and reproductive
fhls;sometimes treatedas a separateorder),Chiro- systems, and, in addition, special patches of chemosen-
teuthidae(e.9..Chiroteufhis), Ommastrephidae (e.9., sory epithelium in many groups, notably the osphra-
Ommastrephes, Dosrdiscus,lllex),Gonatidae(e.9., dia. In aquatic forms, water is circulated through this
Gonatus),Histoteuth idae(e.9.,Hi stioteuthis),Lycoteu- cavlty, passing over the ctenidia, excretory pores, anus/
thidae(e.9.,Lycoteuthis). andOctopoteuthidae (e.9., and other structures.
Octopoteuthis\. Molluscs have a complete, or through-gut that is re-
gionally specialized. The buccal region of the foregut
typically bears a uniquely molluscan structure, the rad-
The Molluscan Body Plan ula, which is a toothed, rasping, tongue-like strap used
in feeding. It is located on a muscular odontophore that
Mollusca is one of the most morphologically diverse moves the radula through its feeding motions. The cir-
phyla in the animal kingdom. Molluscs range in size culatory system usually includes a heart in a pericardial
from microscopic solenogasters,bivalves, snails, and cavity and a few large vessels that empty into or drain
slugs, to whelks attaining 70 cm in length, giant clams hemocoelic spaces.The excretory system consists of one
(Cardiidae) over a 1 m in length, and giant squids (Ar- or more pairs of metanephridial kidneys (here simply
chiteuthis) reaching at least L3 m in overall length (body referred to as nephridia), with openings (nephrostomes)
plus tentacles). The giant Pacific octopus (Octopus to the pericardium via renopericardial canals and to
dofleini) commonly attains an arm span of 3-5 m and the mantle cavity via the n"phridiopore. The nervous
a weight of over 40 kg. It is the largest living octopus, system typically includes a dorsal cerebral ganglion, a
and one particularly large specimen was estimated to nerve ring encircling the buccal area or esophagus, and
have an arm span of nearly 10 m and a weight of over two pairs of longitudinal nerve cords arise from paired
250 kg! Despite their differences, giant squids, cow- pleural ganglia and connecting with the visceral ganglia
ries, garden slugs, eight-plated chitons, and wormlike more posteriorly in the body. Other anterior paired gan-
aplacophorans are all closely related and share a com- glia @uccal and labial) maybe present. Pedal ganglia lie
mon body plan (Box 13A). In fact, the myriad ways in in the foot and may give off pedal nerve cords.
which evolution has shaped the basic molluscan body Gametes are produced by the gonad in the viscer-
plan provide some of the best lessons in homology and al mass and fertilization may be external or internal.
adaptive radiation in the animal kingdom. Development is typically protostomous, with spiral
Molluscs are bilaterally symmetrical, coelomate pro- cleavage and a trochophore larval stage. There is also a
tostomes, but the coelom generally exists only as small secondary larval form unique to gastropod and bivalve
vestiges around the heart (the pericardial chamber), molluscs called the veliger.
the gonads, and parts of the nephridia (kidneys). The Although this general summary describes the basic
principal body cavity is a hemocoel composed of sever- body plan of most molluscs, notable modifications
al large sinuses of the open circulatory system, except occur and are discussed throughout this chapter. The
in some cephalopods thathave a largely closed system. eight classesare characterized above (seeclassification)
In general, the body comprises three distinguishable and are briefly summarized below.
regions: a head, foot, and centrally concentrated viscer- Some of the most bizarre molluscs are the " apla-
al mass, but the configuration differs in different class- cophorans"-Solenogastres and Caudofoveata (Figure
es (Figure 13.13).The head may bear various sensory 13.1C and 13.2).Members of these groups are wormlike
PHYLUMMOLLUSCA 473
(A) Posterior Posterior (c) \-J Funnel
Cartilage
Ink sac
Esophagus
Digestive
gland Anus
:
Funnel Ctenidium
Anterior retractor
surface muscle Intestine
Left Branchial
side NephridioPore
heart
Systemic heart
OlfactorY Nephridium
crest Stomach
TiP
of pen
Eye
Digestive cecum
Anterior
squid'
Figure 13.1 1 The anatomy of a squids' (A-C) The common
morphology (anterior view)' (B) External morphology
titigo.@) External
of a male' The mantle is dissected
lposterior view). (C) Internal anatomy
o p e n a n d p u l | e d a s i d e . ( D ) T h e g i a n t s q u i d ( A r c h i t e u t h i s d u x ) n e tte d o ffth e
coast of New Zealand in 1997'
Polyplacophorans,or chitons,are oval molluscsthat

Ueareifnt (sevenin Paleoloricata)seParatearticulat-
ing shell plates on their backs (Figures13'18 and 13'4)'
Thleyrange in length from about 7 mm to over 35 cm'
Thesemaiine animalsare inhabitantsof deep seato in-
tertidal regionsaround the world, at all latitudes'
Monoplacophoransare limpet-like molluscs with a
single cap-shapedshell ranging from about 1 mm to
aUo'ut+ cm in length (Figures13.14 and 13'3)' Most
live in the deep sei, some at great depths (> 2,000m)'
Their most notable feature is the repetitive arrange-
ment of gills, gonads, and nephridia, a condition that
hasled somebiologiststo speculatethat they must rep-
resenta link to someancientsegmentedancestorof the
Mollusca (an idea no longer deemedreasonable)'
Gastropodsare by far the largest grouP of molluscs
and include some of the best-studied species(Figures
and typically small, and either burrow in sediment
13.1D-G, 13.5,73.6,and73.7).This classincludesthe
their entire lives on the common snailsand slugsin all marine and many fresh-
lCauiofoveutu; ot may spend
branches of various cnidarians (Solenogastres) such as water habitats, and they are the only molluscan class
gorgonians upon which they feed. Caudofoveata lack a to have successfullyinvaded terrestrial environments'
Ioot-,but a reduced one is present in the Solenogastres/ They arethe only molluscsthat undergo torsion during
and neither group has a solid shell. Aplacophorans also early development,a processinvolvinga 90-180'rota-
have no distinct head , eyes, or tentacles' They were tra- tionof the visceral miss relative to the foot (for details
ditionally considered primitive molluscs that evolved seesectionon torsionbelow)'
before the appearance of solid shells, but some molecu-
Bivalves include the clams, oysters,mussels,and
lar and derrilopê.ttal data suggest they may actually
their kin (Figures13.1M-Q and 13.8)'They possesstwo
be highly derived forms that have lost the shell and separateshells,calledvalves. The smallestbivalves are
have iecondarily acquired a simple body morphology'
membersof the marine family Condylocardiidaesome
474 Ch a o te r T h i rre e n
of which are about 1 mm in length; the largest are giant of development of each of these muscle layers differs
tropical clams (Tridacna), one species of which (7. gigas) among the classes(e.g., in solenogasters the diagonal
may weigh over 400 kg! Bivalves inhabit all marine en- layers are frequently absent).
vironments and many freshwater habitats.
Scaphopods, the tusk shells, live in marine surface The Mantle and Mantle Cavity
sediments at various depths. Their distinctive single, The significance of the mantle cavity and its impor-
tubular uncoiled shells are open at both ends and tance in the evolutionary success of molluscs has al-
range from a few millimeters to about 15 cm in length ready been alluded to. Here we offer a brief summary
(Figures 13.1L and 13.9). of the nature of the mantle cavity, and its disposition
The cephalopods are among the most highly modi- in each of the major groups of molluscs.
fied molluscs and include the pearly nautilus, squids, The mantle, as the name implies, is a sheet-like
cuttlefish, octopuses, and a host of extinct forms, in- organ that forms the dorsal body wall, and in most
cluding the ammonites (Figures 13.1I-K, 1.3.10,13.71, molluscs it grows during development to envelop the
13.12, 13.17, and 13.22). This group includes the larg- molluscan body and at its edge there are one or two
est of all living invertebrates, the giant squid, with folds that contain muscle layers and hemocoelic chan-
body and tentacle lengths around 13 m. Among living nels (Figure 13.15C). The outward growth creates a
cephalopods, only the nautilus has retained an exter- space lying between the mantle fold(s) and the body
nal shell. The cephalopods differ markedly from other proper. This space, the mantle cavity, may be in the
molluscs in several ways. For example, they have a form of a groove surrounding the foot or a primitive-
spacious body cavity that includes the pericardium, ly posterior chamber through which water is passed
gonadal cavity, nephriopericardial connections, and by ciliary or, in more derived taxa,by muscular ac-
gonoducts, all of which form an interconnected system tion. Generally, the mantle cavity houses the respi-
representing a highly modified but true coelom. In ad- ratory surface (usually the ctenidia or other gill-like
dition, unlike all other molluscs, many coleoid cepha- structures), and receives the fecal material discharged
lopods have a functionally closed circulatory rysi"tr,. from the anus and excretory waste from the kidney.
The nervous system of cephalopods is the most sophis- Gametes are also primitively discharged into the man-
ticated of all invertebrates, with unparalleled learning tle cavity. Incoming water provides a source of oxygen
and memory abilities. Most of these modifications are for respiration, a means of flushing waste and, in some
associated with the adoption of an active predatory instances, also carries food for suspension feeding.
lifestyle by these remarkable creatures. The mantle cavity of chitons is a groove surround-
ing the foot (Figures 13.44 and 13.13A,8). Water enters
The Body Wall
the groove from the front and sides, passing medial-
The body wall of molluscs typically comprises three ly over the ctenidia and then posteriorly between the
main layers: the cuticle (when present), epidermis, ctenidia and the foot. After passin$ over the gonopores
and muscles (Figure 13.15A). The cuticle is composed and nephridiopores, water exits the back end of the
largely of various amino acids and sclerotized proteins groove and carries away fecal material from the poste-
(called conchin), but it apparently does not contain riorly located anus.
chitin (except in the aplacophorans). The epidermis The aplacophorans have a small mantle cav-
is usually a single layer of cuboidal to columnar cells, ity, with either a pair of ctenidia (Caudofoveata) or
which are ciliated on much of the body. Many of the lamella-like folds or papillae on the mantle cavity wall
epidermal cells participate in secretion of the cuticle. (Solenogastres).The paired coelomoducts and the anus
Other kinds of secretory gland cells can also be pres- also open into the mantle cavity
ent, some of which secretemucus and these canbe very The single mantle cavity of gastropods originates
abundant on external surfaces such as the sole of the during development as a posteriorly located chamber.
foot. Other specialized epidermal cells occur on the As development proceeds, however, most gastropods
dorsal body wall, or mantle. Many of these cells con- undergo a 180orotation of the visceral mass and shell to
stifute the molluscan shell glands, which produce the bring the mantle cavity forward, over the head (Figures
calcareous sclerites or shells characteristic of this phy- 13.5,73.6, and 13.13C) (see section on torsion that fol-
lum. Still other epidermal cells are sensory receptors. lows). The different orientation does not affect the water
The epidermis and outermost muscle layer are often flow, which still passes through this chamber through
separated by a basement membrane and occasionally the ctenidia, and then past the anus, gonopores, and
a dermis-like layer. nephridiopores. A great many secondary modifications
The body wall usually includes three distinct lay- on this plan have evolved in the Gastropoda, includ-
ers of smooth muscle fibers: an outer circular layer, a ing rerouting of current patterns; Ioss or modification
middle diagonal layer, and an inner longitudinal layer. of associated structures such as the gills, hypobranchial
The diagonal muscles are often in two groups with fi- glands and sensory organs; and even "detorsion," as
bers running at right angles to each other. The degree discussed in later sections of this chapter.
PHYLUMMOLLUSCA 475
Figure 13.1 2. The anatomy of Octopus. Dorsal mantle cavitY

(A) General external anatomy. (B) Right- Digestive gland
Mantle muscles
side view of the internal anatomY. Poison gland
(C) Arm and sucker (cross section). Intestinal Cartilagenous cranium
(D) Tip of the hectocotylus arm. (E) The hemocoel
diminutive Eastern Pacific Paroctopus
digueti well camouflaged on a sand Cephalic vein
bottom. (F) The troPical Pacific Stomach Buccal mass
Octopus chierchiae. (G) The remark- Shell rudiment Brachial
able lndo-West Pacific Abdoqus Ceca nerve
horridus.
Systemicheart
Nephridium
Branchial heart
Ctenidium Muscular septum of Anus

the ventral rnantle cavitY
Copulatory
(c) Brachial collector vein (D)
Epidermis
Dermus
Verticalmuscle
Longitudinal muscle
Connective tissue
Brachial artery
Nerve cord
Suction cup
Spermatophore
groove
Figure 13.13 Modifications of the shell, foot, gut, (A)

ctenidia, and mantle cavity in five classes of mol-
luscs. (A,B) Lateral and cross sections of a chiton Stomach
(Polyplacophora). (C) Side view of a snail (Gastropoda).
(D,E) Cutaway side view and cross section of a clam Head
(Bivalvia). (F) Lateral view of a tusk shell (Scaphopoda). Anus
(G) Lateral view of a squid (Cephalopoda). ln cephalopods Foot
the foot is modified fo form_.thefunnel (= siphon) and at Digestive Intestine
Nephridiopore
least parts of the arms. gland
Valve
(B)
Mantle Mantle cavity
lntestine
Ctenidium
Ctenidium
Digestive
Cephalic
gland
tentacle
Stomach
trmu Ctenidia
(F) Shell aperture

I
Ctenidium Mantlecavity
Stornach
Shell (pen) Manfle
cavity
Mantle
Ctenidium
Lrtestine
Mantle
cavity
Bivalves possess a greatLy enlarged mantle cav-

ity that surrounds both sides of the foot and visceral well as accomplishinggas exchange.The water flow
mass (Figures 13.8 and 13.13D,8).The mantle lines the then sweepsacrossthe gonoporesand nephridiopores,
laterally placed shells, and the folds making up the and lastly past the anus asit exits through the exhalant
mantle edges are often fused in various ways posteri- siphon.
orly to form inhalant and exhalant siphons, through Scaphopodshave a tapered,tubular shell (Figures
which water enters and leaves the mantle cavity. The 13.9and 13.13F). Water entersand leavesthe elongate
water passes over and through the ctenidia that, in au- mantle cavity through the small opening in the top of
tobranch bivalves, extract suspended food material as the shell and flushesover the mantle surface,which. in
PHYLUMMOLLUSCA 477
the absence of ctenidia, is the site of gas exchange. The on a protein matrix in layers, and often covered by
anus, nephridiopores, and gonopores also empty into a thin organic surfacecoating called a periostracum
the mantle cavity. (calledthe hypostracum in chitons) (Figure 13.15).The
While no detailed studies of the functioning of the periostracumis composedof a type of conchin (largely
monoplacophoran mantle cavity have been made, ob- quinone-tannedproteins) similar to that found in the
servations of the filst living specimensinl9TT revealed epidermal cuticle. The calcium layers have four crys-
that the gills vibrated, apparently circulating water tal types: prismatic, spherulitic, laminar, and crossed
through the mantle groove. It was also noted that shell structures. All incorporate conchin onto which the
movement was accompanied by an acceleration of gill calcareouscrystals precipitate. The majority of living
beating. Vibrating gills are unknown in other molluscs molluscshave an outer prismatic layer and an inner
where ciliary action, sometimes assisted by muscular porcelaneous,crossedlayer. In monoplacophorans,
contractions, moves water through the mantle cavity. cephalopodsand in somegastropodsand bivalves, an
The anus, nephridiopores, and gonopores also open iridescent,nacre (laminar) layer replacesthe layer of
into the mantle cavity. crossedcrystals.Shellsare often made up of multiple
With the exception of the monoplacophorans, in layers of different crystal gpes.
all of the above cases, water is moved through the Molluscs are noted for their wonderfully intricate
mantle cavity by the action of long lateral cilia on the and often flamboyant shell color patterns and sculp-
ctenidia. But in cephalopods the ctenidial gills are not turing (Figure 13.16),but very little is known about the
ciliated. Instead, in Nautilus a ventilatory current is evolutionary origins and functions of thesefeatures.
passed through the mantle cavity by the unduatory Somemolluscan pigments are metabolic by-products,
movements of two muscular flaps associated with the and thus shell colors might largely representstrategi-
funnel lobes. In the coleoid cephalopods, however, cally depositedfood residues,while othersappearto
well-developed, highly innervated mantle muscles have no relationship to diet. Molluscan shell pigments
perform this function through the regular pulsation include such compounds as pyrroles and porphyrins.
of the mantle wall. The exposed, fleshy body surface Melanins are common in the integument (cuticle and
of squids and octopuses is, in fact, the mantle itself epidermis),the eyes,and internal organs,but they are
(Figures 1,3.1.1.,1.3.72,and 13.13G).Unconstrained by rare in shells.
an external shell, the mantle of these molluscs expands Someshell sculpture patterns are correlatedwith
and contracts to draw water into the mantle cavity and specificbehaviorsor habitats.For example,shellswith
then forces it out through the narrow muscular funnel low spiresaremore stablein areasof heavy wave shock
(= siphon). The forceful expelling of this jet of exhalant or on vertical rock surfaces.Similarly, the low, cap-
water can also provide a means of rapid locomotion shapedshellsof limpets (Figures13.5,4. and 13.16H,I)
for most cephalopods. In the mantle cavity the water are presumably adapted for withstanding exposureto
passes through the ctenidia, and then past the anus, re- strong waves. Heavy ribbing, thick or inflated shells,
productive pores, and excretory openings. and a narrow gape in bivalves are all possibleadap-
The remarkable adaptive qualities of the molluscan tationsto provide protectionfrom predators.In some
body plan are manifested in these variations in the po- gastropods,fluted shell ribs help them land upright
sition and function of the mantle cavity and its associ- when they are dislodged from rocks.Severalgroups of
ated structures. In fact, the nature of many other struc- soft-bottombenthic gastropodsand bivalves have long
tures is also influenced by mantle cavity arrangement, spineson the shell that may help stabilize the animals
as shown schematically in Figure 13.14.That molluscs in loosesedimentsas well as provide someprotection
have been able to successfully exploit an extremely from predators. Many molluscs, particularly clams,
broad range of habitats and lifestyles can be explained have shells coveredwith living epizootic organisms
in part by these variations, which are central to the such as sponges/annelid tube worms, ectoprocts,and
story of molluscan evolution. We will have more to say hydroids. Somestudies suggestthat predators have a
about these matters throughout this chapter. difficult time recognizing such camouflagedmolluscs
aspotential prey.
The Molluscan Shell Molluscsmay have one shell,two shells,eight shells,
Except for the two aplacophoran classes, all molluscs or no shell. In the latter casethe outer body wall may
have solid calcareous shells (either aragonite or calcite) containcalcareousscleritesof various sorts.In the apla-
produced by shell glands in the mantle. In the Caudo- cophorans,for example,the cuticular scleritesvary in
foveata and Solenogastres, aragonite sclerites or scales shapeand range in length from microscopicto about 4
are formed extracellularly in the mantle epidermis mm. Thesescleritesare essentiallycrystals composed
and are embedded in the cuticle. In the other classes almost entirely of calcium carbonateas aragonite.
molluscan shells vary greatly in shape and size, but Caudofoveatesproduceplatelikecuticular scleritesthat
they all adhere to the basic construction plan of cal- give their body surfacea scalytexture and appearance.
cium carbonate produced extracellularly, laid down The scleritesin both taxa appear to be secretedby a
ANTERIOR
posttorsional gonad. The right renopericardial duct serves

both the nephridium and the single gonad and leads to
a urogenital pore. As water enters the manfle cavity from
the front, it passes first over the two bipectinate ctenidia
Key and then over the anus, nephridiopore, and urogenital pore
before exiting through dorsal shell openings (e.g., holes
A Atrium
or slits). (C) The patellogastropod (limpet), Loffra. Here the
AN Anus posttorsional right ctenidium and right atrium are lost, and
CT Ctenidium the nephridiopore, anus, and urogenital pore are shifted
G Gonad to the right side of the mantle cavity, thus allowing a one-
way, left-to-right water flow. A similar configuration and is
N Nephridium
also found in vetigastropods with single gills (e.g., Tricotia\.
PC Pericardium (D) Most caenogastropods have a single, posttorsional left,
V Ventricle monopectinate ctenidium, suspended from the roof of the
mantle cavity. The right renopericardial duct has typically
lost its association with the pericardium and is co-opted
into the genital tract. Such isolation of the tract and gonad
from the excretory plumbing has allowed the evolution of
elaborate reproductive systems among ,,higher,,gastro-
pods (e.9., neritomorphs, caenogastropods, and hetero-
branchs) and probably important in the story of gastropod
Figure 13.14 Variations in the mantle cavity, circulatory success. (E) The condition in monoplacophorans includes
system, ctenidia, nephridia, reproductive system, and the serial repetition of several organs. (F) In polyplacopho-
position of the anus in molluscs (dorsal views). Although rans, the gonoducts and nephridioducts open separately
schematic, these drawings give some idea of the evolu- into the exhalant regions of the lateral pallial grooves. (G)
tionary changes in arrangement of these structures and A generalized bivalve condition. The gonads and nephridia
systems in the phylum Mollusca. (A) A hypothetical, untorl- may share common pores, as shown here, or else open
ed, gastropod-like mollusc with a posterior mantle cavity separately into the lateral mantle chambers. (H) The condi-
and symmetrically paired atria, ctenidia, nephridia, and tion in a generalized cephalopod with a single, isolated
gonads. (B) A posttorsional vetigastropod (e.g., Fissurella) reproductive system and an effectively closed circulatory
wherein all paired organs are retained except the left system.
PHYLUMMOLLUSCA 479
Subepidermal Eosinophilous (B)

gland cell gland cell
Mucocyte
Cuticle
Ciliated
epidermal cells
Circular
muscle layer
Diagonal (oblique) Epidermis
muscle layer Nacreous layer(s)
ffi{;;;;;,
muscle layer Periostracum
Periostracum
Hemolymph spaces (c) Mantle epithelium
Prismaticlayer
Nacreous layer
Outer (secretory)
mantle lobe
Figure 13.15 The body wall and
Mantle muscles
shell of molluscs. (A) A general- Middle (sensory)
ized molluscan body wall (section)" mantle lobe
The cuticle, epidermis, muscle layers, and various
gland cells constitute the body wall. (B) The com- Inner (muscular)
ponents of a generalized molluscan shell (section). mantle lobe
(C) The margin of the shell and the trilobed mantle Mucocyte
of a bivalve (transverse section). muscle Mintle epithelium
diffuse network of specialized groups of cells and dif- the microaesthetes. In some species,megaesthetes may
ferent shapes are found in different regions of the body. be modified as shell eyes, with compound lenses made
The eight transverse plates, or valves (Figures 13.4 of large crystals of araganite. The vertical aesthete ca-
and 13.16A-F), of polyplacophorans are encircled by nals arise from a layer of horizontal canals in the lower
and embedded in a thickened region of the mantle part of the tegmentum and the articulamentum (Figure
called the girdle. The size of the girdle varies from nar- 13.43C) and some pass through the articulamentum to
row to broad and may cover much of the valves. In the join with nerves in the mantle at the lower edge of the
giant Pacific " gumboot" chiton, Cryptochiton stelleri, t!.:te shell valve. The articulamentum is a thick, calcareous,
girdle completely covers the valves. The girdle is thick, porcelaneous layer that differs in certain ways from the
heavily cuticularized, and usually beset with calcare- shell layers of other molluscs.
ous sclerites/ spines, scales/or noncalcareous bristles Monoplacophorans have a single, limpet-like shell
secreted by specialized epidermal cells. These sclerites with the apex situated far forward (Figures 13.14 and
are probably homologous with those in the body wall 13.3).The shell has a distinctive outer prismatic layer
of aplacophorans. and an inner nacreous layer. As in chitons, the mantle
The anterior and posterior valves of chitons are re- encircles the body and foot as a circular fold, forming
ferred to as the end valves, or cephalic (= anterior) and lateral mantle grooves.
anal (= posterior) plates; the six other valves are called The bivalves possesstwo shells, ot valves, that are
the intermediate valves. Some details of chiton valves connected dorsally by an elastic, proteinaceous liga-
are shown in Figure 13.16A-F. The shells of chitons are ment, and enclose the body and spacious mantle cavity
three-layered, with an outer periostracum, a colored (Figures 13.1M-P,13.8, and 13.16J,K).Shells of bivalves
tegmentum, and an inner calcareous layer, or articu- typically have a thin periostracum/ covering two to
lamentum. The periostracum is a very thin, delicate or- four calcareous layers that vary in composition and
ganic membrane and is not easily seen.The tegmentum structure. The calcareous layers are often aragonite or
is composed of organic material (probably a form of an aragonite / calcite mixture, and they usually have a
conchin) and calcium carbonate suffused with various substantial organic framework. The periostracum and
pigments. It is penetrated by vertical canals that lead organic matrix may account for over 70% of the shell's
to minute pores in the surface of the valves. The pores dry weight in some thin-shelled taxa. Each valve has
are of two sizes: the larger ones (megapores) housing a dorsal protuberance called the umbo, which is the
the megaesthetes and the smaller ones (micropores) oldest part of the shell. Concentric growth lines radiate
Anterior
valve Figure 13,16 Shell morphology and terminology.
(A-F) Chiton shells (Polyplacophora): (A) A chiton
Intermediate showing the eight valves (dorsal view). (B) lsolated
valves
valves of Cryptochiton stelleri, the giant "gumboot"
chiton. (C) An anterior valve (ventral view). (D,E) An
intermediate valve (dorsal and ventral views). (F) A
posterior valve (ventral view). (G) lnternal and external
features of a spiral gastropod shell. (H) A lottiid limpet
(Patellogastropoda) (side view). (l) The shell of a veti-
gastropod keyhole limpet (top view). (J) Inside view of
Girdle the left valve of a heterodont clam shell (Bivalvia). (K)
(mantle) Dorsal view of a heterodont clam shell.
(c)
hrsertion
teeth
Posterior Periostracum(folded
(anal) valve onto underside of
(G)
valve from dorsal side)
Posterior
(D) Jugal sinus Apophysis Protoconch
Apex
Diagonal
line
Columella
Lateral Suture
hiangle
Varices
Jugal sinus
Inner lip
Insertion teeth
Insertion plate Perioshacum (folded

onto underside of
valve from dorsal side) Anterior
Foramen
Posterior
Anterior
Dorsal Anterior margin Portion

Umbo of hinge
0) ligament
Posterior retractor Umbo
Hinge teeth exposed
musclescar
Anterior
Lateral hinge retractor Lunule
tooth muscle scar
Right
POSTERIOR ANTERIOR shell
(siphon end) (foot end) valve
Posterior
adductor Anterior
muscle scar adductor
muscle scar
Pallial line
PHYLUMMOLLUSCA 481
Figure 13.17 Two very different kinds of cephalopod

sheffs. (A)The chamberedshell of Nautilus,cut in longitu-
dinal section.(B)The egg case "shell" of the paper nauti- of smooth nacreousor porcelaneousshell. The result
lus, Argonauta.
is a pearl, either free in the extrapallial spaceor partly
embeddedin the growing shell.6
outward from the umbo. \rVhen the valves are closed Scaphopodshells resembleminiature, hollow eI-
by contraction of the adductor muscles, the outer ephant tusks,hencethe vemacular names"tusk shell"
part of the ligament is stretched and the inner part is and "tooth shell" (Figures13.1,Land 13.9).The scaph-
compressed. Thus, when the adductor muscles relax, opod shell is open at both ends,with the smaller open-
the resilient ligament causes the valves to open. The ing at the dorsal end of the body. Most tusk shells are
hinge apparatus comprises various sockets and tooth- slightly curved, the concaveside being the equivalent
like pegs or flanges (hinge teeth) that align the valves to the anterior of other molluscs. The mantle is large
and prevent lateral movement. In most bivalves, the and lines the entire posterior surface of the shell. The
adductor muscles contain both striated and smooth dorsal aperture servesfor both inhalant and exhalant
fibers, facilitating both rapid and sustained closure of water currents.
the valves. This division of labor is apparent in some Most extant cephalopodshave a reduced shell or
bivalves as for example in oysters where the large sin- are shell-less.A completelydevelopedexternal shell
gle adductor muscle is clearly composed of two parts, is found only in fossil forms and the living speciesof
a dark striated region that functions as a rapid closure Nsutilus.In squids and cuttlefishthe shell is reduced
muscle, and a white smoother region that functions to and internal, and in octopusesit is entirely lacking or
hold the shell tightly closed for long periods of time. present only as a small rudiment.
The thin mantle lines the inner valve surfaces in bi- The shell of Nautilus is coiled in a planispiral fash-
valves and separates the visceral mass from the shell. ion (whorls lie in a single plane) and has a thin perio-
The edge of a bivalve mantle bears three longitudinal stracum (Figures 13.10,13.17 A, and 13.228).Nautilus
ridges or folds-the inner, middle, and outer folds shells (and all cephalopod shells) are divided into in-
(Figure 13.15C). The innermost fold is the largest and ternal chambersby transversesepta,and only the last
contains radial and circular muscles, some of which chamberis occupiedby the body of the living animal.
attach the mantle to the shell. The line of mantle at- As the animal grows, it periodically moves forward,
tachment appears on the inner surface of each valve and the posterior part of the mantle secretesa new seP-
as a scar called the pallial line (Figure 13.1.6D,and this tum behind it. Eachseptum is interconnected by a tube
scar is often a useful diagnostic character. The middle through which extendsa cord of tissue called the sip-
mantle fold is sensory in function, and the outer fold is huncle. The siphuncle helps to regulate buoyancy of the
responsible for secreting the shell. The cells of the outer animal by varying the amounts of gas and fluid in the
lobe are special2ed: the medial cells lay down the peri- shell chambers.The shell is composedof an inner nacre-
ostracum, and the lateral cells secrete the first calcareous layer and an outer porcelaneouslayer containing
ous layer. The entire mantle surface is then responsible prisms of calcium carbonateand an organic matrix. The
for secreting the remaining innermost calcareous por- outer surfacemay be pigmented or pearly white. The
tion of the shell. A thin extrapallial space lies between junctions between septaand the shell wall are called su-
the mantle and the shell, and it is into this space that tures, and are simple and straight, or slightly waved (as
materials for shell formation are secreted and mixed. inNautilus), or were highly convoluted (asin the extinct
Should a foreign object, such as a sand grain, lodge
between the mantle and the shell, it may become the 6Pearls are also found in some gastropods with nacreous irrrer
nucleus around which are deposited concentric layers shell layers, such as abalone.
482 ChaPter Thirteen
ammonoids).Lr cuttlefish(orderSepiida),the shellis re- shell form has been derived from coiled ancestorson
duced and intemal, with chambersthat arevery narrow numerous occasionsduring gastropodevolution'
spacesseparatedby thin septa.Like Nautilus,a cuttle- Gastropod shells consist of an outer thin organic
periostracumand two or three calcareouslayers: an
fish can regulate the relative amounts of fluid and gasin
its shell chimbers. The small, coiled, septate,gas-filled buter prismatic (or palisade) layer, and middle and
shells of the deep-watel squid Spirulaare occasionally inner lamellar or crossedlayers.In many vetigastro-
found washedup onbeaches. pods the inner layer is nacreous'In somepatellogastro-
Fossildata suggestthat the first cephalopodshells pods up to six calcareouslayersare distinguishablebut
were probably small curved cones.From theseances- in the great majority of living gastropodsthe shell struc-
tors bbth straight and coiled shells evolved, although ture is primarily one layer composedof crossedcrystals
secondaryunioiling probably occurred in several (crossedlamellar shell structure).Gastropodsin which
groups. Somestraight-shelledcephalopodsfrom the the shell is habitually coveredby mantle lobes lack
Ordovician period exceeded5 m in length, and some a periostracum(e.g.,olives and cowries),but in some
Cretaceouscoited specieshad shell diametersof 3 m' oth"t g.onps the periostracumis very thick and some-
Gastropodshellsare extremelydiversein size and times it is produced into lamellaeor hairs' The prismat-
shape(Figure13.1D,G).The smallestare microscopic ic and lamellate layers consist largely of calcium car-
(lesi than 1 mm) and the largestmay reach 70 cm in bonate,either as calciteor aragonite.Thesetwo forms
length. The "typical" shapeis the familiar conical spi- of calcium are chemicallyidentical,but they crystallize
ral wound around a central axis or columella (Figure differently and canbe identified by microscopicgxami-
13.16G).The turns of the spire form whorls, demar- nation of sectionsof the shell. Small amounts of other
catedby lines calledsutures.The largestwhorl is the inorganic constituentsare incorporated into the calci-
last (or tody) whorl, which bearsthe aperturethrough um Carbonateframework, including chemicalssuch as
which the foot and head protrude. The traditional view phosphate,calcium sulfate,magnesiumcarbonate,and
of a coiled gastropodshell with the spire upPermost, ialts of aluminum, iron, copper,strontium,barium, sili-
is actually';upside down," sincethe lower edge of the con,manganese,iodine, and fluorine.
aperture is anterior and the apex of the shell spire is An intiiguing aspectof gastropod evolution is shell
pbsterior. The first few, very small, whorls at the apex loss and the achievementof the "slug" form. Despite
are the larval shell, or protoconch (or its remnant), the fact that evolution of the coiled shell led to great
which usually differs in sculpturing and color from successfor the gastropods-75% of all living mol-
the rest of the shell' The last whorl and aperture may luscs are snails-secondary loss of the shell occurred
be notched and drawn out into an anterior siphonal many times in this classbut mostly in various groups
canal, to house a siphon when present.A smaller pos- of euthyneurans such as the seaslugs and land slugs'
terior canalmay alsobe presenton the rear edgeof the In forms such as the land and sea slugs, the shell
aperture that houses a siphon-like fold of the mantle may persist as a small vestige coveredby the dorsal
where waste and water are expelled. mantle (e.g.,in the euthyneuranseaslugsAplysiinae
Every imaginable variation on the basic spiraled and Pleurobranchidae,and the caenogastropodfam-
shell occursamong the gastropods(and someunimagi- ily Velutinidae), or as a small external rudiment as
in the carnivorous land slug Testacelln, or it may be
nable):the shell may be long and slender(e'g', auger
lost altogether(e.g.,the nudibranchs, the systellom-
shells)or short and plump (e.g.,trochids),or the shell
matophoransand someterrestrial stylommatophoran
may be flattened, with all whorls more-or-lessin one
plane (e.g.,sundials).In somethe spire may be more slugs, and in the neritimorph slug Titiscania)'In the
br lessiniorporated into the last whorl and eventually nuJibranchs (Nudibranchia) the larval shell is first
disappearfiom view (asin cowries).In somewith a covered,then resorbed,by the mantle during ontoge-
last whorl, the aperturemay be reduced ny. Shell loss occurred numerous times in gastropodt
to an elongatedslit (Figure13.1E(e.g.,cowries,olives,
-rr.h-lu.g"t particularly among the seaslugs ("opisthobranchs")
and cones).In a few groups the shell may coil so loose- and stylommatophoranpulmonates'Shellsare ener-
ly as to form a meanderingwormliketube (e'g',the so- geticaily expensiveto produce and require a reliable
ialed "tube snails,"vermetids and siliquariids; Figure ron."" of calcium in the environment, so it might be
13.19E).In a number of gastropodgroups the shell advantageousto eliminate them if compensatory
may be reduced and overgrown by the mantle, or it mechaniims exist. For example,most, if not all, sea
*uy dituppear entirely resulting in a slug (seebelow)' slugs secretechemicalsthat make them distastefulto
Most gaslropodsspiral clockwise;that is, they show predators.In addition, the bright coloration of many
right-handed, or dextral, coiling. Some-are sinistral nudibranchs may serve a defensivefunction. In some
(left-handed),and somenormally dextral speciesmay species,the color matchesthe animal's background
occasionallyproduce sinistralindividuals' In limpets such as the small red nudibranch, Rostangapulchra,
the shell is iap-shaped, with a low conical shapewith which matches almost perfectly the red sponge on
no or little visible coiling (Figure 13.16H,I)'The limpet which it feeds. Many nudibranchs, are however,
Left cerebral
ganglion
Circumenteric
Left pleural
newe ring
ganglion
Pleurointestinal
Left
connective
ganglion
Mantle edge
Left parietal
(: intestinal) Ventricle
ganglion
Visceral
Left atrium (: abdominal)
ganglion
Left
nephridiopore
Left osphradium Right ctenidium
Left ctenidium cavity
Mantle
(E)
4tt cavitY
Crossing of the
pleurointestinal
connectives
Left ctenidium
Mantle cavity
(behind)
Larval foot
(F)
Buccal
Buccal
ganglion
Figure 13.1 8 Pre- and post-torsional adult gastropods- Pleural

(A-D) Dorsal views. (A) Hypothetical untofted pre-gastro- ganglion-___
pod. (B,C) Stages of torsion. (D) The fully torted condition.
Note that the mantle cavity, gills, anus, and nephridio-
Pedal
pores are moved from a posterior to an anterior orienta-
ganglion
tion, just above and behind the head. Fudhermore, many
structures that were on the right side of the animal in the
pretorsionalcondition (e.9., the right gill, osphradium,
heart atrium, and nephridiopore) are located on the left
side after torsion has taken place (and the pretorsional left
gill, osphradium, atrium, and nephridiopore subsequently Infraparietal
occur on the right side). (E) A gastropod veliger larva, ganglion
before and after torsion (lateral view). Note that after tor-
sion the head can be withdrawn into the anterior mantle
cavity. (F) Configuration of the principal ganglia and con-
nectives of a hypothetical untoded and a torted adult
gastropod.
vent of torsion, a unique synapomolphy of gastropods,

conspicuous in nature. In these cases/ the color may and it is quite unlike anything else in the animal king-
serve to warn predators of the noxious taste of the dom. Torsion takes place during development in all
slug or, as suggestedby Rudman (1991),Predators gastropods, usually during the late veliger larval stage.
may simply ignore such bright "novelties" in their It is a rotation of the visceral mass and its overlying
environment. mantle and shell as much as 180" with respect to the
head and foot (Figures 13.18, and 13.53). The twist-
Torsion, or "How the Gastropod Got lts Twist" ing is always in a counterclockwise direction (viewing
One of the most remarkable and dramatic stepstaken the animal from above), and it is completely different
during the courseof molluscan evolution was the ad- from the phenomenon of coiling (= spiraling). During
torsion, the mantle cavity and anus are moved from a The evolution of asymmetricallv coiled shells had
posterior to an anterior position, somewhat above and the effect of restricting ihe right side of the mantle cavi-
behind the head. Visceral structures and incipient or- ty, a restriction that led to reduction or loss of the struc-
gans that were on the right side of the larval animal end tures it contained on the adult right side (the original
up on the left side of the adult. The gut is twisted into left ctenidium, atrium, and osphradium). At the same
a U-shape, and when the longitudinal nerve cords con- time, these structures on the adult left side (the origi-
necting the pleriral to the visceral ganglia develop, they nal right ctenidium, atrium, and osphradium) tended
are crossed rather like a figure eight. Most veligers have to enlarge. Possibly correlated with torsion and coiling
nephridia, which reverse sides, but the adult gills and was the loss of the left post-torsional gonad. The single
gonads are not fully developed when torsion occurs. remaining gonad opens on the right side via the post-
Torsion is usually a two-step process. During larval torsional right nephridial duct and nephridiopore.
development, an asymmetrical velar or foot retractor Patellogastropods and most vetigastropods retain two
muscle develops. It extends from the shell on the right, functional nephridia, although the post-torsional left
dorsally over the gut, and attaches on the left side of the one is often reduced. In other gastropods the post-tor-
head and foot. At a certain stage in the veliger's devel- sional right nephridium is lost, but its duct and pore
opment, contraction of this muscle causes the shell and remain associated with the reproductive tract in neriti-
enclosed viscera to twist about 90oin a counterclockwise morphs and caenogastropods.
direction. This first 90o twist is usually rapid, taking Such profound changes in spatial relations between
place in a few minutes to a few hours. The second 90o major body regions as those brought about by torsion
twist is tlpically much slower and results from differen- and spiral coiling in gastropods are rare among other
tial tissue growth. By the end of the process, the viscera animals. Several theories on the adaptive significance
have been pulled from above toward the left, ultimately of torsion have been proposed and are still being ar-
leading to the figure-eight arrangement of the adult vis- gued. The great zoologisi Walter Garstang rrrgg"it"d
ceral nerves. But the figure-eight arrangement is not per- that torsion was an adaptation of the veliger larva
fect as the left esophageal ganglion usually comes to lie that served to protect the soft head and larval ciliated
dorsal to the gut and is thus called the supraesophageal velum from predators (see the section on development
(= supraintestinal) ganglion; however, the right esopha- later in this chapter). When disturbed, the immediate
geal ganglion lies ventral to the gut, as a subesophageal reaction of a veliger is to withdraw the head and foot
(= subintestinal) ganglion (Figures 13.18and 13.40). into the larval shell, whereupon the larva begins to sink
Gastropods that retain torsion into adulthood are rapidly. This theory may seem reasonable for evasion
said to be torted; those that have secondarily reverted of very small planktonic predators, but it seems illogi-
back to a partially or fully untorted state in adulthood cal as a means of escape from larger predators in the
are detorted. The torted, figure-eight configuration of sea,which no doubt consume veligers whole-and any
the nervous system is referred to as streptoneury. The adaptive value to adults is not explained. Two zooLo-
detorted condition, in which the visceral nerves are gists finally tested Garstang's theory by offering torted
secondarily untwisted, is referred to as euthyneury. and untorted abalone veligers to various planktonic
Detorted gastropods, such as many heterobranchs, predators; they found that, in general, torted veligers
undergo a postveliger series of changes through which were not consumed any less frequently than untorted
the original torsion is reversed to various degrees. The ones (Pennington and Chia 1985). Garstang first pre-
process shifts the mantle cavity and at least some of sented his theory in verse, in 7928, as he was often
the associatedorgans about 90'back to the right (as in taken to do with his zoological ideas.
many "pulmonates" and some sea slugs), or in some
The Ballad of the Veliger,or
casesall the way back to the rear of the animal (the de- How the Gastropod Got Its Tzuist
torsion seen in some nudibranchs).
TheVeliger'sa liaely tar, thelioeliestafloat,
After torsion the anus lies in front; this means that
the first gastropods could no longer grow in length eas- A zuhirlingutheelon eithersidepropelshis little boaq
ily. Subsequent increase in body size thus occurred by But whenthedangersignalwarnshis bustlingsubmarine,
the development of loops or bulges in the middle por- He stops the engine,shuts the port, and drops below
tion of the gut region, thereby producing the character- unseen.
istic coiled visceral hump. The first signs of torsion and He'switnessedseaeralchangesin pelagicmotorcraft;
coiling occur at about the same time during gastropod Thefirst hesailedu.tasjust a tub, znitha tiny cabinaft.
development. The earliest coiled gastropod shells in An Archi-molluskfashioned it, accordingto hiskind,
the fossil record include both planispiral and conispiral He'd alzlaysstozoed his gills and things in a mantle-sac
forms, and it is possible that coiling predated the ap- behind.
pearance of torsion in gastropods. Once both features YoungArchi-molluskswent to sea with nothing but a
were established, they coevolved in various ways to aeLum-
produce what we see today in living gaskopods. A sort of autocyclinghoop,insteadof pram-to wheel'em;
PHYLUMMOLLUSCA 485
And, spinning round, they one by one acquiredparental theoretically allow for greater growth and elongation
features, of the shell while reducing the tendency of the animal
A shellaboae,afoot below-the queerest little creatures. to topple over sideways.
But when by chancethey brushedagainsttheir neighbors No matter what the evolutionary forceswere that led
in thebriny, to torsion in the earliestgastropods,the resultswere to
Coelenterates with stinging threadsand Arthropodsso move the adult anus, nephridiopores, and gonoPores
spiny, : to a more anterior position, corresponding to the new
By oneweakspotbetrayed, alas,theyfeII an easyprey- position of the mantle cavity. It should be noted how-
Theirsoftpreorallobesin front couldnot betuckedaway! ever, that the actual position and arrangement of the
Theirfeet,you see,amidships,next thecuddly-holeabaft, mantle cavity and its associatedstructuresshow great
Drezuin at once,and left theirheadsexposed to eueryshaft. variation; in many gastropodsthesestructures,while
SoArchi-mollusksdwindled,and theracewassinkingfast, pointing forward, may actually be positioned further
Whenby themerestaccidentsalaationcameat last. towards the posterior region of the animal's body.
A fleet offry turnedout oneday,eoentfulin thesequel, Torsionis not a perfectly s).'mmetricalprocess.
Whoseleft and right retractorson the two sideswere Most of the stories of gastropod evolution focus on
unequar: changesin the mantle cavity and its associatedstruc-
Their starboardhalliardsfixed asternalonesuppliedthe tures, and many of these changesseem to have been
head, driven by some negative impacts of torsion. Many
While thoseset aport werespreadabeamand sentedthe anatomical modifications of gastropods appear to be
backinstead. adaptations to avoid fouling, for without changing
Predaceous foes,still drifting by in numbersunabated, the original flow of water through the mantle cav-
Were bffied now by tactics which their dining plans ity in a primitive gastropod with two ctenidia, waste
frustrated. from the centrally positioned anus (and perhaps the
Their prey upon alarm collapsed,but promptly turned nephridia) would be dumped on top of the head and
about, potentially pollute the mouth and ctenidia. Hence, it
With the tender morselsafe within and the horny foot has long been hypothesized that the first step, sub-
without! sequentto the evolution of torsion, was the develop-
Thismanoeuore (aideLamarck)speeded up with repetition, ment of slits or holes in the shell, thus altering water
llntil thepartsfficted gained a rhythmical condition, flow so that a one-way current passedfirst over the
And torsion,needingnow no morea stimulatingstab, ctenidia, then over the anus and nephridiopore, and
WiIl take its predetermined coursein a watchglassin the finally out the slit or shell holes. This arrangement is
lab. seenin some vetigastropods, such as the slit shells
In this way, then,the Veliger,triumphantlyaskew, (Pleurotomarioidea)and abaloneand keyhole limpets
Acquiredhis cabinfor'ard, holdingall his sailingcrew- (Figures13.1D,1,3.16I, and 13.36).As reasonableas it
A kochospherein armour cased,with a foot to work the sounds, there has been surprisingly little empirical
hatch, evidencein support of this hypothesis.In additiory the
And doublescrewsto driae aheadwith smartnessand adaptive significance of shell holes was examined by
dispatch. Voltzow and Collin (1995),who found that blocking
But whenthefirst new Veligersffimehomeagainto shote, the holes in keyhole limpets did not result in damage
And settleddownas Gastropods with mantle-sac afore, to the organs of the mantle cavity. Thus, the adaptive
The Archi-mollusksoughta cleft,his shameand grief to significanceof torsion in gastropod evolution remains
hide, an open question.
Crunchedhorribly his horny teeth,gaoeup theghost,and Once evolutionary reduction or loss of the gill and
died. osphradium on the right side had taken place, water
flow through the mantle cavity was from left to right,
Other workers have hypothesized that torsion was passing through the left gill and osphradium first, then
an adult adaptation that might have created more acrossthe nephridiopore and anus, and on out the
space for retraction of the head into the shell (perhaps right side. This strategyalso had the effectof allowing
also for protection from predators), or for directing the structureson the left side to enlarge and eventually to
mantle cavity with its gills and water-sensing osphra- develop more control over water flow into and out of
dia anteriorly. Still another theory asserts that torsion the mantle cavity, including the evolution of long si-
evolved in concert with the evolution of a coiled shell- phons. While most gastropodshave retained full or
as a mechanism to align the tall spiraling shells from a partial torsion, many of the heterobranchgastropods,
position in which they stuck out to one side (and were all of which lost the original ctenidium, have under-
presumably poorly balanced and growth limiting), gone various degreesof detorsion, and a host of other
to a position more in alignment with the longitudinal modifications,perhapsin responseto the absenceof
(head-foot) axis of the body. The latter position would constraintsoriginally brought on by torsion.
(zr,)rttti$
Zr,)li rttr'\
?/,)rtttnri '-/trittttlt)
'(,rit,,rl
(D)
Figure 13.1 9 (A,B) Locomotion in a benthic gastropod moving to the right by waves
of contraction of the pedal and foot muscles (solid arrow indicates direction of ani-
mal movement; dashed arrow indicates direction of muscle wave). In (A) the waves of
contraction are moving in the same direction as the animal, from back to front (direct
waves). Muscles at the rear of the animal contract to lift the foot off the substratum:
the foot shortens in the contracted region and then elongates as it is placed back
down on the substratum after the wave passes. In this way, successive sections of the
foot are "pushed" forward. In (B) the animal moves forward as the contraction waves
pass in the opposite direction, from front to back (retrograde waves). In this case, the
pedal muscles lift the anterior part of the foot off the substratum, the foot elongates,
is placed back on the substratum, then contracts to "pull" the animal forward, rather
like "stepping." (C) Calliostoma, a vetigastropod (Calliostomatidae) adapted to crawling
on hard substrata. Note the line separating the right and left sides of the trailing foot;
the line denotes a separation of muscle masses that allows a somewhat "bipedal-like,,
motion as the animal moves (see text for further details). (D) The moon snail, Potinices
(Naticidae), has a huge foot that can be inflated by incorporating water into a network
of channels in its tissue, thus allowing the animal to plow through the surface layer of
soft sediments. (E) Ienagodus (Siliquariidae),a sessile siliquariid worm snail.
Locomotion
The foot in aplacophorans is either rudimentary or lost and Solenogastres are largely symbiotic onvarious cni-
(Figure 13.2). Caudofoveata are mostly infaunal bur- darians. With the exception of these two groups, most
rowers and move by peristaltic movements of the body other molluscs possess a distinct and obvious foot,
wall, using the anterior mouth shield as a burrowing with the exception of the cephalopods where it is very
device and anchor. The foot of solenogasters is only highly modified. In chitons, monoplacophorans and
weakly muscular, and locomotion is primarily by slow most gastropods the foot often forms a flat, ventral,
ciliary gliding movements through or upon the sub- creeping sole (Figures 13.38, 13.48, 13.58, and 13.19).
stratum. Caudofoveata are mostly infaunal burrowers/ The sole is ciliated and imbued with numerous gland
PHYLUM MOLLUSCA 487
cells that produce a mucous trail over which the animal also use their broad girdle for additional adhesion to
glides. In gastropods, enlarged pedal glands supply the substratum by clamping down tightly and rais-
substantial amounts of mucus (slime), this being espe- ing the inner margin to create a slight vacuum. Some
cially important in terrestrial species that must glide snails, such as the Vermetidae and Siliquariidae are en-
on relatively dry surfaces. In most gastropods, there tirely sessile,the former attached to hard substrata, the
is an anterior mucus gland, which opens in a slit on or latter (Figure 13.19E)living in sponges. These gastro-
just behind the antbrior edge of the foot. This anterior pods have typical larval and juvenile shells; but after
lobe is called the propodium, the rest of the foot the they settle and start to grow, the shell whorls become
metapodium. In some caenogastropods, an enlarged increasingly separated from one another, resulting in
metapodial mucus gland opens into the middle of the a corkscrew or twisted shape. Other gastropods, such
sole. Small molluscs may move largely by ciliary pro- as slipper shells, are sedentary. They tend to remain in
pulsion but most move primarily by waves of muscu- one location and feed on organic particles in the sur-
lar contractions that move along the foot. rounding water. The sole of the hipponicid limpets se-
The gastropod foot possessessets of pedal retractor cretes a calcareous plate and the adults are thus oyster-
muscles, which attach to the shell and dorsal mantle at like and deposit feed using their long snout.
various angles. These and smaller muscles in the foot Some limpets and a few chitons exhibithomingbe-
act in concert to raise and lower the sole or to short- haviors. These activities are usually associated with
en it in either a longitudinal or a transverse direction. feeding excursions stimulated by changing tide levels
Contraction waves may move from back to front (di- or darkness, after which the animals return to their
rect waves), or from front to back (retrograde waves) homesites which is seen as a scar or even a depression
(Figure 13.19A,B). Direct waves depend on contraction on the rock surface. Homing behaviors are also seen in
of longitudinal and dorsoventral muscles beginning some land snails and slugs.
at the posterior end of the foo! successive sections of Most bivalves live in soft benthic habitats, where
the foot are thus "pushed" forward. Retrograde waves they burrow to various depths in the substratum
involve contraction of transverse muscles interacting (Figure 13.20E-D. In these infaunal species the foot is
with hemocoelic pressure to extend the anterior part of usually bladelike and laterally compressed (the word
the foot forward, followed by contraction of longitudi- pelecypod means "hatchet foot"), as is the body in
nal muscles. The result is that successive areas of the general. The pedal retractor muscles in bivalves are
foot are "pulled" forward (Figure 13.19A,B). In some somewhat different from those of gastropods, but they
gastropods the muscles of the foot are separated by a still run from the foot to the shell (Figure 13.8D). The
midventral line, so the two sides of the sole operate foot is directed anteriorly and used primarily in bur-
somewhat independently of each other. The right and rowing and anchoring. It operates through a combina-
left sides of the foot alternate in their forward motion, tion of muscle action and hydraulic pressure (Figure
almost in a stepping fashion, resulting in a sort of "bi- 13.20A-D). Extension of the foot is accomplished by
pedal" locomotion. engorgement with hemolymph, coupled with the ac-
Modifications of this general benthic locomotory tion of a pair of pedal protractor muscles. With the
scheme occur in many groups. Some gastropods, such foot extended, the valves are pulled together by the
as moon snails (Figure 13.1,9D),plow through the sedi- shell adductor muscles. More hemolymph is forced
ment, and some even burrow beneath the sediment from the visceral mass hemocoel into the foot hemo-
surface. Such gastropods often possess an enlarged, coel, causing the foot to expand and anchor in the sub-
shield-like propodium that acts like a plough and stratum. Once the foot is anchored, the anterior and
some naticids and cephalaspideans possess a dorsal posterior pairs of pedal retractor muscles contract and
flaplike fold of the foot that covers the head as a pro- pull the shell downward. Withdrawal of the foot into
tective shield. Other burrowers, such as augers, dig the shell is accomplished by contraction of the pedal
by thrusting the foot into the substratum, anchoring retractors coupled with relaxation of the shell adduc-
it by engorgement with hemolymph, and then pull- tor muscles. Many infaunal bivalves burrow upward
ing the body forward by contraction of longitudinal in this same manner, but others back out by using hy-
muscles. In the conch Strombus,the operculum forms a draulic pressure to push against the anchored end of
large " claw" that digs into the substratum and is used the foot. Most motile bivalves possess well-developed
as a pivot point as the animal thrusts itself forward anterior and posterior adductor muscles (the dimyar-
like a pole-vaulter using its muscular, highly modi- ian condition).
fied foot. In some heterobranchs, notably the sea hares There are several groups of bivalves that have epi-
(Aplysidae), lateral flaps of the foot expand dorsally as faunal lifestyles and are permanently attach to the
parapodia and these fuse dorsally in some species. substratum either by cementing one valve to a hard
Some molluscs that inhabit high-energy littoral hab- surface as in the true oysters such as the rock oysters
itats, such as chitons and limpets, have a very broad (Ostreidae) and rock scallops (Spondylidae). Others
foot that can adhere tightly to hard substrata. Chitons use special anchoring threads (byssal threads), as
Exhalant siphon
/ ,.Inhalantslphon
(H)
Figure 13.20 (A*D) Burrowing and life

positions of some infaunal bivalves. (A) Shell
adductor muscle relaxes, causing the shell valves
to push apart and create an anchorage. Pedal retrac-
tor muscles relax. Circular and transverse foot muscles Burrow
cotract, causing the foot to extend into the substratum.
(B) Hemolymph is pumped into the tip of the foot, causing
it to expand and form an anchorage. Siphons close and
withdraw as the shell adductor muscles contract, clos- in mussels (Mytilidae) (Figure 1,3.21A,8),ark shells,
ing the shell and forcing water out between the valves
and a number of other families including winged or
and around the foot. (C) Anterior and posterior pedal
retractor muscles contract, pulling the clam deeper into pearl oysters (Pteriidae), and numerous other pterio-
substratum. (D) The shell adductor muscle relaxes to morphian bivalves including the Pinnidae and many
allow shell valves to push apad and create an anchor- Arcidae and Pectinidae. While the juveniles of many
age in the new position. The foot is withdrawn. (E-l) Five heterodont bivalves produce one or a few temporary
bivalves in soft sediments; arrows indicate direction of byssal threads, a few species, such as the zebra mussel
water flow. (E) A deep burrower with long, fused siphons (Dreissena),remain byssally attached as adults.
(Iresus). (F) A shallow burrower with very short siphons
The true oysters (Ostreidae) (including the edible
(Clinocardium). (G) A deep burrower with long, separate
siphons (Scrobicularia). (H) The razor clam ('Iagelus) lives American and European oysters) initially anchor as
in unstable sands and maintains a burrow into which it a settling veliger larva (called a spat by oyster farm-
can rapidly escape. (l) The pen shell, Atrina, attaches its ers) by secreting a drop of adhesive from the byssus
byssal threads to solid objects buried in soft sediments. gland. Adults, however, have one valve permanently
PHYLUMMOLLUSCA 489
cemented to the substratum, with the cement being by currents(Figure13.1N).In jingle shells(Anomiidae),
produced by the mantle. the byssalthreadsrun from the upper valve through a
Byssal threads are secreted as a liquid by the byssus hole in the lower valve to attach to the substratum after
gland in the foot. The liquid flows along a groove in which they becomesecondarilycalcified.Byssalthreads
the foot to the substratum, where each thread becomes probably representa primitive and persisting larval fea-
tightly affixed. The threads are emplaced by the foot; ture in those groups that retain them into adulthood,
once attached they quicklyhardenby a tann-ingprocess/ and many bivalveslacking byssalthreadsas adults uti-
whereupon the foot is withdrawn. A byssal thread re- lize them for initial attachment during settlement.
tractor muscle may assist the animal in pulling against
its anchorage. Mussels have a small, finger-like foot
whose principal function is generation and placement Figure 13.2'l More bivalves.(A) A "bed" of mussels
(Mytilus californianus) attached by byssal threads (close
of the byssal threads. Giant clams (Cardiidae) initially
up of two mussels). (B) A mussel (lateral view, with left
attach by byssal threads, but usually lose these as they
valve removed). (C) Shell of the Mesozoic rudist clam
mature and become heavy enough not to be cast about Coralliochama. (D) The wood-boring bivalve (shipworm)
Teredo. The pallets (only one is shown) are a pair of shelly
plates that close over the siphons when they are retract-
ed. (E) A shipworm-bored piece of driftwood (notice the
thousands of small holes). (F) A pholad-bored rock. The
oholad can be seen in its bore hole.
490 ChaoterThirteen
In many families of attached bivalves, such as mus- in several different taxa in several different ways, in-
sels and true oysters, the foot and anterior end are re- cluding by valve flapping in scallops. In most others
duced. This often leads to a reduction of the anterior of these groups, the foot is modified as the swimming
adductor muscle (anisomyarian condition) or its com- structure. In the unique caenogastropod group known
plete loss (monomyarian condition). as heteropods, the body is laterally compressed, the
Great variation occurs in shell shape and size shell is greatly reduced, the foot forms a fin, and the an-
among attached 6ivalves. Some of the most remark- imal swims upside down (Figure 13.74{). Swirnming
able were the Mesozoic rudists, in which the lower has evolved several times in the heterobranchs, includ-
valve was horn-like and often curved, and the upper ing the pteropods (seabutterflies), where the parapodi-
valve formed a much smaller hemispherical or curved al extensions of the foot form two long lateral fins that
lid (Figure 73.21C).Rudists were large, heavy creatures are used like oars (Figure 13.7D,8). Some nudibranchs
that often formed massive reef-like aggregations, either also swim by graceful undulations of flaplike parapo-
by somehow attaching to the substratum or by simply dial folds along the body margin or by vigorous un-
accumulating in large numbers on the seabed, in "log dulations of the body. Although not technically swim-
jams." These accumulations of fossil shells provide the ming, violet shells (lanthina) float about the ocean's
spaces in which oil deposits formed in sediments in surface on a raft of bubbles secreted by the foot, and
many parts of the Middle East and Caribbean. some planktonic nudibranchs (e.g., Glaucus,Glaucilla)
Some originally attached bivalves have evolved to stay afloat by use of an air bubble held in the stomach!
live freely upon the sea floor (e.g., some Pectinidae and The champion swimmers are, of course, the cepha-
Limidae) (Figure 13.1M). Some are capable of short lopods (Figures 13.U,K and13.22). These animals have
bursts of " jet-propelled" swimming, which is accom- abandoned the generally sedentary habits of other
plished by quickly clapping the valves together. molluscs and have become highly effective swimming
The habit of boring into hard substrata has evolved predators. Virtually all aspects of their biology have
in several different bivalve lines. In all cases,excava- evolved to exploit this lifestyle. Most cephalopods
tion begins soon after larval settlement. As the animal swim by rapidly expelling water from the mantle cav-
bores deeper, it grows in size and soon becomes per- ity. In the coleoid cephalopods the mantle has both
manently trapped, with only the siphons protruding radial and circular muscle layers. Contraction of the
out of the original small opening. Boring is usually by radial muscles and relaxation of the circular muscles
a mechanical process; the animal uses serrations on the draws water into the mantle cavity while reversal of
anterior region of the shells to abrade or scrape away this muscular action forces water out of the mantle
the substratum. Some species also secrete an acidic cavity. The mantle edge is clamped tightly around the
mucus thatpartially dissolves or weakens hard calcare- head to channel the escaping water through a ventral
ous substrata (limestone, coral,large dead shells). Some tubular funnel, or siphon (Figure 13.11B,C).The fun-
species bore into wood, such as Martesin (Phaladidae), nel is highly mobile and can be manipulated to point in
Xylophaga (Xylophagidae), and nearly all species in nearly any direction, thus allowing the animal to turn
the family Teredinidae (Bankia, Teredo).Teredinids, and steer. Squids attain the greatest swimming speeds
with their long wormlike bodies, are known as ship- of any aquatic invertebrates, and several species can
worms because of the destruction that they can cause even leave the water and propel themselves many feet
to the wooden hulls of ships (or to wood pier pilings). into the air. Most octopuses are benthic and lack the
In the teredinids the shells are reduced to small ante- fins and streamlined bodies characteristic of squids.
rior bulb-like valves that serve as the drilling apparatus Although octopuses still use water-powered jet pro-
(Figure 13.27D,E).Some pholads bore into soft stone pulsion, they more commonly rely on their long suck-
(e.g., Pholas),or into other substrata (e.g., Barnea;Figure ered arms for crawling about the sea floor. Some octo-
73.218). Some species in the family Mytilidae also are puses have even been observed moving about upright
borers, such as Lithophaga,which bores by mechanical on only two tentacles-bipedal locomotion! Cuttlefish
and possibly chemical means into calcareous rocks, are slower than squids, and often use their fins for for-
shells of various other molluscs (including chitons), and ward swimming as well as stabilization and to assist in
corals, and the genus Adula, which bores into soft rocks. steering and propulsion.
Scaphopods are adapted to infaunal habitats, bur- Nnutilus move up and down in the water column on
rowing vertically by the same basic mechanism used by a diurnal cycle, often traveling hundreds of meters in
many bivalves (Figures 13.1L and 13.9).The elongate each direction. They can actively regulate their buoy-
foot is projected downward into soft substrata, where- ancy by secretion and reabsorption of shell chamber
upon a rim in the distal part of the foot is expanded to gases (chiefly nitrogen) by the cells of the siphuncle.
serve as an anchoring device; contraction of the pedal The unoccupied chambers of these shells are filledpart-
retractor muscles pulls the animal downward. ly with gas and partly with a liquid called the cameral
Perhaps the most remarkable locomotory adap- fluid. The septa act as braces, giving the shells enough
tation of molluscs is swimming, which has evolved strength to withstand pressures at depth. As discussed
PHYLUM MOLLUSCA 491
earlier, each septum in nautiloid shells is perforated by The buccal cavity may contain a pair of lateral jaws
a small hole, through which runs the siphuncle, which (or a single dorsal jaw), muscularized regions with chi-
originates in the viscera and is enclosed in a porous tinous plates that can be solid or composed of multiple
calcareous tube. Various ions dissolved in the cameral small units. Molluscan jaws are highly variable. For
fluid can be pumped through the porous outer layers example, in some heterobranchs the jaws can be quite
into the cells of the siphuncular epithelium. \A/hen the complex, with distinct "teeth," in some carnivorous
cellular concentration of ironsis high enough, the diffu- caenogastropods the jaws can be quite large, in ceph-
sion gradient thus created draws fluid from the shell alopods the jaws are modified to form the beak, and
chambers into the cells of the siphuncle while the fluid some lineages have no jaws at all including bivalves,
is replaced with gas. The result is an increase in buoy- whichlackboth jaws and radula.
ancy. By regulating this process,Nautilus may be able The radula is usually a ribbon of recurved chitinous
to remain neutrally buoyant at whatever depth they teeth (Figures 13.23-13.26).The teeth may be simple,
are. It was once thought that this gas-fluid "ptrrr.p" serrate, pectinate, or otherwise modified. The radula
mechanism allowed buoyancy changes sufficient to ex- often functions as a scraper to remove food particles
plain all the large-scale vertical movements of Nautilus, for ingestion, although in many groups it has become
but density changes may not be the sole source of adapted for other actio.tt. A radula is present in the
power for moving great distances up and down in the majority of the most primitive living molluscs and is
water column. Nautilus moves using jet propulsion therefore assumed to have originated in the earliest
by rapidly contracting its head, not by mantle muscle stages of molluscan evolution. In the aplacophoran
contraction. groups the teeth, when present, may not be borne on
a ribbon per se but on a relatively thin cuticle cover-
F eeding ing the foregut epithelium-perhaps the evolutionary
Two basic and fundamentally different types of feed- forerunner of the ribbon-like radula. In some apla-
ing occur among molluscs: the first encompasses the cophorans, the teeth form simple plates embedded in
feeding modes of most molluscs and includes micro- to either side of the lateral foregut wall, while in others
macrophagy involving browsing and scraping, herbiv- they form a transverse row, or up to 50 rows, with as
ory, carnivorous grazing and predation, while the sec- many as 24 teeth per row.
ond is suspension feeding (suspension microphugy).
The basic mechanics of these two feeding modes are
examined in Chapter 4. Here we briefly summarize
the ways in which these feeding behaviors are em-
ployed by molluscs. In this section we also discuss
a uniquely molluscan structure, the radula, which is
used in microphagy, herbivory, and predation, and
has become modified in a variety of unusual and
interesting ways.
Figure 13.22 Swimming cephalopods. (A) Sepra, the

cuttlefish. (B) Naufllus. (C) Vampyroteuthis, a "vampire"
squid, viewed from the side...-
492 ChaoterThirteen
Figure 13.23 A generalized molluscan radula and asso- Esophagus

ciated buccal structures, at three "magnifications" (lon-
gitudinal section).
Radular
retractor
kr gastropodsand other molluscs (exceptbivalves)
an odontophore-projectsfrom the floor of the pharynx Odontophore
rehactor m
or buccal cavity.'It is a muscular structure bearing the
Radular
complex tooth-bearing radular ribbon (Figure 13.23). protractor muscles
The ribbon, calleda radular membrane,is moved back
Muscles
and forth by setsof radular protractor and retractor
musclesover cartilagesencasedin the odontophore
(Figure 13.23).Thesecartilagesare absentin many het- Odontophore Salivary gland
erobranchgastropods.The radula originatesin a radu- protractor muscles
lar sac,in which the radular membraneand new teeth - Radular sac
are continually being produced by specialcells called
odontoblasts,to replacethoselost by erosionduring
feeding.Measurementsof radular growth indicatethat
up to five rows of new teeth may be added daily in
Radular
membrane
with
radular
Muscles in wall
of radular sinus
I, ' l
ular I
some species.The odontophore itself is moved in and teeth

out of the buccalcavity during feedingby setsof odon-
ular
tophore protractor and retractor muscles,which also membrane
assistin applying the radula firmly againstthe substra-
tum (Figures 13.23and 13.24A,8).
The number of radular teeth ranges from a few to
thousandsand seryesasan important taxonomiccharac- Odontophoral
ter in many groups.In somemolluscs,the radular teeth cartilage Subradular pouch
are hardenedwith iron compounds,such as magnetite
(in chitons)and goethite (in patellogastropods).]ust as
in many vertebrates,the radular teeth show adaptations over the bending plane of the odontophore,theseteeth
to the type of food eaten.In vetigastropods(e.g.,keyhole act like stiff brushes, sweeping small particles to the
limpets, abalones,top shells),the rhipidoglossate rad- midline where they are caught on the recurved parts of
ulae bear large numbers of fine marginal teeth in each the centralteeth,which draw the particlesinto the buc-
row (Figures13.254and 13.26A).As the radula is pulled cal cavity. Most vetigastropodsare intertidal foragers
Radular teeth
Figure 13.24 Feeding in macrophagous molluscs.

(A) Cutting and scraping action of a gastropod radula.
(B) A boring gastropod, the moon snail Nafica, with
radula visible in the mouth and the boring gland
exposed (oral view). (C) The Pacific chiton
Placiphorella velata in feeding position, with
raised head flap ready to capture small prey.
PHYLUMMOLLUSCA 493
(c) Central
tooth
Lateral tooth
Marginal teeth
Central tooth
/M\arginal teeth (D)

Central tooth Lateral tooth Marginal tooth
/
Figure 13.25 Various arrangements of radular teeth.

(A) The rhipidoglossan condition of an abalone, Haliotis
(Vetigastropoda). The marginals on the right side are not
shown. (B) The taenioglossan condition of the caeno-
gastropod Viviparus. (C) The taenioglossan condition of
the caenogastropod Littorina. (D) The highly modified
taenioglossan condition of the heteropod Pterotrachea
(Caenogastropoda). Only one transverse row of teeth is
shown. (E) The rachiglossan condition of the neogastro-
pod Buccinum. (F) The toxoglossan condition of the neo-
gastropod Mangelia (a single tooth).
that live on diatoms and other algae and microbes on marginalteethaltogether(Figures13.25Eand13.26c,D).

the substratum.In contrast,patellogastropods(e.g.,lot- They use the remaining (one to three) teeth for rasping,
tiid and patellid limpets)possessa docoglossateradula, tearing, or pulling. Thesesnails are usually carnivores
which is impregnated with iron and bear relatively few or carrion feedersalthough some members of one fam-
teeth in eachtransverserow. Lottiid radulae,for exam- ily, the Columbellidae, are herbivores.Caenogastropods
ple, have only one,two, or no marginal teeth,and only of the families Muricidae and Naticidae eat other mol-
three pairs of lateral teeth per row (Figure 13.268).The luscs by boring through the prey's calcareousshell
mucous trails leftby somelimpets (e.g.,homing species to obtain the underlying flesh. This ability to bore haE
such asthe Pacific Lottingiganteaand Collisellascabra)ac- evolved entirely independently in the two groups. It i3
tually serveas adhesivetraps for the microalgaethat are mainly mechanical;the predator boring with its radula
their primary food resource). while holding the prey with the foot. The boring activity
The radula of many caenogastropodsis the taenio- is complemented by the secretionof an acidic chemical
glossatetype, in which there are only two marginal from a boring gland (alsocalled the "accessoryboring
teeth in eachrow, along with three other teeth (laterals organ"); the chemical is periodically applied to the drill
and central)(Figure 13.258-D).In conjunctionwith the hole to weaken the calcareousmatrix. The boring gland
elaborationof jaws, taenioglossateradulae are capable of the neogastropodmuricids is located on the foot
of powerful rasping,which enablessomelittorid snails while that of the littorinimorphnaticids is located on the
to feedby directly scrapingoff the surfacecell layersof anterior end of the proboscis(Figure 13.248| Boring gas-
algae. tropods such asthe American drill (Urosalpinx)and the
The most derived caenogastropods(Neogastro- |apanesedrt1l(Rapana)causea loss of millions of dollars
poda) usually have rachiglossateradulae, which lack annuallv for ovster farms.
Figure 13.26 Gastropod radulae.

(A) A closeup view of the rhipidoglossate
radufa of the abalone Haliotis rufescens
(Vetigastropoda). Note the many hook-
like marginal teeth. (B) The docoglossate
radula of a lottiid limpet. (C) The serrated
central teeth of a rachiglossate radula
from the muricid whelk Nuce//a emargina-
ta, a neogastropod (Caenogastropoda)
that preys on small mussels and barna-
cles. (D) The worn radular teeth of Nucella.
(E) The radula of the polycerid nudibranch
Triopha, seen here in dorsal view as it
rests in the animal.
Some carnivorous gastropods (e.9., lnnthina) do venom gland. A few Indo-West Pacific cones produce
not gnaw or rasp their prey, but swallow it whole. In a potent neuromuscular toxin that has caused human
these gastropods a ptenoglossate radula forms a cov- deaths.
ering of strongly curved spines over the buccal mass. Among the most unusual gastropod feeding strate-
The prey is seized by the quickly extruded buccal gies are those that involve parasitism on fishes. For ex-
mass and simply pulled whole into the gut. A some- ample, the neogastropod Cancellariacooperiattaches to
what sirnilar feeding method is seen in the camivorous the Pacific electric ray and makes small cuts in the skin
slugTestacella,where the hooked radula catches earth- through which the proboscis is inserted to feed on the
worms that the slug consumes whole. The nudibranch ray's blood and cellular fluids. Several other neogas-
Melibe (chapter opener photo) uses its large hood to tropods parasitize "sleeping" reef fishes by inserting
sweep the water for copepods, amphipods, and other their proboscides into the host and sucking out fluids.
small planktonic prey. Some other gastropods are known to parasitize vari-
A few gastropods have lost the radula altogether ous invertebrate hosts, notably the pyramidellids (on a
and feed by suckingbody fluids from their prey, a habit variety of invertebrates, including other molluscs) and
seery for example, in some nudibranchs. Pyramidellids the Eulimidae on echinoderms, with the latter group
do this with the aid of a hypodermic stylet (a modified including some internal parasites that have lost their
jaw) on the tip of an elongate proboscis. shell and become wormlike.
One of the most specialized feeding modes in gas- Certain euthyneurans also show various radular
tropods is seen in the cone snails (Conus), and rela- modifications. Groups of "opisthobranchs" that feed
tives. Their toxoglossate radula is formed from a few on cnidarians, ectoprocts, and sponges, and those that
harpoon-like/ venom injecting teeth that are probably scrape algae (e.g., aplysiids) usually have typical rasp-
modified marginal teeth. The teeth (Figure 13.25F) are ing radulae. In sacoglossans, however, the radula is
discharged from the end of a long proboscis that can be modified as a single row of lance-like teeth that can
extended very rapidly to capture prey, usually a fish, a pierce the cellulose wall of filamentous algae, allowing
worm, or another gastropod, which is then pulled into the gastropod to suck out the cell contents. A similar
the gut (Figure 73.27). The venom is injected through type of feeding strategy is also seen in the microscopic
the hollow, curved radular teeth by contraction of a lower heterobranch Omalogyra.
PHYLUM MOLLUSCA 495
occur between anemone fishes and their host anemo-

nes), or perhaps that only immature nematocysts sur-
vive, to later undergo maturation in the dorsal cerata.
It may also be that, once the cnidocytes are digested,
the nematocysts' firing threshold is raised, thereby
preventing discharge. In any case, once in the cerata
the nematocysts are stored in structures called cnido-
sacs and presumably help the nudibranch to fend off
attackers. Discharge might even be under the control
of the host nudibranch, perhaps by means of pressure
exerted by circular muscle fibers around each cnidosac,
Some dorid nudibranchs also utilize their prev in
remarkable ways. Many dorids secrete complex toxic
compounds incorporated into mucus released from the
mantle surface. These noxious chemicals act to deter
potential predators. While some of these chemicals
may be manufactured in some dorids, in most cases it
appears that they are obtained from the sponges or ec-
toprocts on which they feed. Some species, such as the
"Spanish dancer" nudibranch (Hex abranchus sangui ne-
us),not only use a chemical from their prey (in this case
a sponge) for its own defense, but deposit some of the
noxious chemical on the egg mass, helping to protect
the embryos until they hatch.
In polyplacophorans there are generally 17 teeth
in each transverse row of the radula (a central tooth
flanked by eight on each side). MoSt chitons are herr:
bivorous gtazers. Notable exceptions are certain mem-
bers of the order Ischnochitonida (family Mopaliidae,
e.g., Mopalia, Placiphorella),which are known to feed on
both algae and sma1l invertebrates. Mopalia consumes
sessile invertebrates, such as barnacles, ectoprocts, and
hydroids. PlaciphoreIIacaptures live microinvertebrates
Figure 13.27 Sequenceof photographsof a cone shell (particularly crustaceans) by trapping them beneath
(Conus)capturing and swallowing a small fish. The pro- its head-flap , alarge anterior extension of the girdle
boscis is extendedand swept back and forth above the
(Figure 73.24C).
substratumin searchof prey; when a fish is encountered,
a poison-chargedtooth of the toxoglossanradulais fired In monoplacophorans the radula consists of a
like a harpoon;and the prey is quickly paralyzedand ribbonlike membrane bearing a succession of trans-
ingested. verse rows of 11 teeth each (a slender central tooth
flanked on each side by five broader lateral teeth).
Monoplacophorans are probably generalized deposit
Aeolid nudibranchs (Figure 1,3.7G)have a well-de- feeders that graze on minute organisms coating the
served reputation for their particular mode of feeding, substratum on which they live.
in which portions of their cnidarian prey are held by Cephalopods are predatory carnivores. Squids are
the jaws while the radula rasps off pieces for ingestion. some of the most voracious creatures in the sea, suc-
A similar mode of feeding has also been observed in cessfully competing with fishes. Octopuses are also
the caenogastropod family Epitoniidae. active camivores, preying primarily on crabs, bivalves
Many aeolid nudibranchs engage in a remarkable and gastropods. Some species of Octopusbore through
phenomenon called kleptocnidae. Some of the prey's the shells of molluscan prey in a fashion similar to that
nematocysts are ingested unfired, passed through the of gastropod drills. Some even drill and prey upon
nudibranch's gut, and eventually transported to lobes their close relatives, the chambered nautiluses. They do
of the digestive gland in dorsal finger-like extensions not use the radula to drill, but instead use a rasp-like
called cerata (singular, ceras) (Figure I3.32D,E). How projection formed from the salivary papilla.
the nematocysts undergo this transport without firing Using their impressive locomotor skills, most
is still a mystery. Popular hypotheses are that mucous cephalopods hunt and catch active prey. Some octo-
secretions by the nudibranch limit the discharge, or puses, however, hunt "blindly," by tasting beneath
that a form of acclimation occurs (like that suspected to stones with their highly sensitive suckers that are both
(A) Visceral mass (B) (c)

Exhalant chamber Afferent Exhalant
blood vessel chamber
Ctenidium
Afferent Efferent
blood vessel Mantle blood vessel
Efferent
Exhalant
blood vessel Ctenidium
chamber
Inhalant
Inhalant
charnber
chamber Inhalant
chamber
Figure 13-28 Arrangementof ctenidia in some bivalves

(transversesections)showingthe followingconditions:
(A) protobranch,(B) lamellibranch,(C)septibranch. e.9., Verffietzs; Turritellidae, e.g., Turritella) and the
freshwater Viviparidae (e.g., Viaipnrus). The ctenidial
filaments in these filter feeders are much elongated
mechano- and chemosensitive. Lr any event, once prey and the mantle waste rejection cilia have evolved into a
is captured and held by the arms, the cephalopod bites food collecting groove that runs to the mouth. The rad-
it with its homy beak (modified jaws) and injects a neu- ula in suspension-feeding gastropods is somewhat re-
rotoxin from modified salivary glands. This ability to duced, serving mainly to pull mucus-bound food into
quickly immobilize prey also helps prevent the soft- the mouth. Some feed entirely by suspension feeding,
bodied cephalopod from being engaged in a potential- others use browsing to supplement this method.
ly dangerous struggle. The wormlike shell of the vermetids is permanently
Suspension feeding evolved in autobranch bivalves, affixed to the substratum and while some adopt cili-
and also several times in gastropods, and in most of ary food collecting, others combihe this with mucolrs
these cases it involves modifications of the ctenidia net collecting or use the latter method exclusively. A
that enabled the animal to trap particulate matter car- special pedal gland in the reduced foot produces copi-
ried into the mantle cavity by incoming respiratory ous amounts of mucus that spreads into the water col-
water curuent. The lamellar nature of molluscan gills umn as a sticky plankton trap. Periodically the net is
exapted them for extracting suspended food particles. hauled in by the foot and pedal tentacles, and a new
Increasing the size of the gills and the degree of fold- one is quickly secreted. Thylacodesarenarius, a large
ing also increased the surface area available for trap- Mediterranean species, casts out individual threads
ping particulate material. In suspension feeders, at least up to 30 cm long, whereas the gregarious California
some of the gill and mantle cilia, which otherwise serve species Thylacodessquamigerus forms a communal net
to remove potentially clogging sediment from the man- shared by many individuals.
tle cavity (as pseudofeces), are preadapted to ftansport The radula apparently disappeared early in the
particulate matter from the gills to the mouth region. course of bivalve evolutiory and in living species there
While collecting food on the gills has been adopted is no trace of this structure or even the buccal cavity
by autobranch bivalves and some groups of gastro- that contained it. Most autobranch bivalves use their
pods, other methods have also been employed. In the large ctenidia for suspension feeding but the more
gltt-tess planktonic sea butterflies (pteropods), the cili- primitive bivalves in the subclass Protobranchia are
ated "wings" or parapodia used for swimming (Figure not suspension feeders but engage in a type of deposit-
13.7D,E) also function as food-collecting surfaces or feeding microphagy. Protobranchs live in soft marine
may cooperate with the mantle to produce large mu- sediments and maintain contact with the overlying
cous sheets that capture small zooplankton. From water either directly (e.g., Nucula) or by means of si-
the foot, ciliary currents carry mucus and food to the phons (e.9., Nuculana, Yoldia). The two ctenidia are
mouth. In some pteropods, the mucous sheet may be small, conforming to the primitive molluscan bipecti-
as large as 2 m across. Mucous sheet feeding is also em- nate plan of an elongated axis carrying a double row
ployed by a few other gastropods including the inter- of lamellae (Figures 13.28A and1,3.29). Protobranchs
tidal limpet-like Trimusculidae and the vermetids (see feed by means of two pairs of large labial palps flank-
below). Flowever, the majority of suspension feeding ing the mouth. The two innermost palps are the short
gastropods, including a few vetigastropods such as the labial palps, and the two outermost palps are formed
sand-beach trochids (Umbonium, Bankiain),the hot vent into tentacular processes called proboscides (each
neomphalid (Neomphnlus),and some marine caenogas- being called a palp proboscis), which can be extend-
tropods (Calyptraeidae, e.g., Crepidula; Vermetidae, ed beyond the shell (Figure 73.29).During feeding the
PHYLUM MOLLUSCA 497
area in the posterior mantle cavity to be discharged; the

Riglt outer labial palp
...'..'.,. spacesventral to the W's are inhalant and communicate
with the inhalant area of the mantle edge. Many other
Position of mouth
bivalves have eulamellibranch ctenidia, which are sim-
ilar to the filibranch design but neighboring filaments
Anterior
adductor are fused to one another by actual tissue junctions at
muscle numerous points along their length. This arrangement
results in interfilament pores that are rows of ostia
rather than the long narrow slits of filibranchs (Figure
13.30B,E,F).In additiorU the ascending and descending
Ctenidium
halves of some filaments may be joined by tissue bridg-
ii!
es thatprovide firmness and strength to the gill.
::ti Both filibranch and eulamellibranch ctenidia are used
to capture food. Water is driven from the inhalant to
the exhalant parts of the mantle cavity by lateral cilia all
along the sides of filaments in filibranchs, or by special
Right palp lateral ostial cilia in eulamellibranchs (Figure 13.30E-F).
proboscis Place where pseudofeces As the water passes through the interfilament spaces
are ejected
it flows through rows of frontolateral cilia, which flick
Figure 13.29 Feeding in the primitive bivalve Nucula particles from the water onto the surface of the filament
(Protobranchia). The clam is seen from the right side, in facing into the current. These feeding cilia are called
its natural position in the substratum (right valve and right compound cirri; they have a pinnate structure that prob-
mantle skirt removed). Arrows show direction of ciliary
ably increases their catching power. Mucus presumably
currents in the mantle cavity and on the palps. Water
plays some part in trapping the particles and keep-
currents are also shown in the (l) inhalant region and
(E) exhalant region. ing them close to the gill surface, although its precise
role is uncertain. Bivalve ctenidia are not covered with
a continuous sheet of mucus, as occurs in many other
proboscides are extended into the bottom sediments. suspension-feeding invertebrates (e.g., gastropods, tu-
Detrital material adheres to the mucus-covered sur- nicates, amphioxus). Once on the filament surface, par-
face of the proboscides and is then transported by cilia ticles are moved by frontal cilia toward a food groove on
to the labial palps, which function as sorting devices. the free edges of the ctenidium, and then anteriorly to
Low-density particles are carried to the mouth; heavy the labial palps. The palps sort the material by size and
particles arl carried to the palp margins and ejectei perhaps also by quality before passing the food to the
into the mantle cavity. mouth. Rejected particles fall off the gill or palp edges
Lr the suspension feeding subclass Autobranchia, lat- into the mantle cavity as pseudofeces. This "hltration"
eral cilia on the ctenidia generate a water current from of waterbybivalves is quite efficient. The American oys-
which suspended particles are gleaned. Increased ef- ter (Crassostreaairginica), for example, can process up to
ficiency is achieved by various ctenidial modifications. 37 liters of water per hour (at24'C), and can capture par-
The primary modification, seen in all living autobranch ticles as small as L pm in size. Studies on the common
bivalves, has been the conversion of the original, small, mussels Mytilus edulis and M. californianus suggest that
triangular plates into V-shaped filaments with exten- these bivalves maintain pumping rates of about 1 liter
sions on either side (Figures 13.288 and 13.308). The per hourper gram of (wet) body weight.
arm of this V-shaped filament that is attached to the cen- Members of the superfamily Tellinoidea (including
tral axis of the ctenidium is called the descending arm; Tellinidae and Semelidae) are deposit feeders, sucking
the arm forming the other half of the V is the ascend- up surface detritus with their long, mobile inhalant si-
ing arm. The ascending arm is usually anchored distally phon (Figure 13.20G) and using the large labial palps
by ciliary contacts or tissue junctions to the roof of the to pre-sort the particles before ingesting them.
mantle, or to the visceral mass. Taken together, the fwo Some members of the order Poromyata (Anomalo-
V-shaped filaments, with their double row of leaflets, desmata) are known as septibranchs and are sessile
form a W-shaped structure when seen in cross section. predators and, unlike other autobranch bivalves, their
Some pteriomorphian autobranch bivalves have gills are not used for feeding. Instead the ctenidia are
filibranch ctenidia (e.9., mussels) wherein adjacent very reduced and modified as a perforated but mus-
filaments are interlocked to one another by periodic cular septum that divides the mantle cavity into dor-
clumps of specialized cilia, leaving long narrow slits in sal and ventral chambers (Figures 13.28Cand 13.314).
between (interfilament spaces) (Figure 13.30C,D). The The muscles are attached to the shell such that the sep-
spaces between the arms of the W's are exhalant su- tum can be raised or lowered within the mantle cavity.
prabranchial chambers, which merge with the exhalant Raising the septum causes water to be sucked into the
(A) Exhalant
(B)
.-\-I i "il
\ t /, Tissue connection
rY
between adjacent Afferent
tl
Lateral ll ! t$ filaments blood vessel
tt vg
Connection of
+ -464
Afferent branchial vessel filament to
mantle
Efferent branchial vessel \Ctenidial
Efferent
Frontolateral Inhalant
blood
cilia sPace
vessel
Frontal cilia Lateral
(to ahium Descending arm of
of heart) ctenidial filament
Ctenidial
filament Inhalant
Ascending arm of sPace
ctenidial filament
Interfilament Exhalant space
sPace A
7-\- Food groove
Tissue connection between
llnr<- ascending and descending
I arms of same filament
Tissue (= interlarnellar junction)
junction
cirri cilia cilia
Figure 13.30 Ctenidial structure in bivalve molluscs. In all

drawings, solid arrows indicate the direction of water flow
(from inhalant space, between ctenidial filaments, to exhalant
space). (A) Section through part of the gill axis in a nuculanid
protobranch, showing four alternating filaments (leaflets) on
each side. Dashed arrows indicate direction of hemolymph
flow in the filament. (B) Highly schematic cutaway view show-
ing four ctenidial filaments, and their interconnections,on one
side of the body of a eulamellibranch. (C) Lateral view of four
ctenidial filaments of a filibranch. (D) Cross section through
ascending and descending arms of four filibranch ctenidial fila-
ments. (E) Lateral view of four filaments of a eulamellibranch. Food
(F) Cross section through ascending and descending arms of groove
four eulamellibranchctenidial filaments. (G) Ctenidial filaments
of the mussel Mytilus californianus showing ciliary junctions
and interfilament spaces. (H) Frontal ciliary tracts on ctenidial
filaments ot Mytilus. (l) Ventral gill edge ol Mytilus showing
food groove.
PHYLUMMOLLUSCA 499
Inhalant
siphon
Adductor muscle
Septal muscle
Sphincter of
Heart exhalant siphon
Stomach
Figure 13.31 Feeding in the septibranch bivalve

'/-Styt"
/ Cuspidaria (Anomalodesmata). (A) General anatomy
sac
of Cuspidaria rostrata. Arrows indicate water flow.
Septal muscle (B) Siphon and sensory siphonal tentacles protruding
from the substratum, but largely contracted. (C) Siphon
tor muscle extended, capturing a microcrustacean. (D) Details of
the siohons and tentacles.
mantle cavity by way of the inhalant siphon; lowering host's nukitional biology. One of the most interesting of
the septum causes water to pass dorsally through the these relationships exists between many molluscs and
pores into the exhalant chamber. These movements also sulfur bacteria. These molluscs appear to derive a por-
force hemolymph from mantle sinuses into the siphonal tion of their nutritional needs from symbiotic, carbon-
sinuses, thereby causing a rapid protrusion of the inhal- fixing sulfur bacteria, which usually reside on the host
ant siphon, which can be directed toward potential prey mollusc's gills. In some monoplacophorans (Laeaipilina
(Figure 13.31B-D). In this fashion, small animals such antartica) and gastropods (Lurifax aitreus, Hirtopeltn) the
as microcrustaceans are sucked into the mantle cavity, bacteria are housed in special cavities called bacterio-
where they are grasped by muscular labial palps and cytes in the mantle cavity. This mollusc-bacteria sym-
thrust into the mouth; at the same time, the mantle tis- biosis has been recently documented from a variety of
sue serves as the gas exchange surface. sulfide-rich anoxic habitats, including deep-sea hydro-
While most pteriomorphians are restricted to thermal vents, where geothermally produced sulfide
epibenthic life, lacking siphons (Figure 1.3.21.A,8), is present, and from other reduced sediments, where
many heterodont bivalves live buried in soft sedi- microbial degradation of organic matter leads to the re-
ments, where long siphons are utilized to maintain duction of sulfate to sulfide (e.g., anoxic marine basins,
contact with the overlying water (Figures 13.8,4.and seagrassbed and mangrove swamp sediments, pulp
13.20D-H). mill effluent sites, sewage outfall areas).
Scaphopods consume foraminifera and other meio- Members of some bivalve families, in particular
farsna taxa, diatoms, zooplankton, and interstitial detri- the Solemyidae and Lucinidae, harbor sulfur bacteria
tus. Two lobes flank the head, each bearing numerous in their enlarged gills which have the ability to direct-
(up to several hundred) long, slender tentacles called ly oxidize sulfide. They do this by means of a special
captacula (Figures 13.9 and 13.13F).The captacula are sulfide oxidase enzyme in the mitochondria. These bi-
extended into the substratum by metachronal beating valves inhabit reduced sediments where free sulfides
of cilia on the small terminal bulb. Within the sediment are abundant. The ability to oxidize sulfide not only
organic particles and microorganisms adhere to the provides the bivalve with a source of energy to drive
sticky terminal bulb; small food particles are transport- ATP synthesis, it also enables them to rid their body of
ed to the mouth by way of ciliary tracts along the ten- toxic sulfide molecules that accumulate in such habi-
tacles, while larger food items are transported directly tats. The nutrients obtained bv this svmbiosis are suffi-
to the mouth by muscular contraction of the captacula. cient for the bivalve so, in solemyids, the gut is reduced
A well-developed, large radula pulls the food into the or, in a few species, absent.
mouth, perhaps partially macerating it in the process. Another notable partnership exists between giant
Several forms of symbiotic relationships have clarns (Tridacna) and their syrnbiotic zooxanthellae (the
evolved within molluscs that are intimately tied to the dinoflagellate Symbiodinium). These clams live with
(A)
Sphincter Outer lip
Radular sac Buccalbulb

Esophagus
Salivary glands Salivary gland duct
Esophagealbulb
Esophagus
Gland of Leiblein
(modified esophageal gland)
Anus
Esophagus
Anal gland
Salivary gland
Gland cells
Digestive
olanrl
Branch of
digestive gland
Intestine Rhinophores
Sphincter
An
Branch of
Foregut gland digestive
gland
Anterior diverticulum
of 'stomach Epidermis
Salivary gland
Digestive Pharynx
-l^-t
6rdrru
their dorsal side against the substratum, and they ex- of these dinoflagellates in "carrier cells" associated with
pose their fleshy mantle to sunlight through the large their digestive glands. Experiments indicate that when
shell gape. The mantle tissues harbor the zooxanthellae. sufficient light is available, host nudibranchs utilize
Many specieshave special lens-like structures that focus photosynthetically fixed organic molecules produced
light on zooxanthellae living in the deeper tissues. A by the alga to supplement their usual diet of prey. The
few other bivalves and certain sea slugs also maintain a dinoflagellates are probably not transmitted with the
symbiotic relationship with Symbiodinium. Several spe- zygotes of the nudibranchs, each new generation thus
cies of Melibe, Pteraeolidia, and Berghia harbor colonies requiring reinfection from the environment. A number
(F) PHYLUMMOLLUSCA 501
Exhalant Intestine Position of internal
siphon opening of duct of (H) Lower jaw U ju*
digestive gland
Esophagus Buccalmass
Mouth Esophagus
Labial palps
Portion of
'intestine
housing style
Gonad
"Liver"
(digestive
gland)
"Pancreatic'l
"Pancreas"
Figure 13.32 Molluscan digestive system. (A) The digestive system of the
neogastropod (Muricidae). (B) The digestive system of the stylommatopho-
ran land snail Cornu. (C) The histology of the intestinal wall of the gastropod
Tonna. (D) A cladobranch nudibranch (Embletonra) in which large branches
of the digestive gland fill the dorsal cerata. (E) A longitudinal section of the
ceras of the aeolidid nudibranch Trinchesia showing the cnidosac where
nematocysts (not shown) from this animal's cnidarian prey are stored.
(F) Diagrammatic lateral view of the digestive tract and nearby organs of the
unionoid dam Anodonfa. (G) The digestive system of the cuttlefish Eledone.
(H) The digestive system of the squid Loligo.
of aeolid nudibranchs accumulatezooxanthellaefrom breaksdown celluloseand makesits oroducts avail-

their cnidarian prey. Some of the dinoflagellates end able to its host.Nitrogen-fixing bacteriaoccur aspart of
up inside cells of the nudibranch's digestivegland, but the gut flora in many animalswhose diet is rich in car-
many othersare releasedin the slug'sfeces,from where bon but deficient in nitrogen (e.g.,termites).However,
they may reinfect cnidarians. An even more remark- shipworms are the only animals known to harbor a ni-
able phenomenonoccursin somemembersof another trogen fixer as a pure (single species)culture in a spe-
group of seaslugs,the Sacoglossa(e.g.,Placobranchus). cialized organ (similar to the host nodule-Rhlzobium
Theseseaslugs obtain functional chloroplastsfrom the symbiosisof leguminousplants).
green algaeupon which they feed and incorporate them In addition to the above and myriad other feeding
into their ornmtissues;the chloroplastsremain active for strategiesof molluscs, some species(notably somebi-
a period of time and produce photosynthetically fixed valves and seaslugs)probably obtain a significantpor-
carbonmoleculesthat are utilized by the hosts. tion of their nutritional needsby direct uptake of dis-
Still another unusual symbiosis occurs between solved organic material from seawater,such as amino
an aerobic bacterium and the wood-boring marine acids.
shipworm bivalves (Teredinidae)(Figure 13.21D).
Shipworms are capable of living on a diet of wood Digestion
alone by harboring this cellulose-decomposing,nitro- Molluscs possesscomplete, or through guts, a few of
gen-fixing bacterium. The bivalve cultures the bacte- which are illustrated in Figure 13.32.The mouth leads
rium in a specialorgan associatedwith ctenidial blood inward to a buccal cavity, within which the radular
vesselscalled the gland of Deshayes.The bacterium apparatusand jaws (when present)are located (Fig-
(B)
(A) Gashic shield (on dorsal
stomach wall) Rotating
mucus-food
string
(protostyle) Intestine Muscle
Crystalline style
layer
Narrow
connection Style sac
of st;lle sac
rystalline style
with
particles intestine Ciliated
'-1;,,l.t;;"'
Ciliated From epidermal cells
sorting mouth
area Connective tissue
Typhlosole Digestive gland
Figure 13.33 The molluscan stomach and style sac. contains a crystalline style (Figure 13.33).This struc-
(A) A generalized stomach and style apparatus of an
ture, which functions to aid in digestion, is a rodlike
autobranch bivalve. The crystalline style rotates against
the gastric shield, releasingdigestive enzymes and wind- matrix of proteins and enzymes (often amylase) that
ing up the mucus-food string to assist in pulling it from are slowly released as the projecting end of the style ro-
the esophagus. Food particles are sorted in the ciliated, tates and grinds against the gastric shield that protects
grooved soding area: small particles are carried (in part by the otherwise delicate stomach wall. The gastric cilia
the typhlosole) to the digestive glands for digestion; large and rotating style wind up the mucus and food into a
particles are carried to the intestine for eventual elimina-
string and draw it along the esophagus to the stomach.
tion. (B) A cross section of a style sac.
The style is produced by special cells of the style sac.
The style of some bivalves is enormous, one-third to
one-half the length of the clam itself. Particulate matter
ure 13.23).The esophagus is generally a straight tube is swept against the stomach's anterior sorting region,
connecting the foregut to the stomach. Various glands which sorts mainly by size. Small particles are carried
are often associated with this anterior gut region, in- into the digestive glands, which arise from the stomach
cluding some that produce enzymes and others that se- wall. Larger particles are passed along ciliated grooves
crete a lubricant over the radula that are usually called of the stomach to the intestine. In more primitive bi-
salivary glands. In many herbivorous species (e.g., cer- valves (Protobranchia) and in many gastropods a crys-
tain eupulmonates, anaspideans lAplysia], and some talline style is absent but a style sac is often present,
cephalaspideans), a muscular gizzard, (unrelated to the which contains a rotating mass of mucus mixed with
jaws) may be present for grinding up tough vegetable particles that is termed a protostyle.
matter. The gizzard may have chitinous, or even cal- Extracellular digestion takes place in the stomach
careous plates or teeth. The stomach usually bears one and lumina of the digestive glands, while absorp-
or more ducts that lead to the large glandular digestion and intracellular digestion occur in the diges-
tive gland (variously called the digestive diverticula, tive gland cells and the intestinal walls. Extracellular
digestive caeca,midgut glands, liver, or other similar digestion is accomplished by enzymes produced in
terms). Several sets of digestive glands may be present. foregut (e.g., salivary glands, esophageal pouches or
The intestine leaves the stomach and terminates as the glands, pharyngeal glands-sometimes called "sugar
anus, which is typically located in the mantle cavity in glands" because they produce amylase), the stomach,
or near the exhalant water flow. and the digestive glands. In primitive groups, intracel-
Once food has entered the buccal cavity of most lular digestion tends to predominate. In Solenogastres
molluscs, it is carried in mucous strings into the esoph- all digestive functions are accomplished in a uniform
agus and then to the stomach. In cephalopods and midgut lined by voluminous digestive and secretory
some predatory gastropods, chunks of food or whole cells. In most molluscs, ciliated tracts line the digestive
prey are swallowed by muscular action of the esopha- glands and carry food particles to minute diverticula,
gus. The food is stored in the stomach or in an expand- where they are engulfed by phagocytic digestive cells
ed region of the esophagus called the " crop," as in oc- of the duct wall. The same cells dump digestive wastes
topuses andNautilus, and many gastropods. In many back into the ducts, to be carried by other ciliary tracts
bivalves and gastropods, the stomach wall bears a chi- back to the stomach, from there to be passed out of the
tinous gastric shield and a ciliated, ridged sorting area gut via the intestine and anus as fecal material. In most
(Figure 13.33).The posterior stomach region (anterior highly-derived groups (e.g., cephalopods and many
in gastropods) is a style sac, which is lined with cilia gastropods), extracellular digestion predominates.
and in autobranch bivalves and in some gastropods Enzymes secreted primarily by the digestive glands
PHYLUMMOLLUSCA 503
Nepfuostome
Figure 13.34 Hemolymph flow in a typical mollusc.

Oxygenated hemolymph is pumped from the ventricle to
the hemocoel, where it bathes the organs; eventually it
Mantle cavity drains into various channels and sinuses and then into
the afferent branchial vessels, which enter the ctenidia.
Ctenidium Oxygen is picked up in the ctenidia, and the hemolymph
is then transported by the efferent branchial vessels to the
left and right atria, through the ventricle, and then back
and stomach digest the food, and absorption occurs in to the hemocoel. Additional auxiliary pumping vessels
the stomach, digestive glands, and intestine. occur in several taxa, particularly in active groups such as
cephalopods.
Circulation and Gas Exchange
Although molluscs are coelomate protostomes, the
coelom is greatly reduced. The main body cavity is, in circulation is shown diagrammatically in Figure 13.34,
most species, an open circulatory space or hemocoel, although it is modified to various degreesin different
which comprises several separate sinuses, and a net- classes(Figure13.35).In somecephalopodsthe circula-
work of vessels in the gills, where gas exchange takes tory systemis secondarilyclosed(Figure 13.35C).
place. The blood of molluscs contains various cells, in- Most molluscshave ctenidia.But many have lost the
cluding amebocytes, and is referred to as hemolymph. ctenidia and rely either on secondarilyderived gills or
It is responsible for picking up the products of diges- on gasexchangeacrossthe mantle or generalbody sur-
tion from the sites of absorption and for delivering face.In the primitive condition the ctenidium is built
these nutrients throughout the body. It usually carries around a long, flattened axis projecting from the wall
in solution the copper-containing respiratory pigment of the mantle cavity (Figure13.304).To eachside of the
hemocyanin. Some molluscs use hemoglobin to bind axis are attachedtriangular or wedge-shapedfilaments
oxygen and many have myoglobin in the active muscle that alternate in position with filaments on the oppo-
tissues, in particular those of the odontophore. site side of the axis (exceptin nucuid protobranchs,
The heart lies dorsally within the pericardial cham- where they are opposite).This arrangement,in which
ber, and includes apaft of atria (often called auricles) filaments project on both sides of the central axis, is
and a single ventricle. In monoplacophorans and in called the bipectinate condition. Thereis one gill on
Nautilus there are two pairs of atria, and in many gas- eachside of the mantle cavity, sometimesheld in posi-
tropods there is only one (the left) that corresponds to tion by membranesthat divide the mantle cavity into
the single gill. The atria receive the efferent branchial upper and lower chambers(Figure 13.28A,8).Lateral
vessels, drawing oxygenated hemolymph from each cilia on the gill draw water into the inhalant (ventral)
ctenidium and passing it into the muscular ventricle, chamber,from which it passesupward between the
which pumps it anteriorly through a large anterior gill filamentsto the exhalant(dorsal)chamberand then
artery (the anterior or cephalic aorta). The anterior out of the mantle cavity (Figure 13.304).
artery branches and eventually opens into various si- Two vesselsrun through eachgill axis. The afferent
nuses within which the tissues are bathed in oxygen- vesselcarriesoxygen-depletedhemolymph into the
ated hemolymph. Return drainage through the sinuses gill, and the efferent vesseldrains freshly oxygenated
eventually funnels the hemolymph back into the affer- hemolymph from the gill to the atria of the heart, as
ent branchial vessels. This basic pattern of molluscan noted above.Hemolymph flows through the filaments
(A) Branchial vessels
Afferent branchial vessel Cephalic aorta Figure 13.35 The circulatory systems
of three molluscs. (4 A neogastropod
Efferent branchial vessel Nephridiocardiac vein
Buccinum (hemolymph sinuses not
shown). (B) A eulamellibranchunionid
Osphradium Digestive bivalve. (C) A squid, Loligo.
Siphon gland
Pallial artery-
Mantle margin
from the afferent to the efferent ves-
sel. Ctenidial cilia move water over the
gill filaments in a direction opposite to
Ventricle
that of the flow of the underlying he-
Visceral aorta
Pedal artery molymph in the branchial vessels. This
Operculum
countercurrent phenomenon enhances
gas exchange between the hemolymph
and water by maximizing the diffusion
Foot gradients of O, and CO, (Figure 13.30,4').
These presumed primitive bipectinate
Ventricle ctenidial gill conditions are expressed
Stomach Pericardialchamber in several living groups, for example, in
Caudofoveata, chitons, protobranch bi-
valves, and some gastropods.
Efferent As a result of torsion, gastropods
branchial evolved novel ways to circulate water
vessel
over the gills before it comes into con-
tact with gut or nephridial discharges.
Ctenidium
Some vetigastropods with two bipec-
tinate ctenidia may accomplish this
Artery to mantle by circulating water in across the gills,
then past the anus an d nephr idio-
Afferent branchial
Hemolymph sinus of toot pore, and away from the body via slits
uJr, from foot vessel or holes in the shell. This circulation pattern is used
(c) by the slit shells (Pleurotomariidae) and the minute
Scissurellidae and Anatomidae (Figure 13.36),abalones
(Haliotidae) (Figure 13.1D), and volcano (or keyhole)
limpets (Fissurellidae) (Figures 13.16H,I and 13.25,{).
Anterior, or Anterior, or cephalic,
artery (= anterior aorta)
Some specialists regard the Pleurotomariidae as "liv-
cephalic,vein ing fossils" that reflect an early gastropod character
state since gastropods bearing slits are found amongst
early gastropod fossils. Most other gastropods have
Efferent lost the right ctenidium and with it the right atrium;
Left branchial vessel
inhalant water enters on the left side of the head and
ctenidium Afferent
then passes through the mantle cavity and straight out
branchial vessel
the right side, where the anus and nephridiopore open.
Other gastropods have lost both ctenidia and utilize
Left branch Anterior
of cephalic mantle vein secondary respiratory regions, either the mantle sur-
vein Ink sac artery face itself, expanded nephridial surfaces, or secondari-
ly derived gills of one kind or another. Limpets of the
genus Patellahave rows of secondary gills in the mantle
Right branchial groove along each side of the body, superficially simi-
(accessory)heart lar to the condition seen in chitons where multiple
Artery to ctenidia are found.
branchial heart
In many gastropods one ctenidium is lost, e.g.,
l- L"ft at ir* Posterior aorta
Heart patellogastropods, some vetigastropods, all neriti-
I Ven Lateral morphs and caenogastropods. In caenogastropods,
mantle artery
Posterior the dorsal and ventral suspensory membranes seen
mantle vein Median in vetigastropod ctenidia are absent and the gill is at-
mantle artery
tached directly to the mantle wall by the gill axis. The
gill filaments on the attached side have been lost, while
PHYLUM MOLLUSCA 505
In chitons the mantle cavity is a groove extending

along the ventral body margin and encircling the foot
(Figure 13.48). A large number of small bipectinate
ctenidial gills lie laterally in this groove. The mantle
is held tight against the substratum, largely enclosing
Pallial tentacle
Epipodial tentacles
Slit band
Epipodial tentacles
Figure 13.36 The slit-bearing vetigastropod Anatoma.
those of the opposite side project freely into the mantle

cavity. This arrangement of filaments on only one side (B)
of the central axis is referred to as the monopectinate
(or pectinobranch) condition (Figure 13.14D). Some
caenogastropods have evolved inhalant siphons by
extension and rolling of the anterior mantle margin
(Figures 13.1E and 13.404). In these casesthe margin
of the shell may be notched, or drawn out as a canal to
house the siphon. The siphon provides accessto sur-
face water in burrowing species, and may also function
as a mobile, directional organ used in conjunction with
the chemosensory osphradium.
All heterobranchs have lost the tvpical ctenidia
but some have a plicate, or folded, giiithat has been
considered by some to be a reduced ctenidium, but is
now considered to be a secondary structure that has
reformed in much the same location as the original
ctenidial gill. Trends toward detorsion, loss of the shell,
and reduction of the mantle cavity occur in many het-
erobranchs, and the process has apparently occurred
several times within this group. Some nudibranchs
have evolved secondary dorsal gas exchange struc-
tures called cerata or, in some nudibranchs, secondary
gills that surround the anus (Figures 13.7F-D.
\A/holly terrestrial gastropods lack gills and exchange
gases directly across a vascularized region of the man-
tle, usually within the mantle cavity, the latter arrange-
ment usually referred to as a lung. In marine, fresh-
water and terrestrial eupulmonates, the edges of the
mantle cavity have become sealed to the back of the ani-
mal except for a small opening on the right side called
a pneumostome (Figure 13.37A) that is controlled by Figure 13.37 Land snails (Heterobranchia:
Eupufmonata). (A) A terrestrial slug (Arion lusitanicus),
a sphincter muscle (except in siphonariid limpets).
showing the pneumostome that opens to the "lung"
Instead of having gills, the roof of the mantle cavity is (Stylommatophora: Limacidae). (B) Terrestrial helicid snails
highly vasculaized. By arching and flattening the man- (Stylommatophora: Helicidae) during summer dormancy in
tle cavity floor, air is moved into and out of the lung. Sicily.
506 C h a p te r T h i rte e n
this groove except on either side at the anterior end to most cephalopods, the pumping of blood into the
form incurrent channels, and in one or two places at ctenidia is assisted by muscular accessory branchial
the posterior end to form excurrent areas. Water enters hearts, which boost the low venous pressure as the he-
the inhalant region of the mantle groove lateral to the molymph enters the gills. The gills are not ciliated and
gills, then passes between the gills into the exhalant their surface is highly folded, increasing their surface
region along the sides of the foot. Moving posteriorly, area for greater gas exchange necessary to meet the de-
the current passes over the gonopores, nephridiopores, mands of their high metabolic rate.
and anus before exiting (Figure 13.48). In the Solenogastres, gills are absent but the mantle
In bivalves the capacious mantle cavity allows the cavity surface may be folded or form respiratory pa-
ctenidia to develop a greatly enlarged surface area, pillae. Caudofoveates have a single pair of bipectinate
serving in most autobranch species for both gas ex- ctenidia in the mantle cavity. Monoplacophoran gills
change and feeding. Maoy of the morphological modi- are well-developed but weakly muscular and ciliated,
fications of bivalve gills are described above in discus- and only have lamellae on one side of the gill axis; they
sion of suspension feeding. In addition to the folded, occur as three to six pairs, aligned bilaterally within
W-shaped ctenidial filarnents seen in many bivalves the mantle groove. The gills of monoplacophorans are
(Figure 1.3.288),some forms (e.g., oysters) have pli- thought to be modified ctenidia that vibrate and venti-
cate ctenidia. A plicate ctenidium has vertical ridges or late the groove where gas exchange occurs.
folds, each ridge consisting of several ordinary ctenid-
ial filaments. So-called "principal filaments" lie in the Excretion and Osmoregulation
grooves between these ridges and their cilia are impor- The basic excretory structures of molluscs are paired
tant in sorting particles from the ventilation and feed- tubular nephridia (often called kidneys) that are primi-
ing currents. The plicate condition gives the ctenidium tively similar to those of annelids. Typical nephridia
a corrugated appearance and further increases the sur- are absent in the aplacophoran groups. Three, six,
face area for feeding and gas exchange. or seven pairs of nephridia occur in monoplacopho-
In spite of these modifications, the basic system of rans, two pairs in the nautiloids, and a single pair in
circulation and gas exchange in bivalves is similar to all other molluscs (except where one is lost in higher
that seen in gastropods (Figure 13.358). In most bi- gastropods) (Figure 73.74). The nephrostome typically
valves, the ventricle of the heart folds around the gut, opens into the pericardial coelom via a renopericardial
so the pericardial cavity encloses not only the heart duct, and the nephridiopore discharges into the mantle
but also a short section of the digestive tract. The large cavity, often near the anus (Figures 13.14and 13.34).In
mantle lines the interior of the valves and provides molluscs, the pericardial fluids (primary urine) pass
an additional surface area for gas exchange, which in through the nephrostome and into the nephridium,
some groups may be as important as the gills in this where selective resorption occurs along the tubule wall
regard. For example, in lucinid bivalves where the gills until the final urine is ready to pass out the nephrid-
are full of symbiotic bacteria, folds on the mantle act as iopore. The pericardial sac and heart wall act as selec-
a secondary gill, and in septibranchs, which have very tive barriers between the open nephrostome and the
reduced gills, the mantle surface is the principal area of hemolymph in the surrounding hemocoel and in the
gas exchange. heart. Mollusc nephridia are rather large and saclike,
Most autobranch bivalves lack respiratory pigments and their walls are often greatly folded. In many spe-
in the hemolymph, although hemoglobin occurs in a cies, afferent and efferent nephridial vessels carry he-
few families and hemocyanin is found in protobranchs. molymph to and from the nephridial tissues (Figure
Scaphopods have lost the ctenidia,heart, and vir- 13.38).Sometimes a bladder is present just before the
tually all vessels. The circulatory system is reduced to nephridiopore and sometimes a ureter forms a duct to
simple hemolymph sinuses, and gas exchange takes carry urine well beyond the nephridiopore.
place mainly across the mantle and body surface. A In many molluscs urine formation involves pres-
few ciliated ridges occur in the mantle cavity that may sure filtration, active secretion, and active resorption.
assist in maintaining water flow. A few tiny gastropods Aquatic molluscs excrete mostly ammonia, and most
and at least one small monoplacophoran species lack a marine species are osmoconformers. In freshwater
heart altogether. species the nephridia are capable of excreting a hypo-
No doubt associated with their large size and active somotic urine by resorbing salts and by passing large
lifestyle, cephalopods have a more developed circu- quantities of water. Terrestrial gastropods conserve
latory system than other molluscs, and in the highly water by converting amrnonia to uric acid. Land snails
active decapodiforms (squid and cuttlefish) it is effec- are capable of surviving a considerable loss of body
tively closed, with many discrete vessels, secondary water, which is brought on in large partby evapora-
pumping structures, and capillaries (Figures 13.11C, tion and the production of the metabolically expensive
13.728, and 13.35C). The result is increased pressure slime trail. They often absorb water from the urine in
and efficiency of hemolymph flow and delivery. In the ureter. In many gastropods (e.g., neritimorphs,
PHYLUMMOLLUSCA 507
Figure 13.38 The nephridium and nearby

organs ol Littorina (cutaway view). The
nephridial sac has been slit open.
way of renopericardial canals and empty

via nephridiopores into the mantle cav-
ity. However, the nephridia bear en-
larged regions called renal sacs. Before
reaching the branchial heart, a large vein
passes through the renal sac, wherein nu-
merous thin-walled evaginations, called
renal appendages, project off the vein. As
the branchial heart beats, hemolymph is
drawn through the renal appendages, and
caenogastropods, and heterobranchs), torsion is ac- wastes are filtered acrosstheir thin walls into the ne-
companied by loss of the adult right nephridium; in phridia. The overall result is an increasein excretoryef-
neritimorphs and caenogastropods, a small remnant ficiency over the simpler arrangementpresentin other
contributes to part of the gonoduct. Some gastropods molluscs.
have lost the direct connection of the nephrostome to The fluid-filled nephridia of cephalopodsare inhab-
the pericardial coelom. In such cases the nephridium ited by a variety of commensalsand parasites.The epi-
is often very glandular and served by afferent and ef- thelium of the convoluted renal appendagesprovides
ferent hemolymph vessels, and wastes are removed an excellentsurfacefor attachment and the renal pores
largely from the circulatory fluid. provide a simple exit to the exterior. Symbiontsidenti-
In bivalves, the two nephridia are located beneath fied from cephalopodnephridia include viruses,fungi,
the pericardial cavity and are folded in a long U-shape. ciliate protists,rhombozoans,trematodes,larval ces-
In autobranch bivalves, one arm of the U is glandular todes,and juvenile nematodes.
and opens into the pericardial cavity; the other arm
often forms a bladder and opens through a nephridio- Nervous System
pore in the suprabranchial cavity. hr protobranchs, the The molluscan nervous system is derived from the
unfolded walls of the tube are glandular throughout. basicprotostome plan of an anterior circumenteric ar-
The nephridiopores may be separate from or joined rangementof ganglia and paired ventral nerve cords.
with the ducts of the reproductive system. In the latter In molluscs,the more ventral and medial of the two
case, the openings are urogenital pores. pairs of nerve cords are called the pedal cords (or
In patellogastropods and vetigastropods and some ventral cords); they innervate the musclesof the foot.
other molluscs, the gonoduct fuses with the renoperi- The more lateral pair of nerves are the visceral cords
cardial canal, and the nephridiopore functions as a uro- (or lateral cords);they servethe mantle and viscera.
genital pore and discharges both excretory wastes and Transversecommissuresinterconnecttheselongitu-
gametes. In some cases/as in one monoplacophoran, dinal nerve cord pairs, creating a ladderlike nervous
a few bivalves and in some vetigastropods, the uro- system. This basic plan is seenin the aplacophorans
genital pore may become glandular. L:r many bivalves and polyplacophorans(Figure 13.39).The molluscan
and chitons the nephridium and gonad have separate nervous systemlacks the segmentally arranged ganglia
ducts. of annelids and arthropods.
In monoplacophorans and chitons, the nephridia In the "simplest" molluscs-such as aplacophorans,
open into the exhalant regions of the mantle grooves; in monoplacophorans,and polyplacophorans-ganglia
scaphopods, the paired nephridia open near the anus. are poorly developed (Figure 13.39).A simple nerve
In most gastropods the nephridiopores open directly ring surrounds the anterior gut, often with small ce-
into the mantle cavity but in some, such as in stylom- rebral ganglia on either side. Each cerebral ganglion,
matophoran pulmonates, there is an elongate ureter or the nerve ring itself, issuessmall nerves to the buc-
that opens outside the enclosed lung (mantle cavity). cal region and gives rise to the pedal and the visceral
Cephalopods retain the basic nephridial plan, in nerve cords. Most other molluscs have more well-de-
which the nephridia drain the pericardial coelom by fined ganglia. Their nervous systemsare built around
(A) (B)
Cerebropedal Cerebral Dorsal buccal comrnissure
Ventral buccal
commlssure
Buccal ganglion
Buccal commissureI Circumenteric
Buccalganglion nerve ring Cerebrobuccalring
_.f
Pedal ganglion Subradular connective
reil 1= venhal) Visceral (: lateral)

nerve cords nerve cord
Visceral (= lateral)
nerve cords Pedal nerve cord
Pedal commissures
Figure 13.39 Aculiferan mollus-
can nervous systems. (A) Class
Pedolateralcommissures Solenogastres, Proneomenia.
(B) Class Polyplacophora,
Terminal ganglion Acanthochitona.
of pedal cords
Pedolateralcommissu
Parietal ganglion
Suprarectalcommissure
three pairs of large ganglia that interconnect to form a to two pairs of nerve cords, one extending posterodor-
partial or complete nerve ring around the gut (Figures sally to the visceral ganglia, the other leading ventrally
L3.40 and 13.41).Two pairs, the cerebral and pleural to the pedal ganglia (Figure 73.47). The two cerebro-
ganglia,lie dorsal or lateral to the esophagus, and one pleural ganglia are joined by a dorsal commissure over
pair, the pedal ganglia, lies ventral to the gut, in the the esophagus. The cerebropleural ganglia send nerves
anterior part of the foot. In cephalopods, bivalves, and to the palps, anterior adductor muscle, and mantle.
advanced gastropods, the cerebral and pleural ganglia The visceral ganglia issue nerves to the gut, heart, gills,
are often fused. From the cerebral ganglia, peripheral mantle, siphon, and posterior adductor muscle.
nerves innervate the tentacles, eyes/ statocysts, and The degree of nervous system development within
general head surface, as well as buccal ganglia, with the Cephalopoda is unequaled among invertebrates.
special centers of control for the buccal region, radu- The paired ganglia seen in other molluscs are not rec-
lar apparatus, and esophagus. The pleural ganglia give ognizable in cephalopods, where extreme cephaliza-
rise to the visceral cords, which extend posteriorly, tion has concentrated ganglia into lobes of a large brain
supplying peripheral nerves to the viscera and mantle. encircling the anterior gut (Figure 13.42A).In addition
The visceral cords eventually join a pair of esophageal to the usual head nerves originating from the dorsal
(= intestinal, = pallial) ganglia and from there pass on part of the brain (more or less equivalent to the cere-
to terminate in paired visceral ganglia. The esophageal bral ganglia), a large optic nerve extends to each eye
ganglia or associated nerves innervate the gills and os- via a massive optic lobe. In most cephalopods, much
phradium, and the visceral ganglia serve organs in the of the brain is enclosed in a cartilaginous cranium. The
visceral mass. The pedal ganglia also give rise to a pair pedal lobes supply nerves to the funnel, and anterior
of pedal nerve cords that extend posteriorly and pro- divisions of the pedal ganglia (called brachial lobes)
vide nerves to muscles of the foot. send nerves to each of the arms and tentacles. an ar-
As described above, due to torsion, the posterior rangement suggesting that the funnel and tentacles
portion of the gastropod nervous system is twisted are derived from the molluscan foot. Octopuses may
into a figure eight, a condition known as streptoneury be the "smartest" invertebrates, for they can be quickly
(Figure 13.404,8). In addition to twisting the nervous taught some rather complex memory-dependent tasks.
system, torsion brings the posterior ganglia forward. In Squid and cuttlefish (Decapodiformes) have a rapid
many advanced gastropods this anterior concentration escape behavior that depends on a system of giant
of the nervous system is accompanied by a shortening motor fibers that control powerful and svnchronous
of some nerve cords and fusion of ganglia. In most de- contractions of the mantle muscles. The command cen-
torted gastropods the nervous system displays a sec- ter of this system is a pair of very large first-order giant
ondarily derived bilateral symmetry and more or less neurons in the lobe of the fused visceral ganglia. Here,
untwisted visceral nerve cords-a condition known as connections are made to second-order giant neurons
euthyneury (Figure 13.40C). that extend to a pair of large stellate ganglia. At the
In bivalves, the nervous system is clearly bilateral, stellate ganglia, connections are made with third-order
and fusion has usually reduced it to three large, dis- giant neurons that innervate the circular muscle fibers
tinct ganglia. Anterior cerebropleural ganglia give rise of the mantle (Figure 73.42D). Other nerves extend
(B) Tentacular nerve
tentacle
ganglion Left cerebral -Cephalic
ganglion
Pleural ganglion Left pleural
Right cerebral ganglion
ganglion
Pedal ganglion t pleural ganglion
Pedal ganglion
Esophaguscut in two places
Supraesophageal
ganglion Cut edge of mantle
Subesophageal
ganglion
Statocyst
Subesophageal
Osphradium
ganglion
Digestive
srp gland Esophagus
Rectum
ganglion
Pallial oviduct
Nephridium
Ovary
Nepfuidiopore
Digestive gland
Figure 13.40 The nervous system of some gas-

Buccal ganglion
tropods. (A) Arrangement of the nervous system
in a toded neogastropod. Note the location of the
Pedal
major ganglia and nerve cords. (B) Nervous system
ganglion of the torted terrestrial caenogastropod Pomatias
(Littorinimorpha) seen in dissection. Note the lack
of a ctenidium. (C) The nervous system of the euo-
pisthobranch, Akera.
aesophageal
ganglion
Visceral ganglion
Subesophageal
ganglion
posteriorly ftom the brain and terminate in various

Stomach
ganglia that innervate the viscera and structures in Anterior Dorsal
adductor commissute
the mantle cavity.
muscle Posterioradductor muscle
For several decades neurobiologists have uti-
Anus
lized the giant axons of Loligo as an experimental
Exhalant siphon
system for the study of nerve physiology and me-
chanics, and much of our fundamental knowledge
of how nerve cells work is based on squid neurol-
Visceral
ogy. The sea hare Aplysia, and some eupulmonate ganglia
snails have also been used in the same fashion and,
although they lack giant axons, they possessexcep-
tionally large neurons and ganglia that canbe eas- siphon
ily impaled with microelectrodes to discover the
physiological secrets of such systems.
Sense Organs
With the exception of the aplacophorans, mol-
luscs possess various combinations of sensory ten- Figure'13.41 The reduced and concentrated nervous sys-
tacles, photoreceptors, statocysts, and osphradia. tem of a typical autobranch bivalve.
510 ChapterThirteen
lobe Nerves to hind

part of body
Optic gland
Optic tract
Vertical lobe
Frontal lobe Supraesophageal
Superior complex
Nerves to
buccal region buccal lobe
Esop
Optic nerves (cut)

complex
Statocystof left side
nerves to arms Brachial lobe

(c) Tentacle
Brachial nerves
Buccal nerves rior buccal ganglion
Optic lobe
Esophagealganglia Brachial ganglion
Optic tract Cerebral ganglion
lobe Olfactory Pallial nerve
Optic nerves (= giant nerve)
Stellatenerve ganglion
Stellateganglion
isceralnerve
Branchial ganglion
isceral ganglion
Figure 13.42 The highly developed

nervous system of cephalopods. (A) The
brain of an octopus. The lobes of the (D)
supraesophageal complex approximately First-order Second-order Third-order
correspond to the cerebral and buccal giant nerve cells giant cells giant cells
ganglia of other molluscs while the sube- Proximal (finer)
Distal (thicker)
.sophageal complex comprises the fused third-order axons
third-order axons
pedal and pleurovisceral ganglia. About
15 structurally and functionally distinct
pairs of lobes have been identified in the
brain of octopuses. (B) Nervous system
of an octopus. (C) Nervous system of a
squid (Lolrgro).(D) Giant fiber system of
a squid. Note that the first-order giant
neurons possess an unusual cross con-
nection, and that the third-order giant
iovisceral Stellate ganglion
neurons are arranged so that motor
impulses can reach all parts of the man-
tle-wall musculature simultaneously (as
a result of the fact that impulses travel
faster in thicker axons).
PHYLUM MOLLUSCA 511
(A) Osphradia are patches of sensory epithelium, located

on or near the gills, or on the mantle wall (Figures
Osphradial
13.408 and 13.43A,B). They are chemoreceptors, and
leaflet
their cilia can also assist in mantle cavity ventilation in
some caenogastropods. Little is known about the biol-
ogy of osphradia, and their morphology and histology
ganglion differs markedly within the phylum and even within
Epithelial cells some classessuch as the gastropods.
In vetigastropods, a small osphradium is present
on each gill; in those gastropods that possess one gill,
there is only one osphradium, and it lies on the mantle
Ciliary tufts--___r_
cavity wall anterior and ventral to the attachment of
the gill itself. Osphradia are reduced or absent in gas-
tropods that have lost both gills, that possess a highly
reduced mantle cavity, or that have taken up a strict-
Lateral ciliated ly pelagic existence. Osphradia are best developed in
zone benthic predators and scavengers, such as neogastro-
pods and some other caenogastropods.
Most gastropods have one pair of sensory cephalic
(c) tentacles, but eupulmonates and many sea slugs pos-
Megaesthetes sesstwo pairs. Many vetigastropods also have epipo-
Shell eye.
dial tentacles on the margin of the foot or mantle and
Microaesthete there are also epipodial sense organs present (Figure
13.5A,C). The cephalic tentacles may bear eyes as well
as tactile and chemoreceptor cells. Many nudibranchs
have a pair of branching or folded anterior dorsal che-
moreceptors called rhinophores (Figure I3.7F,G).
Aesthete
The primitive patellogastropods have simple pig-
canal
ment-cup eyes, while the more advanced gastropods
rmls
(mantle) have more complex eyes with a lens and often a cor-
Articulamentum nea (Figure 13.444,8,D). Most gastropods have a small
eye at the base of each cephalic tentacle, but in some,
(D) such as the conch Strombusand some neogastropods,
Pigment Cornea the eyes are enlarged and elevated on long stalks. The
stylommatophoran and systellommatophoran pulmo-
Lens
nates also have eyes placed on the tips of special optic
Fibrillar area tentacles and, in stylommatophorans, these tentacles
have become olfactory organs.
Gastropods typically produce a mucopolysaccha-
ride slime trail as they crawl. In many species the trail
contains chemical messengers that other members of
the species "read" by means of their excellent chemo-
reception. These chemical messengers may be simple
Fiber bundle trail markers, so one animal can follow or locate anoth-
of aesthetescord er, or they may be alarm substances that serve to warn
others of possible danger on the path ahead. For ex-
ample, when the carnivorous cephalaspidean sea slug
Figure 13.43 Two sensory organs of molluscs: osphra- Naaanax is attacked by a predator, it quickly releases a
dia and aesthetes. (A) Cross section of a bipectinate yellow chemical mixture on its trail that causes other
osphradium of the caenogastropod Ranella showing two members of the species to abort their trail-following ac-
leaflets. (B) Part of the osphradium of a littorinimorph tivity. Laboratory experiments have shown that at least
caenogastropod such as Littorina. (C) One valve of a poly- one nudibranch (Tritonia diomedea)possessesgeomag-
placophoran (Tonicia). The aesthetes extend to the shell
netic orientation to the Earth's magnetic field. Motile
surface through megalopores and micropores. (D) Eye-
gastropods usually possessa pair of closed statocysts
bearing aesthetes (longitudinal section) in a megalopore of
a chiton (Acanthopleura). near the pedal ganglia in the anterior region of the foot
that contain either a single large statolith or several sta-
toconia (much smaller particles).
512 ChaoterThirteen
Scaphopods lack eyes, tentacles, and osphradia typi- ning of optic cup
cal of the epibenthic and motile molluscan groups. The
captacula may function as tactile (as well as feeding)
structures. Sense organs are found in the mantle edge Photoreceptor cells
surrounding the ventral aperture and at the dorsal
water intake opening.
Bivalves have ?nost of their sensory organs along Pigment cells
the middle lobe of the mantle edge where they are in
contact with the external environment (Figure 13.15C).
These receptors may include mantle tentacles, which
can contain both tactile and chemoreceptor cells. Such
tentacles are conunonly restricted to theiiphonal areas,
but in some swimming clams (e.g., Lima, Pecten)they
may line the entire mantle margin. Paired statocysts
usually occur in the foot near the pedal ganglia, and are
(E)
Epidermis
Epidermis
Pigment layer
Ciliary
Optic nerves
Optic
Photoreceptor cells Figure 13.44 Molluscan eyes. (A) The simple pigment-
cup eye of some gastropods. (B-E) Eyes with lenses.
(B) The eye of a garden snail (Cornu)-a heterobranch
Pigment cells gastropod. (C) The eye of a scallop (Pecten), a pteriomor-
phian bivalve. (D) The eye of a marine caenogastropod
(Littorina). (E) The eye of an octopus (Ocfopus). (F) The
queen scalfop Aequipecten opercularis, showing its dark
eyes along the mantle edges.
PHYLUM MOLLUSCA 513
of particular importance in georeception by burrowing coleoids can discriminate among objects by size, shape,
bivalves. Mantle eyes may also be present along the and vertical versus horizontal orientation. The eyes
mantle edge or on the siphons and have evolved inde- of Nautilus are rather primitive relative to the eyes of
pendently in a number of bivalve groups. In the spiny coleoids. They lack a lens, and are open to the water
oyster Spondylus and the swimming scallop Pecten, through the pupil. They are thought to function in the
these eyes are "mirror eyes" with a reflective layer same way that a pinhole camera does.
(the tapeum) behind pairdd retinas. This layer reflects Coleoids have complex statocysts that provide in-
light back into the eye giving these bivalves a separate formation on static body position and on body motion.
focal image on each retina-one from the lens and the Those of Nautilus are relatively simple. hr additioru the
other from the mirror. (Figure 1,3.44C-E).The bivalve arms of coleoids are liberally supplied with chemosen-
osphradium lies in the exhalant chamber, beneath the sory and tactile cells, especially on the suckers of ben-
posterior adductor muscle. thic octopuses, which have extremely good chemical
Chitons lack statocysts, cephalic eyes, and tentacles. and textural discrimination capabilities. Nnutilus is the
Instead, they rely largely on two special sensory struc- only cephalopod with osphradia.
tures. These are the adanal sensory structures in the
posterior portion of the mantle cavity and the aesthetes, Cephalopod Coloration and Ink
which are a specialized system of photoreceptors unique Coleoid cephalopods are noted for their striking pig-
to the class Polyplacophora. Aesthetes occur in high mentation and dramatic color displays. The integu-
numbers across the dorsal surface of the shell plates. ment contains many pigment cells, or chromatophores,
They are mantle cells that extend into the minute vertical most of which are under nervous control. Such chro-
canals (megalopores and micropores) in the upper teg- matophores can be individually rapidly expanded or
mentum of the shell (Figure 13.43C,D). The canals and contracted by means of tiny muscles attached to the pe-
sensory endings terminate beneath a cap on the shell riphery of each cell. Contraction of these muscles pulls
surface. Little is known about the functioning of aes- out the cell and its internal pigment into a flat plate,
thetes, but they apparently mediate light-regulated be- thereby displaying the color; relaxation of the muscles
havior. Lr at least one family (Chitonidae), some of them causes the cell and pigment to concentrate into a tiny,
are modified as simple lensed eyes. The outer mantle inconspicuous dot. Because these chromatophores are
surface of the girdle of many chitons is liberally supplied displayed or concealed by muscle action, their activity
with tactile and photoreceptor cells (Figure 13.43D). is extremely rapid and coleoid cephalopods can change
Like the rest of their nervous system, the sense or- color (and pattern) almost instantaneously. Chromato-
gans of cephalopods are highly developed. The eyes phore pigments are of several colors-black, yellow,
are superficially similar to those of vertebrates (Figure orange, red, and blue. The chromatophore color may be
73.44E), and these two types of eyes are often cited as enhanced by deeper layers of iridocytes that both reflect
a classic example of convergent evolution. The eye of and refract light in a prismatic fashion. Some species,
a coleoid cephalopod such as Octopus sits in a socket such as the cuttlefish Sepiaand many octopuses, are ca-
associated with the cranium. The cornea, iris, and lens pable of closely mimicking their background coloration
arrangement is much like that of vertebrate eyes. Also (Figure 73.72E) as well as producing vivid contrasting
as in vertebrates, the lens is suspended by ciliary mus- colors (Figure 13.12F,G).Many epipelagic squids show
cles but has a fixed shape and focal length. An iris dia- a dark-above, light-below countershading sirnilar to that
phragm controls the amount of light entering the eye, seen in pelagic fishes. Most coleoids also undergo color
and the pupil is a horizontal slit. The retina comprises changes in relation to behavioral rituals, such as court-
closely packed, long, rodlike photoreceptors whose ship and aggression. In octopuses, many color changes
sensory ends point toward the front of the eye; hence are accompanied by modifications in the surface texture
the cephalopod retina is the direct type rather than the of the body, mediated by muscles beneath the skin-
indirect type seen in vertebrates. The rods connect to something like elaborate, controlled "gooseflesh."
retinal cells that supply fibers to the large optic ganglia In addition to the color patterns formed by chro-
at the distal ends of the optic nerves. Unlike the eyes matophores, some coleoids are bioluminescent. When
of vertebrates, the coleoid comea probably contributes present, the light organs, or photophores, are arranged
little to focusing because there is almost no light refrac- in various patterns on the body, and in some cases
tion at the corneal surface (as there is at an air-cornea even occur on the eyeball. The luminescence is some-
interface). The coleoid eye accommodates to varying times due to symbiotic bacteria, but in other cases it
light conditions by changes in the size of the pupil and is intrinsic. The photophores of some species have a
by migration of the retinal pigment. Coleoid eyes form complex reflector and focusing-lens arrangement, and
distinct images (although octopuses are probably quite some even have an overlying color filter or chromato-
nearsighted) and experimental work suggests that they phore shutter to control the color or flashing pattern.
do not see colors other than as different shades oI grey, Most luminescent species are deep-sea forms, and lit-
although they can detect polarized light. In addition, tle is known about the role of light production in their
514 C h a o te r T h i rte e n
lives. Some appear to use the photophores to create a

countershading effect, so as to appear less visible to
predators (and prey) from below and above. Others
livingbelow the photic zone may use their glowing or
flashing pattems as a means of communicatiory the sig- Seminal receptacles
nals serving to keep animals together in schools or to
attract prey. Th6flashir.rg may also play a role in mate
attraction. The fire squid, Lycoteuthis,can produce sev-
eral colors of light: white, blue, yellow, and pink. At
least one genus of squid, Heteroteuthis,secretes a lumi- Nephridioduct
nescent ink. The light comes from luminescent bacteria
cultured in a small gland near the ink sac, from which
ink and bacteria are ejected simultaneously. ricardial
ducts
ln most coleoid cephalopods, alarge ink sac is locat- Seminal vesic
ed near the intestine (Figure 13.32H). An ink-producing Pericardium
gland lies in the wall of the sac, and a duct runs from Nephridium
the sac to a pore into the rectum. The gland secretes a
brown or black fluid that contains a high concentration Urogenitalpores
of melanin pigment and mucus; the fluid is stored in Figure 13.45 A solenogasterurogenitalsystem.
the ink sac. When alarmed, the animal releases the ink
through the anus and mantle cavity and out into the
surrounding water. The cloud of inky material hangs where fertilization occurs. Monoplacophorans pos-
together in the water, forming a"d:urnrrry" image that sess two pairs of gonads, each with a gonoduct con-
serves to confuse predators. The alkaloid nature of the nected to one of the pairs of nephridia (Figures 13.3D
ink may also act to deter predators, particularly fishes, and 13.14E). One tiny monoplacophoran species,
and may interfere with their chemoreception. Micropilina arntzi, is a hermaphrodite and broods its
Like virtually all other aspects of coleoid biology, embryos in its mantle cavity.
the ability to change color and to defend against preda- Most chitons are gonochoristic, although a few her-
tors are part and parcel of their active hunting life- maphroditic species are known. In chitons, the two
styles. In the course of their evolution, coleoid cepha- gonads are fused and situated medially in front of the
lopods abandoned the protection of an external shell, pericardial cavity (Figure 13.4F). Gametes are trans-
becoming more efficient swimmers but also exposing ported directly to the outside by two separate gonod-
their fleshy bodies to predators. The evolution of cam- ucts. The gonopores are located in the exhalant region
ouflage and ink production, coupled with high mobil- of the mantle groove, one in front of each nephridio-
ity and complex behavior, played a major role in the pore. Fertilization is external but can occur in the man-
successof these anirnals in their radical modification of tle cavity of the female. The eggs are enclosed within
the basic molluscan body plan. a spiny, buoyant membrane and are released into the
sea individually or in strings. A few chitons brood
Reproduction
their embryos in the mantle groove, and in one species
Primitively, molluscs are mostly gonochoristic, with (Callistochitonaiaiparous)development takes place en-
a pair of gonads that discharge their gametes to the tirely within the ovary.
outside, either through the nephridial plumbing or In living gastropods, one gonad has been lost and
through separate ducts. In species that free-spawn, the remaining one is usually located with the digestive
fertilization is external and development is indirect. gland in the visceral mass. The gonoduct is developed
Many molluscs with separate gonoducts that store in association with the right nephridium in patello-
and transport the gametes also have various means of gastropods and vetigastropods (Figure 13.46,4)while
internal fertilization. In these forms, direct and mixed in neritimorphs and caenogastropods a vestige of the
Iife history patterns have evolved. right nephridium is incorporated in the oviduct. In
Caudofoveata are gonochoristic with paired gonads, cases where the right nephridium is still functional in
while Solenogastres are hermaphroditic with a pair of transporting excretory products, as in the patellogas-
gonads (Figure 13.45).In both aplacophoran groups the tropods and vetigastropods, the gonoduct is properly
gonads discharge gametes by way of short gonoperi- called a urogenital duct, because it discharges both
cardial ducts into the pericardial chamber, from which gametes and urine.
they pass through gametoducts to the mantle cavity. In Gastropods may be gonochoristic or hermaphro-
the Solengastres fertilization is internal and the young ditic, but even in the latter case usually only a single
are sometimes brooded, while in the Caudofoveata the gonad (an ovotestis) exists, although a few hetero-
gametes are discharged into the surrounding seawater branchs have separate male and female gonads (e.g.,
PHYLUMMOLLUSCA 515
Capsule gland
(A) Gonoduct (B)
Renopericardial duct
Urogenital pore
Right kidne lips Genital duct

Site of Ventral sperm channel
fertilization
(c) (D)
Gonad (= ovotestis) Mucous gland Bursa Gonad
Hermaphroditic Common
seminalis genital Capsule
Oviduct
aperture
Albumen
gland
Mucous gland al Copulatory

Vas
of recePtacle copuiatorybursa bursa
deferens
fertilization
Figure13.46 Reproductive
systems in gastropods,
(A) Female vetigastropod (E)
frochidae). (B) Female Gonad

neogastropod (Muricidae, Hermaphroditic
(ovotestis)
Nucella). (G) Hermaphrodite
system of the euopistho-
branch Aplysia. (D) Herma- Penis
phrodite system of the eupul- complex
monate Cornu. (E) Herma-
phrodite system of the eupul-
monatehygrophi lan Physa. Seminal Fertilization Regions Sperm Prostate Vagina $ Go.ropo.e
vesicles pouch of oviduct duct gland I Gonopore
Omalogyra and in the mathildi d Gegania aalkyrie), while and a mucous or capsule gland. Many heterobranchs
others are protandric. The commihent of the right ne- lay fertilized eggs in jelly-like mucopolysaccharide
phridial plumbing entirely to serving the reproductive masses or strings produced by these glands. Most ter-
system was a major step in gastropod evolution. The restrial pulmonates produce a small number of large,
isolation of the reproductive tract allowed its indepen- individual, yolky eggs, which are often provided with
dent evolution, without which the great variety of re- calcareous shells. Other pulmonates brood their em-
productive and developmental patterns in gastropods bryos internally and give birth to juveniles. Many
may never have been realized. caenogastropods produce egg capsules in the form of
In many gastropods with isolated reproductive leathery or hard cases that are attached to objects in
tracts, the female system bears a ciliated fold or tube the environment, thereby protecting the developing
that forms a vagina and oviduct (or pallial oviduct). embryos. A ciliated groove is often present to conduct
The tube develops inwardly from the mantle wall and the soft egg capsules from the female gonopore down
connects with the genital duct. The oviduct may bear to a gland in the foot, where they are molded and at-
specialized structures for sperm storage or egg case se- tached to the substratum.
cretion. An organ for storing received sperm, the semi- The male genital duct, or vas deferens, may include
nal receptacle often lies near the ovary at the proximal a prostate gland for production of seminal secretions.
end of the oviduct. Eggr are fertilized at or near this In many gastropods the proximal region of the vas
location prior to entering the long secretory portion of deferens functions as a sperm storage area/ or seminal
the oviduct. Many female systems also have a copu- vesicle. In many caenogastropods, neritimorphs, and
latory bursa, usually at the distal end of the oviduct, lower heterobranchs the males have an external penis
where sperm are received during mating. In such to facilitate transfer of sperm (Figures 13.68 and 73.47),
casesthe sperm are later transported along a ciliated and internal fertilization takes place prior to forma-
groove in the oviduct to the seminal receptacle, near tion of the egg case. The penis is a long extension of
where fertilization takes place. The secretory section the body wall usually arising behind the right cephalic
of the oviduct may be modified as an albumin gland tentacle. In these groups with a cephalic penis, most
(A)
Prostategland Sperm groove Mantle edge
Testicular duct acting Penial

as seminal vesicle glands
(D) Vas deferens
Flagellum Epiphallus
Vagina Tentacle (cut)
Buccal mass
Penis
Diverticulum of
Mucous glands
spermathecal d
Radula sac Dart sac
Crop covered by
Spermathecalduct
salivary glands
Figure 13.47 Reproductive systems in some
gastropods. A-C showing animals removed Vas deferens
from their shells. (A) The periwinkle Littorina
Oviduct
(Caenogastropoda). (B,C) Reproductive systems of gland
Albumen
the slipper shell Crepidula (Caenogastropoda) (male
Common
and female). (D) Dissection of the common garden genital duct
snaitlCornu aspersum (Stylommatophora)'
of digestive gland
Stomachleading to
intestine and recfum
Ovotestis
of the glandular parts of the reproductive systemlie the two. Sedentaryspecies,such as territorial limpets
withinlhe mantle cavity or may extendback alongside and slipper shells,are often protanddc hermaphro-
the nephridium. In most euthyneurans theseparts of dites. In slipper shells (Crepidula),individuals may
the reproductive systemhave migrated into the body stack one atop the other (Figure 13.48),with the more
caviti and the Penis has become a retractile, internal recently settied individuals being males on top of
struciure. Sperm transfer in somegastropodsinvolves the staik, femaleson the bottom. Eachmale (Figure
the use of spermatoPhoreseither involving a penis or 1g.478)uses its long penis to inseminate the females
without one in the caseof the cerithiomorph groups/ (Figure 1,9.47C) below. Males that are in association
and someothers.In some,large paraspermare used to wii-h femalestend to remain male for a relatively long
transport the normal sPerm' period of time. Eventually, or if isolated from a fe-
Witfr both simultaneous and sequential hermaph- male, the male developsinto a female.Femaleslipper
rodite gastropods,copulation is the rule-either with shellscannot switch back to males,becausethe mas-
one indlviduil acting as the male and the other as the culine reproductive system degeneratesduring the
female, or with a mutual exchangeof sperm between sexchange.
PHYLUMMOLLUSCA 517
Old male
Transitional individual
Young female
Young male
Old female
Figure 13.48 A stack ot Crepidula fornicata, a slip-

per shell (Caenogastropoda) displaying sequential
hermaphroditism. the body wall of the other,perhapsas a meansof sexu-
ally arousingits parfirer.
Most eupulmonates are simultaneous hermaph- Mostbivalves are gonochoristic and retain the primi-
rodites, although protandric hermaphroditessome- tivelypaired gonads.However, the gonadsare large and
times occur. In most simultaneous hermaphrodite closely invested with the visceraand with eachother, so
euthyneurans a single complex gonad, the ovotestis, an apparently single gonadalmassresults.The gonod-
produces both eggsand sperm (Figures13.46C-Eand ucts are simple tubes, and fertilization is usually exter-
13.47D)with the mature gametesleaving the ovotestis nal, although somemarine and most freshwater species
via the hermaphroditic duct. Euthyneuran reproduc- brood their embryosfor a time. [r primitive bivalves, the
tive systemsare amazingly complex and varied in their gonoductsjoin the nephridia and gametesare released
plumbing and structure, and sometimeshave separate through urogenital pores.L:rmany advancedbivalves,
male and female gonopores,or only a single common the gonoducts open into the mantle cavity separately
gonopore(Figure 13.46D,E). from the nephridiopores. Hermaphroditism occurs in
Distinct precopulatory behaviors occur in a few somebivalves,including shipworms and somespecies
groups of gastropods.Theseprimitive courtship rou- of cockles,oysters,scallops,and others.Oysters of the
tines are best documentedin land pulmonates and in- genus Ostreaare sequentialhermaphrodites,and most
clude behaviors such as oral and tentacular stroking, are capableof switching sexin either direction.
and intertwining of the bodies. In some pulmonates Cephalopodsare almost all gonochoristic,with a
(e.g.,the common garden srrail,Cornu,formerly Helix) single gonad in the posterior region of the visceral
the vagina containsa dart sac,which secretesa calcare- mass (Figures13.11C,13.\28,and 13.49).The testis
ous harpoon. As courtship reachesits crescendo,and a releasessperm to a coiled vas deferens,which leads
pair of snailsis intertwined, one will drive its dart into anteriorly to a seminalvesicle.Here various glands as-
sist in packaging the sperm into elaboratesper-
matophores,which are storedin a large reservoir
called Needham's sac.From there the spermato-
I phores are releasedinto the mantle cavity via a
Gonopore
sperm duct. In femalesthe oviduct terminates
in one oviducal gland in squids, and two in oc-
Accessory topuses.This gland secretesa protective mem-
nidamental brane around eachegg.
gland The highly developed nervous system of
Oviduct cephalopods has facilitated the evolution of
Nidamental some very sophisticated precopulatory behav-
glands iors, which culminate in the transfer of sper-
Oviducal gland matophoresfrom the male to the female.Because
Ovary the oviducal opening of femalesis deep within
the mantle chamber,male coleoids use one of
their arms as an intromittent organ to transfer
the spermatophores.Thesemodified arms are
Figure 13.49 Reproductive systems in a coleoid cephalopod, calledhectocotyli (FiguresI3.12D and 13.498).
the squid Loligo. (A) Female and (B) male. In squids and cuttlefish the right or left fourth
518 Ch a o te r T h i rte e n
arm is used; in octopuses it is the right third arm. In them by flushing the egg mass with jets of water.
Nautilus four small arms form a conical organ, the Octopuses and squids tend to grow quickly to matu-
spadix, that functions in sperm transfer. Hectocotylus rity, reproduce, and then die, usually within a year or
arms have special suckers, spoonlike depressions, or two. The pearly nautilus, however, is long-lived (per-
superficial chambers for holding spermatophores dur- haps to 25-30 years), slow growing, and able to repro-
ing the transfer, wJrich may be a brief or a very lengthy duce for many years after maturity.
process. One of the most astonishing reproductive behaviors
Each spermatophore comprises an elongate sperm among invertebrates occurs in members of the pelagic
mass, a cement body, a coiled, "spring-loaded" ejacu- octopod genus Argonauta, known as the paper nauti-
latory organ, and a cap. The cap is pulled off as the luses. Female argonauts use two specialized arms to
spermatophore is removed from the Needham's sac in secrete and sculpt a beautiful, coiled, calcareous shell
squids or by uptake of seawater in octopuses. Once the into which eggs are deposited (Figure 13.178). The
cap is removed, the ejaculatory organ everts, pulling thin-walled, delicate shell is carried by the female and
the sperm mass out with it. The sperm mass adheres serves as her temporary home and as a brood chamber
by means of the cement body to the seminal receptacle for the embryos. The much smaller male often cohabits
or mantle wall of the female, where it begins to disinte- the shell with the female.
grate and liberate sperm for up to two days.
Precopulatory rituals in coleoid cephalopods usu- Development
ally involve striking changes in coloration, as the male Development in molluscs is similar in many funda-
tries to attract the female (and discourage other males mental ways to that of the other spiralian protostomes.
in the area). Male squids often seize their female part- Most molluscs undergo typical spiral cleavage, with
ner with the tentacles, and the two swim head-to-head the mouth and stomodeum developing from the blas-
through the water. Eventually the male hectocotylus topore, and the anus forming as a new opening on the
grabs a spermatophore and inserts it into the mantle gastrula wall (protostomous). Cell fates are also typi-
chamber of his partner, near or in the oviducal open- cally spiralian, including a 4d mesentoblast.
ing. Mating in octopuses can be a savage affair. The By the end of the 64-cell stage, the distinctive mol-
exuberance of the copulatory embrace may result in luscan cross is formed by a group of apical micromeres
the couple tearing at each other with their sharp beaks, (7a12-16,t2cels and their descendants, with cells 1a112-
or even strangulation of one partner by the other as 1d112forming the angle between the arms of the cross)
the former's arms wrap around the mantle cavity of (Figure 13.50).This configuration of blastomeres ap-
the latter, cutting off ventilation. In many octopuses pears to be unique to the Mollusca. Beyond these gen-
(e.g.,Argonauta, Philonexls)the tip of the hectocotylus eralities, a great deal of variation occurs in molluscan
arm may break off and remain in the female's mantle cleavage. As detailed studies are conducted on more
chamber.' and more species, the phylogenetic implications of
As the eggs pass through the oviduct, they are cov- these variations are being evaluated.
ered with a capsule-like membrane produced by the
oviducal gland. Once in the mantle cavity, various
kinds of nidamental glands may provide additional
layers or coatings on the eggs. In the squid Loligo,
which migrates to shallow water to breed, the nida-
mental glands coat the eggs within an oblong gelati-
nous mass, each containing about 100 eggs. The female
holds these egg casesin her arms and fertilizes them
with sperm ejected from her seminal receptacle. The
egg masses harden as they react with seawater and are
then attached to the substratum. The adults die after
mating and egg laying. Cuttlefish deposit single eggs
and attach them to seaweed or other substrata. Many
open-ocean pelagic coleoids have floating eggs, and
the young develop entirely in the plankton. Octopuses
usually lay grapelike egg clusters in dens in rocky
areas, and many species care for the developing em-
bryos by protecting them, and aerating and cleaning
TThe detached arm was mistakenly first described

as a parasitic
Figure 13.50 The "molluscan cross" of developing
worm and given the genus name Hectocotylus (hence the origin of embryos. (A) Gastropoda (Lymnaea). (B) Polyplacophora
the term). (Stenoplax). (C) Aplacoph ora (Epimenia).
PHYLUM MOLLUSCA 519
Figure13.51 Molluscantrocho-
phore larvae.(A)Generalized
molluscantrochophorelarva.
(B)Trochophoreof a solengaster
aplacophoran.(C) Metamorphosis
of a polyplacophoranfrom
trochophoreto juvenile.
Prototroch
Prototroch
Telotroch
Stomodeum,/blastopore
+ ----------.-->
Development may be direct, mixed, or indirect. (lecithotrophic) and live on yolk reserves. Eventually
During indirect development, the free-swimming eyes and tentacles appear, and the veliger transforms
trochophore larva that develops is remarkably similar into a juvenile, settles to the bottom, and assumes an
to that seen in annelids (Figure 13.51).Like the anne- adult existence.
lid larva, the molluscan trochophore bears an apical Like gastropods, some bivalves have long-lived
sensory plate with a tuft of cilia and a girdle of ciliated planktotrophic veligers, whereas others have short-
cells-the prototroch-just anterior to the mouth. lived lecithotrophic veligers. Many widely distributed
In some free-spawning molluscs (e.9., chitons and species have very long larval lives that allow dispersal
Caudofoveata), the trochophore is the only larval over great distances. A few bivalves have mixed de-
stage, and it metamorphoses directly into the juve- velopment and brood the developing embryos in the
nile (Figure 13.51C).Solenogastersusually have a so- suprabranchial cavity through the trochophore period;
called "test cell lala," where a bell-shaped larval test then the embryos are released as veliger larvae. Some
encloses parts of the developing animal. But in other marine and freshwater clams have direct development,
groups (e.g., gastropods and bivalves), the trocho- as for example in the freshwater family Sphaeriidae
phore is followed by a uniquely molluscan larval stage where embryos are brooded between the gill lamellae
called a veliger (Figure 13.52). The veliger larva may and juveniles shed into the water when development is
possess a foot, shell, operculum, and other adult-like completed. Several unrelated marine groups have in-
structures. The most characteristic feature of the ve- dependently evolved a similar brooding behavior (e.g.,
liger larva is the swimming organ, or velum, which Arca aiaipara, some Carditidae, etc.).
consists of two large ciliated lobes developed from the In the freshwater mussels (Unionida), the embryos
trochophore's prototroch. In some species the velum is are also brooded between the gill lamellae, where they
also a feeding organ and is subdivided into four, five, develop into veligers highly modified for a parasitic life
or even six separate lobes (Figure 13.52C).Feeding on fishes, thereby facilitating dispersal. These parasitic
(planktotrophic) veligers capture particulate food be- larvae are called glochidia (Figure 13.52E).They attach
tween opposed prototrochal and metatrochal bands of to the skin or gills of the host fish by a sticky mucus/
cilia on the edge of the velum, others are non-feeding hooks, or other attachment devices. Most glochidia
lack a gut and absorbnutrients from the host by means caenogastropods and heterobranchs). As with bivalves,
of specialphagocyticmantle cells.The host tissueoften some of these gastropods have planktotrophic veligers
forms a cyst around the glochidium. Eventually the that may have brief or extended (to several months)
larva matures,breaks out of the cyst, drops to the bot- free-swimming lives. Others have lecithotrophic ve-
tom, and assumesits adult life. ligers that remain planktonic only for short periods
Among the gastropods,only the patellogastropods
and vetigastropirdsthit rely on externalfertilization (A)
have retained a free-swimming trochophore larva. All
other gastropods suppressthe trochophore or pass
through it quickly before hatching. In many groups
Hindgut
embryos hatch as veligers (e.g.,many neritimorphs,
Figure 13.52 Molluscan veliger larvae. (A,B)

Side and front views of the veliger larva of a
caenogastropod snail. (C) A caenogastropod
veliger with four velar lobes. (D) Generalized
bivalve veliger. (E) Glochidium larva of a fresh-
water unionoidean bivalve. (F) Late veliger of a
scaphopod (Dentaliu m).
(B) Food groove

Digestive gland
Pedal ganglion
Operculum I"'
Velum
Operculum
Right valve
uctor muscles
PHYLUMMOLLUSCA 521
Remnant of velum
(A) embryo that developswithin the egg case.Early cleav-
age is meroblasticand eventually produces a cap of
cells (a discoblastula)at the animal pole. The embryo
grows in such a w ay that the mouth opens to the yolk
sac,and the yolk is directly "consumed" by the devel-
oping animal (Figure 13.54).
Molluscan Evolution and

Phylogeny
Digestive The phylogenetic details of molluscan evolution have
Mantlecavity Operculum ^l ^.^t
Srarru yet to be thoroughly elucidated.The phylum is highly
diverse, and many named taxa below the class level
are known to be polyphyletic or paraphyletic. The ex-
(B) Mantle cavity
istenceof a good fossil record (primarily of shells)has
been both a blessingand a curse as efforts to trace the
evolutionary history of molluscshave often been frus-
trated by the limited and sometimesconfusing dataset
provided by molluscan shells.
Until fairly recently, the idea of a "hypothetical an-
cestral mollusc" (affectionately known as HAM) was
popular, the nature of which derived largely from early
work of the eminent British biologist and "Darwin's
Bulldog" T. H. Huxley. Detailed and sometimeshighly
imaginative descriptionsof this hypothetical ances-
tral mollusc were proposed by various workets, even
Right shell muscle including speculationson its physiology, ecology and
Figure 13.53 Settled larva of the abalone (Haliotis) behavior (seeLindberg and Ghiseh 2003).The useful-
undergoingtorsion. (A) Left-sideview after about 90' of nessof HAM in molluscanevolutionary studieswas
torsion,with mantlecavity on the right side. (B)Torsion questionedas zoology moved into an era of explicit
continuesas the mantlecavity and its associatedstruc- phylogeneticanalysis(i.e.,cladistics).Thus,most work-
tures twist forward over the head.
ers now avoid the pitfalls of a priori construction of a
(sometime less than a week). Planktotrophic veligers

feed by use of the velar cilia, whose beating drives the
animal forward and draws minute planktonic food par-
ticles into contact with the shorter cilia of a food groove.
Once in the food groove, the particles are trapped in
mucus and carried along ciliary tracts to the mouth.
Almost all pulmonates and many caenogastropods
have direct development, and the veliger stage is passed Optic lobe
in the egg case, or capsule. Upon hatching, tiny snails
crawl out of the capsule into their adult habitat. In some
neogastropods (e.g., certain species of Nucella), the en-
capsulated embryos cannibalize on their siblings, a phe-
nomenon called adelphophagy; consequently, only one
or two juveniles eventually emerge from each capsule.
It is usually during the veliger stage that gastropods
undergo torsion (see previous discussion of torsion),
when the shell and visceral mass twist relative to the
head and foot (Figures 13.18 and 13.53).As we have
seen, this phenomenon is still not fully understood, but
it has played a major role in gastropod evolution.
Cephalopods produce large, yolky, telolecithal eggs.
Development is always direct, the larval stages having Figure 13.54 Juvenile coleoid cephalopod attached to
been lost entirely during evolution of the yolk-laden and consuming its sac of yolk.
hypothetical ancestor, and instead analyze the evolu- are enumerated in the figure legend and briefly sum-
tionary history of molluscs by phylogenetic inference. mafized in the following discussion. The nodes on
Although morphological analyses of molluscan rela- the cladogram have been lettered to facilitate the
tionships have differed in some details, the phyloge- discussion.
netic relationships resulting from this work have been Exactly where the molluscs arose within the spira-
similar. In contrast, more recent molecular analyses of lian clade, and their kinship to other spiralian phyla,
molluscan relationships have produced several alter- are still matters of much debate. While some workers
native trees depending on the molecular data type and treat them as descendant from a segmented ancestor,
analytical methods. Based on these recent phylogenetic most do not. We support the idea that molluscs arose
studies, the probable molluscan common ancestor was from a schizocoelomate/ nonsegmented precursor.
small (-5 mm long), with a dorsal shell or cuticle, and Recent molecular studies (see References: Molluscan
a flattened ventral surface on which the animal moved Evolution and Phylogeny) have suggested different sis-
by ciliary gliding. Our phylogeny (Figure 13.55) sum- ter-group relationships for the molluscs, though com-
marizes some current thinking on molluscan evolu- monly the annelids are nested within these putative
tion. The characters used to construct the cladogram sister clades. However, as with manv of the sr:iralian
Figure 13.55 A cladogram depicting a conservative Solenogastres) defining node e: (10) vermiform body;
view of the phylogeny of the Mollusca based on current (11) foot reduced; (12) gonads empty into pericardial cav-
hypotheses (see Sigwart and Lindberg 2O15tor alterna- ity, exiting to mantle cavity via U-shaped gametoducts;
tive molluscan phylogenies).The numberson the clado- (13) without nephridia.
gram indicatesuitesof synapomorphiesdefiningeach Synapomorphies of Gaudofoveata: (14) calcareous scler-
hypothesized lineor clade. ites of the body wall form imbricating scales; (15) com-
Synapomorphiesof the phylum Molluscadefiningnode plete loss of foot.
a: (1)reductionof the coelomand developmentof an Synapomorphies of Solenogastres: (16) posterior end of
open hemocoeliccirculatorysystem;(2) dorsal body wall reproductive system with copulatory spicules; (17) loss of
forms a mantle;(3) extracellularproductionof calcareous ctenidia.
scferites(and/orshell)by mantleshell glands;(4)ventral Synapomorphies of Polyplacophora: ('18) shell with 8
body wall musclesdevelopas muscularfoot (or foot pre- plates (and with B shell gland regions), articulamentum
cursor);(5) radula;(6) chamberedheart with separateatria layer, and aesthetes; (.19) multiple ctenidia; (20) expanded
and ventricle;(7) increasein gut complexity,with large and highly cuticularized mantle girdle that "fuses" with
digestiveglands;(B)ctenidia. shell olates.
Synapomorphiesof the Aculifera (Aplacophora+ Synapomorphies of the Conchifera defining node b:
Polyplacophora)defining node d: (9) sclerites. (21) presence of a well-defined single shell gland region
Synapomorphiesof the Aplacophora (Caudofoveata+ and larval shell (protoconch); (22) shell univalve (of a
single piece; note: the bivalve shell is derived from the
univalve condition); (23) shell of basically three-layers
Aculifera (periostracum, prismatic layer, lamellar or crossed layer);
(24) mantle margin of three parallel folds, each specialized
Aplacophora Conchifera for specific functions; (25) statocysts; (26) viscera concen-
trated dorsally.
Ê
!9 Synapomorphies of Monoplacophora: (27) 3-6 pairs
!' 6
2 E{j ctenidia; (28) 3-7 pairs nephridia; (29) 8 pairs pedal retrac-
eq 'd
6
9aF " _ E H _ A tor muscles; (30) 2 pairs gonads; (31) 2 pairs heart atria.
a:Y^V> q!- - ,
G Synapomorphies of Gastropoda: (32) torsion; (33) cephal-
a =i.83 q
6 ic tentacles; (34) operculum.
i(Jd> U I
U)
Synapomorphies of Bivalvia: (35) bivalve shell and its
a\
4349 associated mantle and (in autobranch bivalves) ctenidial
tD-l/
z^ /-- . ) ^ a | modifications; (36) loss of radula; (37) byssus (auto-
branchs); (38) Iateral compression of body; (39) adductor
muscles; (40) ligament.
Synapomorphies of the cephalopod-scaphopod line
defining node c: (41) ano-pedal flexure; (42\ new neuro-
anatomical features, including cerebral ganglia fusion and
position.
Synapomorphies of Cephalopoda: (43) expansion of the
coelom and closure of the circulatory system; (44) septate
shell; (45) ink sac (in coleoids); (46) siphuncle; (47) beak-
like jaws; (48) foot modified as prehensile arms/tentacles
and funnel (=siphon); (49) development of large brain.
Synapomorphies of Scaphopoda: (50) tusk-shaped, shell
open at both ends; (51) loss of heart and ctenidia;
(52) captacula.
PHYLUM MOLLUSCA 523
phyla, identification of the mollusc sister group re- Because of their small size, specialized respiratory
mains a work in progress. Molluscs are clearly allied structures were probably not required in the first
with the other spiralian protostomes (Platyhelminthes, molluscs and gas exchange was through the dorsal
Nemertea, Annelida), which are characterized by devel- epidermis. However, with the origin of the cuticle-
opmental features such as spiral cleavage,4d mesento- covered mantle or dorsal shell covering this surface,
blast, and trochop\ore-like lawae. But precisely where posterior, specialized respiratory structures (ctenidia)
in the spiralian linehge they arose remains problematic. originated and became associated with excretory and
The major steps in the evolution of what we gener- reproductive pores in a posterior mantle cavity. This
ally think of as a "typical" mollusc-that is, a shelled arrangement would have been modified at least twice;
mollusc, also remains controversial. Previous scenarios in both the polyplacophorans and monoplacophorans
have often argued that this step took place after the ori- the mantle cavity was lost as it became continuous
gin of the aplacophorans, perhaps as molluscs adapted with the expanded mantle groove alongside the foot
to active epibenthic lifestyles. These steps centered and the ctenidia multiplied and extended anteriorly
largely on the elaboration of the mantle and mantle in the mantle groove. Secondary modifications of the
cavity, the refinement of the ventral surface as a well- shape of the foot and other features in bivalves and
developed muscular foot, and the evolution of a con- scaphopods allowed most of these animals to exploit
solidated dorsal shell gland and solid shell(s) in place infaunal life in soft sediments, and both of these taxa
of independent calcareous sclerites. are highly adapted to sediment burrowing. However,
The description of a solenogaster larva by Pruvot in these modifications are clearly convergent and scaph-
1890,in which the dorsal surface was said to bear seven opods share other characters, including ano-pedal flex-
transverse bands of sclerites (described as "composite ure, with cephalopods. Gastropods also undergo ano-
plates," reminiscent of chitons), led some workers to pedal flexure, but this could be convergent according
postulate that aplacophorans and polyplacophorans to some molecular studies. Scaphopods are also the last
might be sister-groups; a relationships confirmed by class of molluscs to appear in the fossil record (about
several recent phylogenomic studies (e.g.,Kocot et al. 450 Ma, Late Ordovician).
2011). However, the discovery of a possible aplacopho- Monoplacophorans share the character of a single
ran fossil having seven dorsal shell plates from Silurian (univalve) shell with other molluscs (other than bi-
deposits in England (Acaenoplax),as well as "footless" valves and chitons). They also share a similar shell
chitons (Kulindroplax and Phthipodochiton),have further structure and a host of other features. The only syn-
confused the polarity of the aplacophoran-chiton char- apomorphies defining the monoplacophorans seem
acter transformation. Adding to the confusiory there are to be their repetitive organs (multiple gills, nephridia,
fundamental differences between the shells of polypla- pedal muscles, gonads, and heart atria). The ques-
cophorans and those of all other molluscs, an observa- tion of whether this multiplicity arose uniquely in the
tion suggesting that the chitons and aplacophorans may monoplacophorans or represents a symplesiomorphic
stand alone as a unique radiation off the early molluscan retention of ancestral features from some unknown
line. Three hlpotheses have been offered to explain this metameric ancestor (below node a on the cladogram)
"shell problem" in molluscan evolution: (1) The mul- has not been resolved (see discussion below) and will
tiplate shell may have been ancestral, the single-shell likely require developmental studies on monoplacoph-
condition having evolved by coalescence of plates. (2) orans to finally settle the question.
The single shell may have been ancestral, and the mul- The bivalve line in the cladogram is defined by the
tiplate forms arose by subdivision of the single shell. presence of 2 shell valves, adductor muscles, reduction
(3) The single-shell and multishell designs arose inde- of the head region, decentralization of the nervous sys-
pendently from a shell-less ancestor, perhaps by way tem and associated reduction or loss of certain sensory
of sclerite consolidation. The presence of eight pairs of structures, and expansion and deepening of the mantle
pedal retractor muscles in both polyplacophorans and cavity.
monoplacophorans has been taken as evidence in favor C ephal opods are hi ghl y speci al i zed m olluscs
of the first explanation. Acceptance of the first hypoth- and possess a number of complex synapomorphies.
esis suggests that the ancestor at node a in the cladogram Primitive shelled cephalopods are represented today
in Figure 13.55 was a multivalved chiton-like creature. by only six species of Nautilus, although thousands of
Acceptance of the second hypothesis implies that the an- fossil species of shelled nautiloid cephalopods have
cestor at node awas a univalved, monoplacophoran-like been described. This highly successful molluscan
ancestor. The third hypothesis postulates that the ances- class probably arose about 450 million years ago. The
tor at node a lacked a solid shell altogether. nautiloids underwent a series of radiations during
The primitive mantle and foot arrangement was the Paleozoic, but were largely replaced by the am-
probably somewhat similar to that in living polypla- monoids after the Devonian period (325 million years
cophorans or monoplacophorans-that is, a large ago). The ammonoids, in turn, became extinct around
flattened sole was surrounded by a mantle groove. the Cretaceous-Tertiary boundary (65 million years
524 ChaPter Thirteen
ago). The origin of the coleoid cephalopods (octopus-

The various ideas on the origin of the molluscs fall
ei squids, and cuttlefish) is obscure, possibly dating
into three categories:molluscswere derived from (1)
backio the Devonian. They diversified mainly in the
a free-living flatworm (Ptatyhetminthes)ancestor,(2) a
Mesozoic and became a highly successful group by ex-
,rorr"g-"ited coelomateprotostomeancestor,or (3)
ploiting a very new lifestyle, as we have seen'
a segmentedancestor,perhaps even a_commonances-
'
The"issue ofianceslral metamerism in molluscs tor ivith the annelids. The first hypothesis, known as
has been debated since the discovery of the first liv- the "turbellarian theory," was originally basedupon
ing monoplacophoran (Neopilina galatheae)\n 1952' the supposedhomology and similarity in mode of lo-
H6*er r e t, m o n o p l a c o p h o ra n s a r e not the oni y comotion between molluscs and flatworms by means
m ollus cs to e x p re s s s e ri a l re p l i c a ti on or to have
of a "ventral mucociliarygliding surface'"It suggests
r epeat e d o .g u ^ t re m i n i s c e n t o f metameri sm (or
that either the molluscswere the first coelomateproto-
" ps eudo me L me ri s m," a s s o m e p refer to cal l i t)' stomes,or that they sharea common ancestorwith the
Ptlyplacophorans have many serially repeated gills
first coelomates.However,most contemporarywork-
in ti't" tttutttle groove and also typically possess eight ers argue that the large pericardial spacespresentin
pairs of pedal ietractor muscles and eight shell plates'
primitlve molluscs(e.g., aplacophorans,monoplacoph-
'orur-'.r,
fne two pairs of heart atria, nephridia, and-ctenidia in
polyplacophorans) point to a coelomaterather
Nautilus (and two pairs of retractor muscles in some
than an ucoelomite (platyhelminth) ancestry,and the
fossil forms) have ilso been regarded by some work-
turbellarian theory enjoyslittle favor today'
The secondtheory advancedby Scheltemain the
ers as primitive metameric features'
The question is whether or not organ repetition-in
1990ssuggestedthatsipunculans (now placed within
these mtluscs represents vestiges of a true, or funda-
the AnnJida) and molluscsmightbe sistergroups,shar-
mental, metamerism in the phylum' If so, they represent
ing, among other things, the unique"molluscan cross"
remnants of an ancestral metameric body plan and may
a"1it g a"""lopment. Ho-"rret, the idea that sipunculan
indicate a close relationship to annelids' On the other
embryogenyincludesa molluscancrossblastomerecon-
hand, organ repetition in certain molluscan groups may
figuration is no longer strongly supported' Scheltema
be the reiult oi independent convergent evolution and
alio suggestedthat certain features of the sipunculan
pelagos-"p'hera larva may be homologous to somemol-
not an ancestral molluscan attribute at all' And, nothing
like the teloblastic metameric development of arrnelids
irm.o:. structures.Indeed, molluscs sharemost of their
is seen in molluscs. The genetic/evolutionary potential
typical spiralian featureswith the sipunculans,as well
for serial repetition of organs is not uncommon and oc-
as^theechiuridsand other annelids(e'g',spiral cleavage'
curs in other non-annelid bilaterian phyla as well, e'g''
schizocoely,trochophorelarvae)'This leadsto the third
Platyhelminthes, Nemertea, and Chordata' hypothesis,that molluscsand annelidsare closelyrelat-
The origin of molluscs themselves remains enigmat- ed and that molluscsmight have arosetrom a segment-
ic. The exlelent fossil record of this phylum extends ed coelomateancestor.Perhapsthe three most striking
back some 500 million years and suggests that the ori- slmapomorphiesdistinguishingmodern-molluscsfrom
gin of the Mollusca probably lies-{ Jn9 Precambrian'
un ,"iid, and most other spiraliansare:the reduction of
indeed, the late Precimbrian foss\l Kimberella quadratn'
the coelom and the concornitantconversionof the closed
once thought to be a cnidarian, has been argued to have circulatory systemto an open hemocoelicone;the elabo-
molluscan features, including perhaps a shell and mus-
ration of tire-bodywall into a mantlecapableof secreting
cular foot. However, recent examination of hundreds
calcareousscleritesor shell(s);and, the unique mollus-
of specimens now suggests Kimberellamore likely be-
can radula.Identifying the sisterSroup to the Mollusca
longs to an extinct spiralian group' remainsa work in Progress.
Se/ected References
The field of malacology is so large, has had such a countlessshell guides and coffee-tablebooks' Distill-
long history, and has so embraced the mixed blessings ing all of this in-toa small set of key referencesuseful
of Jontributions from amateur shell collectors, that
foientry into the professionalliterature is difficult; the
dealing with the literature is a daunting task' Many
list below is our attemPtto do so.
molus]cs are of commercial importance (e'g', Haliotis, GeneralReferences
Mytilus,Loligo) and for these groups hundreds of stud- Beesley,P. L., G. f. B. Rossand A. Wells (eds')' 1998'Mollusca:
ieJ appear annually; others are importantlabotatoty / Thi Southern Synthesis.Fauna of Australia' VoI' 5' CSIRO
experirnental organisms (e.g., Loligo, Octopus,Aply sia) Publishing, Melbourne, Australia' [Perhaps the best gen-
und papers are also published on these groups' eral reviei of molluscan biology and systematics available'
ûn.y An extraordinary 2-volume text with chapters by leading
New taxonomic monographs on various groups or
specialists.l
geographical regions also appear each year, as do
PHYLUMMOLLUSCA 525
Cheng, T.C.1967. Marine mollusks as hosts for s).'rnbioses.Adv. Todt, C. and L. v. Salvini-Plawen. 2005.The digestive tract of
Mar. Biol. 5:1424. [An exhaustive summary.] Helicoradomenla(Solenogastres,Mollusca), aplacophoran
Falini, G., S. Albeck, S. Weiner and A. Addadi. 1996.Control of molluscs from the hydrothermal vents of the East Pacific Rise.
aragonite or calcite polymorphism by mollusc shell macro- Inver. Biol. 124: 230-253.
molecules. Science27L: 67-59. Todt, C. and A. Wanninger. 2010.Of tests, trochs, shells, and
Giese, A. C. and ]. S. Pearse (eds.). 1977.Reproductionof Marine spicules: Development of the basal mollusk Wirenia argentea
Inaertebrates,Vol. 4, Molluscs: Gastropodsand Cephalopods. (Neomeniomorpha) and its bearing on the evolution of tro-
Academic Press,N.ew Yor\. chozoan larval key features.Front. Zool.7: 6.
Giese,A. C. and J. S. Pearse(eds.).1979.Reproduction of Marine
Inoertebrntes,Vol. 5, Molluscs:Pelecypods and LesserClasses. Monoplacophora
Academic Press,New York. Clarke, A. H. and R. f. Menzies. 1959.Neopilina (Vema)ewingi, a
Harrison, F. W. and A. j. Kohn (eds). 1994.MicroscopicAnatomy second living speciesof the Paleozoic classMonoplacophora.
of Inoertebrates.VoIs.5 and 6. Mollusca.Wiley-Liss, New York. Science 129: L026-1027.
Heller, 1.1,990.Longevity in molluscs. Malacologia 31(2): Haszprunar, G. and Schaefer, K. (1997). Monoplacophora.
259+95. Pp. 41.5-457in F. W. Harrison and A. Kohn, Mollusca 2.
Hochachka, P. W. (ed.). 1983. The Mollusca, VoI. 1, Metabolic Microscopicanatomyof inoertebrates,68.Wiley-Liss, New York.
Biochemistryand Molecular Biomechanics; Vol. 2, Enaironmental Lemche, H.I957. A new living deep-seamollusc of the Cambro-
Biochemistryand Physiology.Academic Press,New York. Devonian class Monoplacophora. Nature L79: 4L3-4L6.
'
Hyman, L. H. 1967. The Inaertebrates,VoI. 6, Mollusca I. [Report of the first discovery of living monoplacophorans.]
Aplacophora,Polyplacophora,Monoplacophora,Gastropoda. Lemche, H. and K. G. Wingstrand. 1959. The anatomy of
McGraw-Hill, New York. [Dated, but still one of the best gen- NeopilinagalatheaeLemche,l95T.Galathea Rpt. 3: 9-71.
eral surveys of molluscan anatomy in English.] Lindberg, D. R. (2009).Monoplacophorans and the origin and re-
Keen, A. M. and E. Coan. 1974. Marine Molluscan Generaof lationshios of mollusks. Evol. Educ. Outreach 2: 191-203.
WesternNorth America:An IllustratedKey.2nd Ed. Stanford Menzies, R.l. and W. Layton. 1962. Anew species of monopla-
University Press,Stanford, Califomia. [Technical keys.] cophoran mollusc, Neopilina (Neopilina)aeleronisfrom the
Kniprath, E. 1981.Ontogeny of the molluscan shell field. Zool. slope of the Cedros Trench, Mexico. Ann. Mag. Nat. Hist.
Scripta t0:61J9. (13)5:401-406.
Lydeard, C. and D. R. Lindberg (eds.). (2003).Molecular system- War6n, A. 1988.Neopilinagoesi,anew Caribbean monoplacoph-
atics and phylogeographyof mollusks.Smithsonian Series in oran mollusk dredged in 1869.Proc. Biol. Soc. Wash. 101:
Comparative Evolutionary Biology. 676481.
Moore, R. C. (ed.). 1957-7l.Treatiseon lnuertebrate Paleontology. Wilson, N. G., D. Huang, M. C. Goldstein, H. Cha, G. Giribet
Mollusca,Parts 1-6 (Vols. I-N). Universify of KansasPressand and G. W. Rouse. 2009.Field collection of Laeaipilinahyalina
Geological Society of America, Lawrence, KA. Mclean, 1979 from southern California, the most accessible
Raven, C. P. 1958. Morphogenesis:The Analysis of Molluscan living monoplacophoran. J. Mollus. Stud. 75: 195-197.
Deaelopment. Pergamon Press,New York. Wingstrand, K. G. 1985.On the anatomy and relationships of re-
Spanier, E. (ed.). 1987. TheRoyalPurpleand BiblicalBIue.Argaman cent Monoplacophora. GalatheaRpt. 1.6:7-94.
andTekhelet.TheStudy of ChiefRabbiDr.IsaacHerzogon theDye
Industries in Ancient lsrael and RecentScientificContributions. Polyplacophora
Keter Publishing House, Jerusalem. Boyle, P. R.1977. The physiology and behavior of chitons. Ann.
Wilbur, K. M. (gen. ed.). 1983-1988.TheMollusca. Vols. L-12. Rev. Oceanogr.Mar. Biol. L5:461-509.
Academic Press,New York. Eernisse, D. J. 1988. Reproductive patterns in six species of
Lepidochitona(Mollusca: Polyplacophora) from the Pacific
Caudofoveata and Solenogastres (the Aplacophorans) Coast of North America. Biol. Bull. 174(3\:287102.
Garcia-Alvarez, O. and L. v. Salvini-Plawen. 2007. Species Fisher, v.-F. P. 1978. Photoreceptor cells in chiton esthetes.
and diagnosis of the families and genera of Solenogastres Spixiana l:2091L3.
(Mollusca). Iberus 25: 73-143. Kass, P. and R. A. VanBelle. \998. Catalogueof Liaing Chitons
Okusu, A. 2002. Embryogenesis and development of Epimenia (Mollusca;Polyplacophora). 2nd Ed. W. Backhuys, Rotterdam.
babai(Mollwsca:Neomeniomorpha). Biol. Bull. 203: 87-103. Nesson, M. H. and H. A. Lowenstam. 1985.Biomineralization
Salvini-Plawen, L., v. (1980).A reconsideration of systemat- processesof the radula teeth of chitons. Pp. 333-363 in f.
ics in the Mollusca (phylogeny and higher classification). L. Kirschvink et al. (eds.), Magnetite Biomineralizationand
MalacoIogia L9: 24917 8. MagnetorecEtionin Organisms.Plenum, New York.
Scheltema,A. H. 1981.Comparative morphology of the radulae Sigwart, J. D. (2008).Gross anatomy and positional homology of
and the alimentary tracts in the Aplacophora. Malacologia 20: gills, gonopores, and nephridiopores in "basal" Iiving chitons
351-383. (Polyplacophora: Lepidopleurina). Am. Malacol. Bull. 25:
Scheltema,A. H. and C. Schander.2000.Discrimination and phy- 4349.
logeny of solenogasterspeciesthrough the morphology of Slieker, F. J. A. 2000. Chitons of the World. An lllustrated Synopsis
hard parts (Mollusca, Aplacophora, Neomeniomorpha). Biol. of RecentPolyplacophora. L'hrformatore Piceno,Ancona, Italy.
BulI.198: 121,-151,. S m i t h , A . G . 1 9 6 6 . T h e l a r v a l d e v e l o p m e n t o f ch i to n s
Scheltema, A. H., M. Tscherkassky and A. M. Kuzirian. L994. (Amphineura). Proc. Calif. Acad. Sci.32:433-M6.
Aplacophora. Pp. 13-54 in F. W. Harrison and E. E. Ruppert Sutton, M. D. and J. D. Sigwart, 1.D.2012. A chiton without a
(eds.),MicroscopicAnatomy of lnaertebrates.VoL 5, Mollusca L foot. Palaeontology 55: 401.41L.
Wiley-Liss,New York.
Scherholz, M., E. Redl, T. Wollesen, C. Todt and A. Wanninger. Gastropoda
201.5.From complex to simple: Myogenesis in an aplacopho- Barker, G. M. (ed.). 2001..The biologyof terrestrial molluscs.CABI
ran mollusk reveals key traits in aculiferan evolution. BMC Publishing, Wallingford, UK.
Evol. Biol. 15:201. Bamhart, M. C. 1986.Respiratory gas tensions and gas exchange
Todt, C. 2013. Aplacophoran mollusks-still obscure and diffi- in active and dormant land snails, Otala lactea.Physiol. Zool.
cult? Am. Malacol.Bull.31: 181-187. 59:733-745.
Bouchet, P. 1989.A marginellid gastropod parasitizes sleeping Havenhand, J. N. 1991.On the behaviour of opisthobranch lar-
fishes. Bull. Mar. Sci. 45(D:76-84. vae. J. Moll. Stud. 57: 119-t31..
Brace, R. C. 1977. Anatomical changes in nervous and vascular Hickmary C. S. 1983.Radular pattems, systematics,diversity and
systems during the transition from prosobranch to opistho- ecology of deep-sealimpets. Yeliger 26:7-92.
branch organization. Trans. Zool. Soc.London 34: 1-26. Hickman, C. S. 1984.Implications of radular tooth-row function-
Branch, G. M. 1981.The biology of limpets: Physical factors, en- aI integration for archaeogastropodsystematics.Malacologia
ergy flow and ecological interactions. Ann. Rev. Oceanogr. 25(1):1,43-1,60.
Mar. Biol. 19:235380. Hickman, C. S. 1992.Reproduction and development of trochean
Brunkhorst, D. l. 1991.Do piryllidiid nudibranchs demonstrate gastropods.Veliger 35: 245-272.
behaviour consistent with their apparent warning color- fudge, J. and G. Haszprttnar.201,4. The anatomy of Lepetellasi-
ations? Some field obsewations. J. Moll. Stud. 57:481489. errai (Y etigastropoda, Lepetelloidea): implications for repro-
Carlton, l. T., G.J. Vermeij, D. R. Lindberg, D.A. Carlton and duction, feeding, and s)'rnbiosis in lepetellid limpets. Invert.
E. C. Dudley. 1991.The first historical extinction of a marine Biol. 133:324-339.
invertebrate in an oceanbasin: The demise of the eelgrasslim- Kempf, S. C. 1984. Symbiosis between the zooxanthella
pet Lottia alaeus.Biol. Bull. 180 72-80. Symbiodinium (= Gymnodinium) mig oadriaticum(Fr atdenthal)
Chase,R. 2002.Behaztior and its neuralcontrolin gastropodmolluscs. and four speciesof nudibranchs. Biol. Bull. 166: L1,0-I26.
Oxford University Press,Oxford. Kempf, S. C. 1991.A "primitive" symbiosis between the aeolid
Collier, I. R. 1.997.Gastropods. The snails. Pp. 189-218 in S. nudibranch Berghiazterrucicornis(A. Costa, 1867) and a zoo-
F. Gilbert and A. M. Raunio, Embryology.Constructingthe xanthella. J. Mol. Stud. 57:75-85.
Organism.Sinauer Associates,Sunderland, MA. Lindberg, D. R. and R. P. Guralnick. 2003. Phyletic patterns of
Conklin, E. J. and R. N. Mariscal. 1977.Feedine behavior, ceras early development in gastropod molluscs. Evol. Dev. 5:
strucfure, and nematocyst storage in the aeolid nudibranch, 494-507.
Spurilla neapolitana.Bull. Mar. Sci.27: 658467. Lindberg, D. R. and W. F. Ponder. 2001.The influence of clas-
Connor, V. M. 1986.The use of mucous trails by intertidal lim- sification on the evolutionary interpretation of structure: a re-
pets to enhancefood resources.Biol. Bull.171:54U564. evaluation of the evolution of the pallial cavity of gastropod
Cook, S. B. L97l. A study in homing behavior in the limpet molluscs. Org. Divers. Evol. 1:273-299.
Siphonariaalternata.Biol. Bull. 141: 449457. Linsley, R. M. 1978.Shell formation and the evolution of gastro-
Croll, R. P. 1983.Gastropod chemoreception.Biol. Rev. 58: pods. Am. Sci. 66: 432441.
293119. Lohmann, K. f. and A. O. D. Willows. 1987.Lunar-modulated
Fretter, V. 7967. The prosobranch veliger. Proc. Malac. Soc. geomagnetic orientation by a marine mollusk. Science 235:
London 37: 357166. 331134.
Fretter, V. and M. A. Graham. 7962.British Prosobranch Molluscs. Marcus, E. and E. Marcus .1967. AmericanOpisthobranch Mollusks.
Their FunctionalAnatomy and Ecology.Ray Society,London. Studiesin TropicalOceanography Series,No.6. University of
Fretter,V. and J. Peake (eds.).1975,1.978. Pulmonates, VoI. 1, Miami Press,Coral Gables, Florida. [Although badly out of
Functionql Anatomy and Physiology; VoL 2A, Systematics, date, this monograph remains a benchmark work for the
Eztolutionand Ecology.Academic Press,New York. American fauna.]
Fursich, F. T. and D. fablonski. 1984. Late Triassic naticid drill Marin, A and J. Ros. 1991.Presenceof intracellular zooxanthellae
holes: Carnivorous gastropods gain a major adaptation but in Mediterranean nudibranchs. |. Moll. Stud. 57: 87-107.
fail to radiate. Science224: 78-80. McDonald, G. and J. Nybakken. 1991.A preliminary report on a
Gaffney, P. M. and B. McGee. 1992. Multiple paternity in worldwide review of the food of nudibranchs. J. Moll. Stud.
Crepidulafornicata (Lirnaeus). Veliger 35(1): 12-15. 57:61-63.
Garstang, W. (1928).Origin and evolution of lawalforms.Nature Miller, S. L.1974. Adaptive design of locomotion and foot form
122:366. in prosobranch gastropods.J. Exp. Mar. Biol. Ecol 14:99-156.
Gilmer, R. W. and G. R. Harbison. 1986.Morphology and field Miller, S. L.1974. The classification, taxonomic distribution, and
behavior of pteropod molluscs: Feeding methods in the fami- evolution of locomotor types among prosobranch gastro-
lies Cavoliniidae, Limacinidae and Peraclididae (Gastropoda: pods. Proc. Malacol. Soc.London4l.:233172.
Thecosomata).Mar. Biol. 9'i.:47-57. Milne-Edwards H. 1848.Note sur la classification naturelle chez
Gosliner, T . M. 7994.Gastropoda: Opisthobranchia. Pp. 253-355 Mollusques Gasteropodes.Annales des SciencesNafuralles,
in F. E. Harrison and A. J. Kohn (eds.),MicroscopicAnatomy of Series3,9: 102-L72.
Inztertebrates,Wiley-Liss, New York. Norton, S. F. 1988.Role of the gastropod shell and operculum in
Gould, S. J. 1985.The consequencesofbeing different: Sinistral inhibiting predation by fishes. Science 241: 92-94.
coiling in Ceriorz. Evolution 39: 1364-1379. Olivera, B. M. et al. 1985.Peptide neurotoxins from fish-hunting
Greenwood, P. G. and R. N. Mariscal. 1984.Immature nemato- cone snails. Science230: 1338-1343.
cyst incorporation by the aeolid nudibranch Spurilla nenpoli- O'Sullivan, J. 8., R. R. McConnaughey and M. E. Huber. 1,987.A
tana.Mar. Biol. 80: 35-38. blood-sucking snail: The cooper's nutmeg, Cancellariacooperi
Haszprunar, G. 1985.The fine morphology of the osphradial Gabb, parasitizes the Califomia electric ray, Torpedocalifornica
senseorgans of the Mollusca. I. Gastropoda, Prosobranchia. Ayres. Biol. B.ull.172: 362166.
Phil. Trans.Roy. Soc.Lond.B307:457496. Pawlik, f. R., M. R. Kernan, T. F. Molinski, M. K. Harper and D.
Haszprunar, G. 1985. The Heterobranchia: New concept of f . Faulkner. 1988.Defensive chemicals of the Spanish danc-
the phylogeny of the higher Gastropoda. Z. Zool. Syst. er nudibranch Hexabranchussanguineusand its egg ribbons:
Evolutionsfors ch. 23: 7517 . Macrolides derived from a sponge diet. J. Exp. Mar. Biol. Ecol.
Haszpntnar, G. 1988.On the origin and evolution of major gas- I79:99-1,09.
tropod groups, with special reference to the Streptoneura. J. Pennington, J. T. and F. Chia. 1985.Gastropod torsion: A test of
Moll. srud. 54:367-441. Garstang's hypothesis. Biol. Bull. 1.69: 39119 6.
Haszprunar, G. 1989.Die Torsion der Gastropoda - ein bio- Perry, D. M. 1985. Function of the shell spine in the preda-
mechanischer Prozess.Z. ZooI. Svst. Evolutionsforsch. 27: ceous rocky intertidal snall Acanthina spirata(Prosobranchia:
1,-7. Muricacea). Mar. Biol. 88: 51-58.
PHYLUMMOLLUSCA 527
Po nd er, W. F. 1973. The or igin and ev o l u t i o n o f t h e and Wtilus edulis (Molhtsca: Bivalvia): The role of mucus in
Neogastropoda. Malacologia 12: 295138. particle transport. Mar. Ecol. Prog. Ser.98:275-282.
Ponder, W. F. (ed.). 1988.Prosobranchphylogeny. Malacol. Rev., Bouchet,P., l.-P.Rocroi, R. Bieler, J. G. Carter and E. V. Coan.
Supp.4. 2010.Nomenclator of bivalve families with a classification of
Ponder, W. F. and D. R. Lindberg. 1997.Towards a phylogeny of bivalve families. Malacologia 52: l-184.
gastropod molluscs: an analysis using morphological charac- Boulding, E. G. 1984.Crab-resistant features of shells of burrow-
ters. Zool. f. Linn. Soc.1L9:83-265. ing bivalves: Decreasing vuL:rerability by increasing handling
Potts, G. W. 1981.The'anatomy of respiratory structures in the time. J. Exp. Mar. Biol. Ecol.76:201-223.
dorid nudibranchs, Onchidorisbilamellataand Archidorispseu- Childress J. J., C. R. Fisher, f. M. Brooks, M. C. Kennicutt II, R. R.
doargus,with details of the epidermal glands. |. Mar. Biol. Bidigare and A. E. Anderson. 1986.A methanotrophic marine
Assoc. U.K. 6L: 959-982. molluscan (Bivalvia, Mytilidae) slnnbiosis: Mussels fueled by
Rudman, W. 8.1,99L.Purpose in pattem: The evolution of colour gas.Science233: 1306-1308.
in chromodorid nudibranchs. J. Moll. Sttd.57: 5-2L Ellis, A. E. 1,978.British FreshwaterBitsaloeMollusks.Academic
Runham, N. W. and P. J. Hunter. 1.970.Terrestrial Slugs. Press,London.
Hutchinson University Library, London. [A lovely review of Goreau, T. F., N. I. Goreau and C. M. Yonge. 1973.On the uti-
slugbiology.l lization of photosynthetic products from zooxanthellae and
Salvini-Plawen, L. v. and G. Haszprunar.1986. The Vetigastro- of a dissolved amino acid inTridacna maxima.l. Zool. 169
poda and the systematics of streptoneurous Gastropoda. f. 4t7454.
Zool.21L:747-770. Harper, E. M., I. D. Taylor and J. A. Crame, J. A. (eds.) 2000.The
Scheltema, R. S. 1989.Planktonic and non-planktonic develop- eaolutionarybiologyof theBiaalaia.Geologkal Society,London,
ment among prosobranch gastropods and its relationship to SpecialPublications.
the geographic range of species.Pp. 183-188 in J. S. Ryland forgensen, C. B.lg74. On gilt function in the musselMytilus edu-
and P. A. Tyler (eds.),Reproduction,Geneticsand Distribution of lls. Ophelia 13: 187132.
Marine Organisms.Olsen and Olsery Denmark. Judd, W. 1979.The secretionsand fine sfructure of bivalve crys-
Seapy, R. and R. E. Young. 1986. Concealment in epipelagic talline style sacs.Ophelia 18:205-234.
pterotracheid heteropods (Gastropoda) and cranchiid squids Kennedy, W. I., I.D. Taylor and A. Hall. 1969.Environmental
(Cephalopoda).J. Zool. 2I0: 137-147. and biological controls on bivalve shell mineralogy. Biol. Rev.
Sleeper,H. L., V. J. Paul and W. Fenical. 1980.Alarm phero- 44:499-530.
mones from the marine opisthobranch Naoanaxinermis.l. Manahan, D. T., S. H. Wright, G. C. Stephensand M. A. Rice.
Chem. Ecol.6:57-70. 1982. Transport of dissolved amino acids by the mussel,
Spengel,J. W. (1881).Die Geruchsorganeund das Nervensystem Mytilus edulis:Demonstration of net uptake from natural sea-
der Mollusken. Z.Wiss. Zool. AbL A. 35: 333-383. water. Science 215: 1253-1255.
Stanley, S. M. 1982.Gastropod torsion: Predation and the oper- Marincovich, L.,lr.1975. Morphology and mode of life of the
cular imperative. Neues. Jahrb. Geol. Palaeontol. Abh. 164: Late Cretaceousrudist, Coralliochamaorcutti'Nh1te (Mollusca:
95-1,07.[Succinctreview of ideas on why and how torsion Bivalvia). J. Paleontol. 49(L):212-223. [Describes the famous
evolved.l Punta Banda deposits of Baja California, Mexico.l
Tardy,J.1991. Types of opisthobranch veligers: Their notum for- Meyhofer, E. and M. P. Morse. 1996.Characterization of the bi-
mation and torsion. f. Moll. Stud. 57:103-112. valve ultrafiltration system in Myti.Ius edulis,Chlamyshastata,
Taylor, I. (ed.). 1996. Origin and EuolutionaryRadiationof the and Mercenariamercenaria.Invert. Biol. 115(\): 2019.
Mollusca.Oxford Univ. Press,Oxford. Morse, M. P., E. Meyhofer, J. J. Otto and A. M. Kuzirian. 1986'
Taylor, J. D., N. ]. Morris and C. N. Taylor. 1980.Food specializa- Hemocyanin respiratory pigments in bivalve mollusks.
tion and the evolution of predatory prosobranch gastropods. Science 231: 1302-L304.
Paleontology 23: 375410. Morton,8.1978. The diurnal rhythm and the Processesof feed-
Thiriot-Quievreux, C. 1973. Heteropoda. Ann. Rev. Oceanogr. ing and digestion in Tr idacnacrocea. J. ZooL 185: 371187 .
Mar. Biol. 11:237-261. Morton, B. 1978.Feeding and digestion in shipworms. Ann. Rev.
Thompson, T.E. t976. Biologyof Opisthobranch Molluscs,VoL 1. Oceanogr.Mar. Biol. t6:107-144.
Ray Society,London. Owen, G. L974.Feeding and digestion in the Bivalvia. Adv.
Thompson, T. E. and G. H. Brown. 1'976.British Opisthobranch Comp. Physiol. Biochem.5: 1-35.
Molluscs.Academic Press,London. Pojeta,J., Jr. and B. Runnegar. 1974. Fordilla troyensis and the
Thompson, T. E. and G. H. Brown. 1984.Biologyof Opisthobranch early history of pelecypod mollusks. Am. Sci. 62: 706-711.
Molluscs,VoL 2. Ray Society,London. Powell, M. A. and G. N. Somero. 1986.Hydrogen sulfide oxida-
van den Biggelaar, I. A. M. and G. Haszprunar. 1996.Cleavage tion is coupled to oxidative phosphorylation in mitochondria
' of Solemyareidl.Science233: 563-566.
patterns and mesentoblast formation in the Gastropoda: An
evolutionary perspective. Evolution 50: 152VL540. Reid, R. G. B. and F. R. Bernard. 1980.Gutless bivalves. Science
Voltzow, J. and R. Collin. 1995.Flow through mantle cavities 208:609410.
revisited: Was sanitation the key to fissurellid evolution? Reid, R. G. B. and A. M. Reid. 1974.The carnivorous habit of
: 145-150.
lnvert. Biol. 11.4(2) members of the septibranch genus Cuspidaria(Mollusca:
Bivalvia). Sarsia56:47-56.
Bivalvia Stanley, S. M. 1970. Relation of shell form to life habits of the
Ansell, A. D. and N. B. Nair. 1969.A comparison of bivalve bor- Bivalvia (Mollusca). Geol. Soc.Am. Mem. \25:I-296.
ing mechanismsby mechanical means. Am. Zool. 9:857-868. Starrley,S. M.1975. \Atrhyclams have the shape they have: An ex-
Bieler, R. 2010.Classification of bivalve families. Malacologia 52: perimental analysis of burrowing. Paleobiology 1:48.
1,t4-t33. Taylor, I.D. L973.The structural evolution of the bivalve shell.
Bieler, R. and 19 others.2014. Investigating the Bivalve Tree of Paleontology 16: 519-534.
Life-an exemplar-basedapproach combining molecular and Trueman, E. R. 1966.Bivalve mollusks: Fluid dynamics of bur-
novel morphological characters.Invertebr. Syst.28: 32-II5. rowing. Science L52: 52T525.
Beninger, P. G., S. St-Jean,Y. Poussart and J. E. Ward. 1993.Gill Vetter, R. D. 1985.Elemental sulfur in the gills of three species
ftrnction and mucocyte distribution in Placopectenmagellanicus of clams containing chemoautotrophic symbiotic bacteria: A
possible inorganic energy storage compound. Mar- Biol. 88: Mutvei, H. 1964.On the shells olNautilus and Spirula with notes
3342. on the shell secretionin non-cephalopod molluscs. Ark. Zool.
Waterbury, 1. 8., C. B. Calloway and R. D' Turner. 1983.A cel- 1.6fl4\:223178.
lulolytic nitrogen-fixing bacterium cultured from the gland of Nixon, M. and J. B. Messenger (eds.). L977. The Biology of
Deshayes in shipworms (Bivalvia: Teredinidae). Science221: CEhalopods.Academic Press,New York.
t40r-1403. Packard, A.1972. Cephalopods and fish: The limits of conver-
Wilkens, L. A. 1986.The visual system of the giantclamTridacna: gence.Biol. Rev. 47:241-307.
Behavioral adaptafions. Biol. Bull' \70: 393408. Roper, C. F. E. and K. J. Boss.1982.The giant squid. Sci.Am.246:
Yonge, C. M. 1953. The monomyarian condition in the 96-1,04.
Lamellibranchia. Trans. R. Soc.Edinburyh62 (p.II): 443478. Saunders, W. B. 1983.Natural rates of growth and longevity of
Yonge,C. M.1973. Giant clams.Sci.Am. 232:96-105. N autilus belauensis. Paleobiology 9: 280-288.
Saunders,W. B., R. L. Knight and P. N. Bond. 199L.Octopus pre-
Scaphopoda dation on Naufllus: Evidence from Papua New Guinea' Bull.
Bilyard, G.R.1974. The feeding habits and ecology of Dentalium Mar. Sci.49(1-2): 280-287.
stimpsoni.Veliger 17: 126-138. Saunders,W. B. and N. H. Landman. 2009.Nautilus: TheBiology
Gainey, L.F.1972. The use of the foot and captacula in the feed- and Paleobiologyof a Lioing Fossil, Reprint with Additions.
ing oI Dentalium.Yeliger 15:2914. Springer Science& BusinessMedia.
Reynolds, P. D. 2002. The Scaphopoda. Adv. Mar. Biol. 42: Sweeney,M. I., C.F. E. Roper, K. A. M. Mangold, M. R. Clarke
L37-236. and S. V. Boletzky (eds.). 1992."Larval" andjuvenile cepha-
Steiner, G.1992. Phylogeny and classification of Scaphopoda.J. lopods: a manual for their identification. Smithsonian Contrb.
Mollus. Stud. 58: 385-400. Zool.5l3:1-282.
Trueman, E. R. 1968. The burrowing Processof Dentalium.l. Ward, P. D. lgST.TheNaturalHistory ofNautilus. Allen & Unwin,
Zool.1.54:19-27. Boston.
Ward, P. D. and W. B. Saunders. 1997.Allonautllus, a new genus
Cephalopoda of living nautiloid cephalopod and its bearing on phylogeny
Aronson, R. B. 1991.Ecology, paleobiology and evolutionary of the Nautilida. f. Paleontol. 7l:L054-L064.
constraint in the octopus. Bull. Mar. Sci.49(1-2): 245+55. Ward, R., L. Greenwald, and O. E. Greenwald. L980.The buoy-
Barber, V. C. and F. Grazialdei . 7967. The fine structure of ancy of the chambered nautilus' Sci.Am. 243:190204.
cephalopod blood vessels. Z. Zellforcch. Mikrosk' Anat. 77: Wells, M. f. 1978.Octopus:PhysiologyandBehaztiour of an Adaanced
162-174.[Also seeearlier papers by these authors in the same Inaertebrate,Chapman and Hall, New York.
joumal.l Wells, M. j. and R. K. O'Dor. 1991.Jet propulsion and the evolu-
Boyle, P. R. (ed.). 7983.Cephalopod Life Cycles,VoIs.1-2. Academic tion of cephalopods.Bull. Mar. Sci. 49(1'-2):419432.
Press,New York. Young, l. Z. 1972.TheAnatomy of the NeroousSystemof Octop'ts
Boyle, P. R. and S. v. Boletzky. 1996.Cephalopod populations: vulgaris. Oxford University Press,New York.
Definition and dynamics. Phil. Trans. Roy. Soc' B, BioI' Sci. Young, R. E. and F. M. Mencher. 1980.Bioluminescencein meso-
351:985-1002. pelagic squid: Diel color change during counterillumination.
Boyle, P. and P. Rodhouse. 2005. Cephalopods: Ecologyand Science208: I28G1288.
Fisheries.Blackwell ScienceLtd., Oxford.
Clarke, M. A. 1966.A review of the systematics and ecology of Molluscan Evolution and Phylogeny
oceanicsquids.Adv. Mar. Biol.4:91-300. Batten, R. L., H. B. Rollins and S. J. Gould. 1967.Comments on
Cloney, R. A. and S. L. Brocco. 1983.Chromatophore organs, re- "The adaptive significance of gastropod torsion." Evolution
flector cells, irridocytes and leucophores in cephalopods.Am. 2l:405406.
Zool.23:581 592. Bieler, R. 1992.Gastropod phylogeny and systematics.Ann. Rev.
Denton, E. f . and f . B. Gilpin-Btown. 1-973.Flotation mecha- Ecol. Syst. 23: 311.138.
nisms in modern and fossil cephalopods. Adv' Mar. Biol' 11: Bieler, R. and 19 others. 2014. Investigating the Bivalve Tree of
r97+64. Life-an exemplar-basedapproach combining molecular and
Fields, W. G. L965.The structure, development, food relations, novel morphological characters.Invertebr. Syst.28: 32-IL5.
reproduction, and life history of the squid Loligo opalescens Dunn, C. W. and 17 others. 2008.Broad phylogenomic sampling
Berry. Calif. Dept. Fish Game Bull. 131:1-108. imoroves resolution of the animal tree of life. Nature 452:745.
Fiorito, G. and P. Scotto. lgg2.Observational learning tnOctopus Ounn, C. W., G. Giribet, G. D. Edgecombeand A. Hejnol.2014.
oulgaris.Science256: 545-547. Animal phylogeny and its evolutionary implications. Ann.
Hanlon, R. T. and J. B. Messenger.1,996.Cephalopod Behattior. Rev.Eco.Evol. S.45:37L195.
Cambridge Univ. Press. Edgecombe,G. D. and 8 others. 2011.Higher-level metazoan re-
Hochberg, F. G. 1983.The parasitesof cephalopods:A review. lationships: recent progress and remaining questions. Org'
Mem. Nat. Mus. Victoria Melboume 44: 109-145. Divers. Evol. 11:I5I-172.
House, M. R. and f . R. Senior. 7981'.The Ammonoidea:The Eldredge, N. and S. M. Stanley (eds.). 1984. Liaing Fossils.
Eaolution,Classification, oJ
Mode of Life and GeologicalUsefulness S p r i n g e r - V e r l a g , N e w Y o r k . [ I n c l u d e s ch a p te r s o n
a Major FossilGroup. Academic Press,New York' Monoplacophora, Pleurotomaria, and N autilus.l
Kier, W. M. 1991.Squid cross-striatedmuscle: The evolution of a Fedonkin, M. A. and B. M. Waggoner. 1997. The Late Precam-
specialized muscle fiber type. Bull' Mar. Sci.49(1'-2):389-403. brian fossil Kimberellais a mollusc-like bilaterian organism.
Kier, W. M. and A. M. Smith. 1990.The morphology and me- Nature 388: 868-871.
chanics of octopus suckers.Biol. Bull. 178:126-136. Garstang, W. [Introduction by Sir A. Hardy]. t95I- Laraal
Lehmann, U. 1981. The Ammonites:Their Life and Their World' Forms,and Other ZoologicalVerses.Basil Blackwell, Oxford.
Cambridge University Press,New York' [Reprinted in 1985by the University of Chicago Press.]
McFell-Ngai, M. and M. K. Montgomery. 1990.The anatomy and Ghisetin, M. T.1966. The adaptive significance of gastropod tor-
morphology of the adult bacterial light organ of Euprymna sion. Evolution 20: 337-348.
scolopesBerry (Cephalopoda: Sepiolidae)' Biol. Bull. 179: Giribet, G. and W. Wheeler. 2002.On bivalve phylogeny: a high-
332139. level analysis of the Bivalvia (Mollusca) based on combined
PHYLUMMOLLUSCA 529
morphology and DNA sequencedata. Invertebr. Biol. 121: Ponder, W. F. and D. R. Lindberg .2008. Phylogenyand Eaolution
27r-324. of theMollusca.Universify of California Press,Berkeley, CA.
Giribet, G., A. Okusu, A. R. Lindgren, S. W. Huff, M. Schrodl Runnegar, B. and f . Pojeta.1974.Molluscan phylogeny: The pale-
and M. K. Nishiguchi. 2006.Evidence for a clade composed ontological viewpoint. ScienceL86:3tl1l7.
of molluscs with serially repeated structures: monoplacopho- Ruppert E. E. and J. Carle. 1983.Morphology of metazoan circu-
rans are related to chitons. PNAS 103:7723-7728. latory systems.Zoomorph. 103:193-208.
Giribet, G.20L4. On Aculifera: a review of hypotheses in tribute Salvini-Plawen, L. v. 1977. On the evolution of photoreceptors
to Christoffer Schander.J. Nat. Hist. 48:2739-2749. alrd eyes.Evol. Biol. 10:207-263.
Gotting, K. 1980. Arguments concerning the descendenceof Salvini-Plawen, L., v. 1990.Origin, phylogeny and classification
Mollusca from metameric ancestors. Zool.Iahrb. Abt. Anat. of the phylum Mollusca. Iberus 9: 1- 33.
L03:21L-218. Scheltema,A.H.1978. Position of the classAplacophora in the
Gcitting, K. 1980.Origin and relationships of the Mollusca. Z. phylum Mollusca. Malacologia 17: 99-109.
Zool. Syst.Evolutionsforsch. 78: 24-27. Scheltema,A. H. 1988.Ancestors and descendants:Relationships
Graham, A. 1979.Gastropoda. Pp. 359-365 in M. R. House (ed.), of the Aplacophora and Polyplacophora. Am. Malacol. Bull.
TheOrigin of Major InztertebrateGroups.Academic Press,New 6:57-68.
York. Scheltema,A. H. 1993.Aplacophora as progenetic aculiferans
Gutmann, W . F. 1974. Die Evolution der Mollusken-Konstruc- and the coelomate origin of mollusks as the sister taxon of
tion: Ein phylogenetisches Modell. Aufsiitze Red. Senckenb. Sinuncula. Biol. Bull. L84:57-78.
Naturf. Ges.25:TJL Scheiholz, M., E. Redl, T. Wollesen, C. Todt and A. Wanningei.
Haas, W. 1981..Evolution of calcareoushardparts in primitive 2013. Aplacophoran mollusks evolved from ancestors with
molluscs. Malacologia 21: 403-418. polyplacophoran-like features.Curr. Biol. 23: 1-5.
Haszprunar, G.1992. The first molluscs-small animals. Boll. Scherholz, M., E. Redl, T. Wollesen, C. Todt and A. Wanninger.
ZooL59:1-76. 2015.From complex to simple: myogenesis in an aplacopho-
Haszprunar, G.2000. Is the Aplacophora monophyletic? A cla- ran mollusk reveals key traits in aculiferan evolution. BMC
distic point of view. Am. Malacol. Bull. 15(2):115-130. Evol.Biol. 15(1):201.
Hejnol, A. 2010. A twist in time-The evolution of spiral cleav- Schrcidl,M. and I. Stoger. 2014.A review on deep molluscan
age in the light of animal phylogeny. Integr. Comp. Biol. 50: phylogeny: old markers, integrative approaches, persistent
695-706. problems. J. Nat. Hist. 48:L-32.
Hickman, C. S. 1988.Archaeogastropod evolution, phylogeny Sharma, P. P. and 12 others. 2012.Phylogenetic analysis of four
and systematics:A re-evaluation. Malacol. Rev., Suppl. 4: nuclear protein-encoding genes largely corroborates the tra-
1714. difional classificationof Bivalvia (Mollusca). Mol. Phylogenet.
Holland, C.H. 1979.Early Cephalopoda. Pp. 367-379in M. R. EvoL 65:64-74.
House (ed.), TheOrigin of Major InaertebrateGroups.Academic Sigwart, J. D., I. Stoeger,T. Knebelsberger and E. Schwabe.2013.
Press,New York. Chiton phylogeny (Mollusca: Polyplacophora) and the place-
Ivantsov, A. 2012. Paleontological data on the possibil- ment of the enigmatic speciesChoriplaxgrayl (H. Adams and
ity of Precambrian existenceof mollusks. Pp. 153-179in A. Angas). Invertebr. Syst.27: 603-621'.
Fyodorov and H. Yakovlev (eds.), Mollusks: Morphology, Sigwart, I. D., C. Todt and A. H. Scheltema. 201.4.Who are the
Behatsior,andEcolo3y.Nova SciencePublishing, New York. 'Aculifera'? J. Nat. Hist. 48:273T2737.
Johnson,C. C.2002. The rise and fall of rudistid reefs.Amer. Sci. Sigwart, J. D. and D. R. Lindberg. 2015.Consensusand confusion
90:148-153. in molluscan trees: evaluating morphological and molecular
Jorger, K. M., I. Stoger,Y. Kano, H. Fukuda, T. Knebelsberger phylogenies. Syst.Biol. 64:384:395.
and M. Schrcidl. 2010.On the origin of Acochlidia and other Smith, S. A. and 7 others. 2011.Resolving the evolutionary rela-
enigmatic euthyneuran gastropods, with implications for the tionships of molluscs with phylogenomic tools. Nature 480:
systematicsof Heterobranchia. BMC Evol. Biol. 10:323. 364-367.
Kocot, K. M. and 10 others. 2011.Phylogenomics reveals deep Stciger,I. and M. Schrodl. 2013.Mitogenomics does not resolve
molluscan relationships. Nature 477: 452456. deep molluscan relationships (yet?). Mol. Phylogenet. Evol.
Kocot, K. M. 2013.Recent advances and unanswered questions 69:376192.
in deep molluscan phylogenetics. Am. Malacol. Bull. 31: Struck, T. H. and 12 others. 2014.Platyzoan paraphyly based
195-208. on phylogenomic data supports a noncoelomate ancestry of
Lauterbach, K.-E. von. 1983. Erorterungen zur Stammes- Spiralia. Mol. Biol. Evol.31: 1,833-1,849.
geschichte der Mollusca, insbesondere der Conchifera. Z. Sutton, M. D. and J. D. Sigwart. 201.2.A chiton without a foot.
Zool. Syst.Evolutionsfors ch. 21: 201-2L6. Palaeontology 55:40'14L1.
Lindberg, D. R. and M. T. Ghiselin. 2003.Fact, theory and tradi- Taylor, J. D. (ed.). L996. Origin and EztolutionaryRadiation of the
tion in the study of molluscan origins. PCAS 54: 663-{,86 Mollusca.New York, Oxford University Press.
Lindgren, A. R., G. Giribet and M. K. Nishiguchi.2004. A com- Trueman, E. R. and M. R. Clarke. 7985.Ettolution. TheMollusca,
bined approach to the phylogeny of Cephalopoda (Mollusca). 10.New York, Academic Press.
Cladistics 20: 454486. Steiner, G. and M. Mriller. 1996.What can L8SrDNA do for bi-
Linsley, R. M. 1978.Shell form and the evolution of gastropods. valve phylogeny? J. Mol. Evol. 43: 58-70.
Am. Sci.66:432441. Vagvolgyi, I. 1967. On the origin of molluscs, the coelom, and
Nesnidal, M. P. and 9 others. 2013. New phylogenomic data coelomic segmentation.Syst. Zool. 16: L53-168.
support the monophyly of Lophophorata and an Ectoproct- Wade, C. M., P. B. Mordan and B. Clarke.2001.A phylogeny of
Phoronid clade and indicate that Polyzoa and Kryptrochozoa the land snails (Gastropoda: Pulmonata). Proc. Royal Soc.
are causedby systematicbias. BMC Evol. Biol. 13:253. London B. 268:4L3422.
Okusu, A., E. Schwabe,D. l.Eernisse and G. Giribet. 2003. Wagner, P. J.2001.Gasfropod phylogenetics: Progress,problems
Towards a phylogeny of chitons (Mollusca, Polyplacophora) and implications. J. Paleontol. 75(6):712U11'40.
based on combined analysis of five molecular loci. Org. Wilson, N. G., G. W. Rouse and G. Giribet. 2010.Assessing
Divers. Evol. 3: 287102. the molluscan hypothesis Serialia (Monoplacophora +
530 ChapterThirteen
Polyplacophora) using novel molecular data. Mol. Phylogene. Yonge, C. M. 1957. Reflections on the monoplacophoran
Evol.54: 187-193. NeopilinagalatheaeLemdne.Nature 179 672473.
Wingstrand, K. G. 1985.On the anatomy and relationships of re- Zapata, F. and 8 others. 2014. Phylogenomic analyses of deep
cent Monoplacophora. Galathea Rpt. 16:7-94. gastropod relationships reject Orthogastropoda. proc. Royal
Yonge, C. M. 1,957.Neopilina:Survival from the paleozoic. Soc.London B. 281. doi: 1,0.1098/ r spb.20l4.I739
Discovery (London), June 1957: 255-256.

Classification of The Animal Kingdom (Metazoa)

Uploaded by

Document Information

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Classification of The Animal Kingdom (Metazoa)

Uploaded by

Copyright:

Available Formats

olluscsinclude someof the best-knowninvertebrates;almost everyone

is familiar with snails,clams,slugs,squids,and octopuses.Molluscan

and bivalves are also burdened with numerous names C OH OR TH Y P S OGA S TR OP OD A

SUBCLASSVETIGASTROPODA "Pdmitive"marinetop- MEGAORDER PALAEOHETERODONTAFTeshwateT

rMultitudes of extinct molluscs have been described. Perhaps the

(c) Radular Dorsal Gonopericardial

Oral shield Mantle cavity

Ventral 1= pedal) Lateral

SYNOPSES OF MOLLUSCAN GROUPS CLASSSOLENOGASTRES (= NEOMENIOMORPHA) (Figure

(L) Cerebral Buccal Lateral

invertebrates. are proba-

Minute preoral tentacle

Lateral nerve cord

Pedal nerve cord

Lateropedal sometimes haveshelleyes(Figure13.43C,D). Marine,inter-

Main nephridial (G)

(A) Mantle edge papillae protruding (c) Right nephridium

Figure 13.5 General anatomy of limpet-like gastropods.

Posterior aorta Figure 13.6 Generalanatomy of coiled gastropods.

Cephalic Mantle edge

lip Buccal mass

Figure 13.7 More gastropod anatomy; some caenogastropods

(B) Esophagus (D) Foot

Median keel on shell

(F) Branchial plume

dividedinto two main groups,Cerithiomorpha

nudibranchs) whichincludesmanyfamilies, someex- freshwater species)andthe ectoparasitic hTramidelli-

Nerve Right atrium

Right visceral ganglion

SUPERORDER NUCULIFORMII (= OPPONO-BRAN- ORDER OSTREIDAThetrueoystersOstreidae(true

Figure 13.9 General anatomy of a scaphopod.

SUBCLASS NEOCEPHALOPODA Includesone fossil

(A) Posterior Posterior (c) \-J Funnel

Polyplacophorans,or chitons,are oval molluscsthat

Figure 13.1 2. The anatomy of Octopus. Dorsal mantle cavitY

Ctenidium Muscular septum of Anus

Figure 13.13 Modifications of the shell, foot, gut, (A)

(F) Shell aperture

Bivalves possess a greatLy enlarged mantle cav-

posttorsional gonad. The right renopericardial duct serves

Subepidermal Eosinophilous (B)

Insertion plate Perioshacum (folded

Dorsal Anterior margin Portion

Figure 13.17 Two very different kinds of cephalopod

Figure 13.1 8 Pre- and post-torsional adult gastropods- Pleural

vent of torsion, a unique synapomolphy of gastropods,

Figure 13.20 (A*D) Burrowing and life

Figure 13.22 Swimming cephalopods. (A) Sepra, the

Figure 13.23 A generalized molluscan radula and asso- Esophagus

some species.The odontophore itself is moved in and teeth

Figure 13.24 Feeding in macrophagous molluscs.

/M\arginal teeth (D)

Figure 13.25 Various arrangements of radular teeth.

that live on diatoms and other algae and microbes on marginalteethaltogether(Figures13.25Eand13.26c,D).

Figure 13.26 Gastropod radulae.

occur between anemone fishes and their host anemo-

(A) Visceral mass (B) (c)

Figure 13-28 Arrangementof ctenidia in some bivalves

area in the posterior mantle cavity to be discharged; the

cirri cilia cilia

Figure 13.30 Ctenidial structure in bivalve molluscs. In all

Figure 13.31 Feeding in the septibranch bivalve

Sphincter Outer lip