Mollusca

Mollusca
Mollusca is t he second-largest phylum of invert ebrat e animals, aft er Art hropoda; members are
known as molluscs or mollusks [a] (/ˈmɒləsks/). Around 76,000 ext ant species of molluscs are
recognized.[3] The number of fossil species is est imat ed bet ween 60,000 and 100,000 addit ional
species.[4] The proport ion of undescribed species is very high. Many t axa remain poorly st udied.[5]
Cornu aspersum (formerly Helix

aspersa) – a common land snail
Molluscs are t he largest marine phylum, comprising about 23% of all t he named marine organisms.
Numerous molluscs also live in freshwat er and t errest rial habit at s. They are highly diverse, not just in
size and anat omical st ruct ure, but also in behaviour and habit at . The phylum is t ypically divided int o 7
or 8[6] t axonomic classes, of which t wo are ent irely ext inct . Cephalopod molluscs, such as squid,
cut t lefish, and oct opuses, are among t he most neurologically advanced of all invert ebrat es—and
eit her t he giant squid or t he colossal squid is t he largest known ext ant invert ebrat e species. The
gast ropods (snails and slugs) are by far t he most diverse molluscs and account for 80% of t he t ot al
classified species.
The four most universal feat ures defining modern molluscs are a body largely consist ing of solid
muscle, a mant le wit h a significant cavit y used for breat hing and excret ion, t he presence of a radula
(except for bivalves), and t he st ruct ure of t he nervous syst em. Ot her t han t hese common element s,
molluscs express great morphological diversit y, so many t ext books base t heir descript ions on a
"hypot het ical ancest ral mollusc" (see image below). This has a single, "limpet -like" shell on t op, which
is made of prot eins and chit in reinforced wit h calcium carbonat e, and is secret ed by a mant le
covering t he whole upper surface. The underside of t he animal consist s of a single muscular "foot ".
Alt hough molluscs are coelomat es, t he coelom t ends t o be small. The main body cavit y is a
hemocoel t hrough which blood circulat es; as such, t heir circulat ory syst ems are mainly open. The
"generalized" mollusc's feeding syst em consist s of a rasping "t ongue", t he radula, and a complex
digest ive syst em in which exuded mucus and microscopic, muscle-powered "hairs" called cilia play
various import ant roles. The generalized mollusc has
Mollusca
t wo paired nerve cords, or t hree in bivalves. The brain,
Temporal range:
in species t hat have one, encircles t he esophagus.
Most molluscs have eyes, and all have sensors t o
det ect chemicals, vibrat ions, and t ouch. The simplest
t ype of molluscan reproduct ive syst em relies on
ext ernal fert ilizat ion, but more complex variat ions
Helix pomatia, a Common
occur. Nearly all produce eggs, from which may
gast ropod oct opus, a
emerge t rochophore larvae, more complex veliger
cephalopod
larvae, or miniat ure adult s. The coelomic cavit y is
reduced. They have an open circulat ory syst em and
kidney-like organs for excret ion.
Good evidence exist s for t he appearance of At lant ic surf clam, Tonicella lineata, a
gast ropods, cephalopods, and bivalves in t he a bivalve chit on
Cambrian period, 541–485.4 million years ago.
However, t he evolut ionary hist ory bot h of molluscs' Scientific classification
emergence from t he ancest ral Lophot rochozoa and
Domain: Eukaryot a
of t heir diversificat ion int o t he well-known living and
fossil forms are st ill subject s of vigorous debat e Kingdom: Animalia
among scient ist s.
Subkingdom: Eumet azoa
Clade: ParaHoxozoa
Clade: Bilat eria
Clade: Nephrozoa
(unranked): Prot ost omia
(unranked): Spiralia
Superphylum: Lophot rochozoa
Fossilized ammonite displayed at the Phylum: Mollusca

National Museum of the Philippines
Linnaeus, 1758
Molluscs have been and st ill are an import ant food
source for anat omically modern humans. Toxins t hat Classes
can accumulat e in cert ain molluscs under specific

condit ions creat e a risk of food poisoning, and many
See t ext .
jurisdict ions have regulat ions t o reduce t his risk.
Molluscs have, for cent uries, also been t he source of
Diversity[1]
import ant luxury goods, not ably pearls, mot her of
pearl, Tyrian purple dye, and sea silk. Their shells have 85,000 recognized living species.
also been used as money in some preindust rial
societ ies.
A handful of mollusc species are somet imes considered hazards or pest s for human act ivit ies. The
bit e of t he blue-ringed oct opus is oft en fat al, and t hat of Octopus apollyon causes inflammat ion
t hat can last over a mont h. St ings from a few species of large t ropical cone shells of t he family
Conidae can also kill, but t heir sophist icat ed, t hough easily produced, venoms have become
import ant t ools in neurological research. Schist osomiasis (also known as bilharzia, bilharziosis, or snail
fever) is t ransmit t ed t o humans by wat er snail host s, and affect s about 200 million people. Snails
and slugs can also be serious agricult ural pest s, and accident al or deliberat e int roduct ion of some
snail species int o new environment s has seriously damaged some ecosyst ems.
Etymology
The words mollusc and mollusk are bot h derived from t he French mollusque, which originat ed from
t he post -classical Lat in mollusca, from mollis, soft , first used by J. Jonst on (Hist oriæ Nat uralis,
1650) t o describe a group comprising cephalopods.[7] Molluscus is used in classical Lat in as an
adject ive only wit h nux (nut) t o describe a part icular t ype of soft nut . The use of mollusca in
biological t axonomy by Jonst on and lat er Linnaeus may have been influenced by Arist ot le's τὰ
μαλάκια ta malákia (t he soft ones; < μαλακός malakós "soft "), which he applied inter alia t o
cut t lefish.[8][9] The scient ific st udy of molluscs is accordingly called malacology.[10]
The name Molluscoida was formerly used t o denot e a division of t he animal kingdom cont aining t he
brachiopods, bryozoans, and t unicat es, t he members of t he t hree groups having been supposed t o
somewhat resemble t he molluscs. As now known, t hese groups have no relat ion t o molluscs, and
very lit t le t o one anot her, so t he name Molluscoida has been abandoned.[11]
Definition
The most universal feat ures of t he body st ruct ure of molluscs are a mant le wit h a significant body
cavit y used for breat hing and excret ion, and t he organizat ion of t he nervous syst em. Many have a
calcareous shell.[12]
Molluscs have developed such a varied range of body st ruct ures, finding synapomorphies (defining
charact erist ics) t o apply t o all modern groups is difficult .[13] The most general charact erist ic of
molluscs is t hey are unsegment ed and bilat erally symmet rical.[14] The following are present in all
modern molluscs:[15][16]
The dorsal part of t he body wall is a mant le (or pallium) which secret es calcareous spicules,
plat es or shells. It overlaps t he body wit h enough spare room t o form a mant le cavit y.
The anus and genit als open int o t he mant le cavit y.
There are t wo pairs of main nerve cords.[17]
Ot her charact erist ics t hat commonly appear in t ext books have significant except ions:
Whet her charact erist ic is found in t hese classes of Molluscs
Supposed
universal Aplacophora Polyplacophora Monoplacophora Gast ropoda Cephalopoda Bivalvia S
[17]: 291–292 [17]: 292–298 [17]: 298–300 [17]: 300–343 [17]: 343–367 [17]: 367–403
Molluscan
charact erist ic [15]
In
Radula, a rasping
Absent in 20% of c
"tongue" wit h Yes Yes Yes Yes No
Neomeniomorpha e
chit inous t eet h
b
Broad, muscular Reduced or Modified into S

Yes Yes Yes Yes
foot absent arms "
Dorsal
concent rat ion of
Not obvious Yes Yes Yes Yes Yes Y
int ernal organs
(visceral mass)
Large digest ive No ceca in some

Yes Yes Yes Yes Yes N
ceca Aplacophora
Large complex
met anephridia None Yes Yes Yes Yes Yes S
("kidneys")
Snails, yes;
Octopuses,
Primitive forms, slugs,
One or more no; cuttlefish,
yes; modern Yes Yes mostly yes Yes Y
valves/ shells nautilus,
forms, no (internal
squid, yes
vestigial)
Odontophore Yes Yes Yes Yes Yes No Y

Diversity
Diversity and variability of shells of

molluscs on display
About 80% of all known mollusc

species are gastropods (snails and
slugs), including this cowry (a sea
snail).[18]
Est imat es of accept ed described living species of molluscs vary from 50,000 t o a maximum of
120,000 species.[1] The t ot al number of described species is difficult t o est imat e because of
unresolved synonymy. In 1969, David Nicol est imat ed t he probable t ot al number of living mollusc
species at 107,000 of which were about 12,000 fresh-wat er gast ropods and 35,000 t errest rial. The
Bivalvia would comprise about 14% of t he t ot al and t he ot her five classes less t han 2% of t he living
molluscs.[19] In 2009, Chapman est imat ed t he number of described living mollusc species at
85,000.[1] Haszprunar in 2001 est imat ed about 93,000 named species,[20] which include 23% of all
named marine organisms.[21] Molluscs are second only t o art hropods in numbers of living animal
species[18] — far behind t he art hropods' 1,113,000 but well ahead of chordat es'
52,000.[17]: Front endpaper About 200,000 living species in t ot al are est imat ed,[1][22] and 70,000 fossil
species,[15] alt hough t he t ot al number of mollusc species ever t o have exist ed, whet her or not
preserved, must be many t imes great er t han t he number alive t oday.[23]
Molluscs have more varied forms t han any ot her animal phylum. They include snails, slugs and ot her
gast ropods; clams and ot her bivalves; squids and ot her cephalopods; and ot her lesser-known but
similarly dist inct ive subgroups. The majorit y of species st ill live in t he oceans, from t he seashores t o
t he abyssal zone, but some form a significant part of t he freshwat er fauna and t he t errest rial
ecosyst ems. Molluscs are ext remely diverse in t ropical and t emperat e regions, but can be found at
all lat it udes.[13] About 80% of all known mollusc species are gast ropods.[18] Cephalopoda such as
squid, cut t lefish, and oct opuses are among t he most neurologically advanced of all invert ebrat es.[24]
The giant squid, which unt il recent ly had not been observed alive in it s adult form,[25] is one of t he
largest invert ebrat es, but a recent ly caught specimen of t he colossal squid, 10 m (33 ft ) long and
weighing 500 kg (1,100 lb), may have overt aken it .[26]
Freshwat er and t errest rial molluscs appear except ionally vulnerable t o ext inct ion. Est imat es of t he
numbers of non-marine molluscs vary widely, part ly because many regions have not been t horoughly
surveyed. There is also a short age of specialist s who can ident ify all t he animals in any one area t o
species. However, in 2004 t he IUCN Red List of Threat ened Species included nearly 2,000
endangered non-marine molluscs. For comparison, t he great majorit y of mollusc species are marine,
but only 41 of t hese appeared on t he 2004 Red List . About 42% of recorded ext inct ions since t he
year 1500 are of molluscs, consist ing almost ent irely of non-marine species.[27]
Anatomy
Anatomical diagram of a hypothetical ancestral mollusc
Because of t he great range of anat omical diversit y among molluscs, many t ext books st art t he
subject of molluscan anat omy by describing what is called an archi-mollusc, hypothetical generalized
mollusc, or hypothetical ancestral mollusc (HAM) t o illust rat e t he most common feat ures found
wit hin t he phylum. The depict ion is visually rat her similar t o modern monoplacophorans.[13][16][28]
The generalized mollusc is an unsegment ed, bilat erally symmet rical animal and has a single, "limpet -
like" shell on t op. The shell is secret ed by a mant le covering t he upper surface. The underside
consist s of a single muscular "foot ".[16] The visceral mass, or visceropallium, is t he soft , nonmuscular
met abolic region of t he mollusc. It cont ains t he body organs.[14]
Mantle and mantle cavity
The mant le cavit y, a fold in t he mant le, encloses a significant amount of space. It is lined wit h
epidermis, and is exposed, according t o habit at , t o sea, fresh wat er or air. The cavit y was at t he rear
in t he earliest molluscs, but it s posit ion now varies from group t o group. The anus, a pair of osphradia
(chemical sensors) in t he incoming "lane", t he hindmost pair of gills and t he exit openings of t he
nephridia (kidneys) known as "Organs of bojanus" and gonads (reproduct ive organs) are in t he mant le
cavit y.[16] The whole soft body of bivalves lies wit hin an enlarged mant le cavit y.[14]
Shell
The mant le edge secret es a shell (secondarily absent in a number of t axonomic groups, such as t he
nudibranchs[14]) t hat consist s of mainly chit in and conchiolin (a prot ein hardened wit h calcium
carbonat e),[16][29] except t he out ermost layer, which in almost all cases is all conchiolin (see
periost racum).[16] Molluscs never use phosphat e t o const ruct t heir hard part s,[30] wit h t he
quest ionable except ion of Cobcrephora.[31] While most mollusc shells are composed mainly of
aragonit e, t hose gast ropods t hat lay eggs wit h a hard shell use calcit e (somet imes wit h t races of
aragonit e) t o const ruct t he eggshells.[32]
The shell consist s of t hree layers: t he out er layer (t he periost racum) made of organic mat t er, a
middle layer made of columnar calcit e, and an inner layer consist ing of laminat ed calcit e, oft en
nacreous.[14]
In some forms t he shell cont ains openings. In abalones t here are holes in t he shell used for
respirat ion and t he release of egg and sperm, in t he naut ilus a st ring of t issue called t he siphuncle
goes t hrough all t he chambers, and t he eight plat es t hat make up t he shell of chit ons are
penet rat ed wit h living t issue wit h nerves and sensory st ruct ures.[33]
Foot
0:51
A 50-second video of snails (most

likely Natica chemnitzi and Cerithium
stercusmuscaram) feeding on the sea
floor in the Gulf of California, Puerto
Peñasco, Mexico
The body of a mollusc has a vent ral muscular foot , which is adapt ed t o different purposes
(locomot ion, grasping t he subst rat um, burrowing or feeding) in different classes.[34] The foot carries
a pair of st at ocyst s, which act as balance sensors. In gast ropods, it secret es mucus as a lubricant
t o aid movement . In forms having only a t op shell, such as limpet s, t he foot act s as a sucker
at t aching t he animal t o a hard surface, and t he vert ical muscles clamp t he shell down over it ; in
ot her molluscs, t he vert ical muscles pull t he foot and ot her exposed soft part s int o t he shell.[16] In
bivalves, t he foot is adapt ed for burrowing int o t he sediment ;[34] in cephalopods it is used for jet
propulsion,[34] and t he t ent acles and arms are derived from t he foot .[35]
Circulatory system
Most molluscs' circulat ory syst ems are mainly open, except for cephalopods, whose circulat ory
syst ems are closed. Alt hough molluscs are coelomat es, t heir coeloms are reduced t o fairly small
spaces enclosing t he heart and gonads. The main body cavit y is a hemocoel t hrough which blood and
coelomic fluid circulat e and which encloses most of t he ot her int ernal organs. These hemocoelic
spaces act as an efficient hydrost at ic skelet on.[14] The blood of t hese molluscs cont ains t he
respirat ory pigment hemocyanin as an oxygen-carrier. The heart consist s of one or more pairs of at ria
(auricles), which receive oxygenat ed blood from t he gills and pump it t o t he vent ricle, which pumps it
int o t he aort a (main art ery), which is fairly short and opens int o t he hemocoel.[16] The at ria of t he
heart also funct ion as part of t he excret ory syst em by filt ering wast e product s out of t he blood and
dumping it int o t he coelom as urine. A pair of met anephridia ("lit t le kidneys") t o t he rear of and
connect ed t o t he coelom ext ract s any re-usable mat erials from t he urine and dumps addit ional
wast e product s int o it , and t hen eject s it via t ubes t hat discharge int o t he mant le cavit y.[16]
Except ions t o t he above are t he molluscs Planorbidae or ram's horn snails, which are air-breat hing
snails t hat use iron-based hemoglobin inst ead of t he copper-based hemocyanin t o carry oxygen
t hrough t heir blood.
Respiration
Most molluscs have only one pair of gills, or even only a singular gill. Generally, t he gills are rat her like
feat hers in shape, alt hough some species have gills wit h filament s on only one side. They divide t he
mant le cavit y so wat er ent ers near t he bot t om and exit s near t he t op. Their filament s have t hree
kinds of cilia, one of which drives t he wat er current t hrough t he mant le cavit y, while t he ot her t wo
help t o keep t he gills clean. If t he osphradia det ect noxious chemicals or possibly sediment ent ering
t he mant le cavit y, t he gills' cilia may st op beat ing unt il t he unwelcome int rusions have ceased. Each
gill has an incoming blood vessel connect ed t o t he hemocoel and an out going one t o t he heart .[16]
Eating, digestion, and excretion
Snail radula at work

= Food = Radula
= Muscles
= Odontophore "belt"
Molluscs use int racellular digest ion. Most molluscs have muscular mout hs wit h radulae, "t ongues",
bearing many rows of chit inous t eet h, which are replaced from t he rear as t hey wear out . The radula
primarily funct ions t o scrape bact eria and algae off rocks, and is associat ed wit h t he odont ophore, a
cart ilaginous support ing organ.[14] The radula is unique t o t he molluscs and has no equivalent in any
ot her animal.
Molluscs' mout hs also cont ain glands t hat secret e slimy mucus, t o which t he food st icks. Beat ing
cilia (t iny "hairs") drive t he mucus t owards t he st omach, so t he mucus forms a long st ring called a
"food st ring".[16]
At t he t apered rear end of t he st omach and project ing slight ly int o t he hindgut is t he prost yle, a
backward-point ing cone of feces and mucus, which is rot at ed by furt her cilia so it act s as a bobbin,
winding t he mucus st ring ont o it self. Before t he mucus st ring reaches t he prost yle, t he acidit y of
t he st omach makes t he mucus less st icky and frees part icles from it .[16]
The part icles are sort ed by yet anot her group of cilia, which send t he smaller part icles, mainly
minerals, t o t he prost yle so event ually t hey are excret ed, while t he larger ones, mainly food, are sent
t o t he st omach's cecum (a pouch wit h no ot her exit ) t o be digest ed. The sort ing process is by no
means perfect .[16]
Periodically, circular muscles at t he hindgut 's ent rance pinch off and excret e a piece of t he prost yle,
prevent ing t he prost yle from growing t oo large. The anus, in t he part of t he mant le cavit y, is swept
by t he out going "lane" of t he current creat ed by t he gills. Carnivorous molluscs usually have simpler
digest ive syst ems.[16]
As t he head has largely disappeared in bivalves, t he mout h has been equipped wit h labial palps (t wo
on each side of t he mout h) t o collect t he det rit us from it s mucus.[14]
Nervous system
Simplified diagram of the mollusc

nervous system
The cephalic molluscs have t wo pairs of main nerve cords organized around a number of paired
ganglia, t he visceral cords serving t he int ernal organs and t he pedal ones serving t he foot . Most pairs
of corresponding ganglia on bot h sides of t he body are linked by commissures (relat ively large
bundles of nerves). The ganglia above t he gut are t he cerebral, t he pleural, and t he visceral, which
are locat ed above t he esophagus (gullet ). The pedal ganglia, which cont rol t he foot , are below t he
esophagus and t heir commissure and connect ives t o t he cerebral and pleural ganglia surround t he
esophagus in a circumesophageal nerve ring or nerve collar.[16]
The acephalic molluscs (i.e., bivalves) also have t his ring but it is less obvious and less import ant . The
bivalves have only t hree pairs of ganglia— cerebral, pedal, and visceral— wit h t he visceral as t he
largest and most import ant of t he t hree funct ioning as t he principal cent er of "t hinking".[36][37] Some
such as t he scallops have eyes around t he edges of t heir shells which connect t o a pair of looped
nerves and which provide t he abilit y t o dist inguish bet ween light and shadow.
Reproduction
Apical t uft (cilia)
Prot ot roch (cilia)

St omach
Mout h
Met at roch (cilia)
Mesoderm
/// = cilia Anus
Trochophore larva[38]
The simplest molluscan reproduct ive syst em relies on ext ernal fert ilizat ion, but wit h more complex
variat ions. All produce eggs, from which may emerge t rochophore larvae, more complex veliger
larvae, or miniat ure adult s. Two gonads sit next t o t he coelom, a small cavit y t hat surrounds t he
heart , int o which t hey shed ova or sperm. The nephridia ext ract t he gamet es from t he coelom and
emit t hem int o t he mant le cavit y. Molluscs t hat use such a syst em remain of one sex all t heir lives
and rely on ext ernal fert ilizat ion. Some molluscs use int ernal fert ilizat ion and/or are hermaphrodit es,
funct ioning as bot h sexes; bot h of t hese met hods require more complex reproduct ive syst ems.[16]
C. obtusus is an endemic snail species of t he East ern Alps. There is st rong evidence for self-
fert ilizat ion in t he east ernmost snail populat ions of t his species.[39]
The most basic molluscan larva is a t rochophore, which is plankt onic and feeds on float ing food
part icles by using t he t wo bands of cilia around it s "equat or" t o sweep food int o t he mout h, which
uses more cilia t o drive t hem int o t he st omach, which uses furt her cilia t o expel undigest ed remains
t hrough t he anus. New t issue grows in t he bands of mesoderm in t he int erior, so t he apical t uft and
anus are pushed furt her apart as t he animal grows. The t rochophore st age is oft en succeeded by a
veliger st age in which t he prot ot roch, t he "equat orial" band of cilia nearest t he apical t uft , develops
int o t he velum ("veil"), a pair of cilia-bearing lobes wit h which t he larva swims. Event ually, t he larva
sinks t o t he seafloor and met amorphoses int o t he adult form. While met amorphosis is t he usual
st at e in molluscs, t he cephalopods differ in exhibit ing direct development : t he hat chling is a
'miniat urized' form of t he adult .[40] The development of molluscs is of part icular int erest in t he field
of ocean acidificat ion as environment al st ress is recognized t o affect t he set t lement ,
met amorphosis, and survival of larvae.[41]
Ecology
Feeding
Most molluscs are herbivorous, grazing on algae or filt er feeders. For t hose grazing, t wo feeding
st rat egies are predominant . Some feed on microscopic, filament ous algae, oft en using t heir radula as
a 'rake' t o comb up filament s from t he sea floor. Ot hers feed on macroscopic 'plant s' such as kelp,
rasping t he plant surface wit h it s radula. To employ t his st rat egy, t he plant has t o be large enough
for t he mollusc t o 'sit ' on, so smaller macroscopic plant s are not as oft en eat en as t heir larger
count erpart s.[42] Filt er feeders are molluscs t hat feed by st raining suspended mat t er and food
part icles from wat er, t ypically by passing t he wat er over t heir gills. Most bivalves are filt er feeders,
which can be measured t hrough clearance rat es. Research has demonst rat ed t hat environment al
st ress can affect t he feeding of bivalves by alt ering t he energy budget of organisms.[41]
Cephalopods are primarily predat ory, and t he radula t akes a secondary role t o t he jaws and t ent acles
in food acquisit ion. The monoplacophoran Neopilina uses it s radula in t he usual fashion, but it s diet
includes prot ist s such as t he xenophyophore Stannophyllum.[43] Sacoglossan sea-slugs suck t he sap
from algae, using t heir one-row radula t o pierce t he cell walls,[44] whereas dorid nudibranchs and
some Vet igast ropoda feed on sponges[45][46] and ot hers feed on hydroids.[47] (An ext ensive list of
molluscs wit h unusual feeding habit s is available in t he appendix of GRAHAM, A. (1955). "Molluscan
diet s" (ht t p://mollus.oxfordjournals.org/cont ent /31/3-4/144.short ) . Journal of Molluscan Studies.
31 (3–4): 144..)
Classification
Opinions vary about t he number of classes of molluscs; for example, t he t able below shows seven
living classes,[20] and t wo ext inct ones. Alt hough t hey are unlikely t o form a clade, some older works
combine t he Caudofoveat a and Solenogast ers int o one class, t he Aplacophora.[28][17]: 291–292 Two of
t he commonly recognized "classes" are known only from fossils.[18]
Described living
Class Major organisms Dist ribut ion
species[20]
all snails and slugs including abalone, limpets, conch, marine,

Gastropoda [17]: 300 70,000
nudibranchs, sea hares, sea butterflies freshwater, land
Bivalvia [17]: 367 clams, oysters, scallops, geoducks, mussels, rudists† 20,000 marine, freshwater
Polyplacophora rocky tidal zone

[17]: 292–298
chitons 1,000
and seabed
Cephalopoda squid, octopuses, cuttlefish, nautiluses, Spirula,

[17]: 343
900 marine
belemnites†, ammonites†
marine 6–7,000
Scaphopoda
[17]: 403–407
tusk shells 500 metres (20–
22,966 ft)
seabed 200–3,000
Aplacophora
[17]: 291–292
worm-like molluscs 320 metres (660–
9,840 ft)
seabed 1,800–
7,000 metres
Monoplacophora
[17]: 298–300
ancient lineage of molluscs with cap-like shells 31 (5,900–23,000 ft);
one species 200
metres (660 ft)
Rostroconchia†[48] fossils; probable ancestors of bivalves extinct marine
Helcionelloida†[49] fossils; snail-like molluscs such as Latouchella extinct marine
Cricoconarida†[50] extinct
Classificat ion int o higher t axa for t hese groups has been and remains problemat ic. A phylogenet ic
st udy suggest s t he Polyplacophora form a clade wit h a monophylet ic Aplacophora.[51] Addit ionally, it
suggest s a sist er t axon relat ionship exist s bet ween t he Bivalvia and t he Gast ropoda. Tent aculit a
may also be in Mollusca (see Tentaculites).
Evolution
The use of love darts by the land snail

Monachoides vicinus is a form of sexual
selection
Fossil record
The enigmatic Kimberella quadrata

(fossil pictured) from the Ediacaran
has been described as being
"mollusc-like" because of its features
which are shared with modern day
molluscs.
Good evidence exist s for t he appearance of gast ropods (e.g., Aldanella), cephalopods (e.g.,
Plectronoceras, Nectocaris?) and bivalves (Pojetaia, Fordilla) t owards t he middle of t he Cambrian
period, c. 500 (ht t ps://geolt ime.git hub.io/?Ma=500) million years ago, t hough arguably each of
t hese may belong only t o t he st em lineage of t heir respect ive classes.[52] However, t he evolut ionary
hist ory bot h of t he emergence of molluscs from t he ancest ral group Lophot rochozoa, and of t heir
diversificat ion int o t he well-known living and fossil forms, is st ill vigorously debat ed.
Debat e occurs about whet her some Ediacaran and Early Cambrian fossils really are molluscs.[53]
Kimberella, from about 555 (ht t ps://geolt ime.git hub.io/?Ma=555) million years ago, has been
described by some paleont ologist s as "mollusc-like",[54][55] but ot hers are unwilling t o go furt her t han
"probable bilat erian",[56][57] if t hat .[58]
There is an even sharper debat e about whet her Wiwaxia, from about
505 (ht t ps://geolt ime.git hub.io/?Ma=505) million years ago, was a mollusc, and much of t his
cent ers on whet her it s feeding apparat us was a t ype of radula or more similar t o t hat of some
polychaet e worms.[56][59] Nicholas But t erfield, who opposes t he idea t hat Wiwaxia was a mollusc,
has writ t en t hat earlier microfossils from
515 t o 510 (ht t ps://geolt ime.git hub.io/?Ma=515%E2%80%93510) million years ago are fragment s
[60]
of a genuinely mollusc-like radula. This appears t o cont radict t he concept t hat t he ancest ral
[61]
molluscan radula was mineralized.
However, t he Helcionellids, which first appear over

540 (ht t ps://geolt ime.git hub.io/?Ma=540) million years ago in
[62][63]
Early Cambrian rocks from Siberia and China, are t hought
t o be early molluscs wit h rat her snail-like shells. Shelled
molluscs t herefore predat e t he earliest t rilobit es.[49] Alt hough
The tiny Helcionellid
most helcionellid fossils are only a few millimet ers long, Spirally coiled shells
fossil Yochelcionella
specimens a few cent imet ers long have also been found, most appear in many
is thought to be an
gastropods.[17]: 300–343
wit h more limpet -like shapes. The t iny specimens have been early mollusc[49]
suggest ed t o be juveniles and t he larger ones adult s.[64]
Some analyses of helcionellids concluded t hese were t he earliest gast ropods.[65] However, ot her
scient ist s are not convinced t hese Early Cambrian fossils show clear signs of t he t orsion t hat
ident ifies modern gast ropods t wist s t he int ernal organs so t he anus lies above t he
head.[17]: 300–343 [66][67]
= Sept a
= Siphuncl
Septa and siphuncle in nautiloid shell
Volborthella, some fossils of which predat e

530 (ht t ps://geolt ime.git hub.io/?Ma=530) million years ago, was long t hought t o be a cephalopod,
but discoveries of more det ailed fossils showed it s shell was not secret ed, but built from grains of
t he mineral silicon dioxide (silica), and it was not divided int o a series of compart ment s by sept a as
t hose of fossil shelled cephalopods and t he living Nautilus are. Volborthella's classificat ion is
uncert ain.[68] The Middle Cambrian fossil Nect ocaris is oft en int erpret ed as a cephalopod wit h 2
arms and no shell, but t he Lat e Cambrian fossil Plectronoceras is now t hought t o be t he earliest
undisput ed cephalopod fossil, as it s shell had sept a and a siphuncle, a st rand of t issue t hat Nautilus
uses t o remove wat er from compart ment s it has vacat ed as it grows, and which is also visible in
fossil ammonit e shells. However, Plectronoceras and ot her early cephalopods crept along t he
seafloor inst ead of swimming, as t heir shells cont ained a "ballast " of st ony deposit s on what is
t hought t o be t he underside, and had st ripes and blot ches on what is t hought t o be t he upper
surface.[69] All cephalopods wit h ext ernal shells except t he naut iloids became ext inct by t he end of
t he Cret aceous period 65 (ht t ps://geolt ime.git hub.io/?Ma=65) million years ago.[70] However, t he
shell-less Coleoidea (squid, oct opus, cut t lefish) are abundant t oday.[71]
The Early Cambrian fossils Fordilla and Pojetaia are regarded as bivalves.[72][73][74][75] "Modern-
looking" bivalves appeared in t he Ordovician period,
488 t o 443 (ht t ps://geolt ime.git hub.io/?Ma=488%E2%80%93443) million years ago.[76] One bivalve
group, t he rudist s, became major reef-builders in t he Cret aceous, but became ext inct in t he
Cret aceous–Paleogene ext inct ion event .[77] Even so, bivalves remain abundant and diverse.
The Hyolit ha are a class of ext inct animals wit h a shell and operculum t hat may be molluscs. Aut hors
who suggest t hey deserve t heir own phylum do not comment on t he posit ion of t his phylum in t he
t ree of life.[78]
Phylogeny
⁠
Brachiopods
⁠
⁠
Bivalves
⁠
⁠Monoplacophorans
⁠("limpet-like", "living fossils")
⁠
Scaphopods (tusk shells)
⁠⁠ ⁠
⁠ ⁠ ⁠Gastropods
⁠ ⁠(snails, slugs, limpets, sea hares)
⁠Mollusca
⁠⁠
⁠ ⁠ ⁠Cephalopods
(nautiloids, ammonites, octopus,
⁠ squid, etc.)
⁠Lophotrochozoa
⁠ ⁠ ⁠Aplacophorans
⁠ ⁠ ⁠(spicule-covered, worm-like)
⁠⁠
Polyplacophorans (chitons)
⁠
⁠
Wiwaxia
⁠† Halwaxiids ⁠
⁠ ⁠ ⁠
Halkieria
⁠ ⁠
⁠ † Orthrozanclus
⁠
⁠
† Odontogriphus
⁠
A possible "family tree" of molluscs (2007).[79][80] Does not include
annelid worms as the analysis concentrated on fossilizable "hard"
features.[79]
The phylogeny (evolut ionary "family t ree") of molluscs is a cont roversial subject . In addit ion t o t he
debat es about whet her Kimberella and any of t he "halwaxiids" were molluscs or closely relat ed t o
molluscs,[55][56][59][60] debat es arise about t he relat ionships bet ween t he classes of living
molluscs.[57] In fact , some groups t radit ionally classified as molluscs may have t o be redefined as
dist inct but relat ed.[81]
Molluscs are generally regarded members of t he Lophot rochozoa,[79] a group defined by having
t rochophore larvae and, in t he case of living Lophophorat a, a feeding st ruct ure called a lophophore.
The ot her members of t he Lophot rochozoa are t he annelid worms and seven marine phyla.[82] The
diagram on t he right summarizes a phylogeny present ed in 2007 wit hout t he annelid worms.
Because t he relat ionships bet ween t he members of t he family t ree are uncert ain, it is difficult t o
ident ify t he feat ures inherit ed from t he last common ancest or of all molluscs.[83] For example, it is
uncert ain whet her t he ancest ral mollusc was met americ (composed of repeat ing unit s)—if it was,
t hat would suggest an origin from an annelid-like worm.[84] Scient ist s disagree about t his: Giribet and
colleagues concluded, in 2006, t he repet it ion of gills and of t he foot 's ret ract or muscles were lat er
development s,[13] while in 2007, Sigwart concluded t he ancest ral mollusc was met americ, and it had
a foot used for creeping and a "shell" t hat was mineralized.[57] In one part icular branch of t he family
t ree, t he shell of conchiferans is t hought t o have evolved from t he spicules (small spines) of
aplacophorans; but t his is difficult t o reconcile wit h t he embryological origins of spicules.[83]
The molluscan shell appears t o have originat ed from a mucus coat ing, which event ually st iffened
int o a cut icle. This would have been impermeable and t hus forced t he development of more
sophist icat ed respirat ory apparat us in t he form of gills.[49] Event ually, t he cut icle would have
become mineralized,[49] using t he same genet ic machinery (engrailed) as most ot her bilat erian
skelet ons.[84] The first mollusc shell almost cert ainly was reinforced wit h t he mineral aragonit e.[29]
The evolut ionary relat ionships wit hin t he molluscs are also debat ed, and t he diagrams below show
t wo widely support ed reconst ruct ions:
⁠ ⁠
Solenogastres Solenogastres
⁠⁠ ⁠
⁠Aculifera ⁠ ⁠Caudofoveata ⁠
Caudofoveata
⁠ ⁠ ⁠
⁠ ⁠
Polyplacophorans Polyplacophorans
⁠ ⁠
⁠ ⁠
Monoplacophorans Monoplacophorans
⁠ ⁠
⁠Molluscs ⁠ ⁠ olluscs ⁠
M ⁠
⁠ Bivalves ⁠ ⁠ Bivalves
⁠ ⁠
⁠ ⁠
T estaria ⁠
⁠Conchifera Scaphopods ⁠ ⁠ Scaphopods
⁠ ⁠ ⁠
⁠ ⁠⁠
⁠ ⁠Gastropods ⁠ ⁠Gastropods
⁠ ⁠
⁠ ⁠
Cephalopods Cephalopods
⁠ ⁠
The "Aculifera" hypothesis[79] The "Testaria" hypothesis[79]
Morphological analyses t end t o recover a conchiferan clade t hat receives less support from
molecular analyses,[85] alt hough t hese result s also lead t o unexpect ed paraphylies, for inst ance
scat t ering t he bivalves t hroughout all ot her mollusc groups.[86]
However, an analysis in 2009 using bot h morphological and molecular phylogenet ics comparisons
concluded t he molluscs are not monophylet ic; in part icular, Scaphopoda and Bivalvia are bot h
separat e, monophylet ic lineages unrelat ed t o t he remaining molluscan classes; t he t radit ional phylum
Mollusca is polyphylet ic, and it can only be made monophylet ic if scaphopods and bivalves are
excluded.[81] A 2010 analysis recovered t he t radit ional conchiferan and aculiferan groups, and
showed molluscs were monophylet ic, demonst rat ing t hat available dat a for solenogast res was
cont aminat ed.[87] Current molecular dat a are insufficient t o const rain t he molluscan phylogeny, and
since t he met hods used t o det ermine t he confidence in clades are prone t o overest imat ion, it is
risky t o place t oo much emphasis even on t he areas of which different st udies agree.[88] Rat her t han
eliminat ing unlikely relat ionships, t he lat est st udies add new permut at ions of int ernal molluscan
relat ionships, even bringing t he conchiferan hypot hesis int o quest ion.[89]
Human interaction
For millennia, molluscs have been a source of food for humans, as well as import ant luxury goods,
not ably pearls, mot her of pearl, Tyrian purple dye, sea silk, and chemical compounds. Their shells
have also been used as a form of currency in some preindust rial societ ies. A number of species of
molluscs can bit e or st ing humans, and some have become agricult ural pest s.
Uses by humans
Molluscs, especially bivalves such as clams and mussels, have been an import ant food source since
at least t he advent of anat omically modern humans, and t his has oft en result ed in overfishing.[90]
Ot her commonly eat en molluscs include oct opuses and squids, whelks, oyst ers, and scallops.[91] In
2005, China account ed for 80% of t he global mollusc cat ch, net t ing almost 11,000,000 t onnes
(11,000,000 long t ons; 12,000,000 short t ons). Wit hin Europe, France remained t he indust ry leader.[92]
Some count ries regulat e import at ion and handling of molluscs and ot her seafood, mainly t o minimize
t he poison risk from t oxins t hat can somet imes accumulat e in t he animals.[93]
Saltwater pearl oyster farm in Seram,

Indonesia
Most molluscs wit h shells can produce pearls, but only t he pearls of bivalves and some gast ropods,
whose shells are lined wit h nacre, are valuable.[17]: 300–343, 367–403 The best nat ural pearls are
produced by marine pearl oyst ers, Pinctada margaritifera and Pinctada mertensi, which live in t he
t ropical and subt ropical wat ers of t he Pacific Ocean. Nat ural pearls form when a small foreign
object get s st uck bet ween t he mant le and shell.
The t wo met hods of cult uring pearls insert eit her "seeds" or beads int o oyst ers. The "seed" met hod
uses grains of ground shell from freshwat er mussels, and overharvest ing for t his purpose has
endangered several freshwat er mussel species in t he sout heast ern Unit ed St at es.[17]: 367–403 The
pearl indust ry is so import ant in some areas, significant sums of money are spent on monit oring t he
healt h of farmed molluscs.[94]
Byzantine Emperor Justinian I
clad in Tyrian purple and
wearing numerous pearls
Ot her luxury and high-st at us product s were made from molluscs. Tyrian purple, made from t he ink
glands of murex shells, "fet ched it s weight in silver" in t he fourt h cent ury BC, according t o
Theopompus.[95] The discovery of large numbers of Murex shells on Cret e suggest s t he Minoans
may have pioneered t he ext ract ion of "imperial purple" during t he Middle Minoan period in t he 20t h–
18t h cent uries BC, cent uries before t he Tyrians.[96][97] Sea silk is a fine, rare, and valuable fabric
produced from t he long silky t hreads (byssus) secret ed by several bivalve molluscs, part icularly
Pinna nobilis, t o at t ach t hemselves t o t he sea bed.[98] Procopius, writ ing on t he Persian wars circa
550 CE, "st at ed t hat t he five heredit ary sat raps (governors) of Armenia who received t heir insignia
from t he Roman Emperor were given chlamys (or cloaks) made from lana pinna. Apparent ly, only t he
ruling classes were allowed t o wear t hese chlamys."[99]
Mollusc shells, including t hose of cowries, were used as a kind of money (shell money) in several
preindust rial societ ies. However, t hese "currencies" generally differed in import ant ways from t he
st andardized government -backed and -cont rolled money familiar t o indust rial societ ies. Some shell
"currencies" were not used for commercial t ransact ions, but mainly as social st at us displays at
import ant occasions, such as weddings.[100] When used for commercial t ransact ions, t hey funct ioned
as commodit y money, as a t radable commodit y whose value differed from place t o place, oft en as a
result of difficult ies in t ransport , and which was vulnerable t o incurable inflat ion if more efficient
t ransport or "goldrush" behavior appeared.[101]
Bioindicators
Bivalve molluscs are used as bioindicat ors t o monit or t he healt h of aquat ic environment s in bot h
fresh wat er and t he marine environment s. Their populat ion st at us or st ruct ure, physiology, behaviour
or t he level of cont aminat ion wit h element s or compounds can indicat e t he st at e of cont aminat ion
st at us of t he ecosyst em. They are part icularly useful since t hey are sessile so t hat t hey are
represent at ive of t he environment where t hey are sampled or placed.[102] Pot amopyrgus
ant ipodarum is used by some wat er t reat ment plant s t o t est for est rogen-mimicking pollut ant s
from indust rial agricult ure. Several species of mollusca have been used as bioindicat ors of
environment al st resses t hat can cause DNA damage. These species include t he American oyst er
Crassostrea virginica,[103] zebra mussels (Dreissena polymorpha)[104][105] and t he blue mussel Mytilus
edulis.[106]
Harm to humans
Stings and bites
The blue-ringed octopus's

rings are a warning signal;
this octopus is alarmed, and
its bite can kill.[107]
Some molluscs st ing or bit e, but deat hs from mollusc venoms t ot al less t han 10% of t hose from
jellyfish st ings.[108]
All oct opuses are venomous,[109] but only a few species pose a significant t hreat t o humans. Blue-
ringed oct opuses in t he genus Hapalochlaena, which live around Aust ralia and New Guinea, bit e
humans only if severely provoked,[107] but t heir venom kills 25% of human vict ims. Anot her t ropical
species, Octopus apollyon, causes severe inflammat ion t hat can last for over a mont h even if
t reat ed correct ly,[110] and t he bit e of Octopus rubescens can cause necrosis t hat last s longer t han
one mont h if unt reat ed, and headaches and weakness persist ing for up t o a week even if
t reat ed.[111]
Live cone snails can be dangerous to
shell collectors, but are useful to
neurology researchers.[112]
All species of cone snails are venomous and can st ing painfully when handled, alt hough many species
are t oo small t o pose much of a risk t o humans, and only a few fat alit ies have been reliably report ed.
Their venom is a complex mixt ure of t oxins, some fast -act ing and ot hers slower but
deadlier.[112][108][113] The effect s of individual cone-shell t oxins on vict ims' nervous syst ems are so
precise as t o be useful t ools for research in neurology, and t he small size of t heir molecules makes it
easy t o synt hesize t hem.[112][114]
Disease vectors
Skin vesicles created by the

penetration of Schistosoma. (Source:
CDC)
Schist osomiasis (also known as bilharzia, bilharziosis or snail fever), a disease caused by t he fluke
worm Schistosoma, is "second only t o malaria as t he most devast at ing parasit ic disease in t ropical
count ries. An est imat ed 200 million people in 74 count ries are infect ed wit h t he disease – 100
million in Africa alone."[115] The parasit e has 13 known species, t wo of which infect humans. The
parasit e it self is not a mollusc, but all t he species have freshwat er snails as int ermediat e host s.[116]
Pests
Some species of molluscs, part icularly cert ain snails and slugs, can be serious crop pest s,[117] and
when int roduced int o new environment s, can unbalance local ecosyst ems. One such pest , t he giant
African snail Achatina fulica, has been int roduced t o many part s of Asia, as well as t o many islands in
t he Indian Ocean and Pacific Ocean. In t he 1990s, t his species reached t he West Indies. At t empt s
t o cont rol it by int roducing t he predat ory snail Euglandina rosea proved disast rous, as t he predat or
ignored Achatina fulica and went on t o ext irpat e several nat ive snail species inst ead.[118]
See also
Land snail
Sea slug
Sea snail
Explanatory notes
a. The formerly dominant U.K. spelling mollusk is still used in the U.S.—see the reasons given by Gary
Rosenberg (1996).[2] For the spelling mollusc, see the reasons given in: Brusca & Brusca. Invertebrates
(2nd ed.)..
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Further reading
St urm, C.; Pearce, T.A. & Valdes, A. The Mollusks: A Guide to Their Study, Collection, and
Preservation. Universal Publishers. 2006. 454 pages. ISBN 1581129300
Trigo, J.E.; Díaz Agras, G.J.; García-Álvarez, O.L.; Guerra, A.; Moreira, J.; Pérez, J.; Rolán, E.; Troncoso,
J.S. & Urgorri, V. (2018). Troncoso, J.S., Trigo, J.E. & Rolán, E., ed. Guía de los Moluscos Marinos de
Galicia. Vigo: Servicio de Publicacións da Universidade de Vigo. 836 pages. ISBN 978-84-8158-
787-6
External links
"Mollusca" (ht t ps://www.eol.org/pages/2195) at t he Wikimedia Commons has

Encyclopedia of Life media related to
Mollusca.
Researchers complet e mollusk evolut ionary t ree; 26 Oct ober
T he Wikibook
2011 (ht t ps://phys.org/news/2011-10-mollusk-evolut ionary-t r Dichotomous Key has a
ee.ht ml) page on the topic of:
Mollusca
Hardy's Int ernet Guide t o Marine Gast ropods (ht t ps://conchol
Wikisource has the text
ogy.be/?t =261) of the 1911
Encyclopædia Britannica
Rot t erdam Nat ural Hist ory Museum (ht t p://www.nmr-pics.n article "Mollusca".
l/) Shell Image Gallery
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0907012135/ht t p://ccma.nos.noaa.gov/about /coast /nsandt / information related to
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musselwat ch.aspx)
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ux1.fr/index.php?rubrique=accueil&lang=en) Archived (ht t ps://web.archive.org/web/2016111317
3444/ht t p://molluscan-eye.epoc.u-bordeaux1.fr/index.php?rubrique=accueil&lang=en) 2016-11-
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Cornu aspersum (formerly Helix

Clade: Bilat eria

(unranked): Prot ost omia

Superphylum: Lophot rochozoa

Fossilized ammonite displayed at the Phylum: Mollusca

can accumulat e in cert ain molluscs under specific

The anus and genit als open int o t he mant le cavit y.

There are t wo pairs of main nerve cords.[17]

Whet her charact erist ic is found in t hese classes of Molluscs

Broad, muscular Reduced or Modified into S

Large digest ive No ceca in some

Odontophore Yes Yes Yes Yes Yes No Y

Diversity and variability of shells of

About 80% of all known mollusc

Anatomical diagram of a hypothetical ancestral mollusc

A 50-second video of snails (most

Eating, digestion, and excretion

Snail radula at work

Simplified diagram of the mollusc

Apical t uft (cilia)

Prot ot roch (cilia)

all snails and slugs including abalone, limpets, conch, marine,

Polyplacophora rocky tidal zone

Cephalopoda squid, octopuses, cuttlefish, nautiluses, Spirula,

Rostroconchia†[48] fossils; probable ancestors of bivalves extinct marine

Helcionelloida†[49] fossils; snail-like molluscs such as Latouchella extinct marine

The use of love darts by the land snail

The enigmatic Kimberella quadrata

However, t he Helcionellids, which first appear over

suggest ed t o be juveniles and t he larger ones adult s.[64]

Septa and siphuncle in nautiloid shell

Volborthella, some fossils of which predat e

Saltwater pearl oyster farm in Seram,

Stings and bites

The blue-ringed octopus's

Skin vesicles created by the

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"Mollusca" (ht t ps://www.eol.org/pages/2195) at t he Wikimedia Commons has

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