Class Bivalvia (Pelecypoda)

• 2nd largest mollusc class

• ~ 20,000 living; 20,000 fossil spp.
• Appear in Cambrian
• Height of diversity ~ 350 mya
 Class Bivalvia (Pelecypoda)
• Sizes: 2 mm to > 1.35 m (Tridachna
gigas, giant clam can weigh 440 lbs.)
• 1. Bivalve shell
• 2. Bilateral symmetry, body compressed
laterally
• 3. Rudimentary head, no eyes, no
tentacles, no radula
 Characters, cont.
• 4. Large blade-like foot
• 5. Pair of large, complex, ciliated gills
– Filter feed on plankton, sediments
• 6. Some sessile species have byssal
threads for attachment to substrate
– Mussels and oysters
– Secreted by gland at based of foot
 Bivalves
• 7. Most are dioecious
• External fertilization, or in mantle
chamber (some marine and most
freshwater spp.)
• 8. Marine species: trochophore + veliger
larvae present
– Freshwater Unionacea veliger = glochidia
• Parasitic on fish gills
• Fishes are host to glochidia
 Subclass Protobranchia
• Primitive, small, marine bivalves
• Nut clams Nucula yoldia
• Cryptodont Solemya

0.45 cm 1.5 cm
 Subclass Lamellibranchia
98% of bivalves
• A. arks, Arca
– Hinge straight w/ numerous fine, regular
teeth
– Bitter sweet clams, Glycemeris
 Subclass Lamellibranchia
• B. Mussels, winged or tree oysters, pen
shells
– Byssal threads for attachment
– Mytilus, Modiolus are edible mussels
– Pearl oysters Pinctada are winged oysters
 Subclass Lamellibranchia
• C. Scallops Pectin, jingle shells, file
shells
– Scallops swim by clapping their valves
– Numerous blue eyes on mantle margin
 Subclass Lamellibranchia
• D. Oysters Osstrea, Crassostrea (Asian)
– Form large attached colonies
– Oyster farming began over 100 years ago in
U.S.
 Subclass Lamellibranchia
• E. Unionacea, freshwater clams
• Small commercial fishery in Indiana and
elsewhere in U.S. produced “mother of
pearl”
• Currently for Japanese pearl industry
Amblema = three-ridge
Actinonaias = mucket
Potamilus = Bleufer
Ptychobranchus =
kidneyshell
 Subclass Lamellibranchia
• F. Lucines, jewel boxes, and cockles
• Cockles are edible and popular in
Europe, Cardium
 Subclass Lamellibranchia
• G. Giant clams, Tridachna are tropical
sessile species
• Rely on commensal algae for much of
nutrition
• Most species are endangered in many
areas by shell collectors
 Subclass Lamellibranchia
• H. Sphaeridae: freshwater fingernail
clams
• Corbicula, Asian clams
• Common, most < 1/4 inch
 Subclass Lamellibranchia
• I. Razor clams - elongate valves

• J. venus clams
– Mercenaria is the tasty cherrystone or
hard-shelled clam
 Subclass Lamellibranchia
• K. Soft-shell clams (in comparison with
Mercenaria)
– Mya is commercial clam in U.S.
– Geoducks
– Panopea
Corbicula fluminea, Asian clam
• Corbiculidae
• Hermaphroditic
• Introduced to North America early
1900’s
• Widespread E., S., and far W. United
States
 Corbicula
• Life history adapted for unstable,
unpredictable habitats
• Highly invasive, replaces native
Sphaeriid populations
– Highest filtration and assimilation rates of
any freshwater bivalve
– Highest growth and production rate
– 3-6 mos. to maturity
– Single adult can produce 68,678 juv/year
Dreissena polymorpha, zebra
mussel
• Widespread colonial, sessile bivalve
from Europe
– Caspian Sea
• Spread through European drainages in
1700’s
 Zebra mussels
• 1985 - Lake St. Claire by ship water
ballast
• 1990 - entire Great Lakes and St.
Lawrence
• Continues to spread through rivers
 Zebra mussels
• Tend to foul pipes - industry, boats, etc.
• Dioecious, trochophore and veliger larvae
• Adults attach by byssal threads
• Very small eggs,
– 30,000 - 40,000 per female
– Kill natives by overcolonization
 Boring clams (Pholas) and
shipworms (Teredo)
• Shipworms cause economic damage by
boring in pilings and submerged
wooden structures
 Subclass Septibranchia
• Watering pot shells
• Elongate tube-shaped with tiny shells
• Live in mud or bore in rock, coral, or
clamshells
Class Cephalopoda - octopus
and squids
• Extremely complex, advanced molluscs
• Nervous and sensory system
development surpasses all other invert’s
• ~ 1000 extant spp.
• ~ 10,500 fossil spp.
• Appear in Cambrian
– Peaks in abundance Paleozoic + Mesozoic
 Cambrian = 500 mya
present
250 mya

65 mya

65 mya

550 mya
250 mya
Class Cephalopoda - octopus
and squids
• Largest invertebrate animal is giant
squid of North Atlantic (Architeuthes)
– to 16 m with long arms
• All marine + carnivores
 Characteristics:
• 1. Primitively - straight or coiled shell
divided into compartments by septa
– Shell used for buoyancy
– Many cephalopod shells are reduced or
absent
– Compartments connected by siphuncle
• Secretes or absorbs nitrogen-rich gas into
chambers
– New chambers secreted w/growth
Characters
• 2. Bilateral symmetry

• 3. Well-developed head w/prominent, well-

developed eyes
• Eyes resemble vertebrate eyes

• 4. Prehensile tentacles - derived from

anterior of foot - suckers (except Nautilus)
 Characters

• 5. Dorsal-ventral axis of body has

become functional posterior-anterior
axis, by elongation
• 6. Mantle encloses body
– Thick and muscular
– Opening to cavity is funnel-shaped siphon
– Pump water out for backwards “jet-
propulsion”
 Characters
• 7. Mouth equipped with pair of chitinous
jaws - resemble hawk’s beak
– Radula present
• 8. Respiration- pair of gills in mantle
cavity.
– No cilia
• 9. Ability to change skin color by
melanophores in most spp.
 Characters
• 10. Brain large + complex.
– Behavior and learning highly developed

• 11.Dioecious: copulation by transfer of

spermatophore by one of tentacles

• 12. Egg development direct - no larva

 Classification
• 1. Subclass Nautiloidea
– Nautiloids, shelled

– Living Nautilus with 3? spp.

• Indo-Pacific
– Many fossil species
– Don’t confuse with “paper nautilus” - an
octopus
 2. Subclass Ammonidea
Ammonoids
• Shell w/ complex sutures
• Silurian - Cretaceous
 3. Subclass Coleoidea - shell
internal or absent
• A. Belemnites - extinct, w/chambered
internal shell. Probably ancestral to
other coleoids

• B. Spirula - coiled chambered internal

shell.
– Deep-water denizen of tropics
 Subclass Coleoidea
• C. Cuttlefish
– straight, chambered, internal shell
– Sepia
– Rossia (bob-tailed squid)
 D. Squids
• Shell chitinous, no chambers, serves as
body skeleton
• Many pelagic and deep-sea spp.; few in
shallow waters
• Loligo
• Lolliguncula
• Architeuthes
 E. Vampire squid
• Octopus-like forms with webbed arms
• Vampyroteuthis infernalis
• Black skin
 F. Octopods
• Eight arms, no fins
• Octopus
• Argonauta (produces shell-like egg-
case, has dwarf males)
 Molluscan phylogeny
• Classic view - group with annelids and
arthropods:
– 1. Similar early embryology (spiral
cleavage, protostomes, schizocoely)
– 2. Trochophore larva - however, in
molluscs mesodermal bands leading to
body segments not formed
 Classic view of molluscan
phylogeny
• 3. Segmentation - certain molluscs
(Neopalina) with repeating structures
show remnants of segmentation
– Rejected by most
• 4.Coelomate bodies - in molluscs
– Consists of pericardial cavity and sinuses
of hemocoel
– Probably not homologous with annelid
coelom
 Classic view of molluscan
phylogeny
• Stacek (1972) - molluscs are
acoelomate - technicality
• Then molluscs get classified with
platyhelminthes and nemerteans
• 5. Chitinous structures (19-21 phyla
have chitin)
 It appears that:
• Molluscs have evolved from ancestors
of flatworms by acquisition of:
– 1. Complete gut
– 2. A shell
– 3. “body cavity” (elements of circulatory
system spaces)

– See online Mollusca Phylogeny paper