Chapter 11

Lecture Outline

Two groups of coelomates animals have been distinguished: the protostomes and the
deuterostomes.

Protostomes are further divided into the groups lophotrochozoa and ecdysozoa.
Lophotrochozoans do not molt a cuticle whereas ecydosozoans do. Lophotrochozoans include
all of the organisms from the previous chapter, molluscs, annelids, and some lesser phyla; both
the annelids and the molluscs have a​ trochophore​ larval stage in development, suggesting an
evolutionary relationship.

The trochophore larval stage includes translucent, free-swimming larva.

The distinction between protostomes and deuterostomes is driven in large part by a variety of
embryological characteristics. Protostomes have spiral, determinate, cleavage and the
blastopore ​eventually develops into the mouth of the organism.

In deuterostomes, the cleavage is radial and indeterminate and the blastopore develops into the
anus.

A fourth distinction between the two groups is the manner in which the coelom develops.
Molluscs have been very successful in a myriad of habitats. Members of this very diverse group
range in size from a few mm to the 18 meter giant squid.

Bivalves and gastropods are the most

successful of the 8 molluscan classes.

Cephalopods (https://www.britannica.com/list/6-reasons-to-love-cephalopods) have declined in

species numbers. Molluscs are triploblastic and have bilateral symmetry, a trochophore larva,
and basic protostome characteristics, but they also have 3 unique features:
1. A body composed of a head-foot and visceral mass.
2. A mantle that encloses the visceral mass, secretes the shell (if present), and forms the
mantle cavity.
3. A radula, a rasping structure used in ​food collection​ that is supported by a cartilaginous
odontophore, is usually present. The radula has been lost in the bivalves.
 Although molluscs are coelomate, the coelom is reduced to cavities surrounding the heart,
nephridia, and gonads. Molluscs have an open circulatory system, in all classes other than
the cephalopods.

The class Gastropoda is the most speciose molluscan class (over 35,000 described species), but it
is characterized by a unique development process called torsion.​ All gastropods undergo
torison​, an 180° counterclockwise (synestral) twisting of the body that causes the gills and anus
to be located behind the head. This position of the anus allows wastes to fall onto the head and
gills, fouling them. However, some adaptations to prevent fouling of the gills include slits in the
mantle cavity, and detorsion, or untwisting, until the mantle cavity opens on the right side of the
body behind the head. The function of torsion is unknown, but has been suggested to relate to
shell coiling. The foot is flattened. Because of torsion, the snail’s head is drawn into the shell
first, and as the foot retracts, an operculum may cover the aperture, so perhaps protecting the
head is a benefit of torsion.

The basic features of gastropods are outlined below. Most gastropods are herbivores, feeding
with a radula, but some are predatory. Once food is ingested, the ​digestive gland​ in the stomach
aids in breaking down food particles. Gastropods have an open circulatory system and the
hydraulic skeleton (where blood or other fluids under pressure are forced into tissues to extent
them) functions in circulation, as well as movement and support. Gastropods have well
developed senses, including eyes on tentacles, statocysts, and osphradia (chemoreceptors).
Nephridium ​are the excretory structures in gastropods.

Members of the Subclass Pulmonata are mostly freshwater and terrestrial snails.​ The terrestrial
forms use a lung, made from the vascularized mantle cavity, in place of gills. The opening to the
lung is the ​pneumostome​.

Class Bivalvia includes clams, mussels, oysters and scallops, and is the second largest molluscan
class (30,000 described species).​ Bivalves are found in almost all aquatic habitats, buried, or
attached to rock or man-made substrata by byssal threads. A pair of ​hinge ligament​ muscles
keeps the shell closed. The oldest part of the shell is called the umbo. Many species are edible
and some form freshwater and marine pearls. Pearls are formed whenever an irritant (such as a
grain of sand) lodges between the shell and mantle cavity and the mantle secretes nacre to
surround the irritant.

Bivalves are typically sedentary filter feeders. The incurrent ​siphon ​is the conduit for providing
the water current. Filter feeding is accomplished by the lamellae of the gills. Gills are used in
both respiration and feeding. Collected food is directed to the cilia-covered labial palp
surrounding the mouth.

Circulation and respiration involve blood vessels in the heart, tissues, sinuses, and gills. The
circulatory system is open. Two nephridia are located below the pericardial cavity. The nervous
system is primitive, including several ganglia, sense organs, around the margin of the mantle,
and may include complex eyes as seen scallops.

Bivalve mollusks help to remove suspended particles from water and thus provide a clarifying
effect. With the loss of populations of bivalve feeders waters often become much more turbid
and less productive. Because bivalves are filter feeders, they are often utilized (even have their
tissues tested by toxicologists) as indicator species of water quality.

Class Cephalopoda, including octopuses, squid, cuttlefish, and the nautilus, contains the most
morphologically complex invertebrates, particularly with respect to the nervous system.
Ancestral cephalopods were shelled; most extant cephalopods have reduced or lost the shell (the

nautiloids are the exception).

The cephalopods move by contraction of longitudinal and circular muscles in the mantle which
produces a rapid water jet from the funnel (a modified foot). Their rapid locomotion aids in
their predatory habits. This increased activity is supported by the unusual closed circulatory
system allowing more efficient blood flow for excretory and respiratory functions. Carnivorous
cephalopods feed using their jaws and radula; digesting food is moved by peristalsis through the
gastrointestinal tract; wastes pass out the anus and exhalent water flushes it out of the mantle
cavity. Discharge of ink from the mantle cavity may also deter predators.

The cephalopod nervous system is the most advanced of any invertebrate. They have large
brains tied to ​chromatophores​, pigment sacs in the skin that allow cephalopods to change colors
rapidly. The prevailing belief among scientists is that intelligence in cephalopods arose as a
result of their predaceous lifestyle. Not only are they very active predators, but they also
consistently avoid predators. Both of these characteristics may have driven the evolution of their
large brains and complex nervous systems. Color changes are utilized for alarms/camoflauge
and for communication and mating displays. The cephalopod eye is complex and
image-forming; it is and example of convergent evolution with the vertebrate eye. Nautiloid
eyes lack lenses and are open to the seawater.

The class Polyplacophora contains the chitons.​ All chitons have 8 plates on their dorsum, and a
ventral food to adhere to the substrate. They are mainly herbivorous grazers,
feeding on algae is accomplished by a radula. Gills in the mantle cavity provide for respiration.
The nervous system is ladder-like with a nerve ring around the esophagus.
Chitons are dioecious. Early Native Americans utilized chitons as a food resource. A subradular
organ (chemoreceptor) extends from the mouth to detect food sources.

The class Scaphopoda contains the tooth or tusk shells.​ Tusk shells are burrowers and feed upon
foraminiferans, while lying partially buried in the marine sediments.
The tubular or cone-shaped shell is open at both ends. The mantle is elongated and extends the
length of the shell. Various sensory structures are found in many places on the body. These
dioecious animals have both a trochophore and veliger larva in the life cycle.

The class Monoplacophora contains primitive molluscs with an extensive fossil record; living
forms have been known only since 1952.​ ​Neopilina i​ s the only extant genus. They have one
limpet-like shell, in spite of the serially repeated retractor muscles and gills that are also present.
They are dioecious.

The class Aplacophora contains 2 subclasses of primitive shell-less molluscs.​ The​ ​subclass
Neomeniomorpha houses the solenogasters (some with a radula). The subclass
Chaetodermomorpha (previously the Caudofoveata) contains animals with scale-like spicules on
the body surface. Most burrow or creep on the substrate. They have a nervous system similar to
that of flatworms.

Phylogenetic studies of the Mollusca indicate that the group is more than 500 million years old
and did not have a segmented ancestor.​ Many characters, such as segmentation, may be
secondarily derived. Although developmental information on Neopilina is incomplete, no
serially repeating structure in any other mollusc develops as do those of arthropods and annelids.
The evolutionary relationships among the classes are not well understood.