CH8: ACOELOMORPHA,

PLATYZOA, AND MESOZOA

FLATWORMS, GASTROTRICHS, GNATHIFERANS, AND MESOZOANS
A. GETTING AHEAD
1. Animals that actively seek for food, shelter, home sites, and reproductive mates,
require a different set of strategies and a new body organization.
2. Two major evolutionary advances
Cephalization
Sense organs and centers for nervous system are located at the anterior end
(head region)
Primary bilateral symmetry
Body can be divided along only 1 plane of symmetry to yield 2 mirror images
of each other
Active, directed movement most efficient with an elongated body form with
anterior (head), posterior (rear), dorsal, and ventral sides.

B. GENERAL CHARACTERISTICS OF WORMS

1. All but one phylum contains bilaterally symmetrical, triploblastic animals.
2. Members of two phyla have acoelomate bodies. Typical acoelomates have only one
internal space, the digestive cavity. The digestive cavity lining is filled with a cellular,
mesodermally derived parenchyma.
o Members of acoelomate phyla, Acoelomporpha, Gastrotricha, Gnathostomulida
and Platyhelminthes, present more internal complexity
3. Phyla Rotifera, Acanthocephala present a pseudocoelomate body.
4. Worms are highly specialized parasites.

C. PHYLUM ACOELOMORPHA
1. Contains worms that previously belong to two orders within classes Turbellaria in
phylum Platyhelminthes.
2. These worms are less than 5mm long and have a cellular ciliated epidermis.
3. Some have a saclike gut without an anus, and others entirely lack a digestive cavity.
4. Acoelomorphs have a radial arrangement of nerves in the body, instead of the ladderlike pattern.
5. Most acoelomorphs are free-living, but some are symbiotic and others parasitic.
6. Typically live in marine sediments; few are pelagic. Some species live in brackish
water
7. Group contains ~350 species

D. CLADE PLATYZOA

E. PHYLUM PLATYHELMINTHES
1. Commonly known as flatworms, their typically flattened bodies may be slender,
broadly leaflike, or long and ribbonlike.
2. Theres 4 classes within Platyhelminthes:
o Nonparasitic forms: Class Turbellaria.
o Parasitic forms: classes Trematoda, Monogenea and Cestoda.

ECOLOGICAL RELATIONSHIPS
1. Free living flatworms occur only in Turbellaria. Most are adapted as bottom-dwellers in
marine or fresh water or live in moist places on land.
2. Most Monogenea are ectoparasites, but all trematodes and cestodes are
endoparasites.

FORM AND FUNCTION

TEGUMENT, MUSCLES

Most turbellatians have a ciliated cellular epidermis resting on a basement

membrane.
The epidermis typically contains rod-shapped rhabdites, composed of fused vesicles
from the Golgi apparatus, that swell to form a protective mucous sheath around the
body when discharged with water; single-celled mucous glands may also be present.

Turbellarians also possess a locomotory system that allows them quickly to attach
and detach from surfaces. In those cases, the epidermis contains dual-gland
adhesive organs consisting of three cell types: viscid and releasing gland cells and
anchor cells.
All members of the parasitic classes possess a nonciliated body covering called a
syncytial tegument (syncytial = many nuclei).
Adults of all members of Tretamoda, Monogenea and Cestoda share a syncytial
covering that entirely lacks cilia and is designated tegument.

NUTRITION AND DIGESTION

The digestive system of most Platyhelminthes includes a mouth, pharynx, and

intestine.
In turbellarians the muscular pharynx opens posteriorly just inside the mouth,
through which it can extend in planarians.
The intestine may be simple or branched.
In Tretamoda and Monogea organisms have an anteriorly opening pharynx that is not
extendable.
Tapeworms (cestodes) lack a digestive system and must absorb all of their nutrients
as small molecules through their tegument.

EXCRETION AND OSMOREGULATION

The osmoregulatory system consists of canals with tubules that end in flame cells
(protonephridia).
In some turbellarians and in other classes of flatworms, the protonephridia form a
weir.
The beat of the flagella provides a negative pressure to draw fluid through the weir
into the space (lumen) enclosed by the tubule cell.
The wall of the duct beyond the flame cells commonly bears folds or microvilli that
probably function in reabsorption of certain ions or molecules.
This osmoregulatory system is absent or reduced in marine turbellarians, which do
not have to expel excess water.
Metabolic wastes are largely removed by diffusion through the body wall.

NERVOUS SYSTEM AND SENSE ORGANS

The simplest system found in some turbellarians, is a subepidermal nerve plexus

resembling the nerve net of cnidarias
Other flatworms have, in addition to a nerve plexus, one to five pairs of longitudinal
nerve cords lying under a muscle layer. Ladder-type pattern.
The neurons are organized into senseory, motor, and association types.
Tactile cells and chemoreceptive cells are abundant over the body, and in planarians
they form definitive organs on the auricles (the earlike lobes on the sides of the
head).
Some also have statocysts for equilibrium and rheoreceptors for sensing water
current direction.
Ocelli or light sensitive eyespots are common in turbellarians, monogeneans and
larval trematodes.

REPRODUCTION

Flatworms reproduce both sexually and asexually.

Turbellarians can reproduce asexually by fission, merely constricting behind the

pharynx and separating into two animals, each of which regenerates the missing
parts.
In forms such as Stenostomum and Microstomum individuals do not separate at once
but remain attached, forming chains of zooids.
Flukes reproduce asexually in their snail intermediate host.
Most flatworms are monoecious and practice cross-fertilization.
In some turbellarians, yolk for nutrition of the developing embryo is contained within
the egg cell itself (endolecithal). The possession of endolecithal egg is considered
ancestral.
Other turbellarians, trematodes, monogeneans and cestodes share a derived
condition in which female gametes contain little or no yolk, and yolk is contributed by
cells release from separate organs (yolk glands). Usually yolk cells surround the
zygote within an eggshell (ectolecithal)

CLASS TURBELLARIA
1. Free living worms
2. Orders within class Turbellaria are distinguished by the form of the gut and the
pharynx.

ORDER MACROSTOMIDA

Small basal free-living flatworms

A simple gut and a simple pharynx
Endolecithal eggs

ORDER POLYCLADIA

Highly diverse clade of free-living marine turbellarians

A folded pharynx and a gut with many branches
Have endolecithal eggs

ORDER TRICADIA

3.
4.

They are also known as planarians

A three-branched intestine
Have ectolecithal eggs
Turbellarians are traditionally recognized as a paraphyletic group.
Several synapomorphies, such as insunk epidermis and ectolecithal development,
show that some turbellarians are phylogenetically closer to Trematoda, Monogenea
and Cestoda than they are to other turbellarians.

CLASS HOMOSCLEROMORPHA
1. Trematodes are all parasitic flukes, and as adults they are almost all endoparasites of
vertebrates.
2. Similar to Turbellaria except for the tegument.
3. Adaptations for parasitism:
a. Various penetration glands or glands that produce cyst material
b. Organs for attachment, such as suckers and hooks.
c. Increased reproductive capacity.
4. Share several characteristics with turbellarians:
a. Well-developed gut-tube
b. Similar reproductive, excretory and nervous systems
c. Musculature and parenchyma that differ only slightly from those of
turbellarians.

d. Poorly developed sense organs

5. Of the three subclasses of Trematoda, two are small and poorly known groups, but
Digenea is a large group that includes many species of medical and economic
importance.

SUBCLASS DIGENEA

Digenetic trematodes have a complex life cycle, the first host (intermediate host)
being a mollusk and the final host (definite host) being a vertebrate. The definite
host is the one in which the parasite reproduces sexually.
In some species, a second and sometimes third, intermediate host intervenes.
Although the cycle varies among different species but the typical stages include
adult, egg (shelled embryo), miracidium, sporocyst, redia, cercaria and metacercaria.
Life cycle of Clonorchis sinesis (liver fluke in humans):
o The shelled embryo or larva usually passes from definite host in excreta and
must reach water to develop further.
o There it hatches to a free swimming ciliated larva, the miracidium.
o The miracidium penetrates the the tissue of the snail, where it transforms into
a saclike sporocyst.
o Sporocysts reproduce asexually to yield either more sporocysts or a number or
rediae.
o Rediae in turn, reproduce asexually to produce more rediae or to produce
cercariae.
o Cercariae emerge from the snail and penetrate the seond intermediate host or
encyst on vegetation or other objects to become metacercariae, which are
juvenile flukes.
o Adults grow from metacercariae when that stage is eaten by the definite host.
Schistosomiasis, infection with blood flukes of genus Schistosoma, is one of the major
infectious diseases in the world. Cercariae that contact human skin penetrate the skin
to enter blood vessels, which they follow to certain favorite regions depending on the
type of fluke.
o Schistosoma mansoni: lives in venules draining large intestine.
o S. japonicum: localizes in venules draining the small intestine.
o S. haematobium: lives in venules draining the urinary bladder.
Blood flukes are different in having separate females and males.
First symptoms are a rash or itch
Within two month; chills, cough, diarrhea, fatigue, fever and muscle aches occur
Cause severe dysentery, anemia, liver enlargement, bladder inflammation, and brain
damage.

CLASS MONOGENEA
1. Sister taxon to Cestoda.
2. Mostly external parasites that clamp onto the gills and external surfaces of fish using
hooked attachment organ called an opisthaptor.
3. The life cycles of monogeneans are simple, having single host as suggested by the
name of the group, which means single descent. The egg hatches a ciliated larva
that attaches to a host, sometimes following a free-swimming phase.

CLASS CESTODA
1. Commonly known as tapeworms, differ from preceding classes in many respects:
a. The usually have long, flat bodies composed of a scolex, for attachment to
the host, followed by many reproductive units or proglottids.

2. Tapeworms entirely lack a digestive system, but they have well-developed muscles,
and their excretory and nervous systems are somewhat similar to those of other
flatworms.
3. They have no special sense organs, but sensory endings in their tegument are
modified cilia.
4. Tapeworms are nearly all monoecious. The main body of a cestode is a chain of
proglottids called strobilla. Typically, new proglottids form at a germinative zones
just behind the scolex.
5. A proglottid is usually fertilized by another proglottid in the same or different strobili.
6. Shelled embryos form in the uterus of the proglottid, and either they are expelled
through a uterine pore, or the entire proglottid detaches from the worm as it reaches
the distal end.
7. Beef tapeworm, Taenia saginata, shelled larva shed from the human host are
ingested by cattle.
8. Pork tapeworm, Taenia solium, uses humans as definite hosts and pigs as
intermediate hosts.

CH9: POLYZOA AND

KRYPTROCHOZOA

1. A lophophore is a crown of tentacles covered with cilia that is borne on a ridge or fold
of the body wall. It is an efficient feeding device.

A. PHYLUM ECTOPROCTA
1. Have long been called bryozoans, or moss animals.
2. All are aquatic, both freshwater and marine, but they largely occur in shallow waters.
3. Each member of a colony occupies a tiny chamber called a zoecium, which is
secreted by its epidermis. Each individual zooid, comprises a feeding polypide and a
case-forming cystid. Polypides include a lophophore, digestive tract, muscles, and
nerve centers.
4. A polypide lives a type of jack-in-the-box existence, popping up to feed and then
quickly withdrawing into its little chamber, which often has a tiny trapdoor
(operculum) that shuts to conceal its inhabitant.
5. Lophophore ridges tend to be circular in marine ectoprocts and U-shaped in
freshwater species.
6. Modified non-feeding zooid resembles a bird beak, and prevents other organisms
from settling on the colony.
7. Ovicell is special zoecium for embryo development

CH10: MOLLUSCS

A. SHELL COLLECTORS DELIGHT

1. The group includes wormlike animals, and giant squids, as well as animals with a
single shell, two shells, eight shell plates, or no shell at all.
2. Most molluscs have an unusual ribbon of teeth called radula.
3. Most have a large muscular foot used in locomotion.
4. Most have a unique tissue layer called mantle.

B. ECONOMIC IMPORTANCE
1.
2.
3.
4.
5.

Extremely important food source for people around the world

Culturing of pearls and natural pearls is an industry
Burrowing shipworms destroy wooden ships and wharfs
Snails and slugs often damage gardens and other vegetation
Some snails are intermediate hosts for parasites

C. FUNCTION
BODY PLAN
1. Reduced to simple dimensions, a Nanglia body consists of a head-foot portion and a
visceral mass portion.
2. The head-foot region contains feeding, cephalic sensory, and locomotor organs. It
depends primarily on muscular action for its function.
3. The visceral mass contains digestive, circulatory, respiratory and reproductive
organs, and it depends primarily on ciliary tracts for its functioning.
4. Two folds of skin, outgrowths of the dorsal body wall, form a protective mantle, which
encloses a space between itself and the body wall called a mantle cavity.
5. The mantle cavity houses gills or a lung, and in many molluscs the mantle secretes a
protective shell over the head-foot and visceral mass.

HEAD-FOOT
RADULA

Rasping, protrusible, tonguelike organ found in molluscs except bivalves and some
gastropods and solenogasters.
Protactor and retractor muscles move the radula and its supporting cartilages
(odontophore) outside and back into the mouth for feeding.

FOOT

The molluscan foot is variously adapted for locomotion, for attachment to a substrate,
or for a combination of functions.
There are many modifications, such as the attachment disc of limpets, the laterally
compressed hatchet foot of bivalves, or the funnel for jet propulsion in squids and
octopuses.

VISCERAL MASS
MANTLE AND MANTLE CAVITY

The mantle is a sheath of skin, extending dorsally from the visceral mass, that wraps
around each side of the body, protecting the soft parts and creating the mantle cavity
between itself and the visceral mass.
The outer surface of the mantle secretes the shell.
The mantle cavity houses respiratory organs (gills or a lung), which develop from the
mantle.
Products from the digestive, excretory, and reproductive systems empty into the
mantle cavity.
In cephalopods (squids and octopuses), the muscular mantle and its cavity create a
jet propulsion used in locomotion.

SHELL

A shell typically has three layers:

The periostracum, the outer organic layer, is composed of a resistant protein
called conchiolin. It is secreted by a fold of the mantle edge, and growth
occurs only at the margin of the shell.
The middle prismatic layer composed of densely packed prisms of calcium
carbonate laid down in a protein matrix. It is secreted by the glandular

margin of the mantle, and increase in shell size occurs at the shell margin as
the animal grows.
The inner nacre, or nacreous layer, of the shell is composed of calcium
carbonate sheets laid down over a thin protein matrix. This layer is secreted
continuously by the mantle surface, so that it becomes thicker during the
animals life.

INTERNAL STRUCTURE AND FUNCTION

RESPIRATORY SYSTEM

Gas exchange occurs through the body surface, particularly the mantle, and in
specialized respiratory organs such as gills or lungs.
Most molluscs have an open circulatory system with a pumping heart, blood vessels,
and blood sinuses.
An open circulatory system is less efficient at supplying oxygen to all tissues in the
body, so it is common slow-moving animals.
In a closed circulatory system, blood moves to and from tissues within blood vessels.
Most members of class Cephalopoda have a closed circulatory system with a heart,
vessels, and capillaries.
Closed systems have two kinds of hearts:
o Ventricle or systemic heart delivers the newly oxygenated blood coming
from the gill to the rest of the body.
o Branchial or gill hearts deliver blood to the oxygen gathering organ, the
gill.

DIGESTIVE SYSTEM

Nephridium (plural nephridia), invertebrate organ which occurs in pairs. It has a

similar function to the vertebrate kidney. There are two basic types: metanephridia
and protonephridia
Most molluscs have a pair of kidneys (metanephridia), a type of nephridium in
which the inner end opens into the coelom; ducts of the kidneys in many forms serve
also for discharge of eggs and sperm.

NERVOUS SYSTEM

Consists of several pairs of ganglia with connecting nerve cords. Cerebral ganglia,
bucaal Nanglia, pedal ganglia, visceral ganglion, etc.

SENSE ORGANS

Sense organs vary and may be highly specialized

Olfactory organs, eyes, statocysts and mechanoreceptors

REPRODUCTIVE SYSTEM

Most molluscs are dioecious, although some gastropods are hermaphroditics

Aquatic molluscs pass through free swimming trochophore and veliger larval
stages.
A veliger is the free-swimming larva of most marine snails, tusk, shells, and bivalves.
It develops from a trochophore and has the beginning of a foot, shell, and mantle.

D.
CLADE
ACUIFERA:
SOLENOGASTRES
CLASS CAUDOFOVEATA

CLASSES

CAUDOFOVEATA

AND

Wormlike marine organisms

Burrowing and deep sea molluscs
Have no shell but body is covered with calcareous scales
Feed primarily on microorganisms and detritus

E. CLASS POLYPLACOPHORA: CHITONS

1. Commonly called chitons.
2. Polyplacophora means bearing many plates. Usually 8, rarely 7.
3. They commonly occur on rocky surfaces in intertidal regions, although some live at
great depths.
4. The mantle forms a girdle around the margin of the plates. On each side of the broad
ventral foot and lying between the foot and the mantle is a row of gills suspended
from the roof of the mantle cavity. Water enters the grooves anteriorly, flows across
the gills, and leaves posteriorly, thus bringing a continuous supply of oxygen to the
gills.
5. Two kidneys carry waste from the pericardial cavity to the exterior.
6. Sexes are separate in most chitons. Eggs are released into the water.

F. CLADE CONCHIFERA: CLASS GASTROPODA

1. Largest and most diverse class, containing over 70,000 living species.
2. They include snails, limpets, slugs, whelks, conchs, periwinkles, sea slugs, sea hares,
sea butterflies, and others.
3. Range from marine to terrestrial habitats.
4. When present, the shell is almost always one piece (univalve) and may be coiled or
uncoiled.
5. Develop through a trochophore and a veliger larval stage.

FORM AND FUNCTION

TORSION

Initially the mouth is anterior and the anus posterior, but through the process of
torsion, the relative positions of body parts change.

ONTOGENETIC TORSION

1st movement
o Shell rotate it between 90 and 180 into a permanent position
o Viscera rotates 90 counterclockwise
o Anus and mantle cavity usually move further to the right and mantle
cavity is remodeled to encompass the anus.
o Mantle cavity develops on the right side of the body near the anus.
o Anus and mantle rotate to the right side of the body.
After torsion
o Digestive tract moves both laterally and dorsally
o Anus and mantle cavity open above mouth and head
o Certain viscera on the left are now on the right side and vice versa
o Nerve cords form a figure eight
Position of anus creates the possibility of wastes washing back over the gills
(fouling).

ADAPTATIONS TO AVOID FOULING

Loss of the right gill: Wastes are expelled to the right

Loss of the right gill: Wastes are expelled to the right
Dorsal slot or hole in the shell: Vent excurrent water
Loss of the right gill: Wastes are expelled to the right
Dorsal slot or hole in the shell: Vent excurrent water
Detorsion: Occurs at post-veliger stage. Anus lies posterior on the right

INTERNAL FORM AND FUNCTION

Respiration in most gastropods is performed by a gill (or two), although some aquatic
forms lack gills and depend on the skin.
Pulmonates (most freshwater and terrestrial snails) have lost their gill altogether,
and the vascularized mantle wall has evolved to function as a lung.
The anus and nephridiopore open near the opening of the lung to the outside, which
is called pneumostome and waste is expelled forcibly with air or water from the
lung.
Most gastropods have a single nephridium (kidney)
There are both hermaphrodites and dioecious gastropods. No self-fertilization,

G. CLASS BIVALVIA (PELECYPODA)

1. Bivalved (two-shelled) molluscs that include mussels, clams, scallops, oysters, and
shipworms.
2. Most bivalves are sedentary suspension feeders that depend on ciliary currents
produced by the gills to collect their food.
3. They have no head, no radula and very little cephalization.

FORM AND FUNCTION

SHELL

Bivalves are laterally compressed, and their two shells (valves) are held together
dorsally by a hinge ligament the causes the valves to gape ventrally.
Umbo is the oldest part of the shell
Valves are drawn together by contraction of strong adductor muscles
Valves are open when adductor muscles are relaxed

BODY AND MANTLE

The visceral mass is suspended from the dorsal midline, and the muscular food
attaches to the visceral mass anteroventrally.

The posterior edges of the mantle folds form dorsal excurrent and ventral incurrent
openings.
Cilia on the gills and inner surface of the mantle direct the flow of water over the gills,
bringing in food and oxygen.

FEEDING AND DIGESTION

Most bivalves are suspension feeders.

Cilia on the gills and inner surface of the mantle direct the flow of water over the gills
Gland cells on gills and labial palps secrete mucus to entangle particles
Food in mucous masses slides to food grooves at lower edge of gills
Ciliary tracts move the mucous mass into mouth
In stomach, extracellular and intracellular digestions occur

REPRODUCTION

Sexes are usually separate and fertilization is usually external.

Marine embryos go through three free-swimming larval stage trochophore, veliger
larva, and young spatbefore becoming an adult.
In freshwater clams, fertilization is internal and some gill tubes become temporary
brood chambers. There, larva develops into specialized veligers called glochidia,
which are discharged with the excurrent flow.
If glochidia come in contact with a passing fish, they hitchhike a ride as parasite in
the fishs gills for the next days before dropping to the bottom to become sedentary
adults.

H. CLASS CEPHALOPODA
1. Cephalopoda include squids, octopuses, nautiluses and cuttlefishes. All are marine
and all are active predators.
2. The foot forms a funnel for expelling water from the mantle cavity.

FORM AND FUNCTION

SHELL

Fossil records of cephalopods extend to the Cambrian. The earliest shells were
straight cones.
Nautiloid and ammonoid shells had gas chambers allowing them to swim
Cuttlefishes also have a smail, coiled or curved shell, but it is entirely enclosed by
mantle.
In squids, most of the shell has disappeared, leaving only a thin, flexible strip called
pen, which the mantle encloses.

BODY AND MANTLE

Camera eye: consists of an iris, a circular lens, vitreous cavity (eye gel), pigment
cells, and photoreceptor cells.
Cephalopoda
o Position of photoreceptor cells
o Focused through movement of lens rather than changing shape as the lens
o Lack a blind spot

I. PHYLOGENY
1. Hypothetical Ancestral Mollusc
Probably lacked a shell or crawling foot
Probably small (about 1 mm)

2.
3.
4.
5.

Likely was a worm-like organism with a ventral gliding surface

Probably had a dorsal mantle, a chitinous cuticle and calcareous scales
Gastropoda and Cephalopoda appear to form a sister group with well-developed head
Bivalvia and Scaphopoda appear to form a sister group with reduced head
Polyplacophora then branched off before the veliger was established as a larva
Caudofoveata and Solenogasters both probably branched off before development of a
solid shell

CH11: ANNELIDS AND ALLIED

TAXA

1. Members of phylum Annelida are segmented worms including marine bristle worms,
leeches, and earthworms
2. Members of phylum Sipuncula are benthic marine animals
with unsegmented
bodies
3. Annelida and Spipuncula: (1)Eucoelomates; mesoderm forms from derivatives of 4d
cell, (2) Coeloms are formed by schizocoely, (3). Develop by spiral mosaic cleavage

A. PHYLUM ANNELIDA
1. Metameric body
2. Biramous parapodia with internal setae
3. Paired epidermal setae

ECOLOGICAL RELATIONSHIP

1. About 15,000 species

2. Annelids occur in the sea, freshwater, and on land
a. 2/3 are more obscure marine worms
b. Marine annelids borrow in the mud or live in tubes
c. Freshwater annelids burrow in the mud, live among vegetation, or swim freely
3. Some feed on organic matter in the mud, some are suspension feeders, many are
predators
a. Leeches are bloodsuckers or are carnivores
4. Annelids are important for drainage, aeration, mixing of soil and distribution of
organic matter

ECONOMIC IMPORTANCE
Play a significant part in marine food chains because fish, crustaceans, hydroids, and
many other predators consume them

BODY PLAN
1. Metamerism (or segmentation) in Annelida
Bodies composed of serially repeated units
Each unit contains components of most organ systems
Metameric body plan occurs in two other animal phyla: Arthropoda and Chordata
Metamerism may have arisen independently in the deuterostome line
2. Evolution of metamerism allowed much greater complexity in structure and function:
Increased burrowing efficiency by permitting
independent movement of
segments
Evolution of a more sophisticated nervous system
Provided a safety factor
If one segment failed, others could still function
3. Body is composed of two-part head, segments, and terminal portion.
4. Head is composed of Prostomium and Peristomium
5. Terminal portion bearing anus is the pygidium. New metameres form just in front of
pygidium.
6. Coelom forms a pair of coelomic compartments in each segment. Gut is not
segmented and longitudinal blood vessels extend through septa
7. Peritoneum
Mesodermal epithelium
Lines the body wall and forms dorsal and ventral mesenteries
Covers all organs
Peritonea of adjacent segments meet to form the septa
8. Coelom is filled with fluid and serves as a hydrostatic skeleton
Fluid volume remains constant
Contraction of longitudinal muscles causes the body to shorten and fatten
Contraction of circular muscles causes the body to narrow and lengthen
By separating this force into septa, peristalsis (widening and elongation) move
worms
9. The outermost layer of body wall is the collagenous cuticle.
10.Setae
Tiny chitinous bristles
Arranged in bunldes in Polychaetes; most four pairs in earthworms
Absent in leeches
Short setae anchor segments in earthworms
Prevent earthworms from slipping backward
Prevent worms from being pulled or washed out of their homes
Long setae help aquatic worms swim

11. Polychaetes have parapodial capillaries and respiration mostly occurs through skin.
Respiration also occur through gills in polychaetes and freshwater oligochaetes.
12. Each body segment has a pair of nephridia. Nephridium occupies parts of two
adjacent segments. Nephrostome draws in wastes and leads through the septum.
Wastes are passed through loops of the nephridium and collected in bladder. Wastes
are expelled through the nephridiopore of next segment

PHYLOGENY

Originally divided into three classes: Polychaeta, Oligochaeta, Hirudinea

Polychaeta and Oligochaeta are paraphyletic groups
Hirudinea is monophyletic

POLYCHAETES

Typically, those with parapodia, and many setae

More than 10,000 species
Most marine; free living or sedentary

OLIGOCHAETES

Have few bristles

Lack paired parapodia
With around 10,000 known species
Contain terrestrial and aquatic worms

HIRUDINEA

Arose within Oligochaeta

Share a clitellum with oligochaetes forming a clade Clitellata
Do not have setae
External segmentation does not correspond with internal segmentation
Coeloms are dense with connective tissues
Most live in freshwater, while some are found in terrestrial and marine environments

SUBPHYLUM PLEISTOANNELIDA
1. Molecular phylogenies supported two classes: Errantia and Sedentaria
a. Errantia contains freely moving polychaetes
b. Sedentaria contains sedentary annelids like oligochaetes leeches, and some
polychaetes

CLASS ERRANTIA
Errant polychaetes (clam worms)

Most are marine

Live under rocks, in coral crevices, or in abandoned shells
Some are planktonic
May be free-moving, burrowing, or crawling
Most are from 5 to 10 cm long
Important in marine food chains because fish, crustaceans, and many other predators
consume them.
Errant polychaetes are well cephalized.
Prostomium
o May or may not be retractile
o Has eyes, antennae(prostomial tentecles), and sensory palps

Peristomium
o Fist segment surrounds the mouth
o Has chitinous jaws, peristomial cirri (tentacles)
The nervous system of polychaetes consists of:
o Suprapharyngeal ganglia control sensory and motor functions
o Segmental ganglia coordinate movements in isolated segments
o Subpharyngeal ganglia coordinate distant segments

CLASS SEDENTARIA
Contains sedentary polychaetes, oligochaetes, leeches, Siboglinidae, and Echiuridae

Most sedentary tube and burrow dwellers are particle feeders.

Some oligochaetes are scavengers, feeding on decayed organic matter and bits of
vegetation.
Leeches are mostly predator.

MATING AND REPRODUCTION

1.
2.
3.
4.
5.
6.
7.
8.

Two worms align in opposite direction

Clitellum secretes mucus to hold worms together
Sperms from each worm are transported to the seminal receptacles of other worm
After copulation, each worm secretes a mucus tube and chitinous band to form a
cocoon
Cocoon passes forward and eggs, albumin, and sperm are added into it
Fertilization occurs in cocoon
Cocoon slides off head end and closes
Young worms emerge from cocoon

FORMER PHYLUMS
1. Former phylum Echiura are placed within Annelida
Have no segments
Have paired epidermal setae and repeated body parts
Segmentation might have been lost in this group
2. Former phylum Pogonophora are placed within Annelida
Belong in Siboglinidae
Have a small segmented body, opisthosoma
Deep-ocean worms living around hydrothermal vents

B. PHYLUM SIPUNCULA
1.
2.
3.
4.
5.

Unsegmented body
Benthic marine worms
Head is in the form of an introvert
Crowned by ciliated tentacles
250 known species

CH13: ARTHROPODS

Ecdysozoa

1.
2.
3.
4.

Many protostomes possess a cuticle

o Non-living outer layer secreted by epidermis
Cuticle restricts growth and must be molted via ecdysis
Members of Ecdysozoa molt cuticle as they grow
Regulation of molting achieved by the hormone ecdysone
Ecdysozoans do not share same body plan
Members of Nematoda, Nematomorpha, and Kinorhyncha are pseudocoelomates
Members of Loricifera are pseudocoelomates and acoelomates
Members of Priapulida presumed to be pseudocoelomates
Members of clade Panarthropoda have coelomate bodies
Coeloms of onychophorans and arthropods develop by schizocoely
Coeloms of tardigrades develop by enterocoely
But coeloms are small
Have ventrolateral appendages
Hemocoel and open circulatory system

A. PHYLUM ARTHROPODA
1. Most abundant group of animals on earth
Includes spiders, scorpions, ticks, mites, crustaceans, millipedes, centipedes,
insects
2. Segments have coalesced into tagmata
3. Wide range of sizes
4. Variety of habitats; all type of life styles; all type of trophic levels
5. Effect human populations
Important source of products
Cause many diseases and economic losses

B. WHY ARE ARTHROPODS SO DIVERSE AND ABUNDANT?

1. Versatile Exoskeleton:
Composed of a cuticle that is highly protective
o Thin between segments, it allows movement at the joints
o Muscles attach to the cuticle
o Folds inward to line the foregut, hindgut, and trachea
Outer thin epicuticle
o Composed of proteins and lipids
o Tanning process stabilizes and hardens the protein
Inner thick procuticle contain chitin bound with protein
o Exocuticle secreted before molting
o Endocuticle secreted after molting
o Provides protection against dehydration
o Very hard in lobsters and crabs by impregnation with calcium salts
2. Segmentation and appendages for more efficient locomotion
Usually each segment bears a pair of jointed appendages
Modified for various adaptive functions
Appendages may function in sensing, food handling, walking or swimming
3. Air piped directly to cells
Terrestrial arthropods use an efficient tracheal system that delivers oxygen
directly to cells
o
Provides gas transport without use of oxygen-carrying pigments
o Tracheae are composed of a single layer of cells, hypodermis
o Taenidia prevent the tracheae from collapsing
o Spiracles open to the tracheal trunks
o Tracheoles, fluid-filled tubules, reach body cells
o A valve cuts down on water loss
Aquatic arthropods respire by various forms of efficient gills
o Gills attach to appendages, and contain blood vessels and sinuses
4. Highly developed sensory organs
Sensory organs vary from a compound mosaic eye to other senses of touch,
smell, hearing, balancing and chemical reception
A compound eye may consist of thousands of individual photoreceptor units or
ommatidia
Possess a very large view angle, and can detect fast movement.
5. Complex behavior patterns
Surpass most other invertebrates in complex and organized activities
Most behavior is innate or unlearned but some is learned
Some social communities are temporary and uncoordinated
Other social groups are highly organized and depend on chemical and tactile
communication
o Hymenoptera and Isoptera have complex societies with division of labor
o Caste differentiation is common in the most organized social groups
o Parents remain with young and share duties in a cooperative manner
6. Trophic breadth through metamorphosis
Many arthropods have metamorphic changes that result in different larval and
adult stages
Larvae and adults eat different foods and exploit diverse resources

Arthropoda is accepted as monophyletic

1. Previous additions divide arthropods into four subphyla; Trilobita, Chelicerata,
Crustacea, and Uniramia
2. Recent works supports the division of Uniramia into subphyla Myriapoda and
Hexapoda
3. Arthropods with particular mouthpart form the clade Mandibulata includes Myriapoda,
Hexapoda, and Crustacea
4. Arthropods without a mandible have chelicerae
5. Current hypothesis of arthropods interrelationships:
a. Mandibulate hypothesis: myriapods, hexapods and crustaceans are more
closely related to each other than any of them are to chelicerates.
b. Myriochelata hypothesis: myriapods and chelicerates are sister taxa.

C. SUBPHYLUM CHELICERATA
1. Ancient group that includes euryterids (extinct), horseshoe crabs, spider, ticks and
mites, scorpions, sea spiders and others.
2. Characterized by the presence of two tagmata and six pairs of cephalothoraic
appendages that include a pair of chelicerae, a pair of pedipalps, and four pairs
of walking legs.
3. They have no mandible and no antennae.
4. Classes Merostomata, Pycnogonida, Arachnida

CLASS ARACHNIDA
1. Includes spiders, scorpions, pseudoscorpions, whip scorpions, ticks, mites, and
others.
2.
3.
4.
1.

Tagamata are the cephalothorax and an abdomen joined by thin pedicel.

Pedipalps contain sensitive chemical detectors
Their chelicerae function as fangs and bear ducts from their venom glands.
Respiration: spiders breathe by book lungs or tracheae or both.

2. Sensory: spiders usualy have eight simple eyes, each provided with a lens, optic
rods, and a retina. A spiders awareness of its environment depends specially on its
hairlike sensory setae, with which they can sense the vibrations of its web and air
currents.
3. Excretion: they have a unique excretory system of Malpighian tubules that work in
conjunction with specialized rectal glands. They also have coxal glands which
regulate potassium and water to be efficient in dry environment.
4. Web-Spinning Habits: Two or three pairs of spinnerets containing hundreds of
microscopic tubes connect to special abdominal silk glands.
5. Ticks are among the world's premier disease vectors, ranking second only to
mosquitos. They carry agents including protozoan, viral, bacterial, and fungal
organisms.

D. SUBPHYLUM MYRIAPODA
1.
2.
3.
4.
5.
6.

2 tagmata
No compound eye
1 pair of antennae
Mandible and 2 pairs of maxillae
Legs are uniramous
Classes Chilopoda, Diplopoda

E. SUBPHYLUM CRUSTACEA
1. Includes lobsters, crayfishes, shrimp, crabs, water fleas, copepods, and barnacles.
2. Crustaceans are the only arthropds with two pairs of antennae. In addition to
antennae and mandibles, they have two pair of maxillae on the head followed by
a pair of appendages on each body segment.
3. All appendages, except perhaps the first antennae (antennules), are primitively
biramous.
4. Major tagmata are head, thorax, and abdomen, but these are not homologous across
the subphylum.
5. The dorsal cuticle of the head extends posteriorly and around the sides of the animal
to cover or fuse some or all of the thoracic and abdominal segments. This covering is
called carapace.
6. Classes Branchiopoda, Maxillopoda, Malacostraca (largest of all classes)

FORM AND FUNCTION

APPENDAGES

Swimmerets, or abdominal appendages, retain the ancestral biramous condition

Such appendages consist of inner and outer branches, called endopod and exopod,
which are attached to one or more basal segments collectively called a protopod.
Abdominal swimmerets serve not only for locomotion; in males the first pair, called
gonopods, are modified for copulation, and in the females they all serve as a nursery
for attached eggs and young.
The last pair of appendages called uropods, are wide and serve as paddles for swift
backwards movements; with the telson, they form a protective device for eggs or
young in the swimmerets.

ECDYSIS

Permits growth despite a restrictive exoskeleton in crustaceans.

Molting occurs often in young animals and may cease in adults
Underlying epidermis secretes cuticle
During premolt:

Epidermis secrete a new epicuticle and then begin secreting a new exocuticle
Enzymes released into the area above new epicuticle dissolve the old
endocuticle
o Old cuticle becomes thinner as inorganic salts are withdrawn from it and
stored in tissue
Molt (ecdysis):
o When only the old exocuticle and epicuticle remain
o Animal swells with water or air to expand and burst the old cuticle
During postmolt:
o Soft new cuticle stretches and then hardens with the deposition of inorganic
salts forming new endocuticle
The molting process is often initiated by a stimulus perceived by the central nervous
system, that decreases production of a molt-inhibiting hormone from neurosecretory
cells in the X-organ of the eyestalk. The sinus gland, also in the eyestalk, releases
the hormone. When the level of molt-inhibiting hormone drops, Y-organs near the
mandibles produce molting hormone, which initiates the processes leading to
premolt.
o
o

F. SUBPHYLUM HEXAPODA
1. Six legs present in its members
2. All legs are uniramous
3. Hexapods have three tagmatahead, thorax, and abdomenwith appendages on
the head and thorax.
4. Classes Entognatha, Insecta. Class Encognatha has no wings and bases of mouth
parts in head capsule

CLASS INSECTA: INSECTS

Largest and most diverse groups of all arthropods.
Differ from other arthropods in having three pairs of legs and usually two pairs of
wings on the thoracic region of the body, although some have one pair of wings or none.
Bases of mouth parts are visible
Winged called pterygotes and wingless called apterygotes.

EXTERNAL FEATURES

Insect tagmata are head, thorax and abdomen.

The head usually bears a pair of relatively large compound eyes, a pair of antennae,
and three ocelli.
Mouth parts typically include a labrum, a pair each of mandibles and maxillae, a
labium, and a tonguelike hypopharynx.
Insect mouthparts developed from grasshopper primitive chewing mouthparts:
o Sucking mouthparts: Have stylets, and are used to penetrate solid tissue
and then suck up liquid food.
o Siphoning mouthparts: Lack stylets and are used to suck liquids.
o Sponging mouthparts: Lack stylets. Used to sponge and suck liquids.
The thorax is composed of three segments: prothorax, mesothorax, and metathorax,
each bearing a pair of legs. The mesothorax and metathorax each bare a pair of
wings

INTERNAL FORM AND FUNCTION

The insect nervous system in general resembles that of larger crsutaceans.

Dorsal ganglia in head and ventral nerve cord with fusion of ganglia

Neurosecretory cells in various parts of the brain have an endocrine function, but
except for their role in molting and metamorphosis.

METAMORPHOSIS AND GROWTH

Approx. 80% of insects undergo holometabolous (complete) metamorphosis,

which separates the physiological processes of growth (larva) from those of
differentiation (pupa) and reproduction (adult).
o After a series of instars, a larva forms a case or cocoon around itself and
becomes a pupa, a nonfeeding stage in which many insects pass the winter.
Some
insects
undergo
hemimetabolous
(gradual
or
incomplete)
metamorphosis, where the young are called nymphs and their wings develop
externally as budlike outgrowths in the early instars and increase in size as the
animal grows by successive molts and becomes winged adult.