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Alex Shayo, D (PhD)

Overview of the phylum

Annelids are triploblastic, coelomate protostome

Phylum annelida consists of the segmented worms
such as earthworms, Nereis and leeches.
Annelids have a hydrostatic skeleton.
Locomotion in most annelids is aided by numerous
fine chitinous hairs, called setae or chaetae. The
annelid gut is a straight tube supplied with its own
The nervous system is concentrated anteriorly
(cephalization) into cerebral ganglia from which
arises a ventral nerve cord with segmental ganglia.

Characteristics of Annelids


The body is segmented (metameric) and

bilaterally symmetrical.
Have paired epidermal setae or chaetae for
movement (Setae absent in Leeches).
The circulatory system is closed and has
segmentally arranged respiratory pigments
and amebocytes in blood plasma.
The excretory system typically consists of a
pair of nephridia for each metamere.
Respiratory gases exchange through the
skin, gills or parapodia.

The Annelid body plan

Annelids have elongated worm-like bodies.
The body is divided into similar rings or segments

arranged in linear series.

The segments are externally marked by circular rings
called annuli (the characteristic to which the name of
this phylum refers).
The division of the body into a series of segments,
each of which contains similar components of all
major organ systems is called metamerism.
In the evolutionary tree, annelida is the first group of
animals to show metamerism.
Metamerism evolved independently in annelids,
arthropods and vertebrates.

Annelid body plan cont

Body structure of an annelid

Annelid body plan cont

The annelid body typically consists of an anterior

Prostomium, followed by a segmented body, and

a terminal portion called Pygidium.
During development, new segments differentiate
just in front of the pygidium, thus the oldest
segments are at the anterior end where as the
youngest are at the posterior end.
Anterior segments usually fuse with the

prostomium to form the head.

Annelids have a body covered by an external cuticle

that is never shed or molted.

The body wall has strong circular and longitudinal
muscles which work antagonistically.

Significance of Metamerism
Metamerism increases the efficiency of
body movement by allowing the effect of
muscle contraction to be extremely
localized. Resultant localized changes in
the shape of groups of segments provide
the basis for swimming, crawling, and
2. The repetition of body parts provides
safety to the organisms because if one
segment fails, it may not necessarily lead
to fatal consequences as there are still
other segments which can complement
the functions of the failed segment.

Significance of Metamerism
3.Metamerism permits modification of certain
regions of the body for specialized functions such
as feeding, locomotion, and reproduction.
The specialization of body regions in a
metameric animal is called tagmatization.
Tagmatization is best developed in arthropods;
annelids only show some advent of this regional
specialization of body segments.


The annelid coelom

Annelids are coelomates.
They have a complete gut and a ventral nerve cord.
The annelid coelom is formed by splitting of the

embryonic mesoderm (Schizocoely).

The coelom is segmented such that coelomic
cavities of adjacent segments are separated by
The septa are perforated by the gut and blood


The annelid coelom cont

Except in leeches, the coelom of most annelids is

filled with fluid which serves as a hydrostatic

Because the volume of fluid in each segment is
essentially constant, the contraction of the
longitudinal muscles causes the body to shorten and
become larger in diameter, where as the contraction
the circular muscles makes the body thinner and
The separation of the hydrostatic skeleton into
segments of coelomic cavities greatly increased its
efficiency because the force of local muscle
contraction within one segment is not transferred and
dumped throughout the length of the animal.


Relationship of Annelids to other

Annelids are protostomes.
Recent reevaluation of the phylogenetic relationship within

the protostome lineage suggests that annelids are more

closely related to Molluscs than to Arthropods.
Based on the evidence from the reevaluated phylogeny,
some zoologists suggest that the protostome lineage should
be split into two major clades (super-phyla) namely;



This is the clade of the moulting protostomes.
This clade includes the Phylum Nematoda, Nematomorpha,

Kinoryncha, Loricifera, Priapulida, and Arthropoda.

The proposed synapomorphies for this clade include a
cuticle, loss of epidermal cilia, and shedding of the cuticle in
a process called ecdysis.
Placement of the Arthropoda and Annelida into separate
clades requires that their common form of metamerism
resulted from convergent evolution.
However, there is still a debate among zoologists on
whether the observed metamerism in Annelids and
Arthropods are homologous or analogous.


This is the clade of the non-moulting protostomes.
This clade includes the Annelida, Mollusca, Rotifera, and



Classification of the Phylum

The phylum annelida was traditionally divided into three

classes; Polychaeta, Oligochaeta and Hirudinea.

However, recent cladistic analysis of the phylum resulted
in other interpretations which suggest the reduction of the
number of classes into just two.
According to this reinterpretation, the former classes
Oligochaeta and Hirudinea have been demoted into
subclasses under a new class Clitellata.


Classification of the Phylum

Annelida cont
It is suggested that the presence of a clitellum which is

used in cocoon formation, monoecious direct development

and a few or no setae are symplesiomorphies for the clade
containing the two former classes.
The class Polychaeta remained intact without changes.
The revised cladistic classification for this phylum will be
adopted in this lecture because it has already gained
universal recognition and acceptance by most Zoologists.


("many bristles)

This is the largest class in the phylum Annelida.

It consists mainly of marine annelids such as
Nereis, Arenicola, Sabela etc.
Members of this class have the following
Have a head with eyes and tentacles.
Have parapodia with numerous setae.
Are either monoecious or dioecious.
Development frequently involves a
trochophore larval stage.


Habits and Habitats of

Most polychaetes live on the ocean floor,

under rocks and shells, and within the

crevices of coral reefs.
Some polychaetes burrow into the
intertidal sand and others construct
tubes of cemented sand or secreted
organic material.
The mucus-lined tubes serve as
protective retreats and feeding stations.


Support and Movement

Polychaetes have numerous lateral paddle-like extensions called

parapodia each of which bears numerous setae.

The parapodia and setae aid in locomotion, digging through the substrate,

holding the worm in its burrow/tube and in gaseous exchange.

During movement, the longitudinal muscles on one side of the body acts

antagonistically to the longitudinal muscles on the other side of the body

so that undulatory waves move along the length of the body from the
posterior end towards the head.

The parapodia and setae act against the substrate or water to propel the


Burrowing polychaetes push their way through the sand by contractions

of the body wall or by eating their way through the substrate.


Feeding and Digestion

Polychaetes may be predators, herbivores,

scavengers, filter feeders or deposit feeders.

The anterior region of the polychaete digestive tract is
modified into an eversible proboscis which when
everted, paired jaws are opened and may be used for
seizing prey.
Digestion in polychaetes occurs extracellularly in the
digestive system.
Polychaetes that inhabit substrates rich in dissolved
organic matter can absorb as much as 20 40% of
their energy requirements across their body wall as
sugars and other soluble organic compounds.


Excretion and Water Balance

Annelids excrete nitrogenous wastes in the form of

ammonia most of which diffuses across the body wall

into the surrounding water.
Most polychaetes posses metanephridia for excretion.
A few primitive polychaetes posses protonephridia.
A metanephridium consists of an open, ciliated funnel
called a nephrostome that projects through an anterior
septum into the coelom of an adjacent segment.
A protonephridium consists of a tubule with a closed
bulb at one end and a connection to the outside of the
body at the other end.
A Protonephridium has a tuft of flagella at the bulbular
end that drives fluid through the tubule.


Gas Exchange and Circulatory

The respiratory gases in most polychaetes simply

diffuse across the body wall and parapodia.

Many polychaetes have parapodial gills which
increase the surface area for gas exchange.
All annelids (including polychaetes) have a closed
circulatory system.
Oxygen is carried with molecules called respiratory
pigments which are dissolved in the blood plasma
rather than contained in blood cells as in vertebrates.
The annelid blood may be colourless, green, or red
depending on the type of respiratory pigment


Gas Exchange and Circulatory

System cont
The blood vessels of polychaetes consist of a dorsal

aorta that lies just above the digestive tract and a

ventral aorta which lies ventral to the gut.
The dorsal aorta propels blood from rear to front and
the ventral aorta conveys blood from front to rear.
Two or three sets of segmental blood vessels run
between the dorsal and ventral aorta.
These segmental vessels receive blood from the
ventral aorta and break into capillary beds in the gut
and the body wall before they coalesce again into
segmental vessels that deliver blood to the dorsal


The Nervous System

The nervous system is similar in all annelids.
It consists of a pair of suprapharyngeal ganglia

which connect to a pair of subpharyngeal ganglia

by circumpharyngeal connectives that run
dorsalventrally along either side of the pharynx.
The rest of the central nervous system is generally
ladder-like, consisting of a pair of nerve cords that
run through the ventral part of the body and have
in each segment paired ganglia linked by a
transverse connection.
Lateral nerves emerge from each segmental
ganglion, supplying the body wall musculature and
other structures in that segment.


The Nervous System cont

Polychaetes have various sensory structures which

include palps, antennae, eyes, statocysts, nuchal organs

and lateral organs.
Palps and antennae are located on the head of many
polychaetes. The palps are used for feeding.
Nuchal organs are ciliated, paired, chemosensory
structures, innervated from the posterior part of the brain.
Two or four pairs of eyes are on the surface of the
prostomium. They vary in complexity from a simple cup of
photoreceptor cells (ocelli) to structures made up of
cornea, lens, and vitreous body.
Most polychaetes react negatively to increased light


Reproduction and Development

Most polychaetes are gonochoric (having the sexes

separate) and reproduce sexually.

However, some primitive polychaetes reproduce

asexually by budding or transverse fission.

In sexually reproducing worms, the gametes are
shed to the coelom where they mature.
Mature female worms are often packed with eggs.
Eggs exit the worm through the nephridiopores by
entering the nephrostomes of metanephridia or by
rupturing of the mature worms.
Fertilization in polychaetes is external, (only few
species copulate).


Reproduction and Development

Spiral cleavage of the zygote results into planktonic

trochophore larvae that later on settle and

metamorphose into juvenile worms.
Most polychaetes show a characteristic reproductive
behaviour called epitoky.
Epitoky is a mode of reproduction unique to
polychaetes in which the worm undergoes a partial or
entire transition into a pelagic, sexually reproductive
form, known as an epitoke.
In many cases, epitoky involves degeneration of
digestive structures and enhancement of swimming,
sensory, and reproductive structures.
Degeneration of the digestive system and other
internal organs makes epitokes to have a short life


Reproduction and Development





During the brief reproductive season, epitokes swarm in large

numbers on the surface of water bodies.
Swarming of the epitokes has the following significances:
Because the non-reproductive individuals remain safe below
the surface waters, predators can not devastate the entire
External fertilization requires that individuals become
reproductively active at the same time and in close proximity
to one another in order to increase the chances for
fertilization. Swarming of epitokes ensures that large numbers
of individuals are in the right place at the proper time.
Swarming of many epitokes for a brief period of time provides
safety against predation since predators can not consume
them all. Predators will eat and satisfy their hunger and yet
leave enough epitokes to yield the next generation (there is
safety in numbers).


Class Polychaeta: Diversity

Although a number of

polychaetes are active

predators, some are sedentary
and burrow into mud or live in
protective tubes in the mud
In several of these species
filter feeding has evolved
A good example is the fan
worm Sabella, with their
feather-like head structures
called radioles


Class Polychaeta: Diversity cont.

Chaetopterus is tube dweller;

lives in a U-shaped tube

Parapodia are highly modified
into 3 fan-like structures that
bring water into the tube
The notopodium secretes a
mucous bag that traps food from
the water flowing through the
tube; the bag is periodically
passed anteriorly toward the


Class Polychaeta: Diversity cont.

Arenicola lives in a J-shaped burrow
It employs peristaltic movements to generate a water

Food is filtered out from the front of the burrow


Members of this class include the earthworms and

They are characterized by the presence of a
clitellum for cocoon formation, monoecious/
hermaphrodism, direct development, and a few or
no setae on the integument.
This class consists of two major subclasses

Subclass Oligochaeta
Subclass Hirudinea


A. Subclass Oligochaeta
("few bristles")

This subclass includes the earthworms and their

There are over three thousand Oligochaete species all
over the world.
Most Oligochaetes inhabit fresh water and terrestrial
A few inhabit estuaries and marine environments.


External structure and

Oligochaetes lack parapodia, and have fewer and

shorter setae than polychaetes as these would

interfere with their burrowing lifestyle.
A series of segments in the anterior half are
modified to form a swollen girdle-like structure called
the clitellum whose function is to secrete cocoon
and mucus during copulation.
Oligochaetes move by peristalsis which involves
antagonism of the circular and longitudinal muscles
in groups of segments.
Neurally controlled waves of contraction move from
rear to front, causing the segments to bulge and
retract sequentially.


Feeding and the Digestive

Most Oligochaetes scavenge on fallen and decaying vegetation

which they drag into their burrows at night.

During burrowing, earthworms swallow considerable quantities of soil
which they egest as soil castes.
Calciferous glands along the oesophagus secrete calcium ions into
the gut and so reduce the calcium ion concentration of their blood.
The pharynx acts as a pump for forcing food down the oesophagus
during ingestion.
The oesophagus is narrow but expands at some portions to form a
crop, gizzard or stomach.
The crop is a temporary store of food, the gizzard grinds the food.
The intestine is a straight tube and is the principal site for digestion
and absorption.
The wall of the intestine is infolded dorsally which greatly increases
the surface area for digestion and absorption.


Earthworm digestive system


Anatomy of the Earthworm


Nervous System and Sense Organs

The nervous system includes a pair of cephalic ganglia

attached to a double nerve cord that run the length of the

animal along the ventral body wall, with ganglia and branches
in each segment.
The ventral nerve cords and all ganglia in Oligochaetes have
undergone a high degree of fusion.
Other aspects of the nervous system are the same as those of
the polychaetes.
However, most lack well developed eyes due to their
burrowing lifestyle.
Oligochaetes are sensitive to a variety of chemical and
mechanical stimuli as they have some combination of tactile
organs, chemoreceptors, balance receptors, and


Respiratory system
Earthworms have no respiratory organs.
Gaseous exchange occurs by diffusion across the skin.
The skin is always moist to allow efficient exchange of

respiratory gases.


Oligochaetes excrete nitrogenous wastes in form of ammonia and

They have metanephridia for excretion and for ion and water balance.
Oligochaetes excrete copious amounts of very dilute urine, although
they retain vital ions which are important for organisms living in an
environment where water is plentiful but essential ions are limited.
Just as with polychaetes, the metanephridia of oligochaetes are
associated with the segment just anterior to the segment containing
the tubule and the nephridiopore.
Oligochaetes (as well as other annelids) possess Chloragogen tissue.
The chloragogen tissue is a site for metabolism of amino acids and
The chloragogen tissue, deaminates amino acids, converts ammonia
to urea, and converts excess carbohydrates into glycogen and fat.


Reproduction and Development

All Oligochaetes are monoecious (Hermaphrodites) but

copulate, with mutual exchange of sperm.

The clitellum secretes mucus, after which the sperm leave
the sperm ducts and travel to the seminal receptacles of
the partner.
The clitellum later produces a slime tube, which is moved
along over the head of the worm by muscular contractions.
Into this tube are deposited eggs from the oviducts and
sperm from the seminal receptacles.
The slime tube forms a cocoon within which the miniature
worms develop.
There is no a larval stage.


Oligochaetes differ from polychaetes in

several ways:
No parapods, fewer setae (if at all)
No larval stages



B. Subclass Hirudinea
This subclass consists of the leeches.
Leeches are primarily freshwater annelids, but some live in

the ocean and some in moist soil or vegetation.

Leeches range in length from about 1 cm20 cm; most are
less than 5 cm long.
They are commonly black, brown, green, or red, and may
have stripes or spots.


Structure of Leeches
Unlike other annelids, Leeches lack parapodia, setae and

head appendages.
They are dorsalventrally flattened and taper anteriorly.
A less conspicuous clitellum is present.
Leeches are equipped with anterior and posterior suckers
used for creeping and feeding.
Leeches are the only annelids with a fixed number (34) of
body segments.
However, these segments are difficult to distinguish
externally because have become divided into secondary
subdivisions known as annuli.
The coelom is not subdivided by septa.
The muscles are complex than that of other annelids.


Locomotion in Leeches
Because of the modifications in the coelom and

body musculature, the pattern of movement in

leeches has been altered accordingly.
Leeches move in a looping type of locomotion by
utilizing the anterior and posterior suckers.
A leech moves forward the small anterior sucker,
attaches it, and draws up the larger posterior
Most leeches can swim by rapid undulations of the
body, using well-developed muscles of the body


Feeding and the Digestive

Most leeches are predators of small invertebrates.
They swallow their prey whole, but some suck the soft parts from their

Many leeches have a proboscis used for swallowing the prey or for
sucking its fluids; others have jaws for biting.
Many parasitic leeches are able to parasitize a wide variety of hosts.
The distinction between predatory and parasitic leeches is not sharp as
many predatory leeches take blood meals on occasion.
Most parasitic leeches attach to the host only while feeding; a single
meal may be 5 or 10 times the weight of the leech and provide it with
food for several months.
The digestive tract of bloodsuckers produces an anticoagulant, hirudin,
which keeps the engorged blood from clotting.
A few leeches attach permanently to the host, leaving only to reproduce.
Predatory leeches are active at night and hide by day.


The salivary glands excrete hirudin which

prevents the blood from coagulating

May also secrete an anaesthetic and
substance to dilate small blood vessels
Blood is broken down by symbiotic
bacteria that is then used by the leeches
Leeches were commonly used in the 19th
century for bloodletting
Recent medical uses are to relieve
pressure after vascular tissue is damaged
Snake bites or the reattachment of a finger
or ear


Gas exchange and Circulation

The coelom of leeches differs from that of other

annelids in that it is largely filled in with tissue and

lacks septa.
Coelomic fluid is contained in a system of sinuses,
which in some leeches functions as a circulatory
system; there is a tendency in this group toward
the loss of true blood vessels.
The blood of some leeches is red. In others the
blood lacks oxygen-carrying pigments and is
therefore colorless.
Gas exchange occurs throughout the body surface
of most leeches, although many fish-parasitizing
leeches have gills.


Nervous and Sensory Function

The sense organs consist of sensory cells of various types,

including photoreceptor cells, scattered over the body

There are also 2 to 10 eyes, consisting of clusters of
photoreceptor cells located toward the front part of the body.


Like the oligochaetes, leeches are hermaphroditic and

cross-fertilizing, but fertilization is internal.

In some Leech species the sperm are enclosed in sacs,
called spermatophores, that are attached to the outside of
the partner; the sperm pass through the body wall to the
ovaries, where the eggs are fertilized.
In other species the sperm are not enclosed and are
transferred directly into the body of the partner during
A courtship display is seen among some leeches at the time
of mating.
The fertilized eggs are deposited in a cocoon, secreted by
the clitellum; the cocoon is buried in mud or affixed to
submerged objects.
The young emerge as small copies of the adults.


Leeches excrete nitrogenous wastes by using 10-17

pairs of highly modified metanephridia.

The leech metanephridium has a capsule believed to
be involved in the production of coelomic fluid.
Chloragogen tissue is present
Chloragogen is a substance in cells which serves as a center

for glycogen and fat synthesis in oligochaetes



Polychaetes (about 12,000 species). As their name suggests, they have

multiple chetae ("hairs") per segment. Polychaetes have parapodia that

function as limbs, and nuchal organs that are thought to be
chemosensors. Most are marine animals, although a few species live in
fresh water and even fewer on land

Clitellates (about 10,000 species). These have few or no chetae per

segment, and no nuchal organs or parapodia. However, they have a

unique reproductive organ, the ring-shaped clitellum around their bodies,
which produces a cocoon that stores and nourishes fertilized eggs until
they hatch or, in moniligastrids, yolky eggs that provide nutrition for the
embyros. The clitellates are sub-divided into:

a) Oligochaetes ("with few hairs"), which includes earthworms.

Oligochaetes have a sticky pad in the roof of the mouth. Most are
burrowers that feed on wholly or partly decomposed organic materials.
b) Hirudinea whose name means "leech-shaped" and whose best known
members are leeches. Marine species are mostly blood-sucking parasites,
mainly on fish, while most freshwater species are predators. They have
suckers at both ends of their bodies, and use these to move rather like


Economic Importance of
1. Earthworms cycle huge quantities of soil through


their guts, a process that speeds the turnover of

nutrients in the soil and increases productivity.
Their burrows help to aerate the soil.
Aquatic annelids are important source of food for
larger invertebrates and fish.
Annelids are commonly used by fishermen as fish
A few annelids are ectoparastic e.g. the leeches.
The European Medical Leech (Hirudo medicinalis)
has been used for clinical bloodletting for
thousands of years.