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THE UNIVERSITY OF DODOMA

SCHOOL OF BIOLOGICAL SCIENCES


BI 112- INVERTEBRATE ZOOLOGY

PHYLUM ANNELIDA
Alex Shayo, D (PhD)

Overview of the phylum


Annelids are triploblastic, coelomate protostome

animals.
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
musculature.
The nervous system is concentrated anteriorly
(cephalization) into cerebral ganglia from which
arises a ventral nerve cord with segmental ganglia.

Characteristics of Annelids
1.
2.
3.

4.
5.

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
burrowing.
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.
1.

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.

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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
septa.
The septa are perforated by the gut and blood
vessels.

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The annelid coelom cont


Except in leeches, the coelom of most annelids is

filled with fluid which serves as a hydrostatic


skeleton.
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
longer.
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.

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Relationship of Annelids to other


Animals
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;

Ecdysozoa
Lophotrochozoa

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Ecdysozoa
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.

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

Acanthocephala.

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Classification of the Phylum


Annelida
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.

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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.

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1. CLASS POLYCHAETA
("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
characteristics;
Have a head with eyes and tentacles.
Have parapodia with numerous setae.
Are either monoecious or dioecious.
Development frequently involves a
trochophore larval stage.

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Habits and Habitats of


Polychaetes
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.

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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

animal.

Burrowing polychaetes push their way through the sand by contractions

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

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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.

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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.

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Gas Exchange and Circulatory


System
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
present.

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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
aorta.

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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.

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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
intensities.

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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).

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Reproduction and Development


cont
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
span.

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Reproduction and Development


cont

1.

2.

3.

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
population.
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).

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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

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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
mouth

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Class Polychaeta: Diversity cont.


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

flow
Food is filtered out from the front of the burrow

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2. CLASS CLITELLATA
Members of this class include the earthworms and

leeches.
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
namely;

Subclass Oligochaeta
Subclass Hirudinea

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A. Subclass Oligochaeta
("few bristles")

This subclass includes the earthworms and their


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

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External structure and


Locomotion
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.

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Feeding and the Digestive


system
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.

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Earthworm digestive system

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Anatomy of the Earthworm

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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
photoreceptors.

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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.

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Excretion
Oligochaetes excrete nitrogenous wastes in form of ammonia and

urea.
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
carbohydrates
The chloragogen tissue, deaminates amino acids, converts ammonia
to urea, and converts excess carbohydrates into glycogen and fat.

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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.

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Note:
Oligochaetes differ from polychaetes in

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

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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.

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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.

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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
sucker.
Most leeches can swim by rapid undulations of the
body, using well-developed muscles of the body
wall.

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Feeding and the Digestive


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

victims.
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.

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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

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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.

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Nervous and Sensory Function


The sense organs consist of sensory cells of various types,

including photoreceptor cells, scattered over the body


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

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Reproduction
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
copulation.
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.

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Excretion
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

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Summary

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
inchworms.

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

2.
3.
4.
5.
6.

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
bait.
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.

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