PHYLUM ANNELIDA

Annelida is a monophyletic assemblage of Marine, freshwater and terrestrial worms. They are
lophotrochozoans and,thus,share common ancestry with Mollusca, Brachiopoda, Bryozoa,
Nemertea and others. This Diverse phylum originated in Precambrian Times (more than six
hundred million years ago).

Characteristics of Phylum Annelida

● Their body is metameric, bilaterally symmetrical and wormlike.
● They show protostome characteristics.
● They have spiral cleavage.
● They develop trochophore larvae.
● They have paired epidermal setae for grip and locomotion.
● They closed circulatory system
● They have dorsal suprapharyngeal ganglia and ventral nerve cords with ganglia.
● They develop metanephridia and protonephridia.
● They have Schizocoelous coelom formation.
● Excretion: by nephridia
● Reproduction: sexual or hermaphroditic.
● Fertilization: internal or external
● Development: direct with no larval stages
● Germ layer: triploblastic
● Body possesses 3 separate sections.
1. A prostomium
2. A trunk
3. A pygidium

Schizocoelous origin:
It suggests that the coelom of the annelids has schizocoelous origin. It means that annelids have
evolved from ancient flatworm stock.
Enterocoelous origin:
It suggests the coelom origin is Enterocoelous. It means that the ancient diploblastic animals are
ancestors of annelids.

Lobatocerebrum:
Recently a worm Lobatcerebrum has been discovered. It shows both annelids and Flatworm
Characteristics.
Therefore, it gives support to the enterocoelous origin hypothesis. Lobatocerebrum is classified
as an annelid.
It has certain Annelids characters:
● They have segmentally arranged Excretory Organs.
● They have Annelid-like body covering.
● They have a complete Digestive Tract.
● They have an Annelid-like nervous system.
But it has a ciliated epidermis and is acoelomate like flatworms. Some zoologists believe that
Lobatocerebrum indicates the triploblastic, acoelomate animals can derive from the annelid
lineage.

Metamerism and Tagmatization:

Segmental arrangement of body parts of an animal is called metamerism (meta,after+mere,part).
● Earthworm Bodies are divided into a series of ring-like segments (metameres). The Body
is also divided Internally.
● There is compartmentalization of the body. Therefore, each segment has its own
excretory, nervous and circulatory structures.
● There are two related functions of metamerism:
1. flexible support
2. efficient locomotion.
● Externally marked by circular rings called Annuli (the name of the phylum refers to this
characteristic).
Origin of coelom:
The body cavity of annelids arises by segmental splitting of mesoderm during the embryonic
stage.
Mesoderm occupies the region between ectoderm and endoderm on both sides of the embryonic
gut. Each cavity is enlarged, and it forms a double membrane septum. This septum separates
the anterior and posterior margin of each coelomic space.

Origin of Muscles:
Muscles also develop from the mesodermal layer of each segment.
A layer of circular muscles lies below the Epidermis.
A layer of longitudinal muscle is present just below–the circular muscles.
These muscles run between the septa that separate each segment.
● Some Polychaetes also have oblique muscles.
● Leeches have dorsoventral muscles.

Advantages of Metamerism:
The coelomic spaces and muscles produce hydrostatic compartments.
1. It has locomotory and supportive functions.
Each segment is controlled independently of other segments.
The muscles can act as antagonistic pairs within segment. The Coelomic fluid provides a
hydrostatic skeleton. The Muscles operate against it. These arrangements cause swimming,
crawling and burrowing.
2. A second advantage is that it lessens the impact of injury. If one of a few segments is
lured, adjacent segments are separated from injured segments by septa. They maintain
nearly normal functions. It increases the chance of survival.
3. Metamerism causes modification in certain regions of the body. They become
specialized for feeding, locomotion and reproduction.

The specialization of body regions in metameric animals is called tagmatization.

Tagmatization is best developed in the Arthropods. But some Annelids also display
tagmatization. Because arthropods are closely related.
Triploblastic Coelomate Organization, bilateral symmetry, a complete digestive tract, and ventral
nerve cord. It means that Arthropods and Annelids have common ancestors. Their ancestors may
be marine, wormlike, bilateral animals.

External Structure and Locomotion

Parapodia And Setae:

Polychaetes develop parapodia but are absent in oligochaetes.
● Parapodia are lateral extensions of the body wall. Chitinous Rods support the parapodia.
Numerous setae projects from the parapodia. The name of the class is derived from the presence
of numerous setae.
● Setae are bristles structures. They are secreted by the invaginations of the distal ends
parapodia. These setae dig into the Substrate. Therefore, they help in locomotion.

Prostomium & Epidermis

● The prostomium of a polychaete is a lobelike structure. It projects dorsally and anteriorly
from the mouth with many sensory structures like eyes, antennae, palps. It also contains
ciliated pitsor grooves called nuchal organs.
● The First body segment is the periosteum, surrounds the mouth, and it bears sensory
tentacles or cirri.
● The epidermis of annelids consists of a single layer of columnar cells. These cells
secrete protective, nonliving cuticles. Some have epidermal glands that secrete
luminescent compounds.

Locomotion
Locomotion takes place by antagonistic contraction of circular and longitudinal muscles. The
waves of contraction move from rear to front.
● The longitudinal muscles contract and segments bulge and setae protrude out. Therefore,
the setae penetrate the soil. Contraction of longitudinal muscles in segments behind a
bulging region pulls those segments forward.
 ● The contraction of circular muscles retracts the setae. It elongates the segments and
pushes them forward.
The waves of muscle contraction move anteriorly on worms. Therefore, the segments move
forward.

Feeding And The Digestive System

Scavengers feed on fallen and decaying vegetation. They drag these decay food into their
burrows at night.
The digestive tract is tubular and straight.
● Mouth: The mouth opens into a muscular pharynx.
● Pharynx: The pharyngeal muscles attach to the body wall. The pharynx acts as a pump
for ingesting food. The pharynx pumps the food into the esophagus.
● Esophagus: The esophagus is narrow and tubular. It expands to form a stomach, crop or
gizzard. Crop and gizzard are common in terrestrial species.
● Gizzard: A gizzard is a muscular structure. Its inner wall is lined with cuticles. Gizzard
is a grinding structure. Calciferous glands are present in the invagination of the
esophageal wall. They are used to remove excess calcium absorbed from food. They also
help to regulate the pH of body fluids.
● Intestine: The intestine is a straight tube. It is the principal site of digestion and
absorption. A dorsal fold of the epithelium is present in the intestine. It is called
typhlosole. It increases the surface area of the intestine. The intestine ends at the anus.

Gas Exchange And Circulation

● Respiration
Respiratory gasses simply diffuse across the body wall. The parapodia increase the surface area
for gaseous exchange.
● Circulation
Closed circulatory system.
Oxygen is carried by respiratory pigments. The respiratory pigments are dissolved in the plasma.
They are present in the blood cells in other animals. Blood is colorless, green or red, depending
on the respiratory pigment.
It has following circulation pattern:
Dorsal aorta: The dorsal aorta acts as contracting elements. Dorsal aorta lies just above the
digestive tract. It propels blood from rear to front. In the front, the blood moves into ventral
aorta.
Ventral aorta: It is present ventral to the digestive tract. It propels blood from front to rear.
Hearts: Some segmental vessels expand, and they are contractile. These are sometimes called
hearts.
Segmental vessels: Two or three sets of segmental vessels are present between dorsal and
ventral vessels. Segmental vessels receive blood from the ventral aorta and break into capillary
beds in the gut and body wall.
Capillaries: Capillaries unite again into segmental vessels. These vessels deliver blood to the
dorsal aorta.

Nervous And Sensory Functions:

● The ventral nerve cords and all ganglia show a high degree of fusion. The giant fibers
produce escape responses. An escape response results from the stimulation of the
anterior or posterior end of a worm.
● Some are burrowing animals. Therefore, they lack well-developed eyes. Animals living
in continuous darkness do not have well-developed eyes.
● Other oligochaetes have simple pigmented cup called ocelli. Oligochaetes are sensitive to
different chemical and mechanical stimuli. Receptors of these stimuli are scattered over
the body surface, especially around the prostomium.
● Photoreceptor cells scattered over the dorsal and lateral surfaces of the body. Therefore,
they all have a “dermal light sense”. Photoreceptor cells produce a negative phototaxis
in strong light and a positive phototaxis in weak light.

Excretion:
Oligochaetes use metanephridium for excretion and osmoregulation.
● The Nephrostome of metanephridium is present in the anterior segment. Their tubule and
nephridiopore are present in the posterior segment. Nitrogenous wastes are ammonia and
urea.
 ● Oligochaetes possess chloragogen tissue. These tissues surround the dorsal blood vessel.
Chloragogen tissue acts as a vertebrate liver. It is a site of amino acid metabolism.
Chloragogen’ tissue deaminates amino acids and converts ammonia to urea. It also
converts excess carbohydrates into glycogen and fat.

Reproduction And Development

● Oligochaetes are monoecious.

● In Lumbricus reproductive systems are found in somites 9 to 15.
● Two pairs of small testes and two pairs of sperm funnels are surrounded by three pairs of
large seminal vesicles. Immature sperm from the testes mature in seminal vesicles, then
pass into sperm funnels and down sperm ducts to the male genital pores in somite 15,
where they are expelled during copulation.
● Eggs are discharged by a pair of small ovaries into the coelomic cavity, where ciliated
funnel of the oviducts carry them outside through female genital pores on somite 14.
Two pairs of seminal receptacles in somites 9 and 10 receive and store sperm from the
mate during copulation.
● When mating, worms extend their anterior ends from their burrows and bring their
ventral surfaces together. They are held together by mucus secreted by their clitellum and
by special ventral setae, which penetrate each other’s bodies in the regions of contact.
● After discharge, sperm travel to seminal receptacles of the other worm in its seminal
grooves.

Copulation:
● Two worms line up facing opposite directions during copulation. Their ventral surfaces
and their anterior ends are in contact with each other. This arrangement lines up the
clitellum of one worm with the genital segments of the other worm.
● Clitellum secretes a mucous sheath. This sheath covers the anterior halves of both
worms. This sheath holds the worms in place.
 ● The Sperm duct releases sperm. Sperms travel in a groove, muscular contractions along
this groove propel the sperm toward the openings of the seminal receptacles. In other
Oligochaetes, there is direct transfer of sperm into seminal receptacles during copulation.
Cocoon Formation:
● After copulation, each worm secretes first a mucous tube and then a tough, chitinlike
band that forms a cocoon around its clitellum. The cocoon consists of mucoid and
chitinous materials.
● As the cocoon passes forward, eggs from the oviducts, albumin from skin glands, and
sperm from the mate (stored in the seminal receptacles) pour into it.
● Fertilization occurs in the cocoon. When the cocoon leaves the worm, its ends close,
producing a lemon- shaped body. Embryogenesis occurs within the cocoon, and the form
that hatches from the egg is a young worm similar to the adult.

Development:
● Spiral cleavage is modified.
● Larva is not formed in them.
● Hatching occurs in one to few weeks. The young worms emerge from one end of the
cocoon.
● Thus, development is direct with no metamorphosis.
● Juveniles do not develop a clitellum until they are sexually mature.

Asexual reproduction:
● Freshwater Oligochaetes also reproduce asexually.
● Asexual reproduction takes place by the transverse division of a worm.
● The missing segments are regenerated.
 Classification

Class polychaeta
● Polychaeta is the largest of the annelid classes.
● It is the class of bristle worms.
● They are 5 to 10 cm long. They have more than 5300 species.
● Polychaetes are adapted to a variety of habitats. Many live on the ocean floor, under
rocks and shells. Some live within the crevices of coral reefs. Other polychaetes are
burrowers.
● They move through their substrate by peristaltic contractions of the body wall.
● True coelomates (schizocoelomates), Dioecious or hermaphroditic.
● Bilaterally symmetry, segmented worms. Lateral epidermal setae with each segment.
● Complete digestive system, Closed circulatory system, Well-developed nervous system.
● Excretory system: Both metanephridia and protonephridia.
● Fertilization: external
● Example: Rag worms, Lugworms, Bloodworms

Class Oligochaeta
● The organisms belonging to this class are monoecious.
● 3000 species. They are found throughout the world in freshwater and terrestrial habitats.
A few oligochaetes are estuarine, some are marine. Aquatic species live in shallow water,
where they burrow in mud and debris. Terrestrial species live in soils, some live in hot,
dry weather. The depths of their burrow are 3 m below the surface.
● The body is segmented metamerically.
● Head, eyes and tentacles are not distinct.
● They are hermaphrodites, but cross-fertilization takes place. Fertilization is external.
● Cocoon formation occurs. They exhibit no free larval stage and the development takes
place inside the cocoons.
● Setae are segmented.
● They do not possess parapodia, but clitellum is present.
● Example: Lumbricus terrestris (Common earthworm)
Class Hirudinea
● The animals are monoecious.
● The class Hirudinea contains 500 species of leeches. Most leeches are fresh water. Some
are marine. Some are completely terrestrial. Leeches prey on small invertebrates or they
feed on the body fluids of vertebrates.
● Setae are absent in most of leeches
● Leeches lack 2 appendages (Parapodia, Head).
● The body is segmented.The body is dorsoventrally or cylindrically flattened.
● They have an anterior and posterior sucker on the ventral side. The mouth is located
ventrally in the anterior sucker, while the anus is present dorsally in the posterior sucker.
● The organisms lay eggs in cocoons.
● There is no larval stage during the development of the organism.
● Fertilization is internal. They are hermaphrodites.
● Example: Hirudinaria