Subkingdom: Metazoa

(Multicellular animals)

It is divided into 31 phyla and comprises large animals (vertebrates) which

have backbone and small animals (invertebrates). The vertebrates are also called "the
higher animals" and the invertebrates are "the lower animals".

Metazoan Phylogeny (Evolution of Metazoa):

 It is an important topic in marine biology.
 Biologists believe that the earliest multicellular animals evolved from one
protozoan type (probably flagellated protozoa).
 The lowest phylum is porifera (sponges) having little evolutionary affinity.
 On the evolutionary tree the phyla cnidaria and ctenophore are with radial
symmetry. Adult echinoderms are radially symmetrical but their larvae are
bilaterally symmetrical, so they are in bilateria.
 From radiate phyla we move to bilateria where Platyhelminthes are the lowest
phylum which are also acoelomates and nemertea (ribbon - like).
 The rest phyla have coelom or body cavity. The body cavities are of two types:
a) Coelom.
b) Pseudocoelom.
 Coelomates including some major phyla as, Annelida, Mollusca, Echinodermata,
Arthropoda and Chordate.
 Pseudocoelomate groups comprises 7 phyla as a group are called Aschelminthes.
They are tube shaped worms.
 The coelomate phyla are divided into two groups: protostomes and
deuterostomes.

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 Protostomes include the animals in which the blastopore forms the mouth of the
adult such as bryozoans, molluscs, annelids and arthropods. They are a major
group with complex anatomy.
 Deuterostomes include the animals in which the blastopore does not form the
mouth such as Chaetognatha, Brachiopoda, Echinodermata and Chordata.
 Classification of coelomates into these two groups is based on the embryology.
The 2nd, 3rd and 4th cleavage may be spiral or radial.
 Cleavage is also being either determinant or indeterminant. Determinant
cleavage. All cells are capable of producing an adult form.
 Indeterminant cleavage. Each cell of the embryo is required to produce some
parts of the adult.
In protostomes the coelom called Schizacoal forms from split in the
mesoderm. In deuterastomes it is called enterocoel forms from gut cavity pouches
that enters the mesoderm.

General Characters of Metazoa:

1. With large size which required, high - food materials and specialized digestive
system. Blood vascular system has been developed in most metazoans for
transportation of nutrients and respiratory gases.
2. Small forms which are aquatic respire and excrete by simple diffusion, while
large forms have branching and laminated respiratory and excretory organs.
3. They have a well-developed nervous system and various kinds of external and
internal supporting structures to their bodies or skeletal system which attain
them to live in a wider variety of habitats.
4. They have reproductive organs or gonads, and all of them develop through
similar steps of embryonic development (cleavage, blastulation, gastrulation
and organogeny).

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 Classification of Metazoa

Metazoa

Series: Eumetazoa Series: Parazoa

Phylum: Porifera

Division: Triploblastica Division: Diploblastica

(Bilateria) Phylum: Cnidaria
Phylum: Ctenophora

Acoelomata Pseudocoelomates coelomates

Phylum: Platyhelminthes Phylum: Nematoda
Phylum: Nemertea Phylum: Rotifera
(Aschelminthes)

Entrocoelomates Schizocoelomates
(Deuterostoma) (protostomes)
Phylum: Brachiopoda Phylum: Bryozoa
Phylum: Chaetognatha Phylum: Annelida
Phylum: Echinodermata Phylum: Arthropoda
Phylum: Ectoprocta Phylum: Mollusca
Phylum: Chordata
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 Subkingdom: Metazoa
Series: Parazoa
Phylum: Porifera
 Mostly marine but the only Family Spongillidae is freshwater.
 They are asymmetrical or radially symmetrical.
 No organization (without organs).
 Colonial and sessile animals.
 Permeated with pores, canals and chambers lined with choanocytes.
 Have a cavity called paragaster or spongocoel.
 The skeleton if present is composed of spongin fibres and spicules. The spicules
may be calcareous or siliceous.
 Their colonies vary in form where they may be cup-shaped, lobed, tubular, rod-
shaped, mushroom- shaped, digitate, globular, branched or irregular.
 Their color may be yellow, orange, purple, red, brown, blue or black.
 The colony size ranges from few millimeters to two meters or more in diameter.

Structure of Sponges
There are three main structural types:
1- Ascon type.
2- Sycon type.
3- Leucon type
1- Ascon (Asconoid) type: found in Ascetta & Leucosolenia
Represents the simplest type in which the body is in the form of a thin-walled
vase with a short stalk, and opening at the free end by a relatively wide aperture,
oscule or osculium, through which the water current passes out. The wall is
perforated by numerous incurrent pores or ostia. These lead from the external surface
to a flagellated central cavity, the paragaster, gastral cavity or spongocoel. Each

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ostium is actually an intracellular canal passing through a tubular cell; porocyte,
capable of dosing its pore in unfavourable conditions.

Water route: Ostia → paragaster → to the outside through the osculune

2- Sycon (Syconoid) type: e.g. in Sycon & Grantia.

The wall of the sponge becomes folded into radially arranged finger-like
outgrowths. The ostia lead to tubular incurrent canals. These canals open by
prosopyles into radial or flagellated canals which are lined by choanocytes. These
radial canals open to the paragaster by apopyles.

Water route:
Ostia incurrent canals prosopyles

osculium paragaster apopyles flagellated canals

(outside)

3- Leucon (Leuconoid) type: e.g. Euspongia (Bath sponge)

In the sponges of this type the flagellated canals of the sycon type are
replaced through a process of outfolding by groups of small flagellated chambers in
each fold lined by choanocytes. Each of these canals has apopyle and the fold has an
excurrent canal leading to the reduced paragaster.
Water route:
Ostia incurrent canals prosopyles radial chambers

osculium paragaster excurrent apopyles

(outside) canals

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Structure of the body wall:
The wall of the sponges consists of two layers:
1. Outer layer called dermal epithelium or pinacoderm.
2. Mesohyl or gelatinous protein matrix.
3. Inner layer called gastric layer or inner epithelium.

1. Pinacoderm: A single layer of thin flat cells; pinacocytes.

2. Mesohyl. This matrix in which embedded the spicules and the amoeboid
archaeocytes (cells) which are primarily phagocytic, but can differentiate into
more specialized cells, mainly sclerocytes or scleroblasts (skeletal secreting
cells).
3. Inner layer: Lining the paragaster, consisting of a single layer of
choanocytes or collared flagellate cells, in loose contact. Each flagellate cell is
composed of a body, collar and flagellum, where the collar is a cytoplasmic
elongation of the base of the flagellum containing microvilli and microfibrils
for engulfing the food.
Skeleton:
Most of the sponges have skeletal structures which are highly significant in
their identification and classification. The skeleton consists of spicules or spongin
fibers, or of a combination of both. Spicules possess an axis of organic material
around which is deposited calcium carbonate (calcareous) or silica (siliceous).
Spicules may be either large (megascleres) making up the primary framework of the
skeleton, or small (microscleres) protecting the outer surface and reinforcing canals
and flagellated chambers.

Types of spicules:
According to the number of axes (-axon) or rays (-actine) the megascleres are
classified into the flowing types:

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 1. Monaxonid (monaxonate or mono-radiate): Needle-like, smooth or
horny, termed diactinal when both ends are pointed, and monactinal if it is
pointed at one end and rounded at the other.
2. Triaxonid (triaxonate ): Formed of three axons:
a. Tri-radiate or triactinal: It is the most common form, with nearly equal
rays. Formed of three meeting axes.
b. Hexa-radiate or hexactinal: Formed of three axes crossing each other at
right angles, thus producing six rays.
3. Tetraxonid or tetraxonate (quadriradiate): May be tetractinal
consisting typically of four radiating rays, with one ray (rhabdome) longer than
the other small rays (cladome) at the top.
4. Polyaxonid or polyradiate: Formed of many axes meeting or crossing
each other resulting polyactinal spicules. It has different shapes.
Microscleres may be:
1. Amphidiscs or birotulates: Which are monaxonid with an umbrella-like cap of
spines at both ends of the horny shaft.
2. Chelas.
3. Asters:
i. Spherasters: with radii meeting in a well-defined center.
ii. Oxyasters: with pointed radili
iii. Diplasters:
iv. Sterrasters which are disc-shaped with no projecting rays.
v. Sirasters
4. Sigmas
5. Toxas.
6. Spheres.
7. Raphides or desmas.

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Formation of monaxonate spicule:
1. It is secreted within a binucleated sclerablast.
2. CaCO3 is deposited around an organic axial thread in the cytoplasm between
the two nuclei.
3. As the spicule lengthens, the two nuclei draw apart until the scleroblast divides
into two.
4. One cell, the founder is situated at the inner end.
5. The other, the thickener at the outer end of the spicule, since monaxon spicules
usually project from the body wall.
6. The spicule is laid down chiefly by the founder cell which moves slowly
inward, establishing the shape and length.
7. The thickener cell deposits additional layers of CaCO3, also moves inward
during this process.
8. When the spicule is completed, both cells wander from its inner end into
mesoglea, the founder first and the thickener late.

Formation of triaxon spicule:

1. This spicule is secreted by three scleroblasts which come together in triangle or
cross each other at night angles and divide into two, each into an inner founder
and an outer thickener.
2. Each pair secretes a minute spicule and these rays are early united into a small
triradiate or hexaradiate spicule.
3. Each ray is then completed in the same manner as monaxon spicule.
4. Later on, three or six rays in union together forming a triradiate or hexaradiate.

Formation or other spicules

In the formation of quadriradiate or tetraxon spicules, four scleroblasts are
meet together as the fourth is perpendicular to the other three ones which are in one
plane. The other types of połyaxon spicules are formed by equal numbers of
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scleroblasts i.e. the demand number of scleroblasts is equal to the number of axons of
each type of spicules.

Formation of spongin fibers:

The spongin skeleton is mostly in the form of fibers in a network or branching
manner. The scleroblasts (sclerosponges) are arranged in a network and each cell
secrete a fiber unite with those of the others forming the network of spongin fibers of
the colony. It was clear that all types of skeleton are secreted by sclerocytes which
are either calcocytes, silicocytes or spongocytes.

Nutrition in sponges
The movement of the flagella of the choanocytes sets up a water anent to flow
into the body through the ostia carrying food particles (bacteria, flagellates and
organic matters). The food particles are entangled and attached to the collars of
choanocytes. Only the very fine particles are thus allowed to pass to the base of the
flagellum where ingestion occurs. Archaeocytes (amoebocytes) also ingest food and
are most probably the site of digestion.
Thus the digestion in the sponges is intracellular. The amoebocytes also
distribute the products of digestion throughout the body and store the excess part.
The undigested and large food particles are collect within the collar to be
rejected with the excurrent water.

Respiration & excretion:

Oxygen is taken from the incurrent water and the excurrent water carries the
carbon dioxide produced in respiration. Excretion takes place through simple
diffusion into the adjacent water. In freshwater sponges some of amoebocytes assist
in the collection of wastes and their elimination.

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Reproduction
It may be asexually or sexually:
A) Asexual reproduction:
-Budding:
i- External budding: in which external winds are formed and grow to
new individuals.
ii- Internal budding: In the freshwater forms of the family Spongillidae
during unfavorable conditions they form internal buds or gemmules.
Gemmule is formed of a pear-shaped or spherical group of amoebocytes
with the central cells filled with food reserve of glycoprotein and
lipoprotein. The peripheral cells are arranged in a columnar layer which
secretes an inner hard cuticular membrane and an outer thin one.
Amphidiscs spicules become then disposed radially between the two
membranes. The columnar cells eventually depart and the gemmule is left as
a small ball pierced by a micropyle. The hard construction of the gemmule
affords resistance to freezing and drying, and thus the species can pass the
adverse conditions. After the decay of the parent, the gemmules are set free
and when the conditions become favorable their contents pass through the
micropyle developing then to the adult form.
iii. Regeneration: Any cut piece of a living sponge is capable of giving
rise to a complete sponge, although the process is very show and may take
years. In this respect, while sponges do represent a favorite food for many
marine animals, the regeneration of those sponge bits, which occasionally
fall down to the sea bottom during predation, acts as an efficient natural
adaptation for species conservation. Experimentally dissociated sponge cells
are capable of reassembling and developing to new Individual, a process
known as somatic embryogenesis.

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B. Sexual reproduction: The majority of sponges are hermaphrodites, although
some are dioeceous. The amoebocytes produce the gametes (sperms and ova).
Sperms come to the colony with the incurrent water and enter the choanocytes. These
cells act as a carrier and transport the sperm to the mature ovum neighboring it. Then
fertilization occurs i.e. internal. The fertilized egg undergoes cleavage forming a
blastula.
The blastula composed of micromeres and macromeres surrounding a
blastocoel. Then the micromeres acquire flagella on their internal ends, and a mouth-
like opening makes its appearance between the micromeres, connecting the blastocoel
with the exterior.
A process of inversion then takes place by which the entire embryo turns
inside out through the mouth, and the flagella become external. The embryo in this
stage is known as the amphiblastula or parenchymula class Calcarea and other classes
of sponges respectively. This larva leaves the parent tissue with the out-following
current of water.
After a short free swimming period, the larva undergoes gastrulation by
epiboly or invagination or by both in contrast with all other metazoans the
macromeres overgrow, the micromeres. The larva then settles down to the bottom
and attach to a substratum growing to the new colony.

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Classification of sponges

Phylum: Porifera
Class calcarea (Calcispongiae)
Subclass: Hetrocoela
Subdass: Meternodelo
Class: Scleropongiae
Class: Hyalospongiae (Hexactinallida) Euplectella
Class: Demospongia (venus flower- basket)
Subclass: Homoscleromorpha (Myxospongia)
e g Oscarella & Plakina
Subclass: Tetractinomorpha (tetractinellida)
eg placospongia, Geodia, Tethya & Cliona
(Coral-boring sponge)
Subclass: Ceractinomorpha (monaxonida)
e.g Ephydatia, Halidona & Spongilla
(Freshwater sponge)
Subclass: Keratosa
e.g Euspongia (bath sponge)

Phylum: Cnidaria (Coelenterata)

 These are all aquatic, mostly marine animals. They are considered to be the most
primitive eumetazoans.
 They are either fixed (sessile) or free swimming, radially symmetrical. The body
wall is composed of an external epidermis and an internal gastrodermis
(diploblastica) and in between them there is mesogloea or mesohyl. The
mesogloea, except in the Class Hydrozoa, contains cellular elements.
 They have a single body cavity called "coelenteron" or "gastrovascular cavity
with a main opening, the mouth, for both ingestion and egestion.
 Digestion is both intracellular and extracellular. No special respiratory or
excretory units exist, while respiration and excretion take place through the body
surface.
 Nervous system is not centralized, being in the form of a network of nerve cells,
with ocelli and statcysts as sensory receptors.

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 Reproduction is sexual and asexual, where the former involves a larval stage
called the planula, which possesses a ciliated ectoderm and a solid core of
endoderm.
 Cnidarians appear in two forms:
1. The polyp which is attached erect stage & may be solitary or colonial.
2. The medusa, which is pelagic, solitary, discoid or bell- shaped, with no skeletal
support and often with very thick mesogloea.
When both of the polyp and the medusa occur in the life history of a species,
the polyp reproduces asexually, budding off medusa, and these sexually give rise to
polyps expressing the phenomenon of metagenesis or alteration of generation
(alternation of the asexual polypoid generation with the sexual medusoid generation).
The polyp may be suppressed in some groups such as in the Schyphozoa and
Cubozoa. Also, the medusa may be adsent as in the Anthozoa.
The most distinctive feature of all Cnidarians is their possession of stinging
cells of cnidocytes or cnidoblasts which function in defense and capture of prey. Each
cell composed of double-walled nematocyst filled with a toxic mixture of proteins
and phenols and from which extends a hollow eversible thread. The thread is coiled
in the rest state and is connected with its base to the operculated neck. The base of the
thread is armed with barbs and barbules. There is a rigid protoplasmic hair or cnidocil
projecting from the neck.
The cnidocil is a modified flagellum with a kinetosome at the base. When it is
irritated by physical or chemical stimuli, the thread is discharged powerfully and
turned inside out under hydrostatic pressure. It then penetrates the predator or prey
injecting the paralyzing toxin.
There are different types of cnidoblasts:
1. Penetrant type as the described before.
2. Volvent or desmoneme functions in only tangling the prey by wrapping around
its body (spirocysts, occurring principally in certain Anthozoa).
3. The glutinant type is sticky and is used in anchoring the animal.
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 Nematocysts may be used but once and released form the cnidocyte after
discharge. New cnidocytes are formed from nearby interstitial cells. The various
nematocysts in a Cnidarian are collectively known as a chidom, which is of
considerable taxonomic value.
The cnidarians have a body wall formed of epidermis or ectoderm and
gastrodermis or endoderm and a mesogloea in between:
1. Epidermis consists of musculo-epithelial cells, interstitial cells, cnidocytes
and sensory cells.
2. Gastrodermis. Consists of nutritive muscle cells (or musculo-nitive cells),
gland cells, interstitial cells, sensory cells and nerve cells.
3. Mesogloea: Non-cellular in all hydrozoars, but in Schyphomedusae it is
enlarged as to constitute most of the size of the body of the animal and
contains numerous cellular structures.

Most of Cnidaria are with generally attached polyps but in some forms as in
Hydra the polyps are freely move.
Cnidarians are mostly carnivorous and some are capable of catching and
devouring large preys as fishes, and other marine animals. The digestion is
extracellular and intracellular. The undigested food is ejected through the mouth on
contraction of the body.
Sensory cells and sense organs (ocelli, tentaculocysts, rhopalia, etc ...) are
responsible for receiving the different stimuli.
These animals reproduce asexually by budding or sexually where a planula
larva present. Many of them characterized by the alternation of generation as stated
before.

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Class: Hydrozoa:
Marine mostly, without gullet in polyps, mesogloea is non cellular.
Gastrodermis without nematocysts. Typically with free or sessile medusae. Medusa
with a velum, nerve ring and ectodermal gonads. With alternation of generation is the
typical type of life history.
Order Hydroid:
 Polyps are fixed and the sense organs are ocelli or statocysts.
 Mostly colonials.
 The colony is anchored to the substratum by hydrorhiza.
 Hydrorhiza is absent in the mobile forms as in Hydra.
 Upright polyps may be attached to the hydrorhiza directly (Hydractinia) or by
caulomes (Tubularia).
 Most of hydroids have hydrorhiza giving stalk or stem of hydrocaulus.
 The stem may has branches bearing the polyps.
 The hydrocaulus may be arborescent e.g. Eudendrium & Obelia or pinnate as
Pinnaria and Plumularia.
 Polyps may be scattered on the branches as in pinnate forms.
 Polyps may be staked as in Obelia or sessile as in Plumularia.
 The coenosárc is covered by perisarc which is absent in Hydra.

A) Suborder: Anthomedusae (Gymnoblastea or Athecata)

 The perisarc covered the hydrocaulus and the stalk of the polyps only.
Colonials and fixed mostly (Bougainvillia) or non-colonial and free (Hydra).
 Colonial forms are hydrarhizal (Hydractinia & Clava).
 Monopodials with terminal polyps (Pirnaria & Eudendrium).
 Most colonies are dimorphic with nutritive polyp (feeding, hydranth,
gastrozooid or trophazooid) and reproductive polyp (gonophore) or defensive
polyp (dactylazooid or machozooid).

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 Hydranth formed of mouth on manubrium or oral cone or hypostome encircled
by a number of tentacles.
 The tentades have different forms and dispositions:
a. They may be hollow or solid (Hydra and Bougainvillia respectively).
b. They may be irregularly scattered (Clava) or arranged in one circlet
(Bougainvillia) or in two circlets (Tubularia & Coryamarpha) as oral and
aboral tentades.
c. The tentades may be filiform (Hydra) or short capitate (with terminal knob
of nematocysts) (Syncoryne).
d. The later two types of tentades are found in Annaria with distal capitate and
proudmal filtform tentades.

 The genophores in hydspids may be budded directly from the hydrocaulus or

from the gastrozooid stalk (Bougainvillia & Clava) or from the gastrozoold
body (Syncoryne & Pmania). They may be arise from modified polyp called
gonozooid or blastostyle as in Eudendrium.
 The blastostyle not have a mouth or tentades and bears the gonophores upon its
sides.
 In Tubularia and related forms the numerous gonophores are borne on long
branching stems just above the aboral tentacles.
 In the majority of anthomedusae the gonophores do not develop into free
medusae (eg. Bougainvillia) bit remain attached as Sensile reduced medusoid
structures or sporosacs as in Eudendrium & Tubularia .
 Medusae are conical, bell - shaped or hemispherical with abular manubrium in
the subumbrella, velum, four redial canals, ring canal and oral and marginal
tentacles.

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  Dactylosoold is a hydranth also but modified to catch prey and generally
defends the colony. Without a mouth and may have no tentades or these may
be reduced to mere knobs (Syncoryne & Podocoryne).
 Reproduction in Athecata is asexually and sexually:
i. Asexual reproduction takes place in the polypoid generation by budding.
ii. Sexual reproduction take place where the modusa gives the gametes (eggs
ar sperms) where the sexes are separate. Fertilization is elther external in
the sea or Internal as the eggs starting development as in Tubularla. The
dilated planula larva results and swim in water as a planktonic stage for a
definite period of free swimming. Then the larva settles down to form a
new polyp.
Examples of Athecata:
Hydra: A cosmopolitan freshwater, solitary, without perisarc at all and with only the
polyp phase present.
Bougainvillea: Polyps of the colony with a single dretet of illform tentacles,
around a conical manubrium. Blastostyles atcent, and medusae tkė their origin
directly from the coenosarc.
Clava: Colonial, polyps similar to those of Bougainvillea but with scattered filiform
tentacles.
Podocoryne: Colonial, polyps are polymorphic: gastrozooids, dactyiozoolds and
blastostyles. The gastrozooids with scattered filiform tentacles. Medusa with gonads
surrounding manubrium, four perradial tentade bulbs at the base of the hollow
marginal tentacles.
Syncoryns: Colonial and polymorphic as in podocoryne exactly, also their medusae
are similar. The only difference between Syncoryne and podocoryne is that the
gastrozoolds of syncoryne have capitate tentacles.

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Tubularia: Colonial, the polyps with oral (distal) and aboral (proximal) drclets of
filform tentacles, and with grape-like bunches of permanently attached sessile
gonophores.
Pinnaria: Colonial, the polyps with distal capitate tentacles and a drdet of prosdmal
filform tentades. Medusae rather elongate with vestigial tentacles.
Eudendrium: Colonial, loosely racemose, with male and female gonophores as it is
monogeneric. The hydranths have globular manubria and a single cirdet of filiform
tentades.

Suborder: Leptomedusae (Calyptobiastea or Thecita)

 Perisarc continued to enclose the polyps forming hydrothecae into which the
nutritive polyps can retrack and gonothecae around the reproductive polyps.
 They are colonial with stalked hydranths and sympodial growth or with sessile
thecae and monopodial branching with terminal growing points (Aglaophenia
& Plumisaria) or associated with dichotomous branching (Sertularia &
Sertestarelia).
 Hydranths are elongate with prominent manubrium and only a single circiet of
solid fliform lentades. The hydrothecae may be bell - shaped and open or its
opening may be covered by a fid, operculum , with one to several pieces
(Sertularia). In many forms a crauias chionous shelf projects inwards from the
hydrotheca at the base of hydranth making as a daphragm preventing the
passage of large food partides into the stalk (Obellia).
 The dactylozoolds of Plumulariidae also named nematophores or Sarcostyles.
Each three nemato or anothecae are located on the stems and on the
hydrothecae of the gastrozoolds. They are tentacle-like, with a solid
gastrodermal core and are often provided with one or more cap-like processes
with nematocysts or adhesive cells.

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  Gonophores are mostly located on a typical blastostyle enclosed in an envelope
called gonotheca opens on the top by gonopore, all forming what is known as a
goangium. Goangia arise directly from the angles between branches:
(Plumularia) or between hurdranth stem and branches (Obelia & Sertularia)
or on hydrocaulus or branches (Campanularia). There are leaf-like outgrowths
arch over the goangia forming a closed basket or corbula in Aglaophenia.
 The majority of Leptomedusae lack free medusae (Plurnularia & Sertularia).
The medusae are generally much flatter and less rigid than in the
anthomedusae. The mouth is commonly four labed. The marginal tentacles are
hollow, numerous and rarely solid (Obelia).
 Sexes are separate and the gonads lie connected to the subumbrellar wall of the
perradial canals which are generally four. The development of the zygote is
nearly the same as in the Anthomedusae.

Example of Thecata:
Campanularia & Obelia: Hydranths of the colony with small globular
manubrium; hydrothecae provided with a characteristic lid.
Aglaophenia & Plumularis: Pinnate colonies with monopodial growth and
terminal growing points, with sessile hydrothecae and nematophores. No free
medusae.
Sertularia: Stiff feathery type of growth i.e. dichotomous branching with terminal
growing point, cirved tubular hydrothecae with a lid of one to four pieces. The
hudranths arrange as each tun members of a pair are exactly opposite (in the same
level).
Sertularella: Similar to Sertularia but the polyps are alternate on both sides of the
branches.

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 Order: Milleporina (Hydrocorallina)

 Fire corals. Comprises a single genus Millepora which is mostly associated

with coral reefs.
 The colony is fixed and consists mainly of a much branching and anastomosing
hydrorhiza.
 The perisarc is secreted in the form of a massive calcareous exoskelecon
coenosteum.
 The perisarc is irregularly lobed or branched in one plane with a minutely
pitted appearance but is covered wit a thin layer of living tissue.
 Polyps which arise from the pits in the surface, Into which they can retract, are
of two types: gastrozoolds and dactylozooids (Dimorphic colony).
 Gastrozooids, each with four short capitate tentades.
 Dactylozooids, are long defensive and more numerous without a moith but
with irregularly disposed capitate tentacles.
 The gastrópores are larger than, and irregularly surrounded by the
dactylopores.
 Medusae are budded directly from the coenosarc and lodged in pits in the
skeleton called "ampullae". They lacks a velum, mouth, tentacles and digestive
cavity. Sex cells develop on the manubrium.

Order: Siphonophora:
 Exclusively marine, pelagic and are polymorphic colonies.
 Not only more than one kind of polyps are possessed but also more than one
kind of medusae. The medusae with various functions not including gamete
formation.
 Siphonophores composed of different types of polyps they are:

21
 1. The float or pneumatophore: is a modified medusa capable of eting
gases thus keeping the animal floating on the water surface.
2. Nectocalyx: medusoid, it is vertically present when the development of the
exumbrellar side is vertically and found after the float. It resembles an
Anthomedusae but without mouth or manubrium and is specialized for
propelling the colony. It is atso termed nectophore or swimming bell. e.g.
Muggiaea. The other members are develop in groups termed cormidia.
3. Bract or hydrophyllium: is leaf - like , thick and gelatinous with à
gastrovascular cavity and protect the rest of the cormidium.
4. Gastrozooid or siphon: resembling the manubrium of a medusa with
mouth, a long hollow bentade arising near the base and bears lateral contractile
branches (tentilla) with dense nematorusts, it is the feeding polyp of the
colony.
5. Dactyizooid or palpon: without mouth but with unbranched tentacle , with
a defensive function or capturing prey.
6. Gonozooid: for reproduction, without tentacle and typically forms a
branched stalk (gonodendron) beating grape-like dusters of gonophores that are
not set free. e.g. Muggiaea without float but instead it has swimming bells.
Whe: the exumbrellar side of the coenosarc cievelops horizontally or it
becomes reduced to a horizontal flat piate. The examples of this type are
Physalia, Velella and Parpita.

Physalia (Portugese man- of- war): With flattened float and it becomes in the
form of a cup-shaped blue or orange colored bladder floating on the water surface.
From the underside hang down large numbers of gastrozoolds, dactylozooids and
gonozooids. Also, there are tangles of long retracte tentades which reach a length of
20 m, and armed with batteries of powerful nematocysts. It is the only unisexual
colony in Siphonophora.
21
Velella: With a rounded or oval fiat disc or float. From its underside hangs a large
central gastrozooid, encircled: by: a large number of gonozooids, followed by a series
of dactylozoolds. Medusae arise as bids from the gonozooids.
Parpita: similar to Velella but its float without a sail.
They float with large crest or sall in Phrysalia, which is small in Velelta, while this
crest is absent in Porpita.

Class: Scyphozoa (Scyphomedusae)

 Jellyfish, marine and the medusoid stage predominates.
 The medusa is without velum; tetramerous and bears endodermal filaments and
gonads: in the gastric cavitys on the septa or in the floor of the gastric pouches.
 The gastric cavity is divided in the adult or larva: by four interraria septa into a
central: stomach and four: perradial gastric pouches.
 The life cycle involves a modified alternation of generation of which the polyp
is inconspicuous as it is represented by scyphistoma (polypoid larva).
 The scyphistoma is either metamorphoses directly into the adult or gives off
inval medusae by transverse fission..
 Without skeleton.
 Some are luminescent.
 They are free-swimmers but may be sessile (Stauromedusae).
 Similar to Hydromedusae but are larger.
 The margin is usually scalloped into lappets and is provided with tentacles..
The tentacles are long or short.
 Rhizostomeae are without tentacles.
 Tentacles are borne on gelatinous expansions called pedalla as in coronatae.
 The tentacles are capitate in the Stauromedusae with heads filled with
nematocysts.
 The tentacles are filiform in other orders.
22
 In the Subumbrellar surface the manubrium terminates by a four - cornered
mouth whose angles are drawn out into four lobes.
 In the semaeostomeae the oral lobes are extend into oral arms with exhalent
grooves.
 In tire Rhizostomeae the oral ams are fused at their bases resulting thousands
of minute suctorial mouths.
 The medusa possesses a system of radial canals much more extensive than in
Hydromedusae. The radial canals are of three types, interradials, perradials and
adradials.
 The interradials are four branched and connected to the gastric pouch.
 The perradials are four branched and connected to the stomach.
 The adradials are eight unbranched in between.
 The mesogloea is thick, gelatinous and fibrous containing amoeboid cells and
is a true cellular layer (ectomesodermal in origin).
 The marginal sensory bodies (tentaculocysts) consists of rhopaloid in the
Stauromedusae or rhopalia in other orders.
 Rhopalia are borne on pedalia in most coronatae and inbetween the marginal
lappets in others.
 Each rhopalium consists of a pigmented ocellus sensitive to light, a hollow sac
(statocyst) containing hard particles & two sensory pits with cells that are
thought to be sensitive to food or other chemicals in water.
 The Schyphomedusae are dioecious and the gonads are found in the
gastrodermis on both sides of gastric septa (8 in number).
 In the Semasostomeae and Rhizostomese where the septa are absent, there are
four brightly coloured horse-shoe-shaped gonads lying in the floor of the
gastric pouches.

23
 Life Cycle In Scyphomedusae

Medusa Fertilization Zygote

on maturation male Sperms
and female + Cleavage
ephyra larva ova
transverse fission or strobilation

Coeloblastula
new scyphistoma

in summer asexuaily
by budding planula
(Scyphoistoma larva
or
Scyphopolyo) Polypoid
larva after a period of freeswimming
settles down

Notice: Strobilation is termed monodisc when one ephyra is formed at a time (as in
Cassiopeia), or polydisc when numerous ephyras are produced (as in Aurelia).

24
Classification of Scyphomedusae

Order: Stauromeichisae (Leucernariidae)

e.g. Halidystus & Leucemaria
Order: Coronate e.g. Nausithoe
Order: Semaeostomeae
eg Aurelia, Cyanea & Pelagia
Order: Rheizostomene
e.g. Rhizostama & Cassiopeia

Class: Anthozoa (Actinozoa)

 Marine, solitary or colonial.

 Without medusa.
 With endodermal gonads and a cellular mesogloea.
 Without oral cone but with a stomodaeum or gullet.
 Stomodaeum is attached to the body wall by mesenteriss.
 The mesenteries divide the coelenteron into chambers.
 The stomodaeum has one or two siphonoglyphs (sulcus or sulculus).
 Mostly with internal or external skeleton.
 This class comprises about two-thirds of Cnidaria.
 This class is divided into two subclasses: Alcyonaria & Zoantharia.

25
 Subclass: Alcyonaria (Octocorallia)

 With superficially octamerous symmetry.

 Usually forming lobed or branched colonies.
 Each polyp has eight hollow pinnate tentacles around an oral disc perforated by
the mouth.
 There is a stomodaeum with one ventral siphonoglyph (sulcus).
 There are 8 mesenteries bearing retractor muscles on their sulcal sides.
 The stomodaeum is lined with ectoderm.
 Below the pharynx or stomodaeum the inner edges of mesenteries are
thickened forming mesenteric filaments which are free but only the two asulcal
septa reach the base of the polyp. The asulcal filaments are heavily flagellated.
 The asulcal or directive filaments create an upward current contrary to the role
of the sulcus which beat downward by its cilia to the gastrovascular cavity.
 Fertilization is external.
 The orders are Stolonifera, Alcyonacea , Gorgonacea & Pennatulacea.

1- Order: Stolonifera Cornularia, clavularia & Tubipora musica, (organ pipe-

like coral)
The polyps arising from a creeping base or stolon in the form of a sheet or
meshwork.
In Cornularia and Clavularia the first formed polyp gives rise to a creeping
stolon with a single gastrodermal tube or solenia from which new polyps are given
off at intervals bý budding. Stolons are further produced from the newly formed
polyps, budding again in the same manner, and so on.
Comularia lacks calcareous spicules but has homy investment on the stolon
and polyp bases into which the distal portion of the polyp can retract. In Clavularia
the bases of polyps have numerous warty elongated spicules.

26
 In Tubipora musica (organ-pipe-coral). The distal ends of the polyps newly
produced grow out in the form of horizontal expansions, all unite together into a
common platform from the external surface of which new buds arise. These grow
into another series parallel polyps which in their turn repeat the process. Skeleton is
in the form of spicules deposited in the mesogloa, but are fused together so as to form
a continuous tube for each polyp. The dull or deep red colour of the skeleton results
from the iron salts.

2- order: Alcyonaces
Soft colonies, polyps are embedded in a gelatinous mass of coenenchyme from
which only the oral part of the polyps (anthocodia) are protruded, e.g. Xenia,
Heteroxenia, Alcyonium , Sarcophyton & Lobophytum.
The colony may be massive as Xenia, Heteroxenia or may be mushroom-
shaped as Sarcophyton or may be branched into blunt lobes as Alcyonlum.
Usually the proximal part of the colony is sterile i.e. devoid of polyps. The
anthocodia may be scattered all over the distal region as Alcyonium or in clusters on
the ends the lobes as Xenia or on the top of the mushroom-like colony as Sarcophyton
and lobophytum.
Polyps are retractile as Alcyonium or non-retractile as Xenia. They are
monomorphic as Alcyonium or dimorphic as Sarcophyton and Heteroxenia. The
autozooids and the siphonozooids are found in the dimorphic colonies.

3- Order: Gorgonacea
 Horny corals, sea fans and the polyps arising as lateral outgrowth from the
coenosarc as a tree-like colony.
 The colony branches and anastomoses in one plane like a fan with a basal trunk
devoid of polyps.
 The skeleton is in the form of an axial rod extending all through the colony and
in case of Gorgonia is horny in nature containing gorgonin (protein and
mucopolysaccharides) or partly calcareous and partly horny in some species.
27
  Corallium rubrum is a precious red coral which is exceptionally dimorphic, the
skeleton is entirely calcareous formed of solid central red axis of spicules
cemented together by calcium carbonate.

4- Order: Pennatulacea
 Sea pens, Fleshy dimorphic colonies having an elongated proximal stalk for
anchoring the clony in soft substrata and a distal rachis typically with lateral
branches bearing only autozooids.
 The back of the rachis in Pennatula carries siphonozooids which function in
maintaining the water circulation in the colony.
 The stem is supplied with a horny or calcareous axis which does not extend in
the branches. Spicules also occur in the mesogloea.

Subclass: Zoantharia (Hexacorallia)

 Soilery or cooral and all are marine.
 The polyps have unbranched smooth tentacles which are arranged in one or
several circlets varying from a few (only 16 in Edwardsia) to a hundred or
even thousand or more in number.
 The stomodaeum has two siphonoglyphs (dorsal sulculus and a ventral sulcus)
 Septa or mesenteries are ususally in pairs. The space between members of a
pair is termed endocoel and the other between pairs is termed exocoel.
 Retractors of paired septa usually face each other in the endocoel except for the
two pairs (directives) connected to the siphonoglyphs, where the muscles are
apart facing the exocoels.
 Some pairs of septa extend from the body wall to the stomodaeum termed
perfect, primary, complete or macrosepta. Others do not reach the stomodaeum
termed, imperfect, secondary or tertiary or quaternary, incomplete or
microsepta.

28
  Commonly septal pairs occur in cycles of six or some multiple of six. Thus
there are six primary pairs of macrosepta. In the six exocoels there are the other
types of microsepta.
 Some or all septa bear filaments which are convoluted and trilobed in cross
section. The middle ridge known as the cnidoglandular band, consists of
digestive gland cells and nematocysts (homologous with six of the alcyonarian
filaments). The lateral wings known as the flagellated bands as they are heavily
flagellated (corresponding to the asulcal alcyonarian filament).
 Each filament thus performs the functions of digestion and water circulation.
 The lower part of each septal filament is exclusively digestive in function, the
wing bands being phagocytic.
 Gonads are brne on all or certain septa adjacent to the septal filaments.
 A skeleton is present in some orders but never in the form of mesogloeal
spicules. It is represented wither by an external calcareous mass
(Madreporaria) or horny axis (Antipatharia).
 The orders are Actiniaria, Madreporaria, Antipatharia and Ceriantharia
(Cerianthidea).

1- Order: Actiniaria
Sea anemones, usually solitary with a cylindrical fleshy body divided into oral disc
with a silt-like mouth, column and base. They may be hermaphrodites as the sperms
produce firstly.
Without hard skeleton, mostly littoral or coral reef dwellers, but some are deep sea
inhabitants, sand burrowers or even pelagic. e.g. Actinia, Edwardsia, Adamsia &
Sagartia.
2- Order. Madreporaila (Scleractinia) (Stony Corals) e.g. Fungia fungites,
Fungia echinata, Acropora, Stylophora, Galaxia, Favia, Porites, ceriatopora,

29
Echinopora, coeloria, Lobophyllia, Lophorylia, Montipora, Oculina, Orbicella,
Goniopora, Montipora & Cyphastraea.

 True or stony corals are the largest zoantharian order.

 The main builders of coral reefs and islands.
 Mostly colonial although some are solitary.
 The most characteristic feature is the possession of a hard caleareous
exoskeleton secreted by the epidermis of the basal disc.
 The skeleton of the colony as a whole is termed corallum, that of each polyp is
termed corallite.
 The corallite consists of a cup containing vertical radiating ridges called
sclerosepta (macro - and micro-). The bottom of the cup which is the first to be
secreted is the basal plate. The wall of the cup that encloses the aboral portion
of the polyp called "theca". The inner edges of the septa are fused in the center
forming a columella.
 Extending from the base of the polyp, especially in colonial forms, there is an
extrathecal portion which will give rise to buds for reproduction.
 The rate of calcium deposition is not the same throughout the year, but
influenced by various ecological conditions.

Examples of Stony Corals:

Fungia: Mushroom - or disc shaped.
Stylophora: Fine branching.
Acropora: Tree-like, extratentacular budding, with separate polyps.
Echinopora: Encrusting and leaf-like.
Seriatopora: Fine branching.
Goniopora: Columnar shape.
Montipora: Warty shape.
Galaxea: Upright discrete tubes or cups, extratentacular budding, with separate
polyps.
31
Cyphastraea: Spherical shape.
Favia: Spherical, the theca are close together as to have common walls and the
corallites too short.
Coeloria: Huge globular masses, budding is incomplete and the new polyps are found
in serpentine course with a big corallum.
Porites: Huge globular masses.
Oculina: With extratentacular budding, with separate polyps & with slender
branching.
Orbicella: With extratentacular budding & with separate polyps.
Lobophyllia: With intratentacular budding, group of buds arise on the oral disk and
grow equally leading to dichotomous branching colony.
Lophohelia: The same manner in Lobophyllia but one of the resultant polyps ceases
to grow for a time while the other is active, grows and undergoes budding and
repeated a lternating between the right & left.
3- Order: Antipatharia (Antipathidea) e.g. Antipathes
 Black or horny corals, typically deep sea animals.
 They form slender branching plant-like colonies with a skeleton in the form of
a brown or black spiny horny rod (similar to Gorgonia) secreted from a basal
invagination of the epidermis.
 Polyps are of short cylindrical form with six simple non retractile tentades.
 In Antipathes the polyps are present along branches.
 The name of the order is derived from the ancient belief that wearing pieces of
its skeleton helps keep the body healthy by providing protection against
diseases.
4. Order: Ceriantharia (Cerianthidea) eg. Cerianthus
 They are tube anemones (live in tubes), long, solitary without pedal disc and
with numerous simple slender tentacles.
 The tentacles are arranged in two whorls; short oral and long marginal.

31
  Cerianthus lives in vertical cylindrical cavities in the sea bottom inside a felt-
like tube of hardened slimy secretion in which are embedded specialized
cnidae, sand grains and other foreign objects.

Phylum: Ctenophora (Comb-Jellies)

Exclusively marine, free, solitary and pelagic or benthic. The body is
gelatinous, bilaterally symmetrical and ranges in shape from spherical to ribbon-like.
They are without any skeleton. Locomotion takes place by the beating of eight
rows of fused cilliated plates called combs possessed by all ctenophores at some
stages in their life cycle.
These animats have a pair of tentacles on which sticky colloblasts or Lasso
cells are present. These cells are used for capturing the prey. Each one of these
adhesive cells has a hemispherical, or pear-shaped, papillated head producing a sticky
secretion, by which the prey becomes entangled and is thus easily caught by the long
tentacles. The head is fastened to the tentacle core by contractile filament, acting as a
spring, which is coiled around a stiff straight axis.
Nematocysts are absent (except in Heckelia rubra, which has no colloblasts).
A cydippid larva forms during the development of all except the Beroida.
Ctenophores are carnivorous and all are intensely bioluminescente.
The structure of a Ctenophore can well be demonstrated in the genus
Pleurorachia. This is a common marine animal of a pyriform gelatinous body. It is
almost transparent and is only revealed in water by the iridescence of eight rows of
rapidly beating combs made of long cilia fused at the base.
The body has oral pole and aboral pole. In the oral pole the mouth is found and
on the other hand there is a tiny statocyst-tike sense organ in the aboral pole.
On the opposite sides of the body there are two long tentacles, each one has a
tentacular sheath. The tentacles with an axial core of mesogloea covered by
epidermis. They are highly extensile and bear a row of lateral branches. Their

32
epidermis consists largely of characteristic adhesive cells or colloblasts or Lasso
cells.
Enteric or Gastrovascular System
It is ramify through the thick gelatinous mass superficially homologous with
that of Scyphomedusae. The mouth aborally leads to stomodaeum or pharynx which
is followed by a short oesophagus. The later leads to a stomach (infundibulum or
funnel) from which arise the canals of the gastrovascular system.
From the aboral surface of the stomach arises an aboral canal (or infundibular
canal) to the underside of the sense organ where it gives off four so-called excretory
canals terminating in little sacs. Only two of these open on the surface by two
excretory pores. They are termed anal canals from which indigestible matters are
ejected.
From the oral surface of the stomach it gives a pair of blind pharyngeal or
paragastric canals along each side of the pharynx.
In the tentacular plane, a large transverse (perradial) canal arises from each
side of the stomach from which arising a tentacular canal and two interradial canals.
The tentacular canal into the tentacular sheath and the two interradials one in each
quadrant of the body.
Each of the interradials bifurcates into two adradial canals. Each adradial intern
joins a long and meridional canal underlying the length of each comb-row.

Body Layers
They are consist of epidermis and gastrodermis with a jelly mass inbetween.
The wall of the stomach and the gastrovascular canals constitute the gastrodermis.
The skin and stomadaeum are of ectodermal origin. In the gelatinous mass there is a
network of delicate smooth muscie fibres. Beneath the spidermits there is a
subaticular layer of muscles and neve Fiores similar to those of turbillarian
flatworms. These muscles are suitable to the locomotory organs; alliatalunb-rows.

33
Reproductive Organs and Development
They are hermaphrodites. The gonads are found in the meridional canals, the
ovary on one side and the testis on the other. The gametes are discharged through the
mouth.
Self-fertilization may occurs and takes place in water. Except in the genus
Gastrodes a planula larva is absent. There is a free swimming cydippid larva which
metamorphoses to the adult animal.

Classification of Ctenophora:
Class: Tentaculata (Micropharyngea) with tentacles, usually globular but some are
dorsoventrally flattened.
1. Order: Cydippida pelagic, globular or ovoid; tentacles retracile into sheaths.
Pleurobrachia & Lampea.
2. Order: Platctenida (Platyctenea) Creeping, flattened in the oral-aboral axis;
capable of sexual reproduction. Coeloplana, Ctenoplana and Castrodes.
3- Order: Cestida Pelagic ribbon-like, compressed in the tentacular plane and
elongated in the stomodaeal plane. e.g. Cestum, with venus girdle, the most common
of all Ctenophores, may be more than 2 m lang.

Class: Nuda (Macropharyngea) without tentades.

Order: Beroida more or less conical, flattened in the tentacular plane, with very
wide mouth and pharynx. The pharynx occupying most of the body. Meridional
canals without numerous side branches. e.g. Beroe

Phylum: Rotifera (Rotatoria)

Whneel animalcules, common freshwater, rarely marine or terrestrial,
unsegmented animals.
They are free swimming or sessile, solitary or colonials. The body is composed
of: 1. Corona or head. 2- Trunk. 3- Foot or tail.

34
 1. Corona: Cliated crown or trochal lobe. It may be composed of two to many
trochal discs or in the form of a simple circumapical band or may be vestigial. It
is used for swimming as well as for food collection. The beating corona gives
the appearance of a rotating wheel and brings about the rotation of the animal.
2. Trunk: Large, transparent and vermiform covered by a thickened article called
lorica.
3. Foot or Tail: The posterior part of the animal, provided with pedal glands
secreting an adhesive material with which the animal can attach to the substrate
while feeding. It ends usually with two to 4 spurs and one to four toes.

Rotifers either creep by the aid of the foot or swim by the beating cilia of
corona and many both creep and swim. Some are sessile forms.
The body is covered by a cutice which is much thickened on the trunk to form
a "lorica“. This is usually ornamented with ridges or spines and may be divided into
distinct plates. The autide is annulated on the foot.
The spacious pseudocoel contains a syncytial network of branching amoeboid
cells which are presumably phagocytic and excretory.

Digestive System
The mouth is surrounded by some part of the corona leads to a buccal tube or
directly to a muscular pharynx or mastax with a number of internal jaws (trophy).
The mastax is followed by oesophagus which joins the sac-like or tubular stomach,
the site of digestion and absorption, The stomach leads to a short intestine which is
sometimes called the cloaca, since it receives the excretory and female genital ducts.
The gut is vestigial or altogether absent in males.
Most rotifers are either suspension or raptorial feeders, feeding on algae,
protozoans, and other rotifers.

35
 In raptorial forms, the corona is modified as a trap for preys. In parasitic
members the corona is reduced and either the foot or mastax becomes modified as an
attachment organ.

Excretory System
Composed of a lateral pair of protonephridia which consist of flame cells of
bulbs join to collecting tubules. These empty into a contractile bladder, or directly
open into the cloaca.

Nervous System

This is consists of an anterodorsal ganglionic mass (cervical) or brain, lying

over the mastax and sending nerves to the anterior sense organs. Also, there are two
main nering to the muscles and other parts of the body.

Sense Organs
They are the dorsal and lateral antennae and a red eye spot or paired eye-spots
(as in Philodina).

Reproductive System
Males are always smaller than the females. They are present at certain times
and in some species they have never been reported.
In males there is a single testis (a pair in the Seisonidea) which leads to a
ciliated sperm duct, assodated with a pair of prostate glands and ending in a
copulatory organ.
In female the ovary and vitellarium unite to form a single syncytial body called
germovitellarium of which one or two may exist in Class Monogononta or in Class

36
Digononta, respectively. This leads to oviduct which opens by genital pore in the
cloaca.
Copulation
Takes place by hypodermic impregnation or through the cloaca.
Development
 Eggs are surrounded by a shell and a number of egg membranes. They are
attached to objects or to the female body, rarely brooded internally.
 In rotfers having no males (Class: Digononta) as in Philodina the
development is parthenogenetic and the diploid eggs hatch to females.
 In Class Seisonidea, females produce haploid eggs which are fertilized,
developing into either males or females.
 In Class Monogonanta, females however lay either of two types of summer
eggs, amictic and mictic eggs.
 Amictic eggs are thrin-shelled large, diploid and develop
parthengenetically, into amictic females.
 Mictic eggs are also thin-shelled but are small and haploid. These hatch
into males in case they are not fertilized. But in fertilized ones they
secrete heavy resistant shells and remain dominant for months until
hatching into females.

Class: Digononta Class: Monogononta Class: Seisonida Miemsi 26b) 29lsm on pnt
diploid haploid eggs (amictic) large (mictic) Ismst of ljeus Parthenogenitically
Fertilization and remain dormant\until giving Parthenogenetically develop into
Fertilization Males Males and Females Females Females (3 & 3) (6)G (
Classification: Class: Seisonidea (Seisonoidea) e.g , Seison parasitic on Nebalia
produce one kind of eggs. Class: Digononta. Order: Bdelloida e.g, Philodina, Rotaria

37
& Adineta. Benthic, freshwater, rarely marine, paired gonads, oviparous or
viviparous. 52

Class: Monogononta (Monogonontidea)

With single gonads, includes most of rotifers, without digestive tract. e.g.
Notommata, Brachionus, Pedalia, Collotheca, Macrochaetus, Wolga and
Stephanoceros.

Phylum: Annelida (Ring-worms)

 Annelids are mostly aquatic, marine or freshwater, burrowing or living in

tubes, some are free living forms.
 They are triploblastic, bilaterally symmetrical, elongated and vermiform. They
are metamerically segmented, externally by transverse grooves and internally
by septa into a number of divisions, each division is called as segment,
metamere or somite. The first segment forms the head where it has anterior
prolongation called prostomium and the rest of the segment called
peristomium.
 Outer covering of the body is cuticle secreted by the underlying epidermis.
Body wall is contractile, consists of an outer epidermis, circular and
longitudinal muscles (dermo-muscular).
 Appendages when present are unjointed. The locomotory organs are
segmentally arranged (appendages or chaetae or setae).
 The true coelom is divided by intersegmental septa into intersegmental
chambers or coelomic chambers.
 The alimentary canal is tube-like extending straight from mouth to anus.
 Respiration takes place through the body surface or by gills in some forms.

38
  The circulatory system is mostly of the closed type. The blood is red due to the
presence of hemoglobin.
 Excretion by paired segmental nephridia which communicate the coelom to the
exterior.
 The nervous system consists of a brain and segmental double ganglia
connected by a double ventral nerve cord. The brain is composed of
suprapharyngeal or supra- oesophageal ganglion and a ventral one connected
with each other by two circumpharyngeal or oesophageal commissures or
connectives.
 Sexes are separate in some forms and united (hermaphrodites) in the others.
The development is direct in the hermaphrodites and indirect in those of
separate sexes due to the presence of a larval stage called trochophore larva.

Phylum: Annelida
Class : Oligochaeta
Order: Neooligochaeta - Pheretima, Lumbricus & Allolobophora,
Order: Archioligochaeta - Tubifex & Aeolosoma Bob
Class: Hirudinea
Order: Acanthobdellida- Acanthobdella
Order: Rhynchobdellida Pontobdella, Piscicola, Glossiphonia & Branchellion
Order: Gnathobdellida (Arynchobdellida) Hirudo, Haemopis, Hirudinaria,
Herpobdella or Nepheles. shunibam
Class: Polychaeta
Order: Errantia - Nereis, Heteronereis, Aphrodite, Glycera, Aphrodite &
Lepidonotus Order: Sedentaria Chaetopterus, Terebella, Sabella & Arenicola.
1- Class: Oligochaeta
These are land and freshwater worms, with the coelom divided into coelomic
chambers by the intersegmental septa. They have a small number of chaetae but
without parapodia. With distinct prostomium bearing no appendages. They are
39
hermaphrodites and the gonads are few and fixed in number and position, with
nephridia for excretion. The reproduction is by copulation and cross fertilization.
Eggs are laid in cocoons and the development is direct without a trochophore larva.

2- Class: Hirudinea
Mostly freshwater forms, sometimes are marine or terrestrial; with fixed
number of segments broken up externally into annuli. Mostly without chaectae or
parapodia. Anteriorly and posteriorly several segments are modified into suckers,
with mouth lying ventrally at the anterior sucker. Anus usually located dorsally
immediately in front of the posterior sucker. Genital pores are median, unpaired and
of a fixed position. Coelomic cavity is reduced being invaded by mesenchyme and
usually broken up into a number of intercommunicating sinuses. All are
hermaphrodites (monoecious). Reproduction is by copulation and fertilization. The
development is direct i.e. without passing through a trochophore larva.

3- Class: Polychaeta
Polychaetes are marine and carnivorous, elongated and segmented where the
head is distinct carrying prostomial and peristomial tentacles, palps and eyes on the
prostomium. Without clitellum. Each segment carrying a pair of parapodia on which
are located numerous setae.
The alimentary canal is provided with an eversible buccal region and
protrusible pharynx. Excretory organs are segmentally paired trochophore larva, is
present. Some forms of these animals reproduce asexually by lateral budding.

Order: Errantia
They are free swimming, often pelagic with well-developed parapodia, while
some are living in tubes. All body segments are similar except the anterior and
posterior ones. Pharynx is usually protrusible and armed with chitinous jaws and
teeth.
41
a) Family: Nereidae → Nereis & Heteronereis: They are many segmented,
elongated, with distinct head, each body segment carries a pair of parapodia. In
Nereis all body segments are similar, while in Heteronereis the body is
differentiated into two regions, atoke and epitoke. Atoke is the anterior portion
having ordinary parapodia and the epitoke is on the posterior portion carrying
modified parapodia. They are carnivorous where the pharynx is eversible
forming a proboscis.
b) Family: Aphroditidae The body is usually compressed having interlocking
pairs of hard scales covering at least a part of the body. These scales are readily
thrown off when the worm is handled. e.g. Aphrodite & Lepidonotus.
1. Aphrodite sp. (Sea-mouse): About 10 cm long, with 39-49-50 pairs of
almost circular scales on the back which are covered by dense short grey or
brown setae. The chaetae leave a space in between collecting the water for
respiration.
2. Lepidonotus sp.: Up to 3 cm long, with 26 body segments, alternating cirri
and 12 pairs of oval overlapping dorsal scales which are not round.
c) Family: Syllidae: e.g Syllis: Small delicate worms from 1-5 cm long. The
head region carries 4 eyes, 2 antennae and 2 pairs of peristomial tentacles.
Body segments bear long setae and antennae or tentacles which are ringed. The
proboscis is characterized by the presence of one black jaw on its top. Budding
is common where some forms can produce heads from almost any segments.
This genus can produce a branching mass of individuals.
d) Family: Glyceridae e.g. Glycera: Worms about 4 cm long, thin pointed at
both ends. The body is elongated and cylindrical consisting of about 150
segments. The prostomium is long and conical bearing 4 small tentacles are
arranged in crosswise at the tip. The proboscis is very large and protrusible, it
is flat at its top carrying 4 small sharp jaws which are black in colour and also
arranged crosswise. Behind the head each of the first 2 segments carries one

41
 tuft of setae, while the other segments bear one tuft on each side of them (2
tufts each). When touched this worm often roll up into coils.

Order: Sedentaria, they are burrowing or tube dwelling forms. They build their
tubes from gathered materials and they may leave them from time to time and build
another tube or may never leave them. Body is divisable into two or more regions
with unlike segments and reduced parapodia. Sexes are separate or may be
hermophrodites. Development is direct when sexes are united and indirect when
sexes are separate.
a) Family: Chaetopteridae e.g Chaetopterus sp. (Parchment worm): It lives in
parchment like U-shaped tubes open at both ends embedded in the mud
encrusted with sand and debris. The body is divided into three distinct regions;
anterior, middle and posterior.
1. The anterior region: is flat and composed of 9 segments where the front
nine segments carrying nine pairs of simple short parapodia formed only of
notopodia. The prostomium is small and the peristomium is a funnel-
shaped peristomial callar with a pair of peristomial cirri.
2. The middle region: formed of five segments, the first anterior most (10th
segment) is produced laterally into great wings directed forwards. The next
segment carries a pair of suckers and the rest three segments carry
membranous folds so-called fans formed by the fusion of the notopodia.
The fan make to draw in water for respiration and feeding.
3. The posterior region: it comprises thirty similar segments which are
devoid of setae, the food of Chaetopterus involves mainly small organisms
which are carried in by the currents of water set up by fans, Chaetopterus
is highly phosphorescent emits blue green light.
b) Family: Terebellidae eg, Terebella, Tubicolous burrowing long worm lives in a
tube of mucous and fine materials. The prostomium is horse-shoe shaped
provided with numerous long tentades which serve as gills. Peristomium without

42
 cirri. Just below the head are found two pairs of branched gills. Parapodia are
reduced.
c) Family: Sabellidae eg, Sabella, Dasychone & Bispira: worms with
semicircular head carrying tentacles which appear only when the animal is
covered with water. They form temporary muddy tubes and the worm can form
more than one tube during its life. Parapodia are rudimentary.
 Sabella pavonina: It is 25-30 cm long, body is cylindrical of orange-brown
colour. It lives in a long membranous tube in the sea mud. The body is
divisible into a head, thorax and abdomen.
1. Head: 4-segmented, in front of which are ten pairs of peristomial tentacles
forming ciliated branchiae. It consists of prostomium, peristomium and
collar region of three segments. It also carries dorsal and ventral lips.
2. Thorax: Consists of 5 segments bearing mucous glands ventrally. These
segments also bear parapodia where the notopodia with setae and the
neuropodia with hooks.
3. Abdomen: Long and composed of about 300 segments bearing parapodia
which are like those on the thorax.
 Dasychone sp.: It is about 5 cm long, body yellowish or green. The head
carries a crown of tentacles or gills consisting of 12-18 tentacles fused in the
lower half into a fan which is strongly marked with semicircular pattern. Each
tentacle is ornamented by pail steps and dark spots. It lives in a slimy tube.
 Bispira: It is up to 7 cm long, flat with about 100 segments. The branchial
crown of tentacles arises from two spiral lobes as 45 to 80 on each side, white
or very pale in color.
d) Family: Serpulidae e.g, Hydroides norvegica & Spirorbis corragatus: These
worms live in hard calcareous tubes, they have a crown of tentacles which are
ciliated for collecting the food and respiration. One of these tentacles is
modified to form an operculum to close the tube when the worm is drown inside

43
 the tube. In many individuals the operculum serves as a brood pouch from which
the larvae are liberated.

 Hydroides norvegica: With elongated, smooth and white tubes attached to

the substratum and is characterized by the lacking of calcareous matter in the
ventral side and hence it becomes arch-shaped in cross section, but when
crowded the worms come up from the substratum. The larvae prefere to settle
near the adults. This phenomenon is called gregariousness, it helps in the
formation of overwhelming number of worms. Each form thick encrustations
to submerged objects in the seas such as buoys, ships and biers.
 Spirorbis corragatus: Very small, spirally twisted with white calcareous
tube. The tube with the animal twisted in clockwise direction (dextral) or in
anticlockwise direction (sinistral). It is usually found attached to the green
algae Coulerpa prolifera. Rarely exceed 1.4 mm in diameter.

e) Family: Cirratulidae e.g , Cirratulus: These animals form slimy tubes in

the mud, without a crown of tentacles. They have gill threads on some parts. The
worm is pointed at both ends. The segments are very short, two bundles of many
tentacles on the segments 4 & 5. These tentacles are red. On the other hand the
other segments the tentacles which serve as gills appear in lesser numbers. The
body length is about 4 cm.

44
Reproduction In Polychaeta
1-Sexual reproduction:
Polychaetes are unisexual (sexes are separate) or dioeceous with the gonads not
fixed in numbers and position. The gametes are formed seasonally from the coelomic
lining. Fertilization is external in the sea water. Metamorphosis involves a
trochophore larva which is free-swimming and planktonic. The life-span of this larva
is about 3-4 days, after which it settles down forming the complete animal. Thus the
development is indirect in polychaetes.
Many Nereids and Syllids exhibit the phenomenon of "epitoke" wich involves
the reproductive individual or epitoke, that differs from atoke. Epitoke (Heteronereis)
or heteroform possesses two large eyes, reduced prostomial palps and tentacles. Also,
it has enlarged modified parapodia for swimming. Epitoke may be not have head
region in some species. Epitoke is the fertile part, where the atoke is not fertile.
All epitokal polychaetes at sexual maturity in the reproductive season, swim to
the surface of the sea in enormous groups for discharging their gametes. The
phenomenon is called "swarming" to ensure fertilization takes place over a relatively
short period.
The epitokal portion breaks and swims freely in a spiral manner to the surface.
By the sun-rise the epitokes rupture and fertilization immediately takes place. The
ciliated trochophore larva is formed in the next day and sinks down after 3 days as
described before.
2-Asexual reproduction:
This is best illustrated in the family syllidae as shown in the following selected
types:
a. By fission: A zone of fission is formed between two definite segments in the
middle of the worm. This region becomes differentiated into anal region for the
worm in front and a cephalic region for the posterior half. Sexual reproduction is
not associated in the worms undergoing such type of fission e.g. Filigrana.

45
 b. By fission after the development of an epitokous phase: After
assuming the epitokous condition, the worm divides by fission into an anterior
non-sexual and posterior sexual parts. Eventually, the posterior part acquires a
new head and becomes an independent male or female. The anterior portion
develops a new anal segment and acquires generative organs. It will again repeat
the same process of formation e.g. Syllis ramosa.
c. By combined fission and budding: A zone of fission or Proliferation is
first formed that may be produce not only the head of the posterior worm but
also a number of segments by budding. These become marked out into a chain
of individuals in which the genital organs appear. The anterior portion is
nonsexual and the posterior zooids are all of the same sex. The later then
separate and reproduce sexually e.g. Autolytus.
d. By budding from any segment: This is not confined to a certain region of
the body, but every segment is capable of budding a new individual, so that at
the end the original worm acquires numerous branches. Each branch with a head
and represents a new individual. The branches themselves may in turn form
secondary buds before their separation. In this manner a branched colony of
which some individuals develop genital organs and probably becomes free e.g.
Syllis ramosa.

Feeding In Polychaeta
1- Carnivorous or raptorial feeders:
Many errant polychaetes such as Nereis, Syllis & Glycera are carnivorous.
They have proboscis supplied with jaws as a result of contractions of muscles. The
proboscis is shoot out and the jaws catch the food and then the pharynx or the
proboscis is retracted by other muscles.

46
2- Sand and mud feeders:
In some polychaetes the proboscis works as a shovel taking sand or mud in the
mouth. The volume of sand or mud às a nutrition depends on its constituent of
organisms or their remains.
U-shaped burrow Arenicola marina inhabits U-shaped tube or burrow taking
the sand and throwing up castings at the hind end and can be seen that there is a
depression in the sand over the head shaft end. Under ideal condition of huge organic
matter within the sand, population of this worm may reach about 82000/Acr.

3- Tentacle feeders:
Some polychaetes feed by means of ciliated tentacles such as Terebella. This
animal has particularly extensive sized tentacles which are capable of reaching a
wide area of surface mud. These ciliated tentacles are grooved, hollow and have
complicated musculature enabling them to be moved.
The tentacles attached to the substratum by mucus and perhaps by sucktion or
by continual release. The reformation of the point of attachment provided that each
tentacle is working independently for food collection. They also collect some grains
for tube making from a wide area.
4- Filter feeders:
Some worms which have ciliated tentacles show tendency towards feeding on
particles suspended in water as well as those on the surface on the substratum.
In the families Sabellidae and Serpulidae, the filter is provided by branched
or pinnate tentacles which form a feathery crown throughout it the water is drained
or filtered. On the upper side of the tentacles and the pinnules there are short cilia
which beat toward the mouth. These cilia derive the suspended particles including
the planktonic organisms towards the ciliary tracts along the tentacles towards the
base of the crown. Here, the largest particles are rejected; medium sized particles are
stored for the use in the tube formation. The finest particles are carried to the mouth
and digested.
47
 Phylum: Arthropoda

 Coelomates, bilaterally symmetrical, metamerically segmented and their body

is covered with chitinous cuticle as exoskeleton.
 Each body segment carries a pair of jointed appendages of which some are
modified as foot-jaws.
 The body is differentiate into, head, thorax and abdomen.
 The coelom or main body cavity is haemocoelic where the vascular system is
of the open type.
 The nervous system is of the annelidan plan. They have well developed sense
organs such as eyes and antennae.
 Excretion takes place by nephridia, green glands, coxal glands and Malpighian
tubules.
 They have a separate muscular system and the body wall is dermal. The
muscles are striated and attached to the inner surface of the procuticle.
 Respiration takes place by gills, tracheae, lung books or body surface.
 The hard cuticular exoskeleton secreted by the hypodermis. It consists
basically of two layers: Outer thin epicuticle and inner thicker procuticle. The
epicuticle is composed of proteins, lipids but no chitin. The procuticle is
mainly made of chitin which sometimes combined with CaCO3 and phosphate;
it is differentiated into outer exocuticle and inner endocuticle.

The two problems of movement and growth result in consequence to the

presence of the exoskeleton have been eliminated by the division of the cuticle into
separate plates on each segment from which are the podomeres of legs and the
skeleton of each segment is connected by arthrodial membrane with that of the other
segments. Thus this thin flexible membrane allows the movement and articulation.
The second problem, that of growth have been solved by the periodic moulting or

48
ecdysis (shedding of the old cuticle and formation of a new one below the old). The
newly formed cuticle is thin enabling the animal to stretch and increase in size.

Subphylum: Crustacea
1- Class: Branchiopoda
Order: Anostraca - Artemia
Order: Notostraca- -Apus (Triops)
Order: Diplostraca
Suborder: Conchostraca Estheria(= Cyzicus) & Limnadia
Suborder: Cladocera -Daphnia & Moina & Bosmina

2- Class: Maxillipoda
a- Subdass: Ostracoda e.g. Cypris
b- Subdass: Copepoda
Order 1: Cycdopoida e.g. Cyclops
Order 2: Calanoida e.g. Calanus & Centropages
Order 3: Harpacticoida e.g. Euterpina & Schizopera
Order 4: Monostrilloida e.g. Monostrilla
Order 5: Caligoida e.g. Caligus
Order 6: Poecilostomatoida e.g. Chondracanthus
Order 7: Lernaeopodoida e.g. Lernaea & Salmincola
c- Subdass: Cirripedia
Order 1: Thoracica
Suborder 1: Lepadomorpha (Pedunculata) e.g. Lepas
Suborder2: Balanomorpha (Operculata) Balanus, Chelonebia & corn barnades
Order 2: Rhizocephala e.g. Sacculina
3- Class: Branchiura e.g. Argulus
4- Class: Malacostraca

49
a-Subclass: Phyllocarida
Order: Leptostraca e.g. Nebalia bipes
b Subclass: Hoplocarida mantis
calumnia
Order: Stomatopoda Squilla empusa
oratoria
c- Subclass: Eumalacostraca
i. Superorder: Syncarida
Order: Anaspidacea e.g. Anaspides
ii. Superorder: Peracarida
Order: Mysidacea e.g. Mysis
Order: Cumacea e.g. Diastylis cornula
Order: Amphipoda
Suborder: Gammaridea e.g. Gammarus & Corophium
Suborder: Hyperiidea e.g. Phronima & Hyperia
Suborder: Caprellidea e.g. Caprella
Order: Isopoda
Suborder: Oniscoidea e.g. Oniscus Ligia & Armadillidium
Suborder: Flobellifera e.g. Limnoria Cirolana & Sphaeroma
Suborder: Valvifera e.g. Idotea
Suborder: Asellota e.g. Asellus & Munna
Suborder: Epicaridea e.g. Pseudione & Stegias

Order: Tanaidacea e.g. Neotanais & Apseudes

iii. Superorder: Eucarida
Order: Euphausiacea e.g. Euphausia
Order: Decapoda
Suborder: Dendrobranchiata e.g. Penaeus
Suborder: Pleocyemata
51
 Infraorder: Caridea e.g. Lebbeus & Palaemonetes
Infraorder: Stenopodidea e.g. Stenopus & Spongicola verwsta
Infraorder: Astacidea e.g. Crayfishes (freshwater)
Infraorder: Palinura e.g. Panulirus & Scyllarus
Infraorder: Anomura e.g. Pagurus (hermit crab)
Infraorder: Thalassinidea e.g. Callianassa & Upogebia
Infraorder: Brachyura e.g. Portunus pelagicus (blue crab)
e.g. Ocypoda aegyptiaca (Sand crab)

Class: Branchiopoda:
The branchiopod animals have at least 4 pairs of leaf-like swimming legs
"phyllopodia". The abdomen is devoid of appendages and provided with a pair of
caudal styles or caudal furca. A cephalic carapace when present it may be either
shield or bivalve. Their development includes the nauplius larva or metanaupliu.

1- Order: Anostraca → Artemia The carapace is absent, the eyes are stalked,
uniramous and reduced antennae are present in female, these antennae are geniculate
in males (claspers). Artemia lives in brime water (brime shrimps), it can reach with
high abundance in brackish bonds and be hatched and rared from dissicated eggs.

2- Order: Notostraca → Apus (Triops) (Tadpole fish):

They have a carapace in a form of a dorsal shield. Eyes are sessile and
antennae are reduced. This animal is also characterized by have 3 long filiform and
many segmented endites of thoracic feet on each side of the body. The anal segment
bears a pair of caudal styles which are long and many segmented. Reproduction by
parthenogenesis.

3- Order: Diplostraca
These animals have a bivalved carapace which is usually encloses the trunk,
compound eyes are sessile and sometimes are fused, antennae are large and biramous,
trunk bears 4-27 pairs of appendages, telson with a pair of curved unjointed claws.

51
 a. Suborder: Conchostraca (Clam shrimps): These animals have a bivalved
carapace covering the whole body and provided with adductor muscles and
hinge like the bivalved mollusca e.g. Estheria & Limnadia.
b.Suborder: Cladocera: With 4-6 pairs of trunk appendages; carapace without
hinge or adductor muscle and covers only the trunk while the head is free and
bent downward. Their eyes are sessile and united together. Without naupliu
larva e.g. Daphnia, Ceriodaphnia, Bosmina, Simocephalus & Moina. They
are reproduce parthemogenetically.

Class: Maxillipoda
a) Subclass: Ostracoda → Cypris: They are called mussel or seed shrimps,
widely distributed in the sea and in freshwater. They are mostly minute (1.2 m)
and the body is indistinctly segmented covered with a bivalved carapace
having dorsal hinge and adductor muscles. The head constitute the major part
of the body and the trunk is reduced. With paired compounds eyes, two pairs of
well developed antennae for swimming, Mandibles with biramous palps. The
trunk appendages are few in number (2-4) and are slender. The abdomen is
short, apodous and ends with a caudal furca. Without gills and respiration is
integumentary. Parthenogenesis takes place in the freshwater genus Cypris
while, the development in others involves nauplius larva escape from the egg
and enclosed in a bivalved carapace.

b) Subclass: Copepoda: These are the most abundant crustaceans in the

world, small in size; free living or parasitic. They form the major part of the
diet for many other animals. Also, they are benthic or planktonic. The body is
divided into head, thorax and apodous abdomen. Without compound eyes but
with a median sessile and simple eye. Antennae are small, the antennules are
long and segmented and they are the principal locomotory organs. The

52
 antennules are geniculate in the male acting as grasping organs during
copulation. The head is fused with one or two thoracic segments to form the
cephalothorax which covers with the carapace. There are four or five free
thoracic segments carrying biramous appendages where each pair is connected
with copula (transverse plate) which bring about the combined movement of
the two limbs in each pair in swimming, after the manner of an oar (and hence
the name copepoda).
The abdomen is narrow, cylindrical and apodous. It is 4- segmented
where the first segment bears the genital openings and fused in the female with
succeeding one. The abdomen terminates with a telson bearing two caudal
rami.
The body of the crustacean copepod is differentiated into metasome and
urosome. There is articulation between the last thoracic and first abdominal
segments where it is markedly distinct and is commonly used to differentiate
between the anterior metasome and the posterior urosome.

A- The free-living copepods:

1- Order: Cyclopoida e. g. Cyclops
The metasome is sharply marked off from the urosome. The cephalothorax is
covered with a dorsal carapace and formed of the fusion of the head with two thoracic
segments. Articulation is between 5th and 6th thoracic segments. Antennae and sixth
thoracic segments are uniramous. There are two egg sacs in the female. They are
benthic and planktonic copepods living in fresh and salt water (but a few are
parasitic). 52 260uGure G Lives in freshwater, articulation occurs ida are 26(96 OU
Cyclops lives in fresh water articulation occurs between 4th & 5th pairs of
swimming legs. Two thoracic segments are fused with the head.
Oithona nana: marine cyclopod, females with their large two egg sacs are
common than males. There are four obvious thoracic segments in addition to the last
(6th) one which is contribute to the urosome.
53
2- Order: Calanoida → Calanus, Centropages, Diaptomus and Acartia.
Free living, the most numerous marine individuals and in the species of all
planktonic copepods. The cephalothorax is formed of the head and one thoracic
segment covered with dorsal carapace. The main feature is the metasome is clearly
distinct from the urosome where articulation is between the sixth thoracic segment
and the first abdominal one. The first to antenna is long and many segmented.
Calanus: Widespread in all oceans especially in the northern hemisphere
where it may be found at about 4000 meters depth. It is also found near the surface of
water and considered the main food of herring (‫)الرنجة‬. The first antennae are
symmetrical and slightly longer than the body carrying two long plumose setae on
their 24th and 25th segments. The female has one egg sac.
Centropages: It is characterized by the strong slightly asymmetrical spine on
the 5th thoracic segment. The forceps formed by the right 5 th thoracic limbs of the
male is particularly large, while the setae on the caudal rami (furca) are also
asymmetrical.
Acartia: The main generic characters of it are that; the body is elongated and
carries 4 obvious thoracic segments and the antennules with setae of very unequal
lengths. The cephalothorax is formed of the head fused with the first thoracic
segment only.
3- Order: Harpacticoida → Euterpina & Schizopera
Most of them are benthic but some are planktonic especially in shallow waters.
Their obvious features are their minute size (most being less than 1 mm long) and the
lack of obvious division between the main regions of the body. The anterior part of
the body is usually little broader than the posterior one. Antennules are short usually
with less than 6 segments. Egg sacs may be single or paired.

54
B- The parasitic copepods:
The four order Poecilostomatoida, Monostrilloida, Caligoida and
Lernaeopodoida are exclusively parasitic and contain over 1000 species. They are
usually called fish-lice as both marine and freshwater fish are the most common
fishes.
Some copepods are commensals or endoparasitic within polychaetes,
echinoderms, tunicates and bivalves. Generally, the female copepods is more highly
modified for parasitic existence than the male and some times the male is entirely
free living and displays the typical copepod appearance.
In the most parasitic copepods the larvae are free-swimming. In the common
Lerraeopodoida fish-lice a special attachment button and thread are produced by a
frontal gland on the head as in Salmincola. The mouth parts of ectoparasites are
adapted for piercing and sucking.
In lernaeopodoida the maxillae are adapted for piercing the skin of the host and
sucking its juices. These animals also have often lost their mouth parts and food is
absorbed directly from the host. For example, the body of the larva of Monostrilla
anglica which is enoparasitic within the blood vessels of polychaetes possesses two
or four large absorptive processes.

Subclass: Cirripedia
The most familiar examples of the cirripedes are the barnacles which are found
as fauling on ship bottom, piles etc... Cirripedes are the only sedentary group in
Crustacea. All of them are marine, approximately 900 described species are free
living. Some barnacles are commensals on wheals, turtles and other animals. The
free-living or thoracican barnacles can be divided into stalked and non-stalked
(sessile types).
Lepas (Gooze barnacle): With long stalk or peduncle covered by rinkled skin
that is attached to the substratum and bears at its distal end the capitulum. The
peduncle carries the vestigial antennule and has a cement gland. The peduncle is
55
formed of the pre-oral region, while the rest of the body form the bilaterally
compressed capitulum. The capitulum is surrounded by 5 calcareous plates; these are
median dorsal carina, two lateral posterior scuta and two lateral scuta anterior terga.
There is large adductor muscle between the two scuta.
Balanus (Rock barnacle): Without peduncle but has instead a basal disc
containing cement gland. The top of the animal is covered by an operculum formed
by the paired movable terga and scuta. The calcareous plates surrounding the animal
overlap one another and may be fused to some extent. These are 6 plates named
carina, rostrum and 4 calcareous lateral plates. The 6 pairs of appendages are directed
upward towards the mantle aperture. The major part of the body composed of the
cephalic and thoracic regions.
Stalked barnacles including the peduncle range form few millimeters to 7.5 cm
in length. The majority of sessile species are a few centimeters in diameter but some
are considerably larger. Some sessile barnacles have become adaptive for life on
other surfaces beside rocks such as on turtles and swimming organisms with hard
exoskeleton.

Parasitic barnacles:
Many of cirripedes are parasitic on other animals such as Sacculina from the
order Rhizocephala which is parasitic on the crabs. It appears as a tumour on the
abdomen of the host showing no sign of segmentation. No appendages, no mouth or
anus. It starts its life as a nauplius of the cirripede type but lacking a mouth or
alimentary canal. It passes into a cypris stage which cling by one of its antennules to
the underside of the abdomen of a crab. The crab is attacked by Sacculina while the
integument is soft after the process of ecdysis. The parasite becomes a cuticular sac
with a dart-like organ which forces its way through the antennule into the body of the
crab. By way of this dart the remnant of the larva, in the form of an undifferentiated
mass of cells, passes to the body cavity of the host. It then sends absorptive rootlets
which extend out invading the whole body of the crab excepting the heart and gills.
56
The presence of Sacculina on the crab prevents the cuticular formation at that spot.
Thus at the next moult of the crab the parasite comes to project freely like a tumour
under the abdomen where it may be found in the adult condition.
The adult parasite does not exhibit any homology to a cirripede. It consists of
an outer mantle with a cloacal opening and the mantle cavity surrounds an internal
visceral mass which contains the genital organs and a nerve ganglion.
Crabs attacked by Sacculina suffer from a great disturbance in growth and
development.

Class: Branchiura → Order: Arguloida Argulus

Argulus is the most familiar example of this class. It referes to as carp-lice as it
is an ectoparasite on this freshwater fish. This parasite has a pair of sessile compound
eyes and a large carapace covering the body. The body is formed of head, thorax and
abdomen. The first pair of antennae is used in attachment to the host. While, the
second antennae are small and minute.
The mandibles and the first maxillae are forming piercing organs enclosed in a
sucking tube or proboscis, in front of which is a median tube ending in a spine. The
second maxillae are divided into two portions, the anterior of which is modified into
sucking discs by which the parasite clings to the surface of the host.
There are four pairs of biramous thoracic legs or swimming feet carrying long
setae, no maxillipeds are found, the abdomen is bilobed and apodous. Argulus is
greatly dorsoventrally flattened.

Class: Malacostraca
This class contains about 3/4 of all crustaceans envolving the large forms such
as crabs, lobsters, shrimps and wood-lice. The body is always distinctly segmented
and typically the thorax is formed of 8 segments and abdomen of 6 segments (some
have 7 segments) ending with a telson. All of segments bear appendages.

57
 The first antennae are often ramous. The mandibles usually bear palps. In most
malacostracans the first one, two or three pairs of thoracic appendages form
maxillipeds. The last five pairs of abdominal appendages are biramous. They may be
used for swimming or carrying eggs in the female and the first one or two are often
modified in the males to form copulatory organs. The last abdominal appendages
form with the telson a tail fan.
A very important feature of malacostracs is the presence of the gastric mill in
the fore gut which helps in chewing the food. The female gonopores are always found
on the coxopodites of the 6th thoracic appendages and these of males on that of the 8th
thoracic ones. Nauplius larva is usually passed within the eggs but when present it
does not feed.

1- Subclass: Phyllocarida
Order: Leptostraca → Nebalia bipes
Nebalia is the comonst genus, it is a little shrimp-like marine animal, about 6-
12 mm in length. It lives in bottom mud and in the sea weeds. The body is formed of
the head, thorax and abdomen. This animal is characterized by some special features
these are; the presence of a large carapace which has adductor muscle in the
maxillary region and covers the cephalothorax and the anterior four abdominal
segments. The carapace also contains a rostral plate anteriorly. The abdomen is
formed of 7 segments where the last segment is apodous. The eyes are large
compound and stalked.
Antennae are well developed, where antennules possess squame and flagellum
and the antennae are uniramous. Mandibles with 3-jointed papls.
The thoracic appendages are foliaceous with endopodites of five undistinct
joints provided with long setae, broad exopodites and large epipodites serving as
gills. As in branchiopods they create a water current from which food particies are
filtered.

58
 The anterior four pairs of abdominal appendages are biramous and used for
swimming. While the 5th and 6th pairs are small and uniramous and the 7th segment is
apodous.
The abdomen ends by a telson which is unlike that of other malacostracans,
where it bears a caudal furca with two rami.
The development is direct, where the embryos being carried on the thoracic
limbs of the female, each hatching as a post-larva called a manca stage.
Finally, it is clear that the leptostracans are thus agree with the advanced
malacostracans in having fixed position of the gonopores and the division of
appendages into cephalic, thoracic and abdominal ones. However, they still retain
some primitive characters as the possession of an adductor muscle, an extra
abdominal segment, a caudal furca and foliaceous appendages.

2- Subclass: Hoplocarida
Order: Stomatopoda → Squilla mantis
Hoplocarida includes only one order, Stomatopoda, which includes marine
crustaceans known as mantis shrimps. These are best examplified by the genera
Squilla and Gonodactylus. The latter lives in burrows excavated in sponges or corals,
with the telson serving as a plug to the entrance of the hole.
Squilla mantis: This is the best known genus of the order. The body is dorso-
ventrally flattened and elongated. The abdomen is very large in proportion to the
cephalothorax. The carapace is reduced, shield-like and fused with the first two
thoracic segments and leaves the posterior clear four segments uncovered. The 3 rd
and 4th thoracic segments are vestigial.
The antennules have 3 flagellae, the antennae have squame and one flagellum.
The first five thoracic appendages act as maxillipeds and subchelate. Particularly the
second maxillipeds are enormously dependents in having large raptorial subchelae
which have a distal segment as a knife with toothed inner edges. The last three pairs
of thoracic appendages are slender, biramous and not chelated.
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 bun iramous bearing gills.
The pleopods are biramous bearing gills. The uropods and telson are very
large.
The surface of the carapace, the free thoracic and abdominal tergites and the
broad telson usually ornamented with keel-like ridges and spines and brilliantly
coloured.
In Squilla the first hatching larva is zooea but with peculiar features,
sometimes termed Erichthus.

3- Subciass: Eumalacostraca
They are typically composed of 20 segments, the abdomen being formed of six
segments, the last of which bear appendages and a telson deprived of caudal rami. An
adductor muscle is absent. Thoracic aprendɔges are rare; similar and are specialized
for special purposes.

(i) Superorder: Syncarida

Order: Anaspidacea → Anaspides
Syncarida is represented by only one order the Anaspidacea. They are
freshwater crustaceans, without carapace, the first thoracic segment is fused with the
head, non of the thoracic appendages is chelate or subchelate, the pleopods have long
fringed exopodites but the endopodites are reduced or absent except for the two pairs
in male. Anaspides is well known genus inhabiting pools and lakes.

(ii) Superorder: Peracarida

These animals are characterized by the fusion of at least one thoracic segment
with the head and the freedom of at least four thoracic segments even in the presence
of the carapace. Also, they have a brood pouch or marsupium in the female. This
pouch is formed by large plate like processes (oostegites) on certain thoracic coxae.
These processes form the floor of the marsupium while the thoracic sternite form its
61
roof. The development is direct and the youngs called post larvae which have most of
the adult features.
Order: Mysidacea → Mysis
The average length 3-5 cm, they are mostly marine and few live in freshwater.
The carapace often extends anteriorly as a rostrum but posteriorly is not united with
the last 4 thoracic segments. The thoracic appendages are similarly biramous. In
Mysis the first and the second thoracic appendages are maxillipeds. The first five
pairs of pleopods are large in male and small in female. The 6 th abdominal legs with
the telson form the tail fan. There are two statocysts at the bases. of the endopodites
of uropods. These animals form an important diet of some fishes.

Order: Cumacea → Diastylis cornuta

It is a small group (about 700 species), most of them inhibit the bottom
(benthic) in sand and mud. All the body regions are greatly enlarged and the uropods
are long and slender.

Order: Amphipoda
There are about 2700 described species, most of them are marine but there are
many freshwater species and some are terrestrials. The body is usually laterally
compressed. Eyes are sessile and compound. No carapace although the first and
second thoracic segments are fused with the head. Thoracic exopodites are
disappeared. The first pair of the thoracic appendages are maxillipeds. Gills are
thoracic and not abdominal. Pleopods are usually divided into two groups; the
anterior three pairs are used for swimming, while the others are directed backward
and used for jumping.

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Suborder: Gammaridea
Family: Gammaridae → Gammarus & Corophium
Gammarus is found in fresh and brackish waters. Body is elongated and
compressed bilaterally compressed. Cephalothorax is formed of the head and one
thoracic segment. No carapace. Head bears a pair of antennules which are longer than
the antennae and having a small branch in half way along it.
Corophium: Live in U-shaped burrows in mud and in muddy sand.

Suborder: Hyperiidea → Phronima & Hyperia

They are similar to those of Gammaridea in having the head united with one
thoracic segment and in having a well developed abdomen. The eyes cover much of
the head and the coxal plates of the thoracic legs are small. They are marine and
planktonic group. They seem to be carnivores.

Phronima
The body is skeleton-like and the fifth legs are chelate. Eyes are divided, so
that half of each one faces lateral and half faces upward.

Hyperia:
Pelagic, eyes usually very long covering great part of the head.
Suborder: Caprellidea
Family: Caprellidae → Caprella
In these members two thoracic segments are fused to the head. The eyes are
small, more nearly like those of gammaridean than of hyperiideans. Some of thoracic
legs may be absent or vestigial, and even the well-developed legs do not have
prominent coxal plates. The abdomen is short (sometimes totally absent) and there
are at most only one or two pairs of reduced pleopods. The family Caprellidae
involves "skeleton shrimps" due to their slender bodies. Their third and fourth legs

62
are vestigial but the oostegites that form the brood pouch of these legs; the legs also
usually give rise to epipodites that serve as gills. The fifth legs are reduced in certain
genera of the family.

Order: Isopoda
This is the largest crustacean order, the most successful in terms of
colonization of terrestrial habitats. There are many freshwater species but mostly they
are marine or terrestrials.
Their body is usually at least slightly flattened dorsoventrally, the first thoracic
segment carrying maxillipeds is fused to the head to form a tagma called the
"cephalon". No carapace, the other seven thoracic pairs of legs are used for walking
or clinging and without exopodites or epipodites. All of these thoracic legs are similar
and nonchelate. The brood pouch is found in the female as usual in peracaridans. The
abdomen is well develòped but not all of its segments are distinct; at least one is
fused to the telson. The first five pairs of abdominal appendages are biramous
pleopods, generally much flattened and functioning as gills. The uropods on the last
segment are also biramous but their form varies greatly, and in one large group of
isopods they are turned under the abdomen and cover the pleopods.
Isopods are benthic animals, 5-50 mm in length, and most are adapted for
crawling, some can move backwards rapidly as for forward.

Idotea: marine, the abdominal segments are fused some degree. No uropods
are visible from above but from below they appear as single elongated plates.

Limnoria: marine, resembles Idotea but the uropods are partially visible from
above (only the exopodites of each uropod).
Sphaeroma: marine, dorsoventrally flattened, rough, the last abdominal
segment is fused with the telson forming a tail fan. The uropods are clearly appeared

63
from both sides of the telson (exopodites and endopodites). It is wood borer, and
construct tunnels through the substratum and can cause extensive damage to it.
Oniscus: called "wood louse" or "Sow bug", terrestrial and found under the
stones, logs and bark. The body is broad, oval and dorso-ventrally flattened. The head
is fused with first thoracic segment forming the cephalothorax which is not covered
dorsally by carapace. Eyes are sessile and compound. Antennules and antennae are
uniramous. It feeds upon decaying vegetation.

Order: Tanaidacea → Aspeudes & Neotanais

Small group of animals (about 400 species), stricktly limited to the sea and
estuarine situations. The typical habitat of them is sediment including that which
accumulates in colonies of clump-forming algae. The general body plan of them is
similar to that of isopods and the same appendages but differ from isopods in having
a large chelate second thoracic appendages and a carapace covering the
cephalothorax which is formed of the head and two thoracic segments. The uropods
are generally slender and do not contribute to a tail fan of the type found in some
isopods.

Superorder: Eucarida
Involves most of the large malacostracans. The carapace is highly developed
and fused with all of the thoracic segments covering the cephalothorax. The eyes are
stalked. The marsupium (thoracic broad chamber) is absent. Usually there are larval
stages. The heart is a short chamber situated in the thorax. This superorder contains
only two orders; Euphausiacea and Decapoda.

Order: Euphausiacea → Euphausia

The most primitive, marine, pelagic and shrimp-like crustaceans.
Approximately 3 cm in length. They have biramous thoracic appendages which none
of them being modified as maxillipeds. The pleopods are used for swimming. They
64
have a single series of gills. Most of them are filter feeders and the filtration takes
place through the thoracic appendages. Most of them are luminescent. They are
considered the main food source of wheales. The life cycle includes nauplius larva,
protozoaea, zoaea and postlarval stage.

Order: Decapoda
Contains the largest and the most familiar crustaceans; prawns, shrimps,
lobsters, cray fishes and crabs. It is also from the largest orders of crustacean. Th are
about 8500 described species. Most of them are marine but some are live in
freshwater.
These animals are characterized by having a carapace covering the whole of
the cephalothorax. The three anterior pairs of the thoracic appendages are maxillipeds
and the remaining five pairs are usually with reduced exopodites and called walking
legs. The decapods are included in two suborders; Dendrobranchiata and
Pleocyemata.

Suborder: Dendrobranchiata
There are about 2000 species of shrimps, but only 350 belong to the
Denrobranchiata; the rest are in the Pleocyemata. In general, shrimps are slightly
laterally compressed, have a well developed abdomen, and use their biramous
pleopods for swimming. By spreading its tail fan and vigorously flexing its abdomen
ventrally, a shrimp can derive itself rapidly backward.
The endopodites of the thoracic appendages serve as legs for walking and
grasping. The exopodites are not often present or reduced. The carapace is usually
(not always) prolonged anteriorly as a pointed rostrum. The second antennae are
distinctive in that the exopodites are modified into squames.
The dendrobranchiate shrimps include those of the family Penaeidae, some of
which are large and of great commercial importance. The first to third legs of

65
penaeids are chelate, a feature not often seen in pleocyemate shrimps. As a rule the
eggs are not carried by the female, they fall away as they are laid.
The only other family of Dendrobranchiata is the Sergestidae. In this family
the first thoracic legs are not chelate but the second and third legs are chelate. The
fourth and fifth pairs of legs are reduced.

Suborder: Pleocyemata
Most shrimps and all non-shrimp decapods belong to this suborder. The
animals of this suborder differ from Dendrobranchiata on the basis of gill structure.
More than 95% of the decapods are pleocyemates, and the group is enormously
diversified, for it includes besides shrimps, the crayfishes, lobsters, crabs, hermit
crabs and many other distinctive types.

Infraorder: Astacidea → Crayfishes

They are freshwater and marine lobsters. The first to third legs are chelate, but
the first legs are especially large and their chaelae are powerful. The abdomen is
about as long as the cephalothorax and has a well developed tail fan. The exopodites
of the uropods are divided transversely into two articles. The female crayfish brood
their eggs until the young hatch as juveniles with the form of the adult where lobsters
brood until the young hatch in the mysis stage. Some crayfishes burrow in marshy
habitats and pastures where there is no standing water. Others spend most of their
lives in lakes and ponds, but dig burrows along the margins and find refuge in these
when the water is low or cold.

Infraorder: Palinura
There are about 130 species of them. They are differ from Astacidean in that
their first legs are not specialized as powerful chelipeds (sometimes one or more pairs
of legs are chelate but are not especially large). The exopodites of the uropods are not
divided into two articles but their tail fan is similar to that of an astacidean.
66
 They are mostly warm-water decapods, the spiny lobster (Family: Palinuridae
→ Panulirus) and shovelnose lobsters (Family: Scyllaridae → Scyllarus).

Infraorder: Anomura
The name Anomura (abnormal tail) refers to the fact that in many members the
abdomen is asymmetrical or otherwise peculiar. They have two special characters;
the second antennae are lateral to the eyes rather than between them and the legs of
the fifth pair are small and sometimes tucked away in the gill chamber beneath the
carapace, where they function as cleaners.
They include hermit crabs and some crablike types, marine animals and the
hermit crabs live in snail shells and sometimes in limpet shells, single valves of clam
shells, hard worm tubes or other hollow structures. Few do not protect themselves.
They have coiled abdomen. Pleopods are used for creating a respiratory current and
for brooding eggs and they are generally developed only on the left side. The left
uropods are larger than the right ones, but both are used to wedge the lip of the
abdomen in the upper part of the spire of the shell. The right chela is larger than the
left one and may be used to błock the aperture of the shell. Most species are detritus
feeders and scavengers, but some are carnivores at least under certain conditions. As
a shell-inhabiting hermit crab grows larger, it must find successively larger shells.

Infraorder: Brachyura
They are the true crabs, the abdomen is almost bent under the cephalothorax
and pressed tightly to it (there are rare exceptions). There are no uropods and the
males have only the first one or two pairs of pleopods which are filamentous and
uniramous function as to transfer the sperms to the females.
The carapace usually has a marginal ridge that separates the dorsal side of the
animal from the ventral side. The second antennae are between the eyes, not lateral to
them. The fifth legs are not smaller than others. As in most decapods other than

67
shrimps the first legs are stout and have powerful chelae. When the true crabs move
quickly, they generally do so sideways, rather than backward or forward.

Development In Crustacea

A part from the cirripedes and certain parasitic Isopoda, the majority of
crustacea are dioecious exhibiting sexual reproduction. The male is usually smaller
than the female. Copulation is usually the general rule. The male is usually having
certain appendages modified for clasping the female.
Eggs are mostly brooded, for different lengths of time, either attached to
certain appendages (many decapods), contained within a brood chamber (Cladocera,
Peracarida), or retained within a sac when the eggs are expelled (Copepoda).
Eggs are centrolecithal and the cleavage is superficial while the holoblastic
cleavage is uncommon and many species have a hollow blastula. Gastrulation by
invagination or by migration.
adult
(decapod) Phyllosoma
in scyllarus (modified mysis)

hatch directly
into Naupluis larva
Fertilized (Planktonic)
egg in Malacostraca

hatach directly
into in cirripedes metanauplus
(decapod) Protozoaea
in Homarus Zoaea
mysis Cypris larva Mysis larva
adult adult animal adult animal

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 Phylum Mollusca

This is invertebrates, including such familiar forms as the clams, oysters

(lamellibranchs), squids, octopods (cephalopods) and snails (gastropods). They
comprise the largest invertebrate phylum aside from the arthropods, and invaded the
sea, freshwater and land.
Although the classes of living molluscs superficially appear as a heterogenous
assemblage (snail, clam & squid), they are all built on the same fundamental plan.
Thus the foot in the snail is sued for creeping, in the clam for ploughing through the
mud and in Octopus for seizing the prey.
They are unsegmented, with a head, ventral foot and dorsal visceral hump,
covered with a mantle which often secretes a calcareous shell of different forms and
usually enclosing partially a mantle cavity. This contains a pair of ctenidia and into
which open the anus and kidneys.
The alimentary canal usually possesses a radula in the floor of the buccal
cavity. The circulatory system is of the open type and consists of a heart, veins and
arteries expanding into an extensive haemocoel with haemocyanin as a respiratory
pigment. The heart typically consists of one ventricle and two symmetrically placed
auricles, but the monoplacophoran Neopilina has two auricles and two ventricles.
Nautilus possesses two pairs of auricles. Lower gastropods have only one auricle and
one ventricle.
The coelom varies in development but is always represented by the
pericardium, the cavity of the kidney (which communicates with the pericardium)
and the cavity of the gonads. The nervous system consists of a circumoesophageal
ring (often concentrated into cerebral and pleural ganglia), pedal cords or ganglia and
visceral loops. Animals are dioecious; some are hermaphrodites. Development often
involves a trochophore larva.

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 Foot in Mollusca

It is a muscular organ, lies ventrally, containing mucous glands and modified

to perform many functions such as adhesion, creeping, burrowing and catching prey:
1. In Polyplacophora: Foot is broad, flat, occupy the whole ventral surface to
facilitate adhesion to the substrata.
2. In Gastropoda: Foot lies behind the head as a result of torsion to close the
aperture of the shell (an adaptation to escape from enemies).
3. In Pelecypoda: Foot is laterally compressed & varies in form (adaptation to the
mode of life): A disc, elongated (rock boring), geniculate (Cardium) wel (boring
Lithophaga), reduced developed (Anodonta), compensated by stick threads or
byssus (Mytilus).
4. In Scaphopoda: Foot as short cone, projected downward to burrowing in sand.
5. In Cephalopoda: Foot represented by 8-10 arms, with suckers (to catch food
and preys) and by a funnel (output of water from the mantle cavity & all
openings of the animals).

2- Class Monoplacophora:
The Monoplacophora is extinct since the Devonian (350 million years ago), but
recently represented by four species belonging to the genus Neopilina of the order
Tryblidiacea. They are bilaterally symmetrical, possess a ventral foot and an external
shell made of one piece. The mantle cavity is a shallow space containing 5 or 6 pairs
of external gills. Coelomic cavities are relatively well developed. The heart has two
auricles and two ventricles. Animals are dioecious, with two pairs of gonads, which
discharge through two of the 6 pairs of nephridia. The nervous system is primitive
with longitudinal pallial and pedal cords. The metamerism of Neopilina may
constitute an additional evidence for the relationship of molluscs and annelids best
shown by their similar embryonic development.

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3- Class Polyplacophora:
Polyplacophorans are commonly considered among the most primitive
molluscan group. They include chitons which live on hard substrata and become
highly adapted for adhering to rocks and shells. The foot is broad and flat occupying
the whole ventral surface to facilitate adhesion to the hard substrata. Partially
embedded in the mantle are eight transverse calcareous plates, sometimes encircled
with scales or spicules which may be so long and dense (Acanthopleura). The shell
plates are exposed to varying degrees of mantle reflection, from the nearly naked
(Chaetopleura) to only the apices being visible (Symmetrogephrus) to the
completely covered by the mantle (Amicula). The shell plates are composed of only
two layers: an upper tegmentum of conchiolin and calcium carbonate and a lower
articulamentum (entirely of calcium carbonate).
All chitons are dioecious. Copulation does not take place and fertilization
mostly occurs in the mantle cavity. The eggs are shed into the sea either singly or in
strings. In some cases the prelarval development or even the entire course of
development takes piace in the mantle cavity and the animal directly gives birth to
young without passing through a free-swimming larva.

4- Class Aplacophora:
These are a small group of aberrant deep-sea molluscs, being worm-like
without distinct mantle fold or shell, although the mantle surface is embedded with
calcareous spicules. The foot is reduced to only a small median ventral ridge lying in
a small longitudinal groove, a vestige of the mantle cavity. The posterior end of the
body contains a cavity called the cloaca, perhaps another remnant of the mantle
cavity into which open the anus and a pair of nephridiopores. Gills are lacking
(Proneomenia) or sometimes secondary gills are present (Chaetoderma). The
aplacophorans are either hermaphroditic (Proneomenia) or dioecious
(Chaetoderma). A typical free-swimming trochopnore larva is present, and in some

71
forms newly metamorphosed individuais possess seven dorsal plates, being evidence
that apiacophorans are derived from chitons.

5- Ciass Gastropoda:
This is the largest and certainly the most successful class of molluscs. Marine
species have become adapted to life on all types of bottoms as well as to a swimming
pelagic existence. They have invaded freshwater and the pulmonate snails have
conquered land.

Origin and Evolution:

The most significant modification of the gastropods, from the general
molluscan plan, is the twisting or torsion of the body. Torsion is entirely distinct from
the coiling of the shell; fossil evidence indicates that coiling of the shell evolved
before torsion.
Torsion actually involved the twisting of most of the body behind the head
including the visceral mass, mantle cavity and foot, 180 degrees counter-clockwise.
The mantle cavity, gills and anal and renal openings were consequently located
anteriorly behind the head. Internally the digestive tract instead of being a straight
tube was thrown into a loop; the auricles became located anterior to the ventricle; the
nervous system was twisted along the pleuro-visceral connectives in the form of
figure of eight (Streptoneurous). There might also be a reduction of the paired organs
such as the kidneys, auricles and even the genital ducts, torsion thus leading to the
asymmetry of the animal.
Torsion is not merely an evolutionary hypothesis, for it appears in the
embryological development of living gastropods. The larva is at first bilaterally
symmetrical and then undergoes twisting.
The advantages of torsion are not clear, although many theories have been
advanced to explain the evolutionary significance of torsion. Garstang maintained
that torsion was an adaptation during larval life, and the twisting anteriorly of the
72
pallial cavity allowed possible the retraction of the velum, which was originally
prohibited while the cavity was blocked by the foot. The larva could then escape
predators by simple retraction of the velum and cessation of its ciliary beating thus
sinking to the bottom. According to Garstang torsion was suggested to be of no
adaptive value to the adult, and on the reverse created for it a considerable number of
problems. However, there is much to state against this theory:
There are many pelagic larvae (e.g. of Lamellibranchs) that are not twisted but
still survive pelagic larval life.
Some forms (e.g. Haliotis) undergo torsion on two stages, a first through 90º in
which the animal is pelagic and unable to retract its head, and a second, after
settlement on the bottom, which bring the shell round the full 180º.
The cilia of some larvae are under nervous control and thus could be simply
stopped without the need of withdrawing the velum into the mantle cavity.
It seems difficult to understand how cessation could greatly increase the
chances of escape of the larvae from some plankton feeding predators.
A more recently advanced hypothesis states that torsion was not a larval
adaptation but rather was of advantage to the adult. The shifting of the mantle cavity
to the anterior side made it possible for the inhalant current to move in the same
posterior direction as the external currents thus increasing the ventilation of the
mantle cavity. A serious objection is that there has been a general tendency among
molluscs to restrict the pailial opening and to rely completely upon ciliary or
muscular action for propelling water through the mantle cavity. On the reverse,
torsion raised a number of disadvantages to the adult gastropod. The chief problem is
that of fouling, in which waste from the anus and nephridia would have got on the
gills in case the water circuit in the pallial cavity had remained as it had been in the
ancestral moliusc. The development of a cleft or a series of holes in the shell and
mantle solved this problem in primitive gastropods, and the anus retracted from the
edge to a position beneath the cleft.

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 The respiratory current as it passes out of the mantle cavity flows up and
carries with it the anal and renal waste through the cleft. As far as the shell itself is
concerned, it is pointed out that all existing gastropods possess asymmetrical, or
secondarily symmetrical shells. The plano-spiral shell had the disadvantage of not
being compact, with the consequence that the diameter of the shell could become
relatively great. The problem was solved with the evolution of asymmetrical coiling,
in which the coils were laid down a central axis, the columella, and each coil lie
beneath the preceding coil. Thus, the conical spiral compact shell of modern
gastropod evolved.
The new position of the shell restricted the mantle cavity to one side of the
body and this resulted in the reduction or complete loss of the gill, auricle and kidney
on that side.
After the evolution of modern conical gastropod shell, solution of fouling
problem involved:
The retension of the primitive cleft or perforated shell (Haliotis); the loss or
reduction of one of the gills and the respiratory current swept laterally through the
new mantle cavity; or detorsion, in which the body underwent a reversal of the
twisting process of torsion. Detorsion placed the mantle cavity and anus again at the
posterior end and the body adopted a secondarily derived bilateral symmetry.

Classification of Gastropoda

The gastropods are classified into three subclasses. The large majority of
gastropods belong to that subclass which exhibits torsion in full development, i.e. the
mantle cavity, gills and anus are located anteriorly. This is the Prosobranchia (gills
anteriorly located) or Streptoneura (coiled visceral loop). From the latter evolved the
two remaining subclasses. The subclass Pulmonata, land snails, has retained the
posterior-anterior position of the anus and mantle cavity has been modified into a

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lung. The Opisthobranchia (gills located posteriorly) display various stages of
detorsion; the shell is usually either reduced or absent.

I- Subclass Prosobranchia (Streptoneura):

Marine and freshwater; gills and shells retained; pallial cavity and organs
located anteriorly: gills in front of heart. The nervous system is twisted along the
pleuro-visceral connectives in the form of 8-shape (streptoneurous).

1. Order Archaeogastropoda (Diotocardia): Primitive forms with two gills,

two auricles and two kidneys; nervous system never concentrated; shell coiled
or secondary symmetrical. Largely marine. Patella (limpets), Fissurella,
Haliotis (abalone or ear shell), Trochus, Nerita, Turbo.

2. Order Mesogastropoda: Possess only one gill, one auricle and one
kidney; operculum may be present, proboscis typical. Chiefly marine. Littorina
(periwinkles), Crepidula (slipper shells), Cypraea (cowrie), Strombus, Natica,
vermetus (worm shells), the pelagic Heteropoda (greatly compressed, so that
the foot is transformed into a powerful fin-like swimming organ), Lambis,
Tonna, Vivipara, Pirenella, Bulinus, Oncomelania.

3. Order Neogastropoda: Differ from the mesogastropods in having a

bipectinate osphradium, a concentrated nervous system and a shell usually with
a siphonal canal. Carnivorous species with a radula containing two or three
large teeth in each row: nearly all have an operculum. All marine. Murex,
Urosalpinx, Purpura, Buccinum (whelk) Fasciolaria. Some genera possess a
poison gland associated with the radula, Conus, Terebra, and Fusus.

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II- Subclass Opisthobranchia:
Have one auricle and one kidney but displays various degrees of detorsion and
reduction of shell and mantle cavity which shows a tendency to occupy a posterior
position; auricle behind ventrical; visceral loop untwisted and symmetrical:
hermaphrodite. Exclusively marine, nervous system concentrated and symmetrical
(euthyneurous).
1- Order Tectibranchia: Shell present, although reduced or covered by mantle;
one true gill still present; some with secondary bilateral symmetry. Aplysia (sea
hare), Dolabella, Philine (bubble shell).

2- Order Pteropoda (Sea butterflies): Pelagic; shell present or absent; anterior

part of foot expanded into powerful swimming fins. Cavolina, Limacina.

3- Order Nudibranchia (Sea slugs): Shell absent and body is secondary

symmetrical; mantle cavity and true gills are absent; respiration takes place
through the general body surface or cerata or secondary gills around anus; the
digestive gland is extensive and the nervous system is concentrated.
Chromodoris, Hexabranchus, Aeolidia.

III-Subclass Pulmonata:
One auricle and one kidney, gills are absent and located anterior; mantle cavity
converted into a vascularized chamber for respiration in air or secondary for water;
nervous system concentrated and symmetrical (euthyneurous); shell usually present
but there is no operculum; hermaphroditic; development is direct. They are terrestrial
and freshwater but few are marine.

1- Order Stylommatophora: Two pairs of tentacles with eyes located at the tip of
the posterior pair; entirely terrestrial. Helix, Eremina, Arion, Limax, Lanistes, Pupa,
Onchidium, Succinea, Achantina.

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2- Order Basommatophora: One pair of tentacles; eyes near the tentacle base;
freshwater forms; require air for respiration although some takes water into mantle
cavity and a few have developed secondary gills. Some are terrestrial but few are
marine. Lymnaea, Biomphalaria, Bulinus, Physa, Cleopatra, Segmentina, Abida.

IV. Class Pelecypoda:

The Pelecypoda, or Bivalvia, or Lamellibranchia comprise clams, oysters and
mussels. They are bilaterally symmetrical and compressed. The body is covered with
a bilobed mantle, which secretes a bivalve shell that is hinged dorsally. One or two
adductor muscles close the two valves. The ventral foot is also laterally compressed,
hence the name Pelecypoda (=hatchet foot). There is no distinct head, and radula is
also absent.
The mantle cavity is the most capacious of any class of molluscs. On each side
of the body there are a pair of labial palps and a pair of large gills which function in
most species in food collection in addition to respiration. Sexes are separate in the
majority of pelecypods although there are some hermaphroditic species (as Cardium,
pecten). Development is accompanied by metamorphosis leading to a trochophore
larva followed by a veliger.
Most of these characteristics represent modifications that enabled
lamellibranchs to leave the hard substratum of the ancestral mollusc for a soft-bottom
habitat, becoming burrowers in mud and sand, although a few have become highly
specialized to bore in sandstone, calcareous rocks or even granite.
The bivalves consist of three subclasses, Protobranchia, Lamellibranchia and
Septibranchia, of which the first is considered the most primitive, and the last the
most highly specialized of all existing bivalves. The majority of bivalves are
lamellibranchs.

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General organization of the shell:
Bivalve shells range in size from the tiny seed shells of the freshwater family
Sphaeridae (2mm long) to the giant clam Tridacna (up to 4 feet long and 500 pounds
in weight). They further exhibit an infinite variety of shapes, surface sculpturing and
colours.
The umbo, the oldest part of the shell, is usually directed anteriorly
(inequilateral) in which case differentiation of right and left valves is made possible.
In most pelecypods the two valves are similar and equal in size (equivalve), but in
some sessile families such as the Ostreidae (oysters) and Spondylidae (Spiny
Oysters), the upper, or left valve is always larger than the right attached valve
(inequivalve). The two valves are attached dorsally by an elastic band, the hinge
ligament, composed of conchiolin and continuous with the periostracum and control
the opening and closing of the valve in combination with the adductor muscles. In
most species there are teeth and opposing sockets located on the hinge line beneath
the ligament to prevent lateral slippering. In some forms such teeth are large and few
(Spondylus), or fine (Nucula, Arca) but totally wanting in still others (Anodonta).
As regards the pallial line, the impression of the pallial muscles on the interior
of the valve, shells are described as being integripalliate (Anodonta) when possessing
a confluent pallial line, and sinupalliate (as in Solen, Meretryx) in case the line is
broken posteriorly by a comparatively large sinus.
Mantle:
Despite the muscular attachment of the mantle shell, it happens occasionally
that some foreign object lodges between the mantle and the shell, becoming then a
nucleus around which are laid concentric layers of nacreous materials. In this manner
a pearl is formed. The pearl oysters, Pinctadas margaritifera and P. mertensi,
inhabiting most of the warm pacific areas produce the finest natural pearls. However
pearls are formed in most bivalves, including freshwater clams.
In most pelecypods, the inhalent current enters anteriorly and ventrally and
passes back out posteriorly and dorsally. This system allows the animal to burrow by
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the anterior end first and to leave the inhalent posterior end projecting above the
surface of the bottom and thus avoids excessive sediment. A further modification is
that of sealing of mantle edges at one point or more; the most frequent point of fusion
is at the hind end between the inhalent and exhalent passages, thus a distinct inhalent
and exhalent apertures are formed. Their edges may become elongated to form
retractile tubular siphons of varying lengths. Mantle fusion has developed in some
species (Cardium & Tridacna), where most of the ventral margin has become sealed
except for a third (pedal) aperture through which the foot protrudes.

Foot:
The foot exhibits various forms and is modified according to the mode of life
of the animal. It presents the most primitive condition in Patella where it assumes the
form of a disc. In some rock-boring forms as Rocellaria, the animal is attached
during burrowing by a mpivot-like columnar foot. In still other boring bivalves as
Lithophaga and Modiolus the foot is no longer a functioning organ and its
construction is compensated by the development of a powerful byssal gland which
secretes byssus threads. These stick the animal to the wall of the burrow thus gaining
purchase during the boring process. In a few forms, such as Cardium, the foot is
geniculate. In forms, like Anodonta and Unio, where the foot prolongates through the
muddy bottom, it is large, well developed and wedge-shaped.

Adductor Muscles:
Only very few pelecypods have equal anterior and posterior adductor muscles,
i.e. isomyarian, and thus two equal impressions are left on the interior of each shell
valve, as in Pectunculus. However, in the majority of lamellibranchs, e.g.
Lithophaga, Mytilus, Nucula, the anterior adductor is much reduced than the
posterior, i.e. heteromyarian. In forms with nearly rounded shell, like Pecten, there is
only one central adductor muscle, i.e. monomyarian.

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Gills (Ctenidia):
Pelecypods have retained the single pair of gills of the ancestral mollusc; the
apparent double set of gills on each side of the body in many groups is actually
derived by the folding of the single gill. Each gill consists of a central axis, which
bears two rows of hollow lamellated filaments. Tufts of intermingled cilia loosely
interconnect the filaments of a single gill. There are ciliary tufts, which lock the ends
of the outer filaments to the inner surface of the mantle and the ends of the inner
filaments to the sides of the foot. Behind the foot ciliary junctions lock together the
gills on each side.
In the Lamellibranchia the gills extend more anteriorly than in the
protobranchia. They project laterally to such an extent that they become folded on
each side of the axis into a U-or V-shaped demibranch. The arm of each demibranch
attached to the gill axis is the descending limb and the free arm is the ascending limb.
In a few lamellibranchs the individual filaments are still discrete, i.e.
Filibranchiate. Interlamellar junctions have grown between the two limbs of each
demibranch (absent in Arca) but adjacent filaments are only attached by tufts of cilia
as in Lithophaga, Mytilus and Pecten. But in most lamellibranchs ciliary junctions
have been replaced by actual tissue and lamellae consist of solid sheets of tissue, i.e.
Eulamellibranchiate. Furthermore, interlamellar junctions have increased in number.
There are many pores in the gill lamellae of Anodonta through which water passes
into the interlamellar spaces.
In the Septibranchia the mantle has completely taken over the function of
respiration and the gills have become degenerate and modified to form a pair of
perforated horizontal muscular septa. By musclar contraction each septum moves up
and down like a piston of a pump and forces water into the inhalent and out of the
suprabranchial chambers.

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 Pearl Formation
Mantle lobe is intimately attached to the shelIl valve by a series of pallial
muscles.
It happens occasionally, that an object (sand grain, bacteria, larval stage) finds
its way between them.
As a defense reaction "Natural defensive phenomenon" the mantle secretes
concentric layers of nacreous layer (hence the name "mother of pearl"). Chemically,
composed of 90% CaCO3 + 4% water + conchiolin).
Pearl industry (for artificial culturing pearls): by inserting an irritant (seed)
such as crystalline guanine, drop of liquid, a particle of nacre, piece of fish scales,
between shell and mantle of a young oyster (genus: Pinctada) and left in its natural
habitat for about three years or more.
A good pearl is spherical in form, with bright luster (shining). Pearls take
different colours (silver, cream, white, pinkish, yellowish ting). Iridescence pearls
(100-150 years).
Irregular pearl shape is called "baroques" & caused by either physiological
action (as secretion of unusual conchiolin mechanically by moving the nucleus).

Classification of Pelecypoda:

Subclass Protobranchia: Primitive; ctenidia posteriorly placed and gill filaments

not folded; always with two adductor muscles; feeding by enlarged labial palps.
Nucula.

Subclass Lamellibranchia: Ctenidia play the main role in feeding, labial palps
reduced; ctenidial filaments elongated and reflected; lamellae united by ciliary
junctions or by vascular tissue junctions. Adductor muscles two, either equal
(isomyarian) unequal (heteromyarian), sometimes reduced to one (posterior;
monomyarian).

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Order Taxodonta: Gills filaments free, lacking interlamellar junctions. Hinge with
numerous uniform teeth. Mantle lobes free throughout. Adductors mostly equal.
Arca.

O. Anisomyaria: Gills usually filibranch and with vascular interlamellar junctions;

adductor musculature heteromyarian or monomyarian; hinge variable; mantle lobes
attached at one point to separate an exhalent aperture; usually no siphons; foot
reduced or absent. Mussels (Mytilus, Modiolus); oysters (Ostrea, Spondylus);
scallops (Pecten); rock-borers (Lithophage).

O. Heterodonta: Gills eulamellibranch; isomyarian adductors; mantle edges usually

united at one or more points posteriorly forming siphons. Cardium, Tridacna, Donax
(razor clams).

O. Schizodonta: Gills eulamellibranch, hinge teeth variable. Unio, Anodonta.

O. Adapedonta: Gills eulamellibranch, mantle margins united except for a pedal

gap; siphons long and united; hinge often lacking teeth. Sand burrowers, (Solen);
rock-borers (Saxicava, Pholas); wood-borers (Martezia, teredo or shipworm).

Subclass Septibranchia: Gills degenerate, inhalent chamber and suprabranchial

cavity separated by a pumping septum. Poromya, Cuspidaria.

V. Class Scaphopoda:
This comprises a small group of molluscs, which stands in some ways between
the Gastropoda and Pelecypoda. They are greatly specialized for burrowing and are
known as tusk or tooth shells, from the fact that the shell, which is an elongated
cylindrical tube open at both ends, had usually the shape of a trumpet or elephant's
tusk. The body is greatly elongated anteroposteriorly. The head and foot emerge from
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the wider anterior end of the shell while the posterior narrower end or apex remains
above the surface of sand, and serves for the circulation of water into and out of the
mantle cavity. The head is reduced to a short conical projection, or proboscis, bearing
the mouth but has none of the usual sense organs. The foot resembles a short cone in
the common genus Dentalium and can be projected downward into the sand during
burrowing. The mantle cavity extends the whole length of the ventral surface and
contains no gills. The sides of the head bear a large number of thread-like tentacles
called captacula. These are provided with adhesive tips to capture the food, which is
then brought back to the mouth. They may well serve a sensory function. There is a
well-developed radula. The heart is absent but there is a system of blood sinuses.
There is a pair of nephridia. Scaphopods are dioecious and genital products are shed
out by way of the right nephridium.

VI. Class Cephalopoda:

The cephalopods (nautili, squids and octopods) are the most specialized and
highly organized of all molluscs. They are bilaterally symmetrical with well-
developed head projecting into a circle, or crown, of large prehensile tentacles, or
arms, which are homologous to the anterior part of the foot of other molluscs. The
remaining part of foot forms the funnel or siphon, which is a muscular organ. In the
mantle cavity there are one or two pairs of typical ctenidia, kidneys and auricles.
They have a chambered shell, in the last chamber of which the animal lives, though is
most modern representatives it is reduced and internal or totally absent. The nervous
system is greatly centralized and eyes are of great size and often complex in structure.
Eggs are heavily yolked. All cepholopods are dioecious (except few).

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Classification of Cephalopoda:

Cephalopods are divided into two subclasses:

Subclass Tetrabranchia (Nautiloidea):
Possess a completely developed external shell and two pairs of gills, kidneys
and auricles. The nautiloids are the only group now existing; all other members are
fossils.
The shell of Nautilus is coiled over the head in a bilaterally symmetrical
plano-spiral. Only the last two whorls are visible since they cover the inner whorls. In
spite of its coiled nature, the nautiloid shell is radically different from the gastropod
shell. The former's cavity is divided by a series of transverse septa into internal
chambers of which the last is the largest and is occupied by the living animal. As the
animal grows, it periodically moves forwards and the posterior part of the mantle
secretes a new septum. From the soft parts of the animal a cord-like prolongation of
the visceral hump, termed the siphuncle, extends through pores in the whole series of
septa up to the apex of the spiral. It contains blood vessels and secretes gases into the
empty chambers making the shell light and swim with considerable speed as well as
maintaining a constant pressure. The head is conical and bears simple eyes. Tentacles
are numerous but without suckers, the funnel is not simple but formed of two lobes.
The trunk is sac-like and in the mantle cavity there are two pairs of ctenidia, kidneys
and auricles.
Subclass Dibranchia (Coleoidea):
This comprises the remaining living cephalopods, in which only a single pair
of gills, kidneys and auricles are present and the shell is either reduced and internal
(squids) or completely lacking (octopods). An external shell may exist but its cavity
is not divided by septa.
There are eight or ten arms bearing suckers. The funnel forms a complete tube.
Eyes are of a complex structure. Chromatophores as well as an ink gland are present.

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Order Decapoda:
Dibranchia with eight arms and a pair of long tentacles provided with stalked
suckers and with well developed internal shell. Body more or less elongated and
bears fins. Loligo, Sepia, Sepiola, Architeuthis (giant squid).

Order Octopoda:
Dibranchia with eight arms of equal length provided with sessile suckers and
with or without slight vestiges of internal shell. Body is globular and without fins.
Octopus, Argonauta.

Development of Mollusca
In existing mollusc species, the zygote undergoes typical holoblastic
determinate and spiral cleavage. Of the first formed blastomeres, the D cell is ususlly
rather larger than the other nearly equal cells. Four quartettes of micromeres are
eventually produced forming a cap over the macromeres. In large yolky eggs the
contrast in size between the micromeres and macromeres of the embryo is great and
gastrulation is effected by epiboly or immigration (as in Patella), by invagination in
small egg with litter deutoplasm and consequently with a comparatively wide
blastocoel as in chitons and most pelecypods, and by both processes in medium-sized
egg. The resulting gastrula quickly develops into a free-swimming trochophore and
the fate of the cleavage quartettes is virtually identical to that of polychaetes. The first
three quartettes produce the ectoderm and the macromeres with the 4a, 4b and 4c
cells of the fourth quartette the entoderm. The 4d-cell becomes the partent cell of the
mesoderm. It divides into two mesodermal bands, one on each side of the gut. Then
instead of segmentation as in polychaete annelids the mesodermal bands break into
single cells, some of which elongate giving rise to muscle cells. Relying on the
absence of any trace of segmentation in the embryonic mesoderm, it is strongly
believed that Mollusca are unlikely to descend from a segmented ancestor.

85
 In embryos undergoing epibolic gastrulation, the original lumen forms the mid-
gut. A shell gland then appears on the dorsal side of the ectoderm and secretes an
embryonic shell. The mid-gut becomes considerably widened and acquires
diverticula. A foot primordium protrudes from the ectoderm on the ventral side just
behind the stomodaeal invagination. This trochophore is distinguished from its
corresponding stage in polychaete annelids by the presence of the shell gland and the
traced shell.

Development of the trochophore stage varies in the different molluscan classes:

In Amphineura, the trochophore metamorphoses directly into the adult form.

The post-trochal region elongates to form the major part of the body. The shell gland
extends in the pretrochal region and the shell plates are formed. The prototroch
degenerates and the animal sinks to the bottom as a young chiton.
A free swimming trochophora larva is found only in the primitive
Archaeogastropoda; in all other gastropds, the trochophore stage is reduced and is
passed through in the egg. More characteristic of marine gastropods is the free-
swimming veliger stage which is derived from the trochophore and is the hatching
stage. The veliger is characterized by its swimming organ, the velum, which consists
mostly of two large semicircular folds, sometimes subdivided into four, bearing long
vibratile cilia. The velum arises as an outward extension of the prototroch of the
trochophore. Not only is it an organ of locomotion, but also the velum serves for
attracting food particles to the mouth. Furthermore, the foot, the operculum, tentacles
and mostly eyes appear. The cup-shaped shell of the trochophore acquires spiral
structure because of unequal growth. The stomodaeum develops and connects with
the mid-gut and the larval retractor muscle is formed.
Torsion is eventually takes place; the shell and visceral mass are twisted 180°
in relation to the head and foot. After a period of wandering and free-swimming, the
veliger larva settles down on the desired substratum. The velum becomes gradually
reduced and the foot on the contrary, extends acquiring a creeping function, thus
86
adopting the adult mode of locomotion. The shell grows in size, and the young
gastropod is formed.
The embryology of opisthobranchs recapitulates several stages in their
evolution. Torsion takes place in the veliger, although it is not complete, and is then
followed by detorsion. Especially in nudibranchs, the veliger shell is cast off during
metamorphosis, and the pretorsional positions of the mantle cavity, gills, anus, and
renal aperture are acquired once again.
Some marine prosobranchs (Conus, Natica), nearly all freshwater
prosobranchs, as well as all pulmonates have no free swimming larvae (passed within
the egg) and at hatching, a tiny snail emerges from the embryonic capsule. A few
gastropods are ovoviviparous.
In marine pelecypods, the development of a free-swimming trochophore
succeeded by a veliger larva is typical. Torsion does not take place and the veliger is
always symmetrical. The shell and shell gland are present at first in the form of a
single dorsal plate, which then grows laterally and ventrally. Consequently the dorsal
plate become folded forming the two halves characteristic of pelecyopds

The loss of the velum during metamorphosis is very sudden being abruptly cast
off.
In the freshwater family Unionidae, the veliger is termed glochidium and has
become highly modified for a parasitic existence. The digestive tract is poorly
developed and there is neither mouth nor anus. The glochidium of Anodonta bears
hooks by which they cling to the skin and fins of the host fish. During a parasitic
period that lasts 10-30 days many of the structures of the encysted larva disappear
and the adult organs begin to develop. Eventually the immature clam breaks out of
the cyst, falls to the bottom and burrows in the mud where the development is
completed.
Scaphopod development as that of marine pelecypods. There is a free-
swimming trochophore larva, succeeded by a bilaterally symmetrical veliger with a
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bilobed mantle and shell as that of pelecypods. The mantle lobes then fuse along the
ventral margin and a cylindrical shell, open at both ends is resulted.
Development of all cephalopods is direct. Eggs are telolecithaly and cleavage
is meroblastic. Cleavage results in the formation of a germinal disc, or cap of cells at
the animal pole where the embryo will form. The margin of the disc grows down and
around the yolk mass and forms a yolk sac; the yolk is gradually absorbed during
development. The eyes arise as ectodermal invaginations. The arms form posteriorly
and then migrate forward to be located around the mouth. The funnel which
represents a part of the foot is located within the mantle cavity being overgrown and
enclosed by the developing mantle.

The Lophophorates
Phylum Bryozoa
(Polyzoa or Ectoprocta) (Sea-mats)

This is the largest and most common of the lophophorate phyla. Bryozoans are
colonial and sessile (although Cristatella is not fixed and capable of slow creeping).
The majorities are marine, attached to rocks, pilings, shells; algae and other animals,
e.g. Bugula. Freshwater forms are widely distributed (e.g. Cristatella, Plumatella).
The body (usually less than 0.5 mm) is box-like, oval, ase-like or tubular in
shape and encased in a protective zooecium made of chitin or cuticle and calcium
carbonate. In some forms however, e.g. Buqula, calcification is very light. There is
an opening, sometimes provided with a lid or operculum, for the protrusion of the
lophophore during feeding.
The lophophore is mostly circular, fan-like when extended (horse-shoe-shaped
in the freshwater forms) and consists of a simple ridge bearing 8-30 or more
tentacles, with the mouth at the base.

88
 The mouth opens into the U-shaped digestive tract. This consists of a pharynx,
stomach (cardia and pylorus) with a caecum and a rectum which opens dorsally by
the anus outside the lophophore (and hence the name Ectoprocta). The gut is
surrounded with a spacious coelom which is divided by a perforated septum into an
anterior mesocoel extending into the lophophore tentacles and a larger posterior
metacoel (trunk coelom) crossed by muscle fibres and one or more tissue cords, the
funiculus.
There are no circulatory, excretory or specialized respiratory or sense organs.
The nervous system consists of a ganglionic mass on the dorsal side of the pharynx
giving a nerve ring surrounding the latter.
Colonies may be stoloniferous (Bowerbankia), plant-like (Bugula) or
encrusting (Membranipora). The zooids of a colony communicate through the
transverse end walls or the lateral walls either by pores (Plumatella) or the transverse
septa are only incompletely formed (Cristatella). Most marine colonies are
polymorphic with normal feeding zooids, autozooids, and reduced heterozooids
modified to serve other functions. Of the common types of heterozooids are the
defensive avicularia in which the operculum is modified as jaws (Bugula).
Freshwater colonies are either lophopodid (Cristatella, Pectinatella) where the
zooids project on one side, or plumatellid (Plumatella) with a more or less plant-like
growth form.

Reproduction:
All freshwater bryozoans and most marine species are hermaphroditic. One to
two ovaries and one to many testes are present without gonoducts. The gametes
rupture into the coelom, or in some cases directly into the sea through a coelomopore
in the region of the lophophore. Eggs are mostly large yolky, few in number and are
brooded in the coelom or externally. Bugula brood its eggs in a special external
chamber called ovicell with a placenta-like connection to provide food for the

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growing embryo. Sperms pass into or out of the zooids through the terminal pores of
a number of tentacles of the lophophore.
Cleavage in marine species is radial and leads to the formation of a larva with a
locomotor ciliated girdle or corona, an anterior tuft of long cilia and a posterior
adhesive sac. Only the larvae of non-brooding bryozoans possess a functional
digestive tract and a long larval life prior to settling. From the first zooid formed
(ancestrula) the whole colony is produced asexually by budding.
The formation of statoblasts throughout summer is another means of asexual
reproduction possessed by freshwater bryozoans. They are special resistant disc-
shaped bodies formed of peritoneal cells that contain stored food and enclosed in
protective chitinous valves. One to several statoblasts develop on the funiculus
bulging into the coelom. They either adhere to the parent colony or fall to the bottom
while others float as they contain air spaces or are armed with hooks (Cristatella).
When environmental conditions become favourable (in spring), the valves separate
and a zooid develop from the internal mass of cells.

Phylum Entoprocta

This is a small phylum (60 species) whose taxonomic status is rather doubtful.
They were formerly included in the bryozoa but were removed in view of the
apparent absence, of a coelom. However some zoologists believe that entoprocts
possess a pseudocoel although their relation with other pseudocoelomates are still
very obscure. Still others are of the opinion that entoprocts are related to other groups
possessing trochophore larvae and probably could have a common ancestry with the
coelomate ectoprocts.
Entoprocts are nearly exclusively marine, mostly colonial, and live attached to
rocks, shell, pilings, or as commensal on sponges, polychaetes, bryozoans and other
animals.

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 The body wall is covered with cuticle. The tentacles and certain areas of the
calyx possess longitudinal muscle fibres. The pseudocoel, which extends also into the
tentacles, is filled with a gelatinous mesenchyme containing both free and fixed cells.
Entoprocts are filter feeders and the digestive tract is U-shaped with a large bulbous
stomach. Excretion takes place by a pair of protonephridia opening with a common
pore just behind the mouth.
The nervous system consists of a single and median ganglion, located between
the stomach and vestibule, and sending nerves to the tentacles, stalk and certain parts
of the calyx. Sensory bristles project over the body surface especially on the
tentacles. Asexual reproduction by budding is common. Buds arise from segments of
the stolon or from the upright branches. In solitary entoprocts, buds develop from the
calyx then separate. Entoprocts include both dioecious and monoecious species with
a common median gonopore just behind the nephridiopore. The vestibule acts as a
brood chamber for the eggs from which trochophore larvae are developed. These
after short free swimming existence settle, attach and metamorphose.

Phylum Brachiopoda

These are coelomic unsegmented marine animals with a bivalve calcareous

shell. Most species live attached to rock or other firm substrata, but some forms
(e.g. Lingula) live in vertical burrows in sand and mud bottoms. Although fossil
species were widely distributed and abundant on reefs, modern forms are largely
inhabitants of cold water.

Shell: Despite the resemblance of brachiopods to molluscs (with which they

were grouped till the middle of the 19th century), this is only superficial, for in
brachiopods the two valves enclose the body dorsally and ventrally instead of
laterally. The vental valve is typically large than the dorsal and is usually attached to
the substratum directly or by means of a cord-like stalk.

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 The apex (beak) of the ventral valve in some groups (e.g. Terebratula) is
drawn out posteriorly and upward as a spout, like a Roman lamp (hence the name
lamp shells). The valves are usually convex but in the burrowing lingulids they are
flattened and more equal in size. The valves may be ornamented with concentric
growth lines and a fluted, ridged or even a spiny surface. The colour of the shell in
most living brachiopods is dull yellow or grey, but some species have orange, red or
brown shells.
The two valves articulate with one another along the posterior line of contact,
called the hinge line and the nature of articulation is the basis for the division of the
phylum into two classes: Inarticulata and Articulata. In the former (e.g. Lingula), the
two valves are only held together by muscles, in the latter the valves bear interlocking
processes (ventral valve with a pair of teeth and oppsing socket on dorsal valve) that
greatly restrict the degree of gape. The two valves are closed by a pair of adductor
muscles and opened by a pair of adductor muscles. The shell is secreted by two
mantle lobes. The inarticulate shell is usually composed of a mixture of calcium
phosphate and chitin (chitinophosphate). The outer surface of the shell is covered by
a thin organic periostracum. As in most molluscs the periostracum and the outer
layers of mineral deposition are secreted by the mantle edge, and the inner layer of
shell is secreted by the entire outer mantle surface. The shells of articulate
brachiopods contain uniformly distributed vertical channels that extend upward
through the shell to a point little below the periostracum. The channels are filled with
mantle tissue, which terminates in a cluster of secretory cells. Such shell structure is
termed punctuate of unknown significance. The anterior space between the mantle
lobes (mantle cavity) is filled by the lophophore, the body proper occupying the
posterior part.
Pedicel: Most brachiopods are attached to the substratum by a cylindrical
extension of the ventral body wall, the pedicel. In Lingula this is long and provided
with muscles, emerging at the posterior end of the animal between the two valves.
When the animal is disturbed the pedical contract and pulls the animal downward into
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the burrow. The peidcel of most other brachiopods is very short, lacks muscles and is
composed primarily of connective tissue. The pedical, moreover, emerges either from
a notch at the hinge line of the ventral valve or through a hole at the upturned apex. A
few brachiopods are deprived of a pedicel.
Lophophore: The lophophore (basically a food-catching organ) surrounds
the mouth, and as a means of increasing the surface area, projects anteriorly as two
arms, or brachia, from which the name brachiopod is derived. In its simplest form,
the lophophore is horseshoe-shaped and the arms are further looped or spiraled in
complicated ways. Each brachium bears a row of hollow tentacles, each containing
an extension of the coelom. The ciliary tracts on the tentacles deive a current of water
through the lophophore, and phytoplankton is collected in this process.
Gut: Mouth leads into an oesophagus, which extends dorsally to join a dilated
stomach surrounded by a digestive gland. In articulates a blind intestine extends
posteriorly from the stomach, but in inarticulates the intestine opens to outside
through a rectum and anus. Digestion in Lingula is intracellular within the digestive
gland.
Circulatory system: Brachiopods possess an open circulatory system with
a contractile vesicle (heart), located over the stomach, and colourless blood.
Respiration: There are no specialized respiratory organs, but the lophophore
and mantle lobes are probably the principal sites of gaseous exchange.
Excretion: One or two pairs of metanephridia are present in brachiopods.
The nephrostome opens in the coelom on each side of the posterior end of the
stomach and the nephridiopore opens in the mantle cavity on each side of the mouth.
Nervous system: There is an oesophageal nerve ring with a dorsal small and
a ventral large ganglion from which nerves extend anteriorly and posteriorly. There is
a pair of statocysts near the anterior adductor muscle in Japanese species of Lingula.
Reproduction: With few exceptions, all are dioecious. Ripe gametes are
shed out through the nephridia. Cleavage is radial and nearly equal. Mesoderm is

93
enterocoelic. There is free-swimming larva, which in inarticulates resembles a minute
brachiopod. Thus there is no metamorphosis. Articulate larvae differ in having a
ciliated anterior lobe (body and Lophophore), a posterior lobe (pedicel) and a mantle
lobe directed backward. The larva undergoes metamorphosis gives the adult form.

Phylum Echinodermata

The echinoderms are exclusively marine, largely bottom dwelling coelomate

invertebrates, which are characterized mainly by their pentamerous radial symmetry.
This, however, has been secondarily derived from a bilateral larva; the echinoderms
are in no way related to the other three radiate phyla (Porifera, Cnidaria and
Ctenophora) of a primary radial symmetry. An internal skeleton composed of
calcareous ossicles is characteristic of all echinoderms. The skeleton typically bears
projecting spines or tubercles, the body thus acquiring a spiny appearance and hence
the name of the phylum.
Echinoderms possess a spacious coelom, well-developed. digestive tract and
nervous system but no excretory organs are present. Nearly all members are
dioecious and fertilization is external. Being the major group of deuterostomes,
cleavage is radial and indeterminate. The blastopore forms the anus; the mouth is a
new structure. Coelom forms within pockets proliferated from the archenteron, i.e.
Enterocoel. Development involves a bilaterally symmetrical larva with ciliated
locomotor bands and complete digestive tract. It later develops long slender arms
which differ in nature and position in the different echinoderm classes. The bilateral
larva after a free-swimming existence metamorphoses into the radially symmetrical
young echinoderm.

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Water-vascular system of Astropecten:
From the madreporite (porous-sieve plate), canals lead down into a vertical
ciliated stone canal and joins with the circular ring. Connected to this ring canal are
five Polian vesicles, as muscular sacs with narrow necks help to regulate the internal
pressure (reservoirs). There are five pairs of blind folded pouches, called Tiedmann's
bodies, producing certain coelomocytes. A ciliated radial canal extends from the ring
canal along each arm. Lateral canals arise from either side of the radial canal to join
with the tube feet, with a valve guarding this connection. The tube foot consists of a
bulbous ampulla, acting as a reservoir, and a hollow highly extensible muscular
podium, which usually terminate in a sucker. The podia project into the ambulacral
groove. The operation of the tube foot is effected by the interaction of the podial
muscles and flow of fluid between the ampulla and podium. The podia serve a variety
of functions, including locomotion, fixation, gas exchange, feeding and sensation.

Classification of Echinodermata:
I- Class: Stelleroidea:
 Star-shaped free-moving echinoderms with the body composed of rays, or
arms, radiating from a central disc.
i- Subclass: Asteroidea (Sea stars or starfish). The members of this
subclass are star-shaped or pentagonal echinoderms with a central disc
and mostly five rays or arms. These vary greatly in length in the
different species and are not sharply marked off from the central disc.
However, a greater number of arms is characteristic of many asteroids,
Leptasterias possesses 6 arms, Luidia may have 7, the sun stars
(Crossaster & Solaster) 7-14, and Pychopodia 15-24 arms. Each arm
contains a prolongation of the coelom and its organs.
 On the aboral (upper) surface there is an inconspicuous anus (absent in some
forms, e.g. Astropecten) and a madreporite. On the oral surface (the ambulacral

95
 grooves. These contain two or four rows of tube feet or podia (with suckers)
which are the locomotory organs and form part of the water vascular system.
The body bears pedicellariae, which are ssessile (O. phanerozonia,
Astropecten) or stalked (Order Forcipulata, Asterias) with two opposing
terminal jaws. Pedicellariae are used in cleaning the body surface or capturing
small prey. Regeneration is exhibited by many asteroids (Linckia, Asterias),
and a number reproduce normally by asexual reproduction.
 With a few exceptions, asteroids are dioeious. Development passes by two
larval stages, a bipennaria larva, which changes into a brachiolaria larva. g.
Iepsld
ii-Subclass Ophiuroidea (Brittle stars or Serpent stars):
 Ophiuroids superficially resemble asteroids but differ in possessing
extremely long arms, which are more sharply demarcated from the
pentagonal or rounded central disc.
 The arms are relatively solid having no prolongation of the ceolom.
There are neither ambulacral grooves nor anus. The mouth and
madreporite are both found on the oral surface. The podia are greatly
reduced, without suckers or ampullae and play little role in locomotion.
Pedicellariae are absent. There are typically five arms, e.g. Ophiura &
Ophiocoma (Ophiactis, with 6) but in basket stars, e.g.
Gorgonocephalus, the arms branch repeatedly to produce a great mass
of coiled tentacles.
 The ophiuroid arms have regenerațing powers but not the disc. The
majorities of ophiuroids are dioecious and possess a larva,
ophiopluteus, with long arms.

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II- Class Echinoidea (Sea urchins, heart urchins, cake urchins):
 The echinoids are typically radial echinoderms, which are deprived of arms but
are globular or greatly flattened along the oral-aboral axis. The skeletal plates
or ossicles are fused to form an internal shell or test.
 In sea urchins (Subclass Regularia), the mouth lies in the middle of the
ambulacral surface while the anus and madreporite are found on the opposite
pole. Amublacral grooves are absent. The body surface is differentiated into
radial areas extending between the oral and aboral poles. Five areas contain
tube feet, and described as ambulacral, alternate with five interambulacral
areas devoid of podia. Movable spines (long or short) are symmetrically
arranged in rows along all areas articulating at their bases with bosses
(tubercles) on the skeletal plates. Pedicellariae are distributed over the general
body surface. They resemble the stalked asteroid pedicellariae but mostly with
three opposing jaws. Some are provided with poison glands. Examples of sea
urchins are Echinus, Echinometra, Arbacia, Paracentrotus.
 The heart urchins and cake urchins (Subclass Irregularia) are all adapted for
burrowing in sand and have much smaller and more numerous spines than in
sea urchins. The heart urchins (O. Spatangoida e.g. Lovenia,
Echinocardium) are more or less oval or roughly heart-shaped are longest
anteroposteriorly. The lower surface is flattened, the upper is convex. The
mouth, with the oral pole has migrated anteriorly. The anus alone has migrated
posteriorly, but the aboral pole remained in the centre of the upper surface. The
oral and aboral ambulacral areas have a flower-like arrangement and are
termed phyllodes and petaloids respectively. Pedicellariae are present but not
of the poisonous type.
 The cake urchins or sand dwellars (O. Clypeasteroida, e.g. Clypeaster) are
typically greatly flattened with a circular ouline. The oral and aboral poles are
central but while the mouth lies in the oral centre, the anus is ventral and
located posteriorly. Sea biscuits are thin and fragile and have a nearly heart-
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 shaped circumference. Sand dwellars (e.g. Mellita) have two to many
elongated holes termed lunules, which are symmetrically arranged. The abroal
surface bears petaloids but the phyllodes are absent and replaced by distinct
radiating grooves. Poison pedicellariae are absent.
 All echinoids are dioecious. The echinoid larva, echinopluteus, differs in
general shape from the asteroid bipennaria but bears great resemblance to the
ophiopluteus of ophiuroids.

III- Class Holothuroidea (Sea-cucumbers):

Like echinoids, the holothuroid body possesses no arms or ambulacral grooves
and has mouth and anus at opposite poles. It is however elongated between the oral
and aboral poles where the ambulacral and interambulacral areas extend
meridionally. Some forms are even worm-like (Synapta). The mouth is bordered by a
circle of tentacles (10-30) which are modified buccal podia (tube feet) (irregularly
branched in Cucumaria). The body wall is highly muscular and the surface is
leathery. The skeleton is reduced to microscopic scattered ossicles; spines and
pedicellariae are absent.
Holothuroids lie with one side of the body against the substratum (but some
burrow in sand and mud). This surface is commonly called the sole comprising three
ambulacral areas (trivium) and two interambulacral areas. In some forms
(Cucumaria) podia are present on both surfaces but suckers are developed on the
sole. In other forms podia, apart from those of the sole, are reduced to water
(Holothuria) or may be completely absent (Psolus). Podia are primarily restricted to
the five ambulacral areas (Cucumaria) but may be scattered allover the surface
(Thyone). They generally tend to become scattered at random. They are totally absent
among burrowing forms as members of the order Apoda (Synapta).
Holothuroids, unlike other echinoderms, possess only a single gonad. They
mostly dioecious and only a few are are hermaphrodites. In the course of
development there is a larval stage, auricularia, which is very similar to the
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bipennaria of asteroids. It is further modified to a barrel-shaped larva, doliolaria,
which through gradual metamorphosis forms the young sea cucumber.
IV- Class: Crinoidea
Sea lilies and sea feathers (or feather stars). The crinoids considered in some
respects to be the primitive echinoderms. They constitute over 650 living species, in
addition to a large number of extinct species. They possess a globoid body which is
formed of crown or corona containing the bulk of the visceral mass which is drawn
into almost solid arms. The aboral side is produced into a cup-like calyx.
The sea lilies are of about 100 species. They are stalked and deep sea
inhabitants. In all crinoids the larval stage has a stalk. The stalk, column or stem may
be attached by a basal disc or root-like branching (in soft sediment) or by basal
adhesion (on hard substrates) or by distally hooked root-like cirri (radicular cirri).
The calyx has an oral flat membranous cover or domed rigid roof called the
tegmen or disc with the mouth is usually located near the centre. The arms arise from
the margin between the calyx and tegmen. They are usually 5-10 cm long, but may
reach 35 cm in some species. In some species the arms are primitively only five and
commonly bifurcate once becoming 10, which may be regarded as the basic pattern
of crinoids, e.g. Antedon, but may, however, divide repeatedly becoming 80-200.
There are pinnules or brachials arise bilaterally from the arms giving them a
feathe-like appearance (and hence the name of the group). The anus located beside
the mouth in the interradius.
Apart from the sessile sea lilies, most extant crinoids are capable of crawling
and intermittent swimming for short distances (e.g. Antedon). Cirri are used for
temporary attachment. During crawling, the arms are bent downward and the animal
moves about on the arm tips. Swimming is accompalished by the alternate upward
and downward sweeps of the arms. Crinoids are suspension feedecs.
Crinoid possess considerable power of regeneration. They swhed arms to
escape predators or when exposed to unfavourable conditions, and these arms are
quickly regenerated. Crinoids are all dioecious, but lack distinct gonads or gonoducts.
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The gametes arise from the peritoneum of special coelomic extensions in the pinnules
called genital canals at the base of the arm. The larval stage is one of vitellaria larvae.

Reproduction and development of echinodersms:

Regeneration following injury, or loss of parts is usually among be most
echinoderms. For regeneration to be take place in starfish, at least a portion of the
central disc must remain intact. In ophiuroids, regeneration of shed arms and asexal
reproduction by fission is common. Echinoids possess a considerable power of
regeneration (in spines, pedicellariae, and pidia), and any damage to the test is
repaired.
While gonads are in pairs in echinoids, holothuroids possess a single gonad,
and no definite gonads in crinoids, where gametes arise from peritoneal extensions.
Sexes are mostly separate. Numerous gametes are shed in the sea and fertilization is
external.
Development is mostly indirect via planktonic larval stages. b Cleavage is
radial and indeterminate (deuterostomia). Gastrulation is very invagination. The
blastopore forms the anus, and the mouth sld develops forward of the blastopore.
Asteroids have two successive larval stages; bipinnaria and brachiolaria. The
latter settles, attaches to the substratum and undergoes metamorphosis. Indirect
development of ophiuroids involves the formation of a vitellaria or ophiopluteus. The
gastrula of Echinoids transforms to an echinopluteus larva ( with 4-6 pairs of long
arms). Holothuroids possess a characteristic auricularia larva, this transforms into a
barrel-shaped doliolaria. Development in crinoids passes by vitellaria which later
settles changing into a stalked pentacrinoid larva.

Phylogeny of the Echinodermata:

The origin of this phylum is abscure, despite the extensive fossil recorded in
the palaeozoic strata. There is common agreement that the echinoderms descended
from a bilateral ancestor, evidenced by this possession of pelagic bilateral larvae,

111
which later metamorphose to the radially symmetrical young. Two theories are
proposed, the dipleurula theory & the pentactula theory. The former theory holds that
the phylum eveolved from a creeping, soft-bodied, bilateral coelomate animal. With
the assumption of a sessile existence, the body of the dipleurula underwent a
clockwise 90° rotation, the animal thus shifting to becomne radial symmetry.
According to the second theory, the common ancestral echinoderm càlled a
pentactula, was a bilateral animal with 5 tentacles around the mouth for capturing
food mus & looked very much like, a lophophore. Each tentacle has an extension of
the coelom, became a radial canal of the water-vasular system. The arms are then
develop, with the addition of protective plates, and then acquired radial symmetry.

Phylum: Chaetognatha
(Arrow worms)...e.g. Sagittai
Chaetognatha comprises about 50 species of common planktonic marine
inhabitants. The body is elongated, unsegmented shape (and hence the name arrow
worms) and is commonly 3cm long (up to 10 cm) the body is divided into head, trunk
(separated by a narrow neck) and a postanal tail. The head is provided on each side
with 4-14 large non-chitinous curved spines, used in siezing the prey (thus the name
of the phylum). A pair of eyes is borne on the dorsal surface. On the posterior half of
the trunk and extending to the tail region there is mostly a pair of horizontal lateral
fins (2 pairs in Sagitta). A large spatula-like caudal fin embraces the end of the tail.
All fins have supporting rays.
In the body wall the stratified epidermis is covered with a thin cuticle and the
longitudinal muscles are arranged in two dorsolateral and two ventrolateral bands
with whose contraction the animal darts swiftly forward, then floats with the help of
the fins.
The compartmented coelom is peculiar in having no peritoneum (resembling a
pseudocoel). It divided by two transverse partitions into three sections. Thus the
single coelomic space of the head is separated from the paired coelomic spaces of the
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trunk (being divided by a ventral and a dorsal mesentery) and these in turn are
separated from the single or paired spaces of the tail.
In the digestive tract the mouth leads to a bulbous pharynx no that penetrates
the head-trunk septum to join the straight intestine with a pair of anterolateral
diverticula. The anus opens at the junction between the trunk and tail. Arrow worms
are carnivorous be voracious feeders. Digestion is probably extracellular.
There are no gas exchange or excretory organs. The coelomic en fluid acts as a
circulatory medium (no circular system). There is a peripharyngeal nerve collar with
a dorsal large cerebral ganglion and his s a number of lateral ganglia.
Chaetognaths are hermaphroditic: There is a pair of elongated ovaries in the
posterior trunk coelom, followed by a pair of testes just behind the trunk-tail septum.
An oviduct runs along the side of each ovary and opens laterally in front of the trunk-
tail septum. The sperm duct leads to a seminal vesicle with which the sperms are
formed into a single spermatophore.
Cross-fertilization is known in some forms (Spadella) where sperm transfer is
reciprocal. As in deuterostomes, cleavage is radial, gastrulation is embolic and the
coelom is enterocoelic in origin. The hatching young's are similar to the adults and
develop without metamorphosis (direct development).

Phylogeny of the Chaetognatha:

The chaetognaths have been linked with the nematodes, molluscs and
brachiopods on no sound bases. Their pattern of embryonic development reflects
their deuterostome position, but unrevealed by their body structure. They, however,
differ from other deuterostomes in the mode of coelom formation (being a modified
form of enterocoely), in lacking circular muscles, in having a subterminal anus, and
in lacking of a larval stage.
Others seek their origin at the roots of the bilateral metazoans.

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 Phylum: Chordata
Subphylum: Urochordata
Ascidians (Tunicates or Sea squirts)

These are almost exclusively marine forms, comprising 90% of the

invertebrate choradates. The most common species are sedentary; some are abyssal
forms and few are pelagic or planktonic. A complex secreted tunic (which contains a
cellulose-related polysaccharide, called tunicin) covers the body. The tunic may be
transparent colourless, purple, red, pink, orange, yellow or green. It may be leathery
or gelatinous, delicate or hard, smooth or hairy, sometimes bearing calcareous
spicules. The body wall is provided with striated longitudinal circular muscle bands,
allowing body contraction. They possess a well-developed perforated pharynx. The
notochord and nerve cord are present only in the larval stage, named tadpole, because
of its superficial resemblance to the amphibian larva.
Ascidians are exist all depths, but are particularly abundant on rocky littoral
habitats. Being the only sessile choradates; they are attached to rock, corals, shells,
ship hulls, often as fouling organisms.
Mostly is solitary (e.g. Ascidia, Ciona, and Styela), attaining a length of few
centimeters. The body varies from spherical to cylindrical in shape; some resembling
peaches or potatoes or is a irregular. They are usually attached to hard substrates and
the free end bears two siphons: anterior buccal, oral or incurrent and anterodorsal
atrial or excurrent. Circular muscles are developed in the siphonal wall, where they
act as ephincters. When disturbed, exposed at low tide or taken out of water,
ascidians contract, becoming turgid, and a jet of water is forced (squirted) out of both
narrowed siphons, and hence the common name, "sea squirts".
Other ascidians live socially as clumps of individuals attached at their bases, or
zooids are joined by stalks. They may form typical colonies or compound forms,
assume flower shaped (e.g. Botryllus).

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 The body can be differentiated into three regions, a pharyngeal, an abdomen
(containing the digestive tract), and a basal postabdomen (contains the heart and
reproductive organs). The coelom is represented by paired epicardial sacs, which
grow posteriorly from the pharynx along each side of the heart. The digestive tract
reflects the filter mode of feeding. The buccal siphon opens by the mouth, which
surrounded by a ring of fleshy tentacles to prevent the passage of large particles.
Water flows through the pharyngeal chamber (basket), pharyngeal slits (stigmata)
into the atrial cavity (or cloaca) which completely surrounds the pharynx. From the
atrium, water is expelled by the atrial siphon. The short oesophagus leads to spacious
stomach (the site of extracellular digestion), into which open the digestive gland. The
intestine, bearing a pyloric gland, forms a U-shape and extends as rectum to open by
the anus in the atrial cavity.
Respiration: Dissolved oxygen diffuses into the body through the pharyngeal
slits.
The heart is U-shaped tube, with no valve, and lies in a pericardial cavity or the
base of the digestive loop. The blood has no respiratory pigments, but containing
several kinds of cells.
Excretion by nephrocytes (or storage cells), where the nitrogenous waste
products are removed by simple diffusion. The nervous system is rather degenerate,
with a brain (cerebral ganglion) located between the siphons.
Regeneration or asexual reproduction by budding, the bud (called blastozooid),
may separate from the parent or remain attached forming complex colony (e.g.
Botryllus).
Most tunicates are hermaphrodites, generally with a single testis and a single
ovary, lying posteriorly near the intestinale loop. The gonoducts run parallel to the
intestine and open in the atrium.
They are mostly oviparous, the fertilization is external (but internal in colonial
forms, where the fertilized eggs are brooded in the atrium till larval hatching). Egg
are coated by special floating membranes.
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 Development is deuterostomous to give a tadpole larva, with a well-developed
muscular tail (which has a dorsal and a ventral fin). The larva bears anteriorly three
adhesive papillae, a dorsal ocellus and a statocyst. The mouth leads to a pharynx with
a few gill slits. A mid-dorsal hollow nerve cord extends through the tail. Three
longitudinal strips form the notochord. The larva settles on the bottom, often about 2
days of swimming, attaching by the anterior papillae and starts metamorphosis
(involving resorption of the neural tube, notochord & tail).

Phylogeny of urochordates:
The urochordates departed early from the chordate line. Most of their adults are
associated with a sedentary mode of life. They, however, differ basically from the
chordates in the entire absence of metamerism and the reduction or loss of the
coelom, subsequent to the development of the large atrium.

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