UNIT- 17

Free Living Larva of Invertebrate

Structure
Objectives
Overview
17.0. Larval forms of free-living invertebrates
17.1. Larval forms of Porifera

17.1.1. Stomoblastula larva

17.1.2. Amphiblastula larva
17.1.3. Coeloblastula and parenchymella larva

17.1.4. Rhagon larva

17.2. Larval forms of Coelenterata
17.2.1. Planula Larva
17.2.2. Scyphistoma Larva
17.2.3. Ephyrae Larva
17.3. Larval forms of Annelida
17.3.1. Trochophore larva
17.4. Larval forms of Arthropoda
17.4.1. Nauplius Larva

17.4.2. Metanauplius Larva

17.4.3. Protozoaea Larva
17.4.4. Zoaea Larva

17.4.5. Cypris Larva

17.4.6. Mysis or Schizopod Larva
17.4.7. Megalopa Larva

17.4.8. Phyllosoma Larva

17.4.9. Alima Larva
17.4.10. Kentrogen Larva

17.4.11. Erichthus & Erichthoidina Larvae

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 17.4.12. Calyptopsis Larva
17.4.13. Epicaridium Larva

17.4.14. Cryptoniscus Larva

17.5. Larval forms of Mollusca
17.5.1. Veliger Larva

17.5.2. Glochidium Larva

17.6. Larval forms of Echinodermata
17.6.1. Dipleurula Larva

17.6.2. Bipinnaria Larva

17.6.3. Brachiolaria Larva
17.6.4. Auricularia Larva

17.6.5. Ophiopluteus Larva

17.6.6. Echinopluteus Larva
17.6.7. Doliolaria Larva

17.6.8. Pentachrinoid Larva

Let Us Sum Up
Check Your Progress

Glossaries
Suggested Readings
Weblink
Answers To Check Your Progress

Objectives

After studying this unit, the students will be able to

• Explain the invertebrate larvae
• List out the invertebrate larva and explain them

Overview

In this unit, we will study about the invertebrate larvae and larval
forms of free-living invertebrates. Here, we will focus on the larval forms
of porifera such as, stomoblastula, amphiblastula, coeloblastula and
rhagon larva, larval forms of coelenterate such as, planula, scyphistoma
and ephyrae larva, larval forms of Annelida such as, trochophore larva,

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larval forms of Arthropoda such as, nauplius, metanauplius, protozoaea,
zoaea, cypris, mysis, megalopa, phyllosoma, alima, kentrogen,
erichthus, erichthoidina, calyptopsis, epicaridium and cryptoniscus larva,
larval forms of Mollusca such as, veliger and glochidium larva, and larval
forms of Echinodermata such as, dipleural, bipinnaria, brachiolaria,
Auricularia, ophiopluteus, echinopluteus, doliolaria and pentachrinoid
larva.

17.0. LARVAL FORMS OF FREE-LIVING INVERTEBRATES

A larva is a distinct juvenile form of many animals undergo before

metamorphosis into adults. Animals with indirect development such as
insects, amphibians, or cnidarians typically have a larval phase in their
life cycle. The larva's appearance is generally very different from the
adult form (e.g. caterpillars and butterflies) including different unique
structures and organs that do not occur in the adult. Their diet may also
be considerably different. Many invertebrates (e.g., cnidarians) have a
simple ciliated larva called a planula. Flukes have several larval stages,
and annelids, mollusks, and crustaceans have various larval forms. The
larval forms of the various insects are called caterpillars, grubs,
maggots, and nymphs.
Pelagic larvae can disperse large distances, colonize new
territory, and move away from habitats that have become overcrowded
or otherwise unsuitable. A long pelagic larval phase can help a species
to break its parasite cycles. Pelagic larvae avoid benthic predators.
Many marine invertebrates undergo indirect development, a kind of life
history that includes a larval stage distinct from the adult. Two types of
larval development are distinguished based on the source of larval
nutrition. Lecithotrophy, meaning “feeding on yolk”, refers to
development with a non-feeding larva, which depends on the egg’s yolk
reserve supplied by the mother. Planktotrophy, meaning “feeding on
plankton” refers to development via a larva that must feed in the
plankton in order to develop to metamorphosis.

17.1. LARVAL FORMS OF PORIFERA

Poriferans are pore bearing animals commonly called sponges,

diploblastic, acoelamate, generally sessile and mostly marine.
Reproduction is by asexual as well as sexual, through a larval form.
Sponges can reproduce asexually. A remarkable property of sponges is
that they regenerate when they are cut into pieces. Each bit of sponge,
however small, will grow into a complete new sponge. Sponges also
reproduce by budding. Sexual reproduction is also common among

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sponges. Most sponges are hermaphrodites, meaning they produce both
eggs and sperm. Since eggs and sperm are produced at different times,
self-fertilization is avoided.

In most species of sponges, sperm cells from one sponge enter

another sponge through its pores, Collar cells on the receiving sponge’s
interior pass the sperm into the mesohyl, where the egg cells reside, and
fertilization occurs. The fertilized eggs develop into larvae and leave the
sponge. Early development occurs in the mesohyl. Cleavage occurs in
zygote. A flagellated larval stage is formed. These larvae may be
parenchymula larva or amphiblastula larva. The larva becomes free. The
water currents carry the larva out of the parent sponge. It freely swims
for two days. Then the larva settles on the substrate and develops into
the adult body form.
Development in syconoid sponges
In syconoid sponge, the larva produced is called stomoblastula,
since it has a mouth and feeds on nurse cells within mesogloea and
grows for a few days. After growing stomoblastula changes into
amphiblastula by inverting inside out bringing the flagellated cells on the
outer surface so that the larva can swim in water. Amphiblastula leaves
the sponge body and swims freely in water feeding on microorganisms.
Gastrulation is by invagination of micromeres, bringing the flagellated
cells again inside the body, lining a cavity which later becomes
spongocoel. Cells on the outer surface transform into pinacocytes.
Gastrula swims about and settles on a rock with blastopore against the
rock and grows to form Olynthus stage that looks like a little sponge. An
osculum is formed later.
Development in asconoid and leuconoid sponges
In asconoid and leuconoid sponges, the blastula is called
coeloblastula as it does not possess a mouth but has a blastocoel and
flagella on the surface of the body. This larva escapes from the sponge
body and swims about freely in water. Gastrulation takes place by
delamination of the archaeocytes which are located on one end of the
blastocoel. The archaeocytes gradually fill the blastocoel completely and
the gastrula becomes solid. This solid gastrula is known as
stereogastrula, parenchymula or parenchymella, which swims about for
some time and then settles on substratum to form olynthus stage. The
inner archaeocytes migrate to form pinacocytes on the surface and the
outer flagellated cells migrate towards inside to form choanocytes lining
the spongocoel.

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 Development with Rhagon larva In Spongilla, the larva is
different from parenchymula and it is called rhagon larva, which has a
tent-like body with a broad flat base called hypophare and a conical
body called spongophare, with a narrow upper end on which is located
the osculum. There are flagellated chambers which open to the outside
by ostia and into the spongocoel by apopyles. The sedentary larva
grows to become adult.
17.1.1. STOMOBLASTULA LARVA

Fig.17.1. Archaeocytes of stomoblastula larva

In this stage 8 rounded and nonflagelated macromers are flanked

over by numerous small, elongated and flagellated micromeres. The
flagella of micromeres face into blastocoel which opens outside through
mouth. Mouth is situated in the centre of macromeres and is used for
engulfing the surrounding amoebocytes for nutrition.
17.1.2. AMPHIBLASTULA LARVA

Fig.17.2. Amphiblastula larva

Amphiblastula is a free-swimming larval stage of Sycon and

many other calcareous sponges that comes out from the mesenchyme
into the spongocoel. It develops from stomoblastula through a process of
inversion. It encloses a small cavity the inside of which is filled with
amoebocytes. It finally passes out from the spongocoel through
osculum. It develops from stomoblastula by inversion, in which it turns
itself out through the mouth, so that the flagella of micromeres become
directed outwards.

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 The body is somewhat ovoid in form and is comprised of an
anterior half of small flagellated micromere cells and a posterior half of
large non-flagellated macromere cells. The embryo is now called
amphiblastula (Gr., amphi, both + blastos, germ,) having two kinds of
blastomeres. Amphiblastula larva swims freely in water for some time.
While swimming, the flagellated pole is directed anteriorly and the force
for swimming is supplied by the beating of flagella. Amphiblastula larva
gives rise to young sponge through embolic gastrulation.
17.1.3. COELOBLASTULA AND PARENCHYMELLA LARVA

Fig.17.3. Parenchymella larva

Coeloblastula does not possess a mouth but has a blastocoel

and flagella on the surface of the body. This larva escapes from the
sponge body and swims about freely in water. Gastrulation takes place
by delamination of the archaeocytes which are located on one end of the
blastocoel. The archaeocytes gradually fill the blastocoel completely and
the gastrula becomes solid. This solid gastrula is known as
stereogastrula, parenchymula or parenchymella, which swims about for
some time and then settles on substratum to form Olynthus stage. The
inner archaeocytes migrate to form pinacocytes on the surface and the
outer flagellated cells migrate towards inside to form choanocytes lining
the spongocoel.

17.1.4. RHAGON LARVA

Fig.17.4. Rhagon larva

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 In Spongilla, the larva is different from parenchymula and
it is called rhagon larva, which has a tent-like body with a broad flat base
called hypophare and a conical body called spongophare, with a narrow
upper end on which is located the osculum. There are flagellated
chambers which open to the outside by ostia and into the spongocoel
by apopyles. The sedentary larva grows to become adult.

17.2. LARVAL FORMS OF COELENTERATA

17.2.1. PLANULA LARVA

Fig.17.5. Planula

Planula is the characteristic larva of phylum coelenterata and

planula is a free-living type of larvae which are produced by the polyp
form and by the medusa form. It is free-swimming or crawling larval type
common in many species of the phylum Cnidaria (e.g., jellyfish, corals,
and sea anemones). The planula body is more or less cylindrical or egg-
shaped and bears numerous cilia (tiny hairlike projections), which are
used for locomotion.
Planula larva may be seen in masses on the oral arms of female
medusae. The planula is set free after a time from the oral arms, after a
brief free-swimming existence it sinks, loses its cilia, the blastopore
closes, and it gets fixed to some object by its aboral end. The planula
will metamorphose into a small polyp or hydra which has no perisarc. In
this metamorphosis, an oral cone or manubrium is formed; the
blastopore opens to become the mouth. Four hollow buds arise per-
radially to become tentacles. Subsequently four interradial and eight
adradial tentacles are formed.
Planula larva of Obelia is similar to that of Aurelia, but there are
some differences like planula of Obeliais without a blastopore and

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coelenteron. It grows into a simple Hydra like hydrula stage which
produces directly by budding a new complete branching Obelia colony,
while planula of Aurelia grows into a trumpet shaped scyphistoma.

17.2.2. SCYPHISTOMA LARVA

Fig.17.6. Scyphistoma larva

The ciliated free swimming planula larva are attached with the
substratum. The cilia are lost and a mouth opens at its free end (distal).
The larva now becomes elongated and metamorphose into a small
trumpet shaped, about 5 mm high, called as hydrula or young
scyphistoma. Its proximal part is narrowed into a stalk like organ,
attached to the substratum by an adhesive basal disc. Tentacles bud out
around the mouth. Thus 16 long and slender tentacles are formed and
the mouth becomes square in shape.
17.2.3. EPHYRAE LARVA

Fig.17.7. Strobila

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 Fig.17.8. Ephyra

The scyphistoma undergoes a remarkable process of budding or

transverse fission called strobilation. The whole body develops a series
of ring like transverse constrictions which gradually become deep and
the organism resembles a pile of minute saucers or discs placed one
above the other. At this stage the scyphistoma with a segmented body is
called a strobila and each of the segments is called ephyra larva. Ephyra
is a young medusoid form with a well developed tetramerous symmetry.
The edge of its umbrella is greatly being produced into 8 arms or 8 bifid
lobes. Each lobe is deeply notched to form a pair of marginal lappets.
Each notch bears a small tentaculocyst between the marginal lappet.
Manubrium with a mouth is present in the middle on the sub umbrellar
surface.

17.3. LARVAL FORMS OF ANNELIDA

Annelidans are protostomate, schizocoelomate, triploblastic,

dorsoventrally flattened and have organ grade of body organization.
Annelid eggs exhibit spiral, and determinate cleavage. In annelids, the
first four cells (blastomeres) give rise, by alternating clockwise and
counterclockwise divisions, to a cap of smaller cells, called micromeres,
at one end of the egg and a cap of larger cells, called macromeres, at
the other end. All cells divide simultaneously during the early stages of
annelid development. A free-swimming immature form called
the trochophore larva develops in the polychaete annelids.
Development in oligochaetes takes place entirely within
the cocoon; there is no free-living larval stage. The cocoon of the aquatic
lower oligochaetes contains large eggs and relatively little albumin. The
cocoon of the terrestrial higher oligochaetes contains small eggs but
large amounts of albumin, which nourishes the developing embryos. The
oligochaetes undergo a highly modified form of spiral cleavage. The
ectoderm, endoderm, and mesoderm, however, arise in the same
manner as in the polychaetes. The elongated gastrula has a ventral
mouth at the front end and a posterior anus. At the gastrula stage,

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earthworms begin to feed on the albumin, the embryo elongates, and the
mesoderm bands break into units to form the walls, or septa, of
individual segments; the worm then leaves the cocoon and begins to
construct a burrow nearby.
17.3.1. TROCHOPHORE LARVA

Fig.17.9. Trochophore larva

The trochophore larva of polychaetes is typically diamond

shaped with a circle of short, hair like projections (cilia), called the
prototroch, around the thickest part of the body. Cells bearing the
prototroch develop from specific micromeres at the 16-cell stage. The
four cells of the annelid cross frequently give rise to a so-called apical
tuft of cilia at the anterior end. A tuft of cilia (the telotroch) may appear
later at the posterior end. The prototroch will become the prostomium
(head) containing the brain, the eyes, and the prostomial appendages, if
present in the adult polychaete. The lower half of the trochophore
contains the digestive tract, excretory organs, and other internal organs;
it is also the site of future segmentation.
The mouth and anus form during the trochophore stage, but the
digestive tract may or may not be functional at that point. The first three
segments form almost simultaneously in the lower half of the
trochophore. The body grows in length by the addition of new segments
from the preanal segment (pygidium or tail), which is the site of all
additional segment formation. The setae generally fall off the first
segment, which becomes the adult peristome.
A fully developed Trochophore larva can be divided into three
regions.

1. Prototrocal region consisting of the apical plate, the protoroch and the
mouth region.

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2. The Pygidium consisting of the telotroch and the anal region behind it.
3. The growth zone.

17.4. LARVAL FORMS OF ARTHROPODA

A few crustaceans are parthenogenetic, either entirely so as

many ostracods, or Cladocera that commonly alternate between
parthenogenetic and sexual means of reproduction. Typically such
partially parthenogenetic forms reproduce asexually during favourable
environmental conditions, and sexually at the onset of unfavourable
conditions. Most crustaceans reproduce sexually. The sexes are
separate, but hermaphroditism occurs occasionally, as for example in
the Cirripedia, Cephalocarida. When the egg hatches, the form which
emerges may be quite like the adult, in which case development is said
to be direct or epimorphic, but, more usually, it is not at all like the adult.
Special terms are applied to the various basic and distinctive stages of
larval development.
Crustaceans are Arthropods whose body is covered with
chitinous exoskeleton for protection. But the same exoskeleton does not
allow body growth and hence must be shed in order to allow growth. The
larval stages feed and grow in order to become adults and must undergo
moulting or ecdysis to grow. After each moulting they change their
structure and size and hence are different from the previous stage.
Therefore, each species of crustaceans demonstrates several
successive larval stages before it becomes adult. Nine important larval
forms are found in Crustacea. They are, Nauplius Larva, Metanauplius
Larva, Protozoaea Larva, Zoaea Larva, Cypris Larva, Mysis or
Schizopod Larva, Megalopa Larva, Phyllosoma Larva and Alima Larva.
17.4.1. NAUPLIUS LARVA

Fig.17.10. Nauplius larva

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 Discovered by Muller in 18th century, the Nauplius larva is the
first fundamental stage in all crustaceans that sometimes hatches from
the egg and sometimes passes inside the egg. Body is oval in shape
and unsegmented with a large cephalothorax and rudimentary abdomen.
It has a broad anterior end with a median eye, large labrum and three
paired appendages. The median eye is characteristic of the nauplius
larva and is often referred to as the nauplius eye. The appendages are
uniramous antennules having two groups of sensory cells forming frontal
organs, a pair of biramous antennae, and a pair of biramous mandibles
for swimming. They have gnathobases directed towards the mouth. It
has a well developed digestive system for feeding on planktons. In
Branchiopoda and Copepoda, Nauplius hatched from eggs.

17.4.2. METANAUPLIUS LARVA

Fig.17.11. Metanauplius larva

Metanauplius is an early larval stage of some crustaceans. In

some Branchiopods, the nauplius larva transforms into metanauplius,
which is slightly larger than nauplius and has cephalothorax and
abdomen and a caudal furca. It also has a single median eye. Antennule
is uniramous and sensory but antenna is large, biramous and
locomotory in function. Mandibles reduce in size and are used for
chewing food. In addition, 2 pairs of maxillae and 2 pairs of maxillipedes
make their appearance in metanauplius for handling food.

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17.4.3. PROTOZOAEA LARVA

Fig.17.12. Protozoaea larva

In the case of marine prawns and lobsters, eggs hatch into

protozoea which has a large cephalothorax and elongated unsegmented
abdomen with a caudal fork and a pair of small uropods. Antennule is
uniramous and segmented while antenna is biramous. There is a single
median eye. Mandibles are small and masticatory in function. There are
2 pairs of maxillipedes for food gathering. Three pairs of thoracic limbs
make their appearance as buds. Cephalothorax is covered by a
carapace. The appendages that appeared in metanauplius become well
developed and functional.

17.4.4. ZOAEA LARVA

Fig.17.13. Zoea larva

Zoea is the common larva of decapods and hence it has

variations in its features in different species. It has a large cephalothorax
that is covered with a helmet-like carapace that also sports spines and it

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protruded into a rostrum in front. It has two stalked compound eyes and
one simple eye. Antennule and antenna are short and are sensory in
function. First and second maxillipedes are large and biramous, used for
swimming. Thorax is un-segmented and thoracic appendages are still in
bud form and non-functional. Abdomen is well formed and six
segmented without appendages and has a caudal furca on the tip along
with a telson. In some Malacostraca, zoea changes in to metazoea,
which grows abdominal appendages for swimming.
17.4.5. CYPRIS LARVA

Fig.17.14. Cypris larva

Cypris larva is covered by a ivalve shell having adductor

muscle. Head has compound eyes, antennules with discs on which
cement glands open, antennae are lost but remaining cephalic
appendages are present, thorax has six pairs of biramous limbs, there is
an abdomen of four segments. It has many adult features. In Cirripedia,
e.g., Lepas, the egg hatches as a nauplius, it changes into a cypris
which gets fixed by discs of antennules with the secretion of cement
glands, then it becomes a pupa which forms shell plates and rotates to
assume the adult form.

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17.4.6. MYSIS OR SCHIZOPOD LARVA

Fig.17.15. Mysis larva

In shrimps and some lobsters zoea transforms into mysis. It has

a cylindrical and elongated body bearing a cephalothorax and six
segmented abdomen. Head and thorax have a carapace Carapace is
produced in front into a pointed rostrum. Antennule and antenna are
sensory in function. There are six pairs of biramous thoracic
appendages for locomotion and six pairs of abdominal appendages for
swimming, out of which the last one is modified as uropod. There is a
pointed telson on the tip of abdomen.
Some crustacean decapods, such as Homarus, Nephrops, have
this larva which is similar to megalopa and mysis. It has cylindrical body
with biramous swimming pleopods on the abdomen and biramous
appendages on cephalothorax.
17.4.7. MEGALOPA LARVA

Fig.17.16. Megalopa larva

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 Megalopa larva has a large un-segmented cephalothorax with all
thirteen pairs of appendages like those of a crab, abdomen is straight
and in line with cephalothorax, it is like the abdomen of prawn with six
pairs of well formed pleopods, In crabs the nauplius is passed in the
egg, it hatches as a zoaea which by moulting forms the megalopa stage,
the megalopa by moulting forms the adult.

17.4.8. PHYLLOSOMA LARVA

In spiny lobsters, the egg hatches into phyllosoma larva in which
body is divisible into head, thorax and abdomen. There is a pair of
stalked compound eyes and a pair each of antennules and antenna as
sense organs. Body is dorsoventrally flattened and transparent. The first
maxillipede is rudimentary and the second one is uniramous. The third
maxillipede is large, biramous and is used for swimming. The abdomen
is small, segmented and does not bear appendages.

Fig.17.17. Phyllosoma larva

Three pairs of thoracic appendages are very long and their tips
are flattened oar-like for swimming near the surface of water.
17.4.9. ALIMA LARVA

Fig.17.18. Alima larva

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 In some lobsters such as Squilla, the zoea is slightly different and
is called Alima larva. It also has a large cephalothorax covered with
carapace and a segmented abdomen bearing paired pleopods and a
telson. Carapace bears spines and rostrum in front. Thoracic limbs are
reduced. The antenna and antennule are uniramous and sensory in
function. The second maxillipedes are prehensile for capturing food. The
larva is pelagic and has a transparent body that makes it invisible to
predators.
17.4.10. KENTROGEN LARVA

Fig.17.19. Kentrogen larva

In Sacculina, cypris larva attaches to the host crab with its

antennule and transforms into kentrogen larva by shedding its bivalve
shell and appendages. Body is sac-like and elongated with
undifferentiated mass of cells and a chitinous tube-like dart within the
body. There are no appendages in this larva.
17.4.11. ERICHTHUS & ERICHTHOIDINA LARVAE

Fig.17.20. Erichthoidina larva

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 These larvae are found in stomatopods of Malacostraca such
as Squilla. They have cylindrical elongated body with cephalothorax and
segmented abdomen. Cephalothorax is covered with a large carapace
that extends forward into a pointed rostrum. Both antennae are
uniramous and sensory in function. Eyes are compound and stalked. In
both these larvae the three anterior thoracic limbs are rudimentary or
reduced and posterior limbs are well-developed for swimming. Pleopods
are very small and hardly functional. In Erichthus maxillipedes are
prehensile and used for capturing food.

17.4.12. CALYPTOPSIS LARVA

Fig.17.21. Calyptopsis larva

This is a larval stage of Euphasia and resembles Erichthoidina in

general shape but does not sport a rostrum on the anterior end of
carapace. Abdomen is segmented but limbless. First antenna is
uniramous and the second antenna is biramous and both of them are
quite well developed. There are 4-5 pairs of biramous thoracic legs for
swimming. Carapace is prominent.
17.4.13. EPICARIDIUM LARVA

Fig.17.22. Epicaridium larva

This is a larva of parasitic isopods such as Bopyrus. Body is oval

in shape with 6 pairs of thoracic limbs hook-like for clinging to the host.
Pleopods are biramous and 6 pairs for swimming. First pair of thoracic
appendage is prehensile. Both the first and second antenna are

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uniramous and sensory in function. One pair of sharp mandibles is
modified for piercing the host. This larva occurs in the gill chambers of
prawns and fishes.

17.4.14. CRYPTONISCUS LARVA

This is the second stage of the larva of Bopyrus that occurs in
the gill chambers of prawns and lobsters. Body is elongated oval with 6
pairs of thoracic appendages modified for clinging on to the host. There
is a single pair of pleopods for swimming. The first pair of antenna is
absent while the second antenna is long and uniramous. Mouth parts
are piercing and sucking type.

17.5. LARVAL FORMS OF MOLLUSCA

In molluscs, the development may be direct or indirect. In the

direct development, there will be no larval stage and the young ones will
hatch out from the eggs which resemble their parents in general
appearance, except the size. The direct development is seen in
cephalopods. In indirect development, a larval stage is seen in majority
of molluscan forms.Three main larval forms are observed in molluscan
forms. There are (i) trochophore (ii) veliger and (iii) glochidium.
17.5.1. VELIGER LARVA

Fig.17.23. Veliger larva

Many gastropod (snails) and bivalve (clams, mussels, oysters

and scallops) molluscs pass through a trochophore larval stage before
developing into veliger larvae. Veligers are planktonic larvae of many
bivalve and gastropod molluscs characterized by a shell, foot,
and velum (a lobed, ciliated structure used for swimming and feeding).
The velum is derived from the prototroch - a pre-oral ciliated band in the
trochophore larva. A dorsal shell gland secretes the shell of the veliger.

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The shell of a bivalve veliger is bivalved while the shell of a gastropod
veliger resembles a spiraled snail shell.
17.5.2. GLOCHIDIUM LARVA

Fig.17.24. Glochidium larva

Giochidium larva of Unio comprises a bivalve shell and a false

mantle lining the shell. Each shell valve is triangular, convex externally
and concave internally and is attached to the other valve on the dorsal
side only. The free ventral ends of each valve are slightly curved inwards
and bear few hooks or spines. The mantle is projected into numerous
protuberances bearing sensory bristles. Between the two valves at the
dorsal side is present a median adductor muscle which helps in closure
of the valves. Below the adductor is present a small median byssus
gland horn which arises a long provisional byssus thread or adhesive
filament. The foot is not yet developed but glandular pouch which
secretes a long sticky thread called larval byssus. With the help of the
byssus it gets attached itself to the skin or gills or fins of a fish and leads
a parasitic life for about 15 weeks to get their nourishment from them
and live as a ectoparasite on fishes.

17.6. LARVAL FORMS OF ECHINODERMATA

Echinoderms are unisexual animals. Sexual dimorphism is

absent. Fertilization takes place in water. The development may be
direct or indirect. If the development is indirect it includes larva stages. In
different classes of echinoderms, different types of larvae complete the
development. The larval form is bilaterally symmetrical. It undergoes
metamorphosis and radial symmetrical adult is developed.

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17.6.1. DIPLEURULA LARVA

Fig.17.25. Dipleurula larva

It is the first larval stage in all echinoderms. It is an egg shaped

and bilaterally symmetrical. An anterior mid ventral ectodermal
invagination called stomodaeum becomes continuous with the
archenteron and form a larval mouth. The blastopore becomes anus
(deuterostomium). The archenteron is differentiated into a digestive tract
which is made up of oesophagus, stomach and intestine. It has perioral
band surrounding the mouth and an adoral band lying inside the mouth.
The larvae feed actively on unicellular algae. The food is collected from
the currents produced by the stomodaeal cilia.
17.6.2. BIPINNARIA LARVA

Fig.17.26. Bipinnaria larva

It is the larva form of Star fish. The fertilised egg is homolecithal.

It undergoes holoblastic cleavage and develops into blastula and
gastrula stages. The gastrula elongates in length and it gives rise to
Bipinnaria larva. It is a bilaterally symmetrical free swimming pelagic
larva. It has elongated pre-oral region and broad post-oral region.
The anterior end forms pre-oral lobe. The ciliated band at the pre-oral
lobe forms into 2 separate bands, Pre-oral band of cilia, and post oral
band of cilia. These 2 bands of cilia are drawn into many arms. They are
ventro-median arm, a pair of pre-oral arm, median dorsal arm,a pair of
antero-dorsai arm, pair of posterio-dorsal arm,a pair of posterio-lateral
arm, and a pair of post oral arm. The digestive system is developed with

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mouth and anus. This larva slowly grows s into the next larval form
called Brachiolaria larva.
17.6.3. BRACHIOLARIA LARVA

Fig.17.27. Brachiolaria larva

Bipinnaria larva swims for few weeks in the sea water.lt finally
transforms into next larval stage called Brachiolaria larva. It is bilaterally
symmetrical larva. It is pelagic larval form, it shows 3 brachiolar arms
with suckers. They are one median and two lateral in position. At the tip
of brachiolar arms adhesive structures will make their appearance and
they are for attachment. The larva shows all the arms that are seen in
the Bipinnaria, but these arms are very long and hanging. These ciliated
arms will be helpful for swimming in the water. The digestive system is
completely developed with definite stomach and intestine. This larva
undergoes metamorphosis and develops into an adult.
17.6.4. AURICULARIA LARVA

Fig.17.28. Auricularia larva

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 This larval form is present In Holothuroidea. It is a free swimming
pelagic larva. Arms are absent. Alimentary canal is developed. It opens
with mouth and ends with anus. Intestine is curved. Usually this larva is
1 mm in length. Ciliated bands are well-developed. Ciliated band
continues through oral loop and anal loop.
17.6.5. OPHIOPLUTEUS LARVA

Fig.17.29. Ophiopluteus larva

This larva is seen in the life history of opuriodea (Brittle star). It

shows many long arms. It is bilaterally symmetrical. It is transparent. It is
Pelagic. The arms are supported by calcareous rods. The arms are
directed upwards. Preoral loop is reduced. Ciliated band is undivided.
The postero-lateral arms are very long and they are directed forwards.
The digestive system is developed. It opens with mouth and ends with
anus. This larva swims for sometime before undergoing metamorphosis.
17.6.6. ECHINOPLUTEUS LARVA

Fig.17.30. Echinopluteus larva

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 It is seen in the life history of Echinoidea. It is a microscopic
larva. It swims in water. This larva shows ciliated bands which are
developed into arms. Fully developed echinopluteus larva has 4 or 5
pairs of arms. Usually 6 pairs of arms should be formed. The arms are
supported by (CaCO3) Calcareous rods. The digestive system is
developed which shows mouth and anus. It develops hydrocoel and
vestibule. These parts grow on the oral side of the animal. From the
hydrocoel five radial canals will develop. This larva undergoes rapid
metamorphosis and develops into an adult.

17.6.7. DOLIOLARIA LARVA

Fig.17.31. Doliolaria larva

Crinoidea group of animals have Doliolaria larva form. It is a free

swimming larval form. It contains an apical tuft of cilia which will be
sensory. On the mid ventral line near apical plate adhesive pit will be
present. The body shows 4 or 5 ciliated bands. In between 3rd and 2nd
ciliated bands vestibule is present.
17.6.8. PENTACHRINOID LARVA
When doliolaria larva attaches with the substratum, internal
organs rotate at an angle of 90 degrees from ventral to posterior
position. Now the larva with a stalk is known as pentachrinoid larva. This
larva is also called as cystidean larva as it has resemblance with the
extinct cystids. At first it can't feed for the stomodaeum is closed but
within a few days it opens up. Finally, after some further months of
sessile life it breaks away from the stalk and begins the free swimming
life characteristic of crinoids.

Let Us Sum Up

In this unit, we studied about the invertebrate larvae and larval

forms of free-living invertebrates. Under this unit, we focused on the
larval forms of porifera such as, stomoblastula, amphiblastula,
coeloblastula and rhagon larva, larval forms of coelenterate such as,
planula, scyphistoma and ephyrae larva, larval forms of Annelida such

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as, trochophore larva, larval forms of Arthropoda such as, nauplius,
metanauplius, protozoaea, zoaea, cypris, mysis, megalopa, phyllosoma,
alima, kentrogen, erichthus, erichthoidina, calyptopsis, epicaridium and
cryptoniscus larva, larval forms of Mollusca such as, veliger and
glochidium larva, and larval forms of Echinodermata such as, dipleural,
bipinnaria, brachiolaria, Auricularia, ophiopluteus, echinopluteus,
doliolaria and pentachrinoid larva.

Check Your Progress

1) A larva is a distinct juvenile form of many animals before

__________into adults.
2) Many invertebrates have a simple ciliated larva called __________.
3) In asconoid and leuconoid sponges the blastula is called
__________.
4) The scyphistoma undergoes a remarkable process of budding called
__________.
5) Crustaceans are arthropods whose body is covered with
__________.

Glossaries

Pelagic : Relating to, or living or occurring in the open sea.

Sessile : Fixed in one place or immotile.

Suggested readings

1. MOORE, R.C., LOLICKER and FISCHER, A.G. (1952), Invertebrate

Paleontology, McGraw Hill Book Co., Inc., New York.
2. HIGHNAM, K.C. and HILL, L. (1979). The Comparitive Endocrinology
of Invertebrates, ELBS & Edward Amold (Publishers) Ltd., London.

Weblink

Invertebrate larvae https://youtu.be/GWIk4FyPRrk

Free living larval forms of invertebrates https://youtu.be/FsFvqhyZQCQ

Answers to check Your Progress

1) Metamorphosis
2) Planula
3) Coeloblastula
4) Strobilation
5) Chitinous exoskeleton

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 Unit- 18
Parasitic Larva of Invertebrate
Structure
Objectives
Overview
18.0. Larval forms of parasitic invertebrates
18.1. Larval stages of Fasciola hepatica

18.1.1. Miracidium Larva

18.1.2. Sporocyst Larva
18.1.3. Redia Larva

18.1.4. Cercaria Larva

18.1.5. Metacercaria
18.2. Larval stage of Taenia solium
18.2.1. Muller’s larva or Protrochula
18.2.2. Hexacanth and Onchosphere larva
18.2.3. Cysticercus or Bladderworm Larva
Let Us Sum Up
Check Your Progress
Glossaries

Suggested Readings
Answers To Check Your Progress

Objectives

After studying this unit, the students will be able to

• Explain the larval forms of parasites
• List out the various forms parasitic larva of invertebrates and
explain them

Overview

In this unit, we will study about the larval forms of parasites in

invertebrates. Here, we will focus on the larval stages of Fasciola
hepatica such as, miracidium, sporocyst, redia, cercaria and

219
metacercaria larva, and larval stage of Taenia solium such as, muller’s,
hexacanth, onchospere and cysticercus larva.

18.0. LARVAL FORMS OF PARASITIC INVERTEBRATES

A parasite is an organism that lives on or in a host organism and

gets its food from or at the expense of its host. There are three main
classes of parasites that can cause disease in humans: protozoa,
helminths, and ectoparasites. The process of larval maturation in the
host can take from about two weeks up to four months, depending on
the helminth species. Ascaris spp., Enterobius, Filarioidea, Onchocerca
spp., Rhabditis spp., Trichuris spp., Necator americanus, Ancylostoma
spp.
The helminths are invertebrates characterized by elongated, flat
or round bodies. In medically oriented schemes the flatworms or
platyhelminths include flukes and tapeworms. Roundworms are
nematodes. These groups are subdivided for convenience according to
the host organ in which they reside, e.g., lung flukes, extraintestinal
tapeworms, and intestinal roundworms. Platyhelminths are acoetomate,
triploblastic, Dorso ventrally flattened organisums and have organ grade
of body organization, mostly parasitic and rarely free living. Due to
endoparasitic in nature, their life cycle is very complicated and
completed through various larval stages.

18.1. LARVAL STAGES OF FASCIBLA HEPATICA

18.1.1. MIRACIDIUM LARVA

The miracidium is a ciliated, non feeding larva. It is the
first larval stage of trematode worms (flukes), which hatches from eggs
excreted in the faeces of the primary host. Its body is conical and flat.
This larva bears uniform ciliation all over. At the anterior end the body is
projected into a small and pointed apical papilla. Body is covered with
epidermal plates arranged in 5 rows. A large pitcher-shaped
multinucleated apical gland is present in the centre of apical papilla and
1 to 3 pairs of penetration glands around it. A tetra radiate brain and a
pair of small and blackish eye spots are present above the penetration
and apical glands.

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 Fig.18.1. Miracidium larva

The body wall is comprised of an outermost ciliated, a glandular

epidermis, a well represented muscular layer and a layer of
mesenchyme cells filling the whole body space. A pair of flame cells is
present in the posterior half of the body, it is connected with long
excretory ducts opening to outside through excretory pores. The inside
region of the larva consists of rounded germ balls in different sizes.
These give rise to further larval stages. The presence of apical gland,
penetration glands and ciliated ectoderm are all larval parasitic
adaptations.
18.1.2. SPOROCYST LARVA

Fig.18.2. Sporocyst larva

Sporocyst is developed from the miracidium larva in the

pulmonary chamber of snail. It looks like an elongated sac about 7mm
long. The hexagonal cells and cilia which cover the body of miracidium
are shed. Apical gland, cephalic glands, brain, eye spots and primitive

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gut of miracidium are degraded. It is covered by a cuticle membrane,
body wall which consists of sub-epithelial cells, muscles and
mesenchyme. Body sacs have flame cells and germ cells. It is a non-
feeding stage. Germ cells within the sporocyst give rise to the next larval
stage known as redia larva which develops within it. One sporocyst
many give rise to 5-6 redia These germs cells multiply and give rise to
next larval stage known as Redia Larva.
18.1.3. REDIA LARVA

Fig.18.3. (A) Redia with daughter Redia (B) Redia with Cercaria

Redia develops from the germ cells of the sporocyst and comes
out of the sporocyst by rupturing the sporocyst wall. Redia then migrates
to the liver of the snail. Each redia measures about 1.3-1.6 mm in length.
Body of redia is elongated, cylindrical and sac-like. Body-wall is
composed of tegument, epithelial layer and delicate mesenchyme.
Anterior end consists of mouth which leads into a muscular pharynx with
pharyngeal glands and sac like intestine. Just behind the pharynx is a
muscular ring-like swelling called collar, which helps the redia in
locomotion. Just posterior to the collar is a permanent aperture called
birth pore through which the next larval stage of the cercaria exits
outside. Posterior region has two stumpy processes called lappets which
help the redia in anchoring to the tissues of the snail and are also helpful
in locomotion. The space between the body wall and intestine contains n
few germ cells. Germ cells often give rise to second generation of
daughter rediae, it gives rise to the next larval stage called cercaria.

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18.1.4. CERCARIA LARVA

Fig.18.4. Cercaria larva

Cercaria larva is the fourth larval stage in the lifecycle of F.

hepatica. It is a free-living stage produced by the redia larva. It has a flat
and oval body about 35 mm in length and a long tadpole like tail.
Cercaria moves by muscular undulations of the tail. Cercaria has two
suckers an anterior, oral sucker surrounding the mouth and a ventral
sucker situated in the middle of the body. Body space is filled with
parenchyma and contains a few cystogenous glands on each side which
form the cyst of future larva.

It has alimentary canal consists of mouth, muscular pharynx,

esophagus and bifurcated inverted Y-shaped intestine. It also possesses
an excretory bladder with a pair of protonephridial canals and a number
of flame cells. Cercaria also has two large non-functional penetration
glands as well as rudiments of reproductive organs which have
originated form germ cells. Cercaria is a young fluke of sexual
generation. Cercaria larva first comes out of the redia through its birth
pore. The most important feature of the larva is the presence of
numerous cytogenous glands, which are distributed among the flame
cells and help in secreting a cyst in order to transform the larva into next
stage called metacercaria.

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18.1.5. METACERCARIA

Fig.18.5. Metacercaria

The metacercaria larva is the infective stage for the final host, the
sheep which gets the infection when it swallows the metacercaria larva
along with its food. Metacercaria is the 5th larval stage of F.hepatica. It
is 0.2 mm in size and is called a juvenile fluke. It is different from the
cercaria larva in having a thick hard cyst and large numbers of flame
cells; it lacks tail and cystogenous gland cells. Its excretory bladder
opens directly through a single pore. Germ cells or the genital rudiments
are present.

The cyst of metacercacia provides protection against short period

of desiccation. Its further development takes place in tlie wall final host
the sheep. Metacercaria can also infect humans. This larva develops
into adult only inside its definitive host. Its cyst wall is dissolved in the
proximal part of the host intestine enzymes and then the larva is
liberated into coelomic cavity. Now it infects the liver, feeds on its tissue
and grows in size in five to six weeks. Then takes up its position in the
bile duct, where it finally attains a sexual maturity and starts laying eggs
(capsules).

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18.2. LARVAL STAGE OF TAENIA SOLIUM

18.2.1. MULLER’S LARVA OR PROTROCHULA

Fig.18.6. Muller’s larva

Fertilization is internal and fertilized, shelled eggs containing

embryos are shed to the exterior. Acoela and Polyclads lack yolk glands
and development is direct. In Polyclads a free-swimming larva is
produced which is supposed to foreshadow the Trochophore.
The larva is called Muller’s larva or Protrochula. It is oval in
shape and bears eight prominent arms which are beset with long cilia
forming one continuous band. General body surface is covered with
small cilia.

The mouth aperture is located in the mid-ventral line and three

eyes exist in the anterior part of the dorsal surface. During development
the ciliated arms are absorbed. In other groups the embryonic
development is greatly modified and complicated.
18.2.2. HEXACANTH AND ONCHOSPHERE LARVA

Fig.18.7. Hexacanth larva

During the embryonic development of Taenia solium, morula at

its morphologically posterior end, develops three pairs of chitinous hooks

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secreted by differentiated cells, called onchoblasts. This six hooked
embryo, called hexacanth, possesses a pair of large penetration glands.
It is surrounded by two hexacanth membranes. The hexacanth, together
with all the membranes surrounding it, is known as onchosphere. By the
time onchosphere formation, the mature proglotid of tape worm detach
(apolysis) and passes out with hosts (man) faeces. When the secondary
host (pig) feeds human excreta, it gets infected with onchosphere larva.
18.2.3. CYSTICERCUS OR BLADDERWORM LARVA

Fig.18.8. (a) Cysticercus larva with invaginated scolex. (b) Cysticercus

with evaginated scolex

Bladder worm consists of a single layered, large ovoid sac or

bladder-like structure with an invaginated proscolex which measures 6-
18 mm in length at the anterior end. Proscolex contains suckers, hooks
and rostellum. Bladder is generally filled with clear watery fluid having
mostly plasma of the host. Bladder wall consists of an outer cuticle and
inner mesenchyme. Further development of cysticercus larva takes
place only when it is eaten by a human host, otherwise it lodges in the
tissue of the host and gets calcified. After entering the stomach of the

226
human host, the proscolex gets evaginated, suckers and rostellum come
out and anchor to the mucous membrane. Then this larva develops
further into the adult by proliferation.

Let Us Sum Up

In this unit, we studied about the larval forms of parasites in

invertebrates. Under this unit, we focused on the larval stages of
Fasciola hepatica such as, miracidium, sporocyst, redia, cercaria and
metacercaria larva, and larval stage of Taenia solium such as, muller’s,
hexacanth, onchospere and cysticercus larva.

Check Your Progress

1) There are __________ main classes of parasites that cause disease

in humans.
2) The __________ is a ciliated, non-feeding larva.
3) The redia develops from the germ cells of the __________.
4) The muller’s larva is also called __________.

Glossaries

Sporocyst : It is a diploid cell that divides by meiosis to

produce four haploid spores.
Tegument : It is the integument of an organism, especially
a parasitic worm.
Cercaria : It is the larval form of the trematode class of
parasites.

Suggested readings

1. MOORE, R.C., LOLICKER and FISCHER, A.G. (1952), Invertebrate

Paleontology, McGraw Hill Book Co., Inc., New York.
2. HYMAN, G.H (1967) the invertebrates, Vol. I to VII, McGraw Hill Book
Co., Inc., New York.

Weblink

larval forms of parasites https://youtu.be/93yLDrShEpQ

Answers to check Your Progress

1) 3
2) Miracidium
3) Sporocyst
4) Protochula

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 Unit- 19
Evolutionary Significance of Larva
Structure
Objectives
Overview
19.0. Evolutionary significance of larval life
19.1. Evolutionary significance of Planula larva in Cnidaria

19.2. Evolutionary Significance of Platyhelminthes larva

19.3. Evolutionary significance of Crustaceae larva
19.4. Evolutionary significance of trochophore larva

19.5. Evolutionary Significance of echinoderm larvae

Let Us Sum Up
Check Your Progress
Glossaries
Suggested Readings
Weblink
Answers To Check Your Progress

Objectives

After studying this unit, the students will be able to

• Describe the strategies of larval forms
• Explain the evolutionary significance of larval forms

Overview

In this unit, we will study about the strategies and evolutionary

significance of larval forms in invertebrates. Here, we will focus on the
evolutionary significance of larval life such as, planula larva in cnidaria,
Platyhelminthes larva, crustacea larva, trochophore larva and
echinoderm larva.

19.0. EVOLUTIONARY SIGNIFICANCE OF LARVAL LIFE

Larvae represents one of the classic problems of evolutionary

biology and may explain how new body plans originate. It has often been
suggested that many unique body plans first originated as larvae of

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ancestral organisms. There is firstly the need of a delicate young
organism to grow in conditions which satisfy its special requirements and
avoid unnecessary competition with the adult, secondly there is the need
to provide for dispersal of the species and thus to avoid overcrowding.
Finally there is the need to select its habitat that is suited to the
requirements of the adults. Because of these varied functions larvae
often become very highly specialized and all of them undergo some
degree of metamorphosis. According to the biogenetic law or
recapitulation theory of Haeckel, every organism during its development
(ontogeny) repeats to some extent its evolutionary history (Phylogeny).
In other words, successive stages of individual development correspond
with successive adult ancestors in the line of evolutionary descent.

19.1. EVOLUTIONARY SIGNIFICANCE OF PLANULA LARVA IN

CNIDARIA

The alternation of a sessile polyp and a planktonic medusa

represents the phylogenetically primary mode of life cycle in the
medusozoa. The polyp is involved in asexual reproduction, whereas the
medusa generates planula larva by reproducing sexually. Thus planula
larva and medusa both contribute to species dispersal.
The origin of the cnidarians and ctenophores is obscure,
although the most widely supported hypothesis today is that the radiate
phyla arose from a radially symmetrical, planulalike ancestor. Such an
ancestor could have been common to the radiate and to the higher
metazoans, the latter having been derived from a branch whose
members habitually crept about on the sea bottom. Such a habit would
select for bilateral symmetry. Others became sessile or free floating,
conditions for which radial symmetry is a selective advantage. A planula
larva in which an invagination formed to become the gastro vascular
cavity would correspond roughly to a cnidarian with an ectoderm and an
endoderm.

Fig.19.1. Ancestor of acoela

Some researchers believe trachyline medusae (an order of class

Hydrozoa) resemble the ancestral cnidarian because of their direct
development from planula and actinula larvae to medusas. The

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trachylinelike ancestor would have given rise to other cnidarian lines
after the evolution of the polyp stage and alternation of sexual (medusa)
and asexual (polyp) generations.
19.2. EVOLUTIONARY SIGNIFICANCE OF PLATYHELMINTHES
LARVA

Polyclad flatworms offer an excellent system with which to

explore the evolution of larval structures and the ecological and
developmental mechanisms driving flatworm and marine invertebrate life
history evolution. Several conflicting views have been put forward about
the origin of the Platyhelminthes, but modern opinion is generally divided
between two main theories, a ciliate theory and a planula theory. The
early embryonic development of some polyclads follows a pattern of
quartet-producing spiral cleavage that is very similar to that in the
Nemertini, Annelida and the Mollusca, and chiefly for these reason
flatworms, nemertines, annelids and molluscs are thought all to have
arisen from early archoophoran ancestors.
In more advanced cestodes, the life cycle would no longer
depend upon the fortuitous hatching of ingested eggs in the gut of the
intermediate host; the hatching may have been controlled by the
parasite losing its tanning system and producing eggs with digestible
shells. At the same time the shell could have become protected from
digestion by the host of the parent worm if the eggs were retained in a
swollen saclike uterus, e.g. in the bothriocephalideans. In a diverging
line (e.g. tetraphyllideans, tetrarhynchideans, protocephalideans and
cyclophyllideans) the uterus even lost its external opening.
In those cestodes that retain the eggs in the uterus, egg-
dispersal became effected by the apolytic habit of the tapeworm.
Subsequently the various tapeworms diverged from each other by
following different lines in host evolution, the bothridiocephalideans
maintaining an association with the line that has led to teleost fishes,
tetraphyllideans and tetrarhynchideans with the line that has led to
elasmobranchs, and protocephalideans and cyclophyllideans
maintaining an association with the line that has led to modern
amphibians, reptiles, birds and mammals.
There is no unanimous view among zoologists regarding the
origin of flatworms. There are several views for the origin of flatworms.

Lang (1881) proposed ctenophore- polyclad theory. This theory

states that polyclad turbellarians might have evolved from ctenophores

230
through Platyctenea. The Muller’s larva of polyclad bears some re-
semblances with ctenophores.
Planula larva theory states that turbellarians bear some
resemblances with the planula larva of cnidarians. So many zoologists
such as Hyman (1951), Jagersten (1955) and Hadzi (1963), consider
that flatworms have evolved from a planula-like ancestral stock and
Acoela is the most primitive group among platyhelminths.
Molecular data and cladistic analyses have been suggested a
monophyletic origin for the three parasitic groups. The three parasitic
classes—Trematoda, Monogenea and Cestoda, have evolved from free-
living turbellarians.

Fig.19.2. Possible phylogenetic tree of platyhelminthes

The enterocoel theory states that all bilateral animals are

basically coelomate, and acoelomate flatworms have evolved from
coelmate ancestors by loss of the cavity secondarily. This theory is not
acceptable to the zoologists because it is difficult to explain some
aspects, such as the change from coelomate to acoelomate condition
and from bilateral to radial symmetry.

19.3. EVOLUTIONARY SIGNIFICANCE OF CRUSTACEAE LARVA

According the biogenetic law proposed by Haeckel, ontogeny

recapitulates phylogeny; the successive stages of individual
development correspond with successive ancestors in the line of
evolutionary descent. Nauplius larva occurs in the development of all the
crustaceans and so it was considered as the ancestral form of
crustaceans. The old idea of recapitulations stands greatly modified
now-a-days and the crustacean larval forms are now regarded to be the
larval reversions of simpler crustacean ancestors. The larval forms are

231
useful for finding out homologies and the affinities among various
groups. The animals which pass through similar stages are closely
related. Larvae are helpful in wide range distribution of species and also
in keeping the food reserves of eggs to a minimum.

19.4. EVOLUTIONARY SIGNIFICANCE OF TROCHOPHORE LARVA

The Mollusca, though a highly diversified group of invertebrates,

nevertheless form an extremely well-defined phylum. In the evolutionary
dynamics of invertebrates the trophophore larva occupies a prominent
status. It shows similarities with many invertebrate groups. The affinities
throw light on the emergence of bilateral groups from the animals having
radial symmetry. If anatomical features of recent Mollusca are to be
used, as they must, as the clue to phyletic ancestry, it is necessary to
choose from three characters which are not unique to the group.
The first character is the controversial one, viz. metamerism.
Even if the view is accepted, based on the studies on Neopilina, that the
Mollusca are basically a segmented group, this achieves little more than
to suggest a reasonably close affinity of the phylum with the Annelida
and Arthropoda; it does not help us with a concept of the ancestral
mollusc. Similarly, if the occurrence of a trochophore larva is evoked as
a character significant as a guide towards the ancestry of the group, it
must not necessarily be regarded as leading back to the Annelida.

Even though the two groups generally share spiral cleavage as

an early embryological feature and their trochophores are superficially
similar, there are basic features of difference, notably the 'molluscan
cross' and the 'annelidan cross', and the immediate posttrochophore
stages. Whereas the annelid post-trochophore shows a posterior
elongation and signs of segmentation, its molluscan counterpart
becomes a characteristic veliger whose early features are a shell-gland
and a velum. And again, even if the trochophores reflect a close affinity
between the two phyla, they do not lead us easily to the common
ancestor.

Fig.19.3. Phylogenetic relation of trochophore larva

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 It is claimed that the trochophore represents a transitional stage
in the line of emergence of the bilateral groups (e.g., Rotifers) from the
radial groups (Ctenophores). Similarities between the trochophore and
the echinoderm larva (Bipinnaria and Pluteus) and Tornaria larva of
Balanoglossus added more weight to this contention. Many workers
stated that the Trochophore larva serves as a bridge between radial and
bilateral symmetry. They have opined that the bilateral symmetry has
evolved from the radial one. Regarding their views; there are many
theories which may be given below,

1. Ctenophore - Polyclad theory (proposed by Lang, 1881)

2. Ctenophore - Trochophore theory, later modified by Hatschek (1878)
3. Planuloid - Coeloid theory (proposed by L. Vongraff, 1882).

Second theory is more or less acceptable by the researchers.

According to this theory Trochophore larva arose from the hypothetical
animal Trochozoon. Again, Salvini Plawen (1973) has stated that
annelids and echiurans are closely related by their larval stage
(trochophore larva) whereas flatworms, nemerteans and entoproct
larvae are unrelated.
Mollusca were derived from turbellarian-like ancestors, the
conversion of a larva like Muller's larva into a molluscan trochophore
must also be postulated. The origins of an anus, coelom and blood
vascular system remain as obscure as they do for the annelids and
arthropods. The most notable Mollusca, from nearly all points of view,
are the Cephalopoda which form a singularly isolated group. Despite
their general plan of organisation and, in particular, the possession of an
odontophore apparatus, they are separated sharply from all other
molluscs in their production of yolk-laden eggs and the subsequent
absence of free larval stages. While their evolutionary history from
Cambrian nautiloids to the present-day coleoids is fairly well
documented, their origin is obscure.

19.5. EVOLUTIONARY SIGNIFICANCE OF ECHINODERM LARVAE

All the larval of echinoderms have a bilateral symmetry. Hence it

is believed that the ancestor of echinoderms was a bilaterally
symmetrical animalThe auricularia larva presents very close and striking
resemblances to the tornaria larva of some entercoelous development
parallels that found in primitive chordates. Hence echinoderms and
chordates have long been regarded as related groups.

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 Bather (1900) claimed common ancestry of hemichordates and
echinoderms from the dipleurula larva. Muller and Bateson again
claimed that the tornaria larva and dipleurula larva had evolved form a
common ancestral source.De Beer and Garstang hold that the tornaria
larva, the dipleurula and pluteus larvae are living relics from a very
remote period when the echinoderms and chordates were not diverged.

Fig.19.4. Phylogeny of chordata

The neotenous theory propounded by Garstang in 1894, 1928

holds that the chordates arose from some neotenous form of auricularia
larvae.
The sequence is—Auricularia larva (Echinodermata) → Tornaria
larva (Hemichordata) → Tadpole (Ascidia) → Neoteny → Free
swimming Chordates.

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 Fig.19.5. Derivation of prochordate from echinoderm

The adoral ciliated band of auricularia presumably functions to

convey food particles in the oral aperture, and for this it has been con-
verted to the floor of the pharyngeal cavity. Garstang suggests that
endostyle was derived from this loop of the adoral band of the auricularia
larva. Most of the modern workers like Berrill (1955), Bone (1979) have
supported the main theory of Garstang. The tornaria larva of
Hemichordate is much more similar to the larvae of echinoderms but
differs in its mode of origin and its structure. Young (1981) supported the
neotenous larval theory of Garstang.
Therefore, from the above discussion it may be concluded that
the phylogenetic relationship between echinoderms, hemichordates and
chordates is more convincing, and chordates may have evolved from
non-chordates, probably the echinoderms. But Fell (1963), Pawson and
others deny the ancestry of chordates from any form of echinoderms
arid they also deny the relationship between the tornaria larva and the
echinoderm larvae. Many counter-arguments have been put forward by
them.
1. Protocoel is unpaired in Balanoglossus, but paired in echinoderms.
2. Extant echinoderms lack pharyngeal gill-slits.
3. Bipinnaria larva lacks telotroch.
4. The tornaria larva has a characteristic apical plate with eye-spots.
The study of larval forms and adults of echinoderms reflects a
case of retrogressive metamorphosis because the adult echinoderms
possess many primitive features than the larval forms. The features of
adults are comparable with lower invertebrates (e.g., Cnidaria), such as
radial symmetry, lacking of distinct head, absence of anterior and
posterior sides, etc. and the advanced larvae of echinoderms
metamorphose into primitive adults. Hence it is an example of
retrogressive metamorphosis.

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VIEWS REGARDING THE LARVAL PHYLOGENY
MACBRIDE (1914):
Dipleurula larva was of fixed type and gave rise to the free-
swimming forms of Antedon or Yolk larva.
HYMAN’S SYNTHETIC VIEW (1955):
Dipleurula was remotely related non- echinoderm forms for their
bilateral symmetry and went through a sessile stage of radial forms to
reach the echinoderm status. This radial form Pentactula stage where
reorganisation took place and gave rise to free- swimming and attached
forms parallely.

Fig.19.6. Hyman’s suggestion

H. B. FELL (1963):
Supports free-swimming origin of echinoderm larvae and their
phylogenetic correlation through Vitellaria.
PARKER AND HASWELL (1972):
Dipleurula larva was free-swimming, gave rise to the free-
swimming forms through the free-swimming Antedon on one hand and
to the fixed type through the fixed Antedon forms, on the other.
Barrington has summarised the work of other workers like Berrill
(1955), Bone (1960), Carter (1957), Marcus (1958) and Whitear (1957)
and has proposed that the echinoderms, the pogonophores, the
hemichordates and the rest of the chordates arose separately but
directly from a common bilaterally symmetrical sessile or semi-sessile
ancestor with tripartite body and coelom, ciliated larval stage and ciliary
mode of feeding from external source.

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 Fig.19.7. Phylogenetic relationship of deuterostomia

Let Us Sum Up

In this unit, we studied about the strategies and evolutionary

significance of larval forms in invertebrates. Under this unit, we focused
on the evolutionary significance of larval life such as, planula larva in
cnidaria, Platyhelminthes larva, crustacea larva, trochophore larva and
echinoderm larva.

Check Your Progress

1) __________ in 1881, proposed ctenophore polycad theory.

2) The three parasitic classes are __________, __________ and
__________.
3) Molluscs were derived from __________ like ancestors.
4) The study of larval forms and adults of echinoderms reflects a case
of __________.

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Glossaries

Apopyles : An opening by which water passes out of radial

canal
Biramous : Diving to form two branches

Stomodeum : A depression between brain and pericardium in an

embryo.

Suggested readings

1. HIGHNAM, K.C. and HILL, L. (1979). The Comparitive

Endocrinology of Invertebrates, ELBS & Edward Amold (Publishers)
Ltd., London.
2. HYMAN, G.H (1967) the invertebrates, Vol. I to VII, McGraw Hill Book
Co., Inc., New York.

Weblink

Strategies of larval forms

https://en.wikipedia.org/wiki/Marine_larval_ecology#:~:text=the%20water
%20column.-
,Larval%20development%20strategies,Direct%2C%20lecithotrophic%2C
%20or%20planktotrophic.
Evolutionary significance of larval forms https://youtu.be/-dLoXnWeexk

Answers to check Your Progress

1) Lang
2) Trematoda, monogenea and cestode
3) Turbellaria
4) Retrogressive metamorphosis

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