CORE-3, UNIT-2

CORE-3 UNIT-I B. Sc.
Respiration in Arthropoda
1. Aquatic Respiration:
The organs associated with aquatic respiration are:
1. Gills or Branchiae
2. Tracheal gills
3. Blood gills
4. Rectal gills
5. Book gills
6. Branchiostegite or gill cover
7. Epipodite and
8. Branchial basket.
1. Gills or Branchiae:
(i) Occurrence:
The gills are the respiratory organs of aquatic arthropods. These are best developed in
crustaceans. In other aquatic arthropods, special types of gills are often encountered.
(ii) Location:
Gills are situated within the gill chamber. The gill chamber is located on each lateral side of
the cephalothorax and covered by the gill cover or branchiostegite.
(iii) Origin of gills in crustacean:
Gills originate as out-pushings of the body wall. In Amphipoda, the gills are outgrowths of
the thoracic limbs and in Isopods the endopodites of second and fifth pleapods are modified
as gills.
(iv) Structure of gills in Crustacea:
A typical gill (Fig. 18.10) is crescent- shaped. It consists of a central axis or rod, on each side
of which are arranged blade-like gill filaments, called lamellae. One end of each filament or
lamella remains connected with the rod or central axis and the other end of the filament is
blind. Through the central axis of each gill runs an afferent and an efferent branchial channel.
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By: Debashish Nayak, Department of Zoology, Dr. R.C. SIST Nayagarh, Utkal University.
(v) Types of gills in Crustacea:
A. Based on the shape of the lamellae, the gills are of the following types:
1. Phyllobranchiate gill:
In this type of gills the lamellae are flat, broad leaf-like and are arranged in two rows. (Fig.
18.127A, B). It is found in crabs and prawns (Palaemon).
2. Trichobranchiate gill:
In this type of gills, the gill filaments are tubular-shaped. It is found in crayfish (e.g.,
Astacus) and rock lobsters. It consists of a central axis with numerous lateral filaments,
formed from the sides of the body or from an outgrowth of skin of the legs (Fig. 18.127C, D).
3. Dendrobranchiate gill:
In this type of gills, the leaf-like lamellae are divided into fine branched filaments (Fig.
18.127E, F). It is found in Penaeus.
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B. Based on the mode of attachment, the gills may be of three types:
1. Podobranch:
Podobranch attached with the coxopodite of the thoracic appendage. In Macrobrachium (=

Palaemon), Penaeus the podobranchs are one pair and remain attached to the second
maxillipedes (Fig. 18.10).
2. Arthrobranch:
Arthrobranch the gills are attached with the arthroidal membrane which connect the
appendages of the thorax. In Palaemon the arthrobranchs are two, attached to the arthroidal
membrane of third maxillipede. In Penaeus there are eleven pairs of arthrobranchs, starting
from second maxilliped to third walking leg, two in each appendage and fourth walking legs
contain single arthrobranch.
3. Pleurobranch:
Pleurobranch—the gills are attached with the lateral wall of the thorax. In Palaemon there are
five pleurobranchs, attached to the lateral side of the thorax. Is Penaeus there are six pairs of
pleurobranchs attached to the last six pairs of thoracic appendages.
(vi) Number of gills in Crustacea:
Number varies in different groups. The Decapods which contain all the types of gills
exhibit extreme variation—in the shrimp, Lucifer, gills are absent; penaeid shrimp has 24;
Homarus has 20; Peacrab contains 6 gills. In Palaemon, there are 8 pairs of gills and the
anterior gills are small and the size increases towards the posterior end.
(vii) Modification of gills in Arthropoda:
The gills are variously modified in Crustaceans and other Arthropods. In Phyllocarida, broad
epipodites of the thoracic appendages work as gills. Similar gills are seen in Cumacea. Gills
are plate-like in Amphipoda and flattened in a Decapod, Palinurus.
In Euphausiacea, the tufted podobranchs are not covered by carapace. The gills appear as a
row of small branchial lamellae on each side of Cyprididae. In Phyllopoda, the leaf-like
pleopods work as gills. Among the Crustaceans only Stomatopods and Isopods have
abdominal gills.
2. Tracheal gills:
In the aqualtic larvae of many insects a series of simple and divided external processes are
attached to the abdominal segments. These are richly supplied with tracheae and are called
the tracheal gills (Fig. 18.128A), help in respiration.
3. Blood gills:
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In certain aquatic insect larvae (mainly chironomidae) the tracheae are replaced by the
branching tubular outgrowths containing blood vessels and are called the blood gills.
Remark:
These were formerly thought to be respiratory organs but recently it has found that the
respiration of these animals takes place throughout the whole body surface.
4. Rectal gills:
In the nymphs of several insects the inner surface of the rectum bears gills. These gills are
called the rectal gills.
5. Book gills:
The most specialised gills are seen in Xiphosurids, where the abdominal appendages bear
plate-like book gills (Fig. 18.128E). These gills are formed by the evagination of the posterior
borders of opisthosoma in segments from ninth to thirteenth. Each gill contains nearly 150
lamellae, which look like the delicate leaves of a book.
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Mechanism of gill respiration:
In most Crustaceans, the gills are not covered within a special gill chamber. But in Decapods,
the carapace extends laterally over the gills to house them in a special chamber.
In such forms with chamber, current of water enters through one end and after bathing the
gills, passes out through another direction. In Crustaceans and Xiphosurids, gaseous
exchange takes place in the gills between the blood and the water. But in Insects, after
diffusion the oxygen passes to the tracheal tubes.
Branchial formula:
The number and arrangement of gills are presented in a form of formula on each side, called
branchial formula. The branchial formula of a freshwater prawn (e.g., Macrobrachium sp. =
Palaemon sp.) is represented in Table 18.17.
Other devices of aquatic respiration:
6. Branchiostegites:
In Crustacea the gill-chamber is covered by the lateral extension of carapace, called gill cover
or branchiostegite. The inner lining of the branchiostegite is thin, membranous and richly
supplied with blood. It is in direct contact with water current and exchanges gases between
the blood and the water.
7. Epipodites:
These are small highly vascularised leaf-like membranous outgrowths of integument on the
outer side of coxa of the maxillipeds in first three thoracic segments. These epipodites being
present in the anterior part of each gill-chamber (e.g., Crustacea) carry out respiratory
functions.
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8. Branchial basket:
The immature Odonates (Insects) have their rectum modified into a branchial basket. Its wall
is contractile and richly supplied with the branches of tracheae. This kind of respiration is
often referred to as anal respiration.
2. Aerial Respiration:
The aerial respiration takes place in terrestrial arthropods.
The organs for respiration are the following:
1. Trachea
2. Lungs
3. Book-lungs
4. Pseudotracheae or air tubes
5. Anal respiration
6. Miscellaneous devices
1. Trachea:
This is the most important organ for aerial respiration. This chitin-lined tube is seen in almost
all land arthropods, such as insects, centipedes, millipedes and many arachnids.
Two types of tracheae are seen:
(i) Ventilation trachea—oval in section and collapses after the exhalation of air and
(ii) Diffused trachea—rigid and does not collapse after the exhalation.
Origin:
The tracheae originate as the invagination of the body wall.
(a) Structures of trachea and associated parts:
(i) Each trachea is a tube with walls made up of polygonal cells.
(ii) The wall of trachea is composed of three layers—these are the internal layer, called
intima, a middle layer of epithelium and an outer layer of basement membrane.
(iii) The intima is lined by spiral cuticular ridges, called taenidea, that prevent collapse.
(iv) The tracheal cuticle contains the same layers as the surface cuticle except the cement
layer and wax layer.
(v) The tracheae open externally by small openings, called spiracles or stigmata.
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(vi) These spiracles are located along the sides of the body.
(vii) Each spiracle opens into a chamber, called atrium and the spiracle is placed on a plate,
called penetrene.
(viii) Each spiracle has two lids for opening and closing.
(ix) Within the chamber foreign particles are eliminated by a filtering apparatus, containing
either special bundles of setae or a kind of sieve-like membrane.
(x) Some parts of tracheae are dilated to form air-sacs. They help as reservoirs of air.
(xi) The finer branches of tracheae are called tracheoles which are without inner taerridial
ridges. A tracheole may be 1µ in diameter and reaches every cell of the body.
(xii) The end of a finer tracheae is immersed in a fluid through which gaseous exchange takes
place.
(b) Classification of tracheae:
(i) In adult insects, the tracheal system is of one kind.
(ii) Two pairs thoracic and eight pairs abdominal spiracles are usually present in all adult
insects. There are 12 pairs in primitive condition.
(iii) In certain forms some spiracles may be secondarily absent but they appear at least in
some stages of development. For example, the queen termite has only six pairs abdominal
spiracles instead of eight pairs. The metathoracic pair of spiracles is absent in Lepidoptera,
Hymenoptera, Coleoptera and a few others.
(iv) In millipedes, a pair of spiracles is present in each thoracic segment and two pairs of
spiracles in each abdominal segment.
(v) During development, spiracles appear in varied ways in different in sects. Thus from the
point of view of embryology the tracheal system is classified on the basis of the number of
functional spiracles. This classification does not denote any special kind of tracheal system in
the adult.
On the basis of functional spiracles the tracheal system in larvae may be classified as:
(i) Polypneustic:
Tracheal system openings to the exterior by 8 or more pairs of functional spiracles.
It may again be subdivided into:
(a) Holopneustic:
When 2 pairs of thoracic and 8 pairs of abdominal spiracles are functional. The term is used
when 10 pairs of functional spiracles are present.
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(b) Peripneustic:
A respiratory system with 1 thoracic and 8 abdominal spiracles are present on each side of the
body. The term is denoted when the abdominal spiracles occur on all the segments of the
abdomen.
(c) Hemipneustic:
A respiratory system with 1 thoracic and 7 abdominal spiracles are present on each side of the
body. The term is used when one or more pairs of spiracles are nonfunctional.
(ii) Oligoneustic:
Here, either one or two pairs of spiracles are functional.
It includes divisions like:
(a) Amphipneustic:
When one pair of thoracic and one pair of post-abdominal spiracles are present. Such
condition is found in the larva of the common house fly.
(b) Metapneustic:
Only one pair of post abdominal spiracles is functional. This condition is seen in the
mosquito larva.
(c) Propneustic:
Only one pair of thoracic spiracles is functional. This condition is seen in the pupae of certain
Diptera.
(iii) Apneustic:
No spiracle is present in functional state. Gaseous exchange takes place through the
integument, seen among aquatic insect larvae.
(c) Mechanism of tracheal respiration:
The trachea ramifies into a number of fine networks of tracheoles which terminate into
tissues where exchange of gases takes place by diffusion. Air is drawn in and forced out
through the spiracles by the alternate contraction and expansion of the body. The spiracles
remain closed most of the time and exchange of gases is probably due to diffusion and
ventilation.
Recent studies indicate that the spiracles open very briefly but not all at a time due to
reduction of haemocoelomic pressure. The spiracles are closed by valves, thus control the
water loss, and opening of the spiracles is related to the high CO2 concentration.
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Gaseous exchange through the tracheae takes place by diffusion primarily and tracheoles are
permeable to water and remain fluid-filled. This fluid is believed to be involved in the final
O2 transport to the tissues.
Again it is reported that the movement of trachea is facilitated by the alternate contraction
and relaxation of the body sclerites. In the bed bugs, rigid and convex sternum does not take
part in the respiratory movement, which is done only by the elastic tergum. In cockroaches
the tergum and sternum of the segments are separated by intersegmental membrane which
bulges out during respiration.
(d) Modifications of the tracheae:
In most Collembola, the tracheae are absent and the respiration is largely cutaneous. In
Machiles, segmental tracheae originate from spiracles but do not have trunks. In the larva of
Musca, dorsal longitudinal trunk is provided with one pair of anterior and one pair of
posterior apertures.
In the larvae of mosquito, a single spiracle is connected to the dorsal trunk. In the Myriapods,
stigmata open within air chamber from where large numbers of tracheae are given off. The
other peculiar features of this group are that in Diplopoda the tracheae are branched and in
Symphyla only two tracheae are present on the head.
Other devices of aerial respiration:
1. Lungs:
In the Crustacea, Birgus, the upper part of the gill-chamber is separated from the rest and
forms a closed chamber within which vascular tufts project and perform aerial respiration.
2. Book-lungs:
The book-lungs are best seen in Scorpionids and spiders (Fig. 18.128F). These are blind sacs
which originate from the evaginations of opisthosoma. These are regarded as the modified
abdominal appendages.
Within the sac the inner lining is raised into numerous delicate folds, like the leaves of a
book. These folds are richly vascularised and thus respiration in Scorpionids is circulation
dependent. Each book-lung communicates to the exterior by a stigma.
3. Pseudotracheae or air tubes:
The only land living Crustacea, Oniscus (wood lice), possess numerous minute tube-like
structures in the abdominal appendages, called Pseudotracheae, help in respiration.
4. Anal respiration:
Many crustaceans perform rhythmical contractions of intestine—taking in and expelling out

water. Such anal respiration is common in lower crustaceans and is especially noticeable in
Cyclops.
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5. Miscellaneous devices:
A combination of book-lung with trachea is seen in spiders. Some aquatic members of

Colleoptera and Hemiptera (e.g., Nepa, Ranatra) carry respiratory tubes, located at the
posterior end of the body which is formed by two cerci. While they dive inside the water,
they cany air with them for respiration. In mosquito larvae, a long siphon draws air from the
surface of the water.
Type # 1. Ametabolic Metamorphosis:
In lower insects (Collembola, Thysanura) the young which hatches from an egg is a miniature
of the adult and is called a nymph, it differs from the adult in having immature reproductive
organs; by several moultings and growth it becomes an adult.
These insects are primitively wingless, they are also called Apterygota, e.g., Lepisma, the
change from young to adult is negligible, such insects are ametabolic because there is no
metamorphosis.
Type # 2. Heterometabolic Metamorphosis:
In winged insects the adult differs in several respects from the young, such insects are said to
undergo metamorphosis in becoming adults. The nymph which hatches from the egg has a
general resemblance to the adult in body form, type of mouth parts and possession of
compound eyes, though these nymphs may have adaptations associated with their particular
habits of being aquatic, swimming or burrowing.
In these the change from nymphs to adults is a gradual process in which appendages, mouth
parts, antennae and legs of the nymph grow directly into those of the adult.
Wings develop gradually as external outgrowths of thorax and are visible externally in the
nymphal instars, because of their external wing development they are also called
exopterygota. The reproductive organs mature gradually. Insects showing this slight change
from nymph to adult are known as heterometabolic (gradual), they include Dictyoptera,
Orthoptera, Isoptera, Hemiptera and Anoplura.
Though nymphs of dragon flies, may flies, etc., are quite different from the adult in having
special nymphal adaptations because their nymphs are aquatic, while the adults are aerial, the
nymphal adaptations are shed in changing into adults, such forms with slightly greater
changes are called hemimetabolic (incomplete), they include Odonata, Plecoptera and
Ephemeroptera.
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Type # 3. Holometabolic Metamorphosis:
In Lepidoptera, Coleoptera, Hymenoptera, Diptera, Siphonoptea, etc., the young which

hatches from the egg is called a larva, the larva is very different from the adult in structure,
body form, mouth parts, legs and in its mode of life, the larva has lateral ocelli in place of
compound eyes, it feeds voraciously, grows, moves about and undergoes ecdyses.
The larva is so different from the adult that it first changes into a resting, quiscent instar
called a pupa which is often enclosed in a cocoon secreted by the labial glands of the larva.
Great transformation occurs in this instar, wings develop internally from pockets of the
hypodermis, and they are not visible from outside.
Because wings develop from internal imaginal discs these insects are also called
endopterygota. Appendages are formed, muscles, tracheae and parts of the alimentary canal
are replaced by corresponding organs of the imago. Such vast changes are called
holometabolic metamorphosis.
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In holometabolic insects there is an internal reconstruction during late larval and pupal
instars. Larval organs, with the exception of central nervous system and developing
reproductive organs, are disrupted, their breaking down is called histolysis, this is brought
about by phagocytes which feed on the organs, and products of their digestion are then used
for building new structures.
The building of new structures is brought about by growth centres called imaginal buds or
discs. Imaginal discs are groups of formative cells which are set aside in the larva, they are
the rudiments of future organs of the imago, they form legs, mouth parts, internal organs and
wings.
This process of formation of organs of an imago from imaginal discs inside the pupa is
known as histogenesis and it results in the formation of the imago.
Thus, two postembryonic processes occur in all insects, the first is growth in the young and
the second is metamorphosis, in both of which moulting takes place; both processes are
controlled by hormones of endocrine glands. Insects have two such endocrine glands, they
are corpora allata and prothoracic glands.
The juvenile hormone of corpora allata controls growth and moulting up to the end of the
larval period. So long as the juvenile hormone of corpora allata is produced the final moulting
into a pupa or into an adult cannot take place.
The prothoracic glands are a pair of small glands in the first thoracic segment, they produce a
hormone called ecdyson which brings about moulting and development of imaginal discs and
reproductive organs.
When both hormones are secreted, then moulting of the larva only will take place. The result
of the two hromones is supression of adult characters from appearing during larval and pupal
instars. When only ecdyson is secreted, and the juvenile hormone is not produced, then the
larva will moult into a pupa, and the pupa into an imago.
Thus, it is seen that ecdyson is essential for each moulting, but its action is modified as long
as the juvenile hormone is present. Removal of the old cuticle in ecdysis is brought about by
an enzyme secreted by the hypodermis, the enzyme erodes the lower surface of the cuticle,
then the hypodermis secretes a new cuticle below the old one.
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1. Features of Termites:
1. Termites are pale-coloured and usually blind except macropterous forms.
2. Termites possess comparatively large head, chewing and biting mouth parts.
3. Well-developed eyes in reproductive castes.
4. Ocelli are either touching or away from the eyes.
5. Soft-bodied females have much enlarged abdomen.
6. Winged (alate) and wingless (apterous) forms are composing the colony.
7. In winged forms, the fore-wings and hind-wings are same sizes.
8. The colonies contain many castes, such as queen, workers, soldiers, etc.
2. Distribution of Termites:
The termites are broadly divided into two categories—lower termites and higher termites.
The lower termites include the following families:
1. Mastotermitidae:
It includes the most primitive living termite, Mastotermes darcviniensis. It is found in

Northern Australia and Papua, New Guinea. No primary queen has ever been recorded in the
field and numerous substitute queens are usually found in the nests.
2. Kalotermitidae:
They are known as dry wood termites because they live in dry wood although some dry wood
termites have subterranean habits. There are 400 species and are distributed throughout the
world.
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Examples:
Kalotermes, Glyptotermes.
3. Termopsidae:
They are known as damp wood termites because they live in wet and rotting wood. There are
about 20 species and are distributed in the forests of America, Eurasia, Africa and Australia.
4. Hodotermitidae:
They are known as grass harvesting termites because they have adapted in grass land where
the trees are scanty. They are found in India and Africa.
Examples:
Hodotermes, Anacanthotermes.
5. Rhinotermitidae:
Known as subterranean termites except Coptotermes, Reticulitermes. Some mound-builders

and aerial nesters are found in this family. About 300 species have been recorded and are
distributed in most of the continents.
The higher termites include a single family, Termitidae with several subfamilies.
6. Termitidae:
They build generally clay nests of great height above ground (also called termitaria) that
occur in tropics including Asia, Africa, Australia and South America. About 1800 species
have been recorded. Examples are Amitermes, Microcerotermes, Termes, Cubitermes,
Capritermes, Nasutitermes, Macrotermes, Microtermes, Odontotermes.
The lower groups of termites contain symbiotic intestinal protozoa and bacteria, and higher
termites contain intestinal bacteria only.
3. Social Organization of Termites:
“Social organisation is a process in which single or both parents live under a shelter and
perform the activities in an organised and co-operative manner for the maintenance of
the community”.
The detailed account of termites was published first in the Royal Society of London in 1781
by Henry Smeathman. The account was based on African termites.
A typical colony possesses nymphs (semi-matured young’s), workers, soldiers and

reproductive individuals of both sexes (Fig. 18.91A, B, C, D, E, and F); sometimes also
several egg laying queens are included. The different types of insects, each having a par-
ticular function live in a colony, called castes.
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4. Caste System of Termites:
A termite colony includes two major castes—sterile castes and reproductive castes. The caste
system and the size of the colony are controlled by a pheromone, called social hormone,
which is secreted by the members of reproductive castes (e.g., king and queen).
A. Sterile castes:
They are incapable of reproduction and include usually workers and soldiers (Fig. 18.91).
(i) Workers:
They are sterile of both sexes and developed from the fertilized eggs. They constitute major
numbers in a colony and occupy 80%-90% of the total number. They are small-sized
individuals measuring about 6 to 8 mm in length which bear chewing mandibles and usually
lack of eyes.
But workers of some harvesting species bear well developed compound eyes. They have no
wings (apterous). They feed upon the wood or fungus products. Advanced termites produce
true workers but in primitive families (e.g., Mastotermitidae, Kalotermitidae, Termopsidae,
Rhinotermitidae) the true workers are not found.
Instead, nymphs (an insect larva hemimetabolous insects, resembling the adult except they
have no wings and reproductive organs) of various stages which are arrested in their
development act all the domestic duties of the nest like the workers, called Pseudergater or
false worker, found in the colonies of Kalotermitidae.
They persist as worker throughout life or can metamorphose into winged reproductive castes
and also act as potential supplementary reproductive castes.
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Division of labour:
Workers perform all the duties for the colony except reproduction. They take care of the eggs
and young, undertake the labours of food gathering, food storage and feed the nymphs, king
and queens.
They take part in the building of nests and in the cultivation of fungus-garden in special
chambers in the subterranean channels. They also maintain the moisture in the interior of a
nest of a colony. They perform defense duties in some species and also clean the other castes.
(ii) Soldiers:
The soldiers of some species of different genera show great diversity in characters. They are
developed from the unfertilized eggs. They are wingless and larger than the workers. Soldiers
are more or less pigmented and possess large head and stout, powerful mandibles.
They are mostly devoid of eyes but occasionally appear in some species (e.g., Mastotermes,
Hodotermes, etc.). They constitute 5-8% in number in a colony. Soldiers for food rely on
workers. They maintain the defense of the colony.
The specialized forms of soldier termites within many species of the family Termitidae,
called nasutes. They are characterized by the reduced mandibles and head which is continued
as a rostrum, at the tip of which opens a frontal gland.
The secretion of this gland is acrid which helps to prevent enemies and at the same time it
dissolves hard substances including concrete structures. In some Rhinotermitidae, the soldiers
are specialized with a pair of large strong mandibles and highly elongated grooved labrum
that contains defensive secretions, called nasutoid soldier (e.g., Nasutitermes).
B. Reproductive castes:
These are fertile and are characterised by the presence of the eyes.
From the functional point of view, the reproductive castes are of three types:
1. Macropterous or large winged or primary reproductive forms (Fig. 18.91):
These sexually matured males and females have two pairs of wings longer than the body and
convert kings and queens. The body is usually pigmented and may be yellow, brown or black.
The widely separated large compound eyes and paired ocelli are usually present. The royal
pair or the sexually active males and females of social insects are the original founders of the
colony.
King:
Usually there is a single king in a colony. It is smaller than the queen. Its function is to mate
intermittently and provide sperm to the queen.
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Queen:
The queen is also a single in a colony which is largest in size measuring 60- 80 mm in length.
It lays eggs after mating and can lay more than 1000 eggs per day. Both king and queen live
in a special chamber, called nuptial chamber.
It is widely believed that the queen or king is the primary source of a pheromone (also called
social pheromone) that maintain the colony integration and the size of the colony. The colony
integration is also maintained by the exchange of food, saliva or some secretions among
colony members. This process is called trophallaxis.
2. Brachypterous neotenics or short winged or substitute reproductive forms:
When the primary or original king or queen dies or gets old, its place is taken by the sexually
mature male or female but in nymph form. To replace them, the nymphs of the suitable sex
(male or female) attain maturity by special feeding without further moulting and become the
substitute queen.
The substitute reproductives are characterized by the presence of short pad-like wings, ocelli
and rudimentary compound eyes and small reproductive organs. The substitute reproductives
vary in number in different species within colony. It is recorded fifty functional substitute
queens in a colony of Hodotermes.
Grassi and Sandias reported the condition of castes in the nests after examination of thousand
colonies in Italy where they found none of the primary reproductives in a single colony and
only substitute reproductives numbering 10-200 per nest.
3. Apterous neotenic reproductives or wingless forms or replacement reproductives:
If queen dies in some adverse conditions or gets old, the apterous neotenic reproductives are
produced. In some primitive subterranean species, like Reticulitermes flavipes,
Reticulitermes malletei, the nymphs or ergates (workers) are without wings and possess less
developed eyes, and reproductive organs become a functional reproductive in the colony with
one or two moults.
The workers and soldiers are usually blind and live deep in the soil of the mound, so they do
not require vision and always remain in close contact of food source. The reproductive adults
have well developed eyes because the vision is required for flight and in search of nest sites.
5. Food and Feeding of Termites:
The eusociality of termites largely depends on food and feeding. After the production of
workers and nymphs, the king and queen are fed by the workers. The mutual exchange of
food, salivary secretion and other secretions among the members of termites are called
trophallaxis.
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The social insects exchange food in two ways:
(i) Stomodaeal trophallaxis and
(ii) Proctodaeal trophallaxis.
Stomodaeal trophallaxis is related to the exchange of food among the members through
mouth to mouth but in proctodaeal trophallaxis, the exchange of food and symbiont through
anus to mouth. In termites and cockroaches, the proctodael trophallaxis is crucial for the
replacement of endosymbionts that are lost after every moult and this behaviour is used to the
origin of sociality in insects.
On the basis of feeding, termites can be divided into wood (cellulose) eating and, or litter
feeding, fungus-growing and soil-feeding types.
(i) Wood (cellulose) eating termites:
The wood (cellulose) eating termites (e.g., Archotermopsis, Zootermopsis) feed on cellulose
of the wood, collect either from living or dead plants. The cellulose is a polysaccharide, each
containing three thousand sugar units and important structural component of the cell wall of
plants.
The termites possess intestinal fauna of symbiotic protozoa (e.g., Trichonympha) which help
to digest the cellulose, secreting cellulase enzyme. These protozoans enter the nymphs which
consume the excrement of the adult. The recent molecular evidence indicates that termites
also use own enzymes for cellulose digestion.
(ii) Soil-feeding Termites:
The soil-feeding termites are Cubitermes, Termes, Capritermes (Subfamily Termitinae,

Family Termitidae) which consume large quantity of soil as a food.
In these higher termites they have lost the symbiotic intestinal protozoa, and only gut bacteria
is found which can degrade soil organic matter that provides the termite host with nutrition.
The excreta contain a mixture of clay and broken down vegetable debris, used for the
construction of mounds and galleries.
(iii) Fungus-growing termites:
Fungus-growing termites are small insects, ranging 4 to 15 mm long and belong to the
subfamily Macrotermitinae, feeding on wood and plant fibres using detritus. These termites
could not digest cellulose and need help digestive capabilities of fungi Termitomyces
(Basidiomycetes). Macrotermitinae cultivates a fungus-garden around the colony and creates
some ventilated structures inside the nest in the form of fungi combs.
The termites consume cellulose-based materials like woods and vegetable debris; they also
eat a certain amount of fungi. These fungi help the digestion of food inside the bodies of
termites that help in the breaking down of ligneous and cellulose based substances.
18
6. Swarming and Formation of a New Colony by Termites:
Swarming is the concerted departure of the winged termites from the main colony to start
new colonies, accompanying by a queen or males or females together, generally occurring
during rainy season. Before mating, the new kings and queens leave the colony and fly for a
while.
The kings and queens that take part in the swarming, called alates or swarmer’s. The mating
flight of the royal pair is known as nuptial flight. But it is not a true nuptial flight in termites
because the copulation does not take place during flight. Generally after flight the males and
females fall on ground, discard their wings, choose their mates, and find a suitable place
nearby for nesting.
Swarming behaviour of the opposite sexes is different in different families and genus. The
behaviour pattern in primitive families like Kalotermitidae is most simple. They are
crepuscular flier. Male and females fall on the ground after flight and a wait for the opposite
sex.
Before mating, the wings are shed and after pairing they excavate a chamber in a suitable
place. In higher termite species the female usually alights first on the ground after flight and
shows an attractive altitude elevating its abdomen.
The female releases a pheromone, a social hormone that attracts the males. Before mating,
they shed their wings and dig a chamber nearby in which they take shelter and where
copulation takes place.
Termitarium:
Generally termites construct colony by digging tunnels underground (e.g., Odontotermes

assumthi, Microtermes obesi, etc.) or make a large mound above ground or nests are inside
wood of trees or buildings. The common termites which build clay nests of great height [Fig.
18.92(i)] above ground are Odontotermes redemani, O. obesus, etc.
A termite nest (termitarium) presents highest form of architecture and within the nest there is
a special chamber where the queen and a few males are confined.
The special chamber is called nuptial chamber. A colony [Fig. 18.92(ii)] includes nursuries,
chambers, royal chambers, special chambers for cultivating fungus-gardens or storing food,
guard rooms, bridges, corridors and subterranean streets and canals [Fig. 18.92(ii)].
19
The period of swarming occurs at different times of the year. The swarming period of
Odontotermes usually occurs during rainy season one hour after and of course half an hour
after rainfall. In dry-wood termites the swarming occurs in hot days during summer season.
During flight most of the termites are eaten by birds, small mammals or destroyed.
7. Development of Termites:
After copulation, the queen produces a large number of eggs, most of which develop into
nymphs. These nymphs develop into sexless workers, soldiers and substitute reproductive
20
forms and may become sexual forms, if necessary. The queen at that time is specially fed and
it grows to a large size.
The large size is due to enormous growth of abdomen in which huge quantities of eggs are
deposited. This abnormal-sized abdomen is known as physogastrous and the termite queen
containing such distended abdomen, called physogastrous female. After copulation the queen
starts laying eggs in a chamber, called royal chamber, but the egg-laying capacity is not equal
in all families or species.
In primitive termites the egg-laying is intermittent and the population is small. In

Coptotermes the queen lays 100 eggs per day, but in advanced termites the queen can pro-
duce 30,000-40,000 eggs per day and 10 million a year.
In the primary stages of the colony the nymphs hatch from the eggs within few days and
moult at least three times to become functional workers, and into soldiers after fourth moults,
and also substitute reproductives develop from nymphs (Fig. 18.93).
But at the end of winter several sexual forms appear. When matured, these winged forms take
colonising flight, which occur during rainy season.
The wings in them extend beyond abdomen and are membranous. After becoming functional
workers and soldiers, they take charge of the nest and take proper care of their parents. They
expand the nest and make adequate arrangements to accommodate the growing population.
Longevity:
Primary queens can survive for 15-20 years and some may live for 50 years. Aborigines in
Australia have reported that the termite queens of mounds can survive up to 70 years.
21
Workers may only live for a few months. Mastotermes kings have been recorded about 16
years.
The longevity of the individuals of the different castes in the colony is an important factor in
establishing the close relationship between successive generations which form the basis of
social organization or eusociality of termites.
Communication:
Termites communicate each other in a colony through chemical, acoustical and tactical
signals. The sexual communication and trail following of the termites are connected each
other by pheromones. There are two pheromones in termites, such as (ZZE)-(3, 6, 8)-
dodecatrienol and (E)-6-cembrene, have been recognised as trail pheromones.
8. Economic Importance of Termites:
1. The workers and soldiers leave their nest at night to attack furniture, woods and books.
2. Thus they damage human properties in several ways.
3. The only way to get rid of termite menace is to destroy the queen.
4. In spite of its destructive role, the termites are considered important from the point of view
of agriculture.
5. Like earthworm, the termites also pulverise the soil and make it fertile.
6. Swarming termites are taken as food by birds and other animals.
HONEY BEE
The social organization of the honey-bees is established by the living of all individuals within
the colony and they show the mutual cooperation among the members of the colony, and
exhibit the overlapping generations.
At least there is a division of labour among the different types of honey-bees in the colony or
hive. The different forms or types of insects having a particular function live in the colony,
called the castes.
Caste System:
Thousands of bees (50,000 to 1,00,000 or more) which live in a hive are of three different
forms:
(1) Workers (infertile females),
(2) Drones (males) and
(3) Queens (fertile females) (Fig. 18.74A).
22
The phenomenon of existence of several morphological forms with separate functions in a

species is known as polymorphism. So honey-bees are well known as social and polymorphic
insects.
The true social organization (eusociality) of honey-bees is well understood in the study of
Apis mellifera. They live in colonies in hives and each bee-colony includes several thousands
of bees which consist of one queen, several hundred drones and tens of thousands of worker
bees (50,000-80,000 or more).
Both queen and workers are female and diploid. Drones are males and haploid. A strong or
healthy colony is called when the maximum number of workers is found in the colony.
Queen bee:
Generally a single matured queen is present in each hive. The size of the queen is nearly 2.5
times longer than that of a worker bee. If is characterized by the long tapering abdomen, well-
proportioned body, short and golden coloured wings and colour of the legs. They are 2.8
times heavier than the worker bees.
23
The queen possesses a curved sting at the tip of the abdomen which is known as ovipositor.
The function of the queen is reproduction and lays about 1000- 2000 eggs every day
depending upon seasonal variation and seasonal factors. The eggs may be either fertilized or
unfertilized.
Depending on the type of food supplied to the newly developed larvae by the nursing workers
the eggs may develop either queen or workers. The drones or males are produced by the
laying of unfertilized eggs (i.e. parthenogenetically). The queen deposits each egg in a cell
prepared by the worker bees (Fig. 18.80).
After three days the eggs hatch into small larvae. The larva which is fed with a special food
called ‘royal jelly’ develops into queen. The royal jelly is a high proteinous substance
produced by the hypopharyngeal glands of the workers.
The larva which is selected to become queen is taken before the third day of development in a
special chamber, called queen’s chamber. The queen lives five to eight years on average and
her fertility decreases with the increase of age. The sting of the queen serves an ovipositor for
lying of eggs and is also used for defence. The sting is used only when she meets another
rival queen.
The queen secretes a kind of chemical substance with hormonal properties from the
mandibular glands, called pheromone or queen substance which inhibits the growth of
ovaries of workers and control the activities of all bees within the hive. She can attract the
workers towards the queen and stimulates the workers to build wax cells for worker bees and
drones but prevents in the building of queen cells.
The Dutch naturalist Swammerdam (1673) remarked that queen is the mother of the entire
colony and has no food collecting organs. Johann Dzierzan, a German scientist reported in
1980 that the queen bee is the centre of the bee family and the mother of all the young bees.
24
When the queen becomes matured she leaves the hive with some drones and takes several
nuptial flights and mates with a drone. After copulation the drone dies soon after and the
queen stores enough sperm in the spermatheca to last her lifetime. The queen after copulation
returns in her old hive and is looked after by nurse workers, known as her retinue.
With the increase of the age the egg laying capacity of the queen loses, the workers choose a
three day old egg. This egg after hatching into larva is fed with royal jelly and it develops a
new queen in about 16 days. At that time the old queen leaves the hive along with some
workers to establish a new colony.
Drones:
The drones are the male members of the bee colony and are haploid each genetically. The
drones take 24 days to develop from the egg to adult. They have no food (pollen and nectar)
collecting organs. So the drones are totally dependent on worker bees for food. They feed on
the honey during spring and summer months provided by worker bees and are driven out in
autumn from the hive.
Function:
The main function is to fertilize the queens. They also help to maintain the warmth of the
hive which is necessary for the hatching of the eggs.
Workers:
The size of worker bee is small but they constitute the majority in a hive. They are produced
by the fertilized eggs laid by the queen. It takes 20 days from egg to adult and life span is
about 6 weeks.
Function:
All the time in their lives is spent for the maintenance of the hives and caring for their
members. Workers are involved in hive construction, clean the cells of the hives collect the
nectar, pollen and water and store within the cell properly. Water is required for the
preparation of royal jelly from pollen for food and for dissolving crystalline honey.
They repair the cracks in the walls of the comb and polish the walls with propolis. The
workers also maintain the optimum temperature within the hive by fanning during summer,
and in severe winter when the temperature falls sharply they gather outside the hive and
reduce the temperature loss of the surface of the hive.
It is assumed that workers always remain busy in their duties but A. I. Root, an American
scientist often pass remark during his lectures that the workers slept in night heavily than
during the day. If any intruder bee happens to fly into the hive, the workers not only kill the
intruder but push the body out from the hive as a sign of warning to others.
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Swarming and Formation of New Colony:
During the early summer swarming happens to avoid the overcrowding in the old hive. The
old queen, several thousand workers and hundreds of drones come out from the old hive and
fly together in search of a suitable place for the formation of new colonies, In the new hives,
the different castes live together, in which division of labour is well pronounced.
Nuptial Flight and Copulation:
About a week after emergence from her chamber, the new queen flies in air with many
drones. The copulation takes place in the air with a drone and the queen receives the
spermatophores from the drone. After copulation, the genital parts of the drone are forced out
and the drone dies immediately.
It was believed that the queen copulates only once in her life but recent observations have
shown that they can mate more than twice in her life time. The flight of one queen with
several drones in air for copulation is called nuptial flight.
The honey bee family is an excellent example of social life. The different castes depend on
each other for survival as well as co-operating with the developing broods in the hive.
Hive or Comb:
The worker bees construct hive with the help of wax secreted from the wax-secreting glands
of the abdomen. They repair the cracks of the walls of hive with propolis (resinous substance
collected by bees from different parts of plants for use as glue) and balm collected from the
plants and is used in the construction of comb.
The propolis is used as a glue to bind broken parts, and balm is taken for polishing inner
walls. Each bee hive contains thousands of hexagonal cells arranged in two vertical rows.
These cells are of 5 types:
1. Queen cells:
These are a very few in number in a hive. They are larger than the other cells and vase-
shaped, and are situated at the margin of the comb. These cells are used for queen rearing.
2. Drone cells:
There are about 200 drone cells in each hive and are smaller than the queen cells. The drones
are reared in these cells.
3. Worker cells:
Majority number of cells is worker cells and each cell is about 5 mm across. The workers are
reared in these cells.
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4. Brood cells:
The larvae of the honey bee are reared in these cells.
5. Storage cells:
These cells are meant for the storage of honey and pollen.
Role of Hormone for Social Organization:
The mandibular glands of queens are situated in the head and open at the base of mandible.
The queen secretes a kind of chemical substance that inhibits the development of ovaries of
worker bees.
A British researcher, C. G. Butler found that the nature of the inhibiting substance was oxy-
deconoic acid, a substance with hormonal properties that controls the activities of all bees in
the hive. The queen bee secretes a pheromone by mandibular gland and attracts drones.
Language of Honey Bee:
Bees are known to have some method of communication amongst themselves.
Karl Von Frisch, a vienna born scientist and great experimenter on the behaviour of honey
bees, discovered the communication mechanism and decoded the ‘language of bees’ in 1946.
He was a Noble Prize winner in 1973. As soon as one bee finds a source of nectar, it
immediately conveys the source and direction of the source to other bees of the same
community.
They perform certain rythmic movements and emit odours that are easily received by other
bees. When the source is nearer to hive (within 100 metres), reporter bee or forager or worker
bee performs a round dance (Fig. 18.75A), turning in a circle, once to left then to right and
repeating the same movement for 1½ minutes in one place.
Round dance informs the distance of source of food which is less than 100 metres but cannot
give the indication of direction.
27
If the source is further away, the reporter bee performs a tail wagging dance (Fig. 18.75B). It
runs towards the direction straight ahead for a short distance, wagging the abdomen, makes a
360° turn towards left, runs ahead once again and turns right. This is repeated for several
times.
These dances are closely watched by other bees in the hive and then immediately they come
out in search of the source. The wagging dance informs their sisters of the hive both direction
and distance of the source of food (nectar or pollen) discovered by the worker bees and is
considered as language of bee. The direction of straight run indicates the direction of source
of food and tempo of the dance also indicates distance.
Odours play a vital role in their communication. Sudden death of queen bee is relayed to
60,000 or more bees of the hive in less than an hour. Healthy queen secretes an aromatic
substance called ‘queen substance’ which is licked off by her nurse bees.
28
When the queen dies the secretion stops and the absence of the queen substance is immedi-
ately relayed to all the members of the colony. The message being conveyed to all members
of the colony; they at once set about the vital task of rearing a new queen.
Anatomical Peculiarities of Onychophora:
A. Primitive features:
1. Onychophora are worm-like body covered with thin, flexible, chitinous cuticle.
2. Onychophora are sluggish in nature.
3. Head segments are comparatively small (3 head segments in onychophores but in true
arthropods head segments are 5 or 6).
4. Presence of segmentally arranged nephridia.
5. Presence of cilia in the reproductive tracts.
B. Sole peculiarities:
1. Segmentation indistinct on external surface.
2. Head appendages include a pair of antennae, a pair of jaws and a pair of oral papillae.
3. Texture of the skin is present.
4. Numerous, un-jointed, stumpy walking legs, terminated into a pair of claws, quite unlike
the parapodia of polychaeta.
5. Tracheae and disposition of the tracheal apertures are not arthropod-like.
6. Presence of a pair of slime glands opening at the ends of the oral papillae that secrete
proteinaceous adhesive substance and helps to capture the prey.
7. Lacking of blood pigments.
8. Subcutaneous haemal channels.
C. Salient features:
1. Caterpillar-like body, ranging from 5 mm to 15 cm in length.
2. Body soft and covered by a thin, flexible, chitinous cuticle which is moulted periodically.
3. Indistinct segmentation externally and marked only by the presence of paired, un-jointed,
hollow stumpy appendages (13 to 43 pairs according to species). These un-jointed walking
legs are called lobopods.
4. Each walking leg terminates in a pair of curved claws.
29
5. Integument with fine transverse wrinkles and with numerous conical large and small
tubercles.
6. Simple eyes, similar with that of an-nelidan polychaetes.
7. Head with 3 pairs of appendages including a pair of annulated antennae, a pair of claw-like
mandibles (jaws) which are the modified 2nd pair of appendages and a pair of oral papillae
(3rd pair of appendage).
8. A pair of slime glands are present inside the body which open to the tip of the oral papillae
that discharge the adhesive material, used for to capture prey and defence.
9. Body wall dermomuscular. Muscles are un-striated.
10. Reduced coelom.
11. Haemocoelomic body cavity.
12. Open circulatory system with lateral valvular ostia on the heart.
13. Elongated tubular heart which is surrounded by pericardial sinus occurring the entire
length of the body.
14. Delicate un-branched, rarely branched tracheal tubes open by means of small spiracles,
scattered irregularly. Spiracles are without any closing device.
15. A single pair of nephridia in each segment except the genital opening bearing segment.
16. Ladder-like nervous system. Brain is large, bilobed and situated dorsal to the pharynx.
17. Reproductive and excretory ducts are ciliated.
18. Dorsal coelomic gonads.
19. Sexes separate (gonochoristic).
20. Fertilization internal.
21. Oviparous or ovoviviparous with yolky or non-yolky eggs.
22. Viviparous with placenta.
23. Cleavage holoblastic in the eggs of viviparous species and superficial in the oviparous
forms which lay their eggs in moist condition.
24. Development direct.
25. Nocturnal and carnivorous in habit.
30
Affinities of Onychophora:
The features of Onychophora have made it difficult to place it within any one of the ten major
phyla. The detailed studies of Onychophores have now confirmed that in addition to its own
peculiar features it has characters common with three other large groups, Annelida,
Arthropoda and Mollusca.
An account of such relationship is given below:
A. Relationship with Annelida:
Similar features (Structural):
1. Segmentation in both is homonymous.
2. Presence of paired nephridia in almost every segment of the body.
3. Reproductive tracts are lined by cilia.
4. Skin is thin and flexible.
5. Dermomuscular body wall like Hirudinea. Body wall musculature smooth and composed
of circular, diagonal and longitudinal muscle fibres.
6. True head is absent.
7. Structure of the eye is same as in polychaetes. Simple eyes (Ocelli) rather than compound
eyes.
8. Hollow and non-jointed appendages like those of parapodia.
9. Slime glands and coxal glands correspond with the similar glands of polychaetes and
oligochaetes.
10. Straight gut with an anus.
11. Vermiform boy.
Embryological similarities:
1. Spermatozoan morphology resembles that of oligochaetes and leeches.
2. Meroblastic cleavage.
3. Gastrula by epiboly.
4. Elongated blastopore.
Dissimilar features:
1. Ventrally placed mouth in Onychophores.
31
2. Heart and ostia present in Onychophores.
3. Clawed appendages in Onychophores.
4. Presence of antennae in Onychophores.
5. Ladder-like nervous system.
6. Tracheal tube for respiration in Onychophora.
7. Absence of true metamerism in Onychophora.
8. Texture on the skin.
9. Disposition of the gonads and
Remarks:
In view of the anatomical peculiarities Grube (1874) placed the group under Annelida and it
appears that onychophora have evolved from the annelids, if not directly from them, from the
ancestral stock from which the annelids have evolved.
B. Affinities with Arthropoda:
Moseley (1874) demonstrated its relation with arthropods by showing the presence of
tracheae.
Structural similarities:
Following features show that Onychophora is more related to Arthropoda:
1. The appendages are provided with claws.
2. Locomotion is not annelid-like but takes place with the help of legs having definite
musculature.
3. Jaws are modified appendages.
4. Heart dorsal and tubular, perforated by lateral ostia.
5. Body cavity is a haemocoel, not a true coelom.
6. Absence of perivisceral part of coelom.
7. Body is covered with chitinous cuticle and is moulted.
8. Jaws are provided with striated muscles.
9. Presence of antennae.
32
10. Brain is large and resembles the brain of typical arthropods.
11. Presence of tracheae as respiratory organs.
12. Excretory organs closely resemble the green glands of Crustacea.
13. Salivary glands formed by the modification of nephridia.
14. Pattern of development is same as in other arthropods.
Dissimilarities:
In spite of these similarities, Onychophores differ in many respect from the Arthropods.
1. Arrangement of tracheae is not arthropod-like. Here in each segment there are numerous
permanently opened spiracles (no closing mechanism).
2. Jaw is the modification of second appendages and the movements of jaws operate from
anterior end and proceed towards posterior end.
3. Formation of skin is not like that of arthropods.
4. Segments behind head are simple and identical.
5. Segmentation not distinct in Onychophora.
6. Absence of malpighian tubules in Onychophores.
7. Simply, un-jointed, numerous stumpy legs in Onychophores.
8. Structure of eye is less complicated.
9. Two ventral nerve cords are widely separated and without true ganglia.
10. Body regions or tagmata are not well developed in Onychophores, which are well
developed in Arthropoda, e.g., in most cases the body is divided into head, thorax and
abdomen.
Remarks:
According to Sedgwick (1908) there is no doubt that the Peripatus belong to the Arthropoda
in all the above mentioned characters which are all of morphological importance. Develop-
mental features of Peripatus confirmed the view.
C. Similarities with Mollusca:
1. Slug-like appearance.
2. Ladder-like nervous system resembling that of chiton and lower prosobranchia.
3. Antennae tentacle-like.
33
Remarks:
Guilding (1826) first discovered a peripatus and considered to be an aberrant mollusc. But
according to many scientists the resemblances with molluscs are only superficial.
Time of origin:
Onychophora evolved from the marine fossil onychophoran-like organism Aysheaia

pedunculata from the Mid-Cambrian period about 520 million years ago.
Probable views regarding the Origin of Onychophores:
Different zoologists have put forward different views regarding the origin of Onychophores.
1. Hills (1930):
Only known fossil resembling Peripatus was found in the Mid-Cambrian period. In the Pre-
Cambrian era only fossils of soft- bodied, segmented worms, annelids were found. Then
according to Hills, in early Pre- Cambrian some tracheate arthropods underwent
specialization while others were less specialized to give rise to the Peripatus in Mid-
Cambrian.
2. Thomson and Ritche (1944):
Thomson and Ritche (1944) Opined that the Peripatus is a survivor of forms that were
ancestral to the tracheate arthropods and closely related to the annelids.
3. Snodgrass and Stromer (1944):
Snodgrass and Stromer (1944)—said that the Onychophora originated from the ancestral
form of both annelids and arthropods.
4. Tiegs and Manton (1958):
According to them, Onychophora evolved from generalised lobo-pod ancestor, such as, this
line of evolution is not followed by any other arthropod. They have again stated that the Mid-
Cambrian Aysheaia pedunculata has generally been accepted more or less as a marine
ancestor of modern terrestrial onychophorans.
34
5. Shrock (1958):
Shrock (1958)—remarked that peripatus is not the ancestral form of annelids nor gives rise to
modern arthropods but is a separate autonomous isolated group evolving from the ancestral
trochophore.
Zoological Importance of Onychophora:
Onychophora show a great zoological importance because:
1. They furnish an example of discontinuous distribution and
2. They represent an example of living connecting link between the two phyla—Annelida and
Arthropoda.
Systematic Position of Onychophora:
The characters of Onychophora have made it most interesting from the point of view of
evolution. It is an oldest terrestrial group which probably originated from some marine
ancestors.
It has attained a number of features for terrestrial life, i.e., internal fertilization, viviparity,
semi-solid excretory product, less permeable skin, etc. But at the same time the structure of
spiracles speaks about its limitation on land life and thus shows its primitiveness.
The resemblances with annelids are probably the examples of convergence. Onychophorans
have a mixture of morphological characteristic features which make them effectively cross
between the Annelid worms and the Arthropods. But our modern understandings suggest they
do not represent a missing link between the annelids and the arthropods.
Presence of variously developed specialized characters in Onychophora, such as trachaea

with open spiracles and the origin of jaws do not support the above view that Onychophores
represent a missing link. Instead, like the Tardigrades they are considered as a separate line of
evolution and arose independently from some forgotten ancestor.
35
Again the common characteristic features of Onychophora regard a common relative to

annelids and arthropods. Hence, it is regarded as a living connecting link between two phyla
and it is undoubtedly an ancient form but not an ancestor of arthropods.
Kaestner (1967) has stated that the Onychophora probably represents an early lateral branch
of the evolutionary line terminating in the arthropods. Peripatus are also called living fossils
because they truly represent archaic animals and have changed little in their body shape for
about 500 million of years.
A cladistic analysis which places the Onychophora in an intermediate position between the
Polychaeta and the Tardigra-Arthropoda clade. The current view is that the Onychophora
represent a sister group to the Arthropods on the basis of morphological, palaeontological and
molecular data.
According to Marshall and Williams (1972)-Onychophora can be rightly described as an

aberrant arthropods or highly modified annelids and can be placed all the species under
twelve genera of Onychophora in a separate subphylum “Onychophora” under the phylum
Arthropoda. But absence of jointed chitinons exoskeleton and jointed segmental appendages
do not support the inclusion of Onychophora within Arthropoda.
Some peculiar features of Onychophores which support neither annelids nor arthropods, and
demand a separate phylum status. Ruppert and Barnes (1994) pass the remark that
Onychophorans are not usually considered arthropods rather a phylum animals closely related
to arthropods.
Classification of Onychophora:
Phylum Onychophora [Gk. onyx or onychos = claws, phoros = bearer] Approx. less than 200
species.
Origin:
Early Cambrian period.
Characters:
1. Commonly known as velvet worms or walking worms. [Peripatus. (Gk. Peripatos =

walking about; Gk. Peripatein = to walk about, stroller).
2. Bilaterally symmetrical, metamerically segmented and protostomous coelomates.
3. Body soft and Caterpillar-like.
4. Head is not clearly differentiated. Tagmatization is not well pronounced.
5. Body may grow from 5 mm to 15 cm in length (e.g., Peripatopsis torquatus).
6. Externally the segmentation is denoted only by the presence of short paired (14-13 pairs)
un-jointed stumpy walking legs (lobopods). The legs are terminated into curved claws.
36
7. The head bears 3 paired appendages:
(i) One pair fleshy annulated antennae
(ii) A single pair of jaws (2nd pair of appendages) and
(iii) a pair of short oral papillae (3rd pair of appendages), situated adjacent to the jaws.
8. Eyes are represented by ocelli rather than compound eyes.
9. Integument is thin and the chitinous cuticle contains varied ring-like striations.
10. Body colouration is blue, green, orange or black with papillae and scales.
11. Muscles are un-striated.
12. A straight gut with an anus.
14. Respiration is carried by tracheal tubes, open through the small spiracles.
15. Spiracles without any closing devise.
16. Slime glands discharge adhesive material through the openings of the oral papillae used
for prey capture or defence.
17. Malpighian tubules are absent.
18. Excretory organs are paired segmental coelomoducts.
19. Sexes separate (gonochoristic).
20. Fertilization internal.
21. Usually viviparous but may be oviparous or ovoviviparous.
22. Reproductive and excretory ducts are ciliated.
23. All are terrestrial and are found in moist habitats.
Classification of the Phylum:
The phylum include approximately about 200 species which are distributed among 49
genera and 2 families:
(i) Peripatopsidae and
(ii) Peripatidae.
The defined categories above the family level are absent due to conservative body features of
the Peripatus.
37
Fossil History of Onychophora:
The most primitive fossil of Onychophora is marine, such as the genus Aysheaia (Fig. 19.9)
of Mid-Cambrian period, whose fossils are found in rocks which are probably 520 million
years old.
Other fossils have been recorded from the Baltic and Myanmer Ambers. Onychophorans
became adapted to land before the Late Ordovician period and the two living families became
distinct by the Late Triassic. Other genera are Hallucigena, Tertiapatus, etc.
Characters of the living families:
I. Peripatopsidae:
Number of legs varies from 14-19 pairs; legs with complete spinous pads are 3; the absence
of a diastema on the inner side of the jaws; primary dermal papillae without a constriction
nephridial opening on 4th and 5th pairs of legs in between third spinous pads; genital opening
between or behind last pair of legs, oviparous or ovoviviparous.
Distribution:
They are found in the chilie, South Africa, Australasia, New Britain and New Guinea.
The family includes 39 genera and the revision of the family is being done by C. Brockmon,
A Reid, R. Gleeson and H. Ruhberg.
38
39
II. Family Peripatidae:
Number of legs varies from 19-43 pairs; legs with complete spinous pads 4-6; the presence of
a diastema on the inner blade of the jaws; primary dermal papillae with a constriction;
nephridial openings on 4th and 5th pairs of legs in between third spinous pad; genital opening
in between the legs of the penultimate pair. Skin pigment brownish, extracted by alcohol;
ovoviviparous or viviparous.
Distribution:
Mexico, Central America, Northern South America, Galapagos Islands, West Indies, West
equatorial Africa and South East Asia.
The family includes 10 genera and these are:
(i) Eoperipatus,
(ii) Epiperipatus,
(iii) Heteroperipatus,
(iv) Macroperipatus,
(v) Mesoperipatus,
(vi) Oroperipatus,
(vii) Peripatus,
(viii) Plicatoperipatus,
(ix) Speleoperipatus and
(x) Typhloperipatus.
Clark (1913) divided the genus Peripatus into 4 sections:
(i) Plicatoperipatus,
(ii) Macroperipatus,
(iii) Peripatus and
(iv) Epiperipatus.
40
41

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CORE-3 UNIT-I B. Sc.

The organs associated with aquatic respiration are:

6. Branchiostegite or gill cover

(iii) Origin of gills in crustacean:

(iv) Structure of gills in Crustacea:

(v) Types of gills in Crustacea:

B. Based on the mode of attachment, the gills may be of three types:

Podobranch attached with the coxopodite of the thoracic appendage. In Macrobrachium (=

(vi) Number of gills in Crustacea:

(vii) Modification of gills in Arthropoda:

Mechanism of gill respiration:

Other devices of aquatic respiration:

The aerial respiration takes place in terrestrial arthropods.

The organs for respiration are the following:

4. Pseudotracheae or air tubes

Two types of tracheae are seen:

The tracheae originate as the invagination of the body wall.

(a) Structures of trachea and associated parts:

(i) Each trachea is a tube with walls made up of polygonal cells.

(b) Classification of tracheae:

(i) In adult insects, the tracheal system is of one kind.

Tracheal system openings to the exterior by 8 or more pairs of functional spiracles.

It may again be subdivided into:

Here, either one or two pairs of spiracles are functional.

It includes divisions like:

(c) Mechanism of tracheal respiration:

(d) Modifications of the tracheae:

Other devices of aerial respiration:

3. Pseudotracheae or air tubes:

Many crustaceans perform rhythmical contractions of intestine—taking in and expelling out

A combination of book-lung with trachea is seen in spiders. Some aquatic members of

Type # 1. Ametabolic Metamorphosis:

Type # 2. Heterometabolic Metamorphosis:

Type # 3. Holometabolic Metamorphosis:

In Lepidoptera, Coleoptera, Hymenoptera, Diptera, Siphonoptea, etc., the young which

1. Termites are pale-coloured and usually blind except macropterous forms.

3. Well-developed eyes in reproductive castes.

4. Ocelli are either touching or away from the eyes.

5. Soft-bodied females have much enlarged abdomen.

7. In winged forms, the fore-wings and hind-wings are same sizes.

The lower termites include the following families:

It includes the most primitive living termite, Mastotermes darcviniensis. It is found in

Known as subterranean termites except Coptotermes, Reticulitermes. Some mound-builders

3. Social Organization of Termites:

A typical colony possesses nymphs (semi-matured young’s), workers, soldiers and

4. Caste System of Termites:

1. Macropterous or large winged or primary reproductive forms (Fig. 18.91):

2. Brachypterous neotenics or short winged or substitute reproductive forms:

3. Apterous neotenic reproductives or wingless forms or replacement reproductives:

5. Food and Feeding of Termites:

The social insects exchange food in two ways:

(i) Stomodaeal trophallaxis and

(ii) Proctodaeal trophallaxis.

(i) Wood (cellulose) eating termites:

(ii) Soil-feeding Termites:

The soil-feeding termites are Cubitermes, Termes, Capritermes (Subfamily Termitinae,

(iii) Fungus-growing termites:

6. Swarming and Formation of a New Colony by Termites: