Dominance of Insects

Chapter 2: Dominance of insects:
Diversity and abundance are the two ways in which one can look at the dominance of insects.
In an everlasting inquisitiveness, pioneered probably by the earliest human beings, man

has been asking the question of what is that organism? As a consequence, all organisms
encountered are being named. This process of collecting and naming organisms has been an
important pastime for man. This modest beginning has been now formalised and procedures
have been set for naming all kinds of organisms. The tenth edition of Systema Naturae by the
great Swedish biologist, Carolus Linnaeus, has been considered as the starting point for these
names. This date is presumed to be the first of January, 1759. However, in the last 250 years are
so, since this cutoff date, scientists have collected and formally described an enormous number
of kinds of organisms. In addition, the process of collection and description of organisms
continues to this day, that it is extremely difficult to tell exactly how many species have been
described. As a consequence, different people have tried to estimate the number of species of
different groups of organisms known to science
Insects clearly out number all groups of organisms and account for an estimated 1
million kinds. As against an estimated 785,000 species of all other kinds, insects as a group thus
represent an imposing 56 per cent of all organisms known to science. That apart, the fact that
every day new species of insects continue to pour in has puzzled many a biologist to ask the
question, how many kinds of insects are there on Earth?
One such estimate was done by Terry Erwin, a tropical biologist. Way back in 1982, he
stirred a hornet’s nest by coming up with an astonishing figure for the diversity of life forms on
earth. Erwin studied the canopy insects of Luehea seemannii, a tropical tree in a Costa Rican
forest. He collected all insects by fumigating 19 trees of this species. Then he looked at the
available number of species of beetles, a sub group of insects. By classifying, the beetles he
encountered, on the basis of an assigned host specificity and functional group, he calculated the
number of species of beetles per species of tree and worked out a figure for the 50,000 species of
trees of the world. The figure was a whopping 32 million insect species, considering that beetles
form an estimated 40 per cent of all insects. This figure, made many a biologist to sit up and
start wondering as to what is the diversity of life forms on earth for more than one reason. This
figure is important for many reasons. First is from the point of view of quenching our own thirst
for knowing how many species are there on earth. Secondly, since the last 250 years, the effort
of thousands of biologists has been successful in enlisting an estimated 1,785 thousand species of
all organisms. If a reasonable estimate of the number of species of organisms available on earth
can be made, that will help us know the requirement of taxonomic input for describing the
remaining fauna and flora. More importantly, by losing forest cover worldwide, due to human
population pressure, we can make a reasonable guess of what is the genetic resource we are
likely to lose in the form of number of species and what efforts are needed to conserve this
biological resource.
Erwin’s estimate of 32 million species of insects might simply mean an impractical

daunting task to describe all species of insects. Further, it also meant that loss of every species
of tree simply means a loss of nearly a 1,500 species of insects. That is an enormous erosion of
our genetic resource! Many a biologist thus became upset with this figure. But, Erwin had made
several mistakes in his calculations and included many sensitive values, which by altering
slightly could be shown to have given out highly variable estimates. Erwin, however, was not
the first to make such an estimate. Long before Erwin, 1974 to be precise, Robert May, a British
1
A.R.V. Kumar : Introduction to Entomology

biomathematician had made a simple estimate. He believed that most of the temperate fauna is
well known and that most of the known species by and large represented the temperate fauna.
Further, he believed that the tropics are at least twice as rich as temperate environments of the
world. Therefore, one million species of insects known represent only one third of all kinds of
insects present on earth. Clearly, adding another million for the other life forms, a total of four
million life forms are there. This simple initial estimate being the guide, Erwin’s estimate was
obviously very startling. Hence, several others set out in search of more comfortable figures for
the number of species.
Two scientists, Hodgkinson and Casson were the next to make an estimate. By
intensively searching for members of another group of insects, the true bugs in North Sulawesi,
they identified the proportion of known and unknown number of species of bugs in Sulawesi.
Using this figure and considering an estimated seven per cent of all possible insects to be bugs,
they calculated the number of species of insects possible on earth. They came up with an
estimate of a possible 2.5 to 3.5 million species of insects and to add another million of other life
forms, insects account for anywhere from 71 to 77 per cent of all life forms on earth. Their
estimate, by far the best to date, was also supported by other estimates by another strong critic of
Erwin, Nigel Stork.
Clearly, these estimates point to a possible 70 to 75 per cent of all extant life forms to be
insects. This might be one important reason to tell why most biologists are entomologists, the
people who study insects. One other field, as an exception, could be medicine, where more
biologists are involved for obvious reasons.
Apart from this staggering diversity, insects are also extremely abundant and widely
distributed. Some soil ecosystems may harbour more of mites than insects. Barring this
exception, insects are the most abundant life forms in almost all terrestrial ecosystems. But then
even in tropical soils it is estimated that up to 1 million individual insects may be found per ha.
Similar estimates of absolute numbers of insects are not really available for many ecosystems.
But alternative measures do suffice to explain their abundance. One such measure made by
Wilson is in relation to a group of insects, social insects. Social insects consist of diverse groups
of insects exhibiting colony life and include, ants, wasps, bees and termites. These insects can be
so very abundant that in Tropical South American forests, they are expected to comprise of 40
per cent of all animals and 70 per cent of all insect by biomass. This amounts to 57 per cent of
all animal biomass to be made up of insects. Being small, compared to many other groups of
animals, this biomass would make their numbers astronomical. While this is one measure,
people have also attempted other measures of abundance of insects in other ways. Locusts, a
group of pestiferous insects, occur in large swarms spanning some times several square
kilometers. One such swarm of locusts has been estimated to weigh around a stunning 70,000
tonnes made of locusts weighing on an average 2 mg apiece.
Another alternative measure of abundance is the colony sizes of social insects. A

colony of a social insect is akin to a human family, where, parents live with their children. Many
social insects especially ants and termites, have colony sizes in excess of a million. In a recent
study, a multi-queen Japanese ant has been shown to contain a stunning 306 million individuals
occupying several tens of hectares. The African driver ant, Anomma wilworthi, is known to have
up to 22 million ants per colony. These figures are simply astronomical compared to the
miniscule family sizes of modern day humans, a supposedly social being.

Apart from their diversity and abundance, their ubiquitous presence, makes them the
organisms of frequent encounter by man, thus attracting the attention. But there are more
reasons for man to bother about insects.
Insects form a dominant group among living organisms. The following table gives an
estimate of known species of organisms in different groups.
Table 1: Worldwide number of species of organisms known to science in different taxa.
Taxa No. of species in thousands
1. Kingdom - Monera 5
2. Kingdom - Fungi 69
3. Kingdom - Plantae 308
Flowering plants 248
Algae 27
Ferns 11
Mossess 14
Liver worts 8
4. Kingdom - Animalia 1387
Invertebrates 1161
Protozoa 31
Platyhelminthes 12
Nematoda 12
Mollusca 50
Echinodermata 6
Annelida 12
Coelenterata 9
Arthropoda (other than insects) 200
Hexapoda 1000
Other invertebrates 13
Vertebrates 42
Lower chordates and fishes 19
Birds 9
Amphibians 4
Reptiles 6
Mammals 4
Total 1764
Above figures indicate the abundance of organisms on the basis of estimates of

described species. However, the recent estimates of described and undescribed species of insects
in the world is expected to be as high as 2.5 million, while all other organisms put together may
be 1.0 to 1.5 million species. This amply demonstrates the enormous diversity of insects.
Why are insects dominant? : The reasons for enormous diversity 3

Close on the heels of appreciating the dominance of insects, biologists have also
wondered about the causal factors that have made insects singularly the largest biological
representatives on earth. Many explanations have come forth. Among them are their : a)
antiquity, b) small size, c) functional wings, d) the possession of exoskeleton, e) metamorphosis,
f) diversity of food habits, g) great ecological adaptability and h) enormous reproductive ability
are some widely cited reasons. Apart from these, certain other explanations such as i)
decentralised nervous system, ii) tracheal respiration, iii) hexapodous nature, and iv) specialized
offence and defense mechanisms are also cited by certain authors. A cursory look at this list
suggests that almost every feature of insects is tooted as possible reason to explain the
dominance of insects. However, the fact that insects are diverse and abundant, any feature that is
common to most insects has the potential to be listed as a causal factor. It is extremely difficult
to separate out the cause and effect relationship and also it is reasonable to guess that no one
factor can be responsible for the great diversity of insects. Nevertheless, it may be worth
examining each of these listed factors.
A. Antiquity :
The planet Earth, it is estimated, is about 4.2 billion years old. However, some recent estimates
suggest it to be much older. The primordial Earth, having been formed some 6 billion years ago
is one such recent estimate. Given that Earth is 6 billion years old, about half way through its
life the primordial life must have originated. A history of 3.2 billion years has been suggested as
the time span for the oldest unicellular fossils, which appeared like beads of bacterial cells.
From then on the evolution of life progressed at a slow pace until the great Cambrian explosion,
around 575 million years Before Present (BP). Somewhere during this period, about 600 million
y BP, the protoarthropods appeared on the scene; almost at about the time when the closest
ancestors of the arthropods, Annelida and Onychophora originated. The disjunctly distributed,
small Phylum Onychophora, it is now established, is the closest to the present day arthropods
through RNA studies. However, the protoannelidans (Protostomes) are possibly the next closest
ancestors.
The oldest hexapodan fossils are that of collembolans and the thysanurans. These
fossils are believed to be around 420 million years old and mark the arrival of hexapodans on
Earth. However, the oldest definitive insect fossil, Rhyniognatha hirsti, is estimated to be 407 to
396 million years old. The extinct Meganeura monyi is another example of an oldest insect.
However, in the recent past, Misof et al. (2014) through studies of a large set of protein coding
genes have unambiguously placed the insects as having Remipedia (Crustacea) as the closest
sister group. Besides, the study also unambiguously suggested the time frame of evolution of
different orders of insects. Study pointed out that first insects possibly evolved around 479
million years before present.
This historical account suggests clearly that there are many extant groups that appeared
much earlier to hexapods. Nevertheless, the 480 million years of history has contributed greatly
for the evolution of insects. By considering the extant families of insects and their trophic
organs, Labandeira and Sepkoski argued that the insects have demonstrated a great evolutionary
resilience unlike other groups of organisms. The insects apparently diversified unhindered
through out this history and their growth pattern, considering the families of insects, was never
affected. This is unusual in that many other groups of animals that evolved much later are near
about that time have gone extinct. The remarkable resilience simply suggests that once evolved,
the insect families continued to survive through time to remain extant to this day. Labandeira
and Sepkoski argued that this is unique to insects and has contributed to the present day diversity
of insects.
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Further, Farrel considering the chrysomeloids, demonstrated the possibility of
diversification of insects in association with the diversification of flowering plants. Considering
that the flowering plants having a history of around 150-175 million years, it is possible that the
greatest degree of speciation in insects must have occurred during this period. Other supports
available also suggest this possibility. First, about 40 per cent of all known insects are
phytophagous and nearly all of them feed on flowering plants such that phytophagy has provided
an important adaptive zone for the insects to occupy and diversify. Mitter et al. demonstrated
this aspect considering the phytophagous and non-phytophagous sister groups in different taxa of
insects, where in 11 out of 13 sister groups verified indicated greater species among the
phytophages. Another aspect of not necessarily phytophagous groups to have evolved is the
evolution of social insects. Different social insects also came on the scene by about the time the
flowering plants evolved. Clearly, indicating that the flowering plants have supported the
evolution of not just the phytophages but also other possibly interdependent species of insects.
As a result, the speciation of insects seem to have been propelled to the greatest extent only
during roughly the last one third of their history on earth. Therefore, the antiquity in itself is
unlikely to have contributed for the observed diversity of insects, considering their species
numbers. If antiquity alone were to be responsible for the great diversity of insects, one can ask
the question of why other much older groups have not become as abundant! For example, the
closer relatives, annelidans or the onychophorans are hardly a match for the arthropods. Thus it
is improbable that the antiquity in itself is a sufficient reason to explain the diversity of insects.
Interestingly, some insect species are also shown to have a great resilience, probably
coupled with enormous adaptability. Two important examples are the domestic cockroach,
Periplaneta americana, which is believed to have remained unchanged for nearly 250 m. years.
Similarly, the primitive Australian ant, Nothomyrmecia macrops, it is suggested has remained
unchanged compared to its 90 million year old fossil. However, the newest entrant to this list is
the Amazonian "Ant from Mars", Martialis heureka, being represented in a 120 m y old fossil.
B. The small size:

Insects demonstrate a great range in their body sizes. However, due to their enormous variation
in form, it is difficult to make proper comparisons. One convenient way of measuring their sizes
would be to look at their lengths. Considering the length, the insect sizes range from 0.2 mm or
less of the minute parasitic Mymar species to the longest known insect, a female stick insect
from China, discovered in 2016, Phryganistria chinensis Zhao, at 62.4 centimeters, is the longest
known insect in the world.
This suggests an enormous range of over 3000 folds. Perhaps no other

group of organisms shows this kind of a difference in size range. Nanosella fungi
(Ptiliidae) is among the smallest insects in the world measuring just about 0.25 mm
and less than 0.4 mg. However, Megasoma, a beetle, is called so because it has a large
body (mega = large + soma = body)! A matching species is the Goliathus goliathus
(Scarabaeidae : Cetoniinae) measuring 8 cm is among the heaviest weighing up to 100 grams, as
also the Megasoma. Ornithoptera alexandrae, the butterfly named after queen Alexandra, has a
wing expanse of 28 cm. Quite meaningfully it is called a bird wing (=Ornitho/ptera). Likewise
Attacus atlas, the atlas moth, has a wing span of 30 cm and Dynastes hercules is 19 cm long and
weighs nearly 35g. Interestingly, the largest of any known insect is a fossil dragonfly,
Meganeura monyi, which had a wing span of 75 cm. These samsonites (= Ghatodgajas)

notwithstanding, majority of the insects are small and less than 2 cm in length. Western Pigmy
Blue from the USA at 1.5 cm across the wings, is considered the smallest butterfly in the world.
These size ranges notwithstanding, insects in general are small (Fig. 1). Majority of the
insects are less than 2 cm in length. Although many other organisms are smaller on an average
than insects, insects have the largest number or even proportion of small animals. This small
size of insects is considered an important factor for the enormous diversity of insects. Many
biological features of organisms are size related. The most important among them are the
reproductive rate, generation time and resource requirement for completing a life cycle. Often
these reasons are cited as the general rules and an explanation is given about the abundance of
insects.
Irrespective of these considerations, size has been an important factor, in explaining the
diversity of organisms in general. For example, the smallest mean size among mammals is that
of rodents and rodents are the most diverse among the mammals. Similarly, Passeriniformes is
the largest order of birds that has the smallest birds in size. On the whole, considering all the
organisms also, the size and diversity relationship holds. Robert May explored this aspect and
demonstrated that the size and diversity of organisms by and large follow an exponential
N o . o f s p e c ie s (lo g n o .s )
relationship.
Distribution of Terre stria l organisms by size
1,000,000
100,000
10,000
1,000 Insects
100
10
1
Size of Animals (log mm)
Fig. 1: The number of species of all terrestrial organisms classified according to length.
The line represents the general pattern of size distribution (equation given in the text).
May showed that a rough estimate of the size of all organisms plotted against their
diversity in a double log plot follows the equation, Y = AX –S, where Y, is the number of species,
X, the mean size in milli meters, A, the constant of proportionality and S, the slope of the curve.
The slope indicates the rate at which the number of species changes with increase in body size
(Fig. 1). May also demonstrated that the rule applies to many groups of organisms. In general
‘S’ takes the value of two. Interestingly, insects occupy the left hand side of the graph. Insect
sizes are expected to range from 0.2 mm to as much as 33 cm. The smallest known insects such
as Mymarid parasites are parasitic and grow inside the eggs of other insects. Large insects such
as the giant stick insects are free living. However, majority of insects are less than 2 cm in
length.

This kind of small size is expected to confer many advantages on insects. Small size
allows insects to live in extremely small spaces throughout their lifetime and on small quantities
of resources. Endoparasitic wasps or plant gall forming thrips remain confined in a few cubic
mm, getting their food and completing the life cycles from within the galls. They also complete
their life cycles in short time spans i.e. faster generation turn over rate. A small parasitic insect
may complete a generation in at the most a few days, while an elephant requires several decades.
Large animals have much lower reproductive capacities than the smaller animals. The maximum
lifetime fecundity of a human female, for example, can be at most 27 babies. The highest known
for a non- social insect is as much as 18,000 offspring for the members of the moth genera
Abantiodes and Xylentus. However, for some social insects such as a termite (white ant) queen
laying so many eggs is just half a day’s work! The generation time multiplied by the
reproductive potential brings in an enormous amount of baby insects per insect per unit time as
compared to many other groups of organisms. Greater the number of individuals produced per
female per unit time, the greater will be the opportunity for speciation and thus, small size is
considered a key causal factor for the greater diversity of insects.
Considering both the short generation time and the high fecundity, one can clearly see
that the insects can potentially achieve much more total number of births per unit time than many
other animals. Given this, one can visualise greater births with greater potential for
accumulation of mutations. In essence, a potential for accumulation of genetic changes thus
providing increased opportunities for speciation.
Why are insects small?

Despite being diverse, why are they so small? Is their a limitation for their body size?
People believe that two factors are responsible for their small size. One is their mode of
respiration and the other the process of molting associated with metamorphosis.
Vertebrates and some invertebrates such as earthworms use blood as the medium of
oxygen transport to different parts of the body. Insects like all other arthropods have a tubular air
supply system that connects the external atmosphere directly to individual cells within the insect
body. In order to meet this requirement, the tubes, tracheae, etc. branch out at different positions
into smaller and smaller tubules called tracheoles. Some of these tracheae have associated
collapsible air storage structures called air sacs. The largest of the tracheae open out into the
atmosphere through gates called spiracles. In order to prevent dust from entering into the system
and also control air movement to a certain extent, these spiracles are provided with filters and
closure mechanisms. The entire system is akin to a water or gas supply system found in cities
and towns, but the difference is in respect of the interconnections. The entire set of tubes and
tubules together form a complex network in majority of the insects, with spiracles, numbering up
to 10 pairs. The exchange of gases is achieved both through active muscular movement and
through passive movement of oxygen and carbon dioxide. Passive movement occurs due to
differences in the concentrations of the two gases resulting in differential partial pressure that
sucks the oxygen in to the system. This is an extremely efficient system under low oxygen
situations, permitting many insects to survive in very unusual conditions such as deep inside a
tree trunk or in a seed/ fruit, etc. This system is also more than a 100,000 times faster in
permitting the insects to grow in sizes that can not be thought of by several other invertebrates
such as flat worms, nematodes etc., that respire through the skin, as dissipation rates of oxygen
through a liquid medium will be extremely slow compared to dissipation through gases.
But the system becomes more and more inefficient as the size keeps increasing due to the longer
distances that oxygen has to travel through the pipes. The separation of the oxygen and the
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carbon dioxide is limited, although, some insects have evolved mechanisms of achieving a fair
degree of efficiency in separating out the two gases. As the distance through which oxygen has to
dissipate keeps increasing, the difficulty in separating the two and also in transporting the
oxygen at a faster rate becomes increasingly difficult. With increasing distance from the
spiracle, the size (diameter) of the tubes keeps reducing so that the movement of air becomes
increasingly difficult. An alternative way of checking this reduction is to keep all the terminal
tracheoles a constant size and then to increase the sizes of connecting tubes towards the spiracle.
Even this has difficulty. As the distance of the spiracle from the cell keeps increasing the sizes of
the tubes at every bifurcation needs to increase, and soon the connecting tube has to be so large
that there will hardly be any space left for other anatomical systems. Thus a system that is
extremely efficient at a certain size of the animal becomes increasingly unmanageable beyond a
size limit. How then do the above-cited large insects constitute to exist in nature. Even here
these organisms have adopted a simple strategy of keeping all tissues within a distance of less
than 1.0 to 1.5 cm from the main trunk line tracheae that are directly connected to the spiracles.
The longest insect is a narrow stick, thus falls in line with this explanation! The heavier beetles
tend to be disproportionately flat.
The tracheal system is therefore considered a great limitation for increasing the size in
arthropod body plans in general and insects in particular.
A second and equally important reason is metamorphosis. Insects pass through a

variety of growing systems. Three major types are recognised. The first is a situation, where the
mother lays the egg and the egg hatches into a baby insect, called a nymph, which is a small
miniature version of the mother, the adult. This kind of growth is particularly seen among
wingless and very primitive insects such as the silverfish. The second type involves the same
three stages, egg, nymph and the adult, but with one difference. This difference is primarily in
respect of adults that are winged and the nymphs are miniature adults, but without fully
developed wings. The latter is the most special, for it has four stages. An egg is followed by a
larva, which in turn is followed by a pupa and the adult. The larval stage is completely unlike
the adult and is mostly vermiform without a trace of the winged nature of the future adult.
Adults are fully winged and the pupa is just a packed sac that transforms the wingless larva into
a fully winged adult. Among these stages, it is only the larva that grows. In some species, the
growth can be enormous and could reach over ten thousand times that of the youngest larva. But
larval growth is not a smooth transition from small to large size. All insects just like other
arthropods are covered by a hard skin made of chitin and strong structural proteins called
sclerotins. The ‘skin’ also serves as a skeleton for supporting all the internal organs such as the
muscles and it determines the shape of the larva. If the larva has to grow, it has only one method
- to expand its hard skeleton. This does not happen smoothly, for the structure is a fixed
structure, but for small areas between the segments, which are to some extent stretchable.
Therefore, the growing insect has the only option of removing the old skeleton and having a new
and enlarged one each time it has to achieve some growth. This process is somewhat similar to
that of snakes shedding their old skin and is called molting. After hatching from the egg a larva
may undergo several molts - most often five - and in extreme cases up to 22 molts before
becoming a pupa. From the larval stage to the pupal stage and then from the pupal stage to the
adult stage two more molts are required for the transition to take place so that the winged adult
emerges from a vermiform larva.
Moulting in insects consists of a series of steps where in, the old skin is separated and
the new skin is secreted below. The separated old skin is then slowly shed and the newly secreted
and larger skin is allowed to harden. At this stage the insect faces the difficulty of supporting its
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entire weight on a fragile yet – to - harden skeleton. Entomologists, people who study insects,
believe that this transition stage puts a great limitation on the size of the insect. For the insect
has to support a great amount of weight and then give it shape. Before hardening, gravity pulls
down much of the weight so that a large insect gets bogged down by its own weight and the
resulting animal will be a pancake or ‘dosa’ of an insect. This may be the reason why many
large extant insects are fairly flat! Gravity is an important limiting factor for terrestrial animals.
This is why for the largest animals, the whales, are aquatic to overcome this difficulty of
supporting their immense weights against gravity. Water, after all, provides the necessary
buoyancy to support their enormous weight.
Knowing why insects have to be small, one can ask the question how did the fossil
dragonflies, achieve their gigantic sizes? A simple answer could probably be that their larvae
like the present day dragonflies were aquatic. Irrespective of these considerations, small size has
helped insects to be winners in the numbers game of species diversity.
C. Functional wings
Flight is a very specialized adaptation seen in insects, birds and bats among the extant groups.
But for these, the only other group that had wings was the extinct Pterosaurus. But several other
animals such as flying fox, flying fish, flying squirrel and the flying frog have the capacity to
soar. All these groups, except the insects are vertebrates. Birds, bats and even the extinct
pterodactyls all have one pair of wings, which are essentially the modified fore limbs that they
have sacrificed for wings. Insects on the other hand have true wings that have evolved
independent of any limbs. Wings of insects have developed over a period of 300 m years, as
against the pterosaurs 225 to 75 m years before present, birds of 125 m years and bats of just
around 50 m years. The oldest winged insect is expected to be at least 350 m years. In essence,
insects are the monarchs of the ‘air’ for a long time, until the first vertebrates developed the
capacity to fly. They still are the largest group of animals with the ability to fly and greatly out
number 8,700 species of birds and 1000 species of bats put together.
In an estimated one million known species of insects, primarily wingless forms just
number around 1 per cent, which includes both the non-insectan hexapods and the true insects of
the orders Archaeognatha and the Thysanura. Essentially all the other groups/ orders of insects
are primarily winged. Surprisingly, a few of these groups have lost their ability to fly
secondarily. These secondarily wingless forms include such groups as Fleas and Phthiraptera. A
point to be noted is that both these groups are basically parasitic on vertebrates. Similarly, an
entire family of insects may be wingless, flies of the family, Nycteribiidae and bedbugs of the
family, Cimicidae form the examples for such cases. Alternatively, wingless forms may be
frequently found in almost all the remaining orders of insects. For example, many female moths
are flightless. Similarly, many beetles are flightless. Despite many such examples, their
proportion among the insects is negligible. Thus, flight is a feature strongly associated with the
insects in general and as many as 99 % of all insects are primarily winged forms. Consequently,
just a comparison of the relative numbers of winged and wingless forms of insects
overwhelmingly suggests that flight must have been an important factor that has possibly
contributed for the enormous diversity of insects.
What advantages do insects get by possessing flight?
Flight provides a means to escape from danger of many kinds. The danger could stem
from such annual feature as changing weather, drying of streams, and other aquatic bodies that
makes the insect impossible to survive under extreme situations. Similarly, the growing larvae
9

might eat up so much food just like Kumbhakarnas that no food might be left for the next
generation of the insects. Only way then is to fly away from the area in search of more food.
Many crop pests that we know of also exhibit similar behaviours. The well-known Brown plant
hopper, Nilaparvata lugens and many species of aphids remain wingless through out the crop
growth period and start multiplying at rapid pace. Once the crop starts senescing, the insects
growing at that time of the season, will start sporting wings and start dispersing. Butterflies and
moths are very well known for this kind of migration in search of better living places. The
famed mass migration of Monarch butterflies, Danaus plexippus, to over come the severe winter
of the US and Canada to Mexico is a celebrated phenomenon seen year after year. Monarchs
migrate over 3,000 miles during this mega journey in search of a living place. Vanessa cardui,
the painted lady, is another species known for marathon journey of over 4,000 miles. But the
most recent discovery is that of a near globe trotter, that the scientists suspect may have a range
from Texas to Tamil Nadu. Globe Skimmer, Pantala flavescens, migrates from Southern India to
Maldives, Seychelles and then to the African continent tracing the rains for pools of water to
breed. The entire distance is a whopping 18,000 km. Large surface area of their wings possibly
enables them to ride the prevailing winds, often at altitudes of more than 1000 m, while dining
on aerial plankton and small insects. However, the Globe skimmer takes many other dragonfly
friends along, on its journey across the old world, albeit in small numbers. These celebrated
cases exemplify the utility of flight in overcoming abiotic stress and also the biotic stress like not
finding the food.
Similarly, flight provides an opportunity to escape from many predatory animals. Birds
and bats are serious predators of insects, that may catch even a flying insect. Insect flight is
much more dynamic and provides a plethora of movement options to overcome even such
predators. Similarly, amphibians, reptiles and many species of mammals all depend heavily on
insects as their major source of diet. Flight is one option that limits the predatory influence of
such species.
Flight also helps in another important way. Majority of the insects reproduce by sexual
means. Flight thus provides an opportunity for the male and female to meet for reproductive
purposes. A courting male butterfly pursuing a female or an Aeshnid dragonfly, patrolling a
waterbody is a treat to watch.
Flight provides another great reason to explain the diversification of insects. Many
airborne insects get carried away by wind currents to thousands of miles. Imagine one such
gravid female landing in an alien place where no possibility of finding another male of the
individual of the species exist. If by chance the food for the species is available to some extent,
the progeny of such a species may start growing there for several generations and form a
population, that over time, becomes completely different from the original stalk from where the
progenitor of this population arose. One can wonder whether such a possibility exist. In fact it
does. Air borne insects, have been trapped at heights of over 10,000 meteres into the
atmosphere. There have been a large number of records of the American Monarchs landing in
Europe during the late 1800s and early 1900s when there populations did not really get
established due to lack of possibly food plants. However, as the food plants became available,
the Monarchs are there every where now in the world!
D. Exoskeleton
Insects are not unique in having Exoskeleton. Many animals, including some mammals also
have exoskeleton. Order Insectivora among mammals has many examples such as Pangolin, that
possesses an exoskeleton. But this exoskeleton is a supplement to the endoskeleton in these
10

animals. Among the invertebrates, all arthropods have an exoskeleton. Similarly, many other
groups of invertebrates also have exoskeleton. The basic arthropod exoskeleton is similar to the
chito-proteinous insect exoskeleton. Primitive arthropods originated nearly 200 million years
before the first insects. Insect exoskeleton is thus a pre-adaptation that existed previously in
other arthropods. To that extent exoskeleton is not unique to insects and therefore, might be an
unlikely cause for the radiation of insects. But there is a caveat. One can draw a conclusion that
arthropods are the most numerous of all animals because they have a chito-proteinous
exoskeleton.
Accepting this possibility, what are the advantages of possessing exoskeleton? The
arthropod exoskeleton is expected to provide many advantages. The exoskeleton can vary
greatly in its elasticity and hardness characteristics. Hence, the chito-proteinous nature of the
exoskeleton provided for the development of wings in insects and also allowed for development
of a compulsive metamorphosis to achieve growth. These properties also served as useful
features in having complex and versatile legs, antennae and other external structures associated
with the skeleton. In addition, through the wax layer, the exoskeleton also provided for the
impervious cover to prevent desiccation. Perhaps this is the greatest innovation during the
course of their evolution. This is an important feature that helped insects to colonise terrestrial
habitats and even to conquer air, where the dangers of desiccation are immense. Further, the
poor attitude control in flight seen in insects would have greatly affected the survival chances of
insects but for the hard exoskeleton that prevents damage to many insects, during collisions. The
anti-microbial property of the Exoskeleton protects the insects from many microbes and the
hardness from many predators and parasitoids.
All these features confer a great advantage for insects to live the way they are, but yet,
exoskeleton in itself is an unlikely character that contributed for the diversity of insects, as it
remains a pre-adaptation seen in other arthropods.
E. Metamorphosis
Significant morphological differences noticed in successive stages of an organism during its
post-embryonic development is termed as metamorphosis. Metamorphosis is again not unique to
insects as it is widely prevalent in arthropods and many other groups of invertebrates and
vertebrates such as amphibians.
The rigid exoskeleton restricts the growth of insects. Therefore, a growing larva moults
repeatedly to achieve growth. The number of molts vary greatly in different groups of insects.
During the course of these molts, many insects show distinct shifts in the morphological states.
Such a shift represents metamorphosis.
Insects exhibit varied levels of metamorphosis. There are insects that do not show any
variation in their morphology from the young that hatch out from the egg to the adult stage
except for the development of gonads. Such insects are referred to as Ametabola, meaning those
without metamorphosis. These insects show three developmental states, egg- larva – adult. The
pre-adult larval state is called a nymph. All non-insectan hexapods, Archaeognatha and
Thysanura, which together constitute roughly about 1 % of all hexapods show this kind of
developmental pattern. Note also, that these insects primarily lack wings.
The second group consists of winged forms. These insects as young ones are also
similar to adults when they hatch out of eggs. However, one major difference is seen. The
difference is in respect of the wing that is only rudimentarily seen in the pre-adult stages of the
11

insect. Such insects are termed as Hemimetabola, in that they show limited metamorphosis.
Even in this group of insects, three developmental states are seen. These include egg-larva and
the adult. However, the hemimetabolan insects are also of two different types. There are insect
species with hemimetabolous features where the larvae live in aquatic habitats and are endowed
with the necessary modifications to suit aquatic life. Larvae in such insects are called naiads.
However, the adults are terrestrial. Mayflies, dragonflies and damselflies fall into this category.
Such insects are called true Hemimetabola. Alternatively, some insects have rudimentary wings
in their pre-adult stages but they live in the same habitat as that of the adults. Such species are
grouped as Paurometabola. The immature stages of the paurometabolous insects are called
nymphs. Grasshoppers, leafhoppers, many species of bugs etc. represent this category.
More amazing is the third category of insects that are called Holometabola. These are
the insects that possess wings in their adult forms and are considered to exhibit complete
metamorphosis. But the sign of the existence of wings is totally lacking in these insects during
their pre-adult stages. Wing development in these is said to take place internally, and are called
Endopterygota, as opposed to the Exopterygota, comprising of hemimetabolous insects. In
addition, the pre-adult stages beyond the egg stage consist of one more stage, a quiescent one,
called pupal stage to facilitate the transformation of the larva into a winged adult form. Thus
there are four stages in the development of these insects viz., egg- larva- pupa and the adult.
In all these insects, the egg is a quiescent stage and from the egg, the larva hatches out.
The larva has the only function of growth and therefore is the only feeding stage in many species
of insects, especially the holometabolous insects. Many moths are known not to feed during
their adult stages. However, many adults are known to feed and live much longer than the
larvae. Adults have the task therefore, of achieving the dispersal, because of their ability to fly
and also due to other additional features which make them very different from the larvae in
holometabolous insects. Of course, in addition, they have the primary task of reproduction. The
pupae in these insects is expected to facilitate the transformation of a wing less vermiform larva
into a winged adult. In the course of achieving this goal, the pupa remains quiescent. This
quiescent stage is excellently adapted to facilitate the extremely low metabolic activity under
adverse environmental conditions. Such that, an insect can remain inactive for long times until
the conditions become suitable.
What then is the advantage of metamorphosis to facilitate the diversity of insects? In

many species of insects, the larvae and adults live in very different conditions. In
Hemimetabolous insects as already said the larvae are aquatic and the adults are terrestrial. That
means the metamorphosis helps the insects to exploit the benefits of two different environmental
conditions. Similarly, adults and larvae also live on different food resources. The dragonfly
naiads prey upon small insects and fishes in the water while their adults feed on insects in
terrestrial systems. Similarly, a butterfly larva feeds on plant leaves and the adults feed on a
liquid diet of nectar from the flowers. The examples like this can keep accumulating. All these
go to show that the metamorphosis helps insects to exploit different resources, both habitat wise
and also food wise. This reduces the competition among the larvae and adults of the same
species and helps broaden the horizon of their resources, to get the best of both the worlds.
Ability to exploit different resources helps better survivorship in terms of surviving at a stage
when the food of the other stage is not available. This is considered as an important factor in the
diversification of insects.
One other feature facilitated by the metamorphosis is the phenological adjustments. We

know that the weather is not the same through out the year. As a consequence, the organisms
12

adjust to the conditions of living in such a way that their morphological states are phenological
states change to suit the prevailing weather. Plants for example, put forth new flush of leaves, or
come to flowering and fruiting at different times of the year. Imagine an insect with larvae
capable of feeding on new flush of leaves and adults capable of feeding on flowers. Knowing
that the plant comes to new flush and flowering at different times of the year helps us appreciate
the importance of Metamorphosis in facilitating the timing of occurrence of these different stages
of the insect. One such example is the whitegrubs. Whitegrub adults commonly referred to as
May beetles or June beetles occur soon after the rains in Summer. These beetles feed on leaves
of perennial trees like neem, ficus, etc. They lay their eggs in the soil and the larvae hatch out
soon from these eggs. The larvae are adopted to feed on roots of annual plants such as ragi,
groundnut, vegetables, etc., many of our cultivated plants. Larval movement, however, requires
some amount of soil moisture. Therefore, the larvae should occur at a time when the soil is
relatively moist and the annual plants are available for them to feed on their roots. Adults
therefore, emerge at a time when the conditions are suitable for the larvae in the first place and
also when the plants have relatively fresh leaves for them to feed on, lest the leaves become
poisonous to adults of white grubs. Similarly a resource that is highly ephemeral and limited in
season such as pongamia flowers that hardly last a month in the field are efficiently exploited by
the midge that depends on it. All this is made possible by metamorphosis!
Therefore, the metamorphosis is considered an important factor for the diversification

of insects.
Additional causes attributed for the diversity of insects:

F. Great ecological adaptability
Exoskeleton that also contributed for the tracheal system of respiration, has contributed
for the colonization of terrestrial habitats by insects. Arthropods were the first animals to
colonise terrestrial habitats. This clearly indicates the importance of Exoskeleton and the
tracheal system of respiration in promoting this possibility. Considering insects, a majority are
terrestrial in their habits.
However, a few species accounting for less than 3% of known insects are aquatic and
all these forms are primarily found in fresh water bodies. Many groups of insects live in aquatic
habitats in their larval stages. Almost all mayflies, damselflies, dragonflies, stoneflies,
megalopterans, and trichopterans belong to this category. However, in many other groups, only a
few species are aquatic either as larvae or both in adult and larval stages. Beetles, bugs, flies,
and in exceptional situations, hymenopterans, lepidopterans, cockroaches and grasshoppers live
in aquatic systems. In aquatic systems, insects are usually adopted to living in lentic conditions,
still water bodies and a few however, are adopted to lotic or flowing water conditions.
Nevertheless, only the midges have managed to occupy the deep layers of water bodies. As all
adults require air for breathing, the insects living in deeper layers may face the problem of
reaching the surface quickly to avoid suffocation. Aquatic insects have developed a large variety
of secondary adaptations to live in water.
Exceptionally, few species of insects are also found in marine habitats, but largely as
surface dwellers and in the inshore marine systems, reefs or coasts. Open seas are rarely
habitable to insects. Halobates spp., a group of scavenging bugs living off shore in the marine
habitats, perhaps, are the only group of organisms to exhibit the greatest degree of adaptability.
One other remarkable insect to live off shore is Pontomyia sp., a midge. Both Halobates and
Pontomyia are uniquely adopted for living on the surface waters of marine habitat. Most other
13

marine forms generally occur in areas where the tidal effects are minimum. Marine forms
account for less than 0.1 percent of insects known. Interestingly, primarily aquatic forms have
rarely colonized the marines systems, indicating that life in fresh water does not help colonise
marine systems. On the other hand it is almost always the basically terrestrial groups that have
taken to marine habitats. Managing the osmotic pressure apparently is not a limitation for
insects to occupy marine habitats for many species live in more brackish salt lakes and swamps.
Difficulty of managing the repeated and unpredictable tidal waves seems to be the major
limitation. This is also possibly explained by the fact that the violent upper reaches of the large
rivers such as Amazon account for extremely small number of insect species.
But insects are the lords of the terrestrial systems. The only group to challenge their
dominance are the mites, in certain soils. Ranging from the Arctic to Antarctica, they are found
in every land mass. Over 300 species, mostly flies live in North of 75° parallel. A few species
live in the Antarctica. Insects are also widely distributed from sea level to over 6,000 M above
mean sea level. in the Himalayas. Similarly, from the most arid desert to the wettest ever green
forests, the insects appear to be the most dominant group of animals.
Temperatures is one important factor expected to influence the distribution of

organisms. Insects have exhibited an amazing ability to survive in conditions as different as –20°
C to as high as +40° C. This being the normal range, anywhere from 40 to 45° C is the range
which actually causes many insects to exhibit heat stupor. However, some Ephydriids, a group
of flies, live in unusual heat conditions such as hot sulphur springs, where the temperatures
might be as high as 50 to 55° C. Another exceptional set of species are those that inhabit the
cold polar habitats. Antarctica is perhaps the land mass with the least number of insects. Mostly
these are the insects that are dependent as parasites on some birds and seals. A louse that infests
seals for example can almost survive through constant – 2° C when the seals are submerged in
water for long days. These insects probably use the body heat of their hosts. Another
exceptional group is the Grylloblattids. Grylloblattodea (Suborder of Notoptera ; rock crawlers)
is an order of insects with about 12 known species worldwide. These species are adapted to live
in such cold conditions that holding a rock crawler by hand would cause their death due to body
temperature of human beings. These are specially adapted to live in temperature ranges of less
than 12° C.
Other extremely unusual habitats are the petroleum pools. Some Ephydriids, such as
Psilopa petrolei, are specially adapted to live in these conditions. The larvae of these insects
feed on insects that fall in to these pools.
But the most striking factor that has contributed for their diversification is the ability to
colonise terrestrial habitats. Few animals have successfully colonized the terrestrial habitats.
This is evident by the fact that as many as fourteen phyla are marine, eleven have their
representatives in fresh water systems while only six phyla have terrestrial forms. Thus the
marine habitats account for greater number of phyla of organisms than the terrestrial habitats.
Obviously, specialized adaptations are required for animals to colonise terrestrial habitats which
pose some serious problems for the survival.
The major limitation of terrestrial habitats is the danger of desiccation. Vertebrates and
Arthropods in general are special in overcoming this problem. In fact, even those organisms that
managed to survive in terrestrial habitats have not been successful in overcoming the danger of
desiccation completely and tend to largely inhabit many habitats that provide a relatively safe
environment such as soil and marshes, that help keep the danger of desiccation to the minimum.
14

Crustacea, Nematoda and Annelida, for example, despite colonizing the terrestrial habitats
cannot live in exposed conditions. Among the Arthropoda except the extinct Trilobita, all others
have successfully adopted to over come the problem of desiccation.
Table 2. Relative distribution of different animal phyla in major ecological habitats.
Sl. No. Phylum Marine Freshwater Terrestrial

1 Porifera Dominant Less
2 Cnidaria Dominant Less
3 Ctenomorpha Purely marine
4 Platyhelminthes Dominant Less
5 Rotifera Not found Largely Not found
6 Nematoda More Common Abundant
7 Nemertea Dominant Found Not found
8 Molluska Dominant Found Found
9 Annelida Largely Abundant Found
10 Onychophora Not found Not found All species
11 Arthropoda Found Found Abundant
12 Bryozoa Dominant Found Not found
13 Brachiopoda All Not found Not found
14 Phoronida All
15 Echinodermata All
16 Chordata Found Found Dominant
But many insects occupy extreme dry weather conditions. They can be found in deserts
or they can survive through hot summer remaining active. They can also remain exposed on
plants and on soil surface, which is unusual among other organisms. How do insects manage
this?
The danger of desiccation is particularly more serious for insects compared to

vertebrates. Being small they have greater surface area per unit volume than the vertebrates.
Having large surface area simply means large area for losing body water, which can be a
constant process. Such a problem is much more serious for even smaller organisms such as
nematodes. Nematodes therefore, have developed a mechanism of surviving despite losing body
water to the driest possible condition. In this condition, all their physiological activities are
stopped completely but yet they can potentially remain alive for several years. Cyst formation is
one such mechanism in plant parasitic nematodes. However, they become active when the
conditions are ideal for resuming physiological activity. Such a mechanism of slowly or rapidly
losing body water to stop all physiological activity but yet retain the ability to become active
once the conditions are ideal is called Cryptobiosis. Cryptobiosis is fairly common in micro-
organisms but less common among the metazoa.
Insects are not known to exhibit cryptobiosis, probably they do not need! However, an
African Midge, Polypedilum vanderplankei, is known to exhibit cryptobiosis (Hinton, 1977) in
its larval stage. Hinton kept the cryptobiotic larvae of the midge in –270° C and at 102° C for
one minute and observed that once the conditions are ideal, the larvae came back to their own.
Further, he also observed that cryptobiotic larvae survived for over three years and became active
once the ideal conditions were provided. This amazing capability is the sole property of this
species among insects.
15

But many insects show a variety of alternative mechanisms of overcoming the adverse
weather conditions. In these conditions, they resort to stopping all kinds of physiological
activity except those that are limited to the bare minimum of sustaining life. Such a situation is
referred to as Diapause. Insects have mastered the art of tiding over adverse ecological
conditions through diapause. A large variety of insects exhibits this feature. Basically, two kinds
of diapausing exists. If the insect has developed a mechanism to overcome high temperature
conditions, then such a diapause is called Aestivation. On the contrary if the mechanism is
aimed at overcoming low temperatures then such a diapause is called Hibernation. An insect
may tide over adverse situations in any of its four growth stages. But it is believed that the
evolution of metamorphosis has favoured the achievement of this phenomenon better through the
pupal stage. Aestivation is seen by and large in the pupal stage. However, the larval tiger beetles
can aestivate when the conditions become too hot and dry and return to their normal activity
once the conditions become ideal. But, hibernation is more commonly seen among the larval
stages than in other life stages of insects. Getting cues mostly through changing durations of
day-night cycle, the insects show diapausing states. Day-night cycle is used as the cue. For the
duration of day and night keep changing with the seasons and the trajectory of these changes
indicate the impending changes in the weather conditions for the insects. Diapausing state helps
them to cut down on energy investment at a time when the resources are likely to become limited
or unavailable. Further, additional adaptive features help the insect to reduce moisture loss.
However, it is important to know that the diapause, although helpful in overcoming adverse
weather conditions, the mechanism of diapause initiation might have taken place much earlier to
the diapausing stage or generation and as such might not necessarily guarantee that at the time of
diapause, the insect is in fact experiencing adverse ecological conditions. Nevertheless,
diapausing evolved as a mechanism to overcome adverse ecological conditions.
Not all insects may have diapausing stages in their life cycle. Then what are the basic
adaptations that make life in terrestrial conditions possible for insects? Insects exhibit an
enormous diversity in their adaptations to suit the terrestrial life. These can be broadly divided
in to morphological, physiological and behavioural features.
The most striking of all the morphological adaptations is the exoskeleton. Insect or
arthropod exoskeleton is characterized by three layers; the inner non-cellular basement
membrane, middle cellular epidermis and the outer cuticle. Based on the structure, the cuticle is
further divisible in to two layers. Inner Procuticle and the outer epicuticle. The Procuticle again
structurally comprises of two layers, the inner Endocuticle and the outer Exocuticle. Exocuticle
is the one that shows the greatest degree of sclerotisation and hence the hardness associated with
the exoskeleton. The outer most layer of the cuticle is the Epicuticle. Typically, Epicuticle
comprises of four layers. Arranged from inner to outer, the layers are protein cuticle, cuticulin,
wax layer and the cement layer. The cuticulin is believed to set the limit of body size for a
molting insect. But the most striking innovation of the arthropods in general and the insects in
particular is the development of the wax layer that made it possible for insects to colonise the
terrestrial habitats. Wax layer comprises of two sub layers of which the inner one has a compact
and systematic arrangement of individual wax molecules such that it forms an impermeable layer
for water and is called the monolayer, due to ordered arrangement of single layer of wax
molecules. Development of this layer is crucial for the colonization of different terrestrial
habitats. Insects that are not facing the danger of desiccation, like those that inhabit soil, aquatic
or semiaquatic conditions are devoid of this layer or have a poorly developed monolayer. On the
other hand, those that live in dry deserts and show arboreal life are generally endowed with a
strong and almost impermeable monolayer. The reinforcing of the monolayer is probably by the
16

less compact outer wax layer. Thus, removing or damaging the wax layer is a technique of
managing insects. If the insect can be made to pick up abrasive material repeatedly, then the wax
layer gets so damaged and the desiccation increases such that the insect dries up! One can even
try it out with cockroaches.
The integument, that also forms the exoskeleton of insects is again a feature that has
contributed indirectly in many ways to facilitate the adaptation of insects to a variety of habitats.
Integument lines a number of internal organ systems. One such organ system of importance in
the present context is the tracheal system. Tracheal system is the respiratory system of insects
that helps them to get the much needed oxygen for their metabolic functions. This system in
insects is very different from the vertebrate system of a lung and the use of blood for the
transport of oxygen to different parts of the body. The tracheal system comprises of
invaginations of the integument into tubes that carry air into the body and the tubes keep
branching off akin to a water or gas supply system found in cities such that ultimately the finest
of these tubes reach every cell of the body and merge (or anastamose) with it to supply the
oxygen. The point of entry of the air is through an opening in the body wall called spiracle and
the tubes are called tracheae, hence the name tracheal system. The system is considered very
efficient when the body size is small as is the case with insects. However, the system also poses
a danger. Because the air moves in and out of the body directly from the atmosphere, in dry
habitats the out going air takes away much of the body water thus increasing the danger of
desiccation. Insects have managed to solve this problem by many mechanisms. One mechanism
is to reduce the number of openings, the spiracles, such that some insects have only one pair of
active spiracles. Normally they may have up to ten pairs of spiracles. Having a small number of
spiracles helps in reducing the total amount of air entering and going out of the body thus
reducing the desiccation rate. Alternatively, the relatively free living insects, have developed a
trap door type of complex spiracles as against the simple opening seen in case of some soil
dwelling insects such as many primitive apterygotans.
Another method was in developing complex spiracles with valve closer mechanisms
and strong muscular control. This mechanism helps in regulating the air traffic in and out of the
body by varying the number of active spiracles. Insects also have evolved much stronger air
traffic regulating methods such as keeping the spiracles closed for long hours so that the required
oxygen keeps dissipating in to the body through the integumentary surface and then the
accumulated carbon dioxide would be released in bursts once in a while. This mechanism is
generally adopted in quiescent stages of the insect such as pupae, which have no mechanism of
garnering the required water.
One other technique of overcoming the danger of desiccation is to resort to behavioural

mechanisms. Insects exhibit a variety of behaviours to overcome desiccation. Just like human
beings do, they may drink water, move to shade, burrow into soil or hide. Drinking water for
example is extremely essential for a flying Uzi fly. Thus, it may be possible that just by reducing
the accumulation of free moisture in the trays, and by keeping a water bowl with a drop of
insecticide in the rearing houses, it is possible to greatly reduce the Uzi fly menace in the silk
worm rearing houses. This can be tried by you!
Certain others exhibit stilting. This behaviour has two functions. A stilting tiger beetle
for instance may drastically reduce the body surface exposed to direct sun light such that the
body temperature does not rise. Secondly, due to this reason, the over all rate of loss of body
water reduces.
17

The evolution of specific feeding habits and associated development of morphological
and behavioural adaptations also contribute for reducing desiccation. The best example for this
is the evolution of suctorial mouth parts so that many insects of the order Hemiptera depend
heavily on plant sap as the source of nutrition. This in fact not only reduces the desiccation
problem but results in excess water availability. To overcome this problem many of these insects
have developed ingenious methods of bypassing excess water in the gut from the food and then
to concentrate what needs to be processed. Others have used this opportunity to cover
themselves perennially with watery froth so that even the harshest sun can not affect their water
balance. If you find some spittle on the grass, be assured that there is a bug inside, the spittle
bug.
Living situations is another mechanism of overcoming the danger of desiccation. Soil

of course, is an excellent habitat that provides a great insulation against a variety of dangers.
Insects have found greater number of habitats. They bore through plant tissues and live inside,
especially in their larval stages. They may live inside leaves, pods, flowers, fruits, stems or roots
of plants. In fact, the dangers of desiccation can best be seen, if we can have a variety of crop
plant with its wax genes silenced. Such plants may not have the problem of leaf miners.
Because, leaf miners use the plant wax as the cover and may get dried up if wax layer is not
found on the plant. But remember that wax is the greatest evolutionary innovation for any life in
terrestrial ecosystem, be it an animal or a plant, human beings not excluded. Therefore, not
having wax layer could also be equally dangerous for the plant to thrive!
Many flies, moths, beetles specialize in occupying these kinds of habitats. Few others
not only enter inside and live inside the plant tissues but also turn it into a cozy home by
modifying the very structure of the tissues. Many gall forming Aphids, psyllids, thrips, and
moths specialize in this kind of behaviours. These well insulated galls made of modified plant
parts provide protection against not only the vagaries of weather but also against many natural
enemies.
If water is not available for drinking, or not adopted for drinking then they have
developed alternative mechanisms of getting water. Many such physiological adaptations exist
among insects. One such method is to get water from the food. Insects extract water from their
food by specialized structural designs found in the gut. Many caterpillars also have the
specialised ability to physically squeeze out water from the defecating material using their
specialised rectal papillae. These can help reduce the water being lost along with the faeces.
They can have great abilities of taking the water out of what they eat. Many dermestids, a kind
of beetles, live in extreme dry conditions feeding on almost dry stored food items such as many
millets. This special ability perhaps contributed for many insects to colonise the food stores
where most others fail to survive due to lack of sufficient moisture. This explains the reason
why we have so many insects affecting our stored material, including the stored grains. It is not
just that they can feed on extreme dry organic matter and live but also they have a special ability
to take out water and dry even a water rich food item. This special ability of tenebrionids comes
as a handy natural technique of managing the poultry filth, just dry it using insects, so that it is
easy to handle.
All organisms defecate nitrogenous waste material. But defecation results in loss of
moisture! If there is too much moisture the simplest way to get rid of extra water is to defecate,
or urinate, which might be a liquid or at the most a paste. Which of course all organisms do!
Wasteful Nitrogen under such situation is lost generally as ammonia. In an extreme dry
condition, defecation can be very costly and life threatening. So, under such conditions, many
18

insects avoid water loss by defecating in the form of Urea, that helps save water. Some others
might even avoid defecation and simply store in some corner of the body! Some bugs and
butterflies have mastered this art.
Further, in case of dire needs, they may simply break down the stored food material and
manage to get the required metabolic water. Lipids among all the stored material are expected to
yield the highest amount of water per molecule broken down.
In essence, various mechanisms have been developed by insects to overcome the

problem posed by desiccation in terrestrial habitats that helped them to colonise this vacant
ecological zone. Once they colonised the terrestrial habitats, there was less competition from
other groups, and great diversity of food available apparently contributed for the extra-ordinary
diversity of insects. Side by side, their amazing feature to occupy every conceivable terrestrial
habitat is also considered to have contributed greatly for the diversity of insects.
But there is a problem in this argument. Ecologically, any two species are not supposed
to be having an identical requirement. Each species is therefore, is expected to have different
"niches". A niche is an 'n' dimensional ecological hypervolume in which a species lives. The
dimensions may include food, environmental conditions such as temperature, moisture
requirement, sunlight, living spaces, etc., that one can go on identifying for a species or a set of
species. Given that considering all the finite dimensions, any two species are expected to be
different in their niche requirement. If the niches are the same for two populations, then the two
populations fall in to the same species. If the niches are different, then, we call them as two
separate species. Given this argument, on the basis of the niche theory, as no two species can
occupy the same niche, the number of niches keep on proliferating as the number of species
increase. In other words, the very essence of speciation hinges on the fact that populations keep
colonising newer niches or ecological zones. As a consequence, the process of expanding
ecological niches is a feature associated with the number of species. Therefore, the explanation
becomes tautological (equivalent to the question of which came first, egg or the chicken?). In
other words, we would have said the same thing if some other group were to be equally
numerous.
Nevertheless, colonisation of terrestrial habitat definitely provided an opportunity for

the insects to evolve at a rapid pace. This was in part facilitated by the specialised abilities to
conserve moisture.
G. Diversity of food habits:

Insects exhibit a great diversity of food habits. They eat virtually everything that is organic by
origin. The following are some examples:
Detritivory - feeding on decaying matter - ex. Collembolans, Thysanurans, cockroaches, Flies,

psocids, beetles, etc. Larvae of rhinoceros beetle feed on decaying dung in dung pits. House fly
larvae also similarly feed on decaying matter.
Dead wood and leaf litter – logs, furniture, carpets, etc. - Beetles, cockroaches, silverfishes,
thrips, etc.
Fungivory - Feeding on fungi - Many species of flies, some species of ants, bugs and beetles,
termites, thrips, etc.
Coprophagy – feeding on animals defecation - dung – Beetles, flies etc.

19

Necrophagy – feeding on carcasses of dead animals – Beetles, flies, etc.
Nectarivores and Pollen feeders – Beetles, flies, bees, wasps, etc.
Phytophagous - Plants feeding - Moths and butterflies, flies, beetles, wasps, bugs, grasshoppers,
etc.
Carnivory - Predatory – Bugs, beetles, wasps, praying matids, etc.

Parasitoids - on other insects - wasps, moths ( exceptionally few), flies etc.
Parasitic on higher animals – Flies (mosquitoes, flies, etc.), Phthirapterans (lice), bugs
(bed bugs and kissing bugs ), etc.
Miscellaneous – cockroaches, silverfishes, flies, etc.
processed material of organic origin – stored food, clothing, furnishings, museum specimens, etc.
– beetles, flies, psocids, silverfishes, moths etc.,
Insects have an extraordinary ability to utilise almost anything that is of organic origin.
But remember that food is a part of the ecological diversity and the earlier tautological
explanation holds good, when food is considered as a part of the ecological unit.
But recent studies have indicated that this is not correct and that the food, if not all, has
played a significant role in the diversity of insects. Nearly 40 per cent of all insects feed on
plants and almost this entire set of insects primarily feed on angiosperms, the flowering plants.
Surprisingly, most families of insects and nearly 32 of the 34 possible types of mouth parts were
already in place by the time the angiosperms originated some 150 m years before present, as the
fossil records indicate. Yet flowering plants have been attributed as the important reason in the
recent past for the enormous diversity of extant (living) insects. Studies have shown that,
considering sister groups of phytophagous and non-phytophagous species, the plant feeding
insects out number those of the non-phytophagous groups. An exceptional group is the group of
wasps and bees (the Hymenoptera) where the plant feeding insects are less numerous than the
non-phytophagous groups. Incidentally, the group consists of large many parasitoids of insects
that primarily utilise the phytophagous insects as their hosts. The other group is that of thrips.
Barring these two groups, the entire community of the sister groups identified show that the
phytophagous groups are larger, or more speciose than the non-phytophagous sister group.
Clearly, this indicates that the phytophagy must have played an important role in the
diversification of insect species and especially the phytophagous group. Further studies have
also indicated that there is a strong correspondence between the phylogenies of some beetle
groups with those of the flowering plants on which these beetles depend as the source of food.
Hence, there is reason to believe that the origin and diversification of flowering plants strongly
contributed for the increase in the species numbers of insects.
In addition, the evolution of angiosperms coincided with the evolution of some of the
groups of insects that are today directly or indirectly depend upon these plants. These are the
social insects. An estimated 3,000 species of termites (150 m y BP), 8,000 species of ants (135
m y BP) and 20,000 species of bees all originated by about the time the angiosperms originated.
The present day insect world, is believed to be ruled by these social insects. They contribute
greatly for the animal biomass in any ecosystem and serve as the important source of food for a
large number of organisms including some plants that are specially adapated to live on them.
Further, considering Hymenoptera, the exceptional group discussed above, one can see that large
many of these species are all parasitoids of insects. Large many of them are parasitic on
20

phytophagous insects. Thus, the diversification of phytophagous insects must have contributed
for the evolution of this group. One among the members of this large group is the bees. The
bees, numbering about 20,000 species worldwide are pollen and nectar feeders. Nearly 75-80 %
of all flowering plants are expected to depend on insects for their pollination and bees play the
major role in this respect. Essentially flowering plants are their energy sources. Consequently,
in a way even the non-phytophagous groups also diversified along with the plant feeders, during
the time when the plant diversity increased.
Plants have many adaptations to limit their exploitation. In fact, the challenge is very
formidable for several reasons. It is necessary that a small insect to remain feeding on the plant
need to have a good anchorage to remain on the plant and then to feed on it. Even among
insects, not all can do that. But insects have over come this challenge by developing specialised
mechanisms exemplified by a great diversity of pre-tarsal structures at the tip of the legs to gain
anchorage on the plants. Larval forms such as caterpillars for example have developed
specialised crochets to get this anchorage. Anchorring on exposed parts of plants may only
increase the dangers of desiccation. This we have already discussed. Plants also have poor
nutritional status compared to other forms of energy sources. For example, the phytophagy is
supposed to have originated from pollen feeding insects. Pollen is a much richer source of
protein than many parts of the plant. Similarly, carnivory provides for higher protein sources
than the plants. The insects that had to adapt for phytophagy were required to readjust their
physiology to suit this poor nutritional status. Insects have succeeded in this aspect. More
formidable is the plant defenses. Plants, having been anchored to a single place, do not have
mechanically defensive structures. Many physical defenses do exist. But the challenge was the
defensive chemicals they possess. In fact, so diverse is the arsenal of their defensive chemicals,
plants now serve as the important guide for developing new medicines to fight many ailments of
human beings and also serve as the sources of insect killing chemicals. But insects have even
managed this imposing challenge to expand their feeding to the newly available food as the
flowering plants originated on earth. Thus, being competition free in the "New ecological
adaptive zone" represented by plants, the insects were able to diversify greatly. The challenges
posed by the plants started posing increased degree of resistance such that over time, the insects
also started diversifying such that each challenge broken provided a new adaptive zone and new
area for colonising. But such plants which last the existing defense started adding more
defenses. Thus, a type of an eternal war has ensued between the phytophagous insects and the
plants – the famed "Coevolutionary arms race". As the plants developed new defenses such
plants probably grew in to newer species and thus it is possible that the diversification of plants
and the insects is taking place simultaneously leading to the 250,000 flowering plants in 150 m
years and the staggering 400,000 phytophagous insects, not to mention the large majority of
predatory and parasitic insects that depend on them.
This is a stuation is similar to the colonisation of terrestrial habitats by the insects. A

new ecological zone once occupied provided greater avenues for diversification.
H. Enormous reproductive ability
Higher reproduction rates lead to higher possibilities for mutations to accumulate. As a

consequence, the chances for speciation increase. This aspect has been discussed above, as part
of the small size.
I. Decentralised nervous system
21

Vertebrate nervous system, where a centralized control system exists, is characterized
by a large brain. But in arthropods, the system has been greatly decentralized. Although they
have a brain, smaller brain like structures, that are formed due to integration of sets of neurons
are found through out the body. Mostly, these ganglia are segmental, meaning found in every
segment. In fact, when ever there is confusion regarding the body segmentation, if we can
clearly recognize, the ganglia, that can be attributed to individual ancestral segments.
Ganglionic nervous system thus provides the opportunity to decentralize the control
system in the functioning of body metabolism. There can be several consequences of this
feature. First, the size of the brain is kept low and consequently the size of the head is kept low.
That solves many mechanical complications that would have arisen. Decentralization allows for
each ganglion assuming different responsibilities of regulation such that the individual segments
function probably better. Decentralisation also allows for reduced investment on nervous tissue
to achieve the same role as a central nervous system plays.
Although, these advantages are envisaged, it is extremely hard to visualize, how such a
mechanism of nervous system really contributes for the diversity of insects. First of all
ganglionic nervous system is an ancestral character, not unique to insects. All Arthropoda have
ganglionic nervous system. More strikingly even their poor cousins, Annelida, Onychophora and
Tardigrada also have an almost identical nervous system. That leads to a simple question of why
the poverty of diversity pervades these cousins of Arthropoda, if ganglionic nervous system is
important for the evolution of this great diversity if insects. Thus, it is very unlikely that the
decentralized nervous system has any significant role to play in contributing for the great
diversity of insects.
J. Tracheal respiration
Tracheal respiration is again a feature that has been attributed by many entomologists as
the reason for diversity of insects. Tracheal system of insects has the important roles of
conservation of moisture. But at the same time, the system becomes less and less and less
efficient as the size of the insect grows. However, for a small insect the system aptly fits and
facilitates greater body size relative to the animals that breathe through the skin.
Exoskeleton being hard does not permit breathing through the skin and therefore, as an
alternative to a basic body plan that is covered by hard exoskeleton, the respiratory function has
been achieved in Arthropoda and the velvet worms (Onychophora) through tracheal system.
Therefore, tracheal system is more an offshoot of the evolution of exoskeleton in Arthropoda and
the ancestral Onychophora. Thus, the trait is also not unique to insects and remains a pre-
adaptation.
K. Hexapodous nature
Hexapodous nature of insects is expected to provide better body balance in movement
and thus has been ascribed by some entomologists as a feature that contributed for great diversity
of insects.
Many animals have different numbers of locomotory organs such as the legs.
Vertebrates can have two just like us or four limbs like all the known tetrapods. The Myriapods
several hundreds to almost a dozen. Arachnids have four pairs of limbs. Similarly, the number
of limbs can vary greatly among animals. Most surprisingly, all these organisms have achieved
superb body balance despite varying numbers of legs. An examination of our own mechanical
devices and furnitures, one can clearly see, reveals that four legs are perhaps the best for
22

achieving a good balance. If anything, having additional legs is only an added cost to the body
design, for there will be greater need for adjusting the co-ordination of different pairs of legs in
movement. As a result, hexapodous nature in itself does not justify a reason for the diversity of
insects.
An important point to note is that the basic body plan of different groups only needs to
suit the number of pairs of legs and their positioning. For example, considering the Arachnids,
the four pairs placed in the cephalothorax region give much less leverage for increasing the size
of the abdomen. The length of the leg also needs to be relatively longer. On the other hand,
more numerous legs placed in the trunk region of Myriapods helps maintain a large trunk
attached to a small head. Insects also due to the placement of legs in the middle tagma need to
have a better balance between the anterior head and the posterior abdomen, thus to compensate
this positional disadvantage have to possess a relatively longer set of legs. Correspondingly, the
thorax has to be much stronger and more complex in structure to support the muscles required
for this balancing act. Caterpillars have solved this problem by having additional legs in the
abdominal region.
L. Specialized offence and defense mechanisms

Insects have an enormous diversity of defense mechanisms against their enemies. Some
mechanisms are quite offensive to the intending predator. Considering that each extant species
definitely has some mechanisms to survive in nature, that is basically red in tooth and claw, it is
not surprising that the great diversity of insects possesses a great diversity of defense
mechanisms. Thus, the explanation of relationship between the repertoire of defenses and the
speciose nature of the group is tautological. A situation similar to the ecological adaptation or
the diversity of food habits. However, a better dissection of the individual sets of defenses
similar to the food plants of insects discussed above might provide an insight whether the
diversity of defense mechanisms really contributed for the diversity of insects.
The summary :
In summary one can see that among the various attributed reasons for the enormous
diversity of insects, a few seem to stand the test and allow for meaningful comparison.
Alternatively, many others do not really provide a sound reason to be accepted as potential
causes for this diversity. Another feature that emanates from this analysis is that there is no one
cause that can be suggested as the outright reason for the observed diversity of insects.
Exoskeleton stands out with many associated features – trachea and the true wings of
insects would not have come about but for the versatile exoskeleton. Metamorphosis probably
would not have been required to grow but for the Exoskeleton that limits the growth.
Exoskeleton also limits the size. Exoskeleton also provides the additional; adaptive features of
providing versatile mouth appendages that provide for differential exploitation of food items.
More importantly provides protection against vagaries of weather.
Similarly, wings would not have been their but for the exoskeleton. Flight did probably
help. Wings also contributed for the evolution of metamorphosis. Metamorphosis is there
because of the exoskeleton and the wings that are found in the adult stage.
Essentially, one can see that every one of these features are not exclusive and are interlinked to
one another.
Exercise: link up the interlinking features of insects that made the it possible for the insects
to be so dominant. Now after having done that. Which is the most important reason that23
contributed for the great diversity of insects in your opinion?
Study questions :
a) Provide ten reasons why insects are dominant on earth. Give an idea of the diversity of insects
in relation to other organisms.
b) What are the most important reasons for the dominance of insects according to you and why?
Explain how important is small size for the observed diversity in insects.
c) Provide ten reasons why we should study insects.
d) Differentiate between insects and other arthropoda
e) Differentiate between Arthropoda and Annelida
f) Culturally how important are insects? Provide an evidence of reference to insects in Hindu
Mythology
g) What are the major reasons that have made insects the most dominant creatures on earth?
h) Give one physiological, one morphological and one behavioral adaptation each seen in insects
for conserving moisture.
i) Write a short note on : a) insect flight, b) food habits of insects c) ecological adaptability d)
tracheal system and its importance in promoting the diversity of insects. e) Metamorphosis and
its importance in promoting insect diversity, f) Insect reproductive abilities.
24

The next chapter : What is an Insect?
Classification of Phylum ARTHROPODA

taxa in FULL UPPER CASE indicate the four arthropod subphyla)
==A========= Arachnida
(spiders, mites, scorpions, etc.)
==C==|
========= CHELICERATA ===========| ==X========= Merostomata
(horseshoe crabs)
| |
| ===P============= Pycnogonida
(sea spiders)
|
<<=| ===== TRILOBITOMORPHA ======================== (extinct trilobites)
| |
| | ====== Symphyla (symphylans)
| | ==S==|
| | | ====== Chilopoda (centipedes)
| | ===Myriapoda======|
=T==| | =========== Diplopoda (millipedes)
| |
| == UNIRAMIA ==| ==Entognatha=== Collembola
| | | |=========== Protura
| | ==Hexapoda====|=========== Diplura
| | (Insects) ==============| Insecta
=M=| |
== =======================================to CRUSTACEA
===========================================================>
TRAITS SUPPORTING EACH CLADE
(** plesiomorphic- a primitive state, not unique to clade):
CHELICERATA:
a) two tagmata (cephalothorax=prosoma, without distinct head; abdomen=opisthosoma)**
b) mouth lies before segment 1 embryologically
c) segment #1 limbs= chelicerae (no antennae; may be pincer-like or fangs)
C:
a) cephalothorax has a carapace shield b) 1st or 2nd abdominal segment modified as
genital somite
P:
a) pre-oral proboscis b) ovigers (unusual 3rd pair limbs)
c) abdomen reduced or absent d) long, 9-segmented walking legs
e) multiple pairs of gonopores on some/all legs f) loss of compound eyes
A:
a) abdominal limbs reduced, lost, or modified as spinnerets b) loss of compound eyes
X:
a) enlarged cephalothorax w/ large carapace b) abdominal limbs modified as book gills
c) long, spiked telson
T:
25

a) mouth lies before segment 2 embryologically b) segment #1 limbs= antennae
c) 4 pair post-oral head segments with biramous limbs similar to thoracic limbs
TRILOBITOMORPHA:
a) 3 tagmata (cephalon, thorax, pygidium) b) body dorso-ventrally flattened
c) limbs of post-oral head segments similar to limbs of thorax (all with two branches,
biramous?)**
M (Mandibulata):
a) 2 tagmata (head, trunk of many similar segs.)**
b) 5 pair head appendages (2 pair antennae, 1 pair mandibles, 2 pair maxillae)
c) body segment cuticle of 4 sclerites (tergum, 2 lateral pleura, sternum)
d) mouth anterior to segment 3 in adult
UNIRAMIA (= Tracheata, = Atelocerata):

a) second antennae lost b) mandible is 'whole-limb'
c) unique tracheal system d) uniramous limbs (exopodite lost)
Myriapoda:
a) 2 tagmata (head, trunk of many similar segs.)** b) loss of compound eyes
c) loss of palps on first & second maxillae d) repugnatorial glands
S:
a) medial fusion of both pairs of maxillae b) maxillipeds modified as raptorial poison fangs
Hexapoda:
a) second maxillae fuse as labium b) 3-segment thorax with limbs
c) 11-segment abdomen, no limbs
What is an Insect ?
The Phylum Euarthropoda can be classified into four major taxa, that can in principle
be referred to as Subphyla. These are : Trilobitomorpha, Chelicerata, Myriapoda and
Pancrustacea. Trilobitomorpha is an extinct marine group. Other three taxa are
extant. The Subphylum Chelicerata includes four Classes Arachnida, Eurypterida,
Xiphosura and Pycnogonida. Among these Arachnida is a dominant group and is
commonly encountered. Arachnids (/əˈræknɪdz/) are a class (Arachnida) of joint-
legged invertebrate animals (arthropods), in the subphylum Chelicerata. Almost all adult
arachnids have eight legs, although the front pair of legs in some species has converted to a
sensory function, while in other species, different appendages can grow large enough to take on
the appearance of extra pairs of legs. The term is derived from
the Greek word ἀράχνη (aráchnē), from the myth of the hubristic human weaver Arachne who
was turned into a spider.[1] Spiders are the largest order in the class, which also
includes scorpions, ticks, mites, harvestmen, and solifuges.[2]
26

Almost all extant arachnids are terrestrial, living mainly on land. However, some inhabit
freshwater environments and, with the exception of the pelagic zone, marine environments as
well. They comprise over 100,000 named species.
The subphylum Myriapoda also includes four Classes. These are Pauropoda, Diplopoda,
Chilopoda and Symphyla. Pauropods are small, pale, millipede-like arthropods. The name is
derived from the Greek roots pauro "few" and podo "foot". Around 830 species in twelve families are
found worldwide that live in soil and leaf mould. They look rather like centipedes, but are probably
the sister group to millipedes.
Millipedes are a group of arthropods that are characterised by having two pairs of
jointed legs on most body segments; they are known scientifically as the class Diplopoda, the
name being derived from this feature. Each double-legged segment is a result of two single
segments fused together. Most millipedes have very elongated cylindrical or flattened bodies with
more than 20 segments, while pill millipedes are shorter and can roll into a ball. Although the
name "millipede" derives from the Latin for "thousand feet", no known species has 1,000; the
record of 750 legs belongs to Illacme plenipes. There are approximately 12,000
named species classified into 16 orders and around 140 families, making Diplopoda the largest
class of myriapods, an arthropod group which also includes centipedesand other multi-legged
creatures.
Most millipedes are slow-moving detritivores, eating decaying leaves and other dead plant
matter. Some eat fungi or suck plant fluids, and a small minority are predatory. Millipedes are
generally harmless to humans, although some can become household or garden pests,
especially in greenhouses where they can cause severe damage to emergent seedlings. Most
millipedes defend themselves with a variety of chemicals secreted from pores along the body,
although the tiny bristle millipedes are covered with tufts of detachable bristles. Reproduction in
most species is carried out by modified male legs called gonopods, which transfer packets of
sperm to females.
First appearing in the Silurian period, millipedes are some of the oldest known land animals.
Some members of prehistoric groups grew to over 2 m (6 ft 7 in); the largest modern species
reach maximum lengths of 27 to 38 cm (11 to 15 in). The longest extant species is the giant
African millipede (Archispirostreptus gigas).
Among myriapods, millipedes have traditionally been considered most closely related to the
tiny pauropods, although some molecular studies challenge this relationship. Millipedes can be
distinguishedfrom the somewhat similar but only distantly related centipedes (class Chilopoda),
which move rapidly, are carnivorous, and have only a single pair of legs on each body segment.
The scientific study of millipedes is known as diplopodology, and a scientist who studies them is
called a diplopodologist.
Centipedes (from Latin prefix centi-, "hundred", and pes, pedis, "foot") are arthropods belonging
to the class Chilopoda of the subphylum Myriapoda, an arthropod group which also
includes Millipedesand other multi-legged creatures. Centipedes are
elongated metameric creatures with one pair of legs per body segment. Centipedes are known to
be highly venomous, and often inject paralyzing venom. Despite the name, centipedes can have
a varying number of legs, ranging from 30 to 354. Centipedes always have an odd number of
pairs of legs.[1][2][3] Therefore, no centipede has exactly 100 legs. A key trait uniting this group is a
pair of venom claws or forcipules formed from a modified first appendage. Centipedes are
predominantly carnivorous.[4]:168
Their size can range from a few millimetres in the smaller lithobiomorphs and geophilomorphs to
about 30 cm (12 in) in the largest scolopendromorphs. Centipedes can be found in a wide variety
of environments. They normally have a drab coloration combining shades of brown and red.
27

Cavernicolous (cave-dwelling) and subterranean species may lack pigmentation, and many
tropical scolopendromorphs have bright aposematic colours.
Worldwide, an estimated 8,000 species of centipedes are thought to exist, [5] of which 3,000 have
been described. Centipedes have a wide geographical range, where they even reach beyond
the Arctic Circle.[4] They are found in an array of terrestrial habitats from tropical
rainforests to deserts. Within these habitats, centipedes require a moist microhabitat because
they lack the waxy cuticle of insectsand arachnids, therefore causing them to rapidly lose water.
[6]
Accordingly, they are found in soil and leaf litter, under stones and dead wood, and inside logs.
Centipedes are among the largest terrestrial invertebrate predators, and often contribute
significantly to the invertebrate predatory biomass in terrestrial ecosystems. Only one
species, Scolopendra cataracta, is known to be amphibious and is believed to hunt aquatic or
amphibious invertebrates.[7]
Some species of centipedes can be hazardous to humans because of their bite. Although a bite
to an adult human is usually very painful and may cause severe swelling, chills, fever, and
weakness, it is unlikely to be fatal. Bites can be dangerous to small children and those with
allergies to bee stings. The venomous bite of larger centipedes can induce anaphylactic shock in
such people. Smaller centipedes are generally incapable of piercing human skin. [22]
Even nonvenomous centipedes are considered frightening by humans due to their dozens of
legs moving at the same time and their tendency to dart swiftly out of the darkness towards one's
feet.[23] A 19th-century Tibetan poet warned his fellow Buddhists, "if you enjoy frightening others,
you will be reborn as a centipede."[24]
As a food item, certain large-sized centipedes are consumed in China, usually skewered and
grilled or deep fried. They are often seen in street vendor's stalls in large cities,
including Donghuamen and Wangfujing markets in Beijing.[18][19]
Also in China, as well as in Laos, Thailand, and Cambodia, large centipedes are kept in liquor for
a period of time. This custom is allegedly part of the traditional Chinese medicine. Said to have
medicinal properties and to be reinvigorating, the liquor with the centipede submerged in it is
consumed as a special drink.
Symphylans, also known as garden centipedes or pseudocentipedes, are soil-

dwelling arthropods of the class Symphyla in the subphylum Myriapoda. Symphylans
28

resemble centipedes, but are smaller and translucent, and only distantly related to true
centipedes. They can move rapidly through the pores between soil particles, and are typically
found from the surface down to a depth of about 50 cm. They consume decaying vegetation, but
can do considerable harm in an agricultural setting by consuming seeds, roots, and root hairs in
cultivated soil.
Juveniles have six pairs of legs, but over a lifetime of several years, they add an additional pair at
each moult so an adult instar has twelve pairs of legs.[1] Symphylans lack eyes. Their
long antennae serve as sense organs. They have several features linking them to early insects,
such as a labium (fused second maxillae), an identical number of head segments and certain
features of their legs.[2]
About 200 species are known worldwide.
The subphylum Pancrustacea is divided into two major Superclasses, viz., Crustacea and
Hexapoda. Superclass Crstacea has six classes in it, Malacostraca, Ostracoda, Maxillopoda,
Chephlocarida, Remipedia and Branchiopoda. Malacostraca includes crabs, lobsters, crayfish,
shrimp, krill, mantis shrimp, woodlice, hooded shrimp, scuds, sandhoppers etc.; Ostracoda
includes seed shrimp ; Maxillopoda includes barnacles and copepods; Cephalocarida
contains horseshoe shrimp ; Remipedia consist of members of the order Nectiopoda and
Branchiopoda include brine shrimp, fairy shrimp, water fleas, tadpole shrimp and clam shrimp.
Crustaceans (Crustacea /krʌˈsteɪʃə/) form a large, diverse arthropod taxon which includes such
familiar animals as crabs, lobsters, crayfish, shrimp, krill, woodlice, and barnacles.[1] The
crustacean group is usually treated as a subphylum, and thanks to recent molecular studies it is
now well accepted that the crustacean group is paraphyletic, and comprises all animals in
the Pancrustacea clade other than hexapods.[2]
The 67,000 described species range in size from Stygotantulus stocki at 0.1 mm (0.004 in), to
the Japanese spider crab with a leg span of up to 3.8 m (12.5 ft) and a mass of 20 kg (44 lb).
Like other arthropods, crustaceans have an exoskeleton, which they moult to grow. They are
distinguished from other groups of arthropods, such as insects, myriapods and chelicerates, by
the possession of biramous (two-parted) limbs, and by their larval forms, such as
the nauplius stage of branchiopods and copepods.
Most crustaceans are free-living aquatic animals, but some are terrestrial (e.g. woodlice), some
are parasitic (e.g. Rhizocephala, fish lice, tongue worms) and some are sessile (e.g. barnacles).
The group has an extensive fossil record, reaching back to the Cambrian, and includes living
fossils such as Triops cancriformis, which has existed apparently unchanged since
the Triassic period. More than 10 million tons of crustaceans are produced by fishery or farming
for human consumption, the majority of it being shrimp and prawns. Krill and copepods are not
as widely fished, but may be the animals with the greatest biomass on the planet, and form a
vital part of the food chain. The scientific study of crustaceans is known
as carcinology (alternatively, malacostracology, crustaceology or crustalogy), and a scientist who
works in carcinology is a carcinologist.
The subphylum Crustacea comprises almost 67,000 described species and many are believed
to be discovered yet. The group is considered a paraphyletic and the classification as such is
therefore imperfect. Studies have indicated Crustacea to be a sister group of Hexapoda. But
at the same time, it is believed that Hexapoda could be potentially nested within the larger
Pancrustacea. However, despite their great diversity of form, the group is mostly united by
29

the special larval form known as the nauplius. Some crustaceans are more closely related
to insects and other hexapods than they are to certain other crustaceans.
The majority of crustaceans are aquatic, living in either marine or freshwater environments, but a
few groups have adapted to life on land, such as terrestrial crabs, terrestrial hermit crabs,
and woodlice. Marine crustaceans are as ubiquitous in the oceans as insects are on land.
Many crustaceans are consumed by humans, and nearly 10,700,000 tons were produced in
2007; the vast majority of this output is of decapod crustaceans: crabs, lobsters, shrimp,
crawfish, and prawns. Over 60% by weight of all crustaceans caught for consumption are shrimp
and prawns, and nearly 80% is produced in Asia, with China alone producing nearly half the
world's total. Non-decapod crustaceans are not widely consumed, with only 118,000 tons of krill
being caught, despite krill having one of the greatest biomasses on the planet.
Malacostraca includes crabs, lobsters, crayfish, shrimp, krill, mantis shrimp, woodlice, hooded
shrimp, scuds, sandhoppers etc.; Ostracoda includes seed shrimp ; Maxillopoda includes
barnacles and copepods; Cephalocarida contains horseshoe shrimp ; Remipedia consist of
members of the order Nectiopoda and Branchiopoda include brine shrimp, fairy shrimp, water
fleas, tadpole shrimp and clam shrimp
30

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Chapter 2: Dominance of insects:

In an everlasting inquisitiveness, pioneered probably by the earliest human beings, man

Erwin’s estimate of 32 million species of insects might simply mean an impractical

A.R.V. Kumar : Introduction to Entomology

Another alternative measure of abundance is the colony sizes of social insects. A

A.R.V. Kumar : Introduction to Entomology

Table 1: Worldwide number of species of organisms known to science in different taxa.

Taxa No. of species in thousands

4. Kingdom - Animalia 1387

Above figures indicate the abundance of organisms on the basis of estimates of

Why are insects dominant? : The reasons for enormous diversity 3

A.R.V. Kumar : Introduction to Entomology

A.R.V. Kumar : Introduction to Entomology

B. The small size:

This suggests an enormous range of over 3000 folds. Perhaps no other

A.R.V. Kumar : Introduction to Entomology

Distribution of Terre stria l organisms by size

Size of Animals (log mm)

A.R.V. Kumar : Introduction to Entomology

Why are insects small?

A.R.V. Kumar : Introduction to Entomology

A second and equally important reason is metamorphosis. Insects pass through a

A.R.V. Kumar : Introduction to Entomology

What advantages do insects get by possessing flight?

A.R.V. Kumar : Introduction to Entomology

A.R.V. Kumar : Introduction to Entomology

A.R.V. Kumar : Introduction to Entomology

What then is the advantage of metamorphosis to facilitate the diversity of insects? In

One other feature facilitated by the metamorphosis is the phenological adjustments. We

A.R.V. Kumar : Introduction to Entomology

Therefore, the metamorphosis is considered an important factor for the diversification

Additional causes attributed for the diversity of insects:

A.R.V. Kumar : Introduction to Entomology

Temperatures is one important factor expected to influence the distribution of

A.R.V. Kumar : Introduction to Entomology

Table 2. Relative distribution of different animal phyla in major ecological habitats.

Sl. No. Phylum Marine Freshwater Terrestrial

The danger of desiccation is particularly more serious for insects compared to

A.R.V. Kumar : Introduction to Entomology

A.R.V. Kumar : Introduction to Entomology

One other technique of overcoming the danger of desiccation is to resort to behavioural

A.R.V. Kumar : Introduction to Entomology

Living situations is another mechanism of overcoming the danger of desiccation. Soil

A.R.V. Kumar : Introduction to Entomology

In essence, various mechanisms have been developed by insects to overcome the

Nevertheless, colonisation of terrestrial habitat definitely provided an opportunity for

G. Diversity of food habits:

Detritivory - feeding on decaying matter - ex. Collembolans, Thysanurans, cockroaches, Flies,

Coprophagy – feeding on animals defecation - dung – Beetles, flies etc.

A.R.V. Kumar : Introduction to Entomology

Nectarivores and Pollen feeders – Beetles, flies, bees, wasps, etc.

Carnivory - Predatory – Bugs, beetles, wasps, praying matids, etc.

A.R.V. Kumar : Introduction to Entomology

This is a stuation is similar to the colonisation of terrestrial habitats by the insects. A

H. Enormous reproductive ability

Higher reproduction rates lead to higher possibilities for mutations to accumulate. As a

I. Decentralised nervous system

A.R.V. Kumar : Introduction to Entomology

A.R.V. Kumar : Introduction to Entomology