DOI: 10.5772/intechopen.

70452

Provisional chapter

Chapter 1

Introductory Chapter: Lepidoptera

Introductory Chapter: Lepidoptera
Farzana Khan Perveen and Anzela Khan
Farzana Khan Perveen and Anzela Khan
Additional information is available at the end of the chapter

Additional information is available at the end of the chapter

http://dx.doi.org/10.5772/intechopen.70452

1. Lepidoptera

The word Lepidoptera comes from the Latin word, equivalent to lepido- and from the ancient
Greek words lepis and pteron mean scales and wings, respectively. Therefore, it stands for
insects with scaly wings. It is the second largest, diverse, widespread, and widely recognized
insect’s order in the class Insecta of phylum Arthropoda. Linnaeus (1707–1778) divides it into
three groups: (1) butterflies, (2) skippers, and (3) micro- and macro-moths. It consists of 126
families and 46 superfamilies (Table 1). They can be differentiated on morphological, ana-
tomical, behavioral, and ecological characteristics [1]. Further, 500,250 species of Lepidoptera
are described, with 70,820 species of butterflies [2, 3] and 3700 species of skippers globally
[4–6]. Furthermore, about 165,000 species of moths, including micro- and macro-moths, are
found up to now [6–9]. In nature, Lepidoptera regard as the symbol of beauty and grace. They
are very beautiful creatures of nature (Figure 1A) [10–12].

1.1. Morphology

The group Rhopalocera is related to butterflies and skippers; however, Heterocera is to micro-
and macro-moths. The Lepidoptera show a great diversity in forms, size, structure, and other
distinctiveness (Figure 1A and B) [13, 14].

1.1.1. Head segment

Lepidoptera’s head capsule is the feeding and sensory center. It is small, round, or elliptical
and sclerotizes organization. The upper-middle portion of the head is called the frons; below
is the clypeus, and below it is the labrum, to both sides of which the edges of the mandibles
with different aspects of the maxillary palps may expand beyond and/or underneath, even
when view them from front [15]. As a whole, the shape and size of the head capsule, color
patterns, and location of hairs on the head are supportive in identifying species of caterpillars
with aid of a microscope (Figure 1C).

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4 Lepidoptera

Kingdom: Animalia

Subkingdom: Invertebrata

Super-division: Eumetazoa

Division: Bilateria

Subdivision: Ecdysozoa

Superphylum: Tactopoda

Phylum: Arthropoda von Siebold, 1848

Subphylum: Atelocerata

Superclass: Hexapoda

Class: Insecta

Infraclass: Neoptera

Subclass: Pterygota

Superorder: Endopterygota

Unranked: Amphiesmenoptera

Unranked: Holometabola

Order: Lepidoptera Linnaeus, 1758

Examples:

• Butterflies
• Skippers
• Micro-moths
• Macro-moths

Table 1. The taxonomic position of the order Lepidoptera [13].

1.1.1.1. Mouthparts

The siphoning-type mouthparts are found in the imago. They are transformed into a long
flexible hollow structure, in which the formation of the suctorial proboscis encompasses a
fluid-tight food tube. Lepidoptera feed on nectar, and their proboscis length may increase
almost 100-folds. Usually, when they do not use them, they keep coil under the head with
the help of small muscles present there. In all Lepidoptera, the basic structure of mouthparts
is the same, which include each one labium, labrum, hypopharynx, or tongue with pairs of
mandibles and maxillae (Figure 1C) [16, 17].

1.1.1.2. Antennae

The antennae show a wide variation in forms, size, structure, and other characteristics among
species and even between different sexes. The basic structure of antennae of Lepidoptera is
usually filiform, which is altered into the capitates of antenna, which are club shaped with
a long shaft and a bulb at the end. In the skippers, most of the antennae’s tips are changed
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Figure 1. Parts of Lepidoptera body: (A and B) butterfly and moth; (C) generalize mouth parts; (D) generalize antennae of
order Lepidoptera: butterflies; (a) skippers; (b) micro- and macro- moths (c, d, e) [18, 19]; (E) compound eye of Lepidoptera;
(F) unit and basic structure, ommatidium of a compound eye, showing its construction; (G) simple eye or ocellus [22].

into a narrow hook-like projection. A moth’s antennae are feathery or saw edged. They also
act as the balancing organ. The shapes of antennae are assisting in identifying species of
Lepidoptera (Figure 1D) [20].

1.1.1.3. Compound eyes

The eyes are usually paired, golden brown, or even red as in some skipper species. They built
quite differently from the vertebrate eye, but like Arthropod, it is made up of repeating units
up to 17,000, the ommatidia, each function as a separate visual receptor, which in combina-
tion provide a broad mosaic view of the image, such type of the eye is a compound eye. Each
is connected to a lens, which is attached to a nerve leading to the brain (Figure 1E and F) [21].

1.1.1.4. Simple eyes or ocelli

In all Lepidoptera, in addition compound eyes, simple eyes, or ocelli (singular: ocellus; simple
photoreceptors) are also present. They made with a single lens and several sensory cells. Only
two ocelli are present in imago, excluding a few moths, one on each side of compound eyes.
In some species, a type of sense organs, which called chaetosemata, is found near the ocelli. In
caterpillars, three pairs of simple eyes are found, which are not homologous to ocelli of imago.
Simple eyes of caterpillars are differently named as stemmata. Lepidoptera are able to per-
ceive ultraviolet (UV) light and observe wing colors and patterns by ocelli (Figure 1G) [22].

1.1.2. Thorax

The thorax is the second part of the body, which composed of three jointed segments, the pro-
thorax, mesothorax, and metathorax, each derives from a primitive segment. They are covered
6 Lepidoptera

dorsally with tergites, ventrally with sternites, and laterally with pleurites (chitinous plates)
[23]. The characteristic like the presence or absence and shape of sclerotized plates; location of
primary setae; and location, color, and shape of the prothoracic spiracle assists in identification
of caterpillar and imago species [24].

1.1.2.1. Jointed legs

Lepidoptera have three pairs of well-developed jointed legs. They are located in each seg-
ment of the thorax and covered with scales. Each leg consists of nine segments, that is, coxa,
trochanter, femur, tibia; five tarsal segments with a pretarsus; and a pair of articulated curved
claws on the fifth segment. Morphology of the legs also aids in identifying caterpillar and
imago species (Figure 2a and b) [25]. The aroliar pad (a pad extending between the tarsal
claws) and pulvillus (plural: pulvilli, pads beneath each tarsal claw) are short or absent in
some families. The tibia of each leg contains a subgenual organ, which detects and amplifies
small vibrations (Figure 2a and b) [26].

1.1.2.2. Wings

The mysterious flight of Lepidoptera depends on their wings, which accomplished several kinds
of difficult tasks like diving, circling, parachuting, equilibrium, etc. all because of their lightness
in nature. Their wings are subjected to considerable variations in shape, size, markings, spots,
and vein patterns, thus reflecting their specific functional differences. Strong muscles in the
thorax move the wings up and down in a digit 8 pattern during the flight. Both pairs of wings
are covered with thousands of bright, colorful, and dull scales. Due to the presence of scales on
the wings, the term for order Lepidoptera has been coined (Figure 2f and g). Wing scales adapt

Figure 2. (a) Generalize structure of Lepidoptera Leg; (b) generalize structure of tarsus; (c) TS of vein from wing
generalized wings venation: (d) forewing; (e) hindwing; with names of different veins with their abbreviations are: C:
Costal; Sc: subcostal vein; R1: 1st radius vein; R2: 2nd radius vein; R3: 3rd radius vein; R4: 4th radius vein; R5: 5th radius
vein; M1: 1st median vein; M2: 2nd median vein; M3: 3rd median vein; Cu1: 1st cubitus vein; CuA1: 1st cubitus anterior;
CuP2: 2nd cubitus posterior; 1A: 1st anal vein; 2A: 2nd anal vein; 3A: 3rd anal vein; H: humeral 2012; bars in photographs
indicate 30 mm; (f) wing color of black and white; (g) wing of a male the monarch butterfly, Danaus plexippus (Linnaeus,
1758) (Nymphalidae: Nymphalinae) [23–26].
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to different environmental conditions affecting flights. They are also found to influence by the
time factor, which affects speed, foraging, calling, finding places for spawning, avoiding preda-
tors, etc. Wing pattern is successful in establishing the correlation in adaptation and adaptive
change. Both changes in morphology of wings and the pattern of flight are closely associated
with the genetic material. Similarly, the relationship has been established between wing vena-
tion and wing pattern in the genus Micropterix Hubner, 1825 (Family: Micropterigidae). Such
type of studies plays a major role in understanding its evolutionary status and highlights its
significance in taxonomical analysis. They are chitinous membranes, nourished and supported
by tubular veins (Figure 2c–e). These veins also function in exchange of oxygen. Further, vena-
tion has aerodynamic importance, plays specific role in flight system, and adapts to differ-
ent surroundings [26]. Bashar et al. [27] prepared pictorial key for identification of the local
nymphalid butterflies of Bangladesh, based on the wing venation of the butterflies. It has also
observed that venation is an important trait in Lepidoptera phylogenetic development [28].

1.1.3. Abdomen

The abdomen is third part of the body. It consists of 10–11 segments tapering to the end.
Each segment provides membranes in between allowing for articulation and movement. In
Lepidoptera systematic, they have been one of the most important sources of character infor-
mation [20]. In some caterpillar, four pairs of prolegs normally located on the third to sixth
segments, and a separate pair of prolegs by the anus, which has a pair of tiny hooks called
crotchets, helps in gripping and walking [29].

1.1.4. Scales

The name of this order Lepidoptera is due to the presence of the scales. The head, thorax,
abdomen, wings, and legs are covered with minute scales, are lamellar or blade-like, and
are attached with a pedicel, while other forms may be hair-like or specialized as the second-
ary sexual characteristics. Either, they give color, by color pigments they contain or through
structural coloration with mechanism that include photonic crystal and diffraction grating.
They are functioned as aiding gliding flight, insulation, producing pheromone, thermoregu-
lation, etc. The most important is the large diversity of their bright or indistinguishable pat-
tern, which aids the organisms to protect itself by camouflage or mimicry including rival and
potential mate (Figure 2f and g) [4].

1.1.5. Sound-producing organs

The sounds of some Lepidoptera are clearly audible, for example, members of Sphingidae
and Pyralidae. Many moths have developed ears on their wings or thorax, which are called
as tympanal organ, which can make them aware of thread, predator, etc. The Neotropical
tiger moth, Bertholdia trigona (Grote, 1879) (Noctuoidea: Erebidae), actively makes out of
function of the bat radar by creating its own ultrasound and by vibrating its tympanum
present on its metathorax. Males of the moth, Symmoracma minoralis (Snellen) (Pyralidae:
Nymphulinae), produce a high-intensity calling song from tymbals like structure found in
the genital segment [30–32].
8 Lepidoptera

1.2. Endocrine control

In Lepidoptera, the growth and metamorphosis are under control by interacting sets of hor-
mones. Ecdysis is initiated by ecdysiotropin, or prothoracicotropic hormone (PTTH) or brain
hormone (BH) is secreted by protocerebrum which acts on ecdysial glands. Eclosion hormone is
secreted by brain median neurosecretory cells; it is stored in the corpora cardiac and is released
into the hemolymph during switchover from pupa to imago. Within the abdominal ganglia, it
acts on neurons to begin the pre-eclosion behavior. The corpora allata secret juvenile hormone
(JH). It is liable to bring juvenile progress and variable species to species. The molting hormone
(MT) or ecdysone hormone (EH) is secreted by the ecdysial gland. Distinctively, all hormones
and neurohormones are involved in management of circadian rhythms, growth, development
of the nervous system, diapause, mating, metabolism, oviposition, pheromone biosynthesis,
regulation of dormancy, regulation of migratory behavior, and other physiological functions in
the body. The EH is responsible for several activities of pupal-imago conversion, with respect
to the behavior associated with ecdysis, following deterioration of abdominal intersegmental
muscles. Ecdysis-triggering hormone which is the most newly discovered hormone shows a
significant role in ecdysis. Bursicon (tanning hormone) is usually synthesized in neurohemal
organs related with the ventral chain ganglia. Its functions are to stimulate sclerotization and
tanning of the cuticle during the course of ecdysis (Figure 3) [33–36].

1.3. Polymorphism

Existence of the morphologically different individuals in the life cycle of the same species is
termed as polymorphism. The sexual dimorphism is very common, in which male and female
are structurally different and are found in families Pieridae, Nymphalidae, Papilionidae, and
Psychidae. The other types are geographical, seasonal, genetic, and environmental polymor-
phism. In some species, the polymorphism is limited to one sex, typically the female [33–36].

1.4. Pheromones

They are biochemicals, meant of communication, secreted into the surroundings, and affect
the activities or functioning in others species. Such communication systems have provided
challenges to scientists in different disciplines including behavior, biochemistry, chemistry,
ecology, genetics, and physiology. They produce in greater varieties. As a result, the great
numbers of researches are conducted focusing on butterflies and moths. Pheromone systems
are species specific for attracting mates. Sex pheromones are used for long-distance biochemi-
cal communication [37, 38].

1.5. Migration

Lepidoptera contribute an essential part as the environmental indicators. If any minute dan-
gerous variations occur in the surroundings, they may affect acutely. Due to unfavorable
environmental affects, they migrate rapidly in long distances, from locations to the area,
which are more suitable for any part of the seasons. Their destination may be tropical and
subtropical areas and all continents, excluding Antarctica. They stay away from undesirable
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Figure 3. Life-cycle of Lepidoptera, the common cutworm, Spodoptera litura (Fabricius, 1775): (a) imago male; (b) imago
female; (c) bunch of egg; (d) newly hatched 1st instar caterpillar; (e) 2nd instar caterpillar; (f) 3rd instar caterpillar; (g)
4th instar caterpillar; (h) 5th instar caterpillar; (i) 6th instar caterpillar; (j) defensive posture of 6th instar caterpillar; (k)
female cocoon; (l) male cocoon [19].

circumstances, including adverse climate, food shortage, overpopulation, weather, etc. In

some conditions, few members migrate, and in other conditions, all migrate [39–41].

1.6. Internal anatomy

1.6.1. Digestive system

In Lepidoptera, the foregut is started from the mouth; the pharynx may be highly modi-
fied into a pump. Posterior to it is the esophagus which opens into a crop or storage organ.
Immediately, posterior to it is the proventriculus, a structure that contains sclerotized toothlike
denticles, aiding in grinding the food. Some fluid-feeding Lepidoptera lack a proventriculus.
The stomodael valve (foregut valve) regulates the flow of materials from the foregut-midgut.
The former is lined with chitinous protective layer called the intima. It prohibits absorption
of nutrients. In many Lepidoptera, the foregut valve has associated with gastric caeca that
produce digestive enzymes and increase surface area. The intima is absent in the midgut, and
most of the absorption of nutrients occurs here. The Malpighian tubules attach to the pylorus
region of the hindgut. Posterior to them is the anterior intestine and the highly muscularized
rectum that terminates in the anus. It functions in removing water from the fecal materials;
therefore, Lepidoptera produce very dry excrements (Figure 4) [42–45].

1.6.2. Circulatory system

Lepidoptera have open circulatory system. The major portion of the hemolymph is found in
open cavities. It bathes the organs within the body cavity, the hemocoel. Hemolymph enters
10 Lepidoptera

Figure 4. Internal anatomy of Lepidoptera viewing imago male (family: Nymphalidae), showing the most of the major
organ systems, with characteristic reduced forelegs of that family and the corpora include the corpus allatum and the
corpus cardiac [41].

the dorsal vessel or heart via small openings called the ostia. It is then pumped toward the
head, where it then returns to the hemocoel. It serves as a lubricant for the movement of inter-
nal structures. It is a hydraulic medium for applying pressure for molting, eversible glands
are extruded via pressure changes, and some muscular contraction is opposed by hydrostatic
pressure within the hemocoel. Hemolymph transports various substances from one tissue to
another (Figure 4) [42–45].

1.6.3. Respiratory or tracheal system

Tracheal system consists of a system of branching tubes (tracheae) and openings to the out-
side called spiracles. Atria spiracles have mechanisms that allow Lepidoptera to close the
opening. It prevents water loss and prevents the entry of pathogens and parasites. Tubes
begin rather large and branch to become successively smaller and smaller as they penetrate
deep within the tissues of the insect. The smallest branches of the tracheae are the tracheoles,
and gaseous exchange occurs here. Air sacs have many potential functions, all rather specula-
tive, including increasing the volume of air in the body for exchange, lowering the specific
gravity for flight, and providing room for the growth of internal organs. Usually, the first pair
of spiracles is found on the mesothorax. Lepidoptera may control the flow by opening and
closing the spiracles (Figure 4) [42–45].

1.6.4. Excretory system

The function of the excretory system is to maintain chemical homeostasis. By this system,
hemolymph is cleaned with metabolic wastes including nitrogenous waste products created
during digestion of food. As well as toxins, concentration of salts, and water are also regulated
in hemolymph by the same. Malpighian tubules and the hindgut comprise the excretory sys-
tem. Malpighian tubules, attached to the gut, float freely within the hemocoel and are bathed
in hemolymph. They vary in number from 2 to 250 or more. Their functions are removing
toxins, nitrogenous wastes, and ions to maintain ionic concentrations within the hemolymph.
Water and other small ions are removed from the gut by the rectum (Figure 4) [42–50].
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1.6.5. Nervous system

In Lepidoptera, the central nervous system is composed of a double chain of ganglia joined by
longitudinal connectives. The anterior ganglion is the brain. The brain connects to the ventral
chain of ganglia via two connectives that travel around the pharynx. The brain connects to the
eyes, ocelli, and antennae. The subesophageal ganglion is highly complex and innervates the
sense organs and muscles associated with the mouthparts, salivary glands, and neck region.
The subesophageal ganglion is the primary excitatory or inhibitory influence on motor activ-
ity of the whole Lepidoptera. The frontal ganglion connects the brain to the stomatogastric
subsystem. The hypocerebral ganglion is associated with two endocrine glands one of which
is the corpus allatum that produces JH. The thoracic ganglia contain the sensory and motor
centers for their respective segments. More derived taxa show a reduction in the number
of abdominal ganglia. In visceral nervous system, nerves associated with the brain, salivary
glands, and foregut are the stomatogastric subsystem. The caudal visceral subsystem is asso-
ciated with the posterior segments of the abdomen including the reproductive system. In
peripheral nervous system, all of the nerves are with synapses to the central and the visceral
nervous systems. These nerves are associated with sensory structures (Figure 4) [51–59].

1.6.6. Reproduction

The adult male Lepidoptera reproductive tract is composed of a pair of testes, vas deferens, acces-
sory glands, ejaculatory duct, and aedeagus. In testes, sperm begin to mature during the third
and fourth larval instars. These divisions take place during the larval and pupal stages. Sperm are
matured through spermiogenesis. All butterflies and moths produce two kinds of sperm: eupy-
rene sperm have a nucleus and can fertilize eggs, while apyrene sperm do not have a nucleus, and
they facilitate the eupyrene sperm. Matured sperm are transferred within a protein-rich ejaculate
called a spermatophore. It forms within the male’s aedeagus and is transferred with sperm at
the very end of copulation into the bursa copulatrix of female which can take up to 16 hours.
The adult female Lepidoptera reproductive tract is composed of the bursa copulatrix, sperm duct,
spermatheca, ovaries with ovarioles, and common oviduct. The end of the ovarioles is called the
germarium, where oocytes are produced from the original germ cells. This process begins during
the larval stage and continues in imago. Oocytes are covered with chorion, which forms in the last
stage of oogenesis. However, male genitals include a valva, which is usually large, as it is used to
grasp the female during mating. In female genitalia, there are three basic arrangements of open-
ings for copulation, fertilization, and egg-laying. Firstly, in exoporian, an external opening that
carries sperm from the copulatory opening of gonopore to the ovipore is found in Hepialidae and
its related families. Secondly, in monotrysian, a single genital aperture near the end of the abdo-
men through which both copulation and egg-laying occur is found in primitive groups. Thirdly,
in ditrysian, an internal duct that carries sperm with two distinct openings each for copulation
and egg-laying is found in all the remaining groups (98%). As the egg passes down the common
oviduct, few sperm are released from the spermatheca [33, 46]. Fertilization occurs just before an
egg is about to be laid. High levels of JH circulating in adult butterflies cause eggs to mature in
females and cause the male reproductive tract to develop. Diapause Lepidoptera, which reach
sexual maturity after the overwintering period, have low levels of JH in their hemolymph (Figure
5a–c) [60–67].
12 Lepidoptera

Figure 5. Anatomy of reproductive system of Lepidotera: (a) the female imago reproductive system: A: corpus
bursae; B: signum, C: ductus bursae; D: ostium bursae; and E: diverticulum of bursa copulatrix; F: ductis seminalis; G:
spermathecal gland; H: utriculus I: lagena of spermatheca; J: ductus receptaculi; K: accessory gland (paired); L: accessory
gland reservoir (paired); M: vestibulum; N: calyx of the unpaired oviductus communis; O: one of four ovarioles of
ovary (paired); P: papillae anales; Q: rectum; X: corpus, Y: collum, and Z: frenum of spermatophore; (b) the male imago
reproductive system: A: testis; B: seminal vesicle (paired); C: vas deferens (paired); D: accessory glands (paired); E:
ductus ejaculatorious duplex; F: primary segment of ductus ejaculatorious simplex; G: muascular area; H: area of frenum
formation; and I: area of collum formation of the cuticular secondary segment of the ductus ejaculatorious simplex; J:
caecum of aedeagus; K: aedeagus (After Etman and Hooper, 1979); (c) the mating pair of the the monarch butterfly,
Danaus plexippus (Linnaeus, 1758) (Nymphalidae: Nymphalinae) [48, 49].

2. Lepidoptera as model taxon

Lepidoptera are ideal for to increase awareness toward environmental issues and educa-
tional purposes. They produce a more positive perspective of the invertebrates to the pub-
lic, mostly due to their esthetic value. As Lepidoptera are the most well-known insects, they
have become flagship organisms for the divulgation of invertebrate conservation plans. Their
ecological significance is massive, not only because of the greatest percentage of species and
biomass they account for in ecosystems, although they act as herbivores, pollinators, and food
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for insectivores. For researchers, scientists, and students, they offer a model taxon for precious
to cram of biodiversity, conservation studies, environmental impact estimates, monitoring
of animal populations, ecology, ethnology, evolution, genetics, systematic, and many other
ecological and genetic studies. They open doors to establishment of chemical ecology as a
scientific discipline for study and research.

Author details

Farzana Khan Perveen1* and Anzela Khan2

*Address all correspondence to: [email protected]

1 Department of Zoology, Shaheed Benazir Bhutto University (SBBU), Main Campus,

Sheringal, Khyber Pakhtunkhwa (KP), Pakistan

2 Roots Millennium College (RMC), Sector I-9, Islamabad, Pakistan

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