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5-21-1997 The Life cycle of moths and Mary Walter

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Recommended Citation Walter, Mary, "The Life cycle of moths and butterflies" (1997). Thesis. Rochester Institute of Technology. Accessed from

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A Thesis submitted to the Faculty of the College of Imaging Aits and Sciences in candidacy for the degree of Master of Fine Arts.

The Life Cycle of Moths and Butterflies

by Mary E. Walter

May 21, 1997 Approvals

Chief Advisor: Robert Wabnitz Date Y"- 30 - q 7 /

Chief Advisor: Dr. N. Wanek

Chief Advisor: Dr. Peter Walter Date. :=...J+~Z_'_.L'__,9~/-"-?.L7---- 7 7

I Mary Walter, hereby grant permission to the Wallace Memorial Library of RIT to reproduce my thesis in whole or in pat1. Any reproduction will not be for commercial use or profit.

Signature r-: _ Date :'::>-f/~\"'_"Jf-+-/-+~/_7.l--.- _ 7 17 . 1

Introduction developed as flat, widely attached flaps that It is estimated that eighty percent of all protruded out from the thorax. It is believed are . There are approxi that wings may have developed to aid in "aeri dispersal" mately 800,000 species of insects, 180,000 of al of minute insects, for sexual dis which are (Arnett & Jacques, play to lure mates, or as moveable winglets to 198 1). In this paper I hope to describe the protect openings to air tubules (Douglas,

attributes of butterflies and moths which make 1989). them unique from other insects. In doing so I The thermoregulatory hypothesis will explain their evolutionary history, physi shows that an increase in the size of winglets cal attributes, stages of development and aids in body temperature elevation by absorb metamorphosis, unique survival mechanisms, ing solar radiation to increase body tempera and colonization and migration habits. ture more quickly. Warming occurs in pockets of air beneath winglets, increasing tempera Evolutionary History tures of leg muscles, enabling winged insects Butterflies and moths, like all insects, to forage longer and further, and increasing are members of the phylum Arthropoda. response rates to aid in escape from predators. , which are believed to have It is likely that these advantages also led to evolved from an annelid (earthworm) type increased dispersal of the species. Basking in ancestor are divided into the three major the sun is a common means of thermoregula

branches; the chelicerates (spiders, ticks, scor tion for winged insects. Butterflies and pions, etc.), the crustaceans (crabs, lobster, moths, which are broad winged insects, are crayfish) and the mandibulates (the believed to have arisen during the Triassic Onychophora, centipedes, millipedes, and period from the Stonefly (Douglas, 1989). insects). All members of the phylum arthro poda are similar in that they exhibit the body Life Cycle segmentation of arthropods; head, thorax and The stages of development abdomen. Butterflies and moths, like other include egg, larva, pupa, and adult. members of the class Insecta possess three Metamorphosis is the process by which the pairs of legs (one on each thoracic segment), immature butterfly is transformed from one one pair of antennae, a pair of maxillae with a stage to the next. Metamorphosis is controlled pair of sensory palpi, a pair of mandibles, a by gene regulation. The changes that occur hypopharynx (a tongue like structure), a labi within each of the immature stages: egg, larva,

pah" um (a specialized of head appendages and chrysalis / pupa, are controlled by various behind the maxillae) and two pair of wings groups of genes, which are turned on during (some insects have one or no pair of wings) the different stages of development.

(Arnett & Jacques, 1981). Therefore, the genes of the organism remain Of the twenty-nine orders of insects in the same throughout the life cycle, but differ the world, Lepidoptera, which includes moths, ent groups of genes are turned on and off butterflies and skippers, is one of the largest. throughout development (Douglas, 1989). There are approximately 13,700 species of Monarch Butterfly: Lepidoptera in North America, and 1 80,000 in the world (Arnett & Jacques, 1981). Lepidoptera, like other orders possess wings. There are several theories to explain the evolution of winged insects. The most accepted view is the paranotal theoiy, fig.l Larva hatching fig. 2 Full grown larva which suggests that the first wings of insects from egg or close to the larval butterfly's food supply Douglas, 1989). For example, the Monarch butterfly lays her eggs on the young, tender leaves of the milkweed plant, while the Black Swallowtail commonly deposits its eggs on garden parsley plants, which the larvae quick ly devour. fig. 3 Pupa fig.4 Adult Butterfly eggs are often white or cream (photos of the Buffalo Museum of courtesy Science) colored, with variations in color, size and shape depending on the species and develop Reproduction mental stage of the egg. The surface structure or chorion of eggs is a common means of Butterflies reproduce sexually, with identification. In the center of each the male contributing the sperm, while the specie or through female contributes the egg. Both sperm and egg is a small hole micropyle, which sperm permeates the chorion to fertil egg are haploid in their chromosome num ize the egg. While the chorion facilitates the ber. That is, each sex cell contains half the exchange of oxygen and carbon dioxide, it is number of chromosomes of the body cells resistant injury. Gas exchange (Douglas, 1989). strong and to takes place via ducts or air pockets called

plastrons, which act as gills to diffuse gases, even when the egg is immersed in water. The micropyle may also participate in respiration (Douglas, 1989).

Fig.5 Pair of mating Cecropia moths. The female moth is on the upper left, identifiable because females have a larger abdomen.

Morphology The Egg Eggs are produced within the tubular

ovaries of the female and travel to the

oviducts which join to form one duct

through which the eggs are deposited to the

outside (oviposition). Just before the is egg Fig. 6 The eggs of the Cecropia moth are it is fertilized a sperm cell that released, by generally laid in small clutches on the was stored in the corpus bursae, which is the leaves of lilac, apple, cherry, boxelder, female butterfly's internal sperm receptacle maple, birch, willow, plum and elderberry 1989). (Douglas, trees (Mitchell & Zim, 1964). The female usually lays her eggs on Fig. 7 Poster of the life cycle of the Cecropia moth.

-third larval instar -a pair of mating moths (actual size 1 1/4 to 1 1/2")

-fourth larval -a female laying a clutch of eggs on a lilac instar (actual size 2 to 2 1/2") leaf (a female lays from 200 to 300 eggs) -fifth larval instar (actual size 4 to 5")

-Pupa within -first larval instar (actual size 1/4 to 5/16 ") its silk cocoon (cut-away view 3/4" -second larval instar (actual size 1/2 to ) is shown) 4

Below the chorion is the vitelline food during consumption, is found behind the membrane which envelopes and protects the mandibles. Lastly, the larval spinneret, from developing larva. Several other layers which silk is released, is found between the develop between the chorion and vitelline maxillae and the labium (Douglas, 1989). membrane as the larva develops. The trans The thirteen segments of the larval formation from single celled egg to the tiny body are divided into three thoracic segments, lava usually takes between three to ten days, each possessing a pair of jointed legs with a depending on the species and weather con terminal claw (for food manipulation and ditions. Some species deposit their eggs in locomotion), and ten abdominal segments, the fall and the eggs over-winter in a state five of which possess non-jointed larval pro- of arrest called diapause, and do not hatch legs with ventral crochets I hooks (for loco

until spring, when conditions are optimal motion and grasping) (Douglas, 1989). (Douglas, 1989). The exoskeleton of the caterpillar, with The larva within the egg increases in the exception of the head, is soft and "worm size as the cells multiply geometrically, uti like". A large puncture to the exoskeleton lizing the egg yolk for nutrition. When could cause collapse of the exoskeleton, ready to emerge from the egg the larva resulting in fluid loss and death. Gas chews through the egg membranes and exchange does not occur through the chorion using its mandibles. The larva fre exoskeleton, but via abdominal and thoracic quently eat the rest of the egg chorion, and spiracles located along the sides of the larva. even that of neighboring unhatched eggs The spiracles are valve regulated air openings (Douglas, 1989). that connect to the tracheae which extend throughout the body to supply all body organs The Larva and muscles with oxygen (Douglas, 1989). Young larva feed on their host plants until they reach full size. Some caterpillars are carnivorous, eating aphids and related insects (ie. the larvae of the North American Harvester butterfly) or ant larvae and pupae (ie. the larvae of the

hairstreak Harkenclenus thus and the

blues Glaucopsyche lygdamus and

Celastrina argiolus pseudargiolus ) (Douglas, 1989). Larvae are composed of a head and thirteen body segments. The head of the larva contains antennae, mandibles for crushing food, and six pairs of simple eyes called ocelli. The ocelli are separated six to each side of the head by a triangular section called the frons. A labrum, which acts as an Instar #3 upper lip, is found just above the mandibles,

while the labia which is located just below Fig. 8 The five larval instars of the Cecropia the mandibles serves as a lower lip. The moth maxilla, which works with the labia to hold Fig. 9 The first larval

instar of the Cecropia moth, actual size of 1/4 to 5/16 inches.

Fig. 10 The second

larval instar of the Cecropia

moth, actual size l/2to 3/4 inches.

Fig. 11 The third larval

instar of the Cecropia moth, actual size 1 1/4 to 1 1/2 inches. Fig. 12 The fourth larval instar of the Cecropia moth, actual size of 2 to 2 1/2 inches.

Fig. 13 The fifth larval instar of the Cecropia moth, actual size of 4 to 5 inches 7

The nervous system of the larva con species, although environmental factors can sists of a tiny brain and a paired ventral nerve increase or decrease the number. The stages cord dotted with pairs of ganglia, that extends between the molts are called instars. Here, the length of the body (Douglas, 1989). growth of the larva is the major characteristic. The circulatory system of the caterpil After the last instar stage, the larva undergoes

system" lar is an "open The larval blood or a metamorphic molt to become a pupa. The haemolymph is circulated via a dorsal, tubular pupa does not feed, and its energy must come heart that receives blood from body cavities from those foods ingested while a from paired holes of the segments called ostia larva"(Gilbert, 1994). and from a posterior opening. The heart in The Cecropia moth undergoes six turn pumps haemolymph anteriorly where it is molts in its life cycle. The embryo develops carried back into the rest of the body into the first-instar larva and then molts to (Douglas, 1989). become the second-instar larva. Subsequent

The digestive system begins with the molts separate the second instar from the third mouth, leads to the esophagus, and to the instar, the third instar from the fourth instar, malpighian tubules, where nitrogenous wastes the fourth instar from the fifth instar, and the are filtered and ion regulation takes place. fifth instar from the pupa, and the pupa from The malpighian tubules lead to the hind intes the adult (Gilbert, 1994). tine and finally to the rectum, through which "The molting process is a very com wastes are eliminated (Douglas, 1989). plex series of biochemical reactions under The caterpillar moves via the dorsal neural and hormonal (neurosecretory) con- longitudinal muscles, the ventral longitudinal trol"(Douglas,1989). Stretch receptors or muscles and the dorsal-ventral muscles, which neural sensors inform the brain when the are located in each segment, just under the exoskeleton has been stretched to capacity. skin. These muscles produce the crawling This and other physiological responses inform motion of the caterpillar through specific con the larva that it is ready to molt. traction and relaxation patterns that produce a Consequently, the brain hormone PTTH (pro- compression wave in the last segment that thoraciocotropic hormone) that stimulates the moves forward toward the head. These mus production of other hormones is released cles also function to maintain the hydrostatic (Gilbert, 1994). PTTH travels from the neu pressure within the larval body. If the larva is rosecretory pars intereereballs down nerves to punctured, the muscles contract around the site the corpora cardiaca, which is the neuro- of injury to seal it off and prevent bodily flu haemal area of the brain (Douglas, 1989). ids from leaking out (Douglas, 1989). PTTH is released from the corpora car- diaca and stimulates the Prothoracic glands Metamorphosis to produce the molting hormone ecdysone Moths and Butterflies undergo a sud (Douglas, 1989). However, ecdysone is not an den transformation from larva to adult. The active hormone, and must be activated by a caterpillar progresses through a series of molts heme-containing oxidase to become the active as it increases in size. This is because the hormone 20-hydroxyecdysone (Gilbert, 1994). caterpillar must produce progressively larger Each molt involves one or more pulses exoskeletons and shed the old ones, to accom of 20-hydroxyecdysone, which stimulates modate growth. "The number of molts before epithelial cells to divide and secrete sub becoming an adult is characteristic of the stances that form the new Brain Neurosecretory cells

Corpus cardium

Juvenile Corpus Prothoracicotropic hormone (JH) allatum hormone (PTTH)

Prothoracic\ JH Binding protein (JHBP) gland

JL JH-JHBP i Mitochondria

20-Hydroxyecdysone

Chromosomes Chromosomes Chromosomes \ RNA (L) RNA (P) RNA (A) Protein Protein Protein Synthesis Synthesis Synthesis

Larval Pupal Adult structures structures structures

Fig. 14 The hormone regulation of metamorphosis. (adapted from diagram found in Gilbert, 1994) exoskeleton (Gilbert, 1994; Douglas, 1989). neously supplied JH into the last-instar can "For a molt from a larva, the first pulse pro elicit an extra larval molt, delaying the pupal

molt" duces a small rise in the hydroxyecdysone (Gilbert, 1994). The giant caterpillar concentration in the larval haemolymph and may then molt to a giant chrysalis that pro elicits a change in cellular commitment. duces a giant adult. These strange changes are

The second, large pulse of hydrox possible because even the first instar caterpil yecdysone initiates the differentiation lars have imaginal discs, which are groups of events associated with molting. The immature cells that must potentially become hydroxyecdysone produced by these pulses adult tissue (Douglas, 1989). commits and stimulates the epidermal cells During the last instar, the larva to synthesize enzymes that digest and recy increases in size and gains much weight. Just

exoskeleton" cle the components of the prior to pupation the caterpillar's color may (Gilbert, 1994). change as the larval exoskeleton forms under

In the case of the Cecropia moth, the skin. Caterpillars often travel significant

environmental conditions can impact molt distances to locate a suitable site for pupation ing. The Cecropia pupa over-winters in a (Douglas, 1989). state of diapause. If the pupa is not

exposed to at least fourteen days of cold The Chrysalis

weather, the pupa will remain in diapause When ready to proceed to the pupal indefinitely (Gilbert, 1994). stage, the larva suspends itself from an appro If the larva is molting to yet another priate substrate like a branch, by spinning a larval instar, the corpora allata, a pair of silk pad or a u-shaped girdle. Most larvae neurosecretory lobes in the brain, secretes spin a silk pad, and hang head downward. juvenile hormone. Juvenile hormone is the Those that produce a silk girdle (ie. pierid, second major effector hormone in butterfly papilionide and lycaenid larva) face horizon and moth development, and stimulates the tally or upward, but seldom downward. In epithelial cells to maintain "the larval status addition, there are a few species of primitive

quo"(Douglas, 1994). butterflies that pupate underground in loose If the larva is undergoing the last soil (Baronia brevicornis) or produce cocoons molt to become the pupa, only a small or in surface leaf litter ( amount of JH is released. Lastly, if the spp)(Douglas, 1986). transition is from pupa to adult, no JH is Silk pad producing larvae "sink a

produced (the medial nerve from the brain series of special hooks on the last abdominal

to the corpora allata inhibits the gland from segments, the cremaster, into the silk pad just

producing JH), and the neurosecretory cells as the last of the larval exoskeleton slides over 'stripping' release only 20-hydroxyecdysone and the abdomen. The actual of the lar another eclosion hormone. Consequently, it val skin at the end of a molt is termed ecdysis.

is the amount of JH that determines what As the larval skin splits over the head of the

the next stage will be (Douglas, 1989; new chrysalis, the wing pads, legs, antennae, Gilbert, 1994). and mouthparts are at first almost free of the Researchers have found that when body. But, within minutes these casings join

the corpora allata is removed from larvae, the pupa proper and hours later the chrysalis

premature metamorphosis occurs, resulting appears as a single cohesive structure, even in the formation of small pupae and small though the various appendages and their

exoge- adults. Conversely, "absorption of individual cases can be easily 10

During pupation the genes for larval charac teristics are turned off, while those controlling the expression of adult characteristics are acti vated. Many characteristics of the larva are broken down, and replaced with adult tissue by the imaginal cells. The cuticle of the adult butterfly is produced by the same epidermal cells that made the larval exoskeleton. The

legs of the larva revert to germative tissue and

give rise to the structures of the adult legs.

There is a progressive replacement of larval

tissue with adult tissue that originates from the imaginal discs, and eventually gives rise to the adult butterfly or imago (Douglas, 1989).

Fig. 15 The pupa of the Cecropia moth. In nature, it is covered and suspended by silk.

distinguished"(Douglas, 1989). Though the chrysalis, or pupal stage may appear as a resting phase, many changes are taking place below the surface. The larval structures are systematically bro ken down, and embryonic cells multiply. It is only a matter of hours before the features of the adult are under way, and mouthparts, wing stubs, thoracic muscles and legs are formed. While this occurs immediately in pupa that do not diapause, diapausing pupa do not undergo these changes until appro priate environmental conditions (ie. increas ing temperature and day length) stimulate development and imaginal discs are "turned Fig. on"(Douglas, 1986). 16 The Cecropia moth emerging Metamorphosis is carried out through from its pupal casing. (The outer silken the selective expression of various genes. cocoon was removed.) 11

Many pupa interact with their environ cles of the thorax expand, it splits the ment by producing creaking, chirping or click exoskeleton of the pupa near the apex. The ing sounds. There are three ways pupae pro butterfly's head emerges first, followed by the duce sound: by rubbing pairs of abdominal legs, which are used to secure it to a perch. segments together, by thumping the body Next the abdomen comes forth and rhythmi against its supporting structure (ie. Mourning cally contracts up and down, pumping Cloak pupae) and by rubbing the ventral haemolymph into the wing veins. The wings abdomen against the case around the pro gradually expand with each contraction until boscis. Dozens of lyaenid and riodinid pupae they are fully erect. The butterfly proceeds to create clicking, chirping, creaking and hum excrete meconium, a liquid waste, clean its ming noises by the first method. Several antennae by pulling them through a space on Bronze Copper species produce both a hum the forelegs, and it also fasten its two halves ming sound and a chirp. It is theorized that of the proboscis together, to produce a func the sounds are in response to external agita tional feeding tool (Douglas, 1989). tion, and may be a defense mechanism, or per haps a way to inform potential mates in the The Adult area that they are getting ready to emerge. Adult butterflies and moths have many

stridulation" One suggests that "pupal theory traits in common with other insects including is part of a symbiotic between relationship three main body divisions (head, thorax and many lycaenid species and attendant ants abdomen), compound eyes, antennae, three which milk the caterpillars to obtain their pair of legs and a hard exoskeleton of chitin.

sweet secretions. The secretions which are All higher Lepidoptera also possess a suction released from glands in the seventh abdominal ing proboscis, long antennae, and two pair of segments may continue to be secreted during scale covered wings (Opler, 1992). The main pupation, and the sounds may serve to attract difference between butterflies and moths is the ants (Douglas, 1989). that most butterflies are diurnal, have brightly Pupation lasts between four to fourteen colored wings, possess clubbed antennae, and summer conditions. But days under optimal lack the wing coupling anatomical feature, the diapausing pupae may remain in a suspended frenulum, which most moths possess. The state for up to two years when subjected to majority of moths are nocturnal, and have harsh environmental conditions. "This delay feather-like antennae (Opler, 1992). in development is induced by environmental conditions such as day length, but cannot be terminated until a pre-determined period has

passed. This ensures that the insects do not

emerge too early during temporary favorable periods only to be caught out by a resumption of harsh (Smart, 1975). Metamorphosis progresses as the imag inal discs differentiate to form the structures of

the adult. The pupa gradually darkens in color of and becomes transparent as it nears the end and pat metamorphosis, so that the wing color

tern is visible (Douglas, 1989). As the imago inhales air and the mus Fig. 17 The adult Cecropia moth. 12

Head The Eve

The eyes of the adult are composed of from two to twenty-thousand transparent facets, the ommatidia. Most insects, butter

flies included, have better vision in the ultravi olet wavelengths (Opler, 1992; Douglas, 1989). Vision is obtained as light passes

through the corneal lens to a crystalline cone

like structure, to an optical rod, the rabdom.

The individual ommatidium sends out infor mation about the light intensity it receives. There are pigment cells located around the crystalline cone that keep ommatidia from Fig. 19 Scanning electron micrograph of the being stimulated by more than one light insect eye source. The butterfly in turn sees a mosaic picture, similar to a pixelated computer print Butterflies (and many bee species) "are out. The resolution of sight is determined by attracted to certain flowers based on their the number of ommatidia, the development of ultraviolet color patterns. Furthermore, some the optic lobe of the brain, and by sex. This is butterfly species are sexually dimorphic when because males generally have more ommatidia viewed under these wavelengths. Males and and consequently they possess better vision females that look alike under visible light dif light" needed for locating a mate (Douglas, 1989). fer considerably under ultraviolet (Douglas, 1989).

The eyes of butterflies also have reflec

tive spots that appear to move as one looks at it from different angles. In 1963, Japanese

antenna scientists Yaki and Kayama developed a clas

sification system for butterflies based on reflective eyespot patterns (Douglas, 1989). labial palps compound, e The Antennae The antennae of the butterfly are made up of several components: a donut shaped proboscis scape, a wrist-like pedicel and a segmented

flagellum which also forms the terminal club

of the antennae (Douglas, 1989). The anten nae are partly covered by scales and hairs. uced fore leg The under portion of the terminal club, called mid leg the nudum is the part of the antennae that does not possess scales or hair (Opler, 1992). True butterflies have rounded antennal clubs, while most skippers have an extension, the Fig. 18 The head of the Pearl-Bordered Fritillary. apiculus (Opler, 1992). (adapted from photograph found in Smart, 1975) 13

Fig. 20 Scanning electron micrograph of the Fig. 22 Scanning electron micrograph of antenna of the Cecropia moth. the antenna of the Cecropia moth showing the detailed sensory cells of the antenna.

The antennae possess sensilla (sensory organs and cells) that are important in detecting olfactory stimuli, such as pheromones. Pheromones, which are insect hormones, are an important means by which insects attract and recognize potential mates.

It is believed that the hairs of the antennae

may be important in pheromone and other chemical reception, but function primarily in distortion," touch reception and "mechanism

like when the antennae bend to touch an

object (Douglas, 1989).

The Proboscis The proboscis or feeding tool is locat ed on the underside of the head, between the two three-segmented labial palpi (Opler, 1992). It is a long suctioning device used to take up liquids such as flower nectar, juices from fruits, flows, or moisture from wet Fig. 21 SEM of the antenna of the Cecropia sap but- mud, sand or (Opler, 1992). Most moth at increased magnification. dung 14 terflies rely mainly on sugars found in nectar femur, tibia, and five segmented tarsus. A for energy (Douglas, 1989). The length of the tarsal claw terminates each leg, and is sur proboscis is indicative of the feeding habits of rounded by a pad called the pulvillus. This the species; for example, nectar eaters have a basic leg structure varies; for example, the long proboscis while those that feed on sap or front legs or prolegs of some families are fruit a have relatively shorter proboscis (Opler, greatly reduced and not functional for walk 1992). Some scientists believe the proboscis ing (Opler, 1992). The prolegs are usually may have evolved first as a lapping tool, or smaller than the midleg and hindleg and those water extracting tool, before becoming a suc of some female nymphalid butterflies have tioning device (Douglas, 1989). pits and hairs that help them identify appro The proboscis functions by pumping priate larval plants. The prolegs are also used fluids through the hollow food channel formed by female Red Admiral butterflies to rapidly

"zippered" "drum" by the two halves, into a cibariwn the surface of the larval plant leaves, or anterior suction cavity. Two sets of muscle to perhaps test the suitability (texture, rough dilate the cibarium and the cavity expands, ness, strength, chemical composition) of the producing a vacuum. Stimulation of chemical plant for oviposition (egg deposition). As receptors in the tarsi of the legs will cause the mentioned earlier, the tarsal segment contains butterfly to probe for nectar using its pro receptors. These are called triclioid sensilla, boscis, which has additional sensors at its tip. and are used to determine sugar composition "If a butterfly is placed near a drop of honey of foods, and the suitability of plants for on a wet towel, the proboscis is used as a fine oviposition (Douglas, 1989). probe, gently tapping near the spot of honey in The thorax houses muscles and several

"tubes" order to find the highest concentration of including: "a hollow dorsal aorta, a sugar that can be efficiently withdrawn with solid ventral nen>e cord, and a hollow esoph out overtaxing the physical limits of the pro agus that leads to a crop in the abdomen. boscis. However, if the honey and water solu The aorta is assisted in its periodic pumping tion is mixed evenly before spreading it on the action by accessory pulsatile organs, which towel, the butterfly will remain in one place force haemolymph to percolate into the heart interior" without extensive probing"(Douglas, 1989). (Douglas, 1989). Recent research Butterflies may spend up to a minute probing shows that the heart, abdominal muscles and an individual flower for nectar (Douglas, accessory pulsatile organs are all involved in 1989). the periodic transport of haemolymph.

Scientists have also determined that the but Thorax terfly's diaphragm is important in The thorax, like the head and abdomen haemolymph distribution throughout the is mostly covered by scales and fine hair abdominal cavity. The abdominal muscles (Douglas, 1989). It is made up of the three function to assist in both heart contraction segments: prothorax, mesothorax and and tracheal ventilation (Douglas, 1989). metathorax. Each thoracic segment has a pair The thorax contains the "structures of

motility" of legs attached, while the wings attach to the and is full of muscles to operate the mesothorax and metathorax (Opler, 1992). legs and wings. These muscles are composed of striated fibers, and are small but very pow The Legs erful. Muscle strength is directly proportion The legs of the adult are composed of al to the cross-sectional area, so the larger the five distinct segments: the coxa, trochanter, cross-sectional area of the muscle, the greater 15 the strength of the muscle, relative to its size. veins.The areas outlined by veins are cells, the The muscle cells receive their energy from the largest being the discal cell. The wings are

sub- mitochondria which produce ATP. Over twen also described by areas: basal, postbasal. ty percent of flight muscle is mitochondria median, median, and postmedian, and by the (Douglas, 1989). margins: costal (the leading margin), outer (the outer edge), and inner (the inner edge). The Wings The apex describes the tip of the forewing Butterflies and moths possess two where the outer and costal margins meet, pairs of wings, the /brewings and the hind- while the anal angle describes the area of the wings (Opler, 1992). They are of course used forewing where the outer and inner margins for flight, but are also important in regulating meet (Opler, 1992). body temperature, courtship, and escaping The wings do not have any internal predation. The wings are composed of veins, muscles, but are moved by huge longitudinal sealed between two layers of transparent bands of muscle that attach to internal plates chitin and covered by scales. The veins are located between the second and third thoracic the branching framework of the wings and segment. They are positioned so that they act house tracheae, associated nerves, and circu like levers during flight. Upstroke is lating haemolymph. The chitin is a strong, achieved when the dorsal-ventral muscles yet flexible supporting layer of the adult contract, pushing the top of the thorax down exoskeleton to which the scales are attached on the bases of the wings, and pushing the by a tormogen socket (Douglas, 1989). sides of the thorax outward, causing the distal ends of the wings to move upward.

Downstroke is achieved when the dorsal-ven

tral muscles relax and the longitudinal mus

cles that run between the segments contract. This pushes the top of the thorax upward, the pleura are forced inward, and the wings fall

downward. The dorsal-ventral muscles and

the longitudinal muscles are termed indirect

flight muscles, and only control the up and down motion of the wings during flight. It is the directflight muscles which allow the but terfly to change the angle of incline of its wings in order to change its flight path 1989). Fig. 23 The veins and cells of the forewing (Douglas, Though wings appear quite and hind wing: SC=subcosta; R=radius; butterfly M=media; CU=cubitus; A=anal; LDC=lower flat, they are slightly curved during flight, much like an airfoil. Most butterflies become discocellular cross-vein; RS=radial sector

(adapted from diagram found in Douglas, 1989) airborne with just a few flaps of their wings,

"lift" as the upward becomes greater than the

Butterflies are classified largely by downward pull of gravity. The butterfly their vein design and wing color pattern. The moves its wings in a slanted figure eight pat

main veins are the casta, subcosta, radius, tern during flight (Douglas, 1989). media, cubitus and one or more anal 16

As mentioned earlier, the wings are covered by scales which give butterflies their unique appearances. The scales are usually finely ribbed, and may vary in size, shape, color and even function (Opler, 1992).

Androconia for example, are modified "sex

scales" that release the pheromones to attract mates. They are located on the legs and body of males, as well the wings (Douglas, 1989).

Scales are formed when the trichogen,

a giant epidermal cell produces a club-

shaped extension which in time flattens out

to become a scale. It is not until the cell

membrane of the scale degrades that the air-

filled, grid-like structure of the scale is

tor- apparent. Each wing has thousands of mogens that overlap like roof shingles, cloaking the wing (Douglas, 1989). The scales have a variety of functions, Fig. 24 Micrograph of the scales of the including: they increase the wings capacity Cecropia moth. to hold heat, insulate the body, are important

Fig. 26 Micrograph of the Fig. 25 Micrograph of a closer view of long scales of the Cecropia moth, which create the moth's the scales of the Cecropia moth. furry appearance. tarsus

Fig. 27 Profile view of the Black (adapted from photograph found in Smart, 1975) in temperature regulation, aid in generating themselves appear startling and unattractive. thrust for flight, may produce sexual attrac- or possessing coloration that indicates that tants, produce the color patterns of the organ they are unpalatable or poisonous (Smart, ism, and may even help butterflies escape 1975). from spider webs by pulling from their The small eggs of butterflies and pedicels to free the butterfly (Douglas, 1989). moths are usually laid on the underside of leaves, where they have less chance of being Abdomen noticed. Yet, the eggs of numerous species are The abdomen is an elongated structure parasitized by wasps and other insects. The that tapers at the ends, and houses the majority slow moving, soft bodied larvae on the other of digestive, excretory and reproductive hand are much more at risk for attack, and organs, as well as the energy storage struc have consequently evolved a variety of protec tures, the fat body (Opler, 1992). There are tive adaptations. Some larvae spin webs of eight pregenital segments composed of a scle- silk, and live in groups within them (ie. the rotized dorsal tergite and a ventral sternite, commonly seen Tent caterpillar). The skip with an unsclerotized pleurite on each side pers produce a shelter of silk and leaves or (Douglas, 1989). grass to hide themselves. Many larvae are The last two segments of the abdomen camouflaged by their host food plant. The contain the genitalia. Insemination occurs early larvae of the White Admiral camouflage when the male grasps the female using its themselves with their own excretions, while paired claspers, and inserts its penis into the the later larval instars rely on protective col female opening, the atrium, and into the duc oration for defense. The young instars of tus bursae. The penis delivers the males swallowtails (Papilionidae) resemble bird sperm package, the spennatophore, into the droppings, while the later instars are brightly ductus bursae, to the corpus bursae of the colored. Some species of Papilionidae will female, where it is stored. The spermatophore also present a pair of fearsome looking false is opened and sperm swim up the ductus semi- eyes when threatened. Swallowtail larvae also nalis, and penetrate the micropyle (outer cov emit foul odors from a gland behind the head ering) of the egg as it passes down the called the osmeteria which are everted when oviduct. The male of a few species secrete a threatened. Other families may have scent sphragis around the female genitalia during glands in other areas of the body. Monarch mating, which prevents the female from mat butterfly larvae are poisonous from consuming ing with other males, thus ensuring that the the poisonous milk weed plant. The eggs are fertilized by his sperm (Douglas, Nymphalidae larvae are frequently brightly 1989). colored and possess an armour covering of sharp spines. "Some gregarious larvae may

Methods of Self Defense jerk their bodies in unison to deter predators.

All of the developmental stages of the Occasionally a palatable larva may copy or butterfly are at risk of predation by other mimic a distasteful one and avoid predation

way" insects and spiders as well as by vertebrates this (Smart, 1975). like birds, reptiles and small mammals. Pupae are frequently protected by their Butterflies have in turn evolved protective shape, for example, the Orange Tip looks methods that reduce the risk of predation, like much like a seed pod, or by their coloration. and moth closely resembling their environment, making Many butterfly species will form a 19 cocoon from dried leaves to camouflage the al and yearly increase and decrease in popula pupae. The larvae will often search out a safe tions, the reason for lack of colonization of place for pupation, consequently pupae are apparently suitable habitats, or the reason for a often found in inconspicuous places (Smart, colony's sudden extinction-often seems more

science" 1975). of ant art than a (Douglas, 1989). Adults are most vulnerable when they Factors that effect population structure first emerge from the pupae, and are still include: climate, microbial pathogens (fungi,

unable to fly. As soon as the wings have bacteria and viruses), avian predators, parasitic expanded with haemolymph and dried, they arthropods (tiny flies and mites) and parasiti- can take flight to avoid many predators. doids (parasitic wasps and flies) and insect Danger often waits on the flowers that butter predators (ie. predacious ants) (Douglas, flies frequent. The hunting spider Misumena 1989). for example, will lie in wait on plants fre Butterfly / moth dispersal is often quented by fritillaries and spring on them as affected by weather conditions, feeding and they alight the flower (Smart, 1975). mating opportunity, competition for food, pre Wing coloration is important in adult dation, and other natural phenomena. "The survival. Some wings have a "brilliantly col range of dispersal is important, because dis ored upper surface but are cryptically colored perses are more likely to contribute their below so that when they are folded together at genes to other colonies, hence affect the

species' rest the insect completely harmonizes with its course of the evolution"(Douglas,

surroundings" (Smart, 1975). The wings of 1989). Also, evolutionists determined that

the Lycaenidae have eyespots at their edges smaller populations of certain species (ie.

head" and long tails which act as a "false to C.periphyle) possess more rare genes in their attract the predator to the wings, allowing the gene pool. This may be particularly important butterfly to escape with its body undamaged in smaller populations, to be carrying these

case" (Smart, 1975). The Io has a pair of owl-like extra alleles "just in (Douglas, 1989). eyespots on the top surface of its wings that it quickly flashes at threatening birds. The trick Migration startles potential predators, allowing the moth Many butterfly species disperse in

extra moments to make its escape. what appears to be a random manner, with

The monarch butterfly's bright colors their range of distribution varying from year to

inform would be predators of its noxious taste. year. Yet, others travel great distances, often All stages of the Monarch are poisonous and thousands of miles, en masse or individually

unpalatable because the larvae feeds on milk on their migration. While dispersion is fre

weed. Once a bird has attempted to eat a quently undirected, migrations are usually pre Monarch, it won't do so again. The Viceroy dictable, directed, and seasonally determined

moth takes advantage of the Monarch's repu (Douglas, 1989). tation, by mimicking its colors, and conse Migration may be in response to changes in light hours and quently avoiding predation. day temperatures, scarcity of adult food sources or because or

stage" Population Structure because of "overcrowding in the larval There is still much that is unknown (Douglas, 1989). The Monarch (Danus plex-

population structure of lepi and Painted concerning the ippus) Lady (Vanessa cardui) doptera and other insect species. "The dynam occupy large expanses of the temperate zone

season- ics of colonization of new habitats, the during summer, then environmental changes 20 stimulate the fall brood to migrate south. adult. The process by which the immature They migrate in large groups whereas many butterfly is transformed from one stage to the other butterflies species, like the Dainty Sulfur next is called metamorphosis, which is a gene migrate as individuals (Douglas, 1989). regulated process.

Monarch's over winter in Mexico's Butterflies reproduce sexually, and the

Central Transvolcanic mountain range, and females deposit eggs on or near suitable larval remain there in a state of reproductive dia food sources. The larvae hatch from between pause until lengthening daylight hours and three to ten days of oviposition (unless in dia warmer temperatures stimulate the butterflies pause) and feed on their host plants until they to leave their repro- tree roosts and to become reach full size. The larvae of many species go ductively active. They make the journey through a number of molts, resulting in a north, depositing eggs on milkweed plants series of larval instars, before becoming a along the way. Researchers do not know just pupae. The two main hormones of metamor how far the spring migrants reach, but believe phosis are PTTH and JH. When ready to pro it is their offspring that continue the journey ceed to the pupal stage, the larvae spins a silk frog" north, in a "leap type manner. They ride pad or girdle which it uses to suspend itself. the Southerly winds northward and orient It then strips the larval skin (ecdysis) to reveal using negative sun orientation and magnetic the new chrysalis. During the pupal stage the compassing. Some Monarchs over winter in larval features are systematically broken down Florida and Texas, where they remain repro- and the adult features are formed. Pupation ductively active. These butterflies may join lasts from four to fourteen days, unless in dia up with the spring migrants in their journey pause. north (Douglas, 1989). When the newly formed butterfly is A few other migratory butterflies ready to emerge, it inhales air, splitting the include: the Gulf Fritillary (Agraulis vanillae) exoskeleton of the pupa. The basic anatomy which migrates south in order to establish of adults includes a three segmented body, breeding colonies, the Great SouthernWhite head, thorax and abdomen, two pair of wings, butterfly (Ascia monuste) which can fly in an three pair of legs (one to each body segment), undulating cylindrical formation of millions of antennae and a suctioning proboscis. individuals, and the noetropical pierid Butterflies differ from moths in that most but Kricogonia castalia which fly in huge groups terflies are diurnal, are brightly colored, have within a few feet of the ground, and the clubbed antennae, and lack the wing coupling Cloudless Sulfers which fly only a few inches frenulum, while most moths are nocturnal, are above water along Florida's northern coast. not as brightly colored, have feather like antennae and possess the frenulum. Summary Moths and butterflies are prone to Butterflies and moths are insects, and numerous natural threats, and have conse possess those characteristics (three pairs of quently evolved unique survival mechanisms. legs, one pair of antennae, a pair of maxillae, The individuals of some species migrate to a hypopharynx, a labium and wings) which warmer or more hospitable climates to over typify the Insecta. They are believed to have winter. Though many have studied the life evolve during the Triassic period from the cycles and habits of various butterflies and Stonefly. moths, there are still many unanswered ques The four developmental stages of the tions, and much more research to be pursued. life cycle include the egg, larva, pupa, and 1

About the Artist and the Art Work working up highlights, and deepening shadows. The art work exhibited in the thesis My biggest challenge was imagining how the show consisted of: nine paintings of the creatures would look with more aggressive

Cecropia moth life cycle, a painted poster lighting, and creating that in my paintings. I depicting the stages of the life cycle and their believe my ability to do this increased with relative proportions in a natural environment, a each painting. Consequently, I frequently "finished" collection of five scanning electron micro returned to pieces and reworked graphs of the antenna and the wing scales of the them. adult, a model of the adult moth at approxi The painting I most enjoyed was the life mately three and a half times it's true size, and cycle poster. I enjoyed the freedom I had in an interactive Director project illustrating the creating the layout and poses of the organisms anatomy, life cycle, and migration of the and background foliage. My primary goal was Monarch butterfly. to create a conclusive piece to tie the others All paintings were completed utilizing together, and to give viewers an idea of the rel the technique first learned from Vichai Malikul. ative sizes of the developmental stages. I had The media used includes transparent and originally planned to do this on the computer, opaque palette water color paints, Dr. Martin's using scans of the first nine paintings. I chose Bleed Proof White, and double-sided to paint this final piece so that it would show Strathmore board. I applied the paints utilizing different perspectives of the organisms, in a a dry brush technique, laying down several lay variety of poses. I feel it also demonstrates the ers of paint (with drying time between applica larvae's natural ability to become camouflaged tions) to achieve the desired color. by its environment.

"furry" As I started working-up the tex The five scanning electron micrographs ture of this moth, and the intricate features of were taken here at RIT. I spent several months all the stages, the bleed proof white was invalu last year learning how to use the scanning elec able. Because it has an acrylic base, I was able tron microscope as an independent study pro to use it to layer on top of the watercolor, and ject, and spent several more months this winter to lighten areas and add tiny details. I often shooting micrographs of the Cecropia moth and tinted it with watercolor or applied a color on Monarch butterfly. top of it. I recommend the double sided The first three micrographs displayed Strathmore board for this kind of media are of the moth's antenna, at progressively because it's thick enough to resist warping, it higher magnifications. At first only the individ normally takes color well, it is strong enough to ual segments of the feather like antenna are pre take some abuse (erasing mistakes), and I found sent, then looking closer, tiny hairs and sensory that what I couldn't remove with a secretary's cells (sensilla) of the antenna are visible. The eraser, I was usually able to pick out with an last two micrographs displayed are of the wing exacto knife. scales of the moth. As the magnification was The nine life stage paintings (Egg & increased, the gridlike pattern of the individual Larval Instar scales becomes apparent. The and Larval Instar # 1 , Larval Instar #2, proximity #3, Larval Instar #4, Larval Instar #5, Pupa, arrangement of the lines that make up the scales Emergence, Profile of Cecropia and Cecropia are largely responsible for the colors they will appear (because color is Moth ) were based largely on photo references largely due to the way acquired from The Buffalo Museum of Science, the striations detract light). to whom I owe many thanks. A great deal of The Micrographs were shot

"detailing" time was spent the paintings and using type 55 polaroid film. They were 22 scanned into Adobe Photoshop where I adjusted contrast and brightness, and cleaned up any imperfections in the original. They were printed out on transparent acetate, and mounted approximately .25 inches from their backing. This was done to facilitate the pas sage of light through the transparency, to achieve a more three dimensional looking image.

""VJ~

v? -.:

ss

Antennae time between applications.

Scales

The model of the Cecropia moth was

constructed using silk fabric, wire coat hang Model ers, masking tape, fabric stiffener, feathers, liquid starch, watercolor and acrylic paints,

silicon gel, pipe cleaners, artificial fur, a glue When the wings were adequately gun, and a drift wood base. I first constructed shaped and structurally sound, I painted the veination of the moth by cutting, shaping, them with acrylic paint. Rough areas were and taping together pieces of wire coat hang sanded down, and I painted the wings with ers. Next, I cut pieces of fabric to the approx a combination of watercolors and bleed imate size of the four wings, and soaked them proof white, in much the same manner as in a mixture of fabric stiffener and liquid the paintings. Several layers of starch. These were placed on top of and polyurethane spray and matte media were beneath the wire frame work and sculpted to applied to make the paint water/smudge the shape of the moths chitinous wing covers resistant. Next the and more layers of forewing hindwing encasing the veins. Several (right forewing with right hindwing & left stiffener and starch were applied, with drying 23 forewing with left hindwing) were attached Finally, the section devoted to with silicon gel, and allowed to set up over Monarch migration contained a video clip and night. Then the right and left sides were an animation which included narration to attached with silicon gel, and allowed to set. explain the Monarch's migratory and repro

Wire legs were built into the infrastructure at ductive patterns. this time. The wing attachments were also There are several reasons I chose but reinforced with wire and a glue gun. terfly / moth metamorphosis as my thesis I created the body of the moth with topic. I have always felt a strong attachment silicon gel, then covered it with colored pipe with nature and have often studied the life cleaners and artificial fur to achieve the stages of the Monarch butterfly. My interest desired look. Acrylic and water color paints in nature compelled me to study biology as an were applied to give the body more natural undergraduate, and became apparent in my art coloration. Beads were applied and painted work as well. Then, last year I had the rare for the eyes, and turkey feathers were cut and opportunity to study water color painting of painted for the antennae. I then stained and butterflies with Vichai Malikul, from the polyurethane coated a piece of drift wood I'd Museum of Natural History at The collected, and drilled holes to insert the wire Smithsonian Institute. During his lessons I extensions of the legs. The legs were inserted was able to pick up the fundamentals of his into the holes and secured using a glue gun. technique, which I practiced for the next sev Lastly, painted pipe cleaners and artificial fur eral months while painting insects for the were applied to the wire legs so that the moth Buffalo Museum of Science. appeared to be grasping the branch. While interning at the museum, I had The final exhibit was a Director based access to an extensive photo-library which interactive projector file on the life of the contained a complete series of photographs of

Monarch butterfly. The three main parts of the developmental stages of the Cecropia the project are anatomy, life cycle and migra moth. I decided the Cecropia would be an tion. interesting subject because of their complex The anatomy section illustrated the pri life cycle, and because they are native to west mary anatomical features of the adult butter ern New York. I'd also hoped to spark interest fly. It included photographs of the specimens, in this quickly vanishing beauty, and help to courtesy of the Buffalo Museum of Science, increase viewers to awareness to the and electron micrographs of the eye and wing Cecropia's plight. scales which I had photographed. Labels and By illustrating the intricate details of descriptions accompanied most photos, and even the smallest of the life stages, (the first mouse roll-overs allowed viewers to access larval instar is approximately one fourth of an definitions of the various body parts. inch long), I hope to increase appreciation and The section devoted to the life cycle respect for the these complicated creatures. was comprised of photographs of the various Considering the dangers and obstacles they life stages, with accompanying videos of the face (poisoning by pesticides, parasitization, life processes (a larva hatching from the egg, predation by birds and small , traffic, larval growth, molting, metamorphosis from loss of habitat and larval food sources), it is larva to chrysalis, the changing chrysalis, amazing that any survive to become adult

crosscon- emergence of the butterfly, and the feeding moths. Even more impressive is the mechanism). The video is not my own, and tinental migration of butterflies like the was utilized for educational purposes only. Monarch, which may travel from as far as Canada to over-winter in Mexico's Central Transvolcanic Mountain Range. I am amazed and intrigued by the com plexity and beauty of life, especially lives that appear as fragile these. I hope somewhere

along the way my work makes a difference and some day does nature justice. Glossary extend vertically within segments. empties the Abdomen The third and posterior segment of Ductus bursae The tube that into the insect body. corpus bursae.

of the old cuticle dur Accessory pulsatile organs Tiny organs that Ecdysis The shedding help pump haemolymph throughout the body. ing molting.

Androconial scales Modified scales of male Ecdysone A hormone of molting. butterflies that secrete pheromones to stimu Eclosion The hatching of the caterpillar from late females. the egg and the emergence of the adult from Antennae A pair of sensory organs found on the chrysalis. the head, above the mouth. Fat bodies The storage sacs for fats, that are Atrium The opening leading to the ductus especially large in diapausing and migratory bursae. butterflies.

Basal area The region near the base of the Femur The third segment of the leg. wing. Flagellum The segments of the antenna after

Chitin A nitrogenous polysaccharide con the scape.

stituent of the exoskeleton of insects. Frenulum A type of wing coupling where

Chorion The outer layer of the eggshell bristles from the top wing fit into areas of the which is produced by the secretions of the bottom wing. ovarian follicular cells. Haemolymph The blood of the insect.

Chrysalis The metamorphic instar of the but Imaginal discs Epidermal cells which are terfly. undifferentiated and embryonic and allow for

Compound eye An eye comprised of numer metamorphosis to take place.

ous ommatidia. Imago The adult insect.

Corneal lens The transparent cuticle of the Immature Eggs, larvae or pupae, but usually

ommatidia. referring to the larvae. Indirect muscles muscles cause Corpora allata A pair of lobes behind the The that changes the shape of the thorax and brain that secrete hormones. in indi control the wings. Corpus bursae The main part of the bursa rectly molts. copulatrix. Instar The larva between Juvenile hormone A hormone produced Costal vein (costa) The wing vein that forms by the corpora allata which supports the juvenile the anterior margin of the butterfly wing. and inhibits the development Coxa The segment of the insect leg closest to characteristics, characteristics. the body. of adult Labia The larva's lower paired lip. Cremaster A structure used by the pupa to a Labrum The upper lip. silk support pad. Mandibles The larva's front pair of Crochets Hooks on the plantar surface of the chewing mouthparts. larval prolegs. Meconium The waste material excreted Crop The area of the foregut behind the by the adult after emerging. esophagus. Medial vein The vein that runs longitu Discal Cell The large cell in the central area wing between the radius and cubitus. of the wing. dinally Mesothorax The second segment of the tho Dorsal longitudinal muscles The muscles rax. that extend dorsally and lengthwise between Metathorax The third segment of the thorax. segments. Micropyle The on the of the egg. Dorsal-ventral muscles The muscles that opening top >6 where sperm penetrate. the wing membrane that carry the tracheae and Molting The process by which a new cuticle nerve branches. is produced, and the old one is shed. Vitelline membrane The internal membrane flagel- Nudum The scaleless portion of the of the egg that originates from the egg cell lum of the antennae. membrane, and lies beneath the chorion.

Ocelli A simple eye.

Ommatidium An individual unit of the insects compound eye. Oviposition The process of egg laying. Pedicel The segment of the antenna that act

"wrist" like a between the scape and the fla- gellum. Pheromones Chemical hormones secreted by individual insects that may influence the behavior of other members of the species. Proboscis The coiled, suctioning feeding tube of the adult butterfly.

Prothoracic glands The glands of the brain that produce ecdysone, the molting hormone. Prothorax The first segment of the thorax.

Pupa The stage between larva and adult.

Radius Vein The wing vein that runs longitu dinally between the subcosta and the media. Rhabdom The light-sensitive rod structure of the ommatidium of the compound eye. Scales The individual structures that overlap to cover the chitin of the wing.

Scape The first segment of the antennae between the head and pedicel. Sensilla Sensory cells, which often belong to larger sense organs. Spermatophore The sperm containing pack age of the male.

Spiracles The air openings to the tracheae.

Subcosta vein The wing vein that runs longi tudinally behind the costa vein. Tarsus The final segment of the walking leg. Thorax The second segment of the body. Tibia The fourth leg segment between the femur and tarsus.

Tracheae The internal air transport tubes of insects. Trochanter The segment of the leg between the coxa and the femur. Veins The thick walled supporting tubes of 27 Bibliography

Arnett and Jacques, R. L. Simon &Schuster's Guide to Insects. New York: Simon & Jacques Inc., 1981.

Douglas, M. M. The Lives of Butterflies. Canada: The University of Michigan Press and John Wiley &Sons., 1989, 20-49, 110-111.

Gilbert, S. Developmental Biology. Sunderland: Sinauer Associates, Inc., 1994,729-738.

Mitchell and Zim, H. S. Butterflies and Moths. New York: Western Publishing Company, Inc., 1964.

Opler, P. A. A Field Guide to Eastern Butterflies. New York: Houghton Mifflin Company, 1992

Smart, P. The Illustrated Encyclopedia of the Butterfly World. Salamander Books Ltd. London. 1975, 18-21, 34-40.