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Rochester Institute of Technology RIT Scholar Works Theses Thesis/Dissertation Collections 5-21-1997 The Life cycle of moths and butterflies Mary Walter Follow this and additional works at: http://scholarworks.rit.edu/theses Recommended Citation Walter, Mary, "The Life cycle of moths and butterflies" (1997). Thesis. Rochester Institute of Technology. Accessed from This Thesis is brought to you for free and open access by the Thesis/Dissertation Collections at RIT Scholar Works. It has been accepted for inclusion in Theses by an authorized administrator of RIT Scholar Works. For more information, please contact [email protected]. Rochester Institute A Thesis of submitted to the Imaging Aits degree and of The Life Cycle of Technology Faculty of the College Sciences in candidacy for Master of Fine Arts. of Moths and by Mary E. Walter May 21, 1997 Butterflies the 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 Date r-: :'::>-f/~\"'_"Jf-+-/-+~/_7.l--.7 17 . _ _ 1 Introduction It is developed animal species are mately 800,000 which are that eighty percent of all insects. There are approxi estimated insects, 180,000 Lepidoptera (Arnett & Jacques, 198 1). In this attributes of species of paper butterflies will explain cal their attributes, doing so unique survival mechanisms, habits. shows aids Butterflies the are members of Arthropods, evolved from ancestor are branches; phylum an annelid divided into the three insects). All solar radiation ture more quickly. arthropods; head, Butterflies and moths, body other on each antennae, a thoracic segment), pair of maxillae with a a pair of a mandibles, sensory palpi, hypopharynx (a tongue like structure), a labi (a tion is for pah" specialized of increasing the species. predators. led to Basking in thermoregula insects. Butterflies broad insects winged and insects, are the Triassic during Stonefly (Douglas, arisen 1989). of head Life Cycle The appendages Of the twenty-nine the world, Lepidoptera, butterflies and orders of which skippers, is one insects in includes of the moths, largest. approximately 13,700 species of Lepidoptera in North America, and 1 80,000 in are stages of butterfly development include egg, larva, pupa, and adult. Metamorphosis is the process by which the immature stage by is transformed from butterfly Metamorphosis is to the next. gene regulation. within each of and chrysalis Therefore, groups of the different which are egg, larva, by various during turned on development. the genes of the organism remain the same throughout the ent groups of genes are throughout stages: are controlled stages of controlled changes that occur immature / pupa, genes, The one life cycle, but differ turned on and off development (Douglas, 1989). Monarch Butterfly: (Arnett & Jacques, 1981). Lepidoptera, like possess wings. explain winged and that these advantages also from the (Arnett & Jacques, 1981). world absorb tempera to aid in escape from winged the the further, and a common means of behind the maxillae) and two pair of wings (some insects have one or no pair of wings) There body muscles, enabling moths, which are thorax and like the class Insecta possess three legs (one pair of um likely by members of the phylum arthro exhibit the one pair of leg to forage longer period centipedes, millipedes, and the to Warming occurs in pockets winglets, increasing tempera believed to have in that they members of scor (crabs, lobster, segmentation of pairs of beneath the sun major dis (Douglas, elevation to increase increased dispersal type poda are similar abdomen. temperature response rates the mandibulates (the Onychophora, to air tubules ing It is (spiders, ticks, pions, etc.), the crustaceans crayfish) Arthropoda. (earthworm) the chelicerates and insects, all believed to have which are body tures of moths, like and sexual The thermoregulatory hypothesis that an increase in the size of winglets in of air Evolutionary History insects, for or as moveable winglets 1989). I evolutionary history, physi development and and colonization and migration of minute al protect openings stages of metamorphosis, that wings may have developed to aid in "aeri play to lure mates, and moths which make In protruded out flaps that attached from the thorax. It is believed dispersal" of I hope to describe the them unique from other insects. flat, widely as There other insect are several evolution of winged most accepted view which suggests is the that the theories to insects. The paranotal first orders theoiy, wings of insects fig.l Larva from egg hatching fig. 2 Full grown larva or close to the larval butterfly's food supply Douglas, 1989). For butterfly leaves lays her example, the Monarch eggs on the young, tender of the milkweed plant, while Swallowtail commonly deposits its garden ly fig. 3 Pupa of the Butterfly Buffalo Museum of Science) colored, with variations mental stage of Butterflies reproduce sexually, with on the the egg. or chorion of eggs is in color, The identification. In the female egg is a small sperm and egg are haploid in their chromosome num ber. That is, each sex cell contains half the number of chromosomes of the body cells ize the While the egg. center of each or micropyle, through which sperm permeates the chorion takes and resistant place via micropyle exchange or air pockets called plastrons, which act as gills to even when dioxide, it is to injury. Gas ducts to fertil facilitates the chorion exchange of oxygen and carbon strong (Douglas, 1989). hole develop surface structure a common means of specie Both size and species and the male contributing the sperm, while the contributes the egg. quick eggs are often white or cream depending shape Reproduction eggs on larvae devour. fig. 4 Adult (photos courtesy parsley plants, the which the Black diffuse gases, egg is immersed in water. The may also participate in respiration the (Douglas, 1989). Fig. 5 Pair female of moth mating Cecropia is on the because females have upper a moths. left, larger The identifiable abdomen. Morphology The Egg Eggs ovaries of are produced within the female oviducts which and travel to the join to form through which the eggs are outside the tubular one duct deposited to the (oviposition). Just before the egg is released, it is fertilized by a sperm cell that was stored in the corpus bursae, which is the female butterfly's internal sperm receptacle (Douglas, 1989). The female usually lays her Fig. 6 The eggs of the moth are generally laid in small clutches on the leaves of lilac, apple, cherry, boxelder, maple, birch, willow, plum and elderberry trees (Mitchell & eggs on Cecropia Zim, 1964). Fig. 7 Poster of the life -a pair of mating -a female laying cycle of the Cecropia moth. moths lilac -fourth leaf (a female lays from 200 to 300 eggs) -first larval instar (actual -second larval instar (actual -third a clutch of eggs on a size larval instar (actual -fifth 1/4 to 5/16 ") size 1/2 to 3/4" larval instar (actual -Pupa ) size larval instar (actual within is shown) its 1 1/4 to 1 1/2") 2 to 2 1/2") size size silk cocoon 4 to 5") (cut-away view 4 Below the is the chorion food vitelline membrane which envelopes and protects the developing larva. Several other layers develop between the chorion and vitelline membrane as the formation from single celled egg to the tiny the species and weather con on ditions. Some deposit their species eggs in the fall and the eggs over-winter in a state diapause, of arrest called until spring, do and not hatch when conditions are optimal body are size as lizing the cells multiply geometrically, the egg yolk for uti When nutrition. using its quently eat even the The larva fre mandibles. rest of the egg chorion, and locomotion), The Larva legs with ventral crochets grasping) full on like". A large American Harvester butterfly) to the "worm exoskeleton exoskeleton, but spiracles via abdominal and located along the sides of thoracic the larva. spiracles are valve regulated air openings body which extend to supply all body organs (Douglas, 1989). or ant (ie. the larvae hairstreak Harkenclenus thus of the and blues Glaucopsyche lygdamus Celastrina puncture soft and related carnivorous, eating aphids and insects (ie. the larvae of the North and pupae the caterpillar, with head, is caterpillars are larvae pro- I hooks (for loco exoskeleton of the exception of the larval their host plants Some size. and (Douglas, 1989). and muscles with oxygen reach with a and ten abdominal segments, of which possess non-jointed that connect to the tracheae Young larva feed the larval five throughout the they segments of possessing a pair of jointed legs terminal claw (for food manipulation The that of neighboring unhatched eggs (Douglas, 1989). until (Douglas, 1989). resulting in fluid loss and death. Gas exchange does not occur through the the egg membranes and chorion the labium could cause collapse of the exoskeleton, ready to emerge from the egg the larva chews through is found between the each motion and the egg increases in released, divided into three thoracic segments, The within the larval spinneret, from The thirteen (Douglas, 1989). The larva is maxillae and larva develops. The trans is found behind the consumption, Lastly, mandibles. which silk lava usually takes between three to ten days, depending during the and ) argiolus pseudargiolus (Douglas, 1989). Larvae thirteen larva body are composed of a segments. The head each side of the head called the frons. by lip, is found just while the labia which of simple eyes separated six to which acts as an above lower lip. The which works with Instar #3 the mandibles, is located just below mandibles serves as a maxilla, for triangular section a A labrum, upper and of the contains antennae, mandibles crushing food, and six pairs called ocelli. The ocelli are the head the labia to hold Fig. 8 The five larval instars moth of the Cecropia Fig. 9 The first larval instar of the actual size of Cecropia moth, 1/4 to 5/16 inches. Fig. 10 The larval instar moth, actual size of the second Cecropia l/2to 3/4 inches. Fig. 11 The third larval instar actual size of the Cecropia moth, 1 1/4 to 1 1/2 inches. Fig. 12 The fourth larval instar Fig. 13 The fifth larval instar of of the Cecropia moth, actual size of 2 to 2 1/2 inches. the Cecropia moth, actual size of 4 to 5 inches 7 The larva con brain and a paired ventral nerve tiny dotted with pairs of ganglia, that extends nervous system of the increase cord between the length the of body (Douglas, the The circulatory lar is an "open 1989). decrease the or growth of system of the caterpil factors species, although environmental sists of a the larva The number. molts are called is the can stages instars. Here, major characteristic. After the last instar stage, the larva undergoes system" haemolymph is heart that from and from holes blood from dorsal, body or tubular cavities of the segments called ostia haemolymph anteriorly back into the rest of the where it is begins system with the mouth, leads to the esophagus, and to the The filtered tubules, ion and where nitrogenous wastes regulation malpighian tubules tine and finally are (Douglas, 1989). These motion of ventral each segment, just muscles produce the caterpillar moves forward cles also longitudinal under which the the crawling through specific con in the last toward the function to pressure within the segment that head. These maintain mus hydrostatic the larval body. If the larva is punctured, the muscles contract around the site of injury ids from to seal leaking The Cecropia molts it off and prevent out the bodily flu transformation it increases in Butterflies size. "The an adult is adult. The through a series of molts exoskeletons and shed becoming embryo develops and then molts from the third instar from the fourth instar, fourth instar from the fifth instar, the adult and the from the pupa, and the pupa (Gilbert, 1994). "The molting neural and process biochemical is a neural sensors exoskeleton reactions under Stretch inform has been con- receptors or brain the com very hormonal (neurosecretory) trol"(Douglas,1989). This to larva. Subsequent the second instar the third when the stretched to capacity. and other physiological responses inform larva that it is ready to molt. Consequently, the brain hormone PTTH the hormone) thoraciocotropic production of other rosecretory hormones is released intere ere balls down pars the corpora cardiac a, which haemal area of the (pro- that stimulates the (Gilbert, 1994). PTTH travels from the PTTH is undergo a sud from larva to This is because the caterpillar must produce modate growth. The is the neu nerves to neuro- brain (Douglas, 1989). released and stimulates from the corpora the Prothoracic car- glands to produce the molting hormone ecdysone and caterpillar progresses as must come while a moth undergoes six cycle. fifth instar from (Douglas, 1989). Metamorphosis den its energy the second-instar molts separate instar, diaca Moths in its life plex series of dorsal traction and relaxation patterns that produce a compression wave and The a pupa. larva"(Gilbert, 1994). which the dorsal-ventral muscles, located in skin. hind intes caterpillar moves via the longitudinal muscles, the muscles and the to the rectum, through wastes are eliminated The takes place. lead to feed, not from those foods ingested become The digestive are does pupa become into the first-instar larva body (Douglas, 1989). malpighian a metamorphic molt to The heart in a posterior opening. turn pumps carried circulated via a receives paired The larval blood (Douglas, 1989). However, ecdysone is not active hormone, and must be activated by a heme-containing oxidase to become the active hormone 20-hydroxy ecdysone (Gilbert, 1994). progressively larger the old ones, to accom number of molts characteristic of before the an Each of molt involves 20-hydroxyecdysone, epithelial cells stances that to divide form the one or more pulses which stimulates and secrete sub new Brain Neurosecretory cells Corpus cardium Juvenile hormone (JH) JH Corpus allatum Binding Prothoracicotropic hormone (PTTH) \ Prothoracic protein gland (JHBP) JL i JH-JHBP Mitochondria 20-Hydroxyecdysone Chromosomes Chromosomes Chromosomes \ RNA (L) Protein Synthesis Larval structures Fig. 14 The hormone RNA (P) Protein Synthesis Synthesis Pupal Adult structures regulation of metamorphosis. (adapted from diagram found in RNA (A) Gilbert, 1994) Protein structures exoskeleton (Gilbert, 1994; Douglas, 1989). "For from a molt duces a a small rise concentration the first pulse pro in the larval haemolymph in elicits a change The second, large yecdysone larva, in the hydroxyecdysone and cellular commitment. hydrox pulse of hydroxyecdysone produced by elicit an extra JH into the last-instar larval molt, because the first instar even adult tissue commits and stimulates the epidermal cells prior (Gilbert, 1994). change as caterpil which are groups of (Douglas, 1989). During the to synthesize enzymes that digest and recy cle the components of the are cells that must potentially become immature these pulses the giant caterpillar may then molt to a giant chrysalis that pro duces a giant adult. These strange changes lars have imaginal discs, The delaying (Gilbert, 1994). The can pupal molt" possible initiates the differentiation events associated with molting. supplied neously increases in last instar, the larva Just size and gains much weight. exoskeleton" In the case of the Cecropia moth, environmental conditions can ing. The Cecropia state of pupa is (Douglas, 1989). a of cold in diapause If the larva is molting to larval instar, the in corpora yet another allata, a pair of neurosecretory lobes in the brain, secretes juvenile hormone. Juvenile hormone is the second major effector hormone in development, and moth to epithelial cells butterfly and stimulates maintain "the larval the status If the larva is undergoing the last to become the pupa, only a small amount of JH is released. transition is from produced (the to the pupa corpora allata producing Lastly, if the to adult, medial nerve JH), and the no is the tally or upward, but addition, there are a soil or (Baronia in surface what be (Douglas, 1989; is downward. In species of primitive pupate underground brevicornis) or produce cocoons 1986). pad series of special the last of the abdomen. producing larvae "sink hooks As the larval new removed when from larvae, occurs, resulting of small pupae and small Conversely, "absorption of exoge- silk pad The 'stripping' of the actual skin splits over first within minutes the pupa proper and hours is termed ecdysis. legs, antennae, free of the these casings join later the chrysalis structure, even though the various appendages and their cases can lar the head of the almost appears as a single cohesive individual just the larval exoskeleton slides over chrysalis, the wing pads, body. But, a the last abdominal on the end of a molt val skin at in loose leaf litter (Parnassius spp)(Douglas, Silk seldom few (ie. pierid, larva) face horizon lycaenid and mouthparts are at corpora allata adults. produce a silk girdle neurosecretory Researchers have found that in the formation pad, and Those that papilionide and a head downward. hang spin a silk segments, the cremaster, into the premature metamorphosis pupation Most larvae as Gilbert, 1994). the for branch, by spinning cells JH that determines will a silk pad or a u-shaped girdle. from only 20-hydroxyecdysone and another eclosion hormone. Consequently, it the next stage like priate substrate gland release amount of travel significant often a suitable site When ready to proceed to the pupal stage, the larva suspends itself from an appro JH is from the brain inhibits the under The Chrysalis butterflies that quo"(Douglas, 1994). molt forms exoskeleton Caterpillars distances to locate 1994). indefinitely (Gilbert, skin. the caterpillar's color may the larval not least fourteen days weather, the pupa will remain the pupation molt pupa over-winters diapause. If the exposed to at impact to 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 broken down, characteristics of the larva are tissue and replaced with adult imaginal cells. The cuticle of the adult by butterfly is produced by the same epidermal the that made the larval exoskeleton. cells legs of give rise to the The the larva revert to germative tissue and There is structures of the adult a progressive replacement of legs. larval tissue with adult tissue that originates from the imaginal discs, adult Fig. 15 The pupa of the Cecropia In moth. nature, it is covered and suspended by butterfly eventually gives rise to the imago (Douglas, 1989). and or silk. distinguished"(Douglas, 1989). Though the chrysalis, may appear as a changes are or pupal stage resting phase, many place below the surface. taking The larval structures are ken down, and embryonic cells multiply. is only of a matter of the adult are systematically bro It hours before the features under way, and mouthparts, wing stubs, thoracic muscles and legs are formed. While this occurs immediately in pupa do that do not not undergo diapause, diapausing these changes until priate environmental conditions ing temperature and development on"(Douglas, and day length) imaginal discs (ie. increas stimulate are "turned Fig. 16 The Cecropia 1986). Metamorphosis is the pupa appro carried out through selective expression of various genes. from its moth pupal casing. cocoon was removed.) (The emerging outer silken 11 Many pupa interact with their environ by producing creaking, chirping or click ment ing There sounds. duce sound: by three ways pupae pro are pairs of abdominal rubbing together, by thumping the body its supporting structure (ie. Mourning segments against Cloak pupae) and abdomen against boscis. Dozens by rubbing the ventral the case around the pro of lyaenid riodinid and pupae ming noises by the first method. Several Bronze Copper species produce both a hum ming It is theorized that sound and a chirp. the sounds in are response to external agita may be a defense mechanism, or per way to inform potential mates in the tion, and haps a area that they are getting ready to exoskeleton of the butterfly's head legs, it splits pupa near to the secure abdomen comes it to forth are and rhythmi fully The erect. butterfly liquid waste, proceeds clean to its pulling them through a space on forelegs, and it also fasten its two halves antennae of wings expand with each contraction until excrete meconium, a the the a perch. cally contracts up and down, pumping haemolymph into the wing veins. The they The the apex. first, followed by emerges which are used Next the gradually clicking, chirping, creaking and hum create cles of the thorax expand, the tional by proboscis feeding together, to produce a func (Douglas, 1989). tool The Adult emerge. Adult butterflies and moths have many stridulation" One theory suggests that "pupal is part of a symbiotic relationship between traits in common many lycaenid abdomen), compound eyes, antennae, three which milk species and attendant ants the caterpillars to The sweet secretions. released from segments glands may their obtain seventh abdominal to be continue secreted pupation, and the sounds may serve to the ants during attract (Douglas, 1989). under optimal summer conditions. diapausing pupae may remain in But a suspended for up to two years when subjected to harsh environmental conditions. "This delay state in development is induced conditions such as by emerge This day length, but ensures too early cannot insects do that the during periods only to be of harsh legs ing divisions (head, and a proboscis, insects hard long exoskeleton of chitin. also possess a suction antennae, and two scale covered wings difference between butterflies butterflies by are and moths diurnal, have lack the wing coupling frenulum, anatomical the majority of feather-like moths are antennae becomes transparent as metamorphosis, so that the tern is visible it nears wing (Opler, 1992). be has not imag the end of color and pat (Douglas, 1989). As the imago inhales air and the mus brightly Fig. 17 The adult The nocturnal, and have inal discs differentiate to form the structures of the adult. The pupa gradually darkens in color and is feature, which most moths possess. a resumption progresses as main colored wings, possess clubbed antennae, and (Smart, 1975). Metamorphosis pair of (Opler, 1992). The temporary favorable caught out including thorax and environmental terminated until a pre-determined period passed. body All higher Lepidoptera that most Pupation lasts between four to fourteen days main pair of secretions which are in the three with other Cecropia moth. the 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 There the light are pigment cells crystalline cone that being intensity located keep it receives. around the ommatidia Fig. 19 from insect by more than one light butterfly in turn sees a mosaic stimulated source. The The resolution of sight is determined by the number of ommatidia, the development the optic lobe because and males of the brain, and generally have by sex. of This is more ommatidia consequently they possess better vision for locating a mate (Douglas, 1989). the eye attracted Furthermore, ultraviolet color patterns. butterfly some sexually dimorphic when these wavelengths. Males and species are viewed under females that look fer considerably needed electron micrograph of Butterflies (and many bee species) "are to certain flowers based on their picture, similar to a pixelated computer print out. Scanning light dif alike under visible light" under ultraviolet (Douglas, 1989). The eyes of butterflies also have looks tive spots that appear to move as one it from different scientists antenna Yaki Kayama developed for butterflies based reflective eyespot patterns labial compound, at In 1963, Japanese angles. and sification system reflec palps a clas on (Douglas, 1989). e The Antennae The antennae of the butterfly of several components: a up donut are made shaped proboscis scape, a wrist-like pedicel and a segmented flagellum of uced mid fore leg leg the antennae nae are The the partly nudum is the of the Pearl-Bordered Fritillary. apiculus found in Smart, 1975) anten hairs. the antennae that not possess scales or while most skippers photograph by scales and the terminal club, called part of True butterflies have (adapted from terminal club (Douglas, 1989). The covered under portion of does Fig. 18 The head forms the which also hair (Opler, 1992). rounded antennal have (Opler, 1992). an clubs, extension, the 13 Fig. 20 Scanning antenna of the electron micrograph of the Cecropia moth. Fig. 22 Scanning electron micrograph of the antenna of the the The Cecropia detailed sensory cells) that are detecting hormones, insects such as Pheromones, are an (sensory important in olfactory stimuli, pheromones. which are important means may be important in chemical reception, which of the antennae pheromone and other but function primarily in touch reception and "mechanism when object by insect attract and recognize potential mates. It is believed that the hairs like showing cells of the antenna. antennae possess sensilla organs and moth distortion," bend to touch the antennae an (Douglas, 1989). The Proboscis The proboscis or feeding ed on the underside of the two three-segmented Fig. 21 SEM moth at of the antenna of the Cecropia increased magnification. labial palpi the (Opler, long suctioning device used to up liquids such as flower nectar, juices 1992). take It is tool is locat head, between a from fruits, sap flows, mud, sand or dung or moisture from (Opler, 1992). Most wet but- 14 terflies rely mainly on sugars found in femur, tibia, nectar for energy (Douglas, 1989). The length of the proboscis is indicative of the feeding habits of for the species; long have example, nectar eaters feed proboscis while those that on sap by rounded basic or front legs leg or prolegs of some may have or smaller than suc of some female pits and hairs that first lapping tool, before becoming a water as a extracting tool, tioning device (Douglas, 1989). The functions proboscis by "zippered" the two halves, into or anterior suction cavity. dilate the boscis, "If to probe has is butterfly a for sets of muscle nectar using its placed near a towel, the proboscis gently tapping to find the highest near sugar that can will cause the pro additional sensors at probe, order legs the drop is of its honey used as a the spot of tip. is mixed towel, the fine honey evenly before spreading it butterfly plant in on the one place without extensive probing"(Douglas, 1989). Butterflies may spend up to a minute probing an individual flower for nectar (Douglas, 1989). for covered by scales and It is (Douglas, 1989). made up and abdomen fine hair of the three metathorax. Each thoracic segment has the receptors. These foods, are called triclioid sensilla, determine and are used to sugar composition and the suitability of (Douglas, 1989). oviposition The thorax houses plants including: "a hollow dorsal aorta, The leads that agus is aorta action for muscles and several hollow solid ventral nen>e cord, and a by interior" in the to a crop in its assisted accessory force haemolymph pulsatile also which into the heart to percolate heart, pumping organs, research abdominal muscles and pulsatile organs are all accessory a esoph abdomen. periodic (Douglas, 1989). Recent shows that the involved in haemolymph. determined that the but diaphragm is important in haemolymph distribution throughout the The abdominal cavity. function to and segments: pro thorax, mesothorax and rough of earlier, the tarsal segment contains terfly's Thorax (texture, (egg deposition). As oviposition Scientists have is mostly prolegs are also used composition) the periodic transport of The thorax, like the head appro mentioned in withdrawn with chemical ness, strength, concentration of will remain identify female Red Admiral butterflies to rapidly the surface of the larval plant leaves, "tubes" be efficiently and those butterflies have to perhaps test the suitability of out overtaxing the physical limits of the pro boscis. However, if the honey and water solu tion by walk usually hindleg them The plants. are "drum" cavity expands, Stimulation of chemical which on a wet Two larval priate a cibariwn cibarium and the producing a vacuum. receptors in the tarsi of butterfly by pumping channel formed fluids through the hollow food nymphalid help example, the families prolegs are the midleg and sur This functional for reduced and not (Opler, 1992). The ing is and pulvillus. for structure varies; greatly evolved leg, the a pad called a relatively shorter proboscis (Opler, 1992). Some scientists believe the proboscis A segmented tarsus. tarsal claw terminates each a fruit have five and assist tracheal ventilation The thorax a pair abdominal muscles in both heart contraction (Douglas, 1989). contains the "structures of motility" of legs attached, while the wings attach to the mesothorax and metathorax (Opler, 1992). and legs of striated erful. The Legs The legs five distinct of the segments: is full and wings. fibers, Muscle of muscles These muscles are composed and are small strength to operate the is but very directly pow proportion the cross-sectional area, so the larger the adult are composed of al to the coxa, trochanter, cross-sectional area of the muscle, the greater 15 the strength of the muscle, relative to The muscle cells receive flight percent of size. their energy from the ATP. Over mitochondria which produce ty its is muscle twen mitochondria (Douglas, 1989). Butterflies pairs of wings, the wings body sealed branching and by of scales. framework by haemolymph. The veins are of the wings and is chitin of a strong, (the leading outer edge), and The apex describes the tip (Douglas, 1989). by forewing edge). meet, area of the outer and inner where the outer forewing the describes the while the anal angle sub- postbasal. margin), of the wings are inner (the inner the margins (Opler, 1992). meet The muscles, do wings but have any internal by huge longitudinal not are moved internal of muscle that attach to located between segment. like levers They are positioned so that they during flight. Upstroke is achieved when the bases on the plates the second and third thoracic dorsal-ventral contract, pushing the the adult to which the scales are attached a tormogen socket basal, areas: (the bands transparent The discal the The median, median, and postmedian, and escaping associated nerves, and circu flexible supporting layer exoskeleton by important in regulating between two layers house tracheae, yet They are of course used wings are composed of veins, chitin and covered lating /brewings are also The being described veins are cells, cell. where the outer and costal margins and the hind- temperature, courtship, predation. the and moths possess two (Opler, 1992). for flight, but largest also margins: costal The Wings by veins.The areas outlined of the muscles down of the thorax top act wings, and pushing the sides of the thorax outward, causing the distal ends of the wings to move upward. Downstroke is achieved when tral muscles relax and the cles that run This between the pushes the longitudinal segments contract. top and the wings downward. The dorsal-ventral the longitudinal flight muscles, down the terfly in and hind veins and cells of the wing: forewing discocellular (adapted from diagram found in order to change Though flat, they much cross-vein; RS=radial sector are like butterfly slightly an airfoil. airborne with Douglas, 1989) just a "lift" as the upward Butterflies their vein design are classified and wing largely by color pattern. The main veins are the casta, subcosta, radius, media, cubitus and one or more anal during flight. It is incline its flight the of but its path (Douglas, 1989). SC=subcosta; R=radius; M=media; CU=cubitus; A=anal; LDC=lower indirect the up and muscles which allow to change the angle of wings Fig. 23 The the wings motion of direct flight only control fall muscles and muscles are termed and mus of the thorax upward, the forced inward, pleura are the dorsal-ven downward moves tern its curved in during flight, Most butterflies become few flaps of becomes greater than pull of gravity. wings during wings appear quite The a slanted their wings, the butterfly figure flight (Douglas, 1989). eight pat 16 As covered mentioned earlier, the wings are by scales which give The unique appearances. finely butterflies their scales are usually may vary in size, shape, function (Opler, 1992). ribbed, and color and even Androconia for example, are modified "sex scales" that release the They mates. body are pheromones located on the to attract legs of males, as well the wings and (Douglas, 1989). Scales are formed when the trichogen, a giant epidermal cell produces a shaped extension which to become a scale. It is membrane of the scale filled, in time club- flattens not until the degrades that grid-like structure of the scale out cell the air- is Each wing has thousands of torthat overlap like roof shingles, apparent. mogens Fig. 24 Micrograph Cecropia of the scales of the moth. Fig. 25 Micrograph of a closer view of the scales of the Cecropia moth. cloaking the wing (Douglas, 1989). The scales have a variety of functions, including: they increase the wings capacity to hold heat, insulate the body, are important Fig. 26 Micrograph the Cecropia moth, furry appearance. of the long which create scales of the moth's tarsus Fig. 27 Profile (adapted view of from the Black Swallowtail photograph butterfly found in Smart, 1975) in temperature regulation, aid in generating thrust for flight, may tants, produce the color patterns of the organ ism, and from attrac- produce sexual butterflies help by pulling from their free the butterfly (Douglas, may even to is abdomen The 1989). an elongated structure that tapers at the ends, and digestive, excretory houses the majority body energy storage struc (Opler, 1992). There are eight pregenital segments composed of a dorsal tergite rotized scle- and a ventral sternite, with an unsclerotized pleurite on each side (Douglas, 1989). segments of the abdomen Insemination contain the genitalia. female using its inserts its penis into the claspers, and female opening, the bursae. The sperm penis delivers duc the the males female, where it is bursae corpus stored. opened and sperm swim The up of the the ductus semi- nalis, and penetrate the micropyle (outer cov ering) of the egg The oviduct. as it male of a sphragis around the few female mating, which prevents the ing down the passes during female from mat with other males, thus ensuring that the eggs are fertilized by his sperm silk, and live in groups seen commonly and Tent them (ie. the within The skip leaves or caterpillar). pers produce a shelter of silk and hide themselves. by camouflaged larvae Many host food their are The plant. White Admiral of the camouflage themselves with their own excretions, while the later larval instars rely on protective col for defense. The young instars of oration (Papilionidae) droppings, while Some species of also present a pair of foul odors from the osmeteria threatened. glands in are bird brightly Papilionidae fearsome looking will false Swallowtail larvae eyes when threatened. emit resemble later instars the a gland also behind the head which are everted when Other families may have scent body. Monarch other areas of the butterfly larvae are poisonous from consuming The the poisonous milk weed plant. Nymphalidae larvae (Douglas, on the other for attack, have consequently evolved a variety of protec tive adaptations. Some larvae spin webs of called species secrete a genitalia and insects. The bodied larvae are much more at risk colored. spermatophore wasps and other slow moving, soft swallowtails package, the spennatophore, into the ductus bursae, to the is into atrium, and butterflies the eggs of numerous species are by early larvae occurs when the male grasps the paired Yet, parasitized grass to The last two small eggs of moths are usually laid on the underside of leaves, where they have less chance of being hand and reproductive organs, as well as the tures, the fat (Smart, are unpalatable or poisonous they noticed. The that indicates that coloration possessing and unattractive. startling 1975). escape Abdomen tus or spider webs pedicels of themselves appear frequently brightly are colored and possess an armour 1989). covering of "Some gregarious larvae may Methods of All butterfly sharp spines. jerk their bodies in Self Defense of the developmental are at risk of predation stages of by the Occasionally mimic a other unison to a palatable distasteful deter predators. larva may copy or one and avoid predation way" insects and spiders as well as like birds, that reduce the Pupae of predation, closely resembling their environment, are frequently shape, for example, evolved protective risk (Smart, 1975). this reptiles and small mammals. Butterflies have in turn methods by vertebrates like making much like a seed Many butterfly the pod, protected Orange or by Tip by their looks their coloration. and moth species will form a 19 cocoon from dried leaves to pupae. The larvae place often camouflage the will often search out a safe for pupation, consequently pupae are found in inconspicuous places (Smart, 1975). to are most vulnerable when from fly. As flight take Danger Wing survival. and dried, they flowers hunting and Misumena fre them as on spring wings is important in ored upper surface but have a adult "brilliantly col are cryptically colored below so that when they are folded together at rest the insect completely harmonizes with its surroundings" (Smart, 1975). The Lycaenidae have long its to escape with body (Smart, 1975). The Io has quickly flashes to allowing the startles potential predators, The inform monarch would be its Monarch, it a bird has won't do attempted so again. on milk to eat a The Viceroy Monarch's mimicking its colors, quently avoiding taste. are poisonous and moth takes advantage of the by colors noxious because the larvae feeds Once repu concerning the doptera ics insect is species. of colonization of new and flies) and wasps and predacious ants) parasiti- insect (Douglas, for food, mating opportunity, competition dation, and other natural phenomena. range of pre "The dispersal is important, because dis perses are more likely to genes to other colonies, contribute hence their affect the species' course of the evolution"(Douglas, 1989). Also, evolutionists determined that smaller populations of certain species C.periphyle) gene pool. in smaller possess more rare genes (ie. in their This may be particularly important populations, to be carrying these extra alleles "just in case" (Douglas, 1989). Migration Many butterfly to their range of year. Yet, be species a random disperse in manner, with distribution varying from others travel great distances, thousands of miles, en masse or to individually While dispersion is fre on their migration. quently undirected, dictable, directed, year often migrations are and usually pre seasonally determined (Douglas, 1989). Migration may be in response to in day light hours and temperatures, changes unknown population structure of and other mites) Butterfly / moth dispersal is often affected by weather conditions, feeding and scarcity because still much that and 1989). and conse predation. Population Structure There is (ie. what appears butterfly's bright its predators moth escape. predators of Monarch stages of the unpalatable undamaged a pair of owl-like at extra moments to make tation, head" top surface of its wings that it threatening birds. The trick the eyespots on weed. "false the predator to the wings, allowing the butterfly All wings of eyespots at their edges which act as a (tiny flies arthropods doids (parasitic butter that spider wait on plants coloration tails (fungi, climate, microbial pathogens flower (Smart, 1975). Some population structure that effect and viruses), avian predators, parasitic lie in will Factors they to avoid many predators. alight the attract (Douglas, 1989). include: by fritillaries quented the colony's sudden extinction-often seems more bacteria often waits on the for example, and a have soon as the wings flies frequent. The they for or the reason the pupae, and are still haemolymph expanded with can habitats, science" emerge unable suitable apparently of ant art than a Adults first yearly increase and decrease in popula tions, the reason for lack of colonization of al and lepi "The dynam habitats, the season- of adult of food sources or "overcrowding because in the larval (Douglas, 1989). The Monarch (Danus ippus) and Painted occupy large during Lady or stage" plex- (Vanessa cardui) expanses of the temperate zone summer, then environmental changes 20 fall brood to stimulate the They in large migrate migrate south. many butterflies species, like the Dainty Sulfur other individuals (Douglas, 1989). Monarch's over winter in Mexico's Central Transvolcanic mountain range, and migrate as in remain there lengthening pause until daylight hours leave their tree roosts and to females deposit food and three to ten become repro- a "leap Southerly type manner. They ride and full reach Some Monarchs they where over winter remain in repro- These butterflies may join ductively up with the spring migrants in their journey active. A few other include: the Gulf migratory butterflies Fritillary which migrates south breeding butterfly colonies, the in order to establish days feed undulating which can formation fly Kricogonia in huge castalia which few feet an groups of the ground, and the Cloudless Sulfers above water fly in of millions of and the noetropical pierid on their number of host of species go many molts, resulting in a becoming hormones main PTTH they plants until larval instars, before a of metamor JH. When ready to stage, the larvae spins a and ceed to the pupal which fly along Florida's only a pad or girdle which the new chrysalis. larval features it pro silk During and the adult those characteristics few inches legs, a hypopharynx, insects, (three and pairs of one pair of antennae, a pair of maxillae, a the Insecta. during labium They and are the Triassic wings) which believed to have period from the the reveal the pupal stage fourteen days, to unless in dia pause. exoskeleton of head, the includes pupa. a The basic anatomy three segmented body, thorax and abdomen, two pair of wings, three pair of legs (one to antennae and a each body segment), proboscis. suctioning Butterflies differ from moths in that terflies are diurnal, antennae, and frenulum, colored, and numerous natural quently nocturnal, are have feather like the frenulum. butterflies threats, and are prone to have conse evolved unique survival mechanisms. The individuals warmer or more winter. but lack the wing coupling while most moths are brightly not as most colored, have brightly are of some species migrate hospitable climates to over Though many have cycles and habits to of various studied the life butterflies and moths, there are still many unanswered ques Stonefly. The four developmental cycle itself. are lasts from four clubbed northern coast. and moths are suspend (ecdysis) to skin systematically broken down features are formed. Pupation Moths Butterflies to uses larval strips the Summary life larval (unless in dia of oviposition antennae and possess evolve sexually, and the reproduce The larvae size. phosis are of adults Great SouthernWhite (Ascia monuste) cylindrical (Agraulis vanillae) individuals, typify a gene When the newly formed butterfly is ready to emerge, it inhales air, splitting the (Douglas, 1989). possess is eggs on or near suitable The two pupae. It then winds northward and orient Texas, and within a pause) series of negative sun orientation and magnetic compassing. north to the frog" in Florida immature one stage The larvae hatch from between sources. through a Researchers do not know just along how far the spring migrants reach, but believe it is their offspring that continue the journey using the which called metamorphosis, which dia the way. the is Butterflies ductively active. They make the journey north, depositing eggs on milkweed plants north, butterfly next by process is transformed from regulated process. temperatures stimulate the butterflies warmer to a state of reproductive The adult. groups whereas stages of include the egg, larva, pupa, the and tions, and much more research to be pursued. 1 About the Artist The and the Art Work moth depicting the stages of the life cycle, in relative proportions collection of My biggest in the thesis art work exhibited show consisted of: nine paintings of Cecropia working up highlights, the life cycle and their scales of the wing an and a half times it's true size, interactive Director life cycle, anatomy, project illustrating and The painting I most enjoyed was the life I enjoyed the freedom I had in cycle poster. to create a conclusive piece to tie the others paintings were completed utilizing the technique first learned from Vichai Malikul. together, The originally includes transparent media used opaque palette water color Bleed Proof White, and Strathmore board. I a dry tions) to (with Martin's double-sided desired all the bleed able. Because it has to use it to layer lighten tinted of it intricate features on an acrylic top areas and add base, I of was able I tiny details. I recommend the double Strathmore board for this kind because it's color well, take some abuse (erasing that what I couldn't eraser, I exacto knife. was usually nine , life enough to I found (Egg & Larval Instar #2, Larval Instar #3, Larval Instar #4, Larval Instar #5, Pupa, Emergence, Profile of Cecropia and Cecropia were acquired to whom based largely on photo references from The Buffalo Museum I owe many thanks. A of great Science, deal of "detailing" time was spent the paintings and on the computer, I that it chose would show organisms, in a the of the last two micrographs moth and displayed progressively At first only the individ the feather like antenna are pre of and displayed As are of the arrangement of apparent. the (because the striations color up the colors largely they polaroid were shot film. and scales will due to the way detract light). The Micrographs using type 55 make for the is individual The proximity lines that responsible wing the magnification was the gridlike pattern of the becomes sensory The the antenna are visible. the moth. largely tiny hairs closer, micrographs increased, appear Cecropia winter antenna, at looking (sensilla) scales this magnifications. ual segments of are spent several months use and spent several more months sent, then cells electron micrographs scanning elec independent study pro tron microscope as an higher to pick out with an stage paintings how to learning year scales of Larval Instar # 1 Moth ) last The five scanning taken here at RIT. I are of the moth's it remove with a secretary's able piece so I had stages. nine paintings. perspectives of the The first three of media mistakes), and final do this of the rel environment. sided it is strong to first the shooting micrographs Monarch butterfly. often thick enough to resist warping, normally takes The its ject, of the watercolor, and developmental idea variety of poses. I feel it also demonstrates the larvae's natural ability to become camouflaged were invalu with watercolor or applied a color on it. different tex proof white was give viewers an planned scans of using "furry" stages, the to ative sizes of the by color. working-up the started and to paint this the paints utilizing laying down several lay drying time between applica ture of this moth, and the top paints, Dr. applied achieve the As I to and brush technique, ers of paint pieces and reworked creating the layout and poses of the organisms and background foliage. My primary goal was the and migration of the Monarch butterfly. All "finished" them. adult, a model of the adult moth at approxi mately three with more aggressive and returned to electron micro graphs of the antenna and the look creating that in my paintings. I believe my ability to do this increased with each painting. Consequently, I frequently lighting, a natural environment, a five scanning challenge was creatures would a painted poster deepening shadows. imagining how the and They were 22 into Adobe scanned Photoshop adjusted contrast and brightness, up any imperfections in were printed out on where I and cleaned the original. They transparent acetate, mounted and inches from their approximately backing. This was done to facilitate the .25 pas light through the transparency, to achieve a more three dimensional looking sage of image. ""VJ~ v? -.: ss time between Antennae applications. Scales The constructed model of using the Cecropia moth was silk fabric, wire coat hang ers, masking tape, fabric stiffener, feathers, liquid starch, watercolor and acrylic paints, silicon gel, pipe cleaners, artificial gun, and a drift the and ers. veination of base. I first the moth by a glue When the size of a mixture of fabric soaked stiffener and them painted them with areas were sanded proof liquid These beneath the wire frame top of and work and sculpted to shape of encasing the the wing covers Several more layers of and white, in polyurethane applied the down, I painted the wings with a combination of watercolors and the paintings. starch. were placed on adequately structurally sound, I acrylic paint. Rough shaped and cutting, shaping, the four wings, and wings were constructed taping together pieces of wire coat hang Next, I cut pieces of fabric to the approx imate in wood fur, Model much Several layers spray bleed the same manner as of and matte media were to make the paint water/smudge moths chitinous veins. stiffener and starch were applied, with drying resistant. (right Next the forewing forewing and hindwing right hindwing & left with 23 forewing with with silicon night. left hindwing) gel, and allowed to set Then the left right and Finally, were attached Monarch over up sides were an animation which explain the Wire legs were ductive this time. The wing infrastructure the at reinforced with wire and a glue gun. I created the body of it silicon gel, then covered terfly / the moth with with colored pipe cleaners and artificial fur to desired look. Acrylic and water color paints to give the were applied Beads coloration. for the eyes, painted for achieve the body were applied and painted I then the antennae. polyurethane coated a piece of collected, and Lastly, stained and drift legs. The legs and secured using I'd wood drilled holes to insert extensions of the into the holes were cut and the wire were inserted a glue gun. fur painted pipe cleaners and artificial appeared legs to the wire were applied so that the moth interactive exhibit was a projector file on the Monarch butterfly. The three the project are anatomy, life I have of the cycle and migra Museum of I had scales which descriptions was able of the various The section body to access parts. devoted to the life cycle was comprised of photographs of the various life stages, life with processes accompanying (a larva videos of hatching from larval growth, molting, the the egg, metamorphosis from larva to chrysalis, the changing chrysalis, emergence of the mechanism). The was utilized for butterfly, video is and the not feeding my own, attachment the life My interest biology as an to study became apparent in my art Then, last year I had the rare History at Institute. During to pick up the technique, I which The his lessons I fundamentals practiced of for of his the next sev painting insects for the Buffalo Museum of Science. eral months while interning access to an extensive I had at the museum, photo-library the developmental stages of the moth. I decided the Cecropia life cycle, and educational purposes only. subject and New York. which because Cecropia would Cecropia's help to intricate details of plight. the the smallest of the life stages, (the first larval instar is approximately inch long), I hope to increase respect for an because they are native to west I'd also hoped to spark interest By illustrating even be of their complex in this quickly vanishing beauty, and increase viewers to awareness to the and accompanied most photos, and mouse roll-overs allowed viewers definitions Science, the eye and wing Labels strong Natural Smithsonian ern photographed. a often studied opportunity to study water color painting butterflies with Vichai Malikul, from the interesting and electron micrographs of felt have but chose my thesis Monarch butterfly. work as well. The anatomy section illustrated the pri anatomical features of the adult butter mary fly. It included photographs of the specimens, courtesy I contained a complete series of photographs of main parts of the Buffalo Museum of and repro undergraduate, and tion. of always While Director based life moth metamorphosis as nature compelled me to be grasping the branch. The final to narration are several reasons with nature and in and clip patterns. stages of the more natural turkey feathers and topic. included Monarch's migratory There attachments were also devoted to migration contained a video attached with silicon gel, and allowed to set. built into the section one fourth of an appreciation and the these complicated creatures. Considering the dangers and obstacles they face (poisoning by pesticides, parasitization, predation by birds and small animals, traffic, loss of habitat and larval food sources), it is amazing that any survive to moths. Even more tinental migration of Monarch, which become impressive is the adult crosscon- butterflies like the may travel from as far as Canada to over-winter in Mexico's Central Transvolcanic Mountain Range. I plexity am amazed and and appear as beauty intrigued day the com nature that somewhere work makes a does by life, especially lives fragile these. I hope along the way my and some of difference justice. Glossary extend Abdomen The third Ductus bursae The tube that the and posterior segment of insect body. corpus within segments. vertically bursae. Accessory pulsatile organs Tiny organs that help pump haemolymph throughout the body. Ecdysis The shedding Androconial Ecdysone A hormone butterflies Modified scales scales of male that secrete pheromones to stimu late females. head, pair of found on The area leading region near the base the of Chitin A Chorion The is which ovarian layer outer produced follicular Chrysalis The by of the eggshell third segment of the migratory leg. the antenna after segments of Haemolymph The blood cells. instar and of the but of the insect. Epidermal Imaginal discs cells which are undifferentiated and embryonic and allow Compound An eye eye comprised of numer Corneal lens The transparent cuticle of the allata brain that A secrete lobes behind pair of the main part of the bursa copulatrix. vein (costa) The wing the anterior margin of Coxa The insect. Immature Eggs, larvae segment of the Indirect changes hormones. Corpus bursae The Costal adult or pupae, but usually referring to the larvae. ommatidia. Corpora for metamorphosis to take place. Imago The ous ommatidia. the diapausing are type of wing coupling where bristles from the top wing fit into areas of the bottom wing. terfly. the Femur The for fats, that storage sacs Frenulum A insects. the secretions of the metamorphic from the scape. nitrogenous polysaccharide con stituent of the exoskeleton of from the chrysalis. Flagellum The wing. of the caterpillar and the emergence of the adult egg especially large in butterflies. ductus to the dur of molting. hatching Fat bodies The above the mouth. Atrium The opening bursae. Basal organs sensory the old cuticle of molting. Eclosion The the Antennae A the ing into the empties that vein butterfly leg insect forms wing. closest to muscles in The muscles that cause the shape of the thorax and indi control the wings. rectly Instar The larva between molts. Juvenile hormone A hormone produced the corpora allata which supports the characteristics, and by juvenile inhibits the development of adult characteristics. body. Cremaster A structure used by the pupa to a Labrum The silk support pad. Crochets Hooks larval prolegs. Crop The on the plantar surface of the upper paired lip. lip. Mandibles The larva's front pair of chewing mouthparts. area of the foregut behind the Meconium The waste material excreted by the adult after emerging. esophagus. Discal Cell The large cell in the central area Dorsal longitudinal dorsally Medial dinally of the wing. that extend Labia The larva's lower muscles and The muscles lengthwise between segments. Dorsal-ventral muscles The muscles that vein The wing vein that runs longitu between the radius and cubitus. Mesothorax The second segment of the tho rax. Metathorax The third segment of the thorax. Micropyle The opening on the top of the egg. >6 the wing membrane that carry the tracheae and where sperm penetrate. Molting The is produced, and the old one is insects the unit of compound eye. Oviposition The Pedicel The process of egg laying. the antenna that act segment of "wrist" 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 of the adult butterfly. The feeding Prothoracic glands that ecdysone, the molting produce Prothorax The first Pupa The glands of the segment of between larva stage Rhabdom The light-sensitive tube brain hormone. the thorax. and adult. Radius Vein The wing vein that dinally between the subcosta and longitu runs the media. 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 cells, which often larger Sensory belong to sense organs. Spermatophore The sperm containing pack age of the male. Spiracles The air openings to the tracheae. The wing vein that Subcosta tudinally behind the costa vein. vein Tarsus The final Thorax The femur segment of leg runs longi the walking second segment of Tibia The fourth segment leg. the body. between the and tarsus. Tracheae The internal air transport tubes of insects. Trochanter The branches. membrane egg that membrane, simple eye. Ommatidium An individual like of the of the antennae. Ocelli A nerve Vitelline shed. scaleless portion of the flagel- Nudum The lum by process which a new cuticle segment of the coxa and the femur. Veins The thick walled the leg supporting between tubes of The internal originates and membrane from the egg lies beneath the cell chorion. 27 Bibliography Arnett Jacques, R. L. Simon &Schuster's Guide Inc., 1981. and Douglas, M. M. The Lives of to Insects. Butterflies. Canada: The Wiley &Sons., 1989, 20-49, New York: Simon & Jacques University of Michigan Press and John 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 London. 1975, 18-21, 34-40. of the Butterfly World. Salamander Books Ltd.