EVERYTHING YOU EVER WANfEDTO KNOW ABOUT BU'I'IERFLIES

& (i9 INTRODUCTION

Over the past several years I have presented numer­ous programs on butterflies for conservation organi­zations, garden clubs, classrooms, and summer camps. Always I have been impressed with the public's tre­mendous interest and concern for what I consider the paragon of the insect world, butterflies. But I also have been alarmed at the widespread lack of.knowledge re­garding even the basics of butterfly biology. It seemed to me that a book that would be inexpensive, easy to read, but with sufficient technical information to sat­isfy even the professional entomologist, was in order. "Everything You Ever Wanted To Know About Butter­flies: 100 + Questions and Answers" is my response. Although much of the information contained within is derived from previously published sources, much is documented from both my lifelong pursuit of butter­flies as a professional entomologist and from my burn­ing passion as a butterfly hobbyist. I hope this unique synthesis proves rewarding to all interested in broad­ening their knowledge of the fascinating world of but­terflies.

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TABLE OF CONTENTS

INTRODUCTION ............................................... 4

TAXONOMY: NAMES AND CLASSIFICATION ............................................... 7

1. What is the origin of the word "butterfly?" 2. How are butterflies classified? 3. What is the difference between a butterfly,

a skipper, and a moth? 4. What do you call people who study butter­

flies? 5. What are the nearest relatives of the

lepidoptera?

FORM AND FUNCTION ................................. 11

6. What is responsible for the colors on the wings of butterflies?

7. Do male and female butterflies differ in color? 8. Do male and female butterflies differ in size? 9. Can butterflies fly without their scales? 10. Can butterflies repair damaged wings? 11. What is the significance of the large eyespots

on the wings of so many butterflies? 12. Do the long tails of swallowtail butterflies and

the short hairlike tails of hairstreak butterflies serve any function?

13. Are there any wingless butterflies? 14. Do all butterflies within the same species look

alike? 15. Which is the largest butterfly in the world? 16. Which is the smallest butterfly in the world? 17. Why do some butterflies seem to have only 2

pairs of legs (4 legs) rather than 3 pairs (6 legs)?

18. How many legs do caterpillars have? 19. How can a caterpillar's skin stretch so much

during its life? 20. How do butterflies feed? 21. How do butterflies see? 22. How do butterflies breathe? 23. Can butterflies make sounds? 24. Can butterflies hear? 25. Can butterflies smell? 26. Can butterflies taste? 27. Do :6utterflies have brains?

28. Do all caterpillars sting? 29. Can a caterpillar bite a human? 30. Can the "horns" of the orange dog caterpillar

and the "horns" of the tomato and tobacco horn worms sting?

METABOLISM: HOW THE BODY WORKS .............................. 19

31. Do butterflies grow? 32. Do butterflies have blood? 33. Do butterflies excrete? 34. Why do butterflies so often perch in the sun? 35. Do butterflies sleep? 36. Do caterpillars feed only on leaves? 37. What do butterflies eat? 38. Do butterflies have to drink water?

LIFE CYCLE: METAMORPHOSIS .................. 23

39. What is a typical butterfly life cycle? 40. What is metamorphosis? 41. What is a cocoon? 42. What is a chrysalis? 43. How long do butterflies live? 44. Can butterfly eggs be seen with the naked eye? 45. How many eggs do butterflies lay? 46. Do all butterflies lay eggs? 47. Do female butterflies watch over their eggs? 48. What is the incubation time for an egg? 49. What is the life span of a caterpillar? 50. What is the life span of a chrysalis? 51. How many generations a year does a butter­

fly complete? 52. How do butterflies survive winter? 53. Should caterpillars be called "worms?"

REPRODUCTION .............................................. 27

54. How do male and female butterflies locate one another?

55. How do butterflies copulate? 56. Do male butterflies have a penis? 57. Do female butterflies have a vagina? 58 .. Why do certain butterflies lay eggs only on

certataplants?

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&,fi9 BEHAVIOR .......................................................... 29

59. How far can butterflies fly? 60. How high can butterflies fly? 61. How fast do butterflies fly? 62. What do butterflies do during rains or other

bad weather? 63. Can butterflies fly at night? 64. Why do butterflies sometimes alight on

people? 65. Are any butterflies territorial? 66. Do any American butterflies migrate? 67. Why does the monarch butterfly migrate? 68. How does the monarch butterfly navigate? 69. Where do monarch butterflies go during

their migration? 70. How do scientists track a migrating

butterfly?

ECOLOGY: INTERRELATIONSHIPS, GEOGRAPHY, AND CONSERVATION ........ 32

71. What p.urpose do butterflies serve? 72. How many different species of butterflies

and skippers are there in the world? 73. How many different species of butterflies

and skippers are there in the United States and Canada?

74. How many different species of butterflies and skippers are there within the various States and provinces of North America?

75. Where in the world are most species of but­terflies?

76. Where in the world are the fewest species of butterflies?

77. Are there any endangered or threatened spe-cies of butterflies within the United States?

78. Which are the best months to see butterflies? 79. What is a "Fourth of July Butterfly Count?" 80. What is the best time of day to observe

butterflies? 81. Are butterflies very sensitive to environmen-

tal disturbances? 82. What is the rarest butterfly in the world? 83. Are butterflies marketed commercially? 84. Are there any fossil butterflies? 85. What are the chief enemies of butterflies?

87. Are some butterflies distasteful to predators? 88. Do some caterpillars produce honeydew as

do aphids, and are these caterpillars attended by ants?

BUTTERFLY GARDENS ................................... 37

89. What is a butterfly garden? 90. Are butterfly gardens difficult to install? 91. Are butterfly gardens difficult to maintain?. 92. What plants do butterflies prefer? 93. What is the difference between a "food plant"

and a "host plant?" 94. Don't caterpillars eventually destroy a butter-

fly garden? 95. What is a "butterfly house?" 96. What is a "butterfly hotel?" 97. What is a "butterfly log pile?" 98. What are butterfly feeders?

RESEARCH AND COLLECTING .................. 39

99. Why do people collect butterflies? 100. Can a person attract butterflies to a spe­

cific location? 101. What kind of equipment do you need to

collect and preserve butterflies? 102. How long can butterflies be preserved? 103. What are trap nets? 104. Are there any butterfly clubs or societies

in North America?

AESTHETICS AND SYMBOLISM ................. 42

105. What is the most beautiful butterfly in the world?

106. What is the most beautiful butterfly in North America?

107. Is the symbol of the butterfly ever used out­side of entomology?

108. What is the significance of the humanoid butterfly motif so common in pre-Hispanic American art?

109. Why are butterflies sometimes referred to as "dancing flowers?"

GLOSSARY .......................................................... 44

86. Can butterflies trallsmit any disease to humans?' BIBLIOGRAPHY ................................................ 50

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TAXONOMY: NAMES AND CLASSIFICATION

1. What is the origin of the word "butterfly?"

No one is really sure. A commonly held idea, how­ever, is that the word's origin lies with the sulphur but­terflies found in abundance throughout England and most of continental Europe. Since these bright yellow butterflies begin flying during the spring months when cattle are birthing their new calves--and fresh raw milk is available for butter churning -- the legend states that the color of the flying insects reminded people of the yellow butter. That association became fixed in the name "butter-colored fly," later "butterfly." However, another legend states that the name originally was "flutterby," referring to the insects' manner of flight. The word was later revised to its present form, butterfly.

Carl Masthay (1991) scanned numerous non-English dictionaries for the word butterfly. The following is a sample: FARASHA (Arabic), PAPALOTL (Aztec), HUDLE or HUTLER (Chinese), PAPILLON (French), SCHMETTERLING (German), PULELEHUA (Hawai­ian), PARPAR (Hebrew), FARFALLA (Italian), CHO (Japanese), PAPILIO (Latin), PEPEN (Mayan), BORBOLETA (Portuguese), BABOCHKA (Russian), and MARIPOSA (Spanish).

2. How are butterflies classified? (See Figure I on page 8. Note: illustrations are not to scale. Numbers refer to relative size of most family members: l=smallest, S=largest.)

Since butterflies are biological entities, they are named according to the BINOMIAL SYSTEM OF NO­MENCLATURE, and classified into standardized bio­logical categories--all based on Latin names. Below is a commonly recognized classification (with annota­tions) for butterflies and skippers (moths are omit­ted). (Sources: Lewis, 1985;.Miller, 1992; Smart, 1989).

KINGDOM: Animalia

PHYLUM: Arthropoda (invertebrates with body re­gions; jointed appendages; exoskeleton)

CLASS: Insecta (bodies divided into 3 parts; 3 pairs of legs; 1 pair antennae; metamorphosis; ter­restrial and aquatic but not marine)

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ORDER: Lepidoptera (scaly wings--butterflies, skippers, moths)

SUBORDER: Rhopalocera (clubbed antennae­butterflies and skippers)

SUPERFAMILY: Papilionoidea (true cl~bbed antennae--butterflies)

FAMILY: Papilionidae--swallowtails, bird wings, a polios {parnassians) FAMILY: Pieridae--whites, sulphurs, jezebels, orangetips, marbles, brimstones FAMILY: Lycaenidae--gossamer wings (harvesters, coppers, hairstreaks, blues, elfins) FAMILY: Riodinidae or Nemeobiidae-­metalmarks, judies FAMILY: Libytheidae--snouts or beaks FAMILY: Heliconiidae--longwings, heli­cons, (gulf fritillary) FAMILY: Acraeidae--acraeas FAMILY: Nymphalidae--brushfoots (frit­illaries, checkerspots, crescents, anglewings, admirals, vanessas, maps, buckeyes, crackers, etc.) FAMILY: Apaturidae--emperors, leaf­wings, rajahs FAMILY: Amathusiidae--fauns, saturns, palm-kings, duffers, jungle- glories, jungle­queens, etc. FAMILY: Morphidae--morphos FAMILY: Brassolidae--owls and relatives FAMILY: Satyridae--browns, satyrs, pearly eyes, ringlets, alpines, arctics, etc. FAMILY: Danaidae--milkweed butterflies (monarch, queens, tigers, crows, etc.) FAMILY: Ithomiidae--ithomiids, clear­wings

SUPERFAMILY: Hesperioidea (antennae with hooked ends, often an apiculus--skip­pers)

FAMILY: Hesperiidae--skippers

SUBORDER: Heterocera (antennae straight and tapered, often feathery; pupa within a silken cocoon, or naked and concealed under soil--moths).

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Papilionidae (4-5) Pieridae ( 1-3) Lycaenidae (1-2)

Acraeidae (2-3) Libytheidae (2) Heliconiidae (3)

Figure 1.

Nyrnphalidae (2-4) Satyridae (1-3) Riodinidae (1-2) Apaturidae (2-3)

Brassolidae (3-5) Amathusiidae (3-4) Morphidae (3-5)

Danaidae (2-3) Ithomiidae (2-3) Hesperiidae (1-3) '

3. What is the difference between a butterfly, a skipper, and a moth? (Figure 2)

There is no one character that distinguishes but-

terflies, moths, and skippers, all members of the order Lepidoptera, "scaly-winged insects." The chart below shows the major comparisons:

Butterflies

A. Antennae clubbed.

B. Wings not hooked together.

C.. Wings held vertically over back when at rest.

D. Mainly diurnal, some crep­uscular.

E. Many brightly colored.

F. Body slender, smooth.

G. Pupa termed chrysalis, sus­pended or with a basal attach­ment and girdle (all of silk); no cocoon.·

A

Skippers

A. Antennae clubbed with hooked ends, often with an apiculus.

B. Wings not hooked together.

C. Wings held vertically, horizon­tally, or split when at rest.

D. Mainly diurnal.

E. Most dull-colored.

F. Body thick, heads large.

G. Pupa with basal attachment and girdle (silk) and often housed within a loosely constructed silken cocoon.

B

Figure 2. A typical butterfly, skipper, and moth. A. Butterfly: cloudless sulphur (Phoebis sennae)

Moths

A. Antennae straight or feathery.

B. Wings hooked together.

C. Wings held horizontally or chevron-like with hindwings covered when at rest.

D. Mainly nocturnal.

E. Most dull-colored.

F. Body thick and "furry."

G. Pupa often within a silken cocoon, or if naked, within soil.

C

B. Skipper: golden-banded skipper (Autochoton cellus) C. Moth: Columbia silk moth (Hyalopl10ra columbia)

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&fi9 4. What do you call people who study butterflies?

Since butterflies (as well as skippers and moths) belong to the insect order LEPIDOPTERA (Greek for "scaly-wings"), the science that deals with their research is termed LEPIDOPTEROLOGY, and the people who do the research, LEPIDOPTERISTS. Incidentally, lepi­dopterology is a branch of ENTOMOLOGY (Greek for "study of insects"), and ENTOMOLOGISTS are insect scientists.

5. What are the nearest relatives of the lepidoptera?

Insects belonging to the ORDER TRICOPTERA , commonly called CADDISFLIES, are considered the closest relatives of the lepidoptera. These are delicate­winged insects that are encountered usually only at lights at night. Their larvae are caterpillar-like and live in self-constructed enclosures within clear fresh water streams. Members of the primitive moth family Micropterygidae bear mouth parts that are similar to those in the larvae of the tricoptera--hence the evolu­tionary relationship.

ab

sp

hw

th

oc

Figure 3. Anatomy of a generalized butterfly. ab (abdomen), an (antenna), ce (compound eye), fw (forewing), he (head), hw (hindwing), le (leg), oc (ocellus or eye spot), pr (proboscis), sp (spiracle), ta (tarsus), th (thorax), wv (wing vein)

FORM AND FUNCTION

6. What is responsible for the colors on the wings of butterflies? (Figures 3, 4)

SCALES give the Order Lepidoptera its distinctive­ness. Scales are tiny, dust-like particles that under microscopic examination appear as thin, flat overlap­ping structrues, often with various surface ridges and grooves. All are outgrowths of the body wall, and are attached to the clear underlying wing membrane and veins by a short stalk. Scales impart color to the wings of the lepidoptera. Some colors are produced by pig­ments, chiefly melanins, that are either ingested or else biochemically synthesized; this type of colora­tion is termed CHEMICAL or PIGMENTAL COLORA­TION. Other colors, termed STRUCTURAL COLORA­TION, are generated by the physical configuration of the scales themselves interacting with light. For ex­ample, the metallic blues of the morpho butterflies and the iridescence of many other tropical species are pro­duced when light is diffracted or scrambled in specific ways by specific surface elements of specific scales (the actual color of the scale is usually grey or brown). Col­ors are important in: (a) mate selection (COURTSHIP), (b) concealment by disguise (CAMOUFLAGE or CRYPTICNESS), (c) creating a conspicuousness that advertises obnoxiousness (APOSEMATIC COLOR­ING), (d) imitation of other obnoxious species (MIM­ICRY), (e) surprising and frightening predators,

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thereby creating a moment for escape (FLASHING), (f) heat regulation by absorption or reflection of sun­light (THERMOREGULATION-see question 34). Some scales are responsible for producing sex-attracting aro­mas (aphrodisiacs) involved in courtship. These spe­cialized scales are termed ANDROCONIA.

7. Do male and female butterflies differ in color?

Male and female butterflies often differ in form. SEXUAL DIMORPHISM. This sexual dimorphism can be dramatic. For example, the gigantic birdwing but­terflies (Ornithoptera) of the Pacific islands show ex­treme sexual dimorphism: males are usually brightly colored with metallic hues whereas females are usu­ally dull brown and nearly twice as large. Within North America there are also striking examples of sexual di­morphism: the female diana butterfly (Speyeria diana)

is pale blue and black whereas the male is a striking orange and brown; the male tiger swallowtail (Pter­

ourus glaucus)is yellow and black whereas females are similar or else black and blue (considered a form of MELANISM); and some sulphur butterflies (usuai'Iy yellow and black) have females that are white and black (considered a form of ALBINISM). Scientists theorize that because butterflies are very visual crea­tures, and their ability to spot one another from a dis­tance i~ crucial, sexual dimorphism is a strategy that

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Figure 4. Typical scales of the lepidoptera.

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2A

Costal

~ C \ :c,, I tr 1'0 \.'i; \~ \(1) \ /~ \

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'" ..:

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Inner or Anal Margin

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Inner

C:

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E .0 :, Cl)

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'" ..: C:

"' .... 'O I!)

E .0 :, Cl)

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'" ..: C:

"'

/ /

C:

"' .... 'O I!)

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0 ~

,,,, /w '" ,:

I ~ .... 0,

I '-"' / :;:

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Costal Margin

C:

"' .... 'O I!) :;:

I c: "' ....

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Figure 5. Wings of butterflies. A. Venation in a swallowtail (Papilionidae) B. Venation in a brushfoot (Nymphalidae)

'" 0

- C: ,,, .... _., 0,

(I)'" .... "' o:;:

C. Major wing areas in a generalized butterfly

'" Cl, .., :, 0

'" 0

- C: ,,, .... _., 0,

(I)'" .... "' Cl :;:

C = Costa vein, Sc = Subcosta vein, R = Radius vein, M = Media vein, Cu = Cubitus vein, A = Anal vein, hv = humeral vein (secondary vein), D = Discal cell

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facilitates mate recognition. By contrast, moths com­municate with each other primarily through odors (pheromones). Therefore, their colors tend to be subtle, and there is little or no sexual dimorphism except in size.

8. Do male and female butterflies differ in size?

Female butterflies are invariably larger than males­as with most insects. Females are the egg bearers. Large bodies, therefore, are a distinct advantage. The func­tion of the male usually is solely for sperm production and transfer. And since mating occurs usually only during a brief period of time immediately after emer­gence, males live only a relatively brief time. As such, their bodies do not have to store significant quantities of fat for a lengthy metabolism and extended life span.

9. Can butterflies fly without their scales? (Figure 4)

Contrary to popular belief, butterflies can fly with­out their scales. In fact, there are species of butterflies that have practically no scales on their wings (many species in the family lthomiidae and some within the family Satyridae) . The myth probably is based on the observation that if the scales are accidentally rubbed from the wings of a butterfly during handling, the butterfly usually does not fly very well. In reality what happened is that in handling the specimen its wing membranes and veins (or perhaps even some other body part) were damaged, and that was responsible for the problem with flight. It is important to remem­ber, that only the ventral (undersurface) of the wings should be touched when handling a butterfly (the ven­tral surface has fewer scales than the dorsal or uppersurface). With caution, butterflies can be handled without causing any damage to the individual. Al­though butterflies can fly without their scales, scales do contribute to the aerodynamics of flight. Scales may also help butterflies resist predators: scales are only loosely attached so that they can easily break free dur­ing an attack or when a butterfly collides with a spider's web. In a sense, scales make butterflies, and to an even greater degree moths, "slippery," and therefore, hard to catch.

10. Can butterflies repair damaged wings? (Figure 5)

No. The veins, tissues, and scales of wings of the lepidoptera are part of the insect's EXOSKELETON. As such, these tissues and organs are of dead, tough ma­terial called CHITIN or SCLEROTIN--a colorless nitrog­enous polysaccharide intermediate between proteins and carbohydrates. As such, damaged parts cannot be regenerated or repaired.

11. What is the significance of the large eyespots on so many butterflies? (Figure 3)

Scientists theorize that eyespots, often called OCELLI, function as warning signals to would-be predators. Supposedly, the sudden flash of large eyes is enough to create a moment of surprise, and there­fore, art instant to escape. The owl butterflies (genus Caligo) of the American tropics are noted for their large, intricately formed ocelli on their underwings.

12. Do the long tails of swallowtail butterflies and the short hairlike tails of hairstreak butterflies serve any function? (Figure 1)

Tails are wing appendages, and are examples of BACK TO FRONT MIMICRY. Supposedly, these struc­tures, commonly found within the families Papilionidae and Lycaenidae, immitate the head and antennae of the insect. By having a dummy set of head and anten­nae on their posterior, these butterflies effectively in­crease their survival rate since predators have a good chance of selecting the wrong end to attack. The lycaenid butterflies even increase this illusion: they call additional attention to the dispensable parts by rub­bing their two hindwings together.

13. Are there any wingless butterflies?

No. All butterflies possess two pairs of wings: l pair of forewings, 1 pair of hindwings. There are, how­ever, several species of moths in which the female is wingless or else has greatly reduced wings, and there­fore, incapable of flight. For example, the silkworm moth, Bombyx mori, can be efficiently cultivated only because of flightless females.

A

B

pr

ab

Figure 6. Sample butterfly larvae (caterpillars) A. Giant swallowtail or "orange dog" (Heraclides cresplwntes) B. Monarch (Danaus plexippus)

ab (abdomen), an (antenna). fi (filament), he (head), le (leg), oc (ocellus), os (osmeterium), pa (pal pus), pr (prolegs), sp (spiracle), th (thorax)

14. Do all butterflies within the same species look alike?

Not necessarily. Many species can be divided into geographical races or subspecies that differ significantly from one another. And just as with any biological spe­cies, individuals within any given species of butterfly disptay a range of variation that follows a typical BELL­SHAPED CURVE or NORMAL DlSTRIBUTION CURVE. Then there are seasonal forms, that is vari­ants that are related to climate (for example, summer vs. winter, wet season vs. dry season). Within some species females seem to have several color variations. The tiger swallowtail, for example, has two female forms: a yellow and black form very similar to the male, and a black or melanistic form. Also, many sulphurs

have female forms that often are nearly white. And finally, there are on occasion aberrant forms such as giants, dwarfs, and gynandromorphs (forms with one side exhibiting male coloration, the other, female).

15. Which is the largest butterfly in the world?

The birdwing butterflies, Ornitlwpcm and Troidcs, of tropical Australia, southeast Asia, and tropical Af­rica are extremely large. Wingspans of females can be 9-11 inches (229-279 mm), males 5-7 inches (127-179 mm). Queen Alexandra's birdwing (Onzitlwptcra a/cxandrac) from the island of New Guinea is generally quoted as the largest species.

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16. Which is the smallest butterfly in the World?

The so-called pygmy blues (Brephidium) of Asia, Africa, and the United States are very tiny, ranging between 3/8-3/4 inches (10-19 mm).

17. Why do some butterflies seem to have only 2 pairs of legs (4 legs) rather than 3 pairs of legs (6 legs)?

Because their first pair of legs is greatly reduced in size, butterflies within the family Nymphalidae are called the BRUSHFOOTED BUTTERFLIES. In fact, members of all families except the Papilionidae, Pieridae, and Hesperiidae have only 2 pairs of func­tional walking legs (however, within the families Libytheidae, and Riodinidae, only males have a re­duced number--females retain the full 3 pairs). Al­though the first pair may not be used in locomotion, they are useful in chemical perception (olfaction and taste).

18. How many legs do caterpillars have? (Figure 6)

The larvae of butterflies have three pairs (6) of TRUE LEGS on the thorax, four pairs (8) of PROLEGS in the middle of the abdomen, and one pair (2) of PRO­LEGS at the extreme posterior of the abdomen. True legs are segmented and equipped with claw-like struc­tures that are capable of shallowly piercing a perch; prolegs are unsegmented and function more like suc­tions. Some species of moths lack various prolegs. These caterpillars loop along, and are commonly called "inch worms," "measuring worms," or "loopers."

19. How can a caterpillar's skin stretch so much during its life?

Although all skin is somewhat elastic, caterpillars are able to achieve their remarkable growth because they replace their skin (cuticle) three or four times dur­ing their development. The process, called MOLTING or ECDYSIS, allows the caterpillar to replace its skin when additional stretching is no longer possible. After each molt, the caterpillar renews its growth.

20. How do butterflies feed? (Figures 3, 6, 7,10)

The lepidoptera feed through a tube-like structure technically termed a PROBOSCIS, and commonly, a TONGUE. The structure can be coiled and uncoiled in much the same manner as a party whistle. The probos­cis actually consists of two elongated, extensile fila­ments (the MAXILLAE of mandibulate insects) joined with a groove in between. Liquified foodstuffs are si­phoned up within the groove by muscles within the PHARYNX. During the long history of evolution, but­terflies and flowers have coevolved so that specific plants and specific butterflies often have become codependent upon each other (plants for pollination, butterflies for nourishment). For example, those spe­cies of butterflies that feed predominantly on flowers with long tubular nectaries, have long proboscises; con­versely, those butterflies that feed on flowers with only short nectaries, have short proboscises.

21. How do butterflies see? ( Figures 3, 7)

All lepidoptera see through a pair of COMPOUND EYES occupying most of the head. Each eye consists of several thousand (approximately 6,000) lenses called OMMATIDIA (singular, OMMATIDIUM). The image consists of a mosaic of patterns lacking in detail. Cater­pillars lack compound eyes. Instead, they have 12 small, simple eyes termed OCELLI (singular, OCELLUS). These are arranged in two semicircles of six, one on each side of the head. Ocelli probably function only in the detection of light and darkness.

22. How do butterflies breathe? (Figures 3, 6 )

As with most other animals, butterflies must ac­quire oxygen and dispose of carbon dioxide to carry out their respiratory needs. All insects have an intri­cate system of tubes called the 'TRACHEAL SYSTEM. The system consists of the tubes proper (TRACHEAE) and openings to the outside of the body called SPI­RACLES. The tracheae branch and extend throughout the body eventually terminating within the tissues. Spi­racles are typically located on the lateral portions of the thorax (one pair on the first thoracic segment) and the abdomen ( eight pairs). This arrangement is the same in butterfly larvae.

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Figure 7. Head details of a typical butterfly.

A. Antennae, compound eyes, palps, and proboscis B. Cross section of a proboscis C. Photomicrograph of a compound eye

Insects have relatively low respiratory rates; hence, they require little oxygen. For this reason butterflies and caterpillars can be housed safely in containers with little ventilation for considerable periods of time. And the cooler the temperature, the lower the rate of me­tabolism. To temporarily store an insect, put it into a container with a moist paper towel (to prevent dehy­dration) and place in a refrigerator.

23. Can butterflies make sounds? (Figure 5)

Typically, no, but members of the genus Hamadr­yas, commonly called CRACKERS or CALICOES and found throughout the American tropics, the Florida Keys, and the lower Rio Grande Valley of Texas, can produce a clicking or cracking sound that is audible to humans from distances up to 100-150 feet (30-50 meters). Although no one is positive as to the mecha­nism involved, recent research indicates that the swol­len base of the SUBCOSTA VEIN in the forewings is

ma

C

B

an (antenna), ce (compound eye), fc (food channel), ma (maxilla), om (ommatidium), pa (palpus) pr (proboscis)

involved. Apparently, the strong venation allows sound production when the insect claps its wings. The sound seems to be involved with territorial displays.

24. Can butterflies hear?

Typically, no. Unlike in many other groups of in­sects, the TYMPANIC MEMBRANE (the sound-sens­ing organ) is only poorly developed or even absent in butterflies. (Many species of moths, however, do have well developed tympanic membranes.) Butterflies do seem, however, to be sensitive to aerial vibrations. These vibrations are probably registered by sensory organs on the antennae and body surfaces.

25. Can butterflies smell? (Figures 3, 6, 7, 8, 9)

Yes. The two ANTENNAE of butterflies are equipped with many sensory structures for detecting chemicals. Under the microscope, an antenna appears

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as a sponge, full of holes. Olfaction is important in lo­cating food, mates, rivals, and avoiding predators.

Caterpillars, too, have one pair of antennae. These, however, are short and are on either side of the MAN­DIBLES. As in butterflies, the antennae in caterpillars are involved with olfaction and taste.

26. Can butterflies taste? (Figures 3, 6, 9)

Yes. The six feet of butterflies are equipped with many sensory structures for detecting various chemi­cal stimuli. This is extremely important to female but­terflies, which must accurately locate specific host plants for egg deposition. Many species of butterflies have extraordinary powers of discrimination. Taste in caterpillars lies with the accessory mouth parts called LABIAL PALPI and MAXILLARY PALPI.

Caterpillars, too, have extraordinary powers of dis­crimination. This is important since many species de­pend on the sequestering of secondary plant substances for specific metabolic processes, some of which infer protection from predation. (See question 58.)

27. Do butterflies have brains?

Strictly speaking, no. As with other insects, butter­flies possess a large GANGLION (nerve bundle) in the head. Commonly called a brain, this nerve center regulates most of the nervous system through the pro­duction of electro-chemical signals and hormones. This simple "brain" is connected to other smaller ganglia and a lengthy NERVE CORD that runs throughout the mid-ventral (underside) of the body. Most butterfly be­havior is genetically programmed through DNA. How­ever, some butterflies can learn simple behaviors. For example, many sped.es do learn the direction and po­sition of favorite nectar sources and host plants, favor­ite flyways, and favorite roosting places.

28. Do all caterpillars sting ?

No. Very few species possess URTICATING or URTICARIAL HAIRS. In Louisiana, only six species are capable of inflicting humans. These are: buck moth caterpillar (Hemileuca maia), saddleback caterpillar (Sibine stinmlae), io moth caterpillar (Automeris io), puss

J

caterpillar (Megalopyge opercularis), crinkled flannel moth caterpillar (Lagoa crispata), and hag moth cater­pillar (Phobetron pithecium). One additional species, the whitemarked tussock moth caterpillar (Orgyia leucostigma) is known to have produced an allergic re­action in a handful of sensitive persons. All species are the larvae of moths; no butterfly larva in North America is capable of stinging.

29. Can a caterpillar bite a human? (Figure 9)

Theoretically yes, but in actual practice, no. The MANDIBLES of a caterpillar are designed for cutting and chewing. However, the feeding response is usu­ally initiated only by chemical recognition of a host plant. And if for some reason a caterpillar did make a mistake and try to nibble human tissue, the result would be virtually imperceptible to us.

30. Can the "horns" of the orange dog caterpillar and the "horns" of the tomato and tobacco horn worms sting? (Figure 6)

No. These structures are bluffs, and are used to deter predators. The horns of the orange dog caterpil­lar (the larva of the giant swallowtail), as well as the horns of all the swallowtail butterflies are called OS­METERIA (singular, OSMETERIUM). They are housed in sacs within the thorax. When annoyed, the caterpil­lar everts these while at the same time releasing a pun­gent cheese-like odor. The display usually is sufficient to call off most attacks by vertebrates. The horns on the tomato and tobacco horn worms (larvae of sphinx moths) are part of the caterpillars exoskeleton. As such, they are permanent external appendages, and they can­not deliver any toxins. {In fact, most caterpillars with horns and spines are harmless to humans.)

1'7

Figure 8. Characteristic antennae of the lepidoptera.

A. Butterflies B. Skippers­

ap (apiculus) C. Moths

A

ep

se

B

/

an ma

A

B

18

ap

I? l 1

e

C

Figure 9. Head of a typical caterpillar.

A. Front View B. Ventral View C. Lateral View

an (antenna), ar (adfrontal region), cl (clypeus), ep (epicranium), fr (frons), la (labium), Ip (labial palpus), Ir (labrum), ma (mandible), me (membrane), mp (maxillary palpus), mx (maxilla), oc (ocellus), se (seta), sp (spinneret)

C

'!"'

METABOLISM: HOW THE BODY WORKS

31. Do butterflies grow?

No. Once a butterfly expands and dries after emerg­ing from its chrysalis, a process called ECLOSION, it is an adult, and hence, fully grown. Within each species, however, there is some variation in size--just as with any biological species. Caterpillars, on the other hand, do molt periodically and grow, often increasing their size from hatching by many fold.

Figure 10. Internal anatomy of the lepidoptera. (After Howe, 1975, and used with permission.)

HEAD

thoracic chamber of heart \ THORAX

32. Do butterflies have blood? (Figure 10)

Yes, but it is called HAEMOLYMPH, and it is clear to pale yellow. The blood is pumped by a DORSAL BLOOD VESSEL located along the dorsal (back) sur­face of the body, extending from the head, through the thorax, and through the abdomen (a similar arrange­ment is found in caterpillars). The actual pump or "heart" of this tubular organ is nothing more than the abdominal section of this blood vessel. Consisting of

A. Butterfly (adult) B. Caterpillar (larva)

heart I I

ABDOMEN

septum \

I I

os ti urn crop I

rectum \

closed end of heart

I I

proboscis

HEAD esophagus

\ \

mandibl~ spinneret

crop I I I

THORAX salivarv (mandibular) glands

I I I

\

I valve I \

midgut

A

source of peritrophic meyibrane

I

'

I

I I

I silk(labial) gland

B

\ source of peritrophic membrane

ABDOMEN

heart ostium I

I \ \ I

I I

uncus I I

~- aedeagus

\ I \ I

Malpighian tubules(6)

clasper hindgut

Malpighian bladder I I I

/ hindgut

\

rectum I

I muscle

Malpighian tubule

I I

I I

several chambers, the heart pulses blood forward. The anterior part of the vessel acts as an AORTA, or the section through which blood flows out into the body cavity or HAEMOCOEL. Since the blood does not transport oxygen as it does in vertebrates, no red blood cells are present, and hence the fluid is relatively color­less. The main function of the blood is to transport nu­trients and waste products. All organs are bathed di­rectly by the blood. The blood flows back into the dor­sal blood vessel through tiny openings called OSTIA (singular, OSTIUM). This type of system is called an OPEN CIRCULATORY SYSTEM. It is common in small animals like invertebrates. On the other hand, verte­brates seem to require a more efficient type termed a CLOSED CIRCULATORY SYSTEM.

33. Do butterflies excrete? (Figure 10)

Yes. Undigested food substances and by-products of metabolism are voided from an ANUS located at the extreme posterior of a butterfly. The lepidoptera do not have kidneys. The excretory system consists of six coil­like tubes termed MALPIGHIAN TUBULES. These are attached to the alimentary canal at the junction of the MIDGUT and the HINDGUT. The tubules float freely within the body cavity (HAEMOCOEL). Butterflies, as with all insects, have an OPEN CIRCULATORY SYS­TEM in which blood (HAEMOLYMPH) is not confined within arteries and veins, but pools within the central body cavity. The blood bathes all tissues directly. Waste products, chiefly solid crystals of URIC ACID (rather than liquid urea as in mammals), are absorbed from the blood, and then transported to the HINDGUT. There the crystals are mixed with the solid wastes of digestion (see question 37), and eliminated through the ANUS. Scientists theorize that the production of solid uric acid rather than liquid urea is a strategy for con­serving water, and hence, reducing dehydration. (Some pigments in the wings of butterflies within the family Pieridae are related to uric acid. The storage of these chemicals in the wings may be another strategy for eliminating wastes while at the same time producing characteristic and useful coloration.)

Caterpillars, too, have Malpighian tubules. But be­cause most caterpillars feed on solid plant material, their excrement, technically termed FRASS, is larger (pellet-size) and produced more frequently (see ques­tion 37).

34. Why do butterflies so often perch in the sun?

Sunshine is crucial to most butterflies. (Some ex­ceptions include those species that inhabit the tropical rainforests where ambient temperatures are always warm. Many of these species fly only under condi­tions of low light as on cloudy days or at dawn or dusk­a CREPUSCULAR habit.) Sunlight is a butterfly's elixir of life. To quote from The Natural History of Butter­flies by John Feltwell (1986):

"Their bright wing colours are enhanced by sun­shine, they recognize each other by their different sexual colours and different absorption of ultraviolet light, they fly in it, they court in it and will readily mate in it ... , their distribution is governed by it, their po­tential attack from parasites is influenced by it, their behavior is governed by the amount and intensity of sunlight energy and light, and indirectly, the release of their plant foods (nectar) is determined by the amount of light."

As with all other animals except birds and mam­mals, butterflies are POIKILOTHERMIC, that is, they are "cold-blooded" since they do not have an internal mechanism for maintaining a constant body tempera­ture (birds and mammals, by contrast, are HOMEOTHERMIC or "warm-blooded.") In essence, butterflies are "solar powered." They depend upon the heat from the sun to catalyze the metabolic processes of life.

When a butterfly basks, it positions its wings at a right angle to the incoming light. Some species bask with their wings held horizontally; these are termed DORSAL BASKERS. Others perch with their wings closed and held vertically with one side angled per­pendicular to the sun; these are termed LATERAL BASKERS. The warmth from basking is transported to the butterfly's thoracic flight muscles. Because dark colors absorb more heat than lighter shades, the bod­ies and wing bases of most butterflies are usually darker in color. In addition, the bodies usually have long hair­like scales that act as heat traps. (Moths, which fly in the cooler hours of darkness, are especially "hairy.') Dif­ferent species require different body temperatures be­fore becoming airborne. The monarch, for example, re­quires a body temperature of 55() F (13() C) By contrast, the black swallowtail requires 90" F (32" C) Butterflies

20 I

I

are truly "sun worshippers." Often while basking, butterflies will slowly open

and close their wings. This behavior is a means of regu­lating heat absorption and dispersal. In addition, many temperate and polar species can rapidly contract their flight muscles to cause their wings to vibrate or tremble. This behavior produces significant quantities of meta­bolic heat and is frequently employed when sunlight is insufficient to raise the body temperature to the de­sired level. The various processes by which butterflies regulate their internal body temperatures is technically termed THERMOREGULATION.

35. Do butterflies sleep?

Yes. Butterflies regularly shut down most of their metabolic processes to rest--particularly during night. Since butterflies have no eyelids, their eyes remain open. Some species such as the longwings and milk­weed butterflies have sleeping assemblages. These so­cial roosts theoretically enhance protection from preda­tors and adverse weather. To sleep, butterflies usually cling to the undersurfaces of vegetation, rocks, or even man-made structures. Sleeping beneath a canopy pro­vides concealment and protection from precipitation. If disturbed when asleep, a butterfly usually will fly just a short distance before alighting (remember, for sustained flight a butterfly's body temperature must be above a critical threshold).

36. Do caterpillars feed only on leaves?

Most caterpillars are HERBIVOROUS or PHY­TOPHAGOUS, that is, they feed exclusively on plant parts: leaves, flowers, flower buds, leaf buds, and ten­der stems. A few spec_ies, for example the North Ameri­can harvester (Feniseca targuinius), are CARNIVO­ROUS, that is, they feed on animal tissue, specifically, aphids--those tiny yellowish sluggish insects of the Order Homoptera that suck juices from plants. Inter­estingly, the duration of the larval stage of the harvester is much shorter (only 7-11 days) than in other phytopha­gous species. Scientists theorize that the high protein diet accelerates development.

37. What do butterflies eat? (Figure 10)

Since the mouthpart or PROBOSCIS of a butterfly

is designed for siphoning liquids (see question 21 ), but­terflies feed mainly on plant nectars (sugary fluids pro­duced by specialized organs called NECTARIES located within the flower). Some species, however, imbibe from over-ripe fruits, carrion, urine, manure and feces, ani­mal perspiration, and soil containing high concentra­tions of minerals and salts. Some tropical butterflies, particularly ithomiids and skippers, follow army ant colonies on the move in order to feed on excrement of antbirds (family Formicoriidae), species specialized for feeding on ants, and which also follow army ants. Longwing butterflies are peculiar in that they actually supplement their nectar diets with pollen (the yellow powderlike substance produced by the anthers (male reproductive structures) of flowers. Small quantities of pollen are combined with saliva to create a cocktail that can be easily siphoned through the proboscis. Scien­tists believe that the proteins, amino acids, minerals, and salts acquired from non-nectar food sources aid the butterflies with reproduction-particularly sperm production-and general body maintenance. Perhaps that is the reason longwings have the greatest longev­ity (up to 11-12 months) of any butterflies in the world.

Digestion in the lepidoptera is not too different from that in mammals. Following ingestion through a MOUTH (PROBOSCIS in adults), food moves through the PHARYNX and ESOPHAGUS and then is tempo­rarily stored in a CROP (analogous to the mammalian stomach). Since butterflies feed on liquids, their crop is much larger than its counterpart in caterpillars, which feed on solid plant material. Next, the food moves to the MIDGUT where it is broken down with the aid of SALIVA from SALIVARY GLANDS. Butterflies usually secrete only one kind of enzyme to digest sugar, whereas caterpillars secrete a compliment of chemical catalysts to handle their varied diet. The midgut en­cases food within a thin, porous, and transparent tis­sue termed a PERITROPHIC MEMBRANE--similar to a sausage skin. After nutrients are absorbed, all undi­gested material is transported to the HINDGUT and RECTUM where water is absorbed. In butterflies, waste material is granular, brownish-black, and relatively in­conspicuous. Combined with URIC ACID crystals from the MALPICHIAN TUBULES (see questioin 33), the material is excreted from the ANUS, usually when the butterfly is at rest.

21

Because caterpillars feed voraciously on plant ma­terial, which is relatively difficult to digest, caterpillar excrement is larger and more conspicuous than that of butterflies. Technically termed FRASS, the waste ma­terial is basically greenish in color, and eliminated as sizable pellets at frequent intervals throughout the day­as anyone who has raised caterpillars in captivity can attest!

38. Do butterflies have to drink water?

Most butterflies secure enough water from their nectar diets to sustain all metabolic processes. Migrat­ing species such as the monarch, however, will imbibe water after a long flight. When you observe butterflies on damp soil or at the edge of a source of water they

Egg (ovum)

0 .

' ' ' '

'

•

Caterpillar (larva)

~ . .. -

IQ,IIH·~I·~

Figure 11. Life cycle of a typical butterfly (sizes are not relative).

probably are imbibing the fluid in order to extract any minerals that it might contain. Often butterflies, par­ticularly swallowtails, sulphurs, and blues, will con­gregate at specific damp locations. Such a phenom­enon is called "PUDDLING." If you closely watch a butterfly at this time, you will notice that the insect fre­quently squirts a liquid from its anus. Chemical analy­sis of this liquid has proven it to be nearly one-hun­dred percent water. Presumably, the butterfly siphons up the mineral-rich water, extracts the trace elements, and then excretes the excess water. Since most butter­flies that puddle are males, scientists theorize that the trace elements are important in the production of sperm and possibly pheromones.

Chrysalis, Chrysa lid (pupa)

Adult (imago)

.,.

I

LIFE CYCLE: METAMORPHOSIS

39. What is a typical butterfly life cycle? (Figure 11)

Butterflies characteristically undergo what is termed COMPLETE METAMORPHOSIS, that is, they change form between each of their four stages (see question 40). A typical life cycle is illustrated below:

Stage I: egg · (ovum-ova)

Stage 2: Caterpillar (larva-larvae)

Stage 3: Chrysalis-Chrysalises, Chrysalid-Chrysalids­Chrysalides (pupa-pupae)

Stage 4: butterfly-butterflies (adult, imago-imagos-imagines)

40. What is metamorphosis? (Figure 11)

The word literally means "changing form." Meta­morphosis is a characteristic of the insect orders Lepi­doptera, Coleoptera (beetles), Hymenoptera (wasps, bees, ants), and Diptera (flies, gnats, mosquitoes). All have life cycles containing four distinct stages: EGG, LARVA, PUPA, ADULT. Such insects are termed HOLOMETABOLOUS since they undergo a complete

A B C

23

change from one stage to the next. (Other insects that do not undergo such drastic changes in their life cycles are termed HEMIMETABOLOUS.)

41. What is a cocoon? (Figure 12)

A cocoon is a structure woven by a caterpillar to encase itself for PUPATION. The cocoon is usually com­posed of hairs and spines from the caterpillar, and all cemented together by silken threads that are secreted from the SPINNERET, a nozzle-like organ located on the LABIUM behind the MANDIBLES. The cocoon is a protective housing for the PUPA. Most larvae of moths as well as many larvae of skippers spin cocoons of various complexity.

42. What is a chrysalis? (Figure 11)

The larvae of true butterflies do not spin a cocoon. Their pupae are usually naked, that is, not encased. Termed a CHRYSALIS or CHRYSALID the pupa bears a posterior point of attachment called a CREMASTER equipped with small hooks or CROCHETS. Silken threads spun from the spinneret fasten the crochets onto a twig or other surface. The chrysalis either hangs sus­pended, or else is supported in an upright position by a single silken girdle.

Figure 12. Moth pupae.

A. Ventral View B. Lateral View C. Lateral View D. Pupa enclosed within its cocoon

D

Aztec butterfly stamp from Zempooala,

from the book Pre-Hispanic

Mexican Stamp Designs.

J.

43. How long do butterflies live?

That depends. The average figure quoted for a typi­cal butterfly is between 2-3 weeks. Some early spring species, however, such as orangetips, marbles, and the spring azure, live for only 2-3 days. Others, such as the various anglewings overwinter (HIBERNATE) as adults, and therefore may live 6-7 months. The migra­tory monarch in the fall has an exceptional longevity: 7-8 months. And some of the longwings and ithomiids of the American tropics live up to one full year.

44. Can butterfly eggs be seen with the naked eye? (Figure 11)

Yes. Although tiny, butterfly eggs are quite visible. Eggs come in a variety of shapes and colors, although most tend to be roundish and yellowish. They are usually attached to the young, tender growth of host plants (buds, new leaves, tendrils) by an adhesive secreted by the female butterfly. Oc­casionally, females confuse their chemical cues and deposit their eggs on objects near the host plant. Some species, for example, the greater fritil­laries (Spcycria), always oviposit on ob­jects near their host, never directly on it. And some skippers and satyrids, which breed on various grasses, have been reported to simply broad­cast their eggs while flying low to the ground.

45. How many eggs do butterflies lay?

An average number is 100-400. Some species, how­ever, ·lay many more. The diana (Speycria diana), for example lays between 1,100 and 1,300 eggs. Fertility or fecundity is closely associated with species mortal­ity: high risk species lay greater numbers of eggs than do those species with lower risks, and therefore a higher potential for survival. Remember, butterflies are low on most food chains. Consequently, butterflies-- in all their stages--are food for many other species of organ­isms. Since the vast majority of butterflies succumb to diseases and predators, each species needs to compen­sate for the losses by producing the number of eggs that will more or less guarantee their reproductive sue-

cess. Incidentally, although the females of most spe­cies lay eggs singly over a period of time, some anglewings, checkerspots, many ithomiids, and a few skippers, all lay their eggs in clusters. After hatching young larvae usually remain together for a short time. As they mature, however, they usually separate and live independent of one another. Scientists theorize that clustering increases the chances for survival. Remem­ber the old adage, "safety in numbers."

46. Do all butterflies lay eggs?

Practically all butterflies are OVIPAROUS, that is, they lay eggs from which develop larvae. A few spe­

cies, however, such as sulphur butterflies in the genus Colias, are reported to retain their eggs

until hatching, and then deposit live lar­vae. These butterflies are OVOVIVIPA-

ROUS. This unusual condition suppos­edly gains the advantage of reduced predation and exposure to inclement weather conditions.

47. Do female butterflies watch over their eggs?

Once eggs are deposited, females move on. Females of some danaid spe­

cies in the Philippines, however, watch over their egg clusters until hatching.

48. What is the incubation time for an egg?

An average time for a temperate species is usua!ly 4-10 days. Some species, however, HIBERNATE in the egg stage. In such cases the life span is approximately 6-7 months.

49. What is the life span of a caterpillar?

An average time for a temperate species is usually 3 to 4 weeks. Some caterpillars of arctic and alpine spe­cies require two full years to mature. Also, the larvae of the greater fritillaries usually require eight months. On the other hand, the larvae of the harvester require just 7-11 days.

24

50. What is the life span of a chrysalis?

An average time for a temperate species is usually 7-14 days. Some species such as the orangtips and marbles, however, enter a long period of rest or sus­pended animation (DIAPAUSE). With them, the chrysa­lis may last 3-4 years! Many butterflies (as well as skip­pers and moths) overwinter or HIBERNATE in the chrysalis stage. With those species the life span is ap­proximately 6-7 months.

51. How many generations a year does a butterfly complete?

The number of life cycles (VOLTINISM) depends usually on climatic conditions: temperature, moisture. For example, the colder the temperatures, the fewer generations, the warmer the temperatures, the greater the number; also, the drier the climate, the fewer gen­erations, and the more rainfall, the greater. Several arc­tic and alpine species require a full two years to com­plete just one generation (BIENNIAL). By contrast, many desert species have only one flight each year (UNIVOLTINE) during the brief rainy season, the only time when host plants are available. These butterflies pass through the remaining hot, dry months in a more or less resting state (AESTIVATION) in one of the three immature stages, awaiting the next rainy period. But for most butterfly species within temperate zones, the average number of generations per year is between 2 (BIVOLTINE) and 3 (TRIVOLTINE). (Common excep­tions are the greater fritillaries, and the early spring flyers such as elfins, orangetips, marbles, and various species of skippers--all of which complete only one.) Through natural selection, each species has so fine­tuned its reproduction that its numbers remain rela­tively stable from year to year. Of course, we know that such is not always the case. Certain species have at times become extremely abundant, or else have fallen by the wayside to b~come extinct.

52. How do butterflies survive winter?

That depends. Some butterflies HIBERNATE, that is, they slow down their metabolism and seek shelter under leaves, rocks, logs, or crevices. Some may even take advantage of man-made structures such as "but­

hotels" or "butterfly log piles" (see questions 96

and 97) that we set out as garden accessories. Most spe­cies, however, endure the inhospitable cold season in an immature stage: egg, caterpillar, or chrysalis. A few species in the frigid arcitc and alpine regions actually require two entire years of immature development be­fore reaching the adult stage.

53. Should caterpillars be called "worms?" (Figure 6)

Although caterpillars are commonly referred to as "worms," caterpillars are not really worms. True worms do not possess legs or leg-like structures. Members of the phyla PLATYHELMINTHES (flatworms), NEMATODA (round worms) and ANNELIDA (seg­mented worms) are all invertebrates that fit the classic definition of "worm." Caterpillars, on the other hand, posses TRUE LEGS and PROLEGS, and are simply an immature stage (larva) of a butterfly, skipper, or moth. They should be accorded their rightful status.

25

A

D

E F

Figure 13. Types of androconia in the lepidoptera (illustrations are not relative in size). (Figures B,D and E after Howe, 1975, and used with permission.)

A. Androconia (Alar) Patch-Family Danaidae (monarch) B. Hairpencil-Family Danaidae (large tiger) C. Androconial Fold-Family Papilionidae D. Androconial Patches-Family Satyridae E. Stink Club-Family Heliconiidae (gulf fritillary) F. Hairpencil-Family Ithomiidae G. Androconial Tuft-Family Mophidae

B +

G

26

\ ,

A

(

' I' ... ~

J

REPRODUCTION

54. How do male and female butterflies locate one another? · (Figure 13)

Male butterflies are usually the active partner in courtship and copulation. Wing colors and patterns initiate recognition. The splendid colors that we see in butterflies may play less a role for butterflies in recog­nition than the ultraviolet wavelengths that are invis­ible to us but quite recognizable to butterflies. After the initial attraction, scent in the form of specific sex pheromones beco!11es important. These chemical mes­sengers are often released from specific scales and or­gans (ANDROCONIA) in males: ANDROCONlAL (ALAR) PATCHES, HAIRPENCILS, ANDROCONIAL FOLDS, ANDROCONIAL TUFTS, and STINK CLUBS. Although most of the pheromones are volatile chemi­cals, some are encapsulated in distinct particles called "love dust" that is actually broadcast from the androconia. Sex pheromones can be detected from con­siderable distances. In some moths, for example, the distance can be several miles (kilometers) downwind. In diurnal butterflies, however, distances usually are no greater than 1,000 feet (330 meters) downwind.

Some females produce pheromones before they

Figure 14. Sex differentiation in butterflies.

A .. Male (dorsal view of posterior abdomen) B. Female (dorsal view of posterior abdomen)

A

emerge from their chrysalis. Often males will detect these odors and converge onto a chrysalis in anticipa­tion of the female's emergence. Then, upon hatching, the female is immediately "raped." (Various species of longwings are notorious for this behavior.)

55. How do butterflies copulate? (Figure 14)

Shortly after a male butterfly emerges from its chrysalis, it seeks out a female of the same species (pheromones play a key role in finding a mate). Once located, a male will initiate an elaborate courtship. De­pending on species, there usually are wing displays, abdomen posturing, and sometimes even a release of sex pheromones from specialized scales and organs. If the female is receptive, the male will grasp her with the CLASPERS (VALVULAE) and UNCUS of his geni­talia. He then inserts the AEDEAGUS (the equivalent of a mammalian penis) into her BURSACOPULATRIX (the equivalent of a mammalian vagina). A sperm packet (SPERMATOPHORE) is transferred into the RECEPTACULUM SEMINIS or sperm chamber. Copu­lation may take a few minutes to several hours. If dis­turbed, the couple may fly in tandem (the female usu­ally flys with the male remaining attached and sus­pended) before securing another perch. After release,

ae (aedeagus), cl (clasper). ov (ovipositor), un (uncus)

B

27

r I

males usually seek out other females for additional cou­plings. Depending on species, females may or may not mate again. With parnassians or apollos (Parnassius), the male glues a hard hood-like structure called a SPHRAGIS, secreted during mating by a gland onto the tip of the female's abdomen. The sphragis acts as a sort of" chastity belt" in that it prevents the female from mating again.

56. Do male butterflies have a penis? (Figures 10, 14)

Yes. It technically is an AEDEAGUS, and it is com­posed of a hard chemical substance called CHITIN (the same substance as in the exoskeleton). The aedeagus is retractable, and housed between the pair of CLASP­ERS at the tip of the abdomen.

57. Do female butterflies have a vagina? (Figure 14)

Yes. It is technically called a BURSA COPULATRIX, and is found on the dorsal part of abdominal segment number eight. Sperm is received and then transferred into the RECEPTACULUM SEMINIS, a blind pouch for storage. As eggs pass down the oviduct, sperm from the receptaculum seminis swim out to fertilize them.

58. Why do certain butterflies lay eggs only on certain plants?

The caterpillars of many species of butterflies are

MONOPHAGOUS, that is, they will eat only one type of plant, or plants within only one family. (In contrast, some species are POLYPHAGOUS, that is, they feed on a variety of unrelated plants.) Most plant species contain secondary plant substances, chemicals that are synthesized by the plant, but which are not utilized by the plant during its metabolism. Scientists theorize that these chemicals act as various protective devices for the plant: deter molds, microbes and predators. Dur­ing the evolutionary history of plants and insects, many insects have circumvented the toxic qualities of these chemicals, and in fact, have devised metabolic path­ways that incorporate the would-be toxins (often types of alkaloids) into their own physiological functions. Of­ten these sequestered compounds then serve to pro­tect the insect from its own potential predators. But the

insects have paid a price for the acquisition of their chemical arsenals: they have become dependent upon their host plants. Monarch butterfly larvae, for example, can only survive on plants within the milkweed fami­lies that supply them with specific alkaloids; and mon­arch butterflies will only oviposit on plants that con­tain these compounds. The compounds are detected by the butterfly's antennae and feet (see questions 25 and 26).

28

Aztec butterfly stamp from Guerrero, from the book Pre-Hispanic Mexican Stamp Designs.

)

)

,, BEHAVIOR

59. How far can butterflies fly?

Most butterflies remain within the same general area from which they emerged; after all, loitering in­sures the presence of host plants. Migratory species, however, periodically move from one location to an­other. Monarchs, for example, engage in extraordinary treks. Each fall, millions of these brightly colored in­sects fly from Canada and the United States southward and westward. Populations west of the Rocky Moun­tains head for the coast of southwestern California, while the eastern populations move to the highlands of central Mexico. A monarch tagged in Ontario was recaptured at the overwintering grounds in Mexico, a straight-line distance of 2,133 miles (3,432 km). Since butterflies do not fly in a direct path, the actual dis­tance the monarch flew could be double the recorded figure. (This flight is recorded in the Guiness Book of Records.)

60. How high can butterflies fly?

Most butterflies do not fly very high since they spend most of their time feeding or laying eggs on plants. Migratory species, however, often exploit wind currents and thermals. They, therefore, may ascend considerable heights. Hang gliders and balloonists have recorded migrating monarchs over 7,000 feet (2,300 meters) above the ground.

61. How fast do butterflies fly?

Some butterflies have heavily muscled bodies and are capable of extremely rapid bursts of flight--so fast, in fact, that they are often mistaken for hummingbirds. Migrating species, such as the monarch, often take ad­vantage of prevailing winds to supplement their own flights. Monarchs have been clocked at over 50 miles per hour (80 km) for extended periods. Most butter­flies, however, do not engage in such aerodynamics. They simply flit from flower to flower in a rather er­ratic fashion.

62. What do butterflies do during rains or other bad weather?

During inclement weather, butterflies usually seek shelter under leaves, tree limbs, rocks, or anchored to

the leeward side of secure objects. Rain drops can act as missles to a butterfly, and so it is imperative that these fragile insects quickly find cover when rain be­gins. Although shingled wing scales can effectively re­pel water for a short time, a butterfly's wings will soon become saturated, and so butterflies cannot tolerate even a slow drizzle for long.

63. Can butterflies fly at night?

Usually no. However, migrating monarchs if caught over bodies of water after dark will continue to fly. In addition, some species if accidentally dislodged from their perches after dark will often fly towards bright lights--even if they are some distance away.

64. Why do butterflies sometimes alight on people?

Butterflies are often attracted to specific odors. Per­spiration, various toiletries, and various foods (particu­larly those containing salts and nitrates) are often at­tractive to some butterflies. If a butterfly alights on you notice that it will extend its proboscis in an effort to feed (see questions 26, 37, and 38).

65. Are any butterflies territorial?

Many species throughout the world engage in some degree of territorial behavior, that is, they adopt certain habitat elements as their "home turf," where they remain to defend pugnaciously against all intrud­ers. Territorial behavior is most frequently displayed by males. An individual's territory may be very small, such as a limited space immediately surrounding a fa­vorite perch (many nymphalid species exemplify this), or relatively large, as with many heliconiids and mor­phos that frequently patrol specific pathways or edges of forest for distances of a-half mile (1 km) or more. (These "beats" usually are associated with specific food sources, and remain stable throughout the flowering or fruiting season. As other food sources become avail­able, the route is changed to take advantage of the new banquet.)

Many of the ithomiid butterflies form LEKS, a com­munal display of males engaged in courtship behavior to attract females for mating. Originally named to de­scribe peculiar mating displays of birds, butterfly leks are fairly con;mon throughout the rainforests in tropi~

&fi9 ECOLOGY: INTERRELATIONSHIPS, GEOGRAPHY, AND CONSERVATION

71. What purpose do butterflies serve?

This is a very subjective question, and stems from our anthropocentricity, that is, man's desire to be the most significant entity in the universe. We should re­member that one of the basic laws of ecology states that everything in the universe is in some way related to everything else. Not only is each living species part of a FOOD CHAIN, it is part of a larger FOOD WEB. And so, even though any given species may not seem to have any direct purpose for us, there is indeed a slot for it in the" grand scheme." When it comes to butterflies, there are indeed direct connections to humankind. As polli­nators of plants, butterflies are directly responsible for plant reproduction, and therefore, fruit and seed de­velopment--distinct resources for us (think about our needs for food and natural fibers). Second, since cater­pillars are the food source for many other species-- and the primary food source for some--lepidoptera are in­directly responsible for many of our flesh foods as well as other animal resources (skins, bones, sinews, horns, etc.). Third, since caterpillars are voracious feeders, they are the natural control agents for many species of plants, thereby keeping the plant kingdom from en­gulfing everything. And fourth, butterflies are power­ful aesthetic and spiritual symbols (see questions 107 and 108). As such, they are important to our psyche.

72. How many different species of butterflies and skippers are there in the world?

The actual number as of 1989 is 17,280. New spe­cies, however, are being catalogued each year (although entomologists theorize that probably 90 percent of the world's butterflies have already been catalogued). The eventual number will probably be close to 20,000, or about one-tenth that of the number of moth species (es­timated at 200,000). Below is a breakdown of species known for each family (modified from Shields, 1989).

FAMILY Papilionidae: Pieridae: Lycaenidae:

NUMBER OF SPECIES 566

1,215 4,089

Riodinidae: 1,366 Libytheidae: 12 Acraeidae: 240 Nymphalidae and

Heliconiidae: 2,671 Apaturidae: 431 Amathusiidae: 100 Morphiidae: 55 Brassolidae: 81 Satyridae: 2,400 Danaidae: 157 Ithomiidae: 305 Hesperiidae: 3,592

TOTAL: 17,280

73. How many different species of butterflies and skippers are there in North America?

To date, 717 species have been recorded within the United States and Canada.

74. How many different species of butterflies and skippers are there according to state and province within North America?

Below is a break down of butterflies (including skippers) along with birds for comparison in various political divisions (modified from Robbins and Opler, 1995). Residents are defined as species that reproduce yearly and can survive all seasons; colonists are spe­cies that do not survive the winter, but which annually immigrate into the region and breed; vagrants are spe­cies that have been reported in the region, but do not usually breed there. ·

Butterfly Residents Butterfly Total Breeding

State &Colonists Vagrants Butterflies Birds

Alabama 132 2 135 145 Alaska 68 0 68 430* Arizona 246 80 326 246 Arkansas 127 25 152 130 California 225 25 250 286 Colorado 230 36 266 235 Connecticut 101 13 114 158 Delaware 91 7 98 160

Florida 163 18 181 Georgia 151 8 159 Hawaii 13 0 13 Idaho 154 6 160 Illinois 121 22 143 Indiana 123 19 142 Iowa 107 19 126

, Kansas 133 50 183 Kentucky 116 17 133 Louisiana 117 15 132 Maine 88 13 101

. Maryland 121 19 140 Massachusetts 93 19 112 Michigan 134 10 144 Minnesota 132 13 145 Mississippi 134 10 144

· Missouri 125 31 156 : Montana 184 3 187 '

Nebraska 170 27 197 · Nevada 181 26 207 . New Hampshire 92 9 101

New Jersey 120 23 143 New Mexico 272 46 318 New York 119 19 138

1 North Carolina 140 11 151 ! North Dakota 132 11 143

Ohio 131 7 138 Oklahoma 146 16 162

· Oregon 159 5 164 · Pennsylvania 114 20 134

Rhode Island 83 4 87 . South Carolina 133 9 142 South Dakota 149 21 171 Tennessee 112 12 124 Texas 290 133 423 Utah 197 18 215 Vermont 66 6 71

Virginia 134 21 155 Washington 140 3 143 West Virginia 112 8 120 Wisconsin 133 12 145

•· Wyoming 197 13 210

:CANADA

: Quebec 140

t~Combination of breeding and vagrant species

.)

160 160

269* 355* 160 151 154 175 153 158 176 192 177 202 224 130 175 224 194 224 175 181 247 220 178 171 180 180 232 185 141 152 207 160 300 220 175 179 235 156 203 222

405*

75. Where in the world are the most species of butterflies?

The tropical rainforests are the richest habitats on earth. Brazil, with the largest tract of rainforest (the Amazon Basin), not suprisingly has the greatest diver­sity of butterflies. As an example, within a 2 square mile (3 km) section of rainforest in the state of Rondonia in western Brazil, nearly 2,000 species of butterflies and skippers have been catalogued within only the last 10 years.

Within North America, four regions are exception­ally rich in species: south Texas, southern California, south-eastern Arizona and south-western New Mexico, and the Rocky Mountain Front Range of Colorado. Sci­entists theorize that this high diversity is due to the presence of a wide range of topographic relief and com­plexity combined with a proximity to Mexico with its very diverse tropical fauna.

76. Where in the world are the fewest species of butterflies?

Since the lepidoptera do not have internal heat regulation, they tend to be poorly represented in those regions of the earth that remain relatively cold through­out the year. Antarctica, for example, has no lepi­doptera, and the arctic regions of both the western and eastern hemispheres have but only a few. Six species have been recorded from Greenland (and all occur on the North American mainland as well), and Alaska has 68 species. The country of Iceland has the lowest for any country: four species have been recorded as va­grants. Tropical Hawaii, with a meager 15 species, at first consideration seems an anomaly. However, Ha­waii is an island isolated by great expanses of water in all directions. Furthermore the island is of recent vol­canic origin. Both factors reduce the potential for colo­nization, and therefore override the state's tropical na­ture.

77. Are there any endangered or threatened species of butterflies or skippers within North America?

Yes. The United States Fish & Wildlife Service lists 16 species (including subspecies) as endangered (few living members, and in danger of becoming extinct in

33

the near future) or threatened (not now endangered, but whose populations are heading in that direction). These are (from U.S. Fish & Wildlife Service, August 20, 1994):

bay checkerspot (Euphydryas cditha baycnsis)--CA ... El Segundo blue (Euphilotcs battoides allyni)--CA Karner blue (Lycacides melissa samuclis)--(IL, IN,

MA, MI, MN, NH, NY, OH, PA, WI ...... and Ont., Canada

Lange's metalmark (Apodemia mornzo langci)--CA Lotis blue (Lycacidcs argyrognomon lotis)--CA mission blue (lcaricia icarioides missioncnsis )--CA Mitchell's satyr (Nconympha mitchcllii mitclzcllii)--IN,

MI, NJ, OH Myrtle's silverspot (Spcycria zcrene myrtleae)--CA Oregon silverspot (Spcycria zcrene hippolyta)--CA,

OR,WA Palos Veredes blue (Glaucopsyclze lygdamus

palosverdcscnsis )--CA Saint Francis' satyr (Neonympha mitchcllii francisci)-­

IN, MI, NJ, OH San Bruno elfin (Callophrys mossii bayensis)--CA Schaus swallowtail (Heraclidcs aristodcmus

ponceanus)--FL Smith's blue (Euplzilotcs enoptes smitlzi)--CA Uncompahgre fritillary (Boloria acrocncma)--CO Pawnee montane skipper (Hesperia lconard

mon tana )--CO

The service includes the following exotic species: Corsican swallowtail (Papilio lzospiton)--Corsica,

Sardinia Homerus swallowtail (Papilio /zomcrus)--Jamaica Luzon peacock swallowtail (Papilio cl1ik11e)--Philip­pines Queen Alexandra's birdwing (Troides=Omithoptcra

11/exa11drac)-- Papua New Guinea

Although these are the only species currently listed, several others are candidates for listing and several have already become extinct. For example, the Xerces blue (Cla11copsycl1e xcrccs) has the unfortunate distinc­tion of being the first butterfly to be recorded as extinct within the U.S. (the species was last seen in California in 1943).

Incidentally, one species of moth, the Kern prim­~

rose sphinx (E11proscrpin11s eutcrpc) in California, is also listed as endangered.

78. Which are the best months to observe butterflies?

That depends upon geographic location. Within the southern United States, usually August through Oc­tober are prime months. Then, populations of most species are highest because of many months of repro­duction. However, in more northern or mountainous regions where winters tend to be long and summers relatively short, prime months are more likely to be June and July. And, in all areas, some species (elfins, orangetips, marbles, harvester, and several skippers) have only one generation per year, and that is usually in early spring, or when temperatures begin to warm (March to April in the south, May to June in regions farther north or at high elevations).

79. What is a "Fourth of July Butterfly Count?"

In 1975, the Xerces Society (a conservation organi­zation concerned with the conservation of inverte­brates) initiated a program to survey butterflies in a given locale on a yearly basis. Modified from the Na­tional Audubon Society's "Christmas Bird Count," the rules are simple: select a location that allows you ac­cess to a circle with a 7.5 mile (13 km) radius; count all butterflies (both species and individuals) during a single day within a three-week window before or after July 4; report your findings, and repeat each year. To­day the count is administered by both the XERCES SO­CIETY and the newly formed NORTH AMERICAN BUTTERFLY ASSOCIATION. The address is below:

Ann B. Swengel, International Co-editor "Fourth of July Butterfly Count" 909 Birch Street Baraboo, Wisconsin 53913

80. What is the best time of day to observe butterflies?

Most butterflies are DIURNAL, that is, they fly dur­ing daylight hours. Since butterflies rely mainly on the sun to warm their bodies, most species are not active until mid morning. Within the southeastern United States generally between the hours of 10 AM and 2 PM

34

C

a C

l b 0

s 0

S]

u

s:

p, m

(CST) is prime time for most species. Some species, however, are active during practically all daylight hours throughout the summer and fall months, and some seem to prefer early morning and/ or late afternoon only. (Peak times for butterflies in regions with rela­tively cool temperatures even during the summer months, will differ.) Cloudy days affect peak times, too. Some species in the tropics are CREPUSCULAR, that is they are active at dusk and dawn.

81. Are butterflies sensitive to environmental disturbances?

Yes. Probably no other animal is more symbolic of a clean, healthy environment than the butterfly. But­terflies are ideal monitors for habitat destruction, pol­lution, and applications of chemicals such as pesticides and herbicides. For this reason, the symbol of a butter­fly is frequently employed to epitomize everything that is good, clean, and healthy.

sands of dollars for very rare species (such as Queen Alexandra's Birdwing of New Guinea). As butterflies become more and more endangered, purchasing but­terflies becomes more and more problematic. When purchasing specimens one must keep in mind that traf­ficking in endangered or threatened species is illegal!

84. Are there any fossil butterflies?

Yes. Scientists theorize that butterflies came into existence about 150 million years ago at about the same time as the flowering plants. Geologists designate this time as the early CRETACEOUS PERIOD. Although the delicate wings and soft bodies of butterflies do not preserve well, fossils have been found in slate, gyp­sum, and limestone deposits of Europe dating to the TERTIARY PERIOD (later than the Cretaceous, and about 65 million years ago). The butterflies resemble

...,._ nymphalids, lycaenids, and swallowtails. Amber (fossilized resin from trees) also

has yielded preserved butterflies. De-~ i,iw~ posits from the Baltic, Dominican Re-

82. What is the rarest butterfly in f. public, and eastern Africa have pro-

the world? f '-"'i,,,. .. rlllll~•!::::~• duced swallowtails, lycaenids and

is is i icu t to answer. ere are Th. · d"ff" 1 Th w riodinid-like butterflies. In Canada, · d b • fb an amber sample has produced the many species an su species o utter-

fl . h k 1 f f head of a caterpillar, probably that of . . . . a moth because of the leaf tunnel pre-

1es t at are nown on y rom a ew ~ specimens m collections of considerable d . h . Th. 1 • h 1. . . . . serve wit 1t. 1s samp e is t e ear 1-age. Likewise, some vanehes are known .... ,.-, .

1 f 11 d • 1 d "ld 1 . h est known lepidoptera. It dates to late Cre-on y rom sma an iso ate wi popu a hons t at . . · · • 1 h df 1 f . d. .d taceous deposits, about 72 million years ago. contam m any given year on y a an u o m ivi u-

als. Within North America the least common species certainly include those listed as "Endangered or Threatened" (see question 79). Included, too, would be several of the hairstreaks, elfins, metalmarks, orangetips, and marbles that occur in extremely re­stricted locales, have very short flight periods, and have only a single generation each year. In fact, some of these species are so uncommon that their life cycles remain undescribed.

83. Are butterflies marketed commercially?

Yes. There are scattered throughout the world com­panies and individuals that market butterflies on a com­mercial basis. Prices range from less than a dollar for common species upwards to hundreds and even thou-

~

85. What are the chief enemies of butterflies?

Natural biological agents such as microbes (bacte­ria and viruses), parasites (wasps, flies, bugs), and parasitoids (wasps and flies) destroy the great major­ity of butterflies in all stages of their development. ln addition, the wholesale broadcasting of insecticides and herbicides take their toll on insects in general. Grass mowing and other forms of habitat alteration also con­siderably reduce breeding habitats for many species. Human collecting, except under extraordinary condi­tions (such as in small isolated populations) rarely causes significant long-term damage. (The biology of invertebrates such as insects is very different from the biology of vertebrates. Invertebrates have extremely high reproductive rates in order to compensate for high

35

Aztec image of a butterfly, from Zempoala From the book Pre-Hispanic Mexican Stamp Designs

levels of mortality.) In fact, butterfly collecting is one of man's oldest hobbies and one of the most successful and enjoyable ways of turning youngsters on to the magical world of biology. The operative words for col­lectors are prudence and respect.

86. Can butterflies transmit any disease to humans? (Figures 3,7)

No. The proboscis of a butterfly is engineered for sucking liquids, not for biting. And since butterflies do not usually alight on the food of humans, the acciden­tal mechanical transmission of pathogens from the feet of butterflies is virtually impossible.

87. Are some butterflies distasteful to predators?

Yes (see question 58). For example, the monarch as well as a myriad of other species found principally within the tropics, contain chemicals within their bod­ies that render them unpalatable to most veterbrate predators. Many of these distasteful butterflies possess striking colors and patterns. Would-be predators soon learn by trial and error these signals and then avoid the distasteful species. This warning coloration is termed APOSEMATIC COLORING (see question 6). Aposematically colored individuals, termed MODELS, are often imitated by others, termed MIMICS, that are not endowed with the potent toxins. Because preda­tors are unable to distinguish model from mimic, both are avoided. This type of mimicry was first investigated in 1862 by the famous English naturalist H.W. Bates while working in Brazil, and is now known as BATE­SIAN MIMICRY after its founder. Examples include the monarch (model) and viceroy (mimic), the pipevine swallo\vtail (model) and red-spotted purple and the dark female tiger swallowtail (mimics).

Another form of mimicry was investigated by the German naturalist, F. Muller in 1878, who also worked in Brazil. Miiller observed that two or more obnoxious species often resemble each other even though they are unrelated. He theorized that this mutual guise speeds up predator education by reducing the number of col­ors and patterns confronting potential predators, and therefore reducing the number of trials necessary for learning. In essence, the mimetic species reinforce each

other by reinforcing the guise. Predators, therefore, are continually confused. This form of mimicry is termed MULLERIAN MIMICRY after its founder. (Within the American tropics, the families Heliconiidae and lthomiidae--and to a lesser extent Papilionidae, Pieridae, and Nymphalidae--are actively involved in Mi.illerian complexes.)

88. Do some caterpillars produce honeydew as do aphids, and are these caterpillars attended by ants?

Yes. Larvae of many of the lycaenid and riodinid butterflies have unique organs that not only produce a sweetish honeydew like substance, but also sounds, and pheroinones that attract ants. These organs arc termed MYRMECOPHILOUS, and the ant-association, MYRMECOPHILY-a type of SYMBIOSIS. Within these two families of butterflies there exists a great variety of relationships with ants. In some, the ants simply "milk" the caterpillars. In others, the ants protect the caterpillars by fending off predators, and in some cases by actually herding the caterpillars to safe havens dur­ing prime hours of predation. These relationships are best developed in the tropical zones, and much research still has to be completed before scientists fully under­stand the implications. (See Balmer & Pratt, 1991; DeVries, 1992; Ross, 1985, 1994b.)

36

Aztec butterfly stamp from Teotihuacan, Mexico, from the book Design Motifs of Ancient Mexico.

a s ti fl

9

BUTTERFLY GARDENS

89. What is a butterfly garden?

"While it is true that no matter how hard we try we cannot confine butterflies to our home turf, we can welcome the most species in the greatest numbers in a given space by creating a butterfly-friendly environ­ment" from Gardening Jcir Butterflies in Louisiana (Ross, 1994a). Simply, butterfly gardens are old-fashioned gar­dens containing lots of flowering plants. These plants serve as sources of food for butterflies as well as places for butterflies to lay their eggs .. Although such gardens were common in the past, we industrious Americans have become more organized and less domestic. As such, we have replaced our hand-tended flower beds with border plantings of evergreen shrubbery. Al­though the new look is more convenient for city folk, it is less seductive to pollinators such as butterflies. (See Ruffin, 1993, for listings of butterfly gardens.) A but­terfly garden is a return to the "cottage garden" of old with its compliment of fragrant, colorful flowers and medicinal and culinary herbs.

90. Are butterfly gardens difficult to install?

No. Butterfly gardens require only meager invest­ments of time, energy, and money. Key factors to con­sider are sunlight, good drainage, loose soil, draft-free location, and plants that have proven to be "butterfly friendly," that is, good sources of nectar and species appropriate for egg laying.

91. Are butterfly gardens difficult to maintain?

No, but they do require more attention than typi­cal plantings of shrubbery and trees. Flower gardens require periol:lic weeding, fertilizing, and watering. And since many of the plants that attract butterflies are annuals, they have to be replaced the following sea­son. But, the resulting joy and peace associated with the ever-changing tableaux of flowers and "dancing flowers" (see question 109) are worth the extra efforts.

92. What plants do butterflies prefer?

There are many species of native and exotic flow­ering plants that attract butterflies. In the southern United States favorites include several forms of Ian-

tana, red pentas, and butterfly bush (Buddleia). See the following sources for detailed discussions: Ajilvsgi (1991), Ortho Books (1991), Ross (1994a,d,e), Sedenko (1991), Stokes and Williams (1991), and Xerces Society (1990).

93. What is the difference between a "food plant" and a "host plant?"

A food plant is a plant utilized by a butterfly for feeding: the plant furnishes the butterfly with nectar and sometimes pollen. A host plant is a plant that is utilized by a butterfly during its reproduction: eggs are laid on these plants. Host plants are the food plants for caterpillars.

94. Do caterpillars eventually destroy a butterfly garden?

No. Butterfly gardens usually contain principa!ly plants that produce nectars for butterflies (food plants). Butterflies generally do not oviposit on these. If host plants are included, butterflies will most likely deposit eggs there. The resulting caterpillars can seriously de­foliate these plants. The caterpillars, however, will not feed on any other plants in the garden (see question 93), and natural predators will usually keep them un­der control. Simply plant a sufficient quantity of host plants so that they can be shared equally between gar­dener and butterfly. Serious garden problems are usu­ally caused by snails, slugs, grasshoppers, crickets, aphids, scale insects, and a variety of true bugs. Occa­sional holes in foliage should not be considered prob­lems in a butterfly garden. After all, a garden is a liv­ing mini ecosystem in which interactions are common events. These observations will increase your avvare­ness of the natural world.

95. What is a "butterfly house?"

"Butterfly house," "butterfly pavilion," ''butterfly conservatory," "butterfly aviary," and "butterfly farm," are all names for an enclosure designed to exhibit liv­ing, free-flying butterflies within an aesthetic, natural­istic setting. In tropical climates, the structure usually has walls and ceiling of simple shade cloth. But in more temperate regions, temperature and humidity must be considered, and so glass is used to contain the insPcts.

37

Aztec image of a butterfly,

from Guerrero From the book

Pre-Hispanic Mexican

Stamp Designs

The first butterfly houses appeared in the United King­dom in the late 1970's. Today, the U.K. alone boasts over 60, and the concept has spread to Australia, Singapore, France, Japan, Canada, and the United States. In North America there are now approximately 15 houses. "But­terfly World" in Coconut Creek, Florida and the "Cecil B. Day Butterfly Center" at Callaway Gardens, Pine Mt. Georgia, both opened in 1988 as the hemisphere's first houses. (See Ruffin, 1993, for others.)

Most butterfly houses are attractions in and of themselves; others represent additions to established zoos or botanic gardens. Some houses breed their own stock, while others import live pupae (costing a few cents to nearly $15 apiece) for hatching and later re­lease. Live pupae are marketed by "butterfly farms" located principally within developing tropical coun­tries. For native peoples, who have traditionally relied on environmentally destructive methods for their live­lihood, these simple enterprises create practical lessons f~ learning the value of natural resources and the in­terdependency of all plants and animals. As such, but­terfly husbandry and butterfly pavilions are now im­portant components of the sweeping global movement to increase mankind's stewardship of planet Earth.

Butterfly houses offer exceptional opportunities to learn not only about insects, but the entire natural world. For this reason, many houses offer extensive educational programming that is available not only to visitors but to schools.

But as important as aesthetics, conservation, and education are, there is an even greater reward from a visit to a butterfly house. Today, many people are searching for some form of spirituality in their lives. Because butterflies have historically been associated with joy, hope, and rebirth (see question 108), butterfly houses are able to provide us with just that magical moment when we can connect with our inner self and refresh our hearts and souls with the philosophy of hope.

96. What is a "butterfly hotel?"

A "butterfly hotel" is a garden accessory con­structed of wood and similar to a bird house except that there are several vertical slits for openings. The hotel is technically a HIBERNACULUM or hibernation box. The structure prcwides shelter for butterflies dur- .

~ <;"' ,,

ing inclement weather and during the winter season. These structures also provide a haven for caterpillars searching for a site to pupate. Butterfly hotels seem to work best in large urban areas with extreme winters. In other areas, butterflies usually are able to locate suf­ficient natural places to hide.

97. What is a "butterfly log pile?"

A number of logs stacked crosswise in log cabin fashion, usually at least 3 feet (1 meter) and 3 feet square, and with a sheet of plastic or canvas to roof is called a "butterfly log pile." The structure acts as a HIBER­NACULUM for butterflies and other small creatures.

98. What are butterfly feeders?

Butterfly feeders basically are dishes containing sugary fluids to attract butterflies. The feeders can be purchased or simply created with a pot saucer to hold the fluid and a sponge or scouring pad to act as a perch. Many enthusiasts have their own recipe for priming their feeder. Ingredients often include molasses, stale beer, brewer's yeast, and an assortment of overripe fruit. In the tropics, feeders are particularly effective, so much so that very often the "cocktail" is simply brushed onto some substrate or else simply poured onto the ground. Very often, too, feeders will include dead animal tissue, feces from carnivores, and even aged human urine. Many feeders will attract specific canopy dwelling butterflies that otherwise are almost impossible to observe.

0

-

/

)

RESEARCH AND COLLECTING

99. Why do people collect butterflies?

Throughout the ages people have collected butter­flies to display their beauty as works of art, for scien­tific research, or simply as a hobby. The great icons of natural history, for example, Carolus Linneaus, Charles Darwin, Henry Bates, and Alfred Wallace, were all avid collectors. The development of all the natural sciences has depended heavily on preserved material housed in museums or private repositories. Even today, col­lections are just as important to contemporary research as they were to the scientists of yesteryear. In reality, collections are tangible references to the myriad of life forms that exist or that have ever existed on our planet.

With the current popularity of butterfly pavilions around the world, collections are often made in the field to obtain females for eggs that are either shipped out for rearing or else hatched, the larvae reared through the pupal stage, and the pupae then shipped. Very of­ten, a percentage of the emergent adults is released into the wild to restock natural populations. (See question 45). Such releases are extremely beneficial since the number released far e«ceeds the number that would have developed normally in nature. As such, collect­ing actually increases the survival potential for many species.

People who collect butterflies usually not only per­sonally collect but also trade specimens with other col­lectors--in much the same manner as collectors of stamps. In addition, some collectors purchase desirable specimens from commercial suppliers (see question 82).

Recently, there has been a growing number of butter­fly enthusiasts (both professional and amateur) pro­moting recreational activities that pivot around obser­vations made through binoculars and cameras. (see Glassberg, 1993). As conservation becomes more and more critical, these alternate forms of study and recre­ation will probably become more and more widespread.

100. Can a person attract butterflies to a specific location?

Yes, by planting butterfly-friendly plants (see ques­tions 92-94). Since some species are particularly at­tracted to various salts, nitrogenous compounds, and

fermenting fruits or vegetables, placing these items in a specific locale can often prove successful.

101. What kind of equipment do you need to collect and preserve butterflies? (Figure 15)

A net, insect spreading board, insect pins, a stor­age box, forcep~ (tweezers), moth crystals, writing pa­per, storage envelopes, and small storage cartons such as plastic food containers, are all essential for making a butterfly collection. First, capture the butterfly with a net (can be handmade or purchased). Second, pinch the thorax of the butterfly to kill it (some collectors pre­fer dropping the specimen in a jar containing a poison­ous gas (for example, fingernail polish remover), while others suggest storing the specimen in an envelope until it can be placed in a freezer for killing). Third, place the butterfly in a temporary storage envelope. Fourth, place the envelope into a field storage container. Fifth, when you have returned home, place the butterfly on a spreading board and pin it for drying. Use narrow strips of paper to hold the wings in place. Also, be care­ful n?t to touch the upper ( dorsal) surfaces of the wings; manipulate the specimen with forceps and insect pins. Sixth, after 5-6 days of drying remove the butterfly, at­tach a data label (listing locality, date of collection, and your name) and store in a container to which you have added an insect repellent and mold deterrent (moth crystals do fine as does a cut section of "Pest Strip"). Seventh, store in a relatively dry, bug-free environment. Check the box at least once a year. When necessary, renew the moth crystals, otherwise, your collection will be destroyed.

If your specimens are to be displayed in wall mounts such as Riker Mounts, you must remove the body pin from the butterfly immediately after pinning, otherwise, body fluids will dry around the pin and make later removal difficult. Below are the names and addresses of supply houses that can furnish all equip­ment for collecting.

Bioquip Products 17803 La Salle Avenue Gardena, California 90248-3602

(310) 324-0620

FAX (310) 324-7931

-

'. ✓--· >•: . '

t ·-··········-····· ··-· 1 2 3 4

A

I I

--·--·--------B

C

1 2 3 4

Figure 15. Preparation of a butterfly specimen.

A. Preparing a tempory storage envelope (steps 1, 2, 3, 4). B. Spreading board with specimen positioned for pinning. C. Pinning a specimen !steps 1, 2, 3, 4). .

Carolina Biological Supply Company 2700 York Road Burlington, North Carolina 27215 (800) 334-5551 FAX (910) 584-3399

Ward's Natural Science Establishment, Inc. P.O. BOX 92912 Rochester, New York 14692-9012 (800) 962-2660 FAX (800) 635-8439

102. How long can butterflies be preserved?

The splendor of butterflies is usually unmarred by death. Because lepidoptera have exoskeletons, their outside coverings are virtually indestructible if kept from gnawing creatures and mold. Even colors, if not exposed to strong light for extended periods, will re­tain their brightness for an indefinite number of years. Many of the specimens of the great 19th century natu­ralists are still in excellent shape. But as described in question 101, care must be taken to make sure that some form of insect repellent and mold retardant sur­round the specimens at all times.

103. What are trap nets?

A net, cylindrical in shape, with one end open, and from which is suspended by an inch or two (2-5 cm) a simple platform to serve as a feeding tray is called a TRAP NET. The apparatus is usually hung from a limb in a tree after it has been baited with a butterfly-friendly "cocktail" (see question 98). Many butterfly species within the tropics find baits alluring. The butterflies fly to the feeding platform, crawl through the open­ing, and feed. When satiated, they fly upward, and are then unable to escape. Trap nets are often the principal means of collecting specific species of butterflies. Of­ten collectors will employ several nets scattered throughout a forested area.

104. Are there any butterfly clubs or societies in the United States?

Yes. Following are the names and addresses of as­sociations for both professional and amature lepidop­terists.

North American Butterfly Association, Inc. 39 Highland Avenue Chappaqua, New York 10514 Publications: A111erican ~utterfiies and The Anglewing

Southern Lepidopterists' Society c/o Mr. Thomas M. Neal, Secretary-Treasurer 1705 NW 23rd Street Gainesville, Florida 32605 Publications: Southern Lepidopterists' News and Southern Lepidopterists' Bulletin

The Association for Tropical Lepidoptera, Inc. c/ o Florida State Collection of Arthropods P.O. Box 141210 Gainesville, Florida 32614-1210 Publications: Tropical Lepidoptera, Ho/arctic Lepi­doptera, and Tropical Lepidoptera News

The Lepidoptera Research Foundation, Inc. 9620 Heather Road Beverly Hills, California 90210 Publications: The Journal of Research on the Lepi­doptera and News Letter

The Lepidopterists' Society c/ o L.A. County Museum of Natural History 900 Exposition Blvd. Los Angeles, California 90007-4057 Publications: Journal of the Lepidopterists' Society and News of the Lepidopterists' Society

The Xerces Society 10 Southwest Ash Street Portland, Oregon 97204 Publications: Wings and Atala

t){i!? AESTHETICS AND SYMBOLISM

105. Which is the most beautiful butterfly in the world?

Beauty depends upon personal aesthetics. How­ever, certainly the enormous and brilliant bird wing but­terflies of the Pacific Islands and the iridescent blue morphos of the American tropical rainforests rank high on the lists of most aficionados. Most lepidopterists con­sider the Madagascan sunset moth (Urania ripheus or Chrysiridia madagascariensis) to be the most beautiful of all the lepidoptera. This species is multi-colored, highly iridiscent, and resembles a swallowtail butterfly both in form and in flight. In addition the moth flies during the day.

106. Which is the most beautiful butterfly in North America?

Once again, personal aesthetics are involved. Many collectors, however, consider the diana (Speycria diana), a medium-sized fritillary found principally in the Ap­palachian Mountains, the most handsome. Males are vibrant orange and brown, females pale blue and black. The monarch (Danaus plexippus), is probably the most venerable butterfly in the western hemisphere.

107. How is the butterfly symbol used?

Throughout history many cultures have been in­spired to accord special status to butterflies. Perhaps it is their extraordinary coloration, their etherial habits, or their remarkable metamorphic life cycle. But what­ever the reason, butterflies appear prominently in the folk talcs, myths, and spiritual teachings of practically all cultures. The symbol has frequently been used to represent spring, freshness, airiness, freedom, rebirth, renewal, love, hope, and change. Some examples: the sighting of a butterfly is often regarded as a good omen, but a black butterfly supposedly brings ill health or even dl'ath; thl' Christian church employs the symbol on Easter Sund:1y to represent the resurrection of Jesus Christ, and i\w Christian church also uses the symbol to denote the <1cccptcincc of Christ into one's liftc'; many 1\meric,n, indigcnl)US cultures use the butterfly-human motif tP rt·\·crv the dc,1d (see question l 08); the Hos­pice Founcbtiun us..:·s the symbol as its logo to repre­sent hope' ,ind love; the Compassionate Friends, Inc.

also utilizes the symbol to represent hope and peace; environmentalists employ the symbol to represent a clean, healthy environment; and the phrase "Butterflies are Free" has become not only a popular adage but the title of a successful play and movie as well.

108. What is the significance of the humanoid butterfly motif so common in pre-Hispanic American art? (Figure 16)

Many indigenous cultures throughout the Ameri­cas revere butterflies because of their supposed link to the dead: butterflies represent the actual spirits of the departed. For that reason, the butterfly is often depicted with humanlike features: perhaps a head, or a hand. These motifs are particularly prominant in the Aztec, Maya, Mixtec, and Zapotec cultures of Mexico and Central America.

109. Why are butterflies sometimes referred to as "dancing flowers?"

The term has its origin with the Papago Indian cul­ture of southwestern Arizona. The Papagos believe that when the world was created, it was without joy. To rem­edy this situation Elder Brother gathered the parts of flowers, leaves, corn pollen, sunshine, and other color­ful things. These were then placed into a bag. Children were summoned, the bag shaken, and then opened. Out flew the most beautiful creatures in all of creation. The children leaped with joy at the dancing bits of colors, similar to the earthbound flowers. They yelled "Danc­ing Flowers." And to this day, the term for butterfly in the Papago language translates as "Dancing Flm:-ers."

42

)

A

(, '

C

t '

Culture) (Aztec

Culture). (Aztec

43

B

c9J & c9J

.• Culture) (Aztec

Figure 16. The butterfly-human motif in pre­Hispanic Mexican Art.

GLOSSARY

aedeagus--the male sex organ (penis) in insects; usu­ally enclosed within a pair of claspers

aestivation--inactivity during summer or periods of continued high temperatures; also during a dry sea­son

alar--relating to wings

albinism--absence of color in a form that is usually col­ored

alimentary canal--the tubular passage that extends from mouth to anus and functions in digestion and ab­sorption of food and elimination of residual waste; in insects consists of mouth, pharynx, esophagus, crop, midgut, hindgut, and rectum

androconia--scent scales found commonly in male but­terflies; produce pheromones involved in courtship; ex­amples include hairpencils, stink clubs, andraconial patches (sometimes referred to as alar patches), andraconial tufts, and andraconial folds

antenna (pl. antennae)--the paired segmented sensory organs (olfactory) borne one on each side of the head of most adult insects; commonly termed horns or feel­ers

aorta--in insects, the anterior, non-chambered, narrow part of the heart or dorsal blood vessel, opening into the head

apiculus--an erect, fleshy, short point; typically, the appendage of the antennae in skippers

aposematic--warning colors

arachnid--a member of the Class Arachnida; charac­terized by 4 pairs of legs, 2 body parts, and no anten­nae; includes spiders, scorpions, pseudo-scorpions, mites, and ticks

arthropod--a member of the Phylum Arthropoda; char­acterized as invertebqtt~ with bodies divided into

"» p ''"'»(

ments bearing jointed appendages, exoskeletons, and an open circulatory system

Binomial System of Nomenclature--a system of nam­ing biological species standardized by Carolus Linnaeus in 1758, and consisting of a two- part (genus and species) name in Latin

bursa copulatrix--in insects, the female reproductive organ that receives the aedeagus of the male; the va­gina in female insects

butterfly (pl. butterflies)--insects in the Order Lepi­doptera, Superfamily Papilionoidea; characterized by having clubbed antennae, scaly wings, and a pupa called a chrysalis

"butterfly hotel" --a hibernation box, usually of wood with vertical slits, employed to provide a safe location for the winter; these also provide secure locations for pupating caterpillars

camouflage--a disguise for concealment

carnivorous--flesh-eating

caterpillar--in general, the larval stage of butterflies, skippers and moths; characterized by having three pairs of true legs and usually five pairs of prolegs; wormlike and often ornamented with hairs and spines, some of which can inflict painful stings

chitin (also sclerotin)-- a tough nitrogenenous polysac­charide intermediate between proteins and carbohy­drates, secreted by the epidermis, and the chief com­ponent of the exoskeleton of arthropods

chrysalis, chrysalid (pl. chrysalises, chrysalids, chry­salides)--the naked pupa of a butterfly that is attached to a substrate by silken threads (often suspended), and frequently with a silken girdle

circulatory system--a pathway for the movement of blood throughout the body of an animal; two types: open, in which the blood pools within the haemocoel,

closed, in which the blood is confined to a net-

-

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' /

~ ~

~ . • i: ,.-(-'. ., ~ ',,. ..

)

work of arteries and veins

claspers--part of the male genitalia used for gripping the female during copulation; synonomous with val­vula and harpe

cocoon--a covering composed wholly or partially of silk fibers spun by larvae of most moths and many skip­pers; attached to a substrate and usually concealed; encloses the pupa

coloration--the state of being colored; two types in the Lepidoptera: pigmental or chemical, due to specific chemicals, and structural, due to the reflection and re­fraction of light (as with iridescence)

copulate--to mate

cremaster--the tip of the last segment of the abdomen; the anal hooks by which many pupae suspend them­selves

crepuscular--active during twilight (dusk and dawn)

crochets--the curved spines or hooks on the prolegs of caterpillars and on the cremaster of pupae

cuticle--the outer covering of an insect, formed of a non-cellular layer of chitin

diapause--a condition of suspended animation

dimorphism--a difference in form or color between in­dividuals of the same species; may be seasonal, sexual, or geographical

diurnal--active during the day

dorsal--the upper or back surface

ecdysis--the process of casting the skin; molting; oc­curs several times during the de\'elopment of larvae

eclosion--emergence of the adult from the pupa; the hatching of an egg

entornology--the branch of zoology that deals with the scientific study of insects (and often other arthropods as well); a person who studies entomology is termed an entomologist

exoskeleton--the tough outside covering of all arthropods; although this is periodically molted in immature forms, in adults the exoskeleton is perma­nent

exuviurn (pl. exuvia, exuviae)--the cast skin of larvae at metamorphosis

food chain--a sequence of organisms in which each uses the next lower member of the sequence as a source of food

food plant--a plant used by an insect for its source of nourishment

food web--a series of interconnected food chains

frass--solid larval insect excrement

ganglion (pl. ganglia)--a nerve center composed of a cell mass and fibers; the brain in butterflies is nothing more than a ganglion

genitalia--the reproductive structures of animals; of­ten very distinctive in the Lepidoptera and used in their classification

haernocoel--the blind body cavity in which the blood pools; common to all arthropods

haernolyrnph--the watery blood (usually clear to yel­lowish) in insects and in most other invertebrates

herbivorous--plant eating

hibernaculurn--any place of hibernation, for example, a curled up leaf, or a man-made "butterfly hotel"

hibernation--a period of lethargy or suspended ani­mation in animals occurring during seasonal low tem­peratures

45

homeotherm--an animal that is "warm-blooded," that is, has an internal mechanism for maintaining a con­stant body temperature; for example, birds and mam­mals

host plant--a plant used by an animal for reproduc­tion; with butterflies it is the plant on which females lay their eggs, and is usually different from the food plants

imago (pl. imagos, imagines)--the adult form of an in­sect; for example, a butterfly; imagos do not grow but are principally involved with reproduction

insect--all members of the Class Insecta; characterized by having three body parts (head, thorax, abdomen), three pairs of legs, one pair of antennae, and undergo some form of metamorphosis

instar--the period or stage between molts in the larva, and numbered to designate the various periods; for example, the first instar is the stage between the egg and the first larval molt

labium (pl. labia)--a compound structure that forms the floor of the mouth in mandibulate insects, behind the maxilla

larva (pl. larvae)--a young insect that hatches from an egg in an early stage of development, and differs fundamentally in form from the adult; an immature insect

lateral--relating to the side surface

lepidoptera--insects within the Order Lepidoptera, from the Latin meaning "scaly wings," referring to the powder-like scales on the wings of butterflies, skippers, and moths

Malpighian tubules--long and slender blind excretory tubes lying in the haemocoel of insects, and opening into the large intestine; the insect's urinary system

mandibles--the first pair of jaws in insects; in caterpil­lars, the primary chewing structrues

maxilla (pl. maxillae)--the second pair of jaws in man­dibulate insects; the most persistent structure when the mouth is modified (such as in the proboscis of a but­terfly)

melanism--an abnormal darkening in a form that is not usually dark

metamorphosis--the series of changes through which an insect passes in its growth from egg through larva, pupa, and finally adult; two types: complete or holom­etabolous, in which there are 4 distinct stages (egg, larva, pupa, adult), and incomplete ( simple) or hemi­metabolous, in which only three stages (egg, nymph, adult) occur (as the nymphs grow and molt they gradu­ally assume more and more of the characteristics of the adult as in grasshoppers, roaches and bugs)

migration--the process of leaving a geographic loca­tion and later returning to that same area; for example, the monarch butterfly is the only true migrant butter­fly in North America; by contrast, the one-way move­ment from a location without a return is termed emi­gration; the one-way movement into a new location is termed immigration; for example, ,,several species of butterflies in North America emigrate each fall but do not return the following spring (examples include the cloudless sulphur, gulf fritillary, buckeye, painted lady, little sulphur)

mimic--an organism that closely resembles another of a different group

mimicry--the superficial resemblance of one organism to another or to a natural object within its environment for concealment, protection, or other advantage; in butterflies many species that are palatable to preda­tors mimic others that are unpalatable and thereby gain protection when predators are unable to make the dis­tinction (this type of mimicry is termed Batesian Mim­icry and is exemplified by the monarch and viceroy); in butterflies many species that are unpalatable con­verge on a single striking coloration that easily rein­forces the mimetic pattern (this type of mimicry is termed Miillerian Mimicry and is exemplified by com­plexes of longwings, clearwings, sulphurs, swallow­tails, and brushfoots within the tropical rainforests)

4:6

r 'I

model--the distasteful species mimicked by an inoffen­sive one

myrmecophily--relating to ants (as in myrmecophil­ous caterpillars)

monophagous--feeding on only one kind of food (as in one species or genus)

moth--insects in the order Lepidoptera, Suborder Heterocera; characterized by having antennae that are either straight and tappered or straight and feathery (no clubs), and a pupa either enclosed within a cocoon or else naked and buried within the soil

nectar--the sweet liquid secreated by nectaries within the flowers of many plants; the primary food of most lepidoptera

nocturnal--active at night

ocellus (pl. ocellae, ocelli)--a minute simple eye or eyespot in an invertebrate; for example, caterpillars have six pairs of ocelli; a spot of color encircled by a band of another color; for example, owl butterflies (ge­nus Caligo) have prominent ocelli

ommatidium (pl. ommatidia)--one of the visual ele­ments that compose the compound eye of an arthro­pod

organism--a living biological entity

osmeterium (pl. osmeteria)--fleshy, tubular, eversible processes producing a penetrating odor, and capable of being projected through a slit in the first thoracic segment of certain caterpillars in the family Papilionidae (swallowtails)

ostium (pl. ostia)--small openings; in insects, the slit­like openings into the heart; also, in the lepidoptera, the external female genitalic opening

oviparous--egg-laying

oviposition--the process of egg laying

ovoviviparous--producing live offspring from eggs that have been retained by the female

ovum (pl. ova)--an egg

palpus (pl. palpi, palps)--a mouth feeler; tactile, seg­mented structures borne by the maxillae (maxillary palpi) and labium (labial palpi)

parasitoid--a parasite that lives in its victim and even­tually devours it

peritrophic membrane-the sausage-like thin tissue that encases the ingested food of all insects; secreted at the proximal end of the midgut and reabsorbed in the rec­tum

pheromone--a chemical substance released by one or­ganism and which stimulates a response in another, usually of the same species; for example, sex phero­mones are released by many male butterflies from androconia, and these act as aphrodisiacs on females, thereby stimulating them to mate

phytophagous--plant-eating

poikilotherm--a "cold-blooded" animal, that is, one that has no internal mechanism for maintaining its body temperature; all animals except birds and mammals are poikilothermic

pollen--the yellowish (usually) granules or powder-like substances produced by the male organ (anther) of a flower, and which contains the male reproductive cells (sperm); used by some longwing butterflies as a food supplement

pollination--the process by which pollen is transferred from one flower to another; butterflies and skippers are excellent daytime pollinators, moths have the same function during the night

polyphagous--eating many kinds of food

proboscis (pl. proboscises, proboscides)--any ex-

47

tended mouth structure such as the coiled feeding tube of the lepidoptera; the tongue of the lepidoptera

proleg--any process or appendage that serves the pur­pose of a leg; for example, most caterpillars have in addition to three pairs of true clawed legs, four pairs of fleshy, unjointed, suction-like prolegs on the ventral surfaces of their mid abdomen and one pair at their extreme posterior

puddling--social behavior in the lepidoptera in which several insects (usually males) congregate at a damp area rich in minerals, salts, or nitrogenous chemicals in order to imbibe

pupa (pl. pupae)--the resting inactive instar in all in­sects with complete metamorphosis; the intermediate stage between a larva and an adult; the chrysalis of a butterfly

receptaculum seminis--in female insects the spermath­eca or chamber for receiving sperm

scale--a flat variously shaped unicellular outgrowth of the exoskeleton of insects; for example, the scales of butterflies give the insects their characteristic colors and designs and are responsible for producing pheromones; most scales are flat but some, particularly those close to the body, are often long and hair-like and act as heat traps; some scales are specifically designed to break up light thereby creating metallic or iridescent hues

skipper--insects of the Superfamily Hesperioidea; char­acterized by having clubbed antennae with hooks, heavy, "furry" bodies, large heads, and a pupa that is often within a loosely woven silken cocoon-like struc­ture; closely allied to the butterfly Superfamily Papilionoidea, and often included with butterflies in research

species--a group of similar organisms that are capable of reproducing among themselves in nature; each bio­logical species has a two-part name in Latin

specimen--a living or dead biological enitity

spermatophore-a covering or capsule about the sperm

sphragis--in certain lepidoptera (parnassians or apollos) the large valves of the male that become de­tached and remain with the female, forming a plug to the female copulatory opening, thus preventing a sec­ond mating

spinneret--in the lepidoptera, the larval apparatus for spinning silk

spiracle--the external opening into the respiratory sys­tem of arthropods; one pair is located on the thorax and eight pair are on the abdomen of a caterpillar or butterfly

stadium (pl. stadia)--the interval between the molts of larvae; instar; any one period in the development of an insect

subcosta vein--the first longitudinal vein on the wings of the lepidoptera just inside the margin

symbiosis--the condition in which different organisms live together in more or less intimate association; em­bodies mutualism (both organisms benefit), commen­salism (one benefits, the other doesn't but is not harmed), and parasitism (one benefits, the other is harmed)

thermoregulation--the physical process by which an organism adjusts its body temperature; butterflies fre­quently vibrate or slowly open and close their wings to adjust their body temperatures

trachea (pl. tracheae)--a spirally ringed internal elas­tic air tube in insects; an element of the respiratory sys­tem of insects

tympanic membrane, also tympanum (pl. tympana­a membrane usually on each side of the third thoracic segment or the base of the abdomen, presumed to be an auditory organ

-

1-

( C

J I'

uncus (pl. unci)--the curved hook directed downward at the posterior of the male abdomen, shielding the penis, and used during mating along with the claspers to secure the female

urticating--causing a stinging or burning sensation to the skin; the larvae of several species of moths possess urticating or urticarial hairs or spines (no species of butterfly in North America is so endowed)

ventral--relating to the undersurface

valvula (pl. valvulae)--the claspers in the male geni­talia of the lepidoptera (often termed valva (pl. valvae) or simply valves

viviparous--giving birth to live young

voltinism--referring to the number of generations or flights in a given year; embodies univoltine (one gen­eration), bivoltine (two generations), trivoltine (three generations) and biennial (two years to complete one generation)

worm--any of numerous relatively small elongated usually naked and soft-bodied animals without legs; examples of true worms are the earthworm, tapeworm and a host of other parasitic species; although caterpil­lars are often referred to as worms, they are not

49

Aztec butterfly stamp from Teotihuacan, Mexico, from the book Design Motifs of Ancient Mexico.

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