California Butterflies
By John S. Garth and J. W. Tilden
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About this ebook
Featuring nearly all species known in California, this beautifully illustrated guide provides detailed insights into the habitat, distribution, flight periods, and life stages of these delicate insects. Readers will find practical tips on observing butterflies in the wild, recording meaningful scientific observations, and even collecting and preserving specimens for further study.
With contributions from renowned illustrators and experts, California Butterflies is perfect for those looking to deepen their understanding of this fascinating group of insects. Whether you’re captivated by the dainty Pygmy Blue or the majestic Two-tailed Swallowtail, this guide opens the door to a vibrant and colorful corner of California’s natural heritage. Complete with a checklist of both scientific and common names, as well as references for further study, this field guide is the definitive resource for exploring the rich variety of butterflies found across the Golden State.
This title is part of UC Press's Voices Revived program, which commemorates University of California Press's mission to seek out and cultivate the brightest minds and give them voice, reach, and impact. Drawing on a backlist dating to 1893, Voices Revived makes high-quality, peer-reviewed scholarship accessible once again using print-on-demand technology. This title was originally published in 1986.
John S. Garth
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California Butterflies - John S. Garth
California
Butterflies
California Natural History Guides: 51
California
Butterflies
John S. Garth
and
J. W. Tilden
Illustrated by
David Mooney and Gene M. Christman
UNIVERSITY OF CALIFORNIA PRESS
Berkeley Los Angeles London
CALIFORNIA NATURAL HISTORY GUIDES
Arthur C. Smith, General Editor
Advisory Editorial Committee:
Raymond F. Dasmann
Mary Lee Jefferds
A. Starker Leopold
C. Don MacNeill
Robert Omduff
Robert C. Stebbins
University of California Press
Berkeley, California
University of California Press, Ltd.
London, England
© 1986 by
The Regents of the University of California
Printed in the United States of America
10 987654321
This book is printed on acid-free paper to ensure
its permanence and durability.
Library of Congress Cataloging in Publication Data
Garth, John S. (John Shrader), 1909- Califomia butterflies.
(California natural history guides; 51) Bibliography: p.
Includes index.
1. Butterflies—California— Identification. 2. Insects—Identification. 3. Insects—California— Identification. I. Tilden, J. W. II. Title. III. Series.
QL551.C3G37 1985 595.78'9'09794
84-28071
ISBN 0-520-05249-8
ISBN 0-520-05389-3 (pbk.)
Dedicated to
the past, present, and future
Lepidopterists of our State
On Discovering a Butterfly
1 found it in a legendary land All rocks and lavender and tufted grass, Where it was settled on some sodden sand Hard by the torrents of a mountain pass.
I found it and I named it, being versed In taxonomic Latin; thus became Godfather to an insect and its first Describer—and I want no other fame.
—Vladimir Nabokov, Quoted in The Magnificent Foragers (Smithsonian Institution)
CONTENTS
CONTENTS
1 • STRUCTURE, BEHAVIOR, AND DISTRIBUTION
Butterfly Structure and Growth
How Butterflies Behave
How Butterflies Depend on Plants
Butterfly Distribution
2 • OBSERVING AND COLLECTING BUTTERFLIES
How to Make a Butterfly Collection
History of Butterfly Collecting in California
Variation and Hybridization in Butterflies
How to Tell a Butterfly from a Moth
3 • BUTTERFLY IDENTIFICATION
How Living Things Are Classified
Butterfly Name Changes
Key to the Butterfly Families of California
4 • CALIFORNIA BUTTERFLIES: SPECIES ACCOUNTS
Abbreviations
Satyrs, Arctics, and Ringlets (Family Satyridae)
Milkweed Butterflies (Family Danaidae)
Long-wings (Family Heliconiidae)
Brush-footed Butterflies (Family Nymphalidae)
Swallowtails and Parnassians (Family Papilionidae)
Whites, Sulfurs, Marbles, and Orange-tips (Family Pieridae)
Snout Butterflies (Family Libytheidae)
Metalmarks (Family Riodinidae)
Hairstreaks, Coppers, and Blues (Family Lycaenidae)
Giant Skippers (Family Megathymidae)
Skippers (Family Hesperiidae)
CHECKLIST OF CALIFORNIA BUTTERFLIES
GLOSSARY
SOURCES
INDEX OF COMMON AND SCIENTIFIC NAMES
GENERAL INDEX Including Food Plants
1 • STRUCTURE, BEHAVIOR,
AND DISTRIBUTION
Butterfly Structure and
Growth
How a Butterfly’s Body is Organized
The bodies of insects, including butterflies, are composed of ringlike sections called segments. The body of a butterfly, like that of other insects, is formed of three sections—head, thorax, and abdomen. The segments of the abdomen are easily visible. Those of the head and thorax are largely fused and are much less easy to distinguish. The appendages of the insect body, including mouthparts, legs, and wings, occur in pairs, one pair to each segment that bears them.
The head bears the mouthparts and the antennae (feelers), as well as the many-faceted compound eyes, which are not considered to be appendages. The mouthparts of a butterfly consist of a coiled tube, the proboscis, a paired structure in which the halves fuse, as shown in Figure 1. This structure, by which the butterfly sucks up liquids, lies between two scaly or hairy protruding structures, the palpi. The eyes occupy much of the sides of the head. The eyes of butterflies are good at perceiving movement and are also able to see color, including ultraviolet, which man cannot see.
The thorax, the section next behind the head, is formed of three segments, each of which bears a pair of legs. The thoracic legs are jointed, as are those of all arthropods, a group of organisms that includes spiders, crabs, scorpions, and others, as well as insects. The parts of a usual butterfly leg are shown
FIG. 1 Head with named parts
FIG. 2 Leg with named parts
and named in Figure 2. The second and third thoracic segments each bear a pair of wings as well. The front wings are usually much larger than the hind wings and have somewhat different venation.
The abdomen of the adult does not bear either legs or wings. At the tip of the abdomen are located the external reproductive organs, which are the claspers (valves) in the male, and a transverse opening in the female. The abdomen of the male is more slender than that of the female, especially if the female is gravid (full of eggs).
Inside the abdomen are located the digestive organs and the internal reproductive organs, the testes in the male, the ovaries in the female.
Along the sides of the body are segmental openings called spiracles, which lead into a series of internal tubes called tracheae, which bring air directly to the internal organs.
The larva, or caterpillar, is very different in form from the adult. The mouthparts are chewing jaws (mandibles) and a lower lip
(labium) containing a spinneret, from which is spun the silk used by the larva at pupation. The eyes consist of several simple eyes on each side of the head; there are no compound eyes. The three thoracic segments bear the true legs, which are quite short. The third, fourth, fifth, sixth, and last abdominal segments bear what are called prolegs. These function as legs, and have at their outer ends small hooks called crochets, which help the caterpillar in climbing. These are lost when the insect sheds its last larval skin and becomes a pupa, or chrysalis. The structures of a caterpillar are shown in Figure 3.
How a Butterfly Grows
While man and other vertebrate animals grow, develop, and reproduce without great change in bodily form, many invertebrates, including butterflies, devote an entire stage of their life to each of these activities, with successive changes in body form. The caterpillar, hatching from the egg, feeds and grows, storing up energy for the next stages. The quiescent pupa, or chrysalis, is a reorganizational stage, from which the adult emerges. The adult is the reproductive stage. It feeds on fluids only, the energy from which supports reproductive activities.
The eggs of butterflies are of many forms. Four are shown
FIG. 3 Caterpillar with named parts
FIG. 4 Types of eggs
in Figure 4. Most butterfly eggs have minute grooves, pits, or ridges on their surface, a condition spoken of as sculpture. The eggs are deposited by the female either singly or in clusters, usually on twigs, leaves, or buds of the plant on which the caterpillar feeds. In some cases, such as that of the fritillaries, which feed in the larval stage on violets, the female may never see the plants on which her larvae will feed, since the tops of the plants will have dried up before she emerges as an adult. She deposits her eggs on the ground where the violets will grow the following spring, and when the larvae hatch from the eggs, they find the new growth by themselves.
The chewing mouthparts of the larva work from side to side, not up and down. The larva eats and fills out its body until it seems it would burst. This is exactly what happens to the tightly packed outer skin, but a new and more flexible skin has been forming under the old one. When the old skin splits, the larva wriggles out of it and expands to a much larger size while the new skin is still able to stretch. This skin soon becomes hardened, and the larva resumes feeding until the next change of skin takes place. Each change of skin is called a molt. The time between each molt is called a stadium, and the larva itself at each stage is called an instar. Caterpillars pass through several instars before reaching the next change in form, pupation.
When the larva has reached the maximum size for its species, it crawls away, often entirely away from the food plant, and finds a sheltered place, where it will change into the pupa, in butterflies also called a chrysalis. Different families of butterflies pupate in different ways. The pupa described here and shown in Figure 5 is of the Monarch or Milkweed Butterfly.
On the lower lip, or labium, of the larva is a tiny tube, the spinneret, from which liquid silk, from a silk gland, can be spun. From the spinneret, the larva first weaves a button of silk. As the last larval skin splits, it frees the tip of the pupal abdomen. This tip has a cluster of small spines, the cremaster, and these spines hook into the silk button, so that the pupa now hangs head down, suspended by the cremaster only.
FIG. 5 Pupation of Monarch
Swallowtails and members of the family Pieridae are also attached to the cremaster at the tip of the abdomen, but with the head up and leaning against a supporting silken belt. Some Satyridae and the pamassians pupate at the surface of the ground or in shallow earthen cells. Lycaenidae and Riodinidae usually pupate flat to a surface, but still have the anal cremaster and often a silken belt also. Skippers usually form a slight silken shelter around themselves, whereas giant skippers usually pupate in burrows in their food plants. The pupa resembles neither the caterpillar from which it developed nor the adult which will emerge from it. It is the reorganizational stage inside of which the larva changes into the adult.
After a period of as little as two weeks in some manybrooded species, or as long as a year or more in certain singlebrooded species, the final change from pupa to adult, or imago, takes place. The chrysalis ruptures, and the winged insect slowly emerges. At first the wings are small; then as blood is pumped through their veins, they lengthen and hang loosely. Further inflation brings them to full size. The flow of blood stops, the wings become firm and stiffen until able to support the butterfly in flight. Meanwhile the butterfly has crawled from its place of concealment to an exposed place from which to leave for sunlit skies.
The process by which a caterpillar transforms into a butterfly is called metamorphosis, a word meaning change in form.
And since a pupa, or chrysalis (resting stage
) is involved, such metamorphosis is said to be complete, as distinguished from incomplete metamorphosis, exemplified by the grasshopper, in which the young resembles the adult except for functional wings.
Butterflies may overwinter in various stages of development. Some overwinter in the egg, hatching as first instar larvae the following spring. In some, the egg hatches during the summer, and the hatchling larva overwinters, becoming active at the end of winter. Others hatch from the egg shortly after the egg is laid, then feed for one or more instars, enter a resting period until the following spring, and then complete development. Some species, such as some of the whites and swallowtails, overwinter as pupae and emerge in the spring. Still others, as the anglewings and the Mourning Cloak, overwinter as adults and lay eggs the next spring. Some species, particularly desert species, may remain dormant in some stage of development for several years until sufficient rain falls to ensure their development through the growth of their food plants.
How to Rear a Butterfly
Butterflies that have flown for some time become worn. To obtain perfect specimens it is desirable to have butterflies that have never flown. This may be done by rearing them. You may follow a female as she flits from place to place, depositing an egg at a time on the chosen food plant. A gravid female may be confined with the food plant and induced to lay her eggs with offerings of flowers or sweetened water. Or you may search a known food plant for the larvae or eggs, or the immediate environs for the pupae, which need only to be kept until the adult emerges. If the larva is to be reared, an assured supply of the food plant is needed.
If a growing plant is used, the caterpillars may be enclosed in a gauze sleeve on the plant. If branches of the plant are used, a variety of cages may be made—shoe boxes or ice cream cartons in which cellophane windows have been made work well. Or a small plant or branch may be placed in a flower pot and covered with a lamp chimney. Examples of rearing cages are shown in Figure 6.
FIG. 6 Rearing cages
If grass or small clippings are to be used, they may be placed in a disposable paper or plastic cup covered with gauze secured by a rubber band. Since cleanliness is important, the cages should be cleaned often, and a fresh cup should be used with each food change. Larvae kept under soiled conditions may develop diseases and die.
Be careful to place your plant cuttings in moist sand, not water, because the caterpillars might crawl into the water and drown.
Small larvae may be transferred from old food to new using a small camel’s hair brush.
Some butterflies do not pupate on the plant and need to be supplied with dead twigs or litter.
Records should be kept of the food plants used, the dates of pupation and emergence, and any unusual observations made.
If you wish to preserve one of the larvae, this may be done by killing it in hot water and preserving it in 70 percent alcohol. The accompanying label should be written in soft lead pencil and inserted into the vial with the specimen. The technique for inflating a caterpillar skin over hot air and mounting it dry on an insect pin is given by Holland (see References at end of book).
How Butterflies Behave
The total knowledge of butterfly behavior is great and is being added to continually. In such a book as this only the barest essentials can be mentioned. It is hoped that the interested student will be stimulated to examine the extensive literature on the subject.
As we have seen, butterflies have four stages of development. Each stage behaves differently from the others, and it has been said that each behaves as though it were an independent organism. The egg produces the larva. The larva feeds and grows to a specific size. The pupa, or chrysalis, reorganizes the larval form into the form of the adult. The adult mates and reproduces, laying eggs at a time and place suitable for the perpetuation of the species. Each stage has its appropriate action.
The egg (see Figure 4). Some butterflies lay their eggs in masses; others deposit a few eggs in one place. Some eggs are laid one to a place on the same plant. Others are laid one to a plant, after which the female flies to another plant before depositing the next egg. In some species the larva develops in the egg in a short time and hatches. In others, the development of the larva is delayed for some time. In yet others, the larva develops in the egg quite soon, but remains inside the egg shell until the conditions are right for it to hatch. A few examples follow.
The Bay Region Checkerspot lays its eggs in masses, on or near the food plants. Fritillaries of the genus Speyeria lay their eggs in the vicinity of the dry food plants, or even drop their eggs from the air.
The Fiery Skipper often lays its eggs on lawn grasses, carefully placing each egg on the underside of a grass leaf. Blues of the genus Euphilotes lay their eggs on the same flower heads from which the adults have nectared, and on which the larvae develop. They are truly one-plant insects, the plants being various species of wild buckwheat (Eriogonum).
Perhaps the most remarkable, some populations of Lindsey’s Skipper lay their eggs on a lichen (Usnea florida) that grows on tree trunks and wooden fences. The larva emerges in the spring, makes its way down to the ground, and finds its own food plants, which are Blue Bunch Grass (Festuca idahoensis) and California Oat Grass (Danthonia calif arnica).
The larva. Larvae (Figure 7) may be brightly colored or inconspicuous. Brightly colored larvae are often distasteful to would-be predators and are said to show warning coloration. Inconspicuous (cryptic) larvae are usually edible, and their dull coloration provides some measure of protection. Some larvae live in shelters of their own making; others hide in duff on the ground. Some feed openly in the daytime; others hide by day and feed by night
Larvae may feed on a single species of food plant, or on several related plants, or on unrelated plants that have a similar chemical makeup, or, as do some species, on a wide variety of unrelated food plants.
The larvae of some species of butterflies develop directly, feeding and growing continuously until full sized. Other spe-
FIG. 7 Representative larvae
cies feed and grow for a while, then enter a state of suspended activity (diapause) that will carry them over an unfavorable part of the year. Usually the diapause is in winter, but some species have a summer diapause, their food being too dry or tough at that time. Many species overwinter as hatchlings and develop the following spring.
The Monarch caterpillar is ringed with yellowish and greenish gray, and is distasteful. It feeds openly on its food plants, various milkweeds (Asclepias), and often pupates on the same plant. These caterpillars usually feed at some distance from one another.
The West Coast Lady caterpillar feeds on many members of the Mallow Family (Malvaceae). It makes a shelter by drawing the edges of the leaf together with silken threads. It does not have definite broods. The larvae can be found on the plants during most of the year. Larvae may be present in moderate climates even in winter and will feed whenever the weather is warm.
The blackish caterpillar of the Common or Chalcedon Checkerspot feeds openly on various members of the Figwort Family (Scrophulariaceae). After hatching in the late spring or early summer, the young larvae feed for a while until their food plants begin to dry up, then enter a long overwintering diapause and resume feeding the following spring.
The caterpillar of the Woodland Skipper hatches from the egg in the fall and overwinters as a hatchling. Ihe caterpillar lives in a silken tube in a rolled-up leaf of tall woodland grasses. It reaches full size in late May, goes into a summer diapause until late July or early August, then pupates and soon emerges as an adult.
The pupa (chrysalis) (Figure 8) is usually cryptically colored, and is often difficult to see. The method of attachment of the members of the various families was described above.
FIG. 8 Representative pupae
The chrysalis of the Anise Swallowtail may be either brown or green. The adult may emerge the same season, or may wait one or two years before emerging. The chrysalis of the Common Checkerspot is light gray marked with many tiny dark lines and dots. It hangs either on its food plant or on nearby vegetation. The chrysalis of the Gulf Fritillary is strange looking, the abdomen slender, the anterior part wide and flattened. It can twist slightly, possibly to present its narrower edge to direct sunlight.
The adult (imago). Adult butterflies recognize one another during courtship and mating by color, some combination of markings, manner of movement, and odor. Different species emit different attractant odors (pheromones). The mating habits of various species differ widely. Some butterflies mate early in the day, other species at times ranging from midmoming to early evening. Of the many strategies by which butterflies meet and mate, five are mentioned here.
Bay Region Checkerspot males course back and forth over a field where females will emerge and discover and mate with them, often before the wings of the females have fully expanded.
Males of Lorquin’s Admiral perch on a stick or branch and at intervals fly out, returning to the same or a nearby perch. The same male will occupy the same territory for days, perhaps for life. Any large insect that flies by will be pursued. Almost any kind of butterfly will be followed, and often touched, but mating is attempted only when a female Lorquin’s Admiral is encountered. A male may pursue a female for some distance. Odor is most likely the final factor in recognition. Incidentally, it is amusing to note that male admirals will follow to the ground a chip, small rock, or clod thrown by an observer.
The male Buckeye is even more aggressive. He will closely pursue any large insect that crosses his territory, often chasing it out of the area. When a receptive female enters his territory, the male follows her very closely. Often the pair may flutter high into the air, returning to the ground to mate.
The female Woodland Skipper sits quite openly, usually on the ground, and vibrates her wings at a fixed rate. A passing male will make several flights over her, approaching closely.
Eventually she tips her abdomen up, and it is possible that a pheromone is released at this time. The mating that ensues lasts for some