Chapter V: Introduction (1)

In popular esteem the butterflies among the insects are what the birds are among the higher animals--the most attractive and beautiful members of the great group to which they belong. They are primarily day fliers and are remarkable for the delicacy and beauty of their membranous wings, covered with myriads of tiny scales that overlap one another like the shingles on a house and show an infinite variety of hue through the coloring of the scales and their arrangement upon the translucent membrane running between the wing veins. It is this characteristic structure of the wings that gives to the great order of butterflies and moths its name Lepidoptera, meaning scale-winged.

In the general structure of the body, the butterflies resemble other insects. There are three chief divisions: head, thorax, and abdomen. The head bears the principal sense organs; the thorax, the organs of locomotion; and the abdomen, the organs of reproduction.

By examining a butterfly's head through a lens it is easy to see the principal appendages which it bears. Projecting forward from the middle of the top is a pair of long feelers or antennae. Each of these consists of short joints which in general may be divided into three groups: first, a few large joints at the base connecting the feeler with the head; second, many rather small joints which make up the principal length; third, several larger joints which make up the outer part or "club" of the antennae. In the case of the Skippers, there are in addition a number of small joints coming to a sharp point at the end of the club. Just below the insertion of the antennae on each side of the head are the large compound eyes, which are almost hemispherical. With a powerful glass, one can see the honeycomb-like facets, of which there are thousands, making up each eye. Just below the eyes there are two hairy projections, called the palpi, between which is the coiled tongue or sucking tube. (_See plate, page 64-65._)

Anatomically the thorax is divided into three parts--the prothorax, the mesathorax, and the metathorax; but the lines of division between these parts are not easily seen without denuding the skin of its hairy covering. The prothorax bears the first pair of legs. The mesathorax bears the front pair of wings and the second pair of legs. The metathorax bears the hind pair of wings and the third pair of legs. In many butterflies, the first pair of legs are so reduced in size that they are not used in walking.

The abdomen is composed of eight or nine distinct rings or segments, most of which have two spiracles or breathing pores, one on each side. It also bears upon the end of the body the ovipositor of the female or the clasping organs of the male.

BUTTERFLY TRANSFORMATIONS

The butterflies furnish the best known examples of insect transformations. The change from the egg to the caterpillar or larva, from the caterpillar to the pupa or chrysalis, and from the chrysalis to the butterfly or imago is doubtless the most generally known fact concerning the life histories of insects. It is a typical example of what are called complete transformations as distinguished from the manner of growth of grasshoppers, crickets, and many other insects in which the young that hatches from the egg bears a general resemblance to the adult and in which there is no quiet chrysalis stage when the little creature is unable to eat or to move about.

_The Growth of the Caterpillars_

Caterpillars are like snakes in at least one respect: in order to provide for their increase in size they shed their skins. When a caterpillar hatches from the egg it is a tiny creature with a soft skin over most of its body but with rather a firm covering for its head. While we might fancy that there could be a considerable increase in size provided for by the stretching of the soft skin it is easy to see that the hard covering of the head will not admit of this. So the story of the growth of a caterpillar may be told in this way:

A butterfly lays an egg upon a leaf. Some days later the egg hatches into a larva, which is the technical name for the second stage of an insect's life. In the case of the butterfly we call this larva a caterpillar. The little caterpillar is likely to take its first meal by eating the empty egg shell. This is a curious habit, and a really satisfactory explanation of it seems not to have been made. Its next meal is likely to be taken from the green tissues of the leaf, commonly the green outer surface only being eaten at this time. The future meals are also taken from the leaf, more and more being eaten as the larva gets older.

After a few days of this feeding upon the leaf tissues the little caterpillar becomes so crowded within the skin with which it was born that it is necessary to have a larger one. So a new skin begins to form beneath the first one. Consequently the latter splits open in a straight line part way down the middle of the back just behind the head. Then the new head covering is withdrawn from the old one and the caterpillar wriggles its way out of the split skin and finds itself clothed in a new one. At first all of the tissues of the new skin are soft and pliable and they easily take on a larger size as the body of the caterpillar expands. A little later these tissues become hardened and no further expansion is possible.

This process of skin-shedding is called moulting. The cast skin is often called the exuviae. The period of the caterpillar's life between the hatching from the egg and this moult is often called a stage or instar--that is, the caterpillar up to the time of this moult is living in the first caterpillar stage or instar.

During the actual moulting the caterpillar is quite active in freeing itself from the exuviae. But as soon as it is free it is likely to rest quietly for some hours while the tissues of the new skin are hardening. Then it begins feeding upon the leaf again and continues taking its meals at more or less regular intervals for several days. By that time it will again have reached its limit of growth within this second skin and the process of moulting must be repeated. It takes place in the same way as before and the caterpillar enters upon the third instar of its larval life.

This process of feeding and moulting is continued for several weeks, the number of moults being usually four. During the later stages the increase in size is more marked each time the skin is shed, until the caterpillar finally reaches its full growth as a larva and is ready for the wonderful change to the quiet chrysalis in which all its caterpillar organs are to be transformed into the very different organs of the butterfly.

In the case of butterfly larvae one of the most interesting features of the growth of the caterpillar is that of the remarkable changes in colors and patterns of marking which the caterpillar undergoes. One who had not followed these changes would often be at a loss to recognize caterpillars of slightly differing sizes as belonging to the same species. These changes commonly show a remarkable adaptation to the conditions of life, and generally tend to the concealment of the caterpillar upon its food plant. The stages of growth of the green-clouded swallowtail caterpillar are illustrated on plate opposite page 80.

Before each moult the caterpillar is likely to spin a silken web upon the leaf surface. It then entangles its claws in the web to hold itself in place while the skin is cast. (_See plate, page 17._)

_The Change to the Chrysalis_
(_See plate, pages 32-33._)

A week or ten days after the last moult of its caterpillar growth the larva commonly becomes full fed and ready to change to the chrysalis state. The details of the way in which this is accomplished vary greatly with different butterflies, as will be noted in the stories of many species later in this book. In general, however, the caterpillar provides a web of silk which it spins against some surface where the chrysalis will be secure and in this web it entangles its hind legs. Sometimes there is the additional protection of a loop of silk over the front end of the body. After the legs have become entangled the caterpillar hangs downward until the skin splits open along the median line of the back and gradually shrinks upward until it is almost free, showing as it comes off a curious creature which has some of the characteristic features of a chrysalis. It is seldom at this stage of the same shape as the chrysalis. When the caterpillar's skin is nearly off this chrysalis-like object usually wriggles its body quickly in a manner to entangle a curious set of hooks attached to the upper end in the web of silken thread. This hook-like projection is called the cremaster, and it serves a very important purpose in holding the chrysalis in position.

As soon as the cremaster is entangled in the web the cast skin usually falls off and for a very short period the creature hanging seems to be neither caterpillar nor chrysalis. It is in fact in a transition stage between the two, and it very soon shortens up and takes on the definite form of the chrysalis, the outer tissues hardening into the characteristic chrysalis skin.

From the fact that this chrysalis skin shows many of the characteristic features of the future butterfly it is evident that the change from the caterpillar to the butterfly really began during the life of the larva. The nature of the process by which this change takes place has long been a puzzle to scientists. For the making of a butterfly is one of the most wonderful phenomena in the outer world, and it has challenged the attention of many acute observers. Some two centuries ago the great Dutch naturalist, Swammerdam, studied very carefully the development of many insects, especially the butterfly. He found that if he placed in boiling water a caterpillar that was ready to pupate or become a chrysalis, the outer skin could easily be removed, revealing beneath the immature butterfly with well-developed legs and antennae. From these observations he was led to believe that the process of growth was simply a process of unfolding; that is, as Professor Packard has expressed it, "That the form of the larva, pupa, and imago preexisted in the egg and even in the ovary; and that the insects in these stages were distinct animals, contained one inside the other, like a nest of boxes or a series of envelopes one within the other." This was called the incasement theory and it was held to be correct by naturalists for nearly a century. It was discredited, however, about a hundred years ago, but not until another fifty years had passed was it definitely replaced by another and much more convincing theory propounded by Weismann.

According to Weismann's theory, which is now well-established, the process of development internally is a much more continuous one than the external changes would indicate. So far as the latter are concerned we simply say that a caterpillar changes to a chrysalis and a chrysalis to a butterfly, the transition in each case requiring but a very short time. Internally, however, it has been going on almost continuously from the early life of the caterpillar. The various organs of the butterfly arise from certain germinal disks or "imaginal" buds, the word "imaginal" in this case being an adjective form of imago, so that the imaginal buds are really simply buds for the starting of growth of the various organs of the imago or adult. As the caterpillars approach the chrysalis period these imaginal buds rapidly develop into the various organs of the butterfly. This process is helped along by the breaking down of many of the tissues of the larva, this broken-down tissue being then utilized for the production of the new organs. About the time the chrysalis is formed this breaking-down process becomes very general, so that the newly formed chrysalis seems largely a mass of creamy material which is soon used to build up the various parts of the butterfly through the growth of the imaginal buds.

_The Change to the Butterfly_
(_See plate, pages 32-33._)

There is probably no phenomenon in the world of living creatures which has attracted more attention than the change of the chrysalis into the butterfly. It is not strange that this is so. We see upon a tree or shrub or wall an inert, apparently lifeless object, having no definite form with which we can compare it with other things, having neither eyes nor ears nor wings nor legs--an object apparently of as little interest as a lifeless piece of rock. A few minutes later we behold it again and note with astonishment that this apparently inanimate being has been suddenly transformed into the most ethereal of the creatures of earth, with an exquisite beauty that cannot fail to attract admiration, with wings of most delicate structure for flying through the air, with eyes of a thousand facets, with organs of smell that baffle the ingenuity of man to explain, with vibrant antennae, and a slender tongue adapted to feeding upon the nectar of flowers--the most ambrosial of natural food. So it is not strange that this emergence of a butterfly has long been the theme both of poets and theologians and that it attracts the admiring attention of childhood, youth, and age.

Fortunately, this change from chrysalis to butterfly may readily be observed by any one who will take a little trouble to rear the caterpillars or to watch chrysalids found outdoors. The precise method of eclosion, as we call this new kind of "hatching," varies somewhat with different species but in general the process is similar in all.

Those chrysalids which have a light colored outer skin are especially desirable if we would watch this process. One can see through the semi-transparent membrane the developing butterfly within, until finally, just before it is ready to break out, the markings of the wings and body show distinctly. If at this time the chrysalis is placed in the sunshine it is likely to come out at once, so that you can observe it readily. It usually breaks apart over the head and the newly released legs quickly grasp hold of the empty skin as well as of the support to which it is attached. It then hangs downward with a very large abdomen and with the wings more or less crumpled up, but decidedly larger than when they were confined within the chrysalis. The wings, however, soon begin to lengthen as they are stretched out, probably through the filling of the space by the body juices. Commonly, the hind pair of wings become full size before the front ones. In a short time the wings attain their full size, the abdomen becomes smaller, through the discharge of a liquid called meconium, and the butterfly is likely to walk a few steps to a better position where it will rest quietly for an hour or two while body and wing tissues harden. After this it is likely to fly away to lead the free life of a butterfly. (_See plate, page 16._)

These changes from larva to chrysalis and from chrysalis to adult in the case of the Monarch Butterfly are illustrated on the plates opposite pages 32-33. A little study of these photographs from life will help greatly to an understanding of the process.

Some very interesting observations have been made by Mr. J. Alston Moffat upon the method of the expansion of the wings. In summarizing his investigations he writes:

"When a wing is fully expanded, and for an hour or two after, the membranes can be easily separated. Entrance for a pin-point between them is to be found at the base of the wing where the subcostal and median nervures come close together. The membranes are united at the costal and inner edges, which have to be cut to get them apart; but they are free at the outer angle. At that time the nervures are in two parts, half in one membrane and half in the other, and open in the centre. The fluid which has been stored up in the pupa enters the winglet at the opening referred to, expanding the membranes as it passes along between them, and the nervures at the same time, and when it has extended to every portion of the wing, then it is fully expanded. The expanding fluid is of a gummy consistency, and as it dries, cements the membranes together, also the edges of the half-nervures, and produces the hollow tubes with which we are so familiar."

BUTTERFLIES AND MOTHS

The butterflies and moths both belong to the great order of scale-winged insects--the Lepidoptera. They are distinguished, however, by certain general characteristics, which hold true for the most part in both groups. The butterflies fly by day; the moths fly by night. All of the higher butterflies go into the chrysalis state without making a silken cocoon, while most of the higher moths make such a cocoon. The bodies of the butterflies are usually slender, while those of the larger moths are stout. The antennae of the butterflies are generally slender and commonly enlarged at the tip into a miniature club. The antennae of the larger moths are commonly feathery or are long and slender, tapering gradually toward the tip.

The characteristic features that distinguish a moth from a butterfly are well illustrated in the plate opposite page 49, which shows one of the largest and most beautiful moths in the world. It is the Cynthia moth. As may be seen, the newly emerged moth is resting upon the silken cocoon in which it spent its period as a pupa or chrysalis. This cocoon was attached by the caterpillar to the twig from which it hangs at the time it spun the cocoon. The feathered antennae, the hairy legs, the thick thorax, and large abdomen--all show very clearly in this side view of the moth. As will be seen, the wings are large and very suggestive of those of a butterfly and have the characteristic eye-spots toward the tip and the crescent marks in the middle, which are so often found on the wings of the larger moths.

Some of these large moths on cloudy days occasionally fly during daylight and, by the uninitiated, they are often mistaken for large butterflies. One who will notice their structure, however, will readily see the characteristic features of the moth.

In the caterpillar stage, there are no hard and fast differences between the larvae of butterflies and those of the higher moths. In each case, the insect consists of a worm-like body, having a small head provided with biting jaws and simple eyes or ocelli. Back of the head are the three rings of the thorax, each of which bears a pair of jointed legs. Back of these three rings there are a considerable number of other body rings making up the abdomen, on the middle of which there are commonly four or five pairs of fleshy prolegs, not jointed but furnished at the tip with fine claws. At the hind end of the body there is another pair of prolegs similar in structure.

THE SCENTS OF BUTTERFLIES

Many students of American butterflies have occasionally mentioned the fact that certain species seem to give off a distinct scent which has frequently been spoken of as a pleasing fragrance, suggesting sandalwood or some other aromatic odor. The general subject as exemplified by butterflies of other lands has been studied for many years by Fritz Mueller; and certain English entomologists have paid considerable attention to it. A translation of the Mueller publications and an excellent summary of our present knowledge of the subject is published in Dr. Longstaff's book on butterfly hunting.

The odors given off by butterflies are divided into two principal kinds, namely: first, those which are repulsive to the senses of man, and evidently for the purpose of protecting the butterflies from birds and other vertebrate enemies--these are found in both sexes; second, those which are evidently for the purpose of sexual attraction and confined to the male butterflies--these scents are usually attractive to the senses of man.

The aromatic scents of the second group are generally produced by means of certain scales or hairs of many curious forms, which are scattered over the surface of the wings or are placed within certain special pockets, generally near the borders of the wings. These scales or hairs are called androconia. Some of them much magnified are represented in the picture above.

Our knowledge of the scents given off by American butterflies is very fragmentary, and it is highly desirable that many more observations should be made upon the subject. If collectors generally would make careful notes, both in the field and upon the freshly killed butterflies at home, we ought soon to be able greatly to extend our knowledge. By holding the butterfly with a pair of forceps, one can often determine whether the fragrance is emitted. It is often helpful also to brush the hairs or tufts where the androconia are attached, using a small, dry camel's hair brush for the purpose.

BUTTERFLY MIGRATIONS

Migration seems to be a general instinct in the animal world, developed when a species becomes enormously abundant. At such times this instinct apparently overcomes all others and the creatures move on regardless of obstacles and conditions that may mean certain death to the vast majority. Such migrations among mammals have often been recorded, one of the most notable examples being that of the little lemmings which migrate at periodical intervals in a way which has often been described. Among the insects such migrations have been frequently noticed, and the phenomenon has apparently been observed oftener among the butterflies than in any other group. Entomological literature during the last hundred years contains a great many records of enormous flights of butterflies over long distances, extending even from Africa into Europe or from one part of America to another far remote. As such migration is likely to happen whenever a species becomes extremely abundant it probably is Nature's way of providing for an extended food supply for the succeeding generations. That it results in the death of the great majority of the migrants is doubtless true, but it must lead to vast experiments in extending the geographic area inhabited by these species. Numerous examples of such migrating swarms will be found in the pages of this little book. (_See plates, pages 17,48-49, 160._)

SWALLOWTAIL BUTTERFLY JUST OUT OF ITS CHRYSALIS.
THE CHRYSALIS SKIN BELOW]

TIGER SWALLOWTAIL]

THE HAMMOCK CATERPILLAR ON ITS HAMMOCK]

The migrations thus considered are only exceptional occurrences. There is, however, a regularly recurring annual migration on the part of some butterflies which is also a phenomenon of extraordinary interest. The most notable example is that of the Monarch which apparently follows the birds southward every autumn and comes northward again in spring. There is much evidence to indicate that in some slight degree other butterflies have a similar habit, although the present observations are inadequate to determine to what extent this habit has become fixed in most of these species.

HIBERNATION OR WINTER LETHARGY

The ways in which butterflies spend the winter are always of peculiar interest to the naturalist. Here are creatures with four distinct stages of existence, each of which has the possibility of carrying the species through the season of cold. It is necessary to learn for each insect which stage has been chosen for the purpose, and if possible to find the reasons for the choice.

As a rule the related members of a group are likely to hibernate in a similar stage. Thus most of the Swallowtails pass the winter as chrysalids while practically all the Angle-wings pass the winter as adults. This rule, however, has many exceptions, for one will often find closely related species which differ in the stage of hibernation.

As one would expect, the conditions of hibernation vary greatly with the latitude. In the severe climate of the far north the conditions are likely to be more uniform than in the South where the milder climate permits greater variation to the insect. In some cases where a butterfly hibernates in only one stage in Canada it may pass the winter in two or more stages in Alabama or Florida.

In many other orders of insects the egg is a favorite stage for hibernation. Even in the closely related moths it is often chosen by many species, but comparatively few butterflies pass the winter in the egg stage. The little Bronze Copper may serve as one example of this limited group.

The conditions as to hibernation by the larvae of butterflies are very different from those of the egg. It has been estimated that probably half of all our species pass the winter in some stage of caterpillar growth. This varies all the way from the newly hatched caterpillar which hibernates without tasting food to the fully grown caterpillar which hibernates full fed and changes to a chrysalis in spring without eating anything at that time. A large proportion, however, feed both in fall and spring, going through the winter when approximately half grown.

The Graylings and the Fritillaries are typical examples of butterflies which hibernate as newly hatched larvae. The eggs are laid in autumn upon or near the food plants and the caterpillars gnaw their way out of the shells and seek seclusion at once, finding such shelter as they may in the materials on the soil surface. In spring they begin to feed as soon as the weather permits and complete their growth from then on.

The half-grown caterpillars may hibernate either as free creatures under boards, stones, or in the turfy grass, or they may be protected by special shelters which they have provided for themselves in their earlier life. In the case of the latter each may have a shelter of its own or there may be a common shelter for a colony of caterpillars. Among the examples of those hibernating in miscellaneous situations without special protection the caterpillars of the Tawny Emperor, the Gray Emperor, the Pearl Crescent, and some of the Graylings are examples. Among those which hibernate in individual shelters the Sovereigns, among which our common Viceroy is most familiar, are good examples. Among those which hibernate in a tent woven by the whole colony for the whole colony the Baltimore or Phaeton butterfly is perhaps the best example.

The caterpillars that hibernate when full grown may be grouped in a way somewhat similar to those which are half grown. Many species simply find such shelter as they may at or near the soil surface. The Clouded Sulphur is a good example of these. Others pass the winter in individual shelters made from a leaf or blade. Several of the larger Skippers are good examples of this condition. So far as I know none of our species pass the winter in colonial shelters when full grown.

It would be natural to suppose that the great majority of butterflies would be likely to hibernate in the chrysalis state. Here is a quiet stage in which the insect is unable to move about or to take any food, in which it seems entirely dormant and as a rule is fairly well hidden from the view of enemies. We find, however, that only a rather small proportion of our butterflies has chosen this stage for survival through the winter. The most conspicuous examples are the Swallowtails, nearly all of which hibernate in the chrysalis stage. Other examples are the various Whites, the Orange-tips, and isolated species like the Wanderer, and the Spring Azure and the American Copper. Practically all the butterflies that pass the winter as chrysalids have a silken loop running around near the middle of the body which helps to hold them securely through the long winter months. Apparently none of those chrysalids which hang straight downward are able to survive the winter.

An adult butterfly seems a fragile creature to endure the long cold months of arctic regions. Yet many of our most beautiful species habitually hibernate as adults, finding shelter in such situations as hollow trees, the crevices in rocks, the openings beneath loose bark or even the outer bark on the under side of a large branch. It is significant that most of the adult-wintering Angle-wings are northern rather than southern species, some of them being found in arctic regions practically around the world. One of the few southern forms that hibernates as an adult is the Goatweed Emperor.

These examples are all cases of true hibernation in a lethargic condition. There are certain butterflies, however, which pass the winter as adults that remain active during this period. Obviously this is impossible in latitudes where the winter is severe, and it involves migration to a warmer climate. The one notable illustration of this is the Monarch butterfly which apparently flies southward to the Gulf states at least and there remains until spring, when individuals come north again. The southward migration may be begun in Canada when the butterflies gather together in enormous flocks that remind one of the gathering of the clans with the migrating birds. This is one of the least understood of insect activities but it has been observed so often and over so long a period of years that there seems to be no questioning the general habit.

Like everything else in relation to living things there are numerous variations in the prevailing modes of hibernation. In the case of many species one can find combinations of two or more stages in which the winter is passed. Probably if we could observe with sufficient care we might be able to find somewhere examples of almost any conceivable double combination--as egg with larva or chrysalis or adult--the insect hibernating in two of these stages. Many examples are known in which both chrysalis and adult of the same species pass the winter and also of those in which young and well-grown larvae pass the winter. As one would expect, the conditions as to such combinations are likely to be more variable in southern than in northern regions.

Notwithstanding all the attention which has been paid to butterfly life-histories there is still some uncertainty in regard to the hibernation of many of our species. One of the most interesting series of observations which a young naturalist could undertake would be to learn positively how each species of butterfly in his locality passes the winter.

AESTIVATION OR SUMMER LETHARGY

In some species of butterflies there is a special adaptation to passing through the hottest part of the summer season in a state of lethargy which is suggestive of the torpor of the hibernating period. This phase of butterfly existence has not been extensively studied and there are indications that it exists more generally than has been commonly supposed. It has been noticed even in northern New England that some of the Angle-wings seek shelter and become lethargic during August. Apparently this is an adaptation to single broodedness, helping to carry the species through the year without the exhaustion incident to the continued activity of the butterfly.

In more southern regions, especially in the hot, dry climates where vegetation withers in midsummer, it is well known that some caterpillars become lethargic, remaining inactive until the fall rains start vegetation into growth. The Orange-sulphur butterfly is a good example of this.

This summer lethargy offers excellent opportunity for careful study. Any observer who finds a butterfly hidden away in summer under boards, the bark of a tree, or in a stone pile should look carefully to see what species it is and how the butterfly behaves. Such observations should be sent to the entomological journals in order that our knowledge of the subject may be increased.

FEIGNING DEATH

The fact has long been noticed that various butterflies have the habit at times of feigning death and dropping to the ground where they may lie motionless for a considerable period. This habit is most easily observed in some of the Angle-wings, especially those which hibernate as adults. Those species have the under surfaces of their wings colored in various bark-picturing patterns and apparently live through the winter to some extent, resting beneath the bark of large branches or upon the trunks of trees. Many of them also secrete themselves in hollow trees or beneath loose bark or in board piles or stone walls. It is probable, however, that during the long ages when these insects were adapting themselves to their life conditions, before man interfered with the natural order and furnished various more or less artificial places for hibernation, these butterflies rested more generally upon the under side of branches than they do now.

Even in warm weather when one of these butterflies is suddenly disturbed it is likely to fold its legs upon its body and drop to the ground, allowing itself to be handled without showing any signs of life. This habit is doubtless of value, especially during hibernation or possibly during the summer lethargy or aestivation, the latter a habit which may be more general among these butterflies than is now supposed. As the insect lies motionless upon the ground it is very likely to blend so thoroughly with its surroundings that it becomes concealed, and any bird which had startled it from the branch above would have difficulty in finding it.

Some very interesting observations have been made upon the death-feigning instincts of various other insects, especially the beetles. But no one so far as I know has yet made an extended study of the subject in relation to our American butterflies. It is an excellent field for investigation and offers unusual opportunities for photographic records. One of the pictures opposite page 32 shows a photograph which I took of a Mourning Cloak as it was thus playing 'possum. This species exhibits the instinct to a marked degree.

COLORATION

The caterpillars of butterflies and moths form a large part of the food of insect-eating birds. These caterpillars are especially adapted for such a purpose and in the economy of nature they play a very important part in keeping alive the feathered tribes. During the long ages through which both birds and insects have been developing side by side, there have been many remarkable inter-relations established which tend on the one hand to prevent the birds from exterminating the insects and on the other to prevent the insects from causing the birds to starve. The most important of these, so far as the caterpillars are concerned, are the various devices by which these insects protect themselves from attack, by hiding away where birds are not likely to find them, by clothing their bodies with spiny hairs, by other methods of rendering themselves distasteful, or by various phases of concealing coloration. On the whole, the examples of the latter are not so numerous or so easily found in the case of the larvae of butterflies as in those of moths.

Perhaps the basal principle of concealing coloration is the law of counter-shading, first partially announced by Prof. E. B. Poulton, and later much more elaborately worked out by Mr. Abbott H. Thayer, and discussed at length by Mr. Gerald H. Thayer in his remarkable volume, "Concealing Coloration in the Animal Kingdom." The law of counter-shading is tersely stated in these words: "Animals are painted by nature darkest on those parts which tend to be most lighted by the sky's light and _vice versa_." As this law works out on most animals that live on or near the ground, the upper part of the body exposed to the direct light from above is dark; and the under part, shut off from the upper light and receiving only the small reflection from below, is enough lighter to make the appearance of the creature in its natural environment of a uniform tone from back to breast.

Nearly all caterpillars illustrate this law of counter-shading. If they are in the habit of feeding or resting with their feet downward the back will be darker and the under side lighter, but if they are in the habit of feeding or resting in the opposite position these color tones will be reversed. One can find examples of such conditions almost any summer's day by a little searching of trees or shrubs.

This law of counter-shading, however, is really only the basis for the coloration of caterpillars or other animals. It tends, chiefly, to make the creature appear as a flat plane when seen from the side, and may be said in a way to prepare the canvas upon which Nature paints her more distinctive pictures. A great many examples of color markings that tend to conceal the caterpillar amid its natural surroundings may be found by any one who will study the subject and it offers one of the most interesting fields for investigation. The chapter on caterpillars in the above-mentioned book by Mr. Thayer should serve as a starting point for any one taking up the subject.

Butterflies differ from caterpillars and from most other animals in the fact that their coloring is chiefly shown upon the flat surfaces of the wings. Consequently, there is less opportunity for the various phases of counter-shading which is so commonly shown in the larger caterpillars. The bodies of nearly all butterflies do exhibit this phenomenon, but these bodies are relatively so small that counter-shading plays but a little part in the general display.

Upon the outstretched membranes of the butterflies' wings Nature through the long ages of development has painted a great variety of pictures. Those which tend to protect the insect by concealment amid its surroundings are most commonly spread on the under surface of the wings. Especially is this true in the case of those species which pass the winter as adults or which have the habit of resting upon the bark of trees, the sides of rocks, or the surface of the ground. We here find some of the most interesting examples of obliterative coloring that occur in nature. Some butterflies have taken on the look of tree bark, others the sombre appearance of weathered rocks, while still others are painted with the images of flowerets and their stems.

_Dazzling and Eclipsing Colors_

Many of the butterflies, especially the Angle-wings, which are marked on the under surface in various protective colors, are admirable examples of that phase of animal coloring which is spoken of as _dazzling coloration_. This is apparently one of the most important protective devices to be found in Nature and the validity of it is now generally conceded by naturalists. One phase of it, which may be called eclipsing coloration, seems to have been first definitely formulated by the late Lord Walsingham, a famous English entomologist who enunciated it in an address as president of the Entomological Society of London. The most significant paragraphs in that address were these:

"My attention was lately drawn to a passage in Herbert Spencer's 'Essay on the Morals of Trade.' He writes: 'As when tasting different foods or wines the palate is disabled by something strongly flavored from appreciating the more delicate flavor of another thing afterward taken, so with the other organs of sense a temporary disability follows an excessive stimulation. This holds not only with the eyes in judging of colors, but also with the fingers in judging of texture.'

"Here, I think, we have an explanation of the principle on which protection is undoubtedly afforded to certain insects by the possession of bright coloring on such parts of their wings or bodies as can be instantly covered and concealed at will. It is an undoubted fact, and one which must have been observed by nearly all collectors of insects abroad, and perhaps also in our own country, that it is more easy to follow with the eye the rapid movements of a more conspicuous insect soberly and uniformly colored than those of an insect capable of changing in an instant the appearance it presents. The eye, having once fixed itself upon an object of a certain form and color, conveys to the mind a corresponding impression, and, if that impression is suddenly found to be unreliable, the instruction which the mind conveys to the eye becomes also unreliable, and the rapidity with which the impression and consequent instruction can be changed cannot always compete successfully with the rapid transformation effected by the insect in its effort to escape."

Lord Walsingham then goes on to suggest that this intermittent play of bright colors probably has as confusing effect upon birds and other predaceous vertebrates as upon man; and that on this hypothesis such colors can be accounted for more satisfactorily than upon any other yet suggested. Since then the significance of this theory has been repeatedly pointed out by Professor Poulton, Mr. Abbott H. Thayer, and various other authorities upon animal coloring. The terms _dazzling_ and _eclipsing_ have been applied to the phenomenon.

Shortly after Lord Walsingham propounded this theory I called attention [A] to its fitness in explaining some of the most interesting color phases shown by American insects, notably the moths and locusts which have brilliantly colored under wings and protectively colored upper wings.

[A] _Popular Science Monthly_, 1898, "A Game of Hide and Seek."
Reprinted in the _Insect World_, 1899.

The animals of the north show numberless color phases of interest. One of the most curious of these is exhibited by several families of insects in which the outer wings are protectively colored in dull hues and the under wings brightly colored. For example, there are many species of moths belonging to the genus Catocala found throughout the United States. These are insects of good size, the larger ones measuring three inches in expanse of wings, and the majority of them being at least two thirds that size. Most of them live during the day on the bark of trees, with their front wings folded together over the back. The colors and markings of these wings, as well as of the rest of the exposed portions of the body, are such as to assimilate closely with the bark of the tree upon which the insect rests. In such a situation it requires a sharp eye to detect the presence of the moth, which, unless disturbed, flies only at night, remaining all day exposed to the attacks of many enemies. Probably the most important of these are the birds, especially species like the woodpeckers, which are constantly exploring all portions of the trunks of trees.

The chief beauty of these Catocalas, as they are seen spread out in the museum cabinet, lies in the fact that the hind wings, which, when the moth is at rest in life, are concealed by the front ones, are brightly colored in contrasting hues of black, red, and white in various brilliant combinations. These colors, in connection with the soft and blended tones of the front wings, make a very handsome insect.

It is easy to see that when one of these Underwing Moths is driven to flight by a woodpecker or other bark-searching bird it would show during its rapid, irregular flight the bright colors of the under wings which would be instantly hidden upon alighting and the very different coloring of the upper wings blending with the bark would be substituted. Consequently, the bird would be very likely to be baffled in its pursuit.

_Coloration of Locusts_

On the rocky hills and sandy plains of New England there are several species of grasshoppers or locusts that also illustrate these principles. If you walk along a strip of sandy land in summer, you start to flight certain locusts which soon alight, and when searched for will be found closely to assimilate in color the sand upon which they rest. On a neighboring granite-ribbed hill you will find few if any of this species of locust, but instead there occur two or three quite different species, which when at rest closely resemble the lichen-covered rocks. This resemblance is very striking, and is found in all stages of the insect's existence. If now you go to a lowland meadow, still another color phase will be found to prevail--the green grass is swarming with the so-called "long-horned" grasshoppers, which are green throughout with linear bodies, and long, slender legs and antennae.

Each of these three groups of insects is adapted to its particular habitat. All are constantly persecuted by birds, and have been so persecuted for unnumbered ages in the past. In every generation the individuals have varied, some toward closer resemblance to environment, others in an opposite direction. The more conspicuous insects have been constantly taken, and the least conspicuous as constantly left to reproduce. Were the three groups to change places to-day, the green grasshoppers from the meadows going to sandy surfaces, the sand-colored locusts going to rocky hills, and the "mossbacks" from the hills to the lowland meadows, each would become conspicuous, and the birds would have such a feast as is seldom spread before them.

The species living on sand and rocks are often "flushed" by birds. Those which flew but a few feet would be likely to be captured by the pursuing bird; those which flew farther would stand a better chance of escaping. Similarly, those which flew slowly and in a straight line would be more likely to be caught than those which flew rapidly and took a zigzag course. As a consequence of the selection thus brought about through the elimination of those which flew slowly along the straight and narrow way that led to death, you will find that most locusts living in exposed situations when startled fly some distance in a rapid, zigzag manner.

But still another element of safety has been introduced by some species of these locusts through the adoption of the color tactics of the Catocala moths. The under wings of the common Carolina locust--the species most abundant along the highway--are black, bordered with yellowish white. The base of the hind wings of a related species living on the Western plains is bluish, while in the large coral-winged locust of the Eastern states the hind wings are red, bordered with black. In nearly all of the species of these locusts frequenting open localities where they are liable to disturbance by birds or other animals, the hind wings exhibit contrasting colors in flight. Most of them also fly in a zigzag line, and alight in a most erratic manner. Many times I have had difficulty in determining the exact landfall of one of these peculiar creatures, and I believe Lord Walsingham's suggestion is well exemplified in them. (_See page 33._)

The most famous example of a combination of this dazzling coloring of the upper wing surface with a definite protective coloring of the under wing surface is the Kallima butterfly which is illustrated in almost every book dealing with animal coloration. The under wing surface bears a striking resemblance to a leaf and the hind wings project to form a tail which looks like the petiole of the leaf, and there is also a mark running across the wings which mimics the midrib. When the butterfly is flying the brilliant colors of the upper surface are visible, but when it alights these are instantly replaced by the sombre tone of the under surface, so that apparently the insect completely disappears and in its place there is only a leaf attached to a branch in a most natural position. In Dr. Longstaff's book there is an illustration of another tropical butterfly, _Eronia cleodora_, which resembles on its under surface a yellow disease-stricken leaf but on its upper surface gives a brilliant combination of black and white. This insect alights upon the leaves which it resembles and is a striking example of both dazzling and mimicking coloration.

Many of our own butterflies, notably the Angle-wings, are excellent examples of a similar combination. In flight they reveal conspicuous colors which are instantly hidden upon alighting and then one only sees the bark-like or dead leaf-like under surface as may be seen in the plate opposite pages 160-161. The iridescence upon the upper wing surface of many butterflies, whose under wing surface is colored in concealing tones, is doubtless also of great use to the insect in a similar way. There is a splendid opportunity here for some observer to study this phase of butterfly activity and to get photographs of the insects amid their natural surroundings.

In their book upon "Concealing Coloration" the Messrs. Thayer have called attention to many interesting phases of dazzling coloration. They show that bright marks like the eye-spots or ocelli, which form so prominent a feature on the wing surfaces of many butterflies, really helped to conceal the insect amid its natural surroundings, by drawing the eye away from the outlines of wings and body so that the latter tend to disappear. Their discussion of this subject opens up another vast field for outdoor observations of absorbing interest, in which there is great need for many active workers.

BUTTERFLY FEIGNING DEATH, HANGING TO BARK BY ONE FOOT]

BUTTERFLY IN HIBERNATING POSITION]

THE MONARCH BUTTERFLY: CHANGE FROM CATERPILLAR TO CHRYSALIS.
Photographs from life. (_See pages 8-10, 233_)]

THE MONARCH BUTTERFLY: THE CHANGE FROM CHRYSALIS TO ADULT.
Photographs from life (_See pages 10-13, 235_)]

THE CAROLINA LOCUST
Above, with wings expanded as in flight]

SELECTIVE COLOR SENSE

One who collects the Underwing moths soon discovers that the light colored species which resemble the bark of birch trees are likely to be found upon the trunks of those trees, and that the dark colored kinds which resemble the bark of maple trees are likely to be found upon the trunks of these. Obviously, were this not true the protective coloring would avail but little and it is evident that these moths are able to select a background which is of advantage in helping to conceal them.

There is much evidence to show that in a similar way the butterflies are able by means of a well-developed color sense to select the places where they alight. One of the most notable examples is that of a South American species, _Peridromia feronia_. This is a silvery gray butterfly which alights head downward upon the bark of certain palm trees that have silvery gray stems and remains there with its wings fully expanded so that it utilizes the background in much the same way that the Underwing moths do. "When disturbed they will return to the same tree again and again."

One who will observe the habits of our Angle-wings and other butterflies which have obliterative coloring of the under wing surface can easily learn that these insects select rather carefully the places where they alight. It will be found that as a rule each species utilizes a background that blends with its own coloring. It is probable that this habit is much more common in other groups of butterflies than has been realized. Much evidence of this sort has been collected regarding the butterflies of Europe and other countries, as well as near our own borders in America.

WARNING COLORATION AND MIMICRY

The colors of a great many animals, including a considerable percentage of American butterflies and their larvae, have been commonly explained by the theory of _warning colors_. According to this theory animals which were for any reason not edible by birds and mammals have developed various striking combinations of color such as black and yellow, red and black, or black and white, in order to advertise to their foes their inedible qualities. This theory has been very generally accepted by naturalists and will be found expounded at length in many books published during the last quarter century.

The whole subject of the validity of warning coloration has recently been brought up for reconsideration by the illuminating investigations of Mr. Abbott H. Thayer and discussed at length in the book upon "Concealing Coloration" already mentioned. In an appendix to this book dated 1908 Mr. Thayer states that he no longer holds the belief that "there must somewhere be warning colors." He has convincingly shown that a large proportion of the animals which were supposed to be examples of this theory are really illustrations of concealing coloration. But there yet remain various facts which have been conclusively proven that apparently require the theory of warning colors to explain them. Here is another field in which there is a real need for much careful investigation under conditions that are rigidly scientific.