This little thorn-like tree-hopper and all of its queer harlequin tribe are near relatives to the buzzing cicada, or harvest-fly, whose whizzing din in the dog-days has won it the popular misnomer of "locust."
To the average listener this insect is a mere "wandering voice and a mystery," and its singular form, wide prominent eyes, glassy wings, and double drums are always a surprise to the tyro who first identifies the grotesque as his well-known "locust." Its musical accomplishments during this brief period of its life are known to all, but few have cared to interest themselves in the early history of the singer, ere it perfected its musical resources "for the delight of man." But the naturalist, and especially the arboriculturist and fruit-grower, know to their cost of other tricks of the cicada, or rather of Mrs. Cicada, immortalized by Zenarchus the Rhodian as his "noiseless wife"—
"Happy the cicadas' lives, Since they all have noiseless wives."
I have alluded to the egg of the cicada "inserted in the bark of a twig." This act is accomplished by a knife-like ovipositor, which literally gouges a deep gash into the tender wood of various twigs, a number of the eggs being implanted in its depths, often causing the death of the branch. Shortly after hatching, the young cicadas leap for the ground, and burrowing beneath the surface, remain for a period varying from three to seventeen years, according to the species, to complete their transformations. Now the habits of my little tree-hopper are somewhat modelled after its big cousin. Knowing that the little insect was provided with a keen-edged ovipositor, and was in the habit of thrusting its tiny eggs beneath the bark, and realizing, too, that these strange tufts were of course in some way connected with the maternal instinct, I was led to investigate. Selecting a branch where the tufts and hoppers seemed most prolific, I brought my magnifying-glass to bear upon them at a respectful distance. Was ever actual thorn more motionless or non-committal than most of these?—their under surfaces hugging close against the bark, their telltale feet closely withdrawn, and all their pointed helmets inclined in the same parallel direction. One after another of the sly little family was examined without a revelation. Not until I had reached the upper limit of the group did I get any encouragement. Here I discovered one of the midgets in a new position, its pointed helmet inclined farther downward, and its other extremity correspondingly raised, so that I could see beneath its body. I now observed what at first appeared to be the hind leg of the farther side of the body protruding beneath, but in another moment noted my error, and saw that its sharp point had penetrated the bark, into which it soon sank quite deeply, and I realized that the ovipositor was now conducting its tiny eggs into the cambium layer of the bark. Without waiting for this particular individual to finish her labors, which might be extended for hours for aught I knew, I turned my glass upon its nearest neighbor, and a most accommodating specimen she proved, disclosing all the mysteries of the little froth house, its strange material, and unique method of construction. What I saw reminded me irresistibly of the technique of the cake-frosting art of the fancy baker, with its flowing tube of white condiment, and its following tracery of questionable design in high relief. This accommodating specimen had apparently just completed her egg-laying, or had perhaps just filled one nest; and while her attitude was precisely similar to that of her neighbor, I noticed a tiny ball of glistening froth at the tip of the ovipositor. This was attached to the bark by a touch, and from this starting-point the construction of the glistening house was continued, the apex of the ovipositor pouring out its endless puffy roll of aerated cement, which seemed to set as soon as laid.
And what a convenient implement this for a froth-house builder who is compelled to work behind her back—mortar-feeder, trowel, darby, compass, and level all in one! Beginning with the first touch of the cement, the flowing point describes a very small half-circle to the right, again meeting the bark. It is now carried inward and upward, describing a very close circle with scarcely any space intervening, a similar circle being repeated on the left side. A new tier is then begun in the same manner, only this time a little larger in the sweep, and leaving a perceptible opening at the right as the central wall is carried upward with slightly decreased material. Returning down the central wall again, the white coil is carried to the left along the bark, and up again on the other outer edge, until it once more meets its fellow at the ridge-pole, where the two coils appear to interlock as in a braid. And thus the little builder continues, enlarging the cavity with each circuit, until the full height is reached, and then decreasing proportionately until the glistening braided dome is tapered off again against the bark.
Now what is the object of this frothy pavilion? The life history of the insect, in contrast to that of the cicada, will perhaps throw a little light on that question. In the cicada, as I have shown, the eggs are inserted in the bark, but the young, hatching about six weeks later, immediately forsake the parent tree and enter the ground. But the young of our bittersweet membracis are not thus fickle, the entire life of the insect being spent on the plant. Moreover, its eggs are laid in late summer, and do not hatch until the following spring. What, then, is this canopy of the tree-hopper but the provision of a thoughtful mother, a pavilion about her offspring as a shelter through the winter storms? In early July the tiny hoppers emerge from their egg-cases, and presumably creep out from their luminous domicile, and later on in the season these broods of varying numbers and all sizes are to be seen among the young stems of the plant, their beaks inserted, their pointed heads invariably in the same direction—towards the top of the branch. Even though in flight one of the midgets is seen to alight in violence to the rule, he instantly recognizes his mistake, and quickly glides round to the orthodox position.
This curious insect is chiefly confined to the bittersweet, though he is occasionally found in the company of a much bigger cousin of his on the branches of the locust, where these same telltale corrugated frothy pavilions are often seen to clothe the young twigs in their white tufts, the similar product of the larger species, which thus also presumably spends its entire life upon the locust-tree.
THE WELCOMES OF THE FLOWERS
It is now some thirty years since the scientific world was startled by the publication of that wonderful volume, "The Fertilization of Orchids," by Charles Darwin; for though slightly anticipated by his previous work, "Origin of Species," this volume was the first important presentation of the theory of cross-fertilization in the vegetable kingdom, and is the one that is primarily associated with the subject in the popular mind. The interpretation and elucidation of the mysteries which had so long lain hidden within those strange flowers, whose eccentric forms had always excited the curiosity and awe alike of the botanical fraternity and the casual observer, came almost like a divine revelation to every thoughtful reader of his remarkable pages. Blossoms heretofore considered as mere caprices and grotesques were now shown to be eloquent of deep divine intention, their curious shapes a demonstrated expression of welcome and hospitality to certain insect counterparts upon whom their very perpetuation depended.
Thus primarily identified with the orchid, it was perhaps natural and excusable that popular prejudice should have associated the subject of cross-fertilization with the orchid alone; for it is even to-day apparently a surprise to the average mind that almost any casual wild flower will reveal a floral mechanism often quite as astonishing as those of the orchids described in Darwin's volume. Let us glance, for instance, at the row of stamens below (Fig. 1), selected at random from different flowers, with one exception wild flowers. Almost everybody knows that the function of the stamen is the secretion of pollen. This function, however, has really no reference whatever to the external form of the stamen. Why, then, this remarkable divergence? Here is an anther with its two cells connected lengthwise, and opening at the sides, perhaps balanced at the centre upon the top of its stalk or filament, or laterally attached and continuous with it; here is another opening by pores at the tip, and armed with two or four long horns; here is one with a feathery tail. In another the twin cells are globular and closely associated, while in its neighbor they are widely divergent. Another is club-shaped, and opens on either side by one or more upraised lids; and here is an example with its two very unequal cells separated by a long curved arm or connective, which is hinged at the tip of its filament; and the procession might be continued across two pages with equal variation.
As far back as botanical history avails us these forms have been the same, each true to its particular species of flower, each with an underlying purpose which has a distinct and often simple reference to its form; and yet, incredible as it now seems to us, the botanist of the past has been content with the simple technical description of the feature, without the slightest conception of its meaning, dismissing it, perhaps, with passing comment upon its "eccentricity" or "curious shape." Indeed, prior to Darwin's time it might be said that the flower was as a voice in the wilderness. In 1735, it is true, faint premonitions of its present message began to be heard through their first though faltering interpreter, Christian Conrad Sprengel, a German botanist and school-master, who upon one occasion, while looking into the chalice of the wild geranium, received an inspiration which led him to consecrate his life thence-forth to the solution of the floral hieroglyphics. Sprengel, it may be said, was the first to exalt the flower from the mere status of a botanical specimen.
This philosophic observer was far in advance of his age, and to his long and arduous researches—a basis built upon successively by Andrew Knight, Koehlreuter, Herbert, Darwin, Lubbock, Mueller, and others—we owe our present divination of the flowers.
In order to fully appreciate this present contrast, it is well to briefly trace the progress, step by step, from the consideration of the mere anatomical and physiological specimen of the earlier botanists to the conscious blossom of to-day, with its embodied hopes, aspirations, and welcome companionships.
Most of my readers are familiar with the general construction of a flower, but in order to insure such comprehension it is well, perhaps, to freshen our memory by reference to the accompanying diagram (Fig. 2) of an abstract flower, the various parts being indexed.
The calyx usually encloses the bud, and may be tubular, or composed of separate leaves or sepals, as in a rose. The corolla, or colored portion, may consist of several petals, as in the rose, or of a single one, as in the morning-glory. At the centre is the pistil, one or more, which forms the ultimate fruit. The pistil is divided into three parts, ovary, style, and stigma. Surrounding the pistil are the stamens, few or many, the anther at the extremity containing the powdery pollen.
Although these physiological features have been familiar to observers for thousands of years, the several functions involved were scarcely dreamed of until within a comparatively recent period.
In the writings of ancient Greeks and Romans we find suggestive references to sexes in flowers, but it was not until the close of the seventeenth century that the existence of sex was generally recognized.
In 1682 Nehemias Grew announced to the scientific world that it was necessary for the pollen of a flower to reach the stigma or summit of the pistil in order to insure the fruit. I have indicated his claim pictorially at A (Fig. 3), in the series of historical progression. So radical was this "theory" considered that it precipitated a lively discussion among the wiseheads, which was prolonged for fifty years, and only finally settled by Linnaeus, who reaffirmed the facts declared by Grew, and verified them by such absolute proof that no further doubts could be entertained. The inference of these early authorities regarding this process of pollination is perfectly clear from their statements. The stamens in most flowers were seen to surround the pistil, "and of course the presumption was that they naturally shed the pollen upon the stigma," as illustrated at B in my series. The construction of most flowers certainly seemed designed to fulfil this end. But there were other considerations which had been ignored, and the existence of color, fragrance, honey, and insect association still continued to challenge the wisdom of the more philosophic seekers. How remarkable were some of those early speculations in regard to "honey," or, more properly, nectar! Patrick Blair, for instance, claimed that "honey absorbed the pollen," and thus fertilized the ovary. Pontidera thought that its office was to keep the ovary in a moist condition. Another botanist argued that it was "useless material thrown off in process of growth." Krunitz noted that "bee-visited meadows were most healthy," and his inference was that "honey was injurious to the flowers, and that bees were useful in carrying it off"! The great Linnaeus confessed himself puzzled as to its function.
For a period of fifty years the progress of interpretation was completely arrested. The flowers remained without a champion until 1787, when Sprengel began his investigations, based upon the unsolved mysteries of color and markings of petals, fragrance, nectar, and visiting insects. The prevalent idea of the insect being a mere idle accessory to the flower found no favor with him. He chose to believe that some deep plan must lie beneath this universal association. At the inception of this conviction he chanced to observe in the flower of the wild geranium (G. sylvaticum) a fact which only an inspired vision could have detected—that the minute hairs at the base of the petal, while disclosing the nectar to insects, completely protected it from rain. Investigation showed the same conditions in many other flowers, and the inference he drew was further strengthened by the remarkable discovery of his "honey-guides" in a long list of blossoms, by which the various decorations of spots, rings, and converging veins upon the petals indicated the location of the nectar.
His labors were now concentrated on the work of interpretation, until at length his researches, covering a period of two or three years, were given to the world. In a volume bearing the following victorious title, "The Secrets of Nature in Forms and Fertilization of Flowers Discovered," he presented a vast chronicle of astonishing facts. The previous discoveries of Grew and Linnaeus were right so far as they went—viz., "the pollen must reach the stigma"—but those learned authorities had missed the true secret of the process. In proof of which Sprengel showed that in a great many flowers, as I have shown at C (Fig. 3), this deposit of pollen is naturally impossible, owing to the relative position of the floral parts, and that the pollen could not reach the stigma except by artificial aid. He then announced his startling theory:
1. "Flowers are fertilized by insects."
2. Insects in approaching the nectar brush the pollen from the anthers with various hairy parts of their bodies, and in their motions convey it to the stigma.
But Sprengel's seeming victory was doomed to be turned to defeat. The true "secret" was yet unrevealed in his pages. He had given a poser to Linnaeus (C), yet his own work abounded with similar strange inconsistencies, which, while being scarcely admitted by himself, or ingeniously explained, were nevertheless fatal to the full recognition of his wonderful researches. For seventy years his book lay almost unnoticed.
"Let us not underrate the value of a fact; it will one day flower in a truth." The defects in Sprengel's work were, after all, not actual defects. The error lay simply in his interpretation of his carefully noted facts. As Hermann Mueller has said, "Sprengel's investigations afford an example of how even work that is rich in acute observation and happy interpretation may remain inoperative if the idea at its foundation is defective." What, then, was the flaw in Sprengel's work? Simply that he had seen but half the "secret" which he claimed to have "discovered." Starting to prove that insects fertilize the flowers, his carefully observed facts only served to demonstrate in many cases the reverse—that insects could not fertilize flowers in the manner he had declared. He was met at every hand, for instance, by floral problems such as are shown at E and F, where the pollen and the stigma in the same flower matured at different periods; and even though he recognized and admitted that the pollen must in many cases be transferred from one flower to another, he failed to divine that such was actually the common vital plan involved. It may readily be imagined that his great work precipitated an intense and prolonged controversy, and incited emulous investigation by the botanists of his time. Though a few of the more advanced of his followers, among them Andrew Knight (1799), Koehlreuter (1811), Herbert (1837), Gaertner (1844), clearly recognized the principle and foreshadowed the later theory of cross-fertilization, it was not until the inspired insight of Darwin, as voiced in his "Origin of Species," contemplated these strange facts and inconsistencies of Sprengel that their full significance and actual value were discovered and demonstrated, and his remarkable book, forgotten for seventy years, at last appreciated for its true worth. Alas for the irony of fate! Under Darwin's interpretation the very "defects" which had rendered Sprengel's work a failure now became the absolute witness of a deeper truth which Sprengel had failed to discern. One more short step and he had reached the goal. But this last step was reserved for the later seer. He took the fatal double problem of Sprengel—as shown at E and F, to express the consummation pictorially—and by the simple drawing of a line, as it were, as indicated between G and H, instantly reconciled all the previous perplexities and inconsistencies, thus demonstrating the fundamental plan involved in floral construction to be not merely "insect fertilization," the fatal postulate assumed by Sprengel, but cross-fertilization—a fact which, singularly enough, the latter's own pages proved without his suspicion.
Thus we see the four successive steps in progressive knowledge, from Grew in 1682, Linnaeus, 1735, Sprengel, 1787, to Darwin, 1857-1858, and realize with astonishment that it has taken over one hundred and seventy-five years for humanity to learn this apparently simple lesson, which for untold centuries has been noised abroad on the murmuring wings of every bee in the meadow, and demonstrated in almost every flower.
This infinite field now open before him, Darwin began his investigations, and the whole world knows his triumphs. He has been followed by a host of disciples, to whom his books have come as an inspiration and ennobling impulse. Hildebrand, Delpino, Axell, Lubbock, and, latest and perhaps most conspicuous, Hermann Mueller, to whom the American reader is especially referred. "The Fertilization of Flowers," by this most scholarly and indefatigable chronicler, presents the most complete compendium and bibliography of the literature on the subject that have yet appeared. Even to the unscientific reader it will prove full of revelations of this awe-inspiring interassociation and interdependence of the flower and the insect.
Many years ago the grangers of Australia determined to introduce our red clover into that country, the plant not being native there. They imported American seed, and sowed it, with the result of a crop luxuriant in foliage and bloom, but not a seed for future sowing! Why? Because the American bumblebee had not been consulted in the transaction. The clover and the bee are inseparable counterparts, and the plant refuses to become reconciled to the separation. Upon the introduction and naturalization of the American bumblebee, however, the transported clover became reconciled to its new habitat, and now flourishes in fruition as well as bloom.
Botany and entomology must henceforth go hand-in-hand. The flower must be considered as an embodied welcome to an insect affinity, and all sorts of courtesies prevail among them in the reception of their invited guests. The banquet awaits, but various singular ceremonies are enjoined between the cup and the lip, the stamens doing the hospitalities in time-honored forms of etiquette. Flora exacts no arbitrary customs. Each flower is a law unto itself. And how expressive, novel, and eccentric are these social customs! The garden salvia, for instance, slaps the burly bumblebee upon the back and marks him for her own as he is ushered in to the feast. The mountain-laurel welcomes the twilight moth with an impulsive multiple embrace. The desmodium and genesta celebrate their hospitality with a joke, as it were, letting their threshold fall beneath the feet of the caller, and startling him with an explosion and a cloud of yellow powder, suggesting the day pyrotechnics of the Chinese. The prickly-pear cactus encloses its buzzing visitor in a golden bower, from which he must emerge at the roof as dusty as a miller. The barberry, in similar vein, lays mischievous hold of the tongue of its sipping bee, and I fancy, in his early acquaintance, before he has learned its ways, gives him more of a welcome than he had bargained for. The evening primrose, with outstretched filaments, hangs a golden necklace about the welcome murmuring noctuid, while the various orchids excel in the ingenuity of their salutations. Here is one which presents a pair of tiny clubs to the sphinx-moth at its threshold, gluing them to its bulging eyes. Another attaches similar tokens to the tongues of butterflies, while the cypripedium speeds its parting guest with a sticking-plaster smeared all over its back. And so we might continue almost indefinitely. From the stand-point of frivolous human etiquette we smile, perhaps, at customs apparently so whimsical and unusual, forgetting that such a smile may partake somewhat of irreverence. For what are they all but the divinely imposed conditions of interassociation? say, rather, interdependence, between the flower and the insect, which is its ordained companion, its faithful messenger, often its sole sponsor—the meadows murmuring with an intricate and eloquent system of intercommunings beside which the most inextricable tangle of metropolitan electrical currents is not a circumstance. What a storied fabric were this murmurous tangle woven day by day, could each one of these insect messengers, like the spider, leave its visible trail behind it!
As a rule, these blossom ceremonies are of the briefest description. Occasionally, however, as in the cypripedium and in certain of the arums, or "jack-in-the-pulpit," and aristolochias, the welcome becomes somewhat aggressive, the guest being forcibly detained awhile after tea, or, as in the case of our milkweed, occasionally entrapped for life.
From this companionable point of view let us now look again at the strange curved stamen of the sage. Why this peculiar formation of the long curved arm pivoted on its stalk? Considered in the abstract, it can have no possible meaning; but taken in association with the insect to which it is shaped, how perfect is its adaptation, how instantly intelligible it becomes! Every one is familiar with the sage of the country garden, its lavender flowers arranged in whorls in a long cluster at the tips of the stems. One of these flowers, a young one from the top of the cluster, is shown at A (Fig. 4), in section, the long thread-like pistil starting from the ovary, and curving upward beneath the arch of the flower, with its forked stigma barely protruding (B). There are two of the queer stamens, one on each side of the opening of the blossom, and situated as shown, their anthers concealed in the hood above, and only their lower extremity appears below, the minute growth near it being one of the rudiments of two former stamens which have become aborted. If we take a flower from the lower portion of the cluster (D), we find that the thread-like pistil has been elongated nearly a third of an inch, its forked stigma now hanging directly at the threshold of the flower. The object of this will be clearly demonstrated if we closely observe this bee upon the blossoms. He has now reached the top of the cluster among the younger blossoms. He creeps up the outstretched platform of the flower, and has barely thrust his head within its tube when down comes the pair of clappers on his back (C). Presently he backs out, bearing a generous dab of yellow pollen, which is further increased from each subsequent flower. He has now finished this cluster, and flies to the next, alighting as usual on the lowermost tier of bloom. In them the elongated stigma now hangs directly in his path, and comes in contact with the pollen on his back as the insect sips the nectar. Cross-fertilization is thus insured; and, moreover, cross-fertilization not only from a distinct flower, but from a separate cluster, or even a separate plant. For in these older stigmatic flowers the anther as it comes down upon his back is seen to be withered, having shed its pollen several days since, the supply of pollen on the bee's body being sufficient to fertilize all the stigmas in the cluster, until a new supply is obtained from the pollen-bearing blossoms above. And thus he continues his rounds.
The sage is a representative of the large botanical order known as the Mint family, the labiates, or gaping two-lipped flowers, the arched hood here answering to the upper lip, the spreading base forming the lower lip, which is usually designed as a convenient threshold for the insects while sipping the nectar deep within the tube. This mechanism of the sage is but one of many curious and various contrivances in the Mint family, all designed for the same end, the intercrossing of the flowers.
While each family of plants is apt to favor some particular general plan, the modifications in the various species seem almost without limit.
Let us now look at the Heath family. The family of the heath, cranberry, pyrola, Andromeda, and mountain-laurel—how do these blossoms welcome their insect friends? This group is particularly distinguished by the unusual exception in the form of its anthers, which open by pores at their tips, instead of the ordinary side fissures. Two or three forms of these anthers are shown in my row of stamens (Fig. 1).
Seen thus in their detached condition, how incomprehensible and grotesque do they appear! And yet, when viewed at home, in their bell-shaped corollas, their hospitable expression and greeting are seen to be quite as expressive and rational as those of the sage. Take the mountain-laurel, for instance; what a singular exhibition is this which we may observe on any twilight evening in the laurel copse, the dense clusters of pink-white bloom waited upon by soft-winged fluttering moths, and ever and anon celebrating its cordial spirit by a mimic display of pyrotechnics as the anthers hurl aloft their tiny showers of pollen!
Every one is familiar with the curious construction of this flower, with its ten radiating stamens, each with its anther snugly tucked away in a pouch at the rim of its saucer-shaped corolla. Thus they appear in the freshly opened flower, and thus will they remain and wither if the flower is brought indoors and placed in a vase upon our mantel. Why? Because the hope of the blossom's life is not fulfilled in these artificial conditions; its natural counterpart, the insect, has failed to respond to its summons.
But the twilight cluster in the woods may tell us a pretty story.
Here a tiny moth hovers above the tempting chalice, and now settles upon it with eager tongue extended for the nectar at its centre. What an immediate and expressive welcome! No sooner has this little feathery body touched the filaments than the eager anthers are released from their pockets, and, springing inwards, clasp their little visitor, at the same time decorating him with their compliments of webby pollen (A, Fig. 5).
The nectary now drained of its sweets, the moth creeps or flutters to a second blossom, and its pollen-dusted body thus coming in contact with its stigma, cross-fertilization is accomplished. The pollen of the laurel differs from that of most of the Heath blooms, its grains being more or less adherent by a cobwebby connective which permeates the mass as indicated in my magnified representation (B, Fig. 5).
It is probable that an accessory cross-fertilization frequently results from a mass of the pollen falling directly upon the stigma of a neighboring blossom, or even upon its own stigma, but even in the latter case, as has been absolutely demonstrated as a general law by the experiments of Darwin, the pollen from a separate flower is almost invariably prepotent, and leads to the most perfect fruition, and thus to the survival of the fittest—the cross-fertilized. And, in any event, the insect is to be credited for the release of the tiny catapults by which the pollen is discharged. But the laurel may be considered as an exceptional example of the Heath family. Let us look at a more perfect type of the order to which it belongs, the globular blossom of the Andromeda (A. ligustrina).
Only a short walk from my studio door in the country I recently observed its singular reception to the tiny black-and-white banded bee, which seems to be its especial companion, none the less constant and forgiving in spite of a hospitality which, from the human stand-point, would certainly seem rather discouraging. Fancy a morning call upon your particular friend. You knock at the door, and are immediately greeted at the threshold with a quart of sulphur thrown into your face. Yet this is precisely the experience of this patient little insect, which manifests no disposition to retaliate with the concealed weapon which on much less provocation he is quick to employ. Here he comes, eager for the fray. He alights upon one of the tiny bells scarce half the size of his body. Creeping down beneath it, he inserts his tongue into the narrowed opening. Instantly a copious shower of dust is poured down upon his face and body. But he has been used to it all his life, and by heredity he knows that this is Andromeda's peculiar whim, and is content to humor it for the sweet recompense which she bestows. The nectar drained, the insect, as dusty as a miller, visits another flower, but before he enters must of necessity first pay his toll of pollen to the drooping stigma which barely protrudes beneath the blossom's throat, and the expectant seed-pod above welcomes the good tidings with visions of fruition.
And how beautiful is the minute mechanical adaptation by which this end is accomplished! This species of Andromeda is a shrub of about four feet in height, its blossoms being borne in close panicled clusters at the summit of the branches. The individual flower is hardly more than an eighth of an inch in diameter. From one of three blossoms I made the accompanying series of three sectional drawings (Fig. 6). The first shows the remarkable interior arrangement of the ten stamens surrounding the pistil. The second presents a sectional view of these stamens, showing their peculiar S-shaped filaments and ring of anthers—one of the latter being shown separate at the right, with its two pores and exposed pollen. The freshly opened blossom discloses the entire ring of anthers in perfect equilibrium, each with its two orifices closed by close contact with the style, thus retaining the pollen. It will readily be seen that an insect's tongue, as indicated by the needle, in probing between them in search for nectar, must needs dislocate one or more of the anthers, and thus release their dusty contents, while the position of the stigma below is such as to escape all contact.
In most flowers, with the exception of the orchids, the stamens and pollen are plainly visible; but who ever sees the anthers of the blue-flag? Surely none but the analytical botanist and the companion insect to whom it is so artfully adjusted and so demonstrative. This insect is likely to be either a bumblebee or a species of large fly. In apt illustration of Sprengel's theory of the "path-finder" or honey-guide, the insect does not alight at the centre of the flower, but upon one of the three large drooping sepals, whose veins, converging to the narrow trough above, indicate the path to the nectar. Closely overarching this portion is a long and narrow curved roof—one of three divisions to the style, each surmounting its veined sepals. Beneath this our visiting bee disappears, and a glance at my sectional drawing shows what happens. Concealed within, against the ridge-pole, as it were, the anther awaits his coming, and in his passage to and from the nectar below spreads its pollen over his head and back. Having backed out of this segment of the blossom (A, Fig. 7), he proceeds to the next; but the shelf-like stigma awaits him at the door, and scrapes off or rubs off a few grains of the pollen from his back (B). Thus he continues until the third segment is reached, from which he carries away a fresh load of pollen to another flower. It will be seen that only the outer side of this appendage is stigmatic, and that it is thus naturally impossible for the blue-flag to self-fertilize—only one instance of thousands in which the anther and stigma, though placed in the closest proximity, and apparently even in contact—seemingly with the design of self-fertilization—are actually more perfectly separated functionally than if in separate flowers, the insect alone consummating their affinity.
In some flowers this separation is effected, as I have shown, by their maturing at different periods; in others, as in the iris, by mere mechanical means; while in a long list of plants, as in the willow, poplar, hemp, oak, and nettle, the cross-fertilization is absolutely necessitated by the fact of the staminate and stigmatic flowers being either separated on the same stalk or on different plants, the pollen being carried by insects or the wind. We may see a pretty illustration of this in the little wild flower known as the devil's-bit (Chamaelirium luteum,), whose long, white, tapering spire of feathery bloom may often be seen rising above the sedges in the swamp. Two years ago I chanced upon a little colony of four or five plants at the edge of a bog. The flowers, all of them, were mere petals and stamens (B, Fig. 8). I looked in vain for a single stigmatic plant or flower; but far across the swamp, a thousand feet distant, I at length discovered a single spire, composed entirely of pistillate flowers, as shown in A (Fig. 8), and my magnifying-glass clearly revealed the pollen upon their stigmas—doubtless a welcome message brought from the isolated affinity afar by some winged sponsor, to whom the peculiar fragrance of the flower offers a special attraction, and thus to whom the fortunes of the devil's-bit have been committed.
The presence of fragrance and honey in a dioecious flower may be accepted in the abstract as almost conclusive of an insect affinity, as in most flowers of this class, notably the beech, pine, dock, grasses, etc., the wind is the fertilizing agent, and there is absence alike of conspicuous color, fragrance, and nectar—attributes which refer alone to insects, or possibly humming-birds in certain species.
Look where we will among the blossoms, we find the same beautiful plan of intercommunion and reciprocity everywhere demonstrated. The means appear without limit in their evolved—rather, I should say, involved—ingenuity. Pluck the first flower that you meet in your stroll to-morrow, and it will tell you a new story.
Only a few days since, while out on a drive, I passed a luxuriant clump of the plant known as "horse-balm." I had known it all my life, and twenty years previously had made a careful analytical drawing of the mere botanical specimen. What could it say to me now in my more questioning mood? Its queer little yellow-fringed flowers hung in profusion from their spreading terminal racemes. I recalled their singular shape, and the two outstretched stamens protruding from their gaping corolla, and could distinctly see them as I sat in the carriage. I had never chanced to read of this flower in the literature of cross-fertilization, and murmuring, half aloud, "What pretty mystery is yours, my Collinsonia?" prepared to investigate.
What I observed is pictured severally at Fig. 9, the flowers being shown from above, showing the two spreading stamens and the decidedly exceptional unsymmetrical position of the long style extending to the side. A small nectar-seeking bumblebee had approached, and in alighting upon the fringed platform grasped the filaments for support, and thus clapped the pollen against his sides. Reasoning from analogy, it would of course be absolutely clear that this pollen has thus been deposited where it will come in contact with the stigma of another flower. So, of course, it proved. In the bee's continual visits to the several flowers he came at length to the younger blooms, where the forked stigmas were turned directly to the front, while the immature stamens were still curled up in the flower tubes. Even the unopened buds showed a number of species where the early matured stigma actually protruded through a tiny orifice in precisely the right position to strike the pollen-dusted body of the bee, as he forced his tongue through the tiny aperture.[A]
[Footnote A: In numerous instances observed since the above was written I have noted the larger bumblebees upon the blossom. These insects have a different method of approach, hanging beneath the flower, the anthers being clapped against their thorax at the juncture of the wings, instead of the abdomen, as in the smaller bee.]
If their dainty mechanism excite our wonder, what shall be said of the revelations in the great order of the Compositae, where each so-called flower, as in the dandelion, daisy, cone-flower, marigold, is really a dense cluster of minute flowers, each as perfect in its construction as in the examples already mentioned, each with its own peculiar plan designed to insure the transfer of its own pollen to the stigma of its neighbor, while excluding it from its own?
All summer long the cone-flower, Fig. 10 (Rudbeckia hirta), blooms in our fields, but how few of us imagine the strange processes which are being enacted in that purple cone! Let us examine it closely. If we pluck one of the blossom's heads and keep it in a vase over-night, we shall probably see on the following morning a tiny yellow ring of pollen encircling the outer edge of the cone. In this way only are we likely to see the ring in its perfection, as in a state of nature the wind and insects rarely permit it to remain.
If we now with a sharp knife make a vertical section, as shown at A (Fig. 3), we may observe the conical receptacle studded with its embryo seeds, each bearing a tiny tubular blossom. Three distinct forms of these flowers are to be seen. The lower and older ones are conspicuous by their double feathery tails, the next by their extended anthers bearing the pollen at their extremity, and above these again the buds in all stages of growth. These various states are indicated in Fig. 11.
As in all the Compositae, the anthers are here united in a tube, the pollen being discharged within. At the base of this anther-tube rises the pistil, which gradually elongates, and like a piston forces out the pollen at the top. Small insects in creeping over the cone quickly dislodge it. In the next stage the anthers have withered, the flower-tube elongated, and the top of the two-parted pistil begins to protrude, and at length expands its tips, disclosing at the centre the stigmatic surface, which has until now been protected by close contact. (See section.)
A glance at Fig. 11 will reveal the plan involved. The ring of pollen is inevitably scattered to the stigmas of the neighboring flowers, and cross-fertilization continually insured. Similar contrivances are to be found in most of the Compositae, through the same method being variously applied.
Perhaps even more remarkable than any of the foregoing, which are more or less automatic in their movements, is the truly astonishing and seemingly conscious mechanism displayed in the wild arum of Great Britain—the "lords and ladies" of the village lanes, the foreign counterpart of our well-known jack-in-the-pulpit, or Indian-turnip, with its purple-streaked canopy, and sleek "preacher" standing erect beneath it. A representation of this arum is shown in Fig. 12, and a cross section at A, properly indexed.
How confidently would the superficial—nay, even careful—examination of one of the old-time botanists have interpreted its structure: "How simple and perfect the structure! Observe how the anthers are placed so that pollen shall naturally fall directly on the stigmas and fertilize them!" Such would indeed appear to be intended, until it is actually discovered that the stigmas have withered when the pollen is shed—a device which, acting in association with the little ring of hairs, tells a strange story. It is not my fortune to have seen one of these singular blossoms, but from the description of the process of fertilization given in Hermann Mueller's wonderful work, aided by a botanical illustration of the structure of the flower, I am readily enabled to picture the progressive stages of the mechanism.
In the first stage (B, Fig. 13) small flies with bodies dusted with pollen from a previous arum blossom (for insects, as a rule, remain faithful or partial to one species of flowers while it is in bloom) are entering the narrowed tube, easily passing through the drooping fringe of hairs. Nectar is secreted by the stigmas, and here the flies assemble, thus dusting them with pollen. Their appetite temporarily satisfied, the insects seek escape, but find their exit effectually barred by the intruding fringe of hairs (C). In this second stage the stigmas, having now been fertilized, have withered, at the same time exuding a fresh supply of nectar, which again attracts the flies, whereupon, as shown at D, the anthers open and discharge their pollen upon the insects. In the fourth stage (E), all the functions of the flower having now been fulfilled, the fringe of hairs withers, and the imprisoned pollen-laden flies are permitted to escape to another flower, where the beautiful scheme is again enacted.
In a paper of this kind it is of course possible only to hint at a few representative examples of floral mechanisms, but these would be indeed incomplete without a closing reference to that wonderful tribe of flowers with which the theory of cross-fertilization will ever be memorably associated. I have previously alluded to the absolute dependence of the red clover upon the bumblebee. This instance may be considered somewhat exceptional, though numerous parallel cases are known. Among ordinary flowers this intervention of the insect is largely a preferable intention, and though almost invariably fulfilled, a large proportion of flowers still retain, as a dernier ressort, the power of at least partial self-fertilization and perpetuity in the absence or neglect of their insect counterpart.
The numerous and conclusive demonstrations of Darwin, however, have proved that in the competition for existence such self-fertilized offspring quickly yield before the progeny of cross-fertilization.
But the distinctive feature of the orchids lies in the fact that this dependence on the insect is wellnigh universally absolute. Here are a great host of plants which are doomed to extinction if for any reason their insect sponsors should permanently neglect them. The principal botanical feature which differentiates the orchid from other plants lies in the construction of the floral organs, the pistil, stigma, and anthers here being united into a distinct part known as the column. The pollen is, moreover, peculiar, being collected into more or less compact masses, and variously concealed in the flower. Some of these are club-shaped, with a viscid extremity, others of the consistency of a sticking-plaster, and all are hidden from external view in pouches and pockets, from which they never emerge unless withdrawn on the body of an insect. The various devices by which this removal is insured are most astonishing and awe-inspiring. Nor is it necessary to go to the conservatory for a tropical specimen, as is commonly supposed. An orchid is an orchid wherever it grows, and our native list of some fifty species will afford examples of as strange mechanical adaptations as are to be found among Darwin's pages. Indeed, a few of our American species are there described. One example will suffice for present illustration—the sweet-pogonia or grass-pink of our sedgy swamps (Pogonia ophioglossoides). Its solitary rosy blossom, nodding on its slender stem above the sedges, is always a welcome episode to the sauntering botanist, and its perfume, suggesting ripe red raspberries, is unique in the wild bouquet. One of these flowers is shown in profile at Fig. 14, its various parts indexed. Concealed behind the petals is the column, elsewhere indicated from various points of view. Attracted by its color and fragrance, the insect seeks the flower; its outstretched fringy lip offers a cordial invitation at its threshold, and conducts its visitor directly to the sweets above. In his entrance, as seen at D (Fig. 15), the narrowed passage compresses his back against the underside of the column, forcing his head and back against the stigma. The effect of this inward pressure, as will be seen, only serves to force the anther more firmly within its pocket; but as the insect, having drained the nectar, now backs out, note the result. The lip of the anther catches upon the back, swings outward on its hinge, and deposits its sticky pollen all over the insect's back, returning to its original position after his departure. In another moment he is seen upon another blossom, as at D again, his pollen-laden back now coming in contact with the stigma, and the intention of the blossom is accomplished; for without this assistance from the insect the little lid remains close within its pocket, and the pollen is thus retained.
What startling disclosures are revealed to the inward eye within the hearts of all these strange orchidaceous flowers! Blossoms whose functions, through long eras of adaptation, have gradually shaped themselves to the forms of certain chosen insect sponsors; blossoms whose chalices are literally fashioned to bees or butterflies; blossoms whose slender, prolonged nectaries invite and reward the murmuring sphinx-moth alone, the floral throat closely embracing his head while it attaches its pollen masses to the bulging eyes, or perchance to the capillary tongue! And thus in endless modifications, evidences all of the same deep vital purpose.
Let us then content ourselves no longer with being mere "botanists"—historians of structural facts. The flowers are not mere comely or curious vegetable creations, with colors, odors, petals, stamens, and innumerable technical attributes. The wonted insight alike of scientist, philosopher, theologian, and dreamer is now repudiated in the new revelation. Beauty is not "its own excuse for being," nor was fragrance ever "wasted on the desert air." The seer has at last heard and interpreted the voice in the wilderness. The flower is no longer a simple passive victim in the busy bee's sweet pillage, but rather a conscious being, with hopes, aspirations, and companionships. The insect is its counterpart. Its fragrance is but a perfumed whisper of welcome, its color is as the wooing blush and rosy lip, its portals are decked for his coming, and its sweet hospitalities humored to his tarrying; and as it finally speeds its parting affinity rests content that its life's consummation has been fulfilled.
A HONEY-DEW PICNIC
Several of our notable as well as notorious human, social, and civic customs find their prehistoric prototypes in the insect kingdom. The monarchical institution sees its singular prophecy in the domestic economy of the bees. War and slavery have always been carried on systematically and effectually by ants, and, according to Huber and other authorities, agriculture, gardening, and an industry very like dairy farming have been time-honored customs among this same wise and thrifty insect tribe, whose claims to thoughtful consideration were so long ago voiced by Solomon of proverbial fame. Thevenot mentions "Solomon's ant" as among the "beasts which shall enter paradise." Indeed, the human saint as well as sluggard may "go to the ant" for many suggestive hints and commentaries.
These are only a few of the more notable parallelisms which suggest themselves. But others are not wanting if we care to follow the subject. In addition to the many models of thrift and virtuous industry, embodying types of many of the trade employments known to humanity, have we not also among these "meadow tribes" our luxurious "idlers" and "exquisites," the butterflies and flower-haunting flies and "dandy" beetles; and, opposed to all these, the suggestive antithesis of the promiscuous marauders, thieves, and brigands everywhere interspersed?
Thus we have our individual insect assassin and assassination organized in war; so, on the other hand, have we our insect merrymakers; why not, then, our picnic or carnival?
Such I am moved to call the singular episode which I observed last summer, and which I have endeavored to picture as true to the life as possible in the accompanying presentment The sceptic will perhaps remark on examination that the scene is characterized by somewhat too free a license to warrant the ideal of a "picnic." But he is hypercritical. There are picnics and picnics—picnics of high and of low degree. Do I not recall more than one notorious festive outing of the "next lower than the angels" in which the personnel seemed about similarly proportioned, and the fun and attraction comparatively related to the license?
One July afternoon a year ago I was returning home from one of my botanizing strolls. I had just emerged from a deep wood, and was skirting its border, when my attention was caught by a small fluttering swarm of butterflies, which started up at my approach and hovered about a blossoming blackberry bush a few yards in advance of me at the side of my path. The diversity of the butterfly species in the swarm struck me as singular, and the mere allurement of the blackberry blossoms—not usually of especial attraction to butterflies—could hardly explain so extensive a gathering. Here was the great yellow swallow-tail (Turnus), red admiral (Atlanta), small yellow butterfly (Philodice), white cabbage-butterfly, comma and semicolon, and numerous small fry, fluttering about me in evident protest against my intrusion. They showed no inclination to vacate the premises, so, in pursuance of one of the first articles of my saunterer's creed, I concluded to retreat softly a few paces and watch for developments. One by one the swarm sought their original haunt, settling on the bramble, and I now noticed that only in occasional instances did the insects seek the flowers, the attraction seeming to be confined to the leaves. I stole up softly for a nearer point of observation, and could now distinctly see the beautiful yellow and black open wings of the swallow-tail softly gliding or gently fluttering as it hung from the edge of a leaf, while it explored its surface with its uncoiled capillary tongue. Just beyond my Turnus, on another leaf, I now noted a new presence, the orange Aphrodite butterfly, silvery spotted, its nether wings being folded over its back, too much absorbed to have been startled by my first approach. Occasionally, without any cause which I could detect from my present position—certainly in no way connected with my presence—a small swarm of the butterflies would rise in a flutter above the bush, as though actuated by a common whim—a brief winged tangle in which a beautiful sprite of velvety black hovering in a globular halo, shot through with two white semicircular arcs, was always a momentary feature.
Carefully stealing through the tall grass, I now approached to within touching distance of the haunt, and was soon lost in mingled wonder, amusement, and surprise at the picnic now disclosed, the occasional butterfly swarm being now easily explained. From my first point of view only the top of the bramble spray was visible above the grass, and by far the most interesting portion of the exercises had been concealed from view. The butterflies, while naturally the most conspicuous element, were now seen to be in a small minority among the insect gathering, the bramble leaves being peopled with a most motley and democratic assemblage of insects. Class distinctions were apparently forgotten in the common enthusiasm; the plebeian bluebottle and blowfly now consorted with Aphrodite and sipped at the same drop. Many a leaf was begemmed with the blue bodies closely set side by side or in a close cluster. The meat-fly, house-fly, and horse-fly made themselves promiscuous in every portion of the spray, and what with the rainbow-eyed and ruby-eyed flies, black and silver-banded flower-flies, and other tiny, restless, iridescent atoms of the fly fraternity, the family of Musca was well represented at the feast.
Nor were these all the guests at the banquet—for banquet there certainly was, judging from the eager sipping and crowding everywhere upon the leaves, the flowers even yet, as I first noticed, seeming to have little attraction.
I have no direct means of knowing as to the social discrimination of the host as shown in the entertainment, for that invitations were issued the subsequent facts would show. But I have good reasons for believing, from the course of events, that the gathering included a number of questionable personages that were not counted upon.
Here, for instance, was an overwhelming contingent of the whole tough gang of wasps and hornets—brown wasps from under the eaves and fences; black hornets from the big paper nests; yellow-jackets from where you please; deep steel-blue wire-waisted wasps from the mud cells in the garret, to say nothing of an occasional longer-waisted digger-wasp, and a host of their allied lesser associates scattered around generously among the assemblage.
Every now and then a big darning-needle took a shimmering circuit about the bush, and doubtless knew what he was about; as did also what at first glimpse appeared to be a big bumblebee, which seemed to find attraction in the neighborhood, although he seldom alighted upon the leaves, preferring to sit upon a neighboring weed and watch his opportunities.
I have thus described a few of the more prominent guests or personages present at the feast. But I have reported little of their "goings on." Doubtless there were appropriate toasts and responses, or what in bug etiquette answered to this seemingly indispensable human fad, while as to that other festive social essential of after-dinner speeches, coupled in this case with most vigorous discussion, I am certain the air was blue with something of this sort, if the eloquent pantomime bore any significance. Here, for instance, is one isolated, but frequent, episode. A peaceable little group of plain bluebottle-flies, with but a single thought, are all sipping at the same drop in contentment. A brief respite, for now the tips of a pair of inquisitive antennae appear from the under edge of the leaf upon which they are sipping, and gingerly explore the upper surface. They are quickly followed by the covetous almond-eyed gaze of a brown wasp, that now steals cautiously around to the upper surface, and appears wholly engrossed in licking the leaf. Nearer and nearer he sidles up to the group of flies, and now with deliberate purpose and open jaws makes a dash among them. But they are too quick for him, and are away in a glittering blue tangle, which finally concentrates itself upon a neighboring leaf, where the eager tippling is immediately resumed. The wasp now holds the fort, and seems in no mood to be trifled with. With head and fore feet upraised and open jaws he seems "spoiling for a fight," and ready to make war upon the first comer. But no, he is evidently expecting a friend that, I now observe, approaches him determinedly down the stem of the leaf. The new-comer, a brown wasp like himself, is now at close range, and in an instant more, without any visible courteous preliminaries, the two set upon each other with a common enthusiasm, and with jaws working and stings fencing the interlocked combatants fall to the ground for a finish. I presume the affair was carried to the fourteenth round without any undue interference.
Another and another of these friendly meetings between them and other wasps took place in the half-hour in which I watched the sport. There were lulls in hostilities, during which an atmosphere of perfect peace and harmony seemed to reign around my bramble-bush. The flies were motionless in their ecstasy, and the hornet element seemed by common consent to keep temporarily shady, and even the butterflies seemed to forget that they had wings. But not for long, for now with a shimmering glitter our darning-needle invades the scene, and retires to a convenient perch with a ruby-eyed fly in his teeth, while a swarm of very startled butterflies tells conspicuously of the demoralization which he has left in his path. Among the butterfly representatives I at length observed one individual which at first had escaped me, an exclusive white cabbage-butterfly which sipped quietly at his leaf in the shade, and seemed to take little interest in the disreputable actions of his associates. Nothing could move him or entice him away from his convivial employment. But, alas! his folly soon found him out, for, on happening to look again, I observed he had found a new acquaintance—a hornet that had evidently been long desirous of meeting him. One by one I saw my butterfly's dismembered wings fall to the grassy jungle below, while a big black wasp proceeded to enjoy the collected sweets which he had doubtless observed were being so carefully stored away there in the shady retreat.
And now my pretty black butterfly—no, it proved to be the little day-flying grape-vine-moth, the eight-spotted black Alypia—appeared from some unseen source, and spun his crapy white-streaked halo among the leaves, at length settling among a little company of flies. Softly behind him creeps a brown wasp (Polistes), with his mouth watering, while from the opposite quarter a steel-blue mud-wasp approaches, with apparently similar designs. Neither invader sees the other. Simultaneously, as though answering to a signal, the two make a dash at the moth; but he is too quick for them. In a twinkling he is off in his pretty halo again, while the two disappointed contestants have clinched, and with stings and jaws vigorously plying fall to the jungle below, and seek satisfaction in mortal combat.
Here is a pretty little yellow and black banded flower-fly, which is having a quiet little picnic all by himself on a bed of yarrow bloom close by. But a big black paper-hornet has suddenly seen an attraction hither also, and is soon creeping stealthily among the blossoms with a wild and hungry look. But the hornets seemed to waste their time on the flies. Seemingly confident in their less complicated wing machinery, the two-winged fly rarely sought escape until within very close range of his enemy, and his resources never seemed to disappoint him at the critical moment.
Among the insect assemblage was a large number of ants of all kinds and sizes, the common large black species being conspicuous. Here is one creeping and sipping along a grass stem. A small digger-wasp likes this grass stem too, but instead of exchanging courtesies on the subject, the wasp proceeds to bite the ant's head off without ceremony, and continues sipping at the stem as though decapitation were a mere casual incident in its daily walk.
On the same stem a big blowfly has alighted. Judging from appearances, he has had his fill of good things, and is now making his leisurely toilet in the peculiar fashion of his kind, rubbing down his back and wings with his hind legs, twisting his front feet into spirals, and ever and anon testing the strength of his elastic neck attachment as he threatens to pull his head from his body.
This worldly act has been progressing for some moments under the gaze of a big black digger-wasp, who now concludes to cut it short. When at close range with his prey, the fly suddenly discovers the unhealthy location which he occupies, and actually protruding his tongue by way of parting salute, he is off with a buzz. He has barely taken wing, however, when a still louder buzz is heard, while a great black bumblebee follows closely in his wake, until the sounds of both are lost in the distance. The hum of this bumblebee is a frequent musical feature of the entertainment, and many is the dance that is set to its minstrelsy, as the burly insect darts in among the merrymakers and is off to his perch near by. It is only as we steal away and observe him closely that we learn the secret of his occasional sorties. There on a clover blossom he sits—sipping honey? Oh no. It is honey-dew that he is enjoying, and second-hand at that, as he devours the satiated bluebottle-fly which is empaled on his black horny beak. For this is only a bumblebee in masquerade—a carnivorous fly, in truth, which, safe in its disguise of respectability, hovers in the flowery haunts of the innocents and, of course, reaps his reward.
And what is this? A yellow-jacket has found an ambrosial attraction here upon the bramble leaf. Meanwhile a great black and white paper-hornet has seen his opportunity, and is soon slyly approaching behind the sipper. That he has designs on that jacket and its contents is apparent. In a moment the onslaught is consummated, and in the struggle which ensues the black assailant relieves his victim—of his watch presumably, for he has captured the entire garment, which he soon rifles and discards with some show of satisfaction.
And so my carnival proceeds. So it began with the dawn; so it will continue till dusk; and through the night, with new revels, for aught I know, and will be prolonged for days or weeks.
* * * * *
Reflective reader, how often, as you have strolled through some nook in the suburban wood, have you paused in philosophic mood at the motley relics of good cheer which sophisticated the retreat, so pathetically eloquent of pristine joys to which you had been a stranger? Here in my present picnic is the suggestive parallel, for even though no such actual episodes as those I have described had been witnessed by me, an examination of the premises beneath my bramble were a sufficient commentary. These were the unimpeachable witnesses of the pleasures which I have pictured. Dismembered butterfly wings strewed the grassy jungle, among which were a fair sprinkling from that black and white halo already noted. Occasional dead wasps and detached members of wasp and hornet anatomy were frequent, while the blue glitter of the bodies of flies lit up a shadowy recess here and there, showing that Musca had not always so correctly gauged his comparative wing resources as my observation had indicated.
It was interesting to discover, too, down deep among the herbage, another suggestive fact in the presence of a shrewd spider that showed a keen eye to the main chance, and had spread his gossamer catch-all beneath the bramble. It was all grist into his mill, and no doubt his charnel-house at the base of his silken tunnel could have borne eloquent testimony alike to his wise sagacity and his epicurean luxury.
I have pictured my picnic, and the question naturally arises, what was it all about—what the occasion for this celebration? There was certainly no distinct visible cause for the social gathering upon this particular bramble-bush. There were a number of other bramble-bushes in the near neighborhood which, it would seem, should possess equal attractions, but which were ignored. In what respect did the one selected differ from the others?
This bramble had become the scene of my carnival simply because it chanced to be directly beneath an overhanging branch of pine some twenty feet above. Here dwelt mine host who had issued the invitations and spread the feast, the limb for about a foot space being surrounded by a colony of aphides, or plant-lice, from whose distilling pipes the rain of sweet honey-dew had fallen ceaselessly upon the leaves below. The flies, butterflies, and ants had been attracted, as always, by its sweets; the preoccupied convivial flies, in turn, were a tempting bait for the wasps and hornets, and my dragon-fly and mock bumblebee found a similar attraction in the neighborhood.
An examination of the trunk of the pine showed the inevitable double procession of ants, both up and down the tree, with the habitual interchange of comment; and could we but have obtained a closer glimpse of the pine branch above, we might certainly have observed the queer spectacle of the small army of ants interspersed everywhere among the swarm of aphides. Not in antagonism; indeed, quite the reverse; herders, in truth, jealously guarding their feeding flock, creeping among them with careful tread, caressing them with their antennae while they sipped at the honeyed pipes everywhere upraised in most expressive and harmonious welcome.
This intimate and friendly association of the ants and aphides has been the subject of much interesting scientific investigation and surprising discovery. Huber and Lubbock have given to the world many startling facts, the significance of which may be gathered from the one statement that certain species of ants carry their devotion so far as literally to cultivate the aphides, carrying them bodily into their tunnels, where they are placed in underground pens, reared and fed and utilized in a manner which might well serve as a pattern for the modern dairy farm. Indeed, after all that we have already seen upon a single bramble-bush, would it be taking too much license with fact to add one more pictorial chronicle—an exhilarated and promiscuous group of butterflies, ants, hornets, wasps, and flies uniting in "a health to the jolly aphis"?
A FEW NATIVE ORCHIDS AND THEIR INSECT SPONSORS
In a previous article I discussed the general subject of the fertilization of flowers, briefly outlining the several historical and chronological steps which ultimately led to Darwin's triumphant revelation of the divine plan of "cross-fertilization" as the mystery which had so long been hidden beneath the forms and faces of the flowers.
In the same paper I presented many illustrative examples among our common wild flowers possessing marvellous evolved devices, mechanisms, and peculiarities of form by which this necessary cross-fertilization was assured.
Prior to Darwin's time the flower was a voice in the wilderness, heard only in faintest whispers, and by the few. But since his day they have bloomed with fresher color and more convincing perfume. Science brought us their message. Demoralizing as it certainly was to humanity's past ideals, philosophic, theologic, and poetic, it bore the spirit of absolute conviction, and must be heard.
What a contrast this winged botany of to-day to that of a hundred years ago! The flower now no longer the mere non-committal, structural, botanical specimen. No longer the example of mere arbitrary, independent creation, reverently and solely referred to the orthodox "delight of man." The blossom whose unhappy fate was bemoaned by the poet because, forsooth, it must needs "blush unseen," or "waste its sweetness on the desert air," is found alone in that musty hortus siccus of a blind and deluded past. From the status of mere arbitrary creation, however "beautiful," "curious," "eccentric," hitherto accepted alone on faith—"it is thus because it is created thus: what need to ask the reason why?"—it has become a part of our inspiring heritage, a reasonable, logical, comprehensible result, a manifestation of a beautiful divine scheme, and is thus an ever-present witness and prophet of divine care and supervision.
The flower of to-day! What an inspiration to our reverential study! What a new revelation is borne upon its perfume! Its forms and hues, what invitations to our devotion! This spot upon the petal; this peculiar quality of perfume or odor; this fringe within the throat; this curving stamen; this slender tube! What a catechism to one who knows that each and all represent an affinity to some insect, towards whose vital companionship the flower has been adapting itself through the ages, looking to its own more certain perpetuation!
The great Linnaeus would doubtless have claimed to "know" the "orchid," which perhaps he named. Indeed, did he not "know" it to the core of its physical, if not of its physiological, being? But could he have solved the riddle of the orchid's persistent refusal to set a pod in the conservatory? Could he have divined why the orchid blossom continues in bloom for weeks and weeks in this artificial glazed tropic—perhaps weeks longer than its more fortunate fellows left behind in their native haunts—and then only to wither and perish without requital? Know the orchid?—without the faintest idea of the veritable divorce which its kidnapping had involved!
Thanks to the new dispensation, we may indeed claim a deeper sympathy with the flower than is implied in a mere recognition of its pretty face. We know that this orchid is but the half of itself, as it were; that its color, its form, however eccentric and incomprehensible, its twisted inverted position on its individual stalk-like ovary, its slender nectary, its carefully concealed pollen—all are anticipations of an insect complement, a long-tongued night-moth perhaps, with whose life its own is mysteriously linked through the sweet bond of perfume and nectar, and in the sole hope of posterity.
And the flower had been stolen from its haunt while its consort slept, and had awakened in a glazed prison—doubtless sufficiently comfortable, save for the absence of that one indispensable counterpart, towards whom we behold in the blossom's very being the embodied expression of welcome.
Blooming day after day in anticipation of his coming, and week after week still hoping against hope, we see the flower fade upon its stalk, and with what one might verily believe to be evidences of disconsolation, were it not that the ultra-scientist objects to such a sentimental assumption with regard to a flower, which is unfortunate enough to show no sign of nerves or gray matter in its composition. Who shall claim to know his orchid who knows not its insect sponsor?
To take one of our own wild species. Here is the Arethusa bulbosa of Linnaeus, for instance. Its pollen must reach its stigma—so he supposed—in order for the flower to become fruitful. But this is clearly impossible, as the pollen never leaves its tightly closed box unless removed by outside aid, which aid must also be required to place it upon the stigma. This problem, which confronted him in practically every orchid he met, Linnaeus, nor none of his contemporaries, nor indeed his followers for many years, ever solved.
Not until the time of Christian Conrad Sprengel (1735) did this and other similar riddles begin to be cleared up, that distinguished observer having been the first to discover in the honey-sipping insect the key to the omnipresent mystery. Many flowers, he discovered, were so constructed or so planned that their pollen could not reach their own stigmas, as previously believed. The insect, according to Sprengel, enjoyed the anomalous distinction of having been called in, in the emergency, to fulfil this apparent default in the plain intentions of nature, as shown in the flower. Attracted by the color and fragrance of the blossom, with their implied invitation to the assured feast of nectar, the insect visited the flower, and thus became dusted with the pollen, and in creeping or flying out from it conveyed the fecundating grains to the receptive stigma, which they could not otherwise reach. Such was Sprengel's belief, which he endeavored to substantiate in an exhaustive volume containing the result of his observations pursuant to this theory.
But Sprengel had divined but half the truth. The insect was necessary, it was true, but the Sprengel idea was concerned only with the individual flower, and the great botanist was soon perplexed and confounded by an opposing array of facts which completely destroyed the authority of his work—facts which showed conclusively that the insect could not thus convey the pollen as described, because the stigma in the flower was either not yet ready to receive it—perhaps tightly closed against it—or was past its receptive period, even decidedly withered.
This radical assumption of fertilization in the individual flower, which lay at the base of Sprengel's theory, thus so completely exposed as false, discredited his entire work. The good was condemned with the bad, and the noble volume was lost in comparative oblivion—only to be finally resurrected and its full value and significance revealed by the keen scientific insight of Darwin (1859). From the new stand-point of evolution through natural selection the facts in Sprengel's work took on a most important significance. Darwin now reaffirmed the Sprengel theory so far as the necessity of the insect was concerned, but showed that all those perplexing floral conditions which had disproved Sprengel's assumption, instead of having for their object the conveying of pollen to the stigma of the same flower, implied its transfer to the stigma of another, cross-fertilization being the evident design, or evolved and perpetuated advantage.
This solution was made logical and tenable only on the assumption that such evolved conditions, insuring cross-fertilization, were of distinct advantage to the flower in the competitive struggle for existence, and that all cross-fertilized flowers were thus the final result of natural selection.
The early ancestors of this flower were self-fertilized; a chance seedling at length, among other continual variations, showed the singular variation of ripening its stigma in advance of its pollen—or other condition insuring cross-fertilization—thus acquiring a strain of fresh vigor. The seedlings of this flower, coming now into competition with the existing weaker self-fertilized forms, by the increased vigor won in the struggle of their immediate surroundings, and inheriting the peculiarity of their parent, showed flowers possessing the same cross-fertilizing device. The seeds from these, again scattering, continued the unequal struggle in a larger and larger field and in increasing numbers, continually crowding out all their less vigorous competitors of the same species, at length to become entire masters of the field and the only representatives left to perpetuate the line of descent.
Thus we find in almost every flower we meet some astonishing development by which this cross-fertilization is effected, by which the transferrence of the pollen from one flower to the stigma of another is assured, largely through the agency of insects, frequently by the wind and water, occasionally by birds. In many cases this is assured by the pollen-bearing flowers and stigmatic flowers being entirely distinct, as in cucumbers and Indian-corn; perhaps on different plants, as in the palms and willows; again by the pollen maturing and disseminating before the stigma is mature, as already mentioned, and vice versa.
From these, the simplest forms, we pass on to more and more complicated conditions, anomalies of form and structure—devices, mechanisms, that are past belief did we not observe them in actuality with our own eyes, as well as the absolutely convincing demonstration of the intention embodied: exploding flowers, shooting flowers, flower-traps, stamen embraces, pollen showers, pollen plasters, pollen necklaces, and floral pyrotechnics—all demonstrations in the floral etiquette of welcome and au revoir to insects.
From the simplest and regular types of flowers, as in the buttercup, we pass on to more and more involved and unsymmetrical forms, as the columbine, monk's-hood, larkspur, aristolochia, and thus finally to the most highly specialized or involved forms of all, as seen in the orchid—the multifarious, multiversant orchid; the beautiful orchid; the ugly orchid; the fragrant orchid; the fetid orchid; the graceful, homely, grotesque, uncanny, mimetic, and, until the year 1859, the absolutely non-committal and inexplicable flower; the blossom which had waited through the ages for Darwin, its chosen interpreter, ere she yielded her secret to humanity.
And what is an orchid? How are we to know that this blossom which we plucked is an orchid? The average reader will exclaim, "Because it is an air-plant"—the essential requisite, it would seem, in the popular mind. Of over 3000 known species of orchids, it is true a great majority are air-plants, or epiphytes—growing upon trees and other plants, obtaining their sustenance from the air, and not truly parasitic; but of the fifty-odd native species of the northeastern United States, not one is of this character, all growing in the ground, like other plants. It is only by the botanical structure of the flowers that the orchid may be readily distinguished, the epiphytic character being of little significance botanically.
A brief glance at this structural peculiarity may properly precede our more elaborate consideration of a few species of these remarkable flowers.
The orchids are usually very irregular, and six-parted. The ovary is one-celled, and becomes a pod containing an enormous yield of minute, almost spore-like, seeds (Fig. 3) in some species, as in the vanilla pod, to the number of a million, and in one species of the maxillaria, as has been carefully computed, 1,750,000.
The pollen, unlike ordinary flowers, is gathered together in waxy masses of varying consistency, variously formed and disposed in the blossom, its grains being connected with elastic cobwebby threads, which occasionally permit the entire mass to be stretched to four or five times its length, and recover its original shape when released. This is noticeable specially in the O. spectabilis, later described. The grains thus united are readily disentangled from their mass when brought into contact with a viscid object, as, for instance, the stigma.
But the most significant botanical contrast and distinction is found in the union of the style and stamens in one organ, called the column (Fig. 2), the stigma and the pollen being thus disposed upon a single common stalk. The contrast to the ordinary flower will be readily appreciated by comparison of the accompanying diagrams (Fig. 1).
When, therefore, we find a blossom with the anthers or pollen receptacle united to a stalk upon which the stigma is also placed, we have an orchid.
The order is further remarkable, as Darwin first demonstrated in his wonderful volume "The Fertilization of Orchids," in that the entire group, with very few exceptions, are absolutely dependent upon insects for their perpetuation through seed. They possess no possible resource for self-fertilization in the neglect of these insect sponsors.
Many of our common wild flowers, as perfectly and effectually planned for cross-fertilization as the orchids, do retain the reserve power of final self-fertilization if unfertilized by foreign pollen.
But the orchid has lost such power, and in the progress of evolution has gradually adapted itself to the insect, often to a particular species of insect, its sole sponsor, which natural selection has again gradually modified in relation to the flower.
The above work by Darwin was mostly concerned with foreign species, generally under artificial cultivation, and so startling were the disclosures concerning these hitherto sphinx-like floral beings that a most extensive bibliography soon attested the widespread inspiration and interest awakened by its pages.
But it is by no means necessary to visit the tropics or the conservatory for examples of these wonders. Our own Asa Gray, one of Darwin's instant proselytes, was prompt to demonstrate that the commonest of our native American species might afford revelations quite as astonishing as those exotic species which Darwin had described.
During a period of many years the writer has devoted much study to our native species of orchids from this evolutionary stand-point of their cross-fertilization tendencies. Of the following examples, selected from his list, some are elaborations of previous descriptions of Gray and others, though pictorially and descriptively the result of direct original study from nature; others are from actual observation of the insects at work on the flowers; and others still, original demonstrations based upon analogy and the obvious intention of the floral construction, the action of the insect—its head or tongue—having been artificially imitated by pins, bristles, or other probe-like bodies.
How many an enthusiastic flower-hunter has plucked his fragrant bouquet of the beautiful Arethusa, in its sedgy haunt, without a suspicion of the beautiful secret which lay beneath its singular form! Indeed, how many a learned botanist, long perfectly familiar with its peculiarities of shape and structure, has been entirely content with this simple fact, nor cared to seek further for its interpretation! But
"All may have the flower now, For all have got the seed."
With Darwin as our guide and the insect as our key—an open sesame—the hidden treasure is revealed. It is now quite possible, as Darwin demonstrated, to look upon a flower for the first time and from its structure foretell the method of its intended cross-fertilization; nay, more, possibly the kind, or even the species, of insect to which this cross-fertilization is intrusted.
Let us look at our Arethusa. The writer has never happened to observe an insect at work upon this flower, but the intention of its structure is so plain that by a mere examination we may safely prophesy not only what must happen when the insect seeks its nectar, but with equal assurance the kind of insect thus invited and expected. I have indicated a group of the orchids in their usual marshy haunt, and in Fig. 4, separately, a series of diagrams presents sections of the flower, natural size and duly indexed, which renders detailed description hardly necessary. The column is here quite elongated, forked at the tip, the space between the forks occupied by the anther, which is hinged to the upper division. This anther lid is closed tightly, with the sticky mass of pollen hidden behind it in the cavity. The stigma is on the external inner side of the lower division, and thus distinctly separated from the pollen. The "lip" is extended forward as a hospitable threshold to the insect. And to what insect might we assume this invitation of color, fragrance, nectar, and threshold to be extended?
Let us consider the flower simply as a device to insure its own cross-fertilization. The insect is welcomed; it must alight and sip the nectar; in departing it must bear away this pollen upon its body, and convey it to the next Arethusa blossom which it visits, and leave it upon its stigma. These are the conditions expressed; and how admirably they are fulfilled we may observe when we examine flower after flower of a group, and find their nectaries drained, their anther cells empty, and pollen upon all their stigmas. The nectar is here secreted in a well—not very deep—and the depth of this nectar from the entrance is of great significance among all the flowers, having distinct reference to the length of the tongue which is expected to sip it. In the Arethusa, it is true, the butterfly or moth might sip at the throat of the flower, but the long tongues of these insects might permit the nectary to be drained without bringing their bodies in contact with the stigma. Smaller insects might creep into the nectary and sip without the intended fulfilment. It is clear that to neither of such visitors is the welcome extended. What, then, are the conditions embodied? The insect must have a tongue of such a length that, when in the act of sipping, its head must pass beyond the anther well into the opening of the flower. Its body must be sufficiently large to come in contact with the anther. Such requisites are perfectly fulfilled by the humblebee, and we may well hazard the prophecy that the Bombus is the welcomed affinity of the flower.
The diagrams (Fig. 4) sufficiently illustrate the efficacy of the beautiful plan involved. At A the bee is seen sipping the nectar. His forward movement thus far to this point has only seemed to press the edge of the anther inward, and thus keep it even more effectually closed. As the bee retires (B), the backward motion opens the lid, and the sticky pollen is thus brought against the insect's back, where it adheres in a solid mass. He now flies to the next Arethusa blossom, enters it as before, and in retiring slides his back against the receptive viscid stigma, which retains a portion of the pollen, and thus effects the cross-fertilization (C). Professor Gray surmised that the pollen was withdrawn on the insect's head, and it might be so withdrawn, but in other allied orchids of the tribe Arethusae, however, in which the structure is very similar, the pollen is deposited on the thorax, and such is probably the fact in this species. In either case cross-fertilization would be effected. Nothing else is possible in the flower, and whether it is Bombus or not that effects it, the method is sufficiently evident.
Having thus had one initiation into this most enticing realm of riddles, each successive orchid whose structure we examine from this stand-point becomes a most interesting, perhaps a fresh, problem, whose assumed solution may often be verified by studying the insect in its haunts. Darwin thus foretold the precise manner of the cross-fertilization of Habenaria mascula, and also the insect agent, simply by the structural prophecy of the flower itself.
Suppose, for example, an unknown orchid blossom to be placed in our hands. Its nectary tube is five inches in length, and as slender as a knitting-needle. The nectar is secreted far within its lip. The evolution of the long nectary implies an adaptation to an insect's tongue of equal length. What insect has a tongue five inches long, and sufficiently slender to probe this nectary? The sphinx-moth only. Hence we infer the sphinx-moth to be the insect complement to the blossom, and we may correctly infer, moreover, that the flower is thus a night-bloomer. Examination of the flower, with the form of this moth in mind, will show other adaptations to the insect's form in the position of pollen and stigma, looking to the flower's cross-fertilization. In some cases this is effected by the aid of the insect's tongue; in others, by its eyes.
In our own native orchids we have a remarkable example of the latter form in the Habenaria orbiculata, whose structure and mechanism have also been admirably described by Asa Gray.
All orchid-hunters know this most exceptional example of our local flora, and the thrill of delight experienced when one first encounters it in the mountain wilderness, its typical haunt, is an event to date from—its two great, glistening, fluted leaves, sometimes as large as a dinner-plate, spreading flat upon the mould, and surmounted by the slender leafless stalk, with its terminal loose raceme of greenish-white bloom.
A single blossom of the species is shown in Fig. 5, the parts indexed. The opening to the nectary is seen just below the stigmatic surface, the nectary itself being nearly two inches in length. The pollen is in two club-like bodies, each hidden within a fissured pouch on either side of the stigma, and coming to the surface at the base in their opposing sticky discs as shown. Many of the group Habenaria or Platanthera, to which this flower belongs, are similarly planned. But mark the peculiarly logical association of the parts here exhibited. The nectary implies a welcome to a tongue two inches long, and will reward none other. This clearly shuts out the bees, butterflies, and smaller moths. What insect, then, is here implied? The sphinx-moth again, one of the lesser of the group. A larger individual might sip the nectar, it is true, but its longer tongue would reach the base of the tube without effecting the slightest contact with the pollen, which is of course the desideratum here embodied, and which has reference to a tongue corresponding to the length of the nectary. There are many of these smaller sphinxes. Let us suppose one to be hovering at the blossom's throat. Its slender capillary tongue enters the opening. Ere it can reach the sweets the insect's head must be forced well into the throat of the blossom, where we now observe a most remarkable special provision, the space between the two pollen discs being exactly adjusted to the diameter of the insect's head. What follows this entrance of the moth is plainly pictured in the progressive series of illustrations (Fig. 6). A represents the insect sipping; the sticky discs are brought in contact with the moth's eyes, to which they adhere, and by which they are withdrawn from their pouches as the moth departs (B). At this time they are in the upright position shown at C, but in a few seconds bend determinedly downward and slightly towards each other to the position D. This change takes place as the moth is flitting from flower to flower. At E we see the moth with its tongue entering the nectary of a subsequent blossom. By the new position of the pollen clubs they are now forced directly against the stigma (E). This surface is viscid, and as the insect leaves the blossom retains the grains in contact (F), which in turn withdraw others from the mass by means of the cobwebby threads by which the pollen grains are continuously attached. At G we see the orchid after the moth's visit—the stigma covered with pollen, and the flower thus cross-fertilized.