Chapter 5

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or non-committal than most of these?--their under surfaces hugging close against the bark, their telltale feet closely withdrawn, and all their pointed helmets inclined in the same parallel direction.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, Köhlreuter, Herbert, Darwin, Lubbock, Müller, 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 Linnæus, 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 Linnæus 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 Linnæus 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 Linnæus (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 Müller 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), Köhlreuter (1811), Herbert (1837), Gärtner (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, Linnæus, 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 Müller, 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.