Tropical nature, and other essays

by Alfred Russel Wallace

Light has a much more decided action on plants than on animals. The green colour of leaves is almost wholly dependent on it; and although some flowers will become fully coloured in the dark, others are decidedly affected by the absence of light, even when the foliage is fully exposed to it. Looking therefore at the numerous colouring matters which are developed in the tissues of plants, the sensitiveness of these pigments to light, the changes they undergo during growth and development, and the facility with which new chemical combinations are effected by the physiological processes of plants as shown by the endless variety in the chemical constitution of vegetable products, we have no difficulty in comprehending the general causes which aid in producing the colours of the vegetable world, or the extreme variability of those colours. We may therefore here confine ourselves to an inquiry into the various uses of colour in the economy of plants; and this will generally enable us to understand how it has become fixed and specialised in the several genera and species of the vegetable kingdom.

Protective Coloration and Mimicry in Plants.—In animals, as we have seen, colour is greatly influenced by the need of protection from, or of warning to, their numerous enemies, and by the necessity for identification and easy recognition. Plants rarely need to be concealed, and obtain protection either by their spines, their hardness, their hairy covering, or their poisonous secretions. A very few cases of what seem to be true protective colouring do, however, exist; the most remarkable being that of the “stone mesembryanthemum,” of the Cape of Good Hope, which, in form and colour closely resembles the stones among which it grows; and Dr. Burchell, who first discovered it, believes that the juicy little plant thus generally escapes the notice of cattle and wild herbivorous animals. Mr. J. P. Mansel Weale also noticed that many plants growing in the stony Karoo have their tuberous roots above the soil; and these so perfectly resemble the stones among which they grow that, when not in leaf, it is almost impossible to distinguish them (Nature, vol. iii. p. 507). A few cases of what seems to be protective mimicry have also been noted; the most curious being that of three very rare British fungi, found by Mr. Worthington Smith, each in company with common species which they so closely resembled that only a minute examination could detect the difference. One of the common species is stated in botanical works to be “bitter and nauseous,” so that it is not improbable that the rare kind may escape being eaten by being mistaken for an uneatable species, though itself palatable. Mr. Mansel Weale also mentions a labiate plant, the Ajuga-ophrydis, of South Africa, as strikingly resembling an orchid. This may be a means of attracting insects to fertilize the flower in the absence of sufficient nectar or other attraction in the flower itself; and the supposition is rendered more probable by this being the only species of the genus Ajuga in South Africa. Many other cases of resemblances between very distinct plants have been noticed—as that of some Euphorbias to Cacti; but these very rarely inhabit the same country or locality, and it has not been proved that there is in any of these cases the amount of inter-relation between the species which is the essential feature of the protective “mimicry” that occurs in the animal world.

The different colours exhibited by the foliage of plants and the changes it undergoes during growth and decay, appear to be due to the general laws already sketched out, and to have little if any relation to the special requirements of each species. But flowers and fruits exhibit definite and well-pronounced tints, often varying from species to species, and more or less clearly related to the habits and functions of the plant. With the few exceptions already pointed out, these may be generally classed as attractive colours.

Attractive Colours of Fruits.—The seeds of plants require to be dispersed, so as to reach places favourable for germination and growth. Some are very minute, and are carried abroad by the wind; or they are violently expelled and scattered by the bursting of the containing capsules. Others are downy or winged, and are carried long distances by the gentlest breeze; or they are hooked and stick to the fur of animals. But there is a large class of seeds which cannot be dispersed in either of these ways, and they are mostly contained in eatable fruits. These fruits are devoured by birds or beasts, and the hard seeds pass through their stomachs undigested, and, owing probably to the gentle heat and moisture to which they have been subjected, in a condition highly favourable for germination. The dry fruits or capsules containing the first two classes of seeds are rarely, if ever, conspicuously coloured; whereas the eatable fruits almost invariably acquire a bright colour as they ripen, while at the same time they become soft and often full of agreeable juices. Our red haws and hips, our black elderberries, our blue sloes, and whortleberries, our white mistletoe and snowberry, and our orange sea-buckthorn, are examples of the colour-sign of edibility; and in every part of the world the same phenomenon is found. Many such fruits are poisonous to man and to some animals, but they are harmless to others; and there is probably nowhere a brightly-coloured pulpy fruit which does not serve as food for some species of bird or mammal.

Protective Colours of Fruits.—The nuts and other hard fruits of large forest-trees, though often greedily eaten by animals, are not rendered attractive to them by colour, because they are not intended to be eaten. This is evident; for the part eaten in these cases is the seed itself, the destruction of which must certainly be injurious to the species. Mr. Grant Allen, in his ingenious work on Physiological Æsthetics, well observes that the colours of all such fruits are protective—green when on the tree, and thus hardly visible among the foliage, but turning brown as they ripen and fall on the ground, as filberts, chestnuts, walnuts, beech-nuts, and many others. It is also to be noted that many of these are specially though imperfectly protected; some by a prickly coat as in the chestnuts, or by a nauseous covering as in the walnut; and the reason why the protection is not carried further is probably because it is not needed, these trees producing such vast quantities of fruit, that however many are eaten, more than enough are always left to produce young plants. In the case of the attractively coloured fruits, it is curious to observe how the seeds are always of such a nature as to escape destruction when the fruit itself is eaten. They are generally very small and comparatively hard, as in the strawberry, gooseberry, and fig; if a little larger, as in the grape, they are still harder and less eatable; in the fruit of the rose (or hip) they are disagreeably hairy; in the orange tribe excessively bitter. When the seeds are larger, softer, and more eatable, they are protected by an excessively hard and stony covering, as in the plum and peach tribe; or they are inclosed in a tough horny core, as with crabs and apples. These last are much eaten by swine, and are probably crushed and swallowed without bruising the core or the seeds, which pass through their bodies undigested. These fruits may also be swallowed by some of the larger frugivorous birds; just as nutmegs are swallowed by pigeons for the sake of the mace which incloses the nut, and which by its brilliant red colour is an attraction as soon as the fruit has split open, which it does upon the tree.

There is, however, one curious case of an attractively coloured seed which has no soft eatable covering. The Abrus precatoria, or “rosary bean,” is a leguminous shrub or small tree growing in many tropical countries, whose pods curl up and split open on the tree, displaying the brilliant red seeds within. It is very hard and glossy, and is said to be, as no doubt it is, “very indigestible.” It may be that birds, attracted by the bright colour of the seeds, swallow them, and that they pass through their bodies undigested, and so get dispersed. If so it would be a case among plants analogous to mimicry among animals—an appearance of edibility put on to deceive birds for the plant’s benefit. Perhaps it succeeds only with young and inexperienced birds, and it would have a better chance of success, because such deceptive appearances are very rare among plants.

The smaller plants whose seeds simply drop upon the ground, as in the grasses, sedges, composites, umbelliferæ, &c., always have dry and obscurely-coloured capsules and small brown seeds. Others whose seeds are ejected by the bursting open of their capsules, as with the oxalis and many of the caryophyllaceæ, scrophulariaceæ, &c., have their seeds very small and rarely or never edible.

It is to be remarked that most of the plants whose large-seeded nuts cannot be eaten without destroying their germinating power—as the oaks, beeches, and chestnuts—are trees of large size which bear great quantities of fruit, and that they are long lived and have a wide geographical range. They belong to what are called dominant groups, and are thus able to endure having a large proportion of their seeds destroyed with impunity. It is a suggestive fact that they are among the most ancient of known dicotyledonous plants—oaks and beeches going back to the Cretaceous period with little change of type, so that it is not improbable that they may be older than any fruit-eating mammal adapted to feed upon their fruits. The attractive coloured fruits on the other hand, having so many special adaptations to dispersal by birds and mammals, are probably of more recent origin.[21] The apple and plum tribes are not known earlier than the Miocene period; and although the record of extinct vegetable life is extremely imperfect, and the real antiquity of these groups is no doubt very much greater, it is not improbable that the comparative antiquity of the fruit-bearing and nut-bearing trees may remain unchanged by further discoveries, as has almost always happened as regards the comparative antiquity of animal groups.

[21] I owe this remark to Mr. Grant Allen, author of Physiological Æsthetics.

Attractive Colours of Flowers.—The colours of flowers serve to render them visible and recognizable by insects, which are attracted by secretions of nectar or pollen. During their visits for the purpose of obtaining these products, insects involuntarily carry the pollen of one flower to the stigma of another, and thus effect cross-fertilization; which, as Mr. Darwin was the first to demonstrate, immensely increases the vigour and fertility of the next generation of plants. This discovery has led to the careful examination of great numbers of flowers; and the result has been that the most wonderful and complex arrangements have been found to exist, all having for their object to secure that flowers shall not be self-fertilized perpetually, but that pollen shall be carried, either constantly or occasionally, from the flowers of one plant to those of another. Mr. Darwin himself first worked out the details in orchids, primulas, and some other groups; and hardly less curious phenomena have since been found to occur even among some of the most regularly-formed flowers. The arrangement, length, and position of all the parts of the flower is now found to have a purpose, and not the least remarkable portion of the phenomenon is the great variety of ways in which the same result is obtained. After the discoveries with regard to orchids, it was to be expected that the irregular, tubular, and spurred flowers should present various curious adaptations for fertilization by insect-agency. But even among the open, cup-shaped, and quite regular flowers, in which it seemed inevitable that the pollen must fall on the stigma and produce constant self-fertilization, it has been found that this is often prevented by a physiological variation—the anthers constantly emitting their pollen either a little earlier or a little later than the stigmas of the same flower, or of other flowers on the same plant, were in the best state to receive it; and as individual plants in different stations, soils, and aspects, differ somewhat in the time of flowering, the pollen of one plant would often be conveyed by insects to the stigmas of some other plant in a condition to be fertilized by it. This mode of securing cross-fertilization seems so simple and easy, that we can hardly help wondering why it did not always come into action, and so obviate the necessity for those elaborate, varied, and highly complex contrivances found perhaps in the majority of coloured flowers. The answer to this of course is, that variation sometimes occurred most freely in one part of a plant’s organization, and sometimes in another; and that the benefit of cross-fertilization was so great that any variation that favoured it was preserved, and then formed the starting-point of a whole series of further variations, resulting in those marvellous adaptations for insect-fertilization, which have given much of their variety, elegance, and beauty, to the floral world. For details of these adaptations we must refer the reader to the works of Darwin, Lubbock, Hermann Müller, and others. We have here only to deal with the part played by colour, and by those floral structures in which colour is most displayed.