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CHAPTER XI

THE SPECIAL COLOURS OF PLANTS: THEIR ORIGIN

AND PURPOSE

The general colour relations of plants-Colours of fruits-The meaning of nuts-Edible or attractive fruits-The colours of flowers-Modes of securing cross-fertilisation-The interpretation of the facts-Summary of additional facts bearing on insect fertilisation-Fertilisation of flowers by birds-Self-fertilisation of flowers-Difficulties and contradictions-Intercrossing not necessarily advantageous-Supposed evil results of close interbreeding-How the struggle for existence acts among flowers-Flowers the product of insect agency-Concluding remarks on colour in nature.

THE colours of plants are both less definite and less complex than are those of animals, and their interpretation on the principle of utility is, on the whole, more direct and more easy. Yet here, too, we find that in our investigation of the uses of the various colours of fruits and flowers, we are introduced to some of the most obscure recesses of nature's workshop, and are confronted with problems of the deepest interest and of the utmost complexity.

So much has been written on this interesting subject since Mr. Darwin first called attention to it, and its main facts have become so generally known by means of lectures, articles, and popular books, that I shall give here a mere outline sketch, for the purpose of leading up to a discussion of some of the more fundamental problems which arise out of the facts, and which have hitherto received less attention than they deserve.

The General Colour Relations of Plants.

The green colour of the foliage of leafy plants is due to the existence of a substance called chlorophyll, which is almost universally developed in the leaves under the action of light. It is subject to definite chemical changes during the processes of growth and of decay, and it is owing to these changes that we have the delicate tints of spring foliage, and the more varied, intense, and gorgeous hues of autumn. But these all belong to the class of intrinsic or normal colours, due to the chemical constitution of the organism; as colours they are unadaptive, and appear to have no more relation to the wellbeing of the plants themselves than do the colours of gems and minerals. We e may also include in the same category those alga and fungi which have bright colours-the "red snow of the arctic regions, the red, green, or purple seaweeds, the brilliant scarlet, yellow, white, or black agarics, and other fungi. All these colours are probably the direct results of chemical composition or molecular structure, and, being thus normal products of the vegetable organism, need no special explanation from our present point of view; and the same remark will apply to the varied tints of the bark of trunks, branches, and twigs, which are often of various shades of brown and green, or even vivid reds or yellows.

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There are, however, a few cases in which the need of protection, which we have found to be so important an agency in modifying the colours of animals, has also determined those of some of the smaller members of the vegetable kingdom. Dr. Burchell found a mesembryanthemum in South Africa like a curiously shaped pebble, closely resembling the stones among which it grew ;1 and Mr. J. P. Mansel Weale states that in the same country one of the Asclepiadeæ has tubers growing above ground among stones which they exactly resemble, and that, when not in leaf, they are for this reason quite invisible.2 It is clear that such resemblances must be highly useful to these plants, inhabiting an arid country abounding in herbivorous mammalia, which, 1 Burchell's Travels, vol. i. p. 10. 2 Nature, vol. iii. p. 507.

in times of drought or scarcity, will devour everything in the shape of a fleshy stem or tuber.

True mimicry is very rare in plants, though adaptation to like conditions often produces in foliage and habit a similarity that is deceiving. Euphorbias growing in deserts often closely resemble cacti. Seaside plants and high alpine plants of different orders are often much alike; and innumerable resemblances of this kind are recorded in the names of plants, as Veronica epacridea (the veronica like an epacris), Limnanthemum nymphæoides (the limnanthemum like a nymphæa), the resembling species in each case belonging to totally distinct families. But in these cases, and in most others that have been observed, the essential features of true mimicry are absent, inasmuch as the one plant cannot be supposed to derive any benefit from its close resemblance to the other, and this is still more certain from the fact that the two

species usually inhabit different localities. A few cases exist, however, in which there does seem to be the necessary accordance and utility. Mr. Mansel Weale mentions a labiate plant (Ajuga ophrydis), the only species of the genus Ajuga in South Africa, which is strikingly like an orchid of the same country; while a balsam (Impatiens capensis), also a solitary species of the genus in that country, is equally like an orchid, growing in the same locality and visited by the same insects. As both these genera of plants are specialised for insect fertilisation, and both of the plants in question are isolated species of their respective genera, we may suppose that, when they first reached South Africa they were neglected by the insects of the country; but, being both remotely like orchids in form of flower, those varieties that approached nearest to the familiar species of the country were visited by insects and cross-fertilised, and thus a closer resemblance would at length be brought about. Another case of close general resemblance, is that of our common white deadnettle (Lamium album) to the stinging-nettle (Urtica dioica); and Sir John Lubbock thinks that this is a case of true mimicry, the dead-nettle being benefited by being mistaken by grazing animals for the stinging-nettle.1

1 Flowers, Fruits, and Leaves, p. 128 (Fig. 79).

Colours of Fruits.

It is when we come to the essential parts of plants on which their perpetuation and distribution depends, that we find colour largely utilised for a distinct purpose in flowers and fruits. In the former we find attractive colours and guiding marks to secure cross - fertilisation by insects; in the latter attractive or protective coloration, the first to attract birds or other animals when the fruits are intended to be eaten, the second to enable them to escape being eaten when it would be injurious to the species. The colour phenomena of fruits being much the most simple will be considered first.

The perpetuation and therefore the very existence of each species of flowering plant depend upon its seeds being preserved from destruction and more or less effectually dispersed over a considerable area. The dispersal is effected either mechanically or by the agency of animals. Mechanical dispersal is chiefly by means of air-currents, and large numbers of seeds are specially adapted to be so carried, either by being clothed with down or pappus, as in the well-known thistle and dandelion seeds; by having wings or other appendages, as in the sycamore, birch, and many other trees; by being thrown to a considerable distance by the splitting of the seed-vessel, and by many other curious devices. Very large numbers of seeds, however, are so small and light that they can be carried enormous distances by gales of wind, more especially as most of this kind are flattened or curved, so as to expose a large surface in proportion to their weight. Those which are carried by animals have their surfaces, or that of the seedvessel, armed with minute hooks, or some prickly covering which attaches itself to the hair of mammalia or the feathers of birds, as in the burdock, cleavers, and many other species. Others again are sticky, as in Plumbago europæa, mistletoe, and many foreign plants.

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All the seeds or seed-vessels which are adapted to be dispersed in any of these ways are of dull protective tints, so that when they fall on the ground they are almost indistinguishable; besides which, they are usually small, hard, and

1 For a popular sketch of these, see Sir J. Lubbock's Flowers, Fruits, and Leaves, or any general botanical work.

altogether unattractive, never having any soft, juicy pulp; while the edible seeds often bear such a small proportion to the hard, dry envelopes or appendages, that few animals would care to eat them.

The Meaning of Nuts.

There is, however, another class of fruits or seeds, usually termed nuts, in which there is a large amount of edible matter, often very agreeable to the taste, and especially attractive and nourishing to a large number of animals. But when eaten, the seed is destroyed and the existence of the species endangered. It is evident, therefore, that it is by a kind of accident that these nuts are eatable; and that they are not intended to be eaten is shown by the special care nature seems to have taken to conceal or to protect them. We see that all our common nuts are green when on the tree, so as not easily to be distinguished from the leaves; but when ripe they turn brown, so that when they fall on to the ground they are equally indistinguishable among the dead leaves and twigs, or on the brown earth. Then they are almost always protected by hard coverings, as in hazel-nuts, which are concealed by the enlarged leafy involucre, and in the large tropical brazil-nuts and cocoanuts by such a hard and tough case as to be safe from almost every animal. Others have an external bitter rind, as in the walnut ; while in the chestnuts and beechnuts two or three fruits are enclosed in a prickly involucre.

Notwithstanding all these precautions, nuts are largely devoured by mammalia and birds; but as they are chiefly the product of trees or shrubs of considerable longevity, and are generally produced in great profusion, the perpetuation of the species is not endangered. In some cases the devourers of nuts may aid in their dispersal, as they probably now and then swallow the seed whole, or not sufficiently crushed to prevent germination; while squirrels have been observed to bury nuts, many of which are forgotten and afterwards grow in places they could not have otherwise reached.1 Nuts, especially the larger kinds which are SO well protected by their hard, nearly globular cases, have their dispersal facilitated by rolling down hill, and more especially 1 Nature, vol. xv. p. 117.

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