the large brilliantly-marked wings of some butterflies and

moths.

Colours are produced or intensified by processes of development, either where the integument or its appendages undergo great extension or modification, or where there is a surplus of vital energy, as in male animals generally, and more especially at the breeding season.

Colours are also more or less influenced by a variety of causes, such as the nature of the food, the photographic or physiological action of light, and also by some unknown local action, probably dependent on chemical peculiarities in the soil or vegetation.

These various causes have acted and reacted in a variety of ways, and have been modified by conditions dependent on age or on sex, on competition with new forms, or on geographical or climatic changes. In so complex a subject, for which experiment and systematic inquiry have done so little, we cannot expect to explain every individual case, or solve every difficulty; but it is believed that all the great features of animal coloration and many of the details become explicable on the principles we have endeavoured to lay down.

It will perhaps be considered presumptuous to put forth this sketch of the subject of colour in animals as a substitute for one of Mr. Darwin's most highly elaborated theories— that of voluntary or perceptive sexual selection; yet I venture to think that it is more in accordance with the whole of the facts, and with the theory of natural selection itself; and I would ask such of my readers as may be sufficiently interested in the subject, to read again chapters xi. to xvi. of the Descent of Man, and consider the whole subject from the point of view here laid down. The explanation of almost all the ornaments and colours of birds and insects as having been produced by the perceptions and choice of the females, has, I believe, staggered many evolutionists, but has been provisionally accepted because it was the only theory that even attempted to explain the facts. It may perhaps be a relief to some of them, as it has been to myself, to find that the phenomena can be conceived as dependent on the general laws of development, and on the action of "natural selection," which theory will, I venture to think, be relieved from an

V

abnormal excrescence and gain additional vitality by the adoption of the views here imperfectly set forth.1

Although we have arrived at the conclusion that tropical light and heat can in no sense be considered as the cause of colour, there remains to be explained the undoubted fact that all the more intense and gorgeous tints are manifested by the animal life of the tropics; while in some groups, such as butterflies and birds,. there is a marked preponderance of highly-coloured species. This is probably due to a variety of causes, some of which we can indicate, while others remain to be discovered. The luxuriant vegetation of the tropics throughout the entire year affords so much concealment that colour may there be safely developed to a much greater extent than in climates where the trees are bare in winter, during which season the struggle for existence is most severe, and even the slightest disadvantage may prove fatal. Equally important, probably, has been the permanence of favourable conditions in the tropics, allowing certain groups to continue dominant for long periods, and thus to carry out in one unbroken line whatever developments of plumage or colour may once have acquired an ascendency. Changes of climatal conditions, and pre-eminently the glacial epoch, probably led to the extinction of a host of highly-developed and finelycoloured insects and birds in temperate zones, just as we know that it led to the extinction of the larger and more powerful mammalia which formerly characterised the temperate zone in both hemispheres; and this view is supported by the fact that it is amongst those groups only which are now exclusively tropical that all the more extraordinary developments of ornament and colour are found. The obscure local causes of colour to which we have referred will also have acted most efficiently in regions where the climatal condition remained constant, and where migration was unnecessary; while whatever direct effect may be produced by light or heat will necessarily have acted more powerfully within the tropics. And lastly, all these causes have been in action over an actually greater area in tropical than in temperate

1 These views have been restated and enforced by much fresh illustration and argument in Darwinism, chap. x.

V

zones; while, estimated potentially, in proportion to their lifesustaining power, the lands which enjoy a practically tropical climate (extending as they do considerably beyond the geographical tropics) are very much larger than the temperate regions of the earth.

Combining the effects of all these various causes, we are quite able to understand the superiority of the tropical parts of the globe, not only in the abundance and variety of their forms of life, but also as regards the ornamental appendages and vivid coloration which these forms present.

VI

THE COLOURS OF PLANTS AND THE ORIGIN OF THE

COLOUR-SENSE

Source of Colouring Matter in Plants-Protective Coloration and Mimicry in Plants-Attractive Colours of Fruits-Protective Colours of Fruits-Attractive Colours of Flowers-Attractive Odours in Flowers -Attractive Grouping of Flowers-Why Alpine Flowers are so beautiful-Why Allied Species of Flowers differ in size and beauty-Absence of Colour in Wind-fertilised Flowers-The same Theory of Colour applicable to Animals and Plants-Relation of the Colours of Flowers and their Geographical Distribution-Recent Views as to Direct Action of Light on the Colours of Flowers and Fruits-On the Origin of the Colour-sense: Supposed increase of Colour-perception within the Historical Period-Concluding Remarks on the Colour-sense.

Source of Colouring Matter in Plants

THE colouring of plants is neither so varied nor so complex as that of animals, and its explanation accordingly offers fewer difficulties. The colours of foliage are, comparatively, little varied, and can be traced in almost all cases to a special pigment termed chlorophyll, to which is due the general green colour of leaves; but the recent investigations of Mr. Sorby and others have shown that chlorophyll is not a simple green pigment, but that it really consists of at least seven distinct substances, varying in colour from blue to yellow and orange. These differ in their proportions in the chlorophyll of different plants; they have different chemical reactions; they are differently affected by light; and they give distinct spectra. Mr. Sorby further states that scores of different colouring matters are found in the leaves and flowers of plants, to some of which appropriate names have been given, as erythrophyll, which is red; and phaiophyll, which is brown; and many of

VI

these differ greatly from each other in their chemical composition. These inquiries are at present in their infancy, but as the original term chlorophyll seems scarcely applicable under the present aspect of the subject, it would perhaps be better to introduce the analogous word chromophyll a general term for the colouring matters of the vegetable kingdom.

as

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