• THE ENGLISH AND FOREIGN PHILOSOPHICAL LIBRARY. VOLUME XIV. Los Angeles, Cal. THE COLOUE-SENSE: Its (25rtgtn anH aDeUelopment. AN ESSAY IN COMPARATIVE PSYCHOLOGY. BY GKANT ALLEN, B.A. AUTHOR OK "physiological AiSTHKTlCS." BOSTON: HOUGHTON, OSGOOD, & COMPANY. 1879. 4tei 1^0 /ra. ^ " The Senses of the Lower 2 "The Ibis," vol. ii. p. 344, i860. Animals," in the Quarterly Journal ^ Gould, Handbook to the Birds of of Science, July 1878. 124 THE COLOUR-SENSE. noticed in Jamaica the unerring certainty Vvith wliich chickens darted from blossom to blossom of a yellow potentilla, for which they have a particular fancy, and which they always snapped up as though they supposed it to be alive. These instances lead us on to those of the fruits, whose development we examined in our last chapter. " Eed," says Mr. Wallace, " being a very common colour of ripe fruits which attract birds to devour them and thus dis- tribute their seeds, we may be sure that the contrast of red and green is to them very marked."^ But this seems to me a somewhat inadequate expression of the real evi- dence on the point. We have seen that almost all those seeds or fruits which would be injured by the interference of birds are protectively coloured green or brown, while almost all those seeds or fruits which would be aided by the interference of birds are attractively coloured red, pink, orange, yellow, purple, blue, lilac, or black. I think these facts fully justify us in concluding that birds are able to distinguish every one of these colours from green, and most likely from one another. Otherwise there would be no reason why succulent fruits should differ in colour from nuts. The single case of the almond and the plum will bring the question at issue into strong relief. As in the case of entomophilous flowers, so in the case of succu- lent fruits, unless we believe that the seemingly attractive organs were developed for the purpose of enticing animals, we must believe that they are a positive waste of energy to the parent plant. The evolution of bright flowers themselves shows that birds as well as insects are attracted by their beautiful petals. Mr. Darwin has collected many instances in which blossoms are fertilised by birds ; and Fritz MuUer notes several species of Ahutilon in Brazil, which he believes depend entirely on humming-birds for the disper- sion of their pollen.^ Mr. Wallace observes that brilliant 1 Tropical Nature, p. 246. 2 Cross-Fertilisation, p. 371. THE COLOUR-SENSE IN VERTEBRATES. 125 flowers witli handsome corollas exist in many Oceanic islands, such as Juan Fernandez, where flying insects are almost unknown ; but their place is supplied by humming- birds, which Mr. Mosely mentions as being " extraordinarily abundant." ^ Mr. Belt believes that a climbing plant of Central America, 3f«'7-c9'ra via nepentlioides, has been specially adapted to the same birds ; while Mr. Wallace thinks that many Australian and Malayan flowers have been similarly specialised for the visits of honey-eaters, lories, and sun- birds. "Only large flowers," says Mr. J. E. Taylor, "can be visited by these birds, or those whose polypetalous corollas allow of the head being thrust into the centre. Hence we have, in some measure, a reason afforded us for the laro^er size of the flowers in resjions where such birds are abundant. The large bushes and trees of such countries usually bear very fine showy flowers in order to attract the birds ; and it is found that the brush-tongued parra- keets are particularly fond of the flowers which grow at a height above the ground." ^ Any one who has watched a humming-bird darting with lightning speed from blossom to blossom could "hardly have a doubt of his acute colour- perception. The proofs afforded by imitation and mimicry are stronger in the case of birds than of any other class. One may say generally that almost all insects which display protective or imitative colouring do so for the sake of escaping birds or lizards. A few cases must suffice to show the general tendency of the evidence. The leaf and stick insects (Phylliidoe and Phasmidse) closely imitate the colours and shapes of leaves and sticks. One in particular, the Ceroxylus laceratus, is apparently overgrown by moss or jungermaunia.^ Sir Emerson Tennent describes the 1 Tropical Nature, p. 273. ^'^^ the omission would have been a 2 Flowers, their Origin, &c., p. serious defect; sol have here departed 294. This is the only instance in from my usual rule, and taken most which I have availed myself of new of the above cases from Mr. Taylor's matter from this interesting little pages. volume, which appeared while I was ' Wallace, Contributions to the \n course of revising my manuscript ; Theory of Natural Selection, p. 64. 126 THE COLOUR-SENSE. leaf-insects as possessing " all varieties of hue, from the pale yellow of an opening bud to the rich green of the full-blown leaf and the withered tint of decay." ^ The Kallima paraleJcta, a leaf-like butterfly of the Malay Archipelago, always rests among dead or dry leaves, which it resembles in all their varying hues, even appearing to be spotted with small fungi. Canon Tristram has noted that almost every insect, bird, or reptile inhabiting the desert of Sahara is coloured exactly like sand,- and Lord George Campbell mentions a butterfly similarly imitative of its background which frequented the sea-shore at Am- boyna.^ A South American Zeptalis so closely resembles an uneatable Ithomia "in every shade and stripe of colour," that Mr. Bates could hardly distinguish them, even with the aid of his minute entomological knowledge. *' One of the Hemiptera (Spiniger luteicornis)," says Mr. Belt, " had every part coloured like the hornet {Priocnemis) that it resembled. In its vibrating coloured wing-cases it departed greatly from the normal character of the Hemiptera and assumed that of the hornets." * The same careful observer gives many similar instances of mimetic resemblance in the Coleoptera,^ and Lepidoptera.^ But perhaps the most astonishing of these imitative forms is that of a moss-like insect, the larva of a Phasma, which is prolonged into curious green filaments, to mimic the moss in which it lives.'^ Of course these creatures could derive no advantage from their minute reproduction of spots, lines, and hues, unless the enemies against which they required protection were capable of distinguishing their colours. Mimicry or imitative devices of this sort are not confined to insects. Many lizards, such as the geckos, have colours like those of the walls on which they creep ; while the 1 Ceylon, j). 251. authority, lie may be confidently relied 2 See also Gurney, Kambles of a on for any question of fact. Naturalist, p. 56. ^ The Naturalist in Nicaragua, p. ^ Log-letters from the Challenger, 319. p. 208. Although this amusing writer ^ Ibid., p. 317. ^ P. 382. can hardly be considered a scientific ' Figured in Belt, ubi supra, p. 382. THE COLOUR-SENSE IN VERTEBRATES. 127 protective green hue of the tree-frogs has already been noticed. Even birds occasionally mimic one another in the same manner. For example, two species of Mimeta (a sort of oriole), in Bouru and Ceram, imitate two Tropi- dorhynclii (honeysuckers) in minute details of colour, thus escaping small birds of prey, as the Tropidorhynchi are strong and pugnacious creatures.-^ For other cases the reader must be referred to Mr. "Wallace's admirable essay on " Mimicry and other Protective Eesemblances among Animals." These various proofs, though indirect, can leave us in little doubt with regard to the general existence of a colour-sense among birds. "When we come to the highest class of vertebrates — the mammalia — strangely enough the evidence of a colour- sense almost entirely fails us. The antipathy of male ruminants for scarlet, and the curiosity which certain monkeys display with regard to bright-coloured objects, are the only facts in point which come under ordinary observation. This result, so contrary to what we might have expected, appears really quite natural when we exa- mine more closely the circumstances of the case. By far the larger part of the mammalia are either herbivorous like the ruminants and pachyderms, or carnivorous like the technical carnivores, insectivores, and whales. Only a small portion of the class subsists upon fruits, w^hile none of them are very specially connected with flowers. Hence a large set of possible tests which we can employ in the case of insects and birds are wholly inapplicable to mammals. Moreover, the want of close relations with the coloured parts of plants has probably resulted in a want of any peculiar love for bright colour, such as we see reason to suspect in the butterflies, humming-birds, and parrots. This absence of a taste for brilliancy is probably marked by the absence of brilliant hues in the animals themselves, the result of sexual selection ; for these hues, •■ Wallace, Malay Archipelago, p. 401. 128 THE COLOUR-SENSE. as we shall see hereafter, only appear among the mam- malia in a few higher arboreal and frugivorous species, such as the mandrill and certain squirrels. For the most part, throughout the mammalian series sexual selection seems only to have exerted itself, if at all, in the produc- tion of elegant shapes, protuberances like horns and dew- laps, and marked contrasts of light and shade, as in the zebra, giraffe, and hysena. Nevertheless we can hardly doubt that mammals do possess a considerable colour-sense, though, owing to the circumstances of their practical environment, their taste for any special hue is probably far from strong. Here once more I must remind the reader that the proofs of a colour-sense throughout the whole infra-human world are necessarily very derivative, and that they owe their chief strength to their cumulative character. The fragmentary evidence collected in this chapter will be much corroborated and supplemented by that which will be detailed in the sequel. Enough will have been done if we succeed in showing that the hypothesis of a general colour-sense is consonant with all the facts of nature, and helps us to understand those facts in a way which no other hypothesis can do. ' For the present it will be sufficient if we bear in mind the one great point hitherto settled — that wherever any part of a plant, be it flower or fruit, will derive any benefit from attracting the eye of an animal, be it insect, bird, or mammal, that part is almost invariably coloured with some pure and brilliant hue, be it blue, red, yellow, pink, orange, violet, or lilac, quite distinctive from the green of ordinary vegetation. This one fact is the great pivot upon which turns our whole knowledge of the animal colour-sense. ( 129 ) CHAPTER VIII. THE COMMUNITY OF TASTE BETWEEN FLOWER- FEEDING AND FRUIT-EATING SPECIES. Before we proceed to consider the secondary reactions of the colour- sense in insects and vertebrates upon their own external appearance, we must glance for a moment at one of the determining causes which give approximate unifor- mity to the general results of such reactions in the animals with which we are most specially concerned. In the next chapter we shall have to examine the production of bright hues in the wings of butterflies, the skins of lizards, the feathers of birds, and the fur of mammals, due to the selective action of sexual preferences. But, as a necessary preliminary to that inquiry, we must first set ourselves to determine the principles which govern the formation of tastes generally among the flower-feeding and fruit-eating animals. Before we can trace to its final effects the action of a sexual preference for bright colouring, we must pre- viously find out with certainty the reasons why a taste for such colouring should exist at all in the animal con- sciousness. People are generally too apt to accept as ultimate and obvious every fact with which they have always been familiar. Seeing that bright colours as a rule attract chil- dren and savages, dogs, birds, fish, and insects, they do not trouble themselves to seek a reason for this preference, but take it for granted as an inherent and natural pro- perty of the animal organism, or, more often and more absurdly still, of the colours themselves. If, however, we I I30 THE COLOUR-SENSE. reflect upon the subject for a moment, we shall see that there is no primitive and self-sufficing reason in the nature of things why anj^ one colour should be more beautiful to us than another. Dull and dingy hues might conceivably have been just as pleasant to our sense, under slightly different conditions of our development, as we know bright and pungent hues to be, under the actual circumstances of humanity. We must get a little deeper into the ground- work of our likes and dislikes if we would really under- stand the origin of our native preference for brilliant tints over mixed or unstimulating colours. Now, after this preamble, most readers will imagine that I mean to explain the liking of flower-feeders and fruit- eaters for bright hues by means of that grand but somewhat vaguely employed shibboleth, the Association Theory. I know that to the mass of loose thinkers associa- tion is a sort of psychological dms ex 7nachinawh.ic]i satisfac- torily accounts for every ill-defined mental problem, just as electricity is a sort of physical deus ex machina which simi- larly gets over every ill-comprehended material problem. Such persons say to themselves at once, " Oh, of course, birds and butterflies feed off bright-coloured objects ; so bridit colours f^et associations with their food, and are consequently pleasant to them." Having thus satisfied their nascent critical doubts by the easy application of an accepted formula, they never pause to translate their vague speculation into thinkable terms, but leave it as they took it up, a mere algebraical expression, incapable of rational statement in a concrete form. For how can association with food make a colour or anything else pleasant in itself ? This is the true crux of the Association Theory, a crux which, as I humbly believe, few of its adherents have ever perceived in its full significance. Until it has been solved, the theory remains a mere verbal explanation, adding no- thing to our real knowledge of the subject, yet deluding us by its specious resemblance to an explanatory truth. The mode of exposition here adopted will be a very COMMUNITY OF TASTE. 131 different one, based upon the known psyclio-pliysical law of pleasure and pain. According to that law, pleasure is the psychical aspect of an ultimate physiological fact, which in its physical aspect may be summed up as the unimpeded activity of a fully nurtured and not over- worked nervous structure in unbroken connection with the cerebro-spinal or other central sentient system. Con- versely, or nearly so, pain is the psychical aspect of an ultimate physiological fact, which in its physical aspect may be summed up as the disintegration, insufficient nutri- tion, or excessive activity of a nervous structure, similarly connected with the sentient organism.^ With the latter half of this important law we have here little or nothing to do ; but the former half so intimately concerns our subject that I shall make no apology for endeavouring briefly to explain its meaning in simpler language than that of the above abstract formula. A pleasure, then, is the feeling which results when any sentient nervous centre receives a stimulation not exces- sive in quantity, nor beyond the existing power of the structures concerned. Every centre undergoes at each stimulation a certain amount of disintegration ; and if that disintegration pass beyond the easy repairing-point of the system, pain sets in. But, on the other hand, so long as the stimulation is moderate, by exercising the structures it promotes their general efficiency, and hence it is accom- panied by a feeling of pleasure. Or to translate our law into still more concrete and ordinary language, we may say that whenever an organ which can feel at all is exer- cised not beyond the due amount, pleasure is the result. Hence the pleasurableness of any activity may be ac- cepted as a rough gauge of its general desirability for the organism as a whole, while conversely its painfulness 1 For a full explanation of the form wards), see my "Physiological ^Es- here given to this law (originally due thetics," chap, ii., "Pleasure and to Mr. Herbert Spencer and Professor Pain," where the grounds upon which Bain, working on the basis of previ- the conclusion is based have been de- OU3 inquirers, from Aristotle down- tailed at length. 132 THE COLOUR-SENSE. may be regarded as a certain proof of its general unde- sirability. ]^ow the more fully-nurtured an organ mayj be, tlie higher is its functional efficiency, and the greater the plea- sure to be derived from its exercise. We all know that the fresher our limbs, our muscles, our nerves, and our eyes, the greater the enjoyment we derive from a country walk or a game of cricket. After long fasting we eat our food with greater relish ; after long confinement we use our less and arms with redoubled delio-ht. But we also know that in order to keep up a state of high efficiency in any organ, frequent exercise is necessary. Only by running, jumping, rowing, and gymnastics can we bring our muscles into a proper condition for hard athletic work. Only by constant practice can we retain any accomplishment which we have learnt by dint of serious effi)rt. And just the same is true of nerves. Their existing structure has been acquired by continuous function in past generations, and continuous function is necessary still if we would prevent them from rusting into obsolescence. Accordingly, whenever we find that any activity is pro- ductive of immediate pleasure in ourselves, we may be sure that the activity in question is one which has long been practised by our human or ante-human ancestors. The greater the pleasure, as I have elsewhere endeavoured to show, the greater the intimacy of connection between the activity and the life of the species. Let us, for example, take the case of colour itself, with which we are here so fully engaged. If in any species the need for distinguish- ing different colours ever arose, and if by its side there also arose a nascent structure for so distinguishing them, then those individuals in which that structure was most fully developed would survive from generation to generation, in virtue of their superior adaptation to the needs of their environment above their less highly endowed compeers. But with each such increment in the structure there would go increased pleasure in the function. Conversely, the COMMUNITY OF TASTE. 133 more fully the function was indulged the more would the structure increase and strengthen by exercise. So from generation to generation, as the power of distinguishing colours became more and more developed, the pleasure arising from their perception would grow more and more acute. Such pleasure forms the first groundwork for that differential preference in individuals or species which we Know as taste. But every colour would not probably prove equally pleasurable. Some, like the ordinary greys, greens, and browns, occur too often in the surrounding world to allow of any marked gratification, derivable only from the in- termittent stimulation of little-worked nerves. Moreover, these common colours would have no special reference to the life of the race, and so would have few structural con- nections with other portions of the central nervous system. But in the case of fruit-eating and flower-feeding species, we may well suppose that the special nerves devoted to the perception of red, yellow, orange, and purple would naturally be much strengthened by constant hereditary use ; while the comparatively intermittent nature of the stimulation would render the accompanying feeling far more pleasurable than in the more familiar instances of green or brown. Furthermore, the close relation of these colours with the food of the species would doubtless give rise to numerous nervous connections in the central sys- tem, whereby the sight of such coloured objects might set up the necessary movements for obtaining the booty. In this manner the central organs of special colour-percep- tion for the brilliant hues of fruits and flowers would in all probability assume unusually large dimensions, and would certainly possess large numbers of concurrent fibres along which waves of discharge might readily travel, thus giving free vent for a considerable volume of pleasure- yielding energy. Such species might fairly be said to possess a taste for red, yellow, and other like pungent hues ; and we might accordingly give a rough definition 134 THE COLOUR-SENSE. of taste as a special preference, in an individual or a race, for one or more out of several similar objective stimuli, depending ultimately upon special variations of nervous development. Of course this hasty definition leaves out of considera- tion the other half of the subject, which we might perhaps sum up as distaste, — the special repugnance to one or more among such stimuli, ultimately due to like diversi- ties of individual or generic organisation. In this case, however, we must distinguish between two widely dif- ferent forms of feeling, which are apt at first to be men- tally confused, — mere neutral indifference, which results from a stimulation too languid or too common to produce pleasure, and positive dislike or disgust, which arises from some actually painful and disintegrative action. Thus, in sight, an ugly colour, like that of mud, is simply neutral and unstimulating ; but in taste, a bitter or acrid substance probably sets up material disintegration, and so of course produces a positively painful sensation. This distinction will become more obvious and more important as we proceed. In this way only, then, as it seems to me, can associa- tion have anything to do with the intrinsic pleasurable- ness of any sensation, namely, by affording outlets for the overflowing nervous energy. But the main pleasure of the sensation itself, as I understand the question, must be due to inherited cahbre of the nerves and nervous centres employed, that calibre being due itself to ances- tral function throughout many previous generations. To put once more the concrete case, fruit-eaters and flower- feeders derive pleasure from brilliant colours (postulating the fact for the time being, argumenti gratia), not because those colours have mental associations with their food, but because the structures which perceive them have been continually exercised and strengthened by hereditary use. The connection with food has given numerous outlets for the nervous energy, and has been the ultimate cause for COMMUNITY OF TASTE. 135 the extra development of colour-perceiving structures, but it has had no direct effect, as I believe, upon the immediate pleasure of sensation. It is true that in highly evolved animals, with whom the emotions have attained an immense and preponderant influence, associations do largely enter into all pleasur- able feelings. But even here the ultimate explanation is equally simple and straightforward. These emotions have their own proper nervous seats, as Terrier's experi- ments sufficiently show ; and we must suppose that the actual sensation, being located in a centre which has con- nections with the seats of such emotions, rouses action in the emotional centres, more or less conspicuously, and so adds a more or less distinct factor to the total resulting consciousness. But it would be very foolish to transfer similar ideas into the simple nervous organisation of birds, and still more into that of bees and butterflies. Taste in these animals must be almost entirely a matter of direct sensation, dependent upon the calibre of the nerves employed, and little influenced by the few possible associated feelings.^ Having premised these few considerations as to the nature of taste in general, let us now go on to examine the special tastes of fruit-eating and flower-feeding ani- mals. We shall find on investigation that these appear to be approximately identical throughout the whole animal series ; while they are more or less strongly marked off' from the opposite tastes of carnivores and carrion-eaters. The very same sweet and sugary substances, the very same etherial and delicate perfumes, the very same bright 1 Lest any reader should imagine that I begin to dissent ; and even that I mean by these remarks to cast here my dissent, as will be seen some doubt upon the whole body of from the above paragraph, extends Associationist Psychology, I hasten only to the very simplest elements of to add, parenthetically, that I fully feeling. I do not for a moment accept that system so long as it con- doubt that the a3sthetic pleasures of fines itself to the relation between cultivated Aryan man depend largely, the senses aud the intellect. It is or even mainly, upon associated emo- only when it is brought forward as a tion. verbal explanation of emotional facts 136 THE COLOUR-SENSE. and dainty colours, seem pleasing in the very same way to butterflies, and humming-birds, and parrots, and apes, and men. The similarity of nervous impressibility which we thus perceive to hold throughout the whole heterogene- ous collection of fruit and flower haunters casts much light upon the nature of sexual selection, and upon the identity of taste between man and so many lower animals. It enables us to see why the flowers which the bee de- veloped for his own delight and guidance should be the joy of children and the envy of artists ; why the hues of the orange and the mango should be as beautiful to man as to the toucans and macaws which gave them origin ; why the wing of the butterfly, the tail of the peacock, and the burnished throat of the sun-bird should be exqui- site to our eyes as they were to those of their fastidious mates ; and why human beings should dye their bodies with the woad of Britain and the ochre of Papua, or tinge their garments with the purple of Tyre and the thou- sand hues of Lyon, to vie with the gorgeous tints of bird and insect in the very self-same profusion of refulgent colours. First, then, let us begin with the sense of tasU. It is a most noteworthy fact that wherever any part of a plant can gain any advantage by attracting the notice of animals, it always effects its purpose by the secretion of sugar, or, as we oftener though more incorrectly call it in this connection, honey. Now sugar, as I have already pointed out, has a special power of acting upon the gustatory nerves of animals, through the great solubility, diffusibility, and crystalline texture of its particles. Ac- cordingly, we find that almost all classes of fruit-eaters and flower-feeders show a decided partiality for this pleasant stimulant — a partiality due, doubtless, to the long habits of their ancestors, which have developed cor- respondingly differentiated structures for the perception of the particular body in question. In flowers, sugar is secreted to attract bees and other insects ; while in fruits, COMMUNITY OF TASTE. 137 it acts as an allurement to birds and mammals. Further- more, certain plants possess organs for the secretion of sugar on their stems or at the base of their leaf-stalks, of which Sir John Lubbock gives the following account : — " Belt and Delpino have, I think, suggested the true func- tion of these extra-floral nectaries. The former of these excellent observers describes a South American species of Acacia, which, if unprotected, is apt to be stripped of its leaves by a leaf-cutting ant, which uses the leaves not directly for food, but, according to Mr. Belt, to grow mushrooms on. The Acacia, however, bears hollow thorns, and each leaflet produces honey in a centre-formed gland at the base, and a small sweet pear-shaped body at the tip. In consequence, it is inhabited by myriads of a small ant which nests in the hollow thorns, and thus finds meat, drink, and lodging all provided for it. These ants are continually roaming over the plant, and constitute a most efficient bodyguard, not only driving off the leaf- cutting ants, but even, in Mr. Belt's opinion, rendering the leaves less liable to be eaten by herbivorous mammalia." ^ Indeed, so universal is the taste for suo'ar among insects, that certain small animal creatures, like the Aphides and Cocci, have themselves acquired the habit of developing nectaries, and so gaining the protection of ants, which may be seen " assiduously running up the stems of plants to milk these curious little cattle." And if we want further proof of the general love for sweet food- stuffs, we need only bethink us how the insects flock about a barrel of treacle in our streets, how the birds cono-re^-ate in fruit-gardens, and how our own children gather around the windows of the confectioner's shop to stare at the tempting wares within — rendered all the more enticing, be it observed, through the very same addition of red, blue, and yellow, which had already been invented by the fruit and the flower. 1 " On Certain Eelations between Plants and Insects," a lecture delivered at Glasgow, January 24, 1878, p. 6. 138 THE COLOUR-SENSE. Equally significant are the changes in habit or mode of life between fruit-eating and flower-feeding classes. Thus, a large number of hymenopterous insects live upon honey extracted from flowers ; but the omnivorous wasps, as we all know, have taken to surreptitious feasting upon the sugary juices of peaches, pears, and nectarines. In like manner, I have often noted lepidopterous species, whose natural food consists of the nectar in summer blossoms, feeding greedily upon fallen fruit. Mr. W. M. Gabb captured the lovely Morplios of Nicaragua by baiting with a piece of over-ripe banana ; ^ and the Eev. J. G. Wood, a most trustworthy recorder in all that concerns the habits of animals, notices on one occasion having seen whole hordes of the Eed Admiral butterfly ( Vanessa atalanta) darkening the ground wdiere a number of egg-plums lay beneath their parent tree.^ So, too, amongst birds ; while most of them take their sugary food in the form of fruit or seeds, a few kinds, like humming-birds and sun-birds, live largely off the nectar of flowers, mixed with the insects which frequent them.^ Mr. Webber, an American natura- list, tried the experiment of taming the pretty little ruby-throats, which he fed on syrup alone, and though he found that they could not thrive without a fair pro- portion of insects as well, he also discovered that they showed a decided partiality for the taste of sugar. " Some which had been thus tamed and set free returned the following year, and at once flew straight to the re- membered little cup of sweets." ^ In certain instances, we find the interchange of habit taking place within the limits of a single tribe. Thus, the true parrots live al- most entirely off sweet fruits, but their congeners the lories are nectar-eaters. These facts, once more, we may cor- relate with well-known human habits ; as when we see children, whose taste for sweets is derived from frugi- ^ " Nature," February 7, 1878. see Wallace, " Tropical Nature," p. 2 "Insects at Home," p. 401. 235. 2 Ou the food of humming-birds, ■^ Ibid., p. 137. COMMUNITY OF TASTE. 139 vorous ancestors, sucking the juices of honeysuckle and clover, or stealing the honey-bag from our domesticated hive-bees. Indeed, we could have no more simificant symbol of the community of nature here pointed out than the fact that we keep these same bees to gather honey for us from the nectaries of flowers. Conversely, whatever parts of a plant would be injured by the interference of animals, secrete a bitter or acrid juice, which acts deleteriously upon the nerves of taste. Thus, as I have already pointed out, the pericarp, which in fruits-proper is provided with sugary secretions, in nuts is commonly stored with a nauseous principle as a deterrent to animal foes. Again, those fruits which have a sweet pulp generally guard against the loss of their actual seeds by filling them with a bitter substance, of which prussic acid often forms a leading constituent : cases occur in the peach-stone, the apple-pip, and the seeds of oranges or mangoes. That animals as a rule dis- like bitter substances is a matter of common observation ; and experiments which I have conducted on a small number of insects and birds have always resulted in marks either of indifference or of positive distaste. Thus we see that both in their likes and dislikes a oreat com- munity of taste runs through all the flower-feeding and fruit-eating animals. If, now, we turn to the carnivores and carrion-feeders, we shall find a totally different set of sympathies and antipathies. It is true that many dogs and flesh-eating flies love sugar ; but they also love numerous other bodies which several of the former class of creatures would never touch. Fresh meat, or still worse, putrid flesh, does not appeal at all to the senses of bees and parrots. Man, of course, forms an intermediate link, a fruGjivorous animal who has partially adopted carnivorous tastes. Hence we have a certain liking for the flavour of roast beef and turkey; some of us eat high game and caviare; and savages even prefer meat in an advanced stage of decom- I40 THE COLOUR-SENSE. position. But these are mere surface-tastes, while the deeper-seated ancestral habits come out strongly in our children and our unsophisticated adults. The liking for strong-tasted meats and half -putrid preparations has to be slowly acquired ; whereas the love for sugar, for honey, for fruits, for all sweet things, is born with us into the world, and taken in with our first draught of mother's milk. And in this connection it is worth while to note that the natural food of the human infant contains 62.3 parts of sugar in 1000, while that of a herbivorous calf possesses only 45.6,^ so that it becomes necessary in giving cow's milk to babies to sweeten it considerably up to the proper point. Secondly, let us look at the sense of smell. Here again we notice that wherever any part of a plant wishes to attract animals, it adds to its sweetness the extra allure- ment of perfume ; and the same perfumes are, as a rule, pleasant to all flow^er-feeders and fruit-eaters alike. The delicate odour of a peach, a pine-apple, or a strawberry scarcely differs in kind from that of a lily, a hyacinth, or a violet. Mankind, whose tastes in this matter are de- rived from the tropical fruits, have equal pleasure with bees or butterflies in the dainty scent of clover and meadow-sweet. Only, as might naturally be expected, the perfumes of fruits (which we have already seen reason to believe are comparatively modern structures) are not so highly developed as the perfumes of flowers ; whence arises the seeming anomaly that our olfactory nerves are more pleasurably stimulated by the stephanotis or the jasmine, which is relatively remote from our practical life, than by the apple or the pear, which is relatively essential. Of course, the explanation here is that the more powerful stimulant naturally affords the greater volume of pleasure, irrespective of its ulterior useful- ness. In this case, too, we see the essential agreement 1 Hermann, Human Physiology, English translation, p. 121. COMMUNITY OF TASTE. 141 between the higher and the lower forms of vegetal- feeders. For just as our taste for sweets corresponds to the insect's taste for honey, so our love for the perfume of flowers is absolutely identical with the pleasure which draws the butterfly towards the luscious blossoms in our English meadows. And it is w^orth while to observe that most of the sweet-smelling flowers appear to be quite late developments of vegetal life; a fact which harmonises well with the correspondingly late development of the bees and other highly-adapted honey-suckers. There is no tribe of plants, for example, more noticeable for their perfume than the family of Labiates, which includes the various species of mint, thyme, balm, sage, marjoram, lavender, rosemary, horehound, calamint, patchouli, hyssop, and basil. The flowers of these plants are almost all very peculiarly shaped and highly scented, and their attractiveness for bees has become proverbial — the honey of poetry is commonly " redolent of thyme." Now the Labiates, so far as known, are tertiary plants of rather late date, which did not make their appearance on the earth until bees and other specialised honey-seekers had reached a high point of evolution. Nor should we omit to notice the fact that many of these plants are now cul- tivated by man for the sake of this very property ; lavender to dry and use for scent, patchouli to extract an essence for the handkerchief, and mint, thyme, or sage to flavour various preparations for the table. The exactly similar cases of nutmeg, cloves, and other spices, whose perfume is famous for its diffusibility, while the mode of their dispersion by nutmeg-pigeons has become classical in the pages of Darwin and Wallace, do not need further comment. In some few instances the pleasure of perfume has been turned into a sexual allurement, as with certain butter- flies, where the two sexes exhibit a different arrangement in the nervures of the wings. "In all cases which I know," says Fritz MuUer, " this difference in neuration is 142 THE COLOUR-SENSE. connected with, and probably caused by, the development in the males of spots of peculiarly formed scales, pencils, or other contrivances, which exhale odours agreeable no doubt to their females. This is the case in the genera Mechanitis, Dircenna, in some species of Thecla, &c." ^ Similar instances occur in the musk-deer and other mam- mals, whose perfumes are used by human beings as pleasurable stimulants. Indeed, I do not think it would be too much to say that almost every substance which we employ as a native scent is derived either from a vegetal product whose natural function is the attraction of ani- mals, or from an animal product whose natural function is the attraction of the opposite sex. On the other hand, we find amongst the carnivores and carrion-feeders a totally different form of olfactory plea- sure. Dogs, wolves, and other predatory mammals, track their prey by scent, while the smell of raw meat renders the larger cats wild with excitement. Vultures and sopi- lotes revel in the hideous smell of putrid animal matter, and flies collect around dung or decaying meat. Curi- ously enough, too, some plants have availed themselves of this special taste, and have laid themselves out, as already noticed, to deceive carrion insects by their like- ness in appearance and smell to putrescent flesh. The Sumatran Rafflcsia and the South African Hydnora have large and lurid blossoms, which thus cunningly induce flies to visit them for the purpose of laying their eggs, and are accordingly fertilised by means of an organised de- ception. To naturally frugivorous man, the scent is, of course, simply disgusting. Yet it is worth notice that many savages, who have acquired for generations the habit of eating half-decomposed meat, positively enjoy those odours which are most distasteful to the nostrils of civilised huma- nity. As Kolben quaintly phrases it, in his old-fashioned style, "Wliat you take for a stink, a Hottentot, if you ^ In "Nature," November 29, 1877. COMMUNITY OF TASTE. 143 will believe him, receives as the most agreeable per- fume." 1 Hence one may see how futile is the argument of Geii:^er, who remarks, as illustratinGr the sensuous in- feriority of the lower animals, that it would be -useless to offer a dog a bouquet to sniff at.^ Of course the dog has no reason for being pleased with a perfume which has no special relation to himself or his ancestors in any way ; but if we offer him a piece of meat, or set him to hunt down game, we shall find how keenly he has been pro- vided by nature with senses to aid him in his ow^n mode of life. In fact, we can only expect pleasure to be felt where ancestral habit has produced a corresponding sensory system. The transference of feeling whereby we are enabled to enjoy the perfume of flowers does not con- tradict this general principle, for it is really analogous to the transference whereby the humming-bird sipped the syrup which resembled the native nectar, or whereby we ourselves enjoy sweetmeats and cakes through our here- ditary liking for fruits and berries. Honey is a more concentrated form of sugar than that which we get in strawberries or oranges, and frangipanni is a more con- centrated form of perfume than that which we get in peaches and pine-apples ; but they probably act in just the same way, though to a greater extent, upon the nerves involved as do the original stimulants, and consequently they need no special explanation. Very different, how- ever, would be the case if a dog or any other animal were to feel pleasure in a stimulation derived from some object which had no kind of relation to his ancestral habits. Such an instance, one might venture to say, would be wholly inexplicable, and opposed to all the known prin- ciples of scientific psychology. The sense of hearing, though interesting in itself through 1 Kolben, "Cape of Good Hope," 2 ''Zur Entwickelungsgeschichte vol. i. p. 231. der Menschheit, iii. p. 50. 144 THE COLOUR-SENSE. its connection with song-birds and the sexual allurements of sound amongst sundry insects, has so little relation with our present subject, that I must reluctantly pass it over here. Lastly, then, we arrive at the sense of siglit, which we must only examine with special reference to the taste for colour. And here, as before, we note at once that those portions of plants which lay themselves out to attract animals are almost without exception conspicuous for their bright colours. Entomophilous flowers, as we have already abundantly observed, have a monopoly of brilliant corollas, while fruits - proper differ from nuts in the startling vividness of their hues. These facts would go by themselves to prove that flower-feeding and fruit-eating animals find an allurement in colour. It is certainly a noticeable fact that just as the sweetness of fruits answers to the sweetness of honey, and just as the scent of fruits answers to the scent of flowers, so do the colours of fruits answer exactly to the colours of flowers. It would seem as though, in every case, nature found a single mode of modifying the nervous substance was amply sufficient (because simplest and easiest) alike for insect and reptile, for bird and ape and human being. Some special facts help to point in the same direction. Thus we find within the limits of a single family, the Eosacese, a large number of fruits -proper, the plum, the apple, the hip, the haw, the strawberry, the raspberry, and the bramble, in which the pericarp or other succulent portion, besides being sweet and scented, is more or less brilliantly coloured ; and again, we also find an aberrant member, the almond, whose seed is enclosed in a nut, and whose pericarp accordingly is hard, dry, and green or brownish, after the usual fashion of nuts. Or once more, we know that in most Oceanic islands there are few flying insects, and that most of the flowers are destitute of bright corollas ; but Mr. Wallace has pointed out that in a few, cases, where honey-sucking birds frequent such islands COMMUNITY OF TASTE. 145 the flowers are extremely large and handsome.^ This fact clearly shows that the birds in question find colour quite as attractive as do bees or butterflies. I do not propose to enter very fully into this question until w^e have seen w^hat light may be cast upon it by the examples collected and the inferences drawn in the suc- ceeding chapter. But I should like to point out here that if our general theory of pleasure be well-founded, it must necessarily result that flower-feeding and fruit- eating animals should derive agreeable sensations from coloured objects. So soon as the eyes of insects or birds have become sufficiently differentiated to discriminate the pinkish or ruddy flower-cases and fruit-vessels from the green leaves around them, and to employ their nascent sense in the quest for food, so soon must the special nerves exercised and strengthened in this process receive some faint pleasure from their due stimulation. And the more developed the nerves become, the more intense must be the resulting enjoyment, till at last an ever-in- creasing gratification would grow up side by side with the growth of entomophilous blossoms and coloured fruit, be- coming stronger and stronger day by day as the structures increased by practice in calibre and power. Two short passages from the works of two leading evolutionists will serve to bring out in strong relief the position here assumed. Mr. A. E. Wallace thus sums up his view with regard to the nature of colour-perception in the lower animals : — " The fact that the higher verte- brates, and even some insects, distinguish what are to us diversities of colour, by no means proves that their sensations of colour bear any resemblance whatever to ours. An insect's capacity to distinguish red from blue or yellow may be (and probably is) due to perceptions of a totally distinct nature, and quite unaccompanied by any of that sense of enjoyment or even of radical distinctness which pure colours excite in us. Mammalia and birds, 1 Tropical Nature, p. 238. 146 THE COLOUR-SENSE. whose structure and emotions are so similar to our own, do probably receive somewhat similar impressions of colour ; but we have no evidence to show that they ex- perience pleasurable emotions from colour itself, when not associated with the satisfaction of their wants or the gratification of their passions." ^ From the whole of this passage, with all due deference to ]\Ir. AVallace (for most of whose work I entertain the deepest respect), I must take leave to differ toto codo. Each of its three sentences appears to me to contain a fallacious position. For the first, the burden of proof lies distinctly with Mr. Wallace, not with his opponents; because, where the external stimulus is the same, and where a general continuity of structure exists, we are not justified in assuming a dif- ference of sensation without some special reason; nor do I believe that so clear a thinker as the author of the *' Malay Archipelago " would have assumed such a differ- ence, were it not for that predisposition to find some eff'ective distinction between man and the lower animals which has so often led him into questionable conclusions. For the second, it seems to me that, since the insect's need for discriminating colour is far greater than our own, analogy would lead us to suppose that his enjoyment would be even deeper, and his sense of distinctness more marked, than in the human subject. For the third, the single instance of the oft-quoted bower-birds, who collect coloured objects to decorate their meeting places, shows that some vertebrates at least possess a liking for brilliant hues in themselves, of a truly aesthetic sort : and the be- haviour of monkeys with regard to flowers and birds, or to red shawls and other strikingly-dyed articles, would seem to point in the same direction. For the rest, Mr. Darwin has gathered together a few isolated instances of disinterested love for colour in a well-known section of his " Descent of Man." ^ It is true that the evidence on this head is still far from satisfactory; but it must be ^ Tropical Nature, p. 243. 2 Yo\. ii. p. no. COMMUNITY OF TASTE. 147 remembered tliat without the assistance of language de- finite information as to tastes cannot be procured except with great difficulty, and that human infants only dis- play the love for colour in the same simple ways as mon- keys or bower-birds. The second passage to which I would refer is one from our great naturahst himself. " How the sense of beauty in its simplest form," says Mr. Darwin, — " that is, the recep- tion of a peculiar kind of pleasure from certain colours, forms, and sounds, — was first developed in the mind of man and of the lower animals, is a very obscure subject. The same sort of difficulty is presented if we inquire how it is that certain flavours and odours give pleasure, and others displeasure. Habit in all these cases appears to have come to a certain extent into play ; but there must be some fundamental cause in the constitution of the ner- vous system in each species."^ Now this fundamental cause I believe to reside in the general law that pleasure accompanies normal stimulation when not excessive in amount; while the influence of ancestral habit, joined with natural selection, has so modified the nervous system in each case that it finds itself normally stimulated by those external agents whicli conduce to the general wel- fare of the organism, and excessively or destructively stimulated by those which conduce to its general detri- ment.2 Accordingly, I infer that in all fruit-eating and flower-feeding species, a taste for sweet flavours, delicate perfumes, and bright colours will have been slowly deve- loped by the hereditary mode of life ; and that the taste so developed will have opportunities for exerting itself in the sexual selection of bright-coloured mates. If this be so, then it must follow that flower-feeding and fruit-eating insects or vertebrates will be specially distinguished from other animals by the exceptional bril- liancy of their colouration. In the next chapter we will 1 Origin of species, 6th edit., p. 162. tics," pas«u7i, where this main prin- 2 See my "Physiological Ji^sthe- cii)le is worked out in detail. 148 THE COLOUR-SENSE, submit our general conclusion to the test tliTis suggested to us, and if we find that bright hues are, as a matter of fact, unusually common amongst those species in which we have inferred a 'priori that a taste for brilliancy would have been evolved by the circumstances of their life, then we shall have added another item to our cumulative proof of the existence and influence of a colour-sense among the lower animals. Just as we saw that the taste for sweets, formed upon flowers and fruits, could be trans- ferred to syrup, sugar, honey, bonbons, cakes, and pud- dings ; just as we saw that the human liking for dainty perfumes, formed upon strawberries and oranges, could be transferred to hyacinths and heliotropes : so, I believe, the love for colour, formed upon the natural food of the various species, can be transferred to the choice of beauti- ful mates, and, strengthened by this transference, can be handed down by heredity to mankind till it results at last in the disinterested delight in the sunset and the autumn hues, in the flowers of our gardens, the varying tints of our landscapes, and the exquisite harmony of our Guides and our Eossettis. Let us see, then, how far the facts of nature will bear out the theory on this subject which we have framed from the analogy of our other senses. ( 149 ) CHAPTER IX. THE DIRECT REACTION OF THE COLOUR-SENSE UPON THE ANIMAL INTEGUMENTS. If any unscientific person were asked to name tlie two most beautiful classes of animals in the world, he would unhe- sitatingly answer, '' Butterflies and humming-birds." It is a significant fact that these are perhaps the most exclu- sively flower-haunting of all invertebrate or vertebrate creatures respectively. And if he were asked to name any other birds, besides the single family above mentioned, which are specially conspicuous for their brilliant colour- ation, he would probably reply, '' Parrots and their allies." It is an equally significant fact that these birds are fruit- eaters. Following up the hint thus given us, we may run through the chief instances of brilliant species in both great divisions of articulates and vertebrates, in order to discover whether there is any constancy of connection between the nature of the food and the colouration. As before, we may narrow down our consideration of the articulates to the great group of insects, because we know too little about the habits of their marine congeners to argue with any certainty as to their traits ; while the other land-articulates are relatively unimportant for our present purpose. Now amongst the insects, the most brilliant order are the Lepidoptera, including both the butterflies and moths, which, it need hardly be said, feed upon flowers. Of course, it may be readily objected that the amount of food eaten by the perfect winged insects is relatively small, and that the caterpillars live for the most part upon the I50 THE COLOUR-SENSE. green portions of plants. Indeed, some butterflies possess no mouths at all, and pass the whole of their short lives by the expenditure of energy laid up in the larval condi- tion. Yet this objection does not really invalidate the general conclusion ; for the eyes of the perfect insects have been evidently adapted to the colours of flowers, and the main object of their winged state is the perpetuation of the species ; so that we can easily understand how the tastes ancestrally formed in their last stage should domi- nate the selection of their mates. Hence we find that the colours of caterpillars are mostly protective, being due to natural -selection alone, while those of butterflies are mostly attractive, being largely due to sexual selection. Furthermore, if we examine the Lepidoptera in detail, we shall find similar conclusions thrust upon us. They may be divided into two great sections, — the moths and the butterflies, — of which the former are mainly nocturnal or crepuscular, while the latter are mainly diurnal. Now there can be no comparison as to brilliancy between the vast majority of these two groups. The moths vary for the most part from dingy grey to dusky black, while the but- terflies revel in every shade of golden yellow, splendid crimson, and metallic blue. Again, the eyes of these two divisions differ in structure in a manner which suggests the inference that the diurnal insects are much better pro- vided with optical discrimination than their nocturnal allies,^ especially when the facts are compared with cer- tain exactly similar or corresponding peculiarities in the nerve-terminals in the eyes of owls and bats.^ Nor does the argument stop here. Certain species or families of moths fly by day, and these [e.g., the crimson- speckled Deiopeia pulcliella, Callimorplia dominula, the Agaristidce, ^gcriidm, Zygmnidm, &c.) are as brightly tinted as any butterflies. Mr. Bates mentions a Brazilian Urania, ^ See a paper by Mr. B. T. Lowne, F.L.S., in Proc. Roy. Soc, No. clxxxvi. p. 26 r, 1878. ^ See below. DIRECT RE A CTION. x 5 1 " a beautiful tailed and gilded moth, whose habits are those of a butterfly ; " ^ and I know by personal experience the Jamaican species of similar tastes, whose wings are exqui- sitely dappled with black, green, and gold. Indeed, it may be stated generally that most brilliant insects are fond of displaying themselves in the open sunlight ; while conversely most insects which frequent dark places or fly by night alone are dusky and ugly. By the side of these facts it is well to remember that diurnal flowers, which appeal to bees and butterflies, have corollas in every variety of red, blue, orange, and purple ; while nocturnal flowers, which appeal to moths, are generally white or pale yellow in hue. If we compare the carrion-feeding and omnivorous flies with the flower-haunting Lepidoptera, we see at once the difference of taste, as exhibited in the presence or absence of sexual selection. The flies are generally dark and inconspicuous, with thin transparent wings; and what- ever beauty they possess is due to mere surface-play of interference-colours, not to the existence of distinct pig- ments. Nothing in the nature or appearance of their ordinary food-stuffs would lead us to credit them with any ancestral love for pure and beautiful hues. There are, however, some striking exceptions amongst the dipterous insects, which fully bear out our general conclusion. The tribe of Brachystomatidce are " large flies, adorned with brilliant colours, which for the most part haunt flowers, living upon honey." ^ The Xotacanthte " are also frequently brilliantly coloured. They generally frequent flowers." The Conopidse, too, " are elegantly variegated in their colours. They may be found in great abundance during the summer, hovering upon their power- ful wings over flowers in gardens and elsewhere." The invariability of this conjunction will hardly allow us to recrard it as accidental. o 1 The Naturalist on the Amazons, ^ Dallas, The Animal Kin^dum, p. 105. p. 191. 152 THE COLOUR-SENSE. Beetles or Coleoptera show us like results. The car- rion-feeders are for the most part black and unattractive, as are also the nocturnal species and those which live in water. But the brilliant species are often flower-feeders, and fly much in the sunlight, exhibiting their exquisite metallic sheen, and displaying their beauty to their mates. If we take the Lamellicorn beetles in particular, we shall find a very instructive difference between two of their closely-allied families. The cockchafers feed on leaves, and they are some of the dingiest creatures of their class ; but the Cetoiiiadcc feed upon flowers, as their English name of " rose-beetles " implies, and they are conspicuous for the beauty of their colouring, including " a vast num- ber of the most brilliant exotic species." " It is a signifi- cant fact, too, that their mandibles have been specially modified, so as to enable them to lick up honey, which clearly shows a long persistence in flower-haunting habits, quite sufi&cient to account for the formation of a definite taste for colour. " Those species," says Latreille, speak- ing of Lamellicorns generally, " which live in the perfect state upon vegetable substances, are remarkable for the brilliancy of the metallic colours with which they are adorned. But the majority of the other species, which subsist on decomposing vegetation, manure, tan, or excre- mentitious matter, are generally of a uniform black or brown hue." The magnificent Buprestidse are also, in many cases at least, flower-haunters. The tetramerous beetles, including the gorgeous Longicorns, may be regarded as mainly flower-feeding or plant-haunting insects, and their colours, as a rule, are very brilliant. Similarly, among heteromerous beetles, the Trachelia of Professor West- wood are active diurnal animals, most of which live upon the leaves or suck the honey of flowers, and they are often adorned with beautiful colours ; but the Atrachelia, nocturnal in their habits and foul feeders, are generally black and dingy in hue. Altogether, though it would be difficult to sum up so very varied a group as the Coleop- DIRECT RE A CTION. 153 tera in a single sentence, I think a careful examination will convince the inquirer that here, too, a general con- nection exists between brilliancy of hue and flower-feed- ing or fruit-eating habits. When we turn to the Hymenoptera, or bee and wasp tribe, a great difficulty at first sight arises in our way. It would seem as though some of these insects ought to be of all others the most gorgeously arrayed, and yet for the most part they are but plain and inconspicuous creatures. However, a closer view dispels the doubt. Only one tribe of the Hymenoptera, that of the Anthophila, or bees, is specially adapted for feeding on flowers. Now these fall into two classes, the social and the solitary ; and the habits of the former class, of course, place them almost entirely outside the sphere of sexual selection. The queen or mother-bee, a prisoner for life, does not herself seek honey among flowers, and those bees which do so have no power of transmitting their tastes or habits to descendants. Indeed, the whole question of heredity in these interest- ing animals remains involved in so much mystery, that it would be useless to base any arguments upon it in either direction. On the other hand, the solitary bees are often beauti- fully coloured, as in the well-known case of the carpenter- bee. The Nomadae, or cuckoo-bees, are also very brilliant insects. The omnivorous wasps do not exhibit equal beauty ; and the almost wingless, highly social, and mainly carnivorous ants are quite inconspicuous animals, pro- bably possessing colour-perception in a very sight degree. But the Chrysidse, a family of lower Hymenoptera, are also solitary flower-haunters, and " in the richness of their colours they vie with humming-birds." Of course, it cannot be denied that a few less notable classes of insects which do not haunt flowers are never- theless more or less brilliant in their colouring. But this does not interfere with the general truth of our inference that flower-feeders are specially noticeable for their bright 154 THE COLOUR-SENSE. lilies. If we can find ground for believing that those spe- cies which habitually seek their food in gay blossoms have developed a peculiar love for colour, which is shown in their choice of mates, we shall have done quite as much as is needful. Besides, other sources exist from which a love for colour may be derived as well as from flowers. For example, the Orthopterous family of Mantidse, or praying- insects, are noticeable in many instances for their bright tints ; but as they live by devouring other insects, a taste of the sort may have been generated indirectly in their case from the nature of their food. Still, most of the Mantidse seem rather to be deceptively coloured like their surroundings, so as to escape the notice both of their prey and of their enemies among birds. A similar explanation must be given in the case of the Phasmidae, or leaf and stick insects, whose colouring, though sometimes compara- tively striking when seen in a cabinet, is purely imitative of the foliage or fallen sticks around them. Many flower- haunting spiders, too (to travel for a moment outside the limits of the true insects), are " exquisite gems " of ruby or sapphire colouration ; yet we must rather attribute their magnificent hues to the need for imitating the petals on which they creep than to sexual selection. Such instances, however, in no way militate against our main conclusion ; they only show that other causes at work have sometimes produced similar results to those which we are contem- plating, though in a different manner. Thus it is quite possible that the beauty of the tiger-beetles may be due to their habit of huntincf other brio^ht- coloured insects in the open sunlight. There still remains a margin of inexplicable cases, as might naturally be expected, for the study of these questions is yet in its infancy, and only a few isolated endeavours have hitherto been made to account at all for the external appearance of animals. Among such may be mentioned the gorgeous tropical locusts, the dragon- flies (which, however, prey upon many brilliant species), DIRECT RE A CTION. 155 and several of the Longicorn beetles. But all these in- stances cannot blind us to the fact that if we look at the flower-haunting insects in the mass they are by far the most conspicuous for beauty of their kind. It is not necessary to explain in detail the colouring of every in- dividual species — an endless task, which would demand far more competent treatment than I could give : it is quite sufficient if we find a general coincidence between bright food and bright hues in the feeder, without pretending at once to account for every apparent ex- ception. And now, before we pass on to examine the vertebrate world in the same manner as we have here examined the articulate, we must pause a moment to meet, or rather to touch lightly, a powerful objection which has been urged against the whole theory of sexual selection by no less a writer than Mr. A. E. Wallace. In his work on " Tropical Nature," that ingenious evolutionist endeavours entirely to overthrow Mr. Darwin's laborious superstruc- ture, raised with so much toil and skill in the " Descent of Man," and to substitute for the doctrine of A'oluntary choice, which the older naturalist there advanced, a number of minor principles, whose joint action may be supposed to have produced the existing colours of the animal world. Mr. Wallace has urged his objections with even more than his usual ingenuity ; and I may frankly confess that he has attacked the theory of sexual selection with such judicious vigour that I felt inclined on first reading his essays to abandon entirely all that part of the present work which was based on the orio-inal doctrine enunciated by Mr. Darwin. On fuller consideration, however, I have determined, though with much hesitation, to retain it, in hopes that the few suggestions which I have to make upon the question may possibly contribute to a clearer comprehension of its issues, and to its ultimate settle- ment in one direction or the other. I cannot for a moment pretend to meet a distinguished specialist like 156 THE COLOUR-SENSE. Mr. "Wallace on liis own ground ; nor do I wish to dis- pute the force and accuracy of many among his criticisms ; yet I trust I may be able to add my small quotum of facts and inferences to the whole data for a final opinion, and I believe that the very generalisation which it is the object of the present chapter roughly to establish, may be useful in showing some additional basis for the theory of sexual selection. For if we find that ^fruit-eating and flower-feeding animals do really exhibit unusually beauti- ful colours, then we shall have some further ground for believing that they do exert some vague sort of choice or preference in the search for mates. Accordingly, I shall jot down in passing, under each head, such points as occur in relation to this vexed ques- tion. In the first place, it is well to remember that sexual selection does not necessarily imply a deliberate exercise of will, or comparison between the rival charms of various possible mates, which seems hardly probable in the case of insects at least. We must guard against the error of transferring our own highly-developed notions of beauty to the simple half- conscious minds of beetles or butter- flies. With us, beauty is a very complex idea, compounded of numerous presentative, representative, and re-represen- tative elements ; and our choice of mates is a conscious selection, guided more or less deliberately by many com- plicated considerations, often too numerous for analysis even in our own minds. But, without attributing to the butterfly any such highly-evolved feelings, w^e may well believe that certain individuals, whose brilliant colours contrasted strikingly with the green foliage about them, might more readily succeed in attracting the attention of mates than their dingier compeers. We know that the eyes of insects are allured by the colours of flowers, which have been developed for this very purpose : and there is there- fore nothing improbable in the supposition that they are also allured by bright hues in their fellows. In short, I am DIRECT RE A CTION. 1 57 inclined to suggest that conspicuousncss rather than leauty in our human sense is aimed at by the butterfly's wing. To some this will doubtless appear equivalent to a sur- render of the whole position ; but a little reflection will probably remove such an apprehension. For the sense of beauty in its simplest form, as Mr. Darwin rightly puts it, is nothing more than " the reception of a peculiar kind of pleasure from certain colours, forms, and sounds." ^ Now we have seen reason to believe that the insect feels some slight pleasure in the perception of colour in flowers ; and we may also conclude that the pleasure is equally felt from the similar stimulation of a brilliant mate. In either case, it seems probable that a semi-automatic action is set up by the sight of the bright hue, which leads on the insect instinctively to the blossom or the opposite sex alike. Such a selective process does not seem to me at all to transcend the narrow faculties of a beetle or a butterfly. Mates on this theory are not chosen on account of their brilliancy, but their brilliancy renders them the most natural objects to choose. The familiar instance of the moth and the candle shows us this automatic tendency in its fullest form. In that case it w^ould seem as though the intensity of the visual stim- ulus set up a motor activity of the wings, which w^ould become more and more powerful the more directly the eyes of the insect were turned towards the light. Accordingly, any random movements in that direction would be fol- lowed by more and more rapid gyrations, ending, as we know, in the central flame, whenever the eyes both pointed straight toward that quarter. We must conclude here that in the natural circumstances of moths few bright objects would occur around them, except flowers, and so the eye has probably been connected with the motor system in such a manner that the reception of a light- stimulus acts immediately upon the wings. In the presence of such a rare and unpremeditated object as a candle, the hereditary ^ Origin of SiJecies, sixth edition, p. 162. 158 THE COLOUR-SENSE. instinct or organised habit becomes a bad guide, finds itself at fault, and finally results in the insect's death. But the case is very much like that of a human child, who knowing that bright red berries are usually sweet and innocuous, poisons itself with those of the cuckoo-pint or the yam. The instinctive pleasure can only have been adapted to the usual environment of the race, not to special and excep- tional circumstances like artificial lights and poisonous berries. It is possible that the light of fire-flies and glow-worms may be similarly of use for the guidance of the sexes, though Mr. Belt believes it to be a warning mark of inedi- bility. However this may be, it is certain that butterflies and insects generally are so constituted that they can dis- tinguish their own mates unerringly from all others, even though the difference between the species be almost microscopical. Whether this discrimination be due to sight, or, as some authors maintain, to smell, it shows equally that a minute correspondence exists between the senses of the insect and its habits of life. And as this correspondence, so far as each separate species is con- cerned, must have had a beginning at some time or other, and consequently a cause, there seems no sufficient ground for doubting that conspicuousness of colour formed one of the determining conditions. Indeed, I cannot myself see why Mr. Wallace, who allows the attractive nature of colouring in flowers, should deny its attractive nature in the question of sex. It is allowed on all hands, I believe, that the special odours of insects,! as well as their stridulatingj noises,^ are guides to the sexes in their search for one another. It does not appear that colour stands on any different footing in this respect. Furthermore, many insects have two sets of colours, apparently for different purposes, the one set protective 1 See r. Miiller in " Nature," November 29, 1877. 2 Darwin, Descent of Man, passim. DIRECT RE A CTIOX. 1 59 from tlie attacks of enemies, the other set attractive for the opposite sex. Thus several butterflies have the lower sides of their wings protectively coloured, so as to preserve them from the notice of birds or lizards, while they sit with folded wings on a flower or leaf ; whereas the upper sides are attractively coloured, and displayed in the open sunlight as they flit about in search of mates. Moths, again, whose habits of folding the wings are exactly opposite, often have their upper surfaces imitative or pro- tective, while tlie under sides are bright and beautiful Sometimes the union of protective and attractive features in the same insect is very striking. Thus Mr. WaEace himself mentions a leaflike butterfly, Kallima jparalekta, whose w4ngs are purple and orange above, but exactly mimic dead foliage when closed ;^ this insect always rests among dead or dry leaves, and imitates every stage of decay, being even apparently spotted with small fungi. So, too, Mr. Bates tells us of a grasshopper, Pterochroza, whose sheath-like fore-wings similarly resemble a green leaf, while its hind-wings, usually covered except during flight, are " decorated with gaily-coloured eyelike spots." ^ Again, Mr. Belt observes that the males of some butterflies which mimic the Heliconidae are coloured with black and white, quite unlike the mimicking females,^ while some South African species show perfectly marvellous differ- ences in this respect. In all these cases, one cannot but believe that while the one form of colouring has been acquired for the sake of protection, the other must differ from it for some sufficient functional purpose. Once more, there seems to be a pretty constant connec- tion between the general beauty of the flora in any particular district and the general beauty of its insect 1 Malay Archipelago, p. 131. vious truth that white and pale yel- 2 The Naturalist ou the Amazons, low are really very brilliant colours r- 145- when comi)ared with the green or 3 Mr. Wallace seems to me to have brown of ordinary life. See " Tropi- quite perverted the simple explana- cal Nature," p. 204. tion of this fact by neglecting the ob- i6o THE COLOUR-SENSE. inhabitants. Of course, it lias long been noted that where few or no flying insects exist, few or no bright-coloured flowers are found. But what I wish to point out here is the converse fact that where bright blossoms are common, insects are brilliant, while where most blossoms are inconspicuous, most insects are dingy. On this head the mass of evidence, though difficult to quote, is over- whelming. A few of the more striking instances may, however, be briefly given here. On the whole, the brightest flowers grow among the tropics, and on the whole, tropical insects are unusually beautiful. The flora and the minor fauna of Madagascar are equally remarkable for their splendid hues. Sir Joseph Hooker notices the extreme magnificence of the Himalayan flora; and he also mentions the singular loveliness of the butterflies.^ Sir Emerson Tennant speaks continually of the beauty of the " brilliant flowering shrubs " in Ceylon,^ and he likewise speaks of the beauty of its butterflies in large numbers.^ In Mr. Bates's work on Brazil, I notice almost on every page the conjunction of pretty insects with striking flowers, or the absence of both together.* Especially does he note the beauty of the Longicorn beetles found on flowers^ at Caripi, while a few pages before he remarks upon the abundance of exquisite blossoms at the same place.^ I cannot help interpolating here, though out of proper order, the remark that, just in like manner, he seldom mentions the capture of a handsome bird without adding that it was shot in a fruit-tree. Almost the only bright butterfly which I ever observed in large numbers in Jamaica was the Calliclryas eubule, feeding on the abundant yellow cactus blossom, whose hue it exactly resembled, and which is the only common and conspicuous large entomo- philous flower in the colony. Both Mr. Wallace himself 1 Himalayan Journal, i. 152, and ^ See the Naturalist on the Ama- ii. 98. zons, pp. 20, 274, 2 Ceylon, pp. 87, 88, 92. ^ Ibid, p, no. 2 Ibid, pp. 247, 248. 6 Ibid, p. loi. DIRECT RE A CTION. 1 6 1 and Lord George Campbell,^ an excellent non-scientific observer, remark upon the beauty of the insects and flowers of the Ke Islands. Similarly, the flowers of Amboyna are paralleled by its gorgeous beetles, butterflies, and birds.^ On the other hand, in ISTew Zealand " there are scarcely any gay flowers and blossoms ; but few her- baceous plants, nothing but shrubs and trees; shrubs with obscure green flowers ; " ^ while at the same time " the butterflies are distinguished neither by size nor by richness of colour." * Oceanic islands, which have few or no bright flowers, are remarkable for the absence of bright insects ; and Mr. Darwin mentions of the Galapagos group both the fact that he " did not see one beautiful flower," and also the universal dinginess of the whole fauna. But this question is one on which it is difficult to quote positive authorities : it must rather suffice to mention that a con- siderable search into the general impressions of travellers — the best evidence, after all, on so indefinite a point — has convinced me that such a general relation does actually obtain. It is no answer to say that the insects are neces- sary for the production of the flowers : the real point at issue is this — why are insects bright where bright flowers exist in numbers, and dull where flowers are rare or in- conspicuous ? We can hardly explain this wide coinci- dence otherwise than by supposing that a taste for colour is produced through the constant search for food among entomophilous blossoms, and that this taste has reacted upon its possessors through the action of unconscious sexual selection. Finally, it would seem that Mr. Wallace's own theory of " typical colours " really allows all that is here required. For Mr. Wallace speaks distinctly of the " need of recog- nition and identification by others of the same species " ^ 1 Log-Letters from the Challenger, 133. See also Six- Joseph Hooker's p. 187. "Flora of New Zealand," p. 28. 2 Ibid., p. 208. "* Hochstetter, p. 170. 2 Hochstetter's New Zealand, p. ^ Tropical Nature, p. 215. L i62 THE COLOUR-SENSE. as one among the determining causes of sucli colours, not only in the case of birds, but also in that of butterflies, four families of which he specifies by name. ISTow, con- spicuousness of hue is certainly a very simple means of identification : and I think we must allow that it acts as an allurement to the eye in the case of flower-feeding species. I am quite disposed to accept Mr. Wallace's belief that the actual disposition of the stripes, spots, and lines, is a matter of special typical arrangement ; but even here one would naturally suppose that some minute cause must at first have led to the preference for one arrange- ment over another. Briefly, to sum up the whole question, after making full allowance for warning colours and for mimetic or other protective colours, there seems to remain a large margin of cases in which brilliancy exists for purely attractive purposes : while often the attractive function is combined with more or less of protective device. Anybody who watches our own English butter- flies on a sunny day can hardly doubt that display forms a part of the object for which their yellow, orange, or crimson-spotted wings have been developed, and that such display makes them an easier mark for their scattered mates. Above all, it is necessary to remember that the winged condition in these insects is hardly more itself than a sexual device for the perpetuation of the various species. And now let us pass on to consider the fuller evidence afforded us by vertebrates. To begin with fishes, it must be allowed that our present knowledge of their habits scarcely justifies us in making any distinct inferences from their colouring. ISTevertheless, a few facts may perhaps be gleaned even here. The mass of lower marine animals are brilliantly coloured with what seem to be purely adventitious or protective colours ; and we had occasion already to remark upon the similarity between their hues and those of fungi, saprophytes, and other like vegetal organisms. As a few examples may DIRECT RE A CTION. 163 be mentioned sea-anemones, star-fisli, ecliini, medusae, ascidians, sea-slugs, and corals. When we reach the moUusca, the colouring begins to assume a different type, but it would be difficult to assign any sufficient cause ibr its occurrence. Amongst the marine articulata, and espe- cially the crustaceans, many species exhibit a regularity of hue, and minute arrangement of spots and lines, which seems to bespeak a certain amount of sexual selection. These doubtful instances, however, we must pass over, not because they are less interesting, but because they are so very uncertain ; and in a brief examination like the present, we must necessarily confine our attention to the most salient points. Now, the abundance of coloured organisms, both animal and vegetal, in the sea, affords a fair ground for belief that fishes may have acquired o^colour-sense, and a taste for bright hues. We know that they (as well as the crustaceans) can be attracted by crimson or scarlet rags, and that glistening objects like rnetals or artificial baits rapidly seize upon their attention. As to the brilliancy, beauty, and regularity of their colouring, the reader must be referred to Mr. Darwin's description in the " Descent of Man," where he will find a full account of the principal facts which go to prove the existence among them of sexual selection. I must content myself here by saying that for gorgeous colouring and variety of patterns they are nowhere surpassed in the whole animal kingdom; and that metallic sheen is especially conspicuous among the devices whereby they insure the attention of their mates. As regards the special question upon which we are now engaged, a few facts may be shortly set down. In the first place, the lower animals of tropical seas are on the whole much more brilliantly coloured than those of temperate climates, and the same remark holds good of the fishes. Mr. Darwin has noticed the extreme beauty of the shoals which played in and out among the brilliant organisms of the coral lagoons; and though Mr. Wallace objects that 1 64 THE COLOUR-SENSE. tliis may be due to protective causes, in order tliat they may escape notice among the bright creatures about them, y^et as Mr. Darwin pertinently replies, he was struck by the obvious conspicuousness of their appearance rather than by their resemblance to environing objects. I have myself observed the same point frequently in the West Indian harbours, where the fish and the neioiibourincj creatures, seen through clear still water, all appeared equally beautiful and noticeable. But I cannot do better than quote Mr. Wallace's own description of the harbour of Amboyna. " The bottom," he says, " was absolutely hidden by a continuous series of corals, sponges, actiniae, and other marine productions of magnificent dimensions, varied forms, and brilliant colours. ... In and out among them moved numbers of blue and red and yellow fishes, spotted and banded and striped in the most striking manner, while great orange or rosy transparent medusa3 floated along near the surface." ^ And elsewhere he ob- serves, " The fishes (of Amboyna) are perhaps unrivalled for variety and beauty by those of any one spot on the earth." These facts at least tend to show that our theory does not receive any active contradiction from the condi- tions of marine existence ; and they are confirmed by numerous other like passages in several authors whom I think it superfluous to quote. On the other hand, the larger predatory species, such as sharks and pikes, together with the majority of temperate fishes, are decidedly wanting in brilliant hues. Mr. Wallace observes that river fish, even of the tropics, rarely if ever have gay or conspicuous markings ; ^ and this is just what we would expect from the nature of their food, consisting as it does of worms, small flies, and other inconspicuous objects. However, we must allow that in this case Mr. Wallace has witnessed against himself with excessive fervour ; for many river fish undoubtedly liave very 1 Malay Archipelago, p. 295. " Contributions to the Theory of Natural Selection, p. 55. DIRECT REACTION. 165 brilliant colours, as I can especially testify from having caught in my youth numbers of the Canadian sun-fish, perhaps the most exquisite creature of its class which I have ever seen. But this particular species lives in shallow marshy water, and maij have derived its tastes from butter- flies and dragon-flies. Professor Agassiz speaks much, too, concerning the beauty of the Amazonian fishes,^ which he describes as having a quasi-marine character ; ^ and in spite of a possible accusation for grotesqueness, I would venture to suggest that their colouring may perhaps be due to those numerous butterflies which Mr. Bates so often describes as flitting in numbers along the banks of that mighty river. However this may be, we must leave these somewhat inconclusive cases, and hurry on to the terrestrial vertebrates. The amphibia yield little evidence in either direction. The beautiful colouring of tree-frogs, when compared with toads and common frogs, is more probably protective than attractive. Still, the newts show us very unmistakable signs of sexual selection in their crests ; and we cannot say that the habits of amphibia may not have generated a love for colour. Among the reptiles, however, a good many facts may be quoted to our purpose. In the first place, the large water-haunting crocodiles and alligators are peculiarly dull and unsightly objects; while the whole order of Chelonia, including the turtles and tortoises, are as incon- spicuous for colour as they well can be. On the other hand, the smaller saurians, many of which lead an ar- boreal life, and feed off varied food, sometimes fruit, some- times insects and other small animals, are often noticeable for their beauty. Most iguanas, great jungle lizards, con- trast strongly in hue with the crocodiles ; but one species, AmUyrlujnclius cristatus, which inhabits the dull-coloured Galapagos Islands, and has taken to a strangely abnormal marine life, is remarkable for the same sombre tints ^ A Journey in Brazil, p. 184. - Ibid., p. 238. 1 66 THE COLOUR-SENSE. wliicli characterise the other animals of that singular archipelago, being described by Mr. Darwin as of " a dirty- black colour." The lesser lizards give unmistakable proofs of sexual selection in the brilliant pouches which they protrude when sunning themselves, and whose exqui- site colours have struck every observer. Their mechanism for changing the hue of their skin, by compressing or spreading the layers of pigment cells, has already been noticed ; and it seems to betray a considerable sensitive- ness to colour. Dr. Giinther also sees reason to believe that the frugivorous lizards have the tongue as an organ of taste, while in the insectivorous species he considers it merely an organ for the prehension of prey.^ Now, a large number of all lizards are noticeable for their exqui- site hues. Green, as might naturally be supposed, from protective reasons, forms the general groundwork of their colouring; but, as often happens under similar circum- stances, many other shades are intermingled, apparently to perform the attractive function. Especially is this noticeable in the Anolis and other like genera, whose beautiful orange pouches consist of folds of the skin, which are concealed under ordinary circumstances, but protruded for display when the animal feels himself secure, and can sun himself at leisure on a dead branch. The family of Agamidm include many of the most bril- liant species, especially the exquisite Draco, whose beauty Mr. Darwin extols so highly ;2 and these, says Dr. Giinther, are arboreal in their mode of life, while the dull- coloured genera inhabit rocks or plains.^ It is true that Draco feeds on insects; but when we remember the beauty of many among these little tropical creatures, we see fair grounds for believing that its habits may have led it to form a taste for colour. The Geckos also eat moths ; * ^ Eeptiles of British India, p. 56. amongst butterflies, birds, these liz- - I shiill note hereafter the singular ards, and even the flying squirrels, coincidence betM^een the possession of &c. a flying apparatus and general bril- ^ Reptiles of British India, p. 120. liancy of hue, which seems to obtain ■* Giinther, p. 100. DIRECT REACTION. 167 while Mr. Gosse found on dissection many pretty insects in the stomach of the lovely Yenus lizard of the West Indies.^ Here, too, we may even see the indirect effect of flowers and fruits ; for Mr. Wallace, after noticing the " abundance and varied colours of the little jumping spiders w^hich abound on flowers and foliage (in the Aru Islands), and are often perfect gems of beauty," 2 goes on to say about the lizards in the same place, " Every rotten trunk or dead branch served as a station for some of these active little insect-hunters, who, I fear, to satisfy their gross appetites, destroy many gems of the insect world, which would feast the eyes and delight the heart of our more discriminatincj entomoloofists." ^ Among snakes we find somewhat similar facts. W^hile the arboreal species, still having green for their ground- work, " are characterised by their vivid colouration," ^ the ground-snakes, burrowing-snakes, and water-snakes are mostly dull and inconspicuous. Of the ground-colubrides, in particular. Dr. Glinther says, " They live on the ground, and are generally of not brilliant colouration ; only a few, which frequent grassy plains, are of a bright green colour."^ The Dendrophidse eat lizards and like prey, and are usually very bright ; their colours sometimes, as in the magnificent Chrysopelea ornata, being decidedly not protective. Of course, almost all arboreal snakes feed upon various foods, such as birds, smaller reptiles, or other brilliant animals, whose colours may have served to give them a taste in that direction. I confess I attach little importance to any of these cases ; still I think it worth while prominently to call attention to the fact that most arboreal creatures are conspicuous for their excep- tional brilliancy. Doubtless much of the bridit colouration in all these o animals is more or less warning or protective. Thus, one 1 The Naturalist in Jamaica, p, 145. ■* Gimtber, p. 166. - Malay Archipelago, \). 432. 5 ibid., p. 221. 3 Ibid., p. 433. i68 THE COLOUR-SENSE. may mention Mr. Belt's "little frog that hops about in the daytime, dressed in a bright livery of red and blue." ^ Mr. Belt suspected this species to be inedible, from the staring nature of its hues ; accordingly, he offered one to some ducks, but only succeeded in making one young duck bite it amongst some meat ; and the bird " instantly threw it out of its mouth, and went about jerking its head as if trying to throw off some unpleasant taste." Then, again, there are the coral-snakes of South America, for which Mr. Darwin has fully accounted. Once more Mr. "Wallace, in his " Contributions to the Theory of Natural Selection," has pointed out that green lizards frequent trees, while many Geckos are so marbled as to resemble the bark on which they crawl.^ Mr. Belt speaks also of a green Mcaraguan species, which looks exactly like- the herbage among which it lurks, and has actually acquired leaf-like expansions to deceive its prey.^ Mr. Bates, too, notices a pale-green snake {Dryopliis fulgida) so perfectly imitating the stem of a liana that it deceived even his practised eye at first sight ; ^ and we must, doubtless, refer to the same cause the verdant colour of the grass-snakes mentioned by Dr. Gtinther. But the noticeable point here, as in the case of the butterflies, is thi.s, that while we find a prevailing imitative greenness, apparently for protective purposes, we so often find a mixture of crimson, blue, yellow, orange, or metallic iridescence, whose function seems to me purely attractive. We shall notice similarly, when we come to look at the parrots, that their prevailing ground-tint is likewise green, but that they indulge in every variety of brighter pigments in a decidedly conspicuous manner. My own observation of West Indian lizards would certainly lead me to say that their colours were far more likely to betray them than to protect them, even in their native haunts. ^ Naturalist in Nicaragua, p. 321. ^ Naturalist in Nicaragua, p. 12. 2 See also Tropical Nature, p. iii, ^ Naturalist on the Amazons, p. seq. 99. DIRECT REACTION. 169 Birds, however, offer the best evidence of alh It will be desirable to mark briefly the most conspicuous instances, and then to give the minor cases in detail. The birds of prey — eagles, hawks, and falcons — and the carrion birds — vultures, condors, and (since we are speak- ing of habits only) ravens or adjutant storks — are all dull greyish or blackish birds,^ and their colouring may be com- pared to that of the flies and crocodiles. Nocturnal birds, again, such as owls and goat-suckers, show considerable analogies to moths and other night-flying insects.^ In fact, aU. the raptores, of whatever habit, and almost all birds of similar habit elsewhere, are nearly or quite desti- tute of decorative colouring. On the other hand, among the insessores we find an immense number of the most brilliant of all organic creatures. Especially remarkable are the humming-birds and the sun-birds. Now, Prince Lucien Bonaparte has abundantly shown that the former family are really the allies of our dingy northern swifts, and that the sun-birds are not at all connected with them genetically. But both families feed upon the mixed nectar and insects which they catch in bright-coloured entomophilous flowers, and both are equally noticeable for their exquisite metallic gloss, their varied hues, and the profusion of their decora- tive devices, such as crests, ruffs, feather lappets, and long tail-plumes. Is it not a significant fact that these two families, one in the western hemisphere and the other in the eastern, separately developed from dingy ancestors, should have acquired exactly the same exquisite plumage under exactly like conditions of food ? We can hardly resist the inference that a taste for colour has been aroused 1 I am aware that all sucli general- its ensemble, and not to look at single isatious will be attacked by bringing cases. up isolated instances, such as the - While following approximately king- vulture, who may lay some claim the real biological order in these cases, to be moderately coloured ; but I must I do not scruple to introduce analo- beg the reader to take each group in gous instances from any other tribe when necessary. 170 THE COLOUR-SENSE, in their constant search after flowers, and that this taste has reacted through sexual selection upon their own appearance. JSText in importance to these two families come the parrot group. These are either fruit-eaters, or else, as in the case of the lories, they feed upon nectar. And here I venture to borrow Mr. Wallace's words. "No group of birds," says he — "perhaps no other group of animals — exhibits within the same limited number of genera and species so wide a range and such an endless variety of colour. As a rule parrots may be termed green birds, the majority of the species having this colour as the basis of their plumage, relieved by caps, gorgets, bands, and wing- spots of other and brighter hues. Yet this general green tint sometimes changes into light or deep blue, as in some macaws; into pure yellow or rich orange, as in some of the American macaw-parrots (Conurus) ; into purple, grey, or dove-colour, as in some American, African, and Indian species ; into the purest crimson, as in some of the lories ; into rosy-white and pure white, as in the cockatoos ; and into a deep purple, ashy or black, as in several Papuan, Australian, and Mascarene species. There is in fact hardly a single distinct and definable colour that cannot be fairly matched among the 390 species of known parrots. Their habits, too, are such as to bring them prominently before the eye. They usually feed in flocks ; they are noisy, and so attract attention ; they love gardens, orchards, and open sunny places ; they wander about far in search of food, and towards sunset return homewards in noisy flocks, or in constant pairs. Their forms and motions are often beautiful and attractive. The immensely long tails of the macaws, and the more slender tails of the Indian parra- quets ; the fine crest of the cockatoos ; the swift flight of many of the smaller species, and the graceful motions of the little love-birds and allied forms ; together with their affectionate natures, aptitude for domestication, and powers of mimicry — combine to render them at once the most DIREC T RE A CTION. j 7 1 conspicuous and the most attractive of all tlie specially tropical forms of bird-life." ^ Even the minor variations of these three great groups — the humming-birds, the sun-birds, and the parrots — show us a like result. For there is one sub-family of the former group — the Phaethornidas — which have not taken to flower haunting, but which catch minute insects on exposed situations; and these are described by Mr. Wallace, not in that language of sapphire, ruby, and amethyst which is lavished on their congeners, but simply as " small brown humming-birds." 2 "The members of all these genera," says Mr. Gould in his magnificent work on the Trochilidse, " are remarkable for being destitute of metallic brilliancy, and, as their trivial name of ' hermits ' implies, for affect- ing dark and gloomy situations. They constitute perhaps the only group of the great family of humming-birds which frequent the interior of the forests, and there obtain their insect food — some from the underside of the leaves of the great trees, while others assiduously explore their stems in search of such lurking insects as may be concealed in the crevices of the bark. It has been said that spiders constitute the food of many species of this group." And he adds significantly a few lines further down, " in the colouration of their plumage both sexes are generally alike." ^ Then, again, we learn of the Arachnotherae, or spider-hunters, " which are sun-birds without any metallic or other brilliant colouring," that they hunt for food among the anemophilous and uncoloured blossoms of the palm- trees.* So, too, among the sombre vegetation of New Zea- land an anomalous night-parrot {Strigaps hctbroptilus) is found, which lives in crevices of the ground, or in rocks 1 Tropical Nature, p. 100. argument here employed, for the - Ibid., p. 136. brilliant mass of colour would natu- 2 JMonograph. of the Trochilidse, rally he to them the empirical symbol Introduction, p. 36. of food, and they cannot possibly dis- ^ Of course the other humming- tinguish between the circumstances birds and sun-birds also live mainly which lead to the presence of honey upon insects found in flowers, but and of insects in the blossoms they this does not militate against the suck. 172 THE COLOUR-SENSE. and tree-roots, only coming out after dark, and its colour is spoken of as " dull yellowish green." ^ Bfere one may feel almost certain that the primitive bright hue has be- come faded and dingy owing to the altered habits of the bird, which would effectually prevent the action of sexual selection. Scarcely less interesting are the group of pigeons, which fall under two principal heads, so far as our present purpose is concerned, the fruit-pigeons and the ground- pigeons. The former class are extremely brilliant in their colouring, comprising a large number of the most beauti- ful knoAvn birds ; while the latter almost always display sombre dove-colours, slates, and browns.^ ]N"ow, the fruit- pigeons are "especially arboreal in their habits," and " their nourishment consists for the most part of fruits :" while the ground-pigeons feed almost entirely on seeds. The toucans form another group in which like adapta- tions occur. They live nearly altogether upon fruits, though they also devour birds' eggs, fish, reptiles, and insects, to a slight extent. The exquisite colours wdiich adorn their large bills, besides the varied black, white, green, red, and yellow of their plumage, are well enough known to call for no further detail.^ Several other families, allied to one or other of the preceding groups, are almost equally noticeable for their magnificent colouration. First on the list may come the Australian honey-suckers, and the plantain-eaters of Africa, whose name sufficiently proclaims their habits. I^ext, we may place the allied genus of Touracos, " gene- rally of a green colour, with the quill feathers of the wing and tail violet or red." The birds-of-paradise, too well known to need description, feed on fruits, though some species are flower-suckers. The barbets, known by such 1 Hochstetter's New Zealand, p. 3 gee on the colours and food of 167. toucans, Gould, "Monograph of the - Jerdon, Bii'ds of India, vol. ii. p. Eamphastid*,"i9