Pigeons and doves offer a very curious case of the protection of exposed eggs. They usually build very slight and loose nests of sticks and twigs, so open that light can be seen through them from below, while they are generally well concealed by foliage above. Their eggs are white and shining; yet it is a difficult matter to discover, from beneath, whether there are eggs in the nest or not, while they are well hidden by the thick foliage above. The Australian podargihuge goatsuckers—build very similar nests, and their white eggs are protected in the same manner. Some large and powerful birds, as the swans, herons, pelicans, cormorants, and storks, lay white eggs in open nests; but they keep careful watch over them, and are able to drive away intruders. On the whole, then, we see that, while white eggs are conspicuous, and therefore especially liable to attack by egg-eating animals, they are concealed from observation in many and various ways. We may, therefore, assume that, in cases where there seems to be no such concealment, we are too ignorant of the whole of the conditions to form a correct judgment.
We now come to the large class of coloured or richly spotted eggs, and here we have a more difficult task, though many of them decidedly exhibit protective tints or markings. There are two birds which nest on sandy shores—the lesser tern and the ringed plover,—and both lay sand-coloured eggs, the former spotted so as to harmonise with coarse shingle, the latter minutely speckled like fine sand, which are the kinds of ground the two birds choose respectively for their nests. "The common sandpipers' eggs assimilate so closely with the tints around them as to make their discovery a matter of no small difficulty, as every oologist can testify who has searched for them. The pewits' eggs, dark in ground colour and boldly marked, are in strict harmony with the sober tints of moor and fallow, and on this circumstance alone their concealment and safety depend. The divers' eggs furnish another example of protective colour; they are generally laid close to the water's edge, amongst drift and shingle, where their dark tints and black spots conceal them by harmonising closely with surrounding objects. The snipes and the great army of sandpipers furnish innumerable instances of protectively coloured eggs. In all the instances given the sitting-bird invariably leaves the eggs uncovered when it quits them, and consequently their safety depends solely on the colours which adorn them."[81] The wonderful range of colour and marking in the eggs of the guillemot may be imputed to the inaccessible rocks on which it breeds, giving it complete protection from enemies. Thus the pale or bluish ground colour of the eggs of its allies, the auks and puffins, has become intensified and blotched and spotted in the most marvellous variety of patterns, owing to there being no selective agency to prevent individual variation having full sway.
The common black coot (Fulica atra) has eggs which are coloured in a specially protective manner. Dr. William Marshall writes, that it only breeds in certain localities where a large water reed (Phragmites arundinacea) abounds. The eggs of the coot are stained and spotted with black on a yellowish-gray ground, and the dead leaves of the reed are of the same colour, and are stained black by small parasitic fungi of the Uredo family; and these leaves form the bed on which the eggs are laid. The eggs and the leaves agree so closely in colour and markings that it is a difficult thing to distinguish the eggs at any distance. It is to be noted that the coot never covers up its eggs, as its ally the moor-hen usually does.
The beautiful blue or greenish eggs of the hedge-sparrow, the song-thrush, and sometimes those of the blackbird, seem at first sight especially calculated to attract attention, but it is very doubtful whether they are really so conspicuous when seen at a little distance among their usual surroundings. For the nests of these birds are either in evergreens, as holly or ivy, or surrounded by the delicate green tints of our early spring vegetation, and may thus harmonise very well with the colours around them. The great majority of the eggs of our smaller birds are so spotted or streaked with brown or black on variously tinted grounds that, when lying in the shadow of the nest and surrounded by the many colours and tints of bark and moss, of purple buds and tender green or yellow foliage, with all the complex glittering lights and mottled shades produced among these by the spring sunshine and by sparkling raindrops, they must have a quite different aspect from that which they possess when we observe them torn from their natural surroundings. We have here, probably, a similar case of general protective harmony to that of the green caterpillars with beautiful white or purple bands and spots, which, though gaudily conspicuous when seen alone, become practically invisible among the complex lights and shadows of the foliage they feed upon.
In the case of the cuckoo, which lays its eggs in the nests of a variety of other birds, the eggs themselves are subject to considerable variations of colour, the most common type, however, resembling those of the pipits, wagtails, or warblers, in whose nests they are most frequently laid. It also often lays in the nest of the hedge-sparrow, whose bright blue eggs are usually not at all nearly matched, although they are sometimes said to be so on the Continent. It is the opinion of many ornithologists that each female cuckoo lays the same coloured eggs, and that it usually chooses a nest the owners of which lay somewhat similar eggs, though this is by no means universally the case. Although birds which have cuckoos' eggs imposed upon them do not seem to neglect them on account of any difference of colour, yet they probably do so occasionally; and if, as seems probable, each bird's eggs are to some extent protected by their harmony of colour with their surroundings, the presence of a larger and very differently coloured egg in the nest might be dangerous, and lead to the destruction of the whole set. Those cuckoos, therefore, which most frequently placed their eggs among the kinds which they resembled, would in the long run leave most progeny, and thus the very frequent accord in colour might have been brought about.
Some writers have suggested that the varied colours of birds' eggs are primarily due to the effect of surrounding coloured objects on the female bird during the period preceding incubation; and have expended much ingenuity in suggesting the objects that may have caused the eggs of one bird to be blue, another brown, and another pink.[82] But no evidence has been presented to prove that any effects whatever are produced by this cause, while there seems no difficulty in accounting for the facts by individual variability and the action of natural selection. The changes that occur in the conditions of existence of birds must sometimes render the concealment less perfect than it may once have been; and when any danger arises from this cause, it may be met either by some change in the colour of the eggs, or in the structure or position of the nest, or by the increased care which the parents bestow upon the eggs. In this way the various divergences which now so often puzzle us may have arisen.
Colour as a Means of Recognition.
If we consider the habits and life-histories of those animals which are more or less gregarious, comprising a large proportion of the herbivora, some carnivora, and a considerable number of all orders of birds, we shall see that a means of ready recognition of its own kind, at a distance or during rapid motion, in the dusk of twilight or in partial cover, must be of the greatest advantage and often lead to the preservation of life. Animals of this kind will not usually receive a stranger into their midst. While they keep together they are generally safe from attack, but a solitary straggler becomes an easy prey to the enemy; it is, therefore, of the highest importance that, in such a case, the wanderer should have every facility for discovering its companions with certainty at any distance within the range of vision.
Some means of easy recognition must be of vital importance to the young and inexperienced of each flock, and it also enables the sexes to recognise their kind and thus avoid the evils of infertile crosses; and I am inclined to believe that its necessity has had a more widespread influence in determining the diversities of animal coloration than any other cause whatever. To it may probably be imputed the singular fact that, whereas bilateral symmetry of coloration is very frequently lost among domesticated animals, it almost universally prevails in a state of nature; for if the two sides of an animal were unlike, and the diversity of coloration among domestic animals occurred in a wild state, easy recognition would be impossible among numerous closely allied forms.[83] The wonderful diversity of colour and of marking that prevails, especially in birds and insects, may be due to the fact that one of the first needs of a new species would be, to keep separate from its nearest allies, and this could be most readily done by some easily seen external mark of difference. A few illustrations will serve to show how this principle acts in nature.
My attention was first called to the subject by a remark of Mr. Darwin's that, though, "the hare on her form is a familiar instance of concealment through colour, yet the principle partly fails in a closely allied species, the rabbit; for when running to its burrow it is made conspicuous to the sportsman, and no doubt to all beasts of prey, by its upturned white tail."[84] But a little consideration of the habits of the animal will show that the white upturned tail is of the greatest value, and is really, as it has been termed by a writer in The Field, a "signal flag of danger." For the rabbit is usually a crepuscular animal, feeding soon after sunset or on moonlight nights. When disturbed or alarmed it makes for its burrow, and the white upturned tails of those in front serve as guides and signals to those more remote from home, to the young and the feeble; and thus each following the one or two before it, all are able with the least possible delay to regain a place of comparative safety. The apparent danger, therefore, becomes a most important means of security.