Tropical nature, and other essays

by Alfred Russel Wallace

We find, then, that all the chief peculiarities of the three allied species of humming-birds which inhabit the Juan Fernandez group of islands, may be fairly traced to the action of those general laws which Mr. Darwin and others have shown to determine the variations of animals and the perpetuation of those variations. It is also instructive to note, that where the variations of colour and size have been greatest they are accompanied by several lesser variations in other characters. In the Juan Fernandez bird the bill has become a little shorter, the tail-feathers somewhat broader, and the fiery cap on the head somewhat smaller; all these peculiarities being less developed or absent in the birds inhabiting Mas-afuera. These coincident changes may be due, either to what Mr. Darwin has termed correlation of growth, or to the partial reappearance of ancestral characters under more favourable conditions, or to the direct action of changes of climate and of food; but they show us how varied and unaccountable are the changes in specific forms that may be effected in a comparatively short time, and by means of very slight changes of locality.

If now we consider the enormously varied conditions presented by the whole continent of America—the hot, moist, and uniform forest-plains of the Amazon; the open llanos of the Orinoco; the dry uplands of Brazil; the sheltered valleys and forest slopes of the Eastern Andes; the verdant plateaus, the barren paramos, the countless volcanic cones with their peculiar Alpine vegetation; the contrasts of the East and West coasts; the isolation of the West Indian islands, and to a less extent of Central America and Mexico which we know have been several times separated from South America; and when we further consider that all these characteristically distinct areas have been subject to cosmical and local changes, to elevations and depressions, to diminution and increase of size, to greater extremes and greater uniformity of temperature, to increase or decrease of rainfall; and that with these changes there have been coincident changes of vegetation and of animal life, all affecting in countless ways the growth and development, the forms and colours, of these wonderful little birds—if we consider all these varied and complex influences, we shall be less surprised at their strange forms, their infinite variety, their wondrous beauty. For how many ages the causes above enumerated may have acted upon them we cannot say; but their extreme isolation from all other birds, no less than the abundance and variety of their generic and specific forms, clearly point to a very high antiquity.

The Relations and Affinities of Humming-birds.—The question of the position of this family in the class of birds and its affinities or resemblances to other groups, is so interesting, and affords such good opportunities for explaining some of the best-established principles of classification in natural history in a popular way, that we propose to discuss it at some length, but without entering into technical details.

There is in the Eastern hemisphere, especially in tropical Africa and Asia, a family of small birds called Sun-birds, which are adorned with brilliant metallic colours, and which, in shape and general appearance, much resemble humming-birds. They frequent flowers in the same way, feeding on honey and insects; and all the older naturalists placed the two families side by side as undoubtedly allied. In the year 1850, in a general catalogue of birds, Prince Lucien Bonaparte, a learned ornithologist, placed the humming-birds next to the swifts, and far removed from the Nectarinidæ or sun-birds; and this view of their position has gained ground with increasing knowledge, so that now all the more advanced ornithologists have adopted it. Before proceeding to point out the reasons for this change of view, it will be well to discuss a few of the general principles which guide naturalists in the solution of such problems.

How to Determine Doubtful Affinities.—It is now generally admitted that, for the purpose of determining obscure and doubtful affinities, we must examine by preference those parts of an animal which have little or no direct influence on its habits and general economy. The value of an organ, or of any detail of structure, for purposes of classification, is generally in inverse proportion to its adaptability to special uses. And the reason of this is apparent, when we consider that similarities of food and habits are often accompanied by similarities of external form or of special organs, in totally distinct animals. Porpoises, for example, are modified externally so as to resemble fishes; yet they are really mammalia. Some marsupials are carnivorous, and are so like true carnivora that it is only by minute peculiarities of structure that the skeleton of the one can be distinguished from that of the other. Many of the hornbills and toucans have the same general form, and resemble each other in habits, in food, and in their enormous bills; yet peculiarities in the structure of the feet, in the form of the breast-bone, in the cranium, and in the texture and arrangement of the plumage, show that they have no real affinity, the former approaching the kingfishers, the latter the cuckoos. Such structural peculiarities as these have no direct relation to habits; and they are therefore little liable to change, when from any cause a portion of the group may have been driven to adopt a new mode of life. Thus all the Old World apes, however much they may differ in size or habits, and whether we class them as baboons, monkeys, or gorillas, have the same number of teeth; while the American monkeys all have an additional premolar tooth. This difference can have no relation to the habits of the two groups, because each group exhibits differences of habits greater than often occur between American and Asiatic species; and it thus becomes a valuable character indicating the radical distinctness of the two groups, a distinctness confirmed by other anatomical characters.

On the other hand, peculiarities of organization which seem specially adapted to certain modes of life, are often diminished or altogether lost in a few species of the group, showing their essential unimportance to the type, as well as their small value for classification. Thus, the woodpeckers are most strikingly characterised by a very long and highly extensible tongue, with the muscles attached to the tongue-bone prolonged backward over the head so as to enable the tongue to be suddenly darted out; and also by the rigid and pointed tail which is a great help in climbing up the vertical trunks of trees. But in one group (the Picumni), the tail becomes quite soft, while the tongue remains fully developed; and in another (Meiglyptes) the characteristic tail remains, while the prolonged hyoid muscles have almost entirely disappeared, and the tongue has consequently lost its peculiar extensile power; yet in both these cases the form of the breast-bone and the character of the feet, the skeleton, and the plumage, show that the birds are really woodpeckers; while even the habits and the food are very little altered. In like manner the bill may undergo great changes; as from the short crow-like bill of the true birds-of-paradise to the long slender bills of Epimachinæ, which latter were on that account long classed apart in the tribe of Tenuirostres, or slender-billed birds, but whose entire structure shows them to be closely allied to the paradise-birds. So, the long feathery tongue of the toucans differs from that of every other bird; yet it is not held to overbalance the weight of anatomical peculiarities which show that these birds are allied to the barbets and the cuckoos.

The skeleton, therefore, and especially the sternum or breast-bone, affords us an almost infallible guide in doubtful cases; because it appears to change its form with extreme slowness, and thus indicates deeper-seated affinities than those shown by organs which are in direct connection with the outside world, and are readily modified in accordance with varying conditions of existence. Another, though less valuable guide is afforded, in the case of birds, by the eggs. These often have a characteristic form and colour, and a peculiar texture of surface, running unchanged through whole genera and families which are nearly related to each other, however much they may differ in outward form and habits. Another detail of structure which has no direct connection with habits and economy, is the manner in which the plumage is arranged on the body. The feathers of birds are by no means set uniformly over their skin, but grow in certain definite lines and patches, which vary considerably in shape and size in the more important orders and tribes, while the mode of arrangement agrees in all which are known to be closely related to each other; and thus the form of the feather-tracts or the “pterylography” as it is termed, of a bird, is a valuable aid in doubtful cases of affinity.

Now, if we apply these three tests to the humming-birds, we find them all pointing in the same direction. The sternum or breast-bone is not notched behind; and this agrees with the swifts, and not with the sun-birds, whose sternum has two deep notches behind, as in all the families of the vast order of Passeres to which the latter belong. The eggs of both swifts and humming-birds are white, only two in number, and resembling each other in texture. And in the arrangement of the feather-tracts the humming-birds approach more nearly to the swifts than they do to any other birds; and altogether differ from the sun-birds, which, in this respect as in so many others, resemble the honey-suckers of Australia and other true passerine birds.

Resemblances of Swifts and Humming-birds.—Having this clue to their affinities, we shall find other peculiarities common to these two groups, the swifts and the humming-birds. They have both ten tail-feathers, while the sun-birds have twelve. They have both only sixteen true quill-feathers, and they are the only birds which have so small a number. The humming-birds are remarkable for having, in almost all the species, the first quill the longest of all, the only other birds resembling them in this respect being a few species of swifts; and, lastly, in both groups the plumage is remarkably compact and closely pressed to the body. Yet, with all these points of agreement, we find an extreme diversity in the bills and tongues of the two groups. The swifts have a short, broad, flat bill, with a flat horny-tipped tongue of the usual character; while the humming-birds have a very long, narrow, almost cylindrical bill, containing a tubular and highly extensible tongue. The essential point however is, that whereas hardly any of the other characters we have adduced are adaptive, or strictly correlated with habits and economy, this character is pre-eminently so; for the swifts are pure aërial insect-hunters, and their short, broad bills, and wide gape, are essential to their mode of life. The humming-birds, on the other hand, are floral insect-hunters, and for this purpose their peculiarly long bills and extensile tongues are especially adapted; while they are at the same time honey-suckers, and for this purpose have acquired the tubular tongue. The formation of such a tubular tongue out of one of the ordinary kind is easily conceivable, as it only requires to be lengthened, and the two laminæ of which it is composed curled in at the sides; and these changes it probably goes through in the young birds.

When on the Amazon I once had a nest brought me containing two little unfledged humming-birds, apparently not long hatched. Their beaks were not at all like those of their parents, but short, triangular, and broad at the base; just the form of the beak of a swallow or swift slightly lengthened. Thinking (erroneously) that the young birds were fed by their parents on honey, I tried to feed them with a syrup made of honey and water, but though they kept their mouths constantly open as if ravenously hungry, they would not swallow the liquid, but threw it out again and sometimes nearly choked themselves in the effort. At length I caught some minute flies, and on dropping one of these into the open mouth it instantly closed, the fly was gulped down and the mouth opened again for more; and each took in this way fifteen or twenty little flies in succession before it was satisfied. They lived thus three or four days, but required more constant care than I could give them. These little birds were in the “swift” stage; they were pure insect-eaters, with a bill and mouth adapted for insect-eating only. At that time I was not aware of the importance of the observation of the tongue; but as the bill was so short and the tubular tongue not required, there can be little doubt that the organ was, at that early stage of growth, short and flat, as it is in the birds most nearly allied to them.