# Popular Science Monthly/Volume 30/December 1886/The Wings of Birds

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* THE WINGS OF BIRDS.
By Professor W. H. FLOWER, F. R. S.,
DIRECTOR OF THE BRITISH NATURAL HISTORY MUSEUM.
THE power of flying through the air is one of the principal characteristics of the class of birds. Although some members of the other great divisions of the vertebrates—the bats among mammals, the extinct pterodactyl among reptiles, the flying-fishes among pisces—possess this power in a greater or less degree, these are all exceptional forms, whereas in birds the faculty of flight is the rule, its absence the exception. Among invertebrates this power is possessed in a very complete degree by the greater number of insects.
In the normal structure of the vertebrate animals there are two pairs of limbs, anterior and posterior, never more. It often happens, however, that one pair, and sometimes both, are suppressed, being rudimentary, functionless, or entirely absent. Flight is always performed by the anterior or pectoral pair, more or less modified for the purpose. The superaddition of wings to arms, as in the pictorial representations of angels, has no counterpart in nature. The wings of the bird, the bat, the pterodactyl, and flying-fish, are the homologues of the arms of man, the fore-legs of beasts. In the flying-fish the power is gained simply by an enlargement of the pectoral fin, and the function is very imperfect; in the pterodactyl, by immense elongation of one (the outer) finger, and extension of the skin between it and the side of the body; in the bats, by elongation of the four outer fingers, and extension of a web of skin between them and the body. In the bird the flying organ is constructed mainly of epidermic structures, peculiar outgrowths from the surface, called feathers—modifications of the same tissue which constitutes the hair, horns, scales, or nails of other animals. Feathers are met with only in birds, and are found in all the existing members of the class, constituting the general covering of the surface of the body.
The framework to which the broad expanse formed by the feathers is attached is composed of bones, essentially resembling those of the fore-limbs of other vertebrates. The distal segment, manus, or hand, in the vast majority of birds, has three metacarpal bones and digits, the former being more or less united together in the adult state. The digits appear to correspond with the pollex, index, and medius of the typical pentadactyl manus; the second is always the longest. Both it and the pollex frequently bear small horny claws at their extremity, concealed among the feathers and functionless, but very significant in relation to the probable original condition of the avian wing. These ​claws are altogether distinct from the large, and often functional, spurs developed in many species from the edge of the metacarpal bones, resembling both in use and situation the corresponding weapons in the hind-feet. The third digit does not bear a second phalanx or claw in any existing bird.
The quills, remiges, or flight-feathers attached to the bones of the manus (called "primaries"), never exceed twelve in number, and are (as has been recently shown by Mr. Wray) in the very great majority of birds distributed as follows: Six, or in some few cases (flamingo, storks, grebes, etc.) seven, to the metacarpus; of the remainder or digital feathers, one (ad-digital) is attached close to the metacarpo-phalangeal articulation, and rests on the phalanx of the third digit; two (mid-digital) have their bases attached to the broad dorsal surface of the basal phalanx of the second digit, which is grooved to receive them; the remainder (præ-digital) are attached to the second phalanx of the same digit. These last vary greatly in development; in fact, their variations constitute the most important structural differences of the wing. In most birds there are two: the proximal one well developed, the distal always rudimentary; but the former may show every degree of shortening, until it becomes quite rudimentary, or even altogether absent, as in Fringillidæ, and other "nine-primaried" birds, in which there are six metacarpal remiges, one ad-digital, two mid-digital, and no prædigitals, or only a very rudimentary one. The smaller feathers at the base of the quills, called upper and under coverts, have an equally regular arrangement. The webs or vanes of all the flight-feathers are made up of a series of parallel "barbs" which cohere together by means of minute hooklets, and so present a continuous, solid, resisting surface to the air.
Such is the characteristic structure of the wing in almost all carinate birds, whether powerfully developed for flight, as in the eagles, albatrosses, or swifts, or whether reduced in size and power to practically useless organs, as in the extinct great auk, the dodo and its kindred, weka rail, notornis, cnemiornis, etc., most of which, being inhabitants of islands containing no destructive land mammals, appear to have lost the principal inducement, and with it the power, to fly.
In the penguins (Speniscomorphæ) the feathery covering of the wing entirely departs from the normal type. Each feather is like a flattened scale frayed out at the edges, the barbs are non-coherent and have no hooklets. They form an imbricated covering of both surfaces of the wing, including the broad patagium which extends from the cubital side of the limb, but appear to have no definite relation to the bones, and can not be divided into distinct groups, corresponding to those described above. The structure of the wing separates the penguins sharply from all the other carinate birds.
The Ratitæ, or birds without keel to the sternum, form another very distinct group, distinguished by the rudimentary or imperfect ​condition of the remiges or quills, which never have coherent barbs, and are therefore unfitted to the purpose of flight. In the ostrich and rhea the bones, though comparatively small, are distinct and complete, and the feathers large and definitely arranged. The emu, cassowary, and apteryx show various degrees of degeneration, which apparently culminated in the dinornis, no trace of a wing-bone of which bird has ever been found. The question which naturally presents itself with regard to these birds is, whether they represent a stage through which all have passed before acquiring perfect wings, or whether they are descendants of birds which had once such wings, but which have become degraded by want of use. In the absence of paleontological evidence it is difficult to decide this point. The complete structure of the bony framework of the ostrich's wing, with its two distinct claws, rather points to its direct descent from the reptilian hand, without ever having passed through the stage of a flying organ. The function of locomotion being entirely performed by powerfully developed hind-legs, and the beak, mounted on the long, flexible neck, being sufficient for the offices commonly performed by hands, the fore-limbs appear to have degenerated or disappeared, just as the hind-limbs of the whales disappeared when their locomotory functions were transferred to the tail. This view is strengthened by the great light that has been thrown on the origin of the wings of the flying birds by the fortunate discovery of the Archæopteryx of the Solenhofen beds of Jurassic age, as in this most remarkable animal, half lizard and half bird, the process of modification from hand to perfect flying bird is clearly demonstrated. The three digits, which in the existing forms are more or less pressed together and imperfect, still retain their freedom and complete number of phalanges, and are each armed with terminal claws, while the flight-feathers and remiges of the cubital, metacarpal, and digital series are fully developed and evidently functional. The earlier stages in which the outer digits were still present, and the feathers imperfectly formed or merely altered scales, are not yet in evidence.
Some conception of the process by which a wing may have been formed may also be derived from the study of the growth of feathers on the feet of some domestic varieties of pigeons and poultry.