of the upper surface of the feather, thus revealing the hidden tints.
According to some authorities, however, some birds acquire a change of colour without a moult by the ascent of pigment from the base of the feather. The black head assumed by many gulls in the spring is, for example, said to be gained in this way. There is, however, not only no good evidence in support of the contention, but the whole structure of the feather is against the probability of any such change taking place.
Feathers correspond with the scales of reptiles rather than with the hairs of mammals, as is shown by their development. They make their first appearance in the developing chick at about the sixth day of incubation, in the shape of small papillae. In section each papilla is found to be made upThe development of feathers. of a cluster of dermal cells—that is to say, of cells of the deeper layer of the skin—capped by cells of the epidermis. These last form a single superficial layer of flattened cells—the epitrichium—overlaying the cells of the Malpighian layer, which are cylindrical in shape and rapidly increase to form several layers. As development proceeds the papillae assume a cone-shape with its apex directed backwards, while the base of this cone sinks down into the skin, or rather is carried down by the growth of the Malpighian cells, so that the cone is now sunk in a deep pit. Thereby these Malpighian cells become divided into two portions: (1) those taking part in the formation of the walls of the pit or “feather follicle,” and (2) those enclosed within the cone. These last surround the central mass or core formed by the dermis. This mass constitutes the nutritive pulp for the development of the growing feather, and is highly vascular. The cells of the Malpighian layer within the cone now become differentiated into three layers. (1) An inner, extremely thin, forming a delicate sheath for the pulp, and found in the fully developed feather in the form of a series of hollow, transparent caps enclosed within the calamus; (2) a thick layer which forms the feather itself; and (3) a thin layer which forms the investing sheath of the feather. It is this sheath which gives the curious spine-covered character to many nestling birds and birds in moult. As growth proceeds the cells of this middle layer arrange themselves in longitudinal rows to form the barbs, while the barbules are formed by a secondary splitting. At their bases these rudimentary barbs meet to form the calamus. Finally the tips of the barbs break through the investing sheath and the fully formed down-feather emerges.
A part of the pulp and Malpighian cells remains over after the complete growth of the down-feather, and from this succeeding generations of feathers are developed. The process of this development differs from that just outlined chiefly in this: that of the longitudinal rows which in the down-feather form the barbs, two on the dorsal and two on the ventral aspect of the interior of the cylinder become stronger than the rest, combining to form the main- and after-shaft respectively. The remainder of the rods form the barbs and barbules as in the down-feather.
The reproductive power of the feather follicle appears to be almost inexhaustible, since it is not diminished appreciably by age, nor restricted to definite moulting periods, as is shown by the cruel and now obsolete custom of plucking geese alive, no less than three times annually, for the sake of their feathers. The growth of the feathers is, however, certainly affected by the general health of the bird, mal-nutrition causing the appearance of peculiar transverse V-shaped grooves, at more or less regular intervals, along the whole length of the feather. These are known as “hunger-marks,” a name given by falconers, to whom this defect was well known.
It would seem that while the feather germ may be artificially stimulated to produce three successive generations of feathers within a year, it may, on the other hand, be induced artificially to maintain a continuous activity extending over long periods. That is to say, the normal quiescent period, and periodic moult, may be suspended, so that the feather maintains a steady and continuous growth till it attains a length of several feet. The only known instance of this kind is that furnished by a domesticated breed of jungle-fowl known as the “Japanese long-tailed fowls” or as “Yokohamas.” In this breed the upper tail coverts are in some way, as yet unknown to Europeans, induced to go on growing until they have attained a length of from 12 to 18 or even 20 ft.! In this abnormal growth the “hackles” of the lower part of the back also share, though they do not attain a similar length.
The feathers of birds are not uniformly distributed over the body, but grow only along certain definite tracts known as pterylae, leaving bare spaces or apteria. These pterylae differ considerably in their conformation in different groups of birds, and hence are of service in systematic ornithology.
The principal pterylae are as follows:—
(1) The head tract (pt. capitis), which embraces the head only.
(2) The spinal tract (pt. spinalis), which extends the whole length of the vertical column. It is one of the most variable in its modifications, especially in so far as the region from the base of the neck to the tail is concerned. In its simplest form it runs down the back in the form of a band of almost uniform width, but generally it expands considerably in the lumbar region, as in Passeres. Frequently it is divided into two portions; an upper, terminating in the region of the middle of the back in a fork, and a lower, which commences either as a fork, e.g. plover, barbet, or as a median band, e.g. swallow. Very commonly the dorsal region of this tract encloses a more or less extensive featherless space (apterion), e.g. swift, auk. While, as a rule, the dorsal region of this tract is relatively narrow, it is in some of great breadth, e.g. grebe, pigeon, coly.
(3) The ventral tract (pt. ventralis), which presents almost as many variations as the spinal tract.
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| Fig. 2.—Pterylosis of the plover. |
In its simplest form it runs from the throat backwards in the form of a median band as far as the base of the neck where it divides, sending a branch to each side of the breast. This branch commonly again divides into a short, broad outer branch which lodges the “flank” feathers, and a long, narrow, inner branch which runs backwards to join its fellow of the opposite side in front of the cloacal aperture. This branch lodges the abdominal feathers. The median space which divides the inner branches of the tract may be continued forwards as far as the middle of the neck, or even up to the throat, e.g. plover. Only in a few cases is the neck continuously covered by the fusion of the dorsal and ventral tracts, e.g. flamingo, Anseres, Ciconidae, Pygopodes.
For convenience sake the cervical portions of the spinal and ventral tracts are generally regarded as separate tracts, the pt. colli dorsalis and pt. colli ventralis respectively.
(4) The humeral tract (pt. humeralis), which gives rise to the “scapular” feathers.
(5) The femoral tract (pt. femoralis), which forms an oblique band across the thigh.
(6) The crural tract (pt. cruralis), which clothes the rest of the leg.
(7) The tail tract (pt. caudalis), including the tail feathers and their coverts; and
(8) The wing tract (pt. alaris). The wing tract presents many peculiar features. Each segment—arm, forearm and hand—bears feathers essential to flight, and these are divided into remiges, or “quill” feathers, and tectrices, or “coverts.”
The remiges of the arm, more commonly described as “tertiaries,” are, technically, collectively known as the parapteron and hypopteron, and are composed respectively of long, quill-like feathers forming a double series, the former arranged along the upper, and the latter along the lower aspect of the humerus. They serve to fill up the gap which, in long-winged birds, would otherwise occur during flight between the quill-feathers of the forearm and the body, a gap which would make flight impossible. In short-winged birds these two series are extremely reduced.
The remiges range in number from 16, as in humming-birds, to 48 as in the albatross, according, in short, to the length of the wing. But these numerical differences depend, in flying birds, rather upon the length of the forearm, since the quills of the hand never exceed 12 and never fall below 10, though the tenth may be reduced to a mere vestige.
The quills of the forearm are known as “secondaries,” those of the hand as “primaries.” The former are attached by their bases at relatively wide distances apart to the ulna, while the primaries are crowded close together and attached to the skeleton of the hand. The six or seven which rest upon the fused metacarpals II.-III. are known as “metacarpals.” The next succeeding feather is borne by the phalanx of digit III. and hence is known as the addigital. Phalanx i. of digit II. always supports two quills, the “middigitals,” while the remaining feathers—one or two—are borne by the last phalanx of digit II. and are known as pre-digitals, while the whole series of primaries are known as the metacarpo-digitals.
In their relation one to another the remiges, it must be noted, are always so placed that they overlap one another, the free edge of each, when the wing is seen from its upper surface, being turned towards the tip of the wing. Thus, in flight, the air passes through the wing as it is raised, while in the downstroke the feathers are forced together to form a homogeneous surface.
Birds which fly much have the outer primaries of great length, giving the wing a pointed shape, as in swifts, while in species which fly but little, or frequent thickets, the outer primaries are very short, giving the wing a rounded appearance. This adaptation to environment is commonly lost sight of by taxonomers, who not infrequently use the form of the wing as a factor in classification.
The tectrices, or covert feathers of the wing, are arranged in several series, decreasing in size from behind forwards. The number of rows on the dorsal aspect and the method of their overlap, afford characters of general importance in classification.
