In all the wings which I have examined, whether in the insect, bat, or bird, the wing is recovered, flexed, or drawn towards the body by the action of elastic ligaments, these structures, by their mere contraction, causing the wing, when fully extended and presenting its maximum of surface, to resume its position of rest and plane of least resistance. The principal effort required in flight is, therefore, made during extension, and at the beginning of the down stroke. The elastic ligaments are variously formed, and the amount of contraction which they undergo is in all cases accurately adapted to the size and form of the wing, and the rapidity with which it is worked; the contraction being greatest in the short-winged and heavy-bodied insects and birds, and

FIG. 62.-Left wing of the albatross. d, e, f Anterior or thick margin of pinion. b, a, c Posterior or thin margin, composed of the primary (b), secondary (a), and tertiary (c) feathers. In this wing the first primary is the longest, the primary coverts and subcoverts being unusually long and strong. The secondary coverts and subcoverts occupy the body of the wing (e,d), and are so numerous as effectually to prevent any escape of air between them during the return or up stroke. This wing, which I have in my possession, measures over six feet in length.-Original.

least in the light-bodied and ample-winged ones, particularly such as skim or glide. The mechanical action of the elastic ligaments, I need scarcely remark, insures an additional period of repose to the wing at each stroke; and this is a point of some importance, as showing that the lengthened and laborious flights of insects and birds are not without their stated intervals of rest.

All wings are furnished at their roots with some form of universal joint which enables them to move not only in an

little shorter; and in the swallows this is still more the case, the first quill being the longest, the rest rapidly diminishing in length."- Macgillivray, Hist. Brit. Birds, vol. i. p. 82. "The hawks have been classed as noble or ignoble, according to the length and sharpness of their wings; and the falcons, or long-winged hawks, are distinguished from the short-winged ones by the second feather of the wing being either the longest or equal in length to the third, and by the nature of the stoop made in pursuit of their prey."Falconry in the British Isles, by F. H. Salvin and W. Brodrick. Lond. 1855, p. 28.

upward, downward, forward, or backward direction, but also at various intermediate degrees of obliquity. All wings obtain their leverage by presenting oblique surfaces to the air, the degree of obliquity gradually increasing in a direction from behind forwards and downwards during extension and the down stroke, and gradually decreasing in an opposite direction during flexion and the up stroke.

In the insect the oblique surfaces are due to the conformation of the shoulder-joint, this being furnished with a system of check-ligaments, and with horny prominences or stops, set, f

FIG. 63. The Lapwing, or Green Plover (Vanellus cristatus, Meyer), with one wing (c b, d' e' f') fully extended, and forming a long lever; the other (de f, c b) being in a flexed condition and forming a short lever. In the extended wing the anterior or thick margin (d' e' f') is directed upwards and forwards (vide arrow), the posterior or thin margin (c, b) downwards and backwards. The reverse of this happens during flexion, the anterior or thick margin (d, e, f) being directed downwards and forwards (vide arrow), the posterior or thin margin (c b) bearing the rowing-feathers upwards and backwards. The wings therefore twist in opposite directions during extension and flexion; and this is a point of the utmost importance in the action of all wings, as it enables the volant animal to rotate the wings on and off the air, and to present at one time (in extension) resisting, kite-like surfaces, and at another (in flexion) knife-like and comparatively non-resisting surfaces. It rarely happens in flight that the wing (d e f, c b) is so fully flexed as in the figure. As a consequence, the under surface of the wing is, as a rule, inclined upwards and forwards, even in flexion, so that it acts as a kite in extension and flexion, and during the up and down strokes.-Original.

as nearly as may be, at right angles to each other. The check-ligaments and horny prominences are so arranged that when the wing is made to vibrate, it is also made to rotate in the direction of its length, in the manner explained.

In the bat and bird the oblique surfaces are produced by the spiral configuration of the articular surfaces of the bones of

the wing, and by the rotation of the bones of the arm, forearm, and hand, upon their long axes. The reaction of the air also assists in the production of the oblique surfaces.

That the wing twists upon itself structurally, not only in the insect, but also in the bat and bird, any one may readily satisfy himself by a careful examination; and that it twists upon itself during its action I have had the most convincing and repeated proofs (figs. 64, 65, and 66). The twisting in question.

FIG. 64 shows left wing (a, b) of wasp in the act of twisting upon itself, the tip of the wing describing a figure-of-8 track (a, c, b). From nature.-Original. FIGS. 65 and 66 show right wing of blue-bottle fly rotating on its anterior margin, and twisting to form double or figure-of-8 curves (a b, c d). From nature.-Original.

is most marked in the posterior or thin margin of the wing, the anterior and thicker margin performing more the part of an axis. As a result of this arrangement, the anterior or thick margin cuts into the air quietly, and as it were by stealth, the posterior one producing on all occasions a violent commotion, especially perceptible if a flame be exposed behind the vibrating wing. Indeed, it is a matter for surprise that the spiral conformation of the pinion, and its spiral mode of action, should have eluded observation so long; and I shall be pardoned for dilating upon the subject when I state my conviction that it

forms the fundamental and distinguishing feature in flight, and must be taken into account by all who seek to solve this most involved and interesting problem by artificial means. The importance of the twisted configuration or screw-like form of the wing cannot be over-estimated. That this shape is intimately associated with flight is apparent from the fact that the rowing feathers of the wing of the bird are every one of them distinctly spiral in their nature; in fact, one entire rowing feather is equivalent morphologically and physiologically-to one entire insect wing. In the wing of the martin, where the bones of the pinion are short and in some respects rudimentary, the primary and secondary feathers are greatly developed, and banked up in such a manner that the wing as a whole presents the same curves as those displayed by the insect's wing, or by the wing of the eagle where the bones, muscles, and feathers have attained a maximum development. The conformation of the wing is such that it presents a waved appearance in every direction-the waves running longitudinally, transversely, and obliquely greater portion of the pinion may consequently be removed without materially affecting either its form or its functions. This is proved by making sections in various directions, and by finding, as has been already shown, that in some instances as much as two-thirds of the wing may be lopped off without visibly impairing the power of flight. The spiral nature of the pinion is most readily recognised when the wing is seen from behind and from beneath, and when it is foreshortened. It is also well marked in some of the long-winged oceanic birds when viewed from before (figs. 82 and 83, p. 158), and cannot escape detection under any circumstances, if sought for,—the wing being essentially composed of a congeries of curves, remarkable alike for their apparent simplicity and the subtlety of their detail.

The

The Wing during its action reverses its Planes, and describes a Figure-of-8 track in space. The twisting or rotating of the wing on its long axis is particularly observable during extension and flexion in the bat and bird, and likewise in the insect, especially the beetle, cockroach, and such as fold their wings during repose. In these in extreme flexion

the anterior or thick margin of the wing is directed downwards, and the posterior or thin one upwards. In the act of extension, the margins, in virtue of the wing rotating upon its long axis, reverse their positions, the anterior or thick margins describing a spiral course from below upwards, the posterior or thin margin describing a similar but opposite course from above downwards. These conditions, I need scarcely observe, are reversed during flexion. The movements of the margins during flexion and extension may be represented with a considerable degree of accuracy by a figure-of-8 laid horizontally.

In the bat and bird the wing, when it ascends and descends, describes a nearly vertical figure-of-8. In the insect, the wing, from the more oblique direction of the stroke,

FIGS. 67, 68, 69, and 70 show the area mapped out by the left wing of the wasp when the insect is fixed and the wing made to vibrate. These figures illustrate the various angles made by the wing as it hastens to and fro, how the wing reverses and reciprocates, and how it twists upon itself and describes a figure-of-8 track in space. Figs. 67. and 69 represent the forward or down stroke; figs. 68 and 70 the backward or up stroke. The terms forward and back stroke are here employed with reference to the head of the insect.-Original.

describes a nearly horizontal figure-of-8. In either case the wing reciprocates, and, as a rule, reverses its planes. The down and up strokes, as will be seen from this account, cross each other, as shown more particularly at figs. 67, 68, 69, and 70.

In the wasp the wing commences the down or forward stroke at a of figs. 67 and 69, and makes an angle of some