.-. Frank M. Chapman FOR THE PEOPLE FOR EDVCATION FOR SCIENCE LIBRARY OF THE AMERICAN MUSEUM OF NATURAL HISTORY *? THE FLIGHT OF BIRDS THE FLIGHT OF BIRDS By F. W. HEADLEY, M.B.O.U. Author of " The Structure and Life of Birds " " Life and Evolution" Sec. WITH SIXTEEN PLATES AND MANY TEXT-FIGURES WITHERBY & CO. 326 HIGH HOLBORN LONDON 191 2 11 n I I «P LIBRARY OF THE AMERICAN MUSEUM Printed by WITHERBY & CO. at their Printing Press in Middle Row Place London IS- Mi>0<(.2fc-3w~O PREFACE. The flight of birds cannot but be of interest to men who sail the air on their biplanes or monoplanes, for the bird is, as yet at any rate, peerless among aviators, and in describing his methods I have kept in view the methods and the difficulties of those who are striving to rival him. It is possible, there- fore, that this little book may find readers among those who not only study flight but fly. But, since I have tried as far as possible to avoid technical terms and make it easily intelligible, I hope it may appeal to the larger class who, whether scientific ornithologists or not, take a delight in birds and their doings. I have to thank many friends for their help ; Mr. J. A. Tregelles for three drawings which can speak for themselves (the frontispiece and figures 13 and 14) and also for reading a typewritten copy of the book : Mr. C. W. Adams for reading the first five chapters : four Haileyburians past or present for drawings and a photograph ; W. T. Hichens for figures 18-23 ; G. G. Nanson for the diagrams ; G. H. G. Perry for figure 27 ; A. P. "Whitehead for Plate xrv, d. Mr. P. Disney has been kind enough to read the proofs. In addition I have to thank Mr. P. C. Gilson for some valuable criticisms, Mr. F. H. Jeffery for help in some small experiments : Messrs. Duckworth have very kindly allowed me to reproduce several of the illustrations of my Life and Evolution. The photographs (with the one exception I have mentioned) are my own. Most of them are now published for the first time, but several have appeared in Britisli Birds, and three in the Journal of the Aeronautical Society. The Editors of British Birds and the Secretary of the Aeronautical Society kindly permit me to use them. F. W. HEADLEY. Haileybury, March, 1912. CONTENTS. CHAPTER I. Gliding. pagb Resistance of Air — Lift and Drift — Curve of Wings — Area of Supporting Surface . . . . . . . . . . 1 CHAPTER II. Stability. Centre of Gravity — Points that make for Automatic Stability — Voluntary Adjustments . . . . . . . . 23 CHAPTER III. Motive Power. Leverage — Propulsion — Phases of the Wing-stroke . . 38 CHAPTER IV. Starting. The Preliminary Jump — Loss of Altitude between Wing- strokes — The Wing's Freedom to Rotate — Aeroplanes — Muscles — Big Birds and Small . . . . 48 CHAPTER V. Steering. A Variety of Methods — Good Steerers and Bad . . 59 CHAPTER VI. Stopping and Alighting . . . . 64 CHAPTER VII. The Machinery of Flight. The Breastbone and the Connected Bones — Muscles and Quality of Muscle — The Scaffolding of the Wing — Pneu- matic Bones — Stiffness of Wing — Expanse of Bone — The Spreading of the Wing — Structure of a Flight- Feather — Moulting — Legs . . . . . . 67 CHAPTER VIII. Varieties of Wing and of Flight. Curve — Narrow and Broad Wings — Styles of Flight — Flight in Flocks— The Whir of Wings 91 CHAPTER IX. Pace and Last. Experiments and Observations — Wind — Velocity of Migra- tory Flights — Endurance . . . . . . . . 103 CHAPTER X. Wind and Flight. Rising — Flight with the Wind — Undulating Flight without Movement of the Wings — Advance in a Direct Line without Movement of Wing — Advance Sideways in a Direct Line — Soaring — Soaring in a Horizontal Wind Impossible.. .. .. .. .. .. 120 CHAPTER XL Some Accessories. Digestion — Circulation, Breathing, Temperature — Repair of the Machine — Call-notes and Song . . . . . . 154 LIST OF PLATES. I. — Eagles soaring . . . . . . . . Frontispiece Facing page II. — Photographs of Pigeons and Gull, showing right and left wings giving different strokes . . . . 30 III. — Photographs of — A : Pigeon using tail for mainten- ance of equilibrium. B : Pigeon gliding. C : Gulls gliding 36 IV., V., and VI. — Photographs of Pigeons showing the phases of the wing-stroke in series . . 40, 42, 45 VII. — Photographs of Gulls flying . . . . . . 46 VIII. — A and B : Photographs of Pigeons starting to fly. C : of Pigeon rising with body steeply inclined. D : of Herring-Gulls starting . . . . . . 49 IX. — Photographs showing different methods of steer- ing — A and B : of Gulls. C and D : of Pigeons 60 X. — A and B : Photographs of Pigeons alighting. C : of Pigeon checking speed . . . . . . 64 XI. — A : Photograph of wing, showing elastic liga- ment. B : of flight-feathers and tail-feather . . 85 XII. — Drawing by I. C. Maclean (from Life and Evolu- tion), showing minute structure of flight -feather 86 XIII., XIV., XV.— Photographs of different types of wing 91, 92, 94 XVI. — Photographs of Gulls with motionless wings fol- lowing steamer . . . . . . . . . . 136 IX LIST OF FIGURES IN TEXT. Page 1. — Glass balls, an illustration of Sir I. Newton's experi- ment . . . . . . . . . . . . . . 3 2. — Diagram illustrating gliding . . . . . . . . 6 3. — Parallelogram of forces . . . . . . . . . . 7 4. — Diagram and drawing of boat tacking . . . . 8 5. — Illustrating kite-flying . . . . . . . . . . 9 6. — Showing the proportion of resistance to support . . 10 7. — Further illustrating the subject of resistance and support . . . . . . . . . . . . . . 11 8. — Planes set at different angles to the horizon . . . . 12 9. — Showing the advantage of a curved surface . . 16 10. — Two cubes 19 11. — Showing the disadvantage of an excessive curve .. 25 12. — Showing the shifting of the centre of pressure . . 26 13. — Swift gliding with wings fully extended . . . . 35 14. — House-Martin gliding with wings partly flexed . . 35 15. — Showing the velocity with which the extremity of the wing moves . . . . . . . . . . . . 39 16. — Showing how the downstroke propels as well as lifts 42 17.— (After Marey) Gulls flying 43 18. — Breastbones of Guillemot and Falcon . . . . . . 68 19. — Breastbone and connected bones of Adjutant . . 69 20.— Clavicles of Tern and Eagle 70 21. — Humerus of Eagle . . . . . . . . . . 71 22. — Humerus of (a) Skua ; (6) Hornbill ; (c) Sea-Eagle. Drawn to scale . . . . . . . . . . . . 78 23. — Skeleton of wing of Adjutant . . . . . . . . 80 24. — (After Alix) showing elastic ligament, anterior wing- membrane, eto . . . . . . . . . . . . 82 25. — Illustrating advance with motionless wings at right angles to the wind . . . . . . . . . . 131 26. — Illustrating advance in the teeth of the wind with wings held rigid . . . . . . . . . . 136 27. — Gull gliding sideways with wings held rigid . . 140 THE FLIGHT OF BIRDS. CHAPTER I. GLIDING. RESISTANCE OF AIR LIFT AND DRIFT CURVE OF WINGS AREA OF SUPPORTING SURFACE. Not long ago the sight of a sparrow on the wing, or even the sight of a lark rising in air and singing as he rose, excited but little interest or wonder in the mind of the average man. It came natural to birds to fly, and to fly so perfectly that they concealed their art. If they had had difficulty in flying, there would have been matter for astonish- ment. But why waste any wonder on the easy flight of a lark or a sparrow ? Such familiar things were taken for granted and seemed to call for no explanation. As soon, however, as men began to emulate birds — gliding downwards first from some elevated position and, later, by means of powerful engines, rising from the ground — then there was less disposition to take the bird's flight as a matter of course. The bird too, it was felt, had once been a beginner. He too had had difficult problems to solve and had solved them long ago, the very perfection of his methods making it difficult to 2 THE FLIGHT OF BIRDS understand fully what his methods are. But what- ever the difficulty of learning from him, the bird had certainly much to teach. Such a past master in the art of flying must needs be able to give some hints to the man whose ambition it was to discover the ideal design for an aeroplane. And so the question " How does a bird fly ? " became one that had an interest not only for ornithologists. Those who gave it any thought soon found that it involved a number of problems. First comes the question how the yielding air can support a heavy body when gravity is tugging it downwards. Then, of no less practical interest is the question, how the bird maintains his equi- librium, or recovers it if for the moment he happens to lose it. How does he move his wings so that they may at once propel and support him ? How does he steer ? How is it that the small bird is able to start so easily from the level ground ? What of the easier but important problem of alighting without jar ? How is it that the bird, big or small, is able to treat with contempt the sudden gusts and eddies that the boldest aviator fears ? Does the bird ever gain advantage from the unequal velocity of the wind ? Does he search for up-currents and get them to lift him ? Resistance of Air. The first of these problems was solved by Sir Isaac Newton. By means of an experiment, that may well astonish us if we bear in mind how imperfect were the appliances that he had at his disposal, he demonstrated the peculiar property of GLIDING 3 air that makes flight a possibility. The resistance of air to a moving body may be little, may be great. That it may sometimes be considerable many a bicyclist has found out to his cost when he has tried to double and redouble his speed. There comes at last a time when the " yielding air " almost refuses to yield. In fact the resistance it offers to a body moving through it increases as the square of the velocity. Of the ingenious experi- ment by which Newton proved this I must give a brief account. He took glass globes of equal size but unequal weights, corresponding to the figures 1, 4, 9, 16. These he let fall from the dome of St. Paul's and measured the velocities when they had settled down to a uniform pace. Since there was no gain or loss of velocity, the resistance of the air must have been in each case equal to the weight of the falling globe. But it turned out that the relative velocities corresponded to the figures 1, 2, 3, 4, whereas the weights of the globes are represented by the squares of these numbers, viz. 1, 4, 9, 16. From this he concluded that the resistance of the air increases as the square of the velocity. Recent experiments have shown that Newton's law is not absolutely accurate. Up B 2 4 THE FLIGHT OF BIRDS to 10 metres per second (about 22 miles per hour) the increase in resistance is rather less than the square of the velocity, Avhereas for velocities greater than 10 metres per second the resistance increases at a still more rapid rate.* But to resist, to give effective support, the air must be in proper condition. If it has just been pounded and battered, it is useless to trust to it. And so perpetual forward movement to ever fresh fields is necessary. A bird cannot mark time in the air. It is true that one often sees a Kestrel Hawk hovering, apparently without any forward motion, his wings, one might imagine, pounding the same air over and over again. But there is good reason to believe that the Kestrel never hovers except when there is a fair breeze to bring fresh unbattered air to his wings. If a long string be tied to a Pigeon's leg, he will fly perfectly well till he reaches the end of his tether. But as soon as he feels the pull of the string he will drop to the ground. When a flock of birds are travelling across the sky, it is easy to see that not one of them puts himself immediately behind any other. Were any individual to do so, he would not have at his disposal the fresh undilapidated columns of air that are essential, and moreover he would feel all the back- wash from the bird in front. At least one aviator has lost his life through getting into the wash of an aeroplane that was a little ahead of him. In building a biplane the question of finding air that has not been tumbled is one that cannot be neglected. A writer who appears to speak with authority says * Marey, Vol des Oiseaux, p; 218. GLIDING 5 that its two surfaces must have a space of four feet between them, or else they will interfere with one another ; one will tumble the other's air. It is true that I have seen a triplane with its " planes " at a distance from one another of three feet only. But whatever the minimum interspace, it must be considerable. And yet when birds are flying low over water they do not ruffle its surface. Pelicans flying over the Nile, not more than a foot from the water, so a good observer says, left the surface undisturbed.* Still, I cannot help thinking that the space between the birds and the water was under-estimated. At any rate a pigeon's first strong wing-strokes when he rises scatter straws, dust, and feathers from the ground where he takes off. Enough has now been said to show how effective is the support that the air can give to a rapidly- moving body, and how essential it is that the bird or the aeroplane should be unceasingly moving onward to columns of air that are fresh and undisturbed. Lift and Drift. It is probable that the power of flight was first attained by terrestrial birds which jumped from tree to tree, their wings aiding them to float through the air. If this floating through the air had been their crowning achievement, they would have done nothing remarkable, for mere gliding is not flight. Since, however, gliding seems to have been their first attainment, just as it has been with human flyers, I shall begin by investigating this compara- tively simple performance. At the outset there * Quoted by Marey, Vol des Oiseaux, p. 30. 6 THE FLIGHT OF BIRDS is a very elementary matter to be made clear. Let us picture the bird to ourselves as gliding in an exactly horizontal direction. The resistance of the air to his forward progress is equivalent to a horizontal wind blowing against him, and what is wanted is support, to counteract the downward pull of gravity. From a horizontal wind he must somehow get this support. The question presents no difficulty to anyone who has studied elementary mechanics, and in the application of mathematical principles the bird is a wonderful proficient. He inclines his aeroplane (his expanded wings and his W Fig. body) slightly upward, and the result is that the air supports him more than it resists him. It acts at right angles to the plane that he opposes to it. In fig. 2, b represents the gliding bird, w the rush of air against his expanded surface. If the air is still, there will, nevertheless, be the wind due to the bird's own velocity. W, at right angles to the bird, shows the direction in which the wind acts ; its lifting power, when the bird inclines himself thus, being greater than its resistance. The action of the air at right angles to a plane moving through it, is illustrated by an experiment sometimes made by very unscientific persons. A stone is thrown GLIDING 7 slanting- wise at a window — thrown by a dexterous, mischievous urchin standing far off beneath the wall of the house — and the fragments of glass as they go flying into the room make a right angle with the plane of the window. The air, then, striking against the plane presented by the bird acts at right angles to it, and there comes into play a resolution of one force into two. This introduces what is known as the parallelogram of forces. There is a force acting along a b (fig. 3), and if resistance is in this direction it may resolve itself C ^. Fig. 3. into two forces represented in magnitude and direc- tion by the lines c b, d b. Take the case of a boat tacking. The wind acts at right angles to the sail, but the boat refuses to move much in that direction (i.e. broadside on) ; she makes only a little leeway. The force of the wind, therefore, acting towards x (fig. 4), is broken up into two forces acting towards d and l, and that towards l, as I have said, does not count for much, since the boat will not readily move broadside on. So there is much headway and a little leeway. The principle at work is the same when a bird is gliding horizontally. His body is inclined slightly upward. The force of the air acts 8 THE FLIGHT OF BIRDS at right angles to the surface he presents to it. And this force is broken up into two, the one supporting him, the other tending to drive him backward, or, as it is now briefly and clearly put, the force is resolved into lift and drift. Fig. 4. Boat tacking. — w, the wind which acts at right angles to sl (the sail), towards x. The force is broken up into two forces, acting towards d and l. A paper kite supplies us with another and perhaps still more apt illustration. If the air is still, the kite-flyer may supply a force by running with the string ; if there is a wind, he has merely to hold the string. Let us imagine that the wind is blowing horizontally. It will, nevertheless, lift the kite, as long as the string is held firmly. It strikes against the oblique surface which the kite presents, and GLIDING 9 acts at right angles to it. Drift is prevented by the firmly-held string, and lift is the sole result (fig. 5). It is only when there is resistance that there is any force to be thus resolved into two. If the string breaks, gravity at once begins to pull the kite to the ground. We have now a further question to investigate and, to simplify it, we must consider not the gliding bird, but a flat plane set at a slight upward incline and driven horizontally through the air. A flat plane, having none of the curves and concavities of the bird's wing, is far inferior for purposes of flight, but its simplicity recommends it when the W Fig. 5. To Illustrate Kite-Flying. w, horizontal wind blowing against kite, k t. w' (at right angles to k t), line along which the force of the wind acts. object is to explain elementary principles. Set at an incline and moving horizontally it will tend to rise. The resistance of the air is equivalent to a wind blowing against it. The wind would act in 10 THE FLIGHT OF BIRDS a direction at right angles to the plane, and the force so acting is, of course, resolved into two, one tending to raise the plane, the other resisting its horizontal progress. The question, then, which we wish to decide is : In what proportion is the force of the wind divided between the two components, between lift and drift ? Now, supposing that b d (fig. 6) represents the plane set with an upward incline and driven horizontally through the air, it can be shown that line d c represents the resistance of the Fig. 6. air to its onward progress and b c, a much longer line, the support given by the air. In fact, when the plane is inclined but slightly upward the support it gets from the air is far greater than the resistance, a fact that can be proved by experiment. The mathematical proof that d c represents the drift (or resistance), and b c the lift, I give in fig. 7. It is now apparent that, as the angle of inclination to the horizon is more and more reduced, the pro- portion of lift to drift becomes greater and greater. Why not, then, reduce the angle till the resistance of the air to horizontal progress becomes a negli- gible quantity ? But obviously there is a limit to the process. If the plane has so slight an incline that it is almost horizontal, the air will offer but little resistance, and however big a proportion of this we may allot to lift and however small a one GLIDING 11 to drift, yet the actual amount of lift will be but small. If the strongest man among the survivors w Fig. 7. b d is the plane, w the wind blowing horizontally against it. b c is the cosine of the angle at b, d c the sine of the angle. Pro- duce c d to e, making d e = b c. From d draw d p at right angles to b d. From e draw e f at right angles to d e. (This decides the length of the line d f). Draw f g parallel to e d and d g parallel to f e. Let d f represent the force of the wind acting at right angles to b d. It can be resolved into two forces, f e and fg(=ed). e d we know= b c, and it can be shown that f e = d c. In the triangles d e f, b c d, b c = d e, and the angles at e and c are each right angles. If we could prove a second angle = a second angle, the triangles would be equal in every respect. Now the angles at d together = two right angles, and the angle b d f is a right angle. Therefore b d c with f d e makes one right angle. But d f e with f d e makes one right angle. There- fore angle d f e = angle b d c. The triangles then are equal, and the two sides f e and d e = respectively d c and b c. But f e represents drift and d e lift. Therefore d c, the sine of the angle at b, represents the drift, while b c, the cosine, represents the lift. of a starving ship's crew is able to take for himself nineteen-twentieths of the last biscuit, nevertheless he gets but a poor meal ; and it is obvious that if 12 THE FLIGHT OF BIRDS the plane were not inclined at all, but presented its edge to the air, there would be practically no support. There is, therefore, beyond all dispute a limit somewhere to the possible reduction in the incline of the aeroplane. The question is where, for practical purposes, that limit comes in. Newton formulated a law with regard to this, a law which is now quoted only to be condemned, and some- times quoted with expressions of contempt for him and mathematicians in general.* Newton held that the resistance of the air increases as the square of the sine of the angle of inclination. Fig. 8. Thus, if we take angles of 5°, 10°, 20°, the resistance would increase, from 25 to 100, to 400. Instead of being grateful to Newton for his great contribution to our understanding of flight, his discovery that the resistance of the air increases as the square of the velocity of bodies moving through it, some writers have depreciated him and his work because he has come to a wrong conclusion on this further question. As a matter of fact the resistance of the air varies as the angle, i.e. as its sine, not as the square of its sine. Therefore, as you diminish the angle, you still have a considerable amount of resistance, and it is divided up largely in favour * See Sir H. Maxim's Artificial and Natural Flight, pp. 2-G. The question is well dealt with in Prof. Langley's Experiments in Aero-dynamics ; see especially pp. 24 and 25. GLIDING 13 of support, of lift as opposed to drift. If Newton's law held good, if the angle of inclination were more and more reduced till it amounted only to, say, 5°, then the resistance offered by the air would be too small to be worth dividing up between lift and drift. Even the lion's share would be worth next to nothing. But experiment shows that an aeroplane set at an angle of 5° can, if it travels fast, find support in the air. But, although as we reduce the angle of inclina- tion the resistance of the air does not diminish at the rapid rate that Newton imagined, nevertheless there must obviously be a point beyond which the fining down of the angle cannot go, since the air will at last cease to give the required support. But before we reach the lowest possible limit another factor comes in which checks us as we are making successive reductions. As we continue to cut down the angle there comes at last a point at which the question of friction obtrudes itself in very unpleasant fashion. Imagine the aeroplane driven through the air at a very minute angle. If it is to find support, it must travel at a very great pace, else the resistance of the air will be too small. With every diminution of the angle there must be an increase of pace, and it might be thought that, if only the pace were increased sufficiently to make up for the diminution of the angle, all would go well. Professor Langley made some most valuable experiments which showed the great advantage of a small angle of inclination, and, emboldened by this great and important discovery, he proceeded to frame a formula and to speak 14 THE FLIGHT OF BIRDS of a law. By experiment he had discovered an indubitable principle, but subsequent experiments have discovered an equally indubitable fact, which at a certain point interferes with its operation. The angle cannot be reduced below, approximately, 5° without bad results. At a less angle, with the necessary increase of pace, the friction of the plane against the air increases so rapidly that, so far from there being any gain from the further reduction of the inclination, there is an actual loss.* On this subject a writer in Flight has some very interesting calculations, founded partly, it is true, on theory, and requiring further verification by experiment, but probably representing the facts without any considerable deviation. f Indeed experi- ment has already proved his main thesis, so that it is only detail that requires further testing. He imagines aeroplanes having the ideal camber or curve for their planes, a large curve if they are to travel slowly at a great angle of inclination, a slight curve if they are to travel fast. They are to carry a weight of 100 lb. Let us imagine them driven by 3 horse-power at the angles 30°, 25°, 20°, 15°, 10°, 5°. The velocity will increase rapidly as the angle is reduced. At 30° it will be 38 miles per hour. With the successive reductions of the angle it will increase to 47, 60, 78, 106, 134. The pace at 5° and even at 10° is greater than most people would wish to travel at, so the power applied might with advantage be reduced. Two horse-power at 10° ♦See The Aero Manual, p. 21 ; Flight, July 9, 1910; and Langley's Aero-dynamics, p. 37. t Loc. cit. GLIDING 15 and 5° respectively would mean 70 and 90 miles per hour. Even 1 horse-power, with an angle of 5° gives a velocity of 43 ; in other words, 1 horse- power, if the plane is set at an angle of 5°, is more effective than 3 horse-power when the angle is 30°. In fact, if you have a good aeroplane and the skill to use it well, you require less power to fly fast than to fly slow. But if you reduce the angle of inclina- tion below 5° you find that the tables are turned upon you. So far from economising power, you would have to use it in lavish style, to overcome friction. Curve of Wings. We must now turn to the question of curves. The curved surface is undoubtedly superior to the flat. A toy paper glider, being a mere feather- weight, can dispense with curves, but without curved surfaces — cleverly designed ones too — aviation would be out of the question. An aeroplane presents concave surfaces to the air ; its " planes " curve from front to back, and a bird's wings have concavities that are probably better adapted for flight than anything that human ingenuity has designed. On this subject even an umbrella can tell us something. When its ample concavity is turned towards the wind its efficiency in catching and holding the air is so great that it speedily becomes a wreck of wires. Lilienthal, a distin- guished flight pioneer, whose experiments in gliding did much to make aviation practicable, fully ap- preciated the value of the curved surface. Some- times when he was carrying his glider to a little 16 THE FLIGHT OF BIRDS hillock, his jumping-off place, outside Berlin, he was cheered and emboldened by the way the wind would catch his glider's well-designed concavities and nearly lift him from the ground. When he reached his little hillock he would jump from it and make glides of 150 yards and more. Lilienthal found that the air that met his glider's concave surface did not act at right angles to the chord of the arc (fig. 9), along f, but along a line occupying somewhat the position of f'. Fia. 9. The arrow shows the direction in which the glider is travelling. Experiments made by Mr. Wilbur Wright and his brother have shown that Lilienthal had called attention to a principle on which the aviator can base his calculations. In short, with a properly rounded surface the lift is greater and the drift is less than with a flat one. In order to obtain the maximum gain, the best possible curve must be discovered, and this can only be done by experiment. The writer in Flight already quoted is of opinion that each velocity has its appropriate curve. An aeroplane that is to be a racer should, he maintains, have its surfaces much less curved than a slowly- flying one, and undoubtedly the wings of birds suggest that he has enunciated a true principle. GLIDING 17 Very fast flyers, such as the Swift, have very little concavity. There is another fact, too, that points to the same conclusion : in a bird's wing the part that is near the body is the most concave, while towards the extremity it becomes nearly flat, in some cases quite flat. Now the extremity of the wing, when a stroke is given, moves with great velocity forward as well as downward, the near part comparatively slowly. Thus birds seem to be exponents of the principle that the curve should vary inversely as the speed. And here I must point out a great advantage the bird has over his upstart rival, the aeroplane. The aeroplane is rigid, with at most the rear edge only of its plane flexible. If the principle to which I have called attention is sound, it must be built either with a curve appropriate to great speed or with one which suits comparatively slow travelling. It cannot be varied during flight according as the pace is varied. But when the bird takes a very hard stroke, the front-to-back curve of his wings, which is mainly the curve of the feathers, is much reduced, so that by plying his wings with great vigour he to some extent modifies his configuration and adapts it to his high velocity. (For birds in rapid flight, see Pis. iv-vn.) It is always well to ask " Why is it so ? " when- ever experiment discovers a fact. Why, then, with a curved surface is the lift more and the drift less ? It is very difficult to see how the air does its work, and theory on such a subject may prove to be only random guesswork. But the lift would seem to be greater from the simple fact that the 18 THE FLIGHT OF BIRDS air cannot so easily escape. When the moving surface is a plane, the air is unconfined and the force is dissipated. The reduction of the drift (i.e. of the resistance to the forward movement of the surface) may be illustrated by the passage of a boat, built on good lines, through the water. The resistance of the water to the bow is balanced, or nearly so, by the shove given to the stern by the water closing behind it, so that there is little, except friction, to retard the boat. In the same way the air closes upon the hinder part of the curved surfaces of an aeroplane ; there is no region of " dead air " behind it. The matter is well explained in a little book on Model Flying Machines (pp. 19 and 20) by Mr. W. G. Aston. There is a region of " dead air " behind a flat surface when it is given an upward incline and is driven forward horizontally. The amount of " dead air " is the measure of the amount of resistance. Area of Supporting Surface. We can hardly leave the subject of gliding without touching on this important question : What area of supporting surface is required for, say, one pound weight ? Obviously no hard and fast rule can be laid down. Since it is the front part of the plane on which the wind mainly impinges, the back part is of less importance ; indeed it may be quite superfluous, a useless encumbrance, if the breadth from front to back is excessive. Besides this, as I shall show, we cannot frame a formula that will apply equally to big aeroplanes and small, to the big bird and the small bird. The Aero Manual GLIDING 19 says, " Nearly one square foot for each pound weight." But it has in view big machines only. When we go to the various creatures that fly, weigh them and measure their surfaces, we get most diverse results. The big flyers, we find, have small wings, the small flyers big ones, if difference in weight is allowed for. Compared with a gnat or a butterfly a Stork has a very small supporting surface, a small one even when he is compared with a Swallow. When M. de Lucy discovered these facts and pub- lished them the astonishment was great. Here are some of his figures : — Gnat Surface per lib. sq. yds. ft. ..4 6 weight, ins. 105 Butterfly . . Swallow . . .. 3 .. 8 4 87 18 Pigeon Stork .. .. 1 14 122* ^ / / Fig. 10. But before long mathematicians hit upon a plan by which they were able (or thought they were able) to rob the figures of their startling character, and * See Marey, Animal Mechanism, p. 222, and Pettigrew, Animal Locomotion, p. 133. C 2 20 THE FLIGHT OF BIRDS put big and small birds in this respect more or less on a level. They appealed to the elementary principles of geometry. If you take two cubes, a side of one of which is twice the length of the other, the larger one is in bulk eight times as great as the smaller one, but its surface area is only four times as great. This holds true of other figures of three dimensions that are not cubes. Magnify a bird till it is eight times its former bulk and you will only have multiplied its surface area by four. In order, then, to compare bird with bird correctly, you should take (so say these theorists), the cube root of its weight (for the weight is practically the bulk, i.e. three dimensions multiplied together) and the square root of its surface area (since that is two dimensions multiplied together). When we adopt this method we find the preponderance of the small bird in point of wing-area per pound weight not so very great. And we may, if we are so constituted, derive a certain comfort from feeling that we are following out geometrical principles. But these principles have, it must be owned, in the case of some species been widely departed from. What are we to make of the legs and neck of the Flamingo ? If any small bird of ordinary build were symmetrically enlarged, should we ever arrive at elongations so enormous ? It is, of course, true that if the bulk of a small bird were multiplied many times, and the area (not the bulk) of his wings increased in proportion, he would have a greater expanse of wing than his muscles could possibly work. If the Stork had a wing-area as great in proportion to his weight as the Swallow, then GLIDING 21 (taking his weight as four pounds and four-fifths) his wings would together measure twenty square feet ! This would be a monstrous acreage of wing to raise and lower. But when we have pronounced it monstrous, we have still to answer the question why it is that the big flyer requires, in proportion to his weight, a comparatively very small support- ing surface ? Let us imagine the Swallow supplied with wing- area at no more liberal rate per pound weight than that at which the Stork is supplied. Then, taking the Swallow's weight to be about five-sevenths of an ounce, he would have five square inches of wing-surface — two and a half on either side — a miserably poor allowance. A wing so small would be largely made up of margin, and the air would escape at the edges. The gnat has over four and a half square yards of wing for one pound weight. His actual allowance for his almost imponderable insignificance is considerable, but if we make pro- vision for him at the rate at which the Stork is supplied, his wing-surface becomes a mere point, "without parts and without magnitude," to quote Euclid's familiar definition. The air would offer no resistance to so near an approximation to the theoretical point. Here, no doubt, we are getting at the main fact that explains the comparatively small size (when weight is allowed for) of the wings of the great flyers. Since the wing-surface is much larger absolutely the air does not escape so easily at the margins ; each square inch is more effective, since there is less waste. Later on I shall make a further comparison of big birds and small, but 22 THE FLIGHT OF BIRDS one important difference and its explanation have, I hope, been made clear (see Chap. iv). It will be more convenient to consider the question of upward and downward gliding in the next chapter. CHAPTER II. STABILITY. CENTEE OF GRAVITY — POINTS THAT MAKE FOR AUTOMATIC STABILITY VOLUNTARY ADJUSTMENTS . There are two things that we must carefully distinguish — stability and the maintenance of equi- librium. An aeroplane, when it has once completely lost its balance, cannot recover it, though some swaying or pitching can, no doubt, be corrected. The stability of a bird is a very different thing from the aviator's careful maintenance of equilibrium. A very strong and sudden gust may throw the bird on his side, or even on his back, and yet he will very quickly right himself. Centre of Gravity. But we want to know whether a bird in ordinary flight, when there are no very sudden gusts, is auto- matically stable, or whether he has to be perpetually making small adjustments. It is sometimes main- tained that a bird need take no trouble about the question of balance, since his centre of gravity is low down. The great flight muscles, which are massed upon the breast, form a great part of his weight. I have weighed the three pairs of breast muscles of two Wood-Pigeons and those of two 24 THE FLIGHT OF BIRDS domestic Pigeons, and found that they account for about one-fifth, or even more than one-fifth, of the weight of the whole bird. Indeed, in one Wood- Pigeon they were equal to three thirteenths of the total weight — a little less than a quarter.* Hence the centre of gravity lies considerably below, though not so very far below, the shoulder joints. But the idea that it is good to have the centre of gravity of a flying machine low down is altogether a miscon- ception, a fact that can easily be put beyond all doubt by experiments with small gliders. Weight, placed low, tends to make the machine oscillate and even swing right round. It is best that it should be on a level with the supporting surface. In the case of the bird, the chief supporting surfaces, the wings, are always changing their relative position as they are raised or lowered, and, of course, as they move they to some extent raise or lower the centre of gravity. This must make voluntary adjustments for the purpose of recovering balance still more necessary. But Professor Marey has pointed out a curious fact which may make these oscillations less difficult for the bird to deal with. When the down-stroke takes place and the wings are lowered, the centre of gravity occupies a lower position in the bird, but the bird as a whole, except during very rapid flight, rises. With the up-stroke, on the other hand, there is a raising of the centre of gravity, but a lowering of the bird ; hence, though the bird's * Legal and Reichel, in the Jahresberichte der Schlesischen Gesellschaft (1879), give ¥ .|^ ( = i^§ = more nearly | than A) as the proportion of the total weight of the pigeon accounted for by the three pairs of breast muscles; But this would seem to be a mistake: Charadrius, according to them, comes next with f. STABILITY, 25 body rises and falls, its centre of gravity travels onward almost in a straight line.* Points that make for Automatic Stability. Though the position of the centre of gravity is of no avail, the general build of the bird and the elasticity of the feathers make for automatic stabil- ity. To begin with, there is the question of curves. I have already pointed out that a curved surface gives more lift and less drift than a flat one. But anyone who experiments with gliders soon finds out that an excessive curve is fatal to stability. In the preface to the Aero Manual (1910) it is stated that the curve (the depth of the concavity) should not be more than one-twelfth of the breadth. Imagine a surface with a more considerable curve. The wind W — — ^ Fig. 11. Diagram to show the disadvantage of an excessive curve. will impinge upon its upper side and the glider will duck and descend rapidly to earth. But if the curve is only slight, though the tendency to duck and dive may, no doubt, arise, yet it tends to correct itself. Imagine the glider launched on its way and develop- ing a tendency to lower its head and raise its tail unduly. In proportion as the tendency develops, in * There is no need to discuss the question of pendulum stability. It is possible to suspend a heavy weight from an aeroplane, and there are theorists who hold that automatic stability may be thus attained. It is obvious that in the case of a bird nothing of the nature of a pendulum is possible. See Flight, Feb. 25, March 18, and March 25, 1911. 26 THE FLIGHT OF BIRDS proportion as the glider makes a smaller and smaller angle with the horizon, the point at which the air acts will be progressively nearer to the front margin, and this will obviously tend to prevent a dive downward. Fig. 12 Diagrams showing the shifting of the centre of pressure. This point requires some explanation. Take as an illustration a sailing boat when it is tacking. When it sails close to the wind, the air which strikes against the forepart of the sail is deflected from its course, and rushing towards the stern of the boat forms a buffer which shields the rest of the sail from the air, which would otherwise have impinged upon it. The drawing of a boat tacking, on page 8, and the accompanying diagram, will make this clear. The principle is called the law of Avanzini. The smaller the angle made by the wind with the sail, the nearer the point of impact approaches to the front edge. This is true no less of the slightly curved surfaces of aeroplanes. As the angle of incline to the horizontal is more and more reduced, the wind acts at a point closer and closer to the front edge, and thus the aeroplane may, possibly, correct the dangerous tendency automatically, without the aviator having to make readjustments. To illustrate this principle I have made experi- ments with a catapult which, working within runners, threw pieces of cardboard horizontally, STABILITY 27 the cardboard being set so as to have an upward incline ; in fact as an aeroplane is set when it is travelling horizontally. This was managed by means of small wire carriers having various inclines, on which the cardboard rested, and which were themselves thrown with the cardboard. So far from pitching head downwards, the cardboard missiles would, even when the angle of deflection from the horizon was small, rise in the air, and some- times even turn over backwards, so strong was the action of the air on the front margin. In the bird's wing there is a further automatic safeguard. It curves downwards at the back, at any rate that part of it that is nearer to the body. The wind acting over-strongly upon the downward- curving back part of the wing might, if the whole wing-surface were rigid, capsize the bird and send him diving head-foremost downward. But the elasticity of the feathers prevents an excessive lift. They yield to pressure, and the reduction of their curvature relieves the wing of any excess of pressure on its hinder part. The same principle holds good with regard to lateral stability. If a strong gust or eddy of air strikes the right wing while the left is struck with less violence, the feathers of the right wing yield to the rush of air and bend upward, so that the very force of the gust to some extent reduces its effect. Such elasticity would seem to be impossible in the case of an aeroplane without dangerously reducing its strength. And yet an aeroplane, from its enormous breadth, is far more liable than a bird to suffer from gusts falling with unequal force on its 28 THE FLIGHT OF BIRDS two extremities. At present the aviator is at a great disadvantage as compared with a bird. The great expanse of his planes is in itself a danger, and his machine has less power of automatic adjust- ment. Moreover, the most experienced pilot is a mere novice compared with a bird that has flown many times every day since he left the nest. Still, birds as flyers have already reached their zenith; for aviators greater things are still possible. Leaving the possibilities or impossibilities of the future, I must return to the bird as he is, and call attention to another point which promotes stability. The extremities of the wings of the best flyers have very little downward curve. If they are travelling very fast through the air, the primary wing-feathers may even bend slightly upwards, and aviators have found that the upward incline of their " planes " reduces the amount of rolling. Thus inclined they give less support than if curved downwards at the extremities, but the very fact that the air is allowed to escape easily to right and left is favourable to equilibrium. If an attempt is made by bending the extremities downward to check its escape, to coop it up, it may make a sudden rush from the right or from the left con- cavity and cause risk of a capsize. The wing, amply concave as it is till it begins to taper, and shallowing as it tapers, keeps in view the questions of lift and equilibrium ; it holds the air, yet provides for its escape. Sometimes when a bird wishes to glide downward he will point his wings steeply upward — a method often adopted by pigeons — much more steeply than the right and left surfaces STABILITY 29 of a monoplane. This attitude must tend to ensure a comfortable equilibrium (PI. in, b). The question of the escape of air from beneath the curved surfaces seems to be of the utmost importance. Mr. Pilcher's celebrated glider was punctured with many small holes which were intended to encourage a steady escape instead of a sudden upsetting rush on one side. I cannot judge whether this was a good plan for dealing with the difficulty. At any rate, the wing of even a large bird presents too small a surface to be treated in this way. But it is probable that the notches between the feathers at the wing's posterior margin tend to prevent irregular escapes of air from below. The great soaring birds, whose steadiness as they circle with motionless wings is so marvellous, have the great flight-feathers parted like outspread fingers. Moreover, the force of the wind sometimes bends them conspicuously upward, thus giving them an incline that is recognized as favourable to equilibrium. Voluntary Adjustments. It cannot be gainsaid that there is much in the bird's build that makes for automatic stability. Nevertheless, since the wind, at low levels at any rate, is a chartered libertine, full of capricious un- expected eddies, such automatic adjustments are altogether inadequate. The bird must be ready at a moment's notice to give his mind to the question of balance, and make conscious voluntary adjust- ments. But even we ourselves find that movements we are perpetually making tend, through force of habit, to become automatic, and that in some cases 30 THE FLIGHT OF BIRDS we are not conscious of any difference between the voluntary and the involuntary. In the case of a bird the line is still harder to draw, not only because we cannot get within his mind, but because the movements which we are bound to label voluntary, since they cannot be mere unconscious reflexes, are so largely instinctive. When the young Swallow takes the first plunge from the parental nest and trusts himself to the air, he finds at once that he can fly ; the power of flight is instinctive. A child has to learn by much practice to co-ordinate his muscles ; no other young creature is so devoid of instinctive skill. The young Swallow, though he can co-ordinate his muscular activities well enough to fly with some success, has, of course, much to learn. At the outset he misses most of the flies and gnats, and his parents have to come alongside and put their captures in his mouth. But he seems by instinct to spread his tail when it should be spread, and, no doubt, though it is hard to see this, he takes a harder stroke with one wing than the other, when a harder stroke is required. For fore-and-aft balance he depends largely on his tail. He has not a long neck that he can bend or straighten out, and his legs are so short and light that no movement of them can have much effect. For lateral stability he must, as far as voluntary adjustments are needed, depend mainly on unequal wing-strokes.* That birds do take unequal strokes has been clearly proved by the camera in the case of Pigeons, Owls, Gannets and some others (see PI. n). This inequality, sometimes so conspicuous in a photograph, the human eye has * See Chap. V. PLATE II. Right and left wings giving different strokes. A and B : Pigeons. C : Young Gull ; the right wing only is flexed at the wrist. (See Chap. II.) To face p. 30.] STABILITY 31 great difficulty in detecting, and but for the camera we should have had to own that, though the bird could hardly dispense with this method of balancing and steering, we could get no absolutely conclu- sive optical evidence. For fore-and-aft balance the means employed vary very much according to the build of the particular bird. Long-necked birds can move their centre of gravity forward and backward by extending or bending their necks. Ducks and Geese usually carry theirs stretched out, Herons habitually make a crook in theirs. Long necks are usually correlated with long legs, and long legs are no less serviceable in the matter of balance. This is shown by the fact that the long-legged have usually small, short tails, the long legs to some extent taking the place of the tail as balancers. A Flamingo, to take a conspicuous example, having legs of enormous length and a neck to match, has no need of much tail to regulate his fore-and-aft balance. A very little extension or retraction of legs or neck will set matters right if they are going wrong. But not only do long legs make a great expanse of tail unnecessary for balancing purposes, they must in- evitably hamper its movements if it is to be pulled downward with a view to checking speed or steering to right or left. It may well be that this necessary inefficiency of tail is in part the cause of the compara- tively clumsy steering of these big, heavy-legged birds. It can be shown, too, that for purposes of balance webbed feet very probably play a part of some importance. Not only are they fairly heavy, but they can be used as the tail is used, though not so 32 THE FLIGHT OF BIRDS effectively. The following table brings out very clearly the poverty of the long-legged and the web- footed in point of tail. I give first the actual length of tail, then the approximate weight of the bird. In two cases I have had to take the weight from a learned paper by two German ornithologists, and I suspect that their hen Sparrow-Hawk was far from plump. Lastly I have given the length of tail which each of the birds has for one pound of its weight. The results are very striking. Were I able to give the area of the tails, they would be much Bird. Length of tail. Weight of bird. Length of tail for each lb. weight of bird. Notes. Sparrow-Hawk (hen) Sparro w-Hawk ( cock ) Ins. 8 6-5 5-3 oz. (Legal & Reichel)* 5-2 oz. Ins. 24- 1 20 Tail very broad. Lapwing 41 6-7 oz. (Legal & Reichel)* 9-8 — Wood-Pigeon 6 1 lb. 4-25 oz. 4-7 Tail very- broad. Common Wild-Duck 4-75 lib. 15-7 oz. 23 Tail pointed, small in area, weak. Common Heron 6-5 3 1b. 0-6 oz. 21 — Curlew 462 2 lb 4 oz. 2-05 — * Jahresbericht der Schlesischen Gesellschaft, 1879. STABILITY 33 more so. But the area varies so much, according as they are expanded or not, that I have found it very difficult to give measurements. However, we can bear in mind that the tail of the Sparrow- Hawk, for example, is not only very long but very broad, whereas the tail of the Duck is not only short, but narrow and weak, its inefficiency being even more marked than its small size. It is noteworthy that the bird which stands at the head of the list as being remarkable for size of tail is not a particularly short-legged bird. Two cock Sparrow-Hawks, of which I took measurements, had each legs 7 inches long (the toes being included). However, the legs are very thin, as if not intended for standing, but, armed as they are with long, efficient claws, for seizing a victim. It is often maintained that Hawks hang their legs down during flight, but this is certainly not usually the case. However, they could on occasion be lowered, to give the tail freer play, without much affecting the bird's equilibrium. The main function of the tail is to prevent loss of equilibrium, and when large it plays its part wonderfully well. In spite of its great expanse, its weight is a negligible quantity and its working is splendidly prompt. Evolution has done wonders in thus metamorphosing the long, heavy tail of the bird's reptilian ancestor. A number of vertebrae have been compressed to form the pygostyle, the small bony base of this wonderful piece of machinery. What weight there is is almost all accounted for by this little bone and the muscles that move the large spread of feathers. The muscles have the 34 THE FLIGHT OF BIRDS tail completely under command. At a moment's notice they spread it, make it concave, lower it or raise it, lower this side or that. If the tail is spread and lowered, at once the hinder part of the body is lifted (see Pis. in and ix). The opposite effect will follow from the raising of the tail, so that it may catch the wind but little. If a Lark be watched through a field-glass as he rises facing a fair breeze, he will be seen to be perpetually busy with the work of correcting his fore-and-aft balance ; and it is to his tail that he trusts. A Chaffinch, perched on a rail with a fairly high wind blowing in his face, keeps his tail perpetually at work. The tail is, in fact, a fine balancer, even more important in this capacity than it is as a rudder. Were it not for the perfection of this balancer, a small bird could hardly land without danger to his eyes in the chevaux de frise of a furze bush. There are other balancing movements still to be considered. From our present point of view, the way the wings are held is of great importance. I have already pointed out that when a boat is tacking (see fig. 4, p. 8) it is the front part of the sail that does most of the work, and that similarly when the gliding bird inclines his wings at a small angle to the horizon, it is the front margin that gives him most support. If, then, he holds his wings fully expanded, so as to have as wide-stretching a front as possible, not only will the lift be greater but the centre of pressure (the point at which we may consider the force of the wind as being, so to speak, focussed) will move forward, and this will tend to give the bird's body an upward incline. STABILITY 35 And so, if he wishes to maintain his level or rise, he must extend his wings to the utmost; if, on the other hand, he wishes to descend rapidly, a partial flexing is the measure to adopt. To the aviator such a method is only to a very limited Fig. 13. Swift, gliding with wings fully expanded. Fig. 14. House-Martin, gliding with wings partly flexed. extent possible ; he cannot half-flex his planes. Nevertheless, he is able to slide down a steep incline, much as a bird does. By means of his fore-and- aft balancing apparatus, he sets his aeroplane at the proper angle and shuts off steam. If he is D 2 36 THE FLIGHT OF BIRDS a skilled and experienced pilot, he is, apparently, as steady as he rushes down towards the earth as a bird could be. But, if he did lose his balance, it would be fatal, whereas to the bird a momentary loss of equilibrium is of no consequence. Leaving aviators and their splendid achievements, I must describe another attitude sometimes adopted by a bird in gliding downward. He will extend his wings to the full, but hold them slanting upward (see PI. in). Obviously in this position the wings give less support, and so he descends. But he descends slowly, not with the rush that is character- istic of the head-foremost downward glide. The wings do not travel edgeways through the air and so they check his pace. Their upward slant is, as I have before remarked, advantageous to balance. I must now conclude this brief investigation of the bird's stability when on the wing. What we see in the flight of birds — I am not now speaking of soaring — is not a steady, careful maintenance of equilibrium, but an instantaneous recovery of balance whenever it is lost. The bird can afford to be indifferent to the difficult problems which this subject presents. He has something much better than the power of maintaining equilibrium. However the gusts and vagaries of the wind may upset him, he can right himself at once. He owes his wonderful stability to some extent to his fine build and the elasticity of his feathers, but mainly to manoeuvres and adjustments that cannot be mere reflexes. The flying machine which he pilots is admirably built : still it can never dispense with a pilot. But his voluntary adjustments are largely PLATE ill A : Pigeon using tail for maintenance of equilibrium. (See Chap. II. and Plates IV., V., VI. and IX.). B : Pigeon gliding. C : Gulls gliding : (See Chap. II. and Chap. III.) _To face p. 36.] STABILITY 37 instinctive, and even the niceties of adjustment that he has to learn must become through habit almost automatic. And the result of it all — of his fine build, his instinct, and his art — is the perfect stability of the machine ; a thing to make the most skilful aviator envious : for who ever heard of a bird losing his balance and falling to the ground ? There are, of course, trials to which he may prove unequal. He may lose his way in a cloud, become exhausted when crossing a wide stretch of sea, or even fly stupidly into a telegraph wire. But an accident from want of balance would be evidence of morbid condition. CHAPTER III. MOTIVE POWER. LEVERAGE — PROPULSION — PHASES OF THE WING-STROKE. In describing gliding we have taken the motive power for granted, assuming that the bird already has momentum. We must now investigate his method of lifting and propelling himself. The lifting is the hard work, for the bird is so shaped that when he sets himself at a suitable angle the air offers but little resistance to his movement onward in a hori- zontal direction. To lift himself he must put great force into the downward beat of his wings, making their extremities move with such velocity that the air, the resistance of which increases as the square of the velocity, will very soon offer effective support. They will, in fact, become levers, each having its fulcrum mainly near the extremity, the weight to be lifted being, of course, the bird's body, or, more correctly, the whole bird. The power is applied quite close to the body (see fig. 21, Chap. vn). The depressor muscle, the great muscle that springs from the breastbone and covers it with its great expanse, attaches to the humerus (or upper- arm bone) close to the nearer end. With a lever like this there is no economy of power — very far from it. What is gained is rapidity of movement. A quick movement MOTIVE POWER 39 of the near part of the wing will produce a movement of astonishing rapidity towards the further end, near the tips of the great primary- feathers. If x (see the figure) moves half an inch, x will move two inches. Preparatory to the down-stroke, the wing is lifted till it points straight upward, its anterior margin being turned in the direction of the bird's flight. During the first part of its descent it X Diagram to show the velocity with which the extremity of the wing moves. cuts edgeways through the air. But soon it turns face downwards and, the air opposing its descent, it finds a fulcrum. But, of course, there is some give. The fulcrum that the air supplies to the wing is, like the fulcrum that the oar finds in water, an imperfect one. When, however, the bird is taking full-length strokes, the wings appear to move with a far longer sweep than is really the case. With each stroke the 40 THE FLIGHT OF BIRDS body rises, and this means a relative lowering of the wings. It is a wonderful thing that the air can supply a tolerably firm support, something that will do duty as a fixed point. Archimedes undertook to lift anything if he could find a fulcrum for his lever. The bird finds a fulcrum where Archimedes would not have thought it worth while to look for one. The bird when flying is, in fact, taking a number of jumps. Often when he appears to be travelling in a horizontal line he is really, as Professor Marey has shown in his wonderful photographs that give successive phases of the process, rising and sinking with each down-stroke and up-stroke. Thus the bird's apparently horizontal line of progress is often an undulating one. But when he is travelling with great velocity, then there is no drop between the strokes ; of this Professor Marey has obtained evidence. But there is probably some reduction of pace. Even though there is no rise with each stroke, the bird is nevertheless taking a series of jumps. And the marvel of these jumps, with no better take-off than the air, no amount of thinking can do away with. A man who is accounted a good high jumper can do very little if he has a poor take- off — if the ground is spongy. We all find it very hard work walking over soft snow when at each step we sink up to our knees before we find anything firm and resistant beneath our feet. We walk slowly and with labour along a beach where the small pebbles let our feet sink in. We climb with effort up a volcanic cone where, each step that we take, the small rounded ashes let us slide downward PLATE IV Phases of the wing-stroke : photographs of Pigeons. A : Ready for the down-stroke. The series is continued on Plates V. and VI. (See Chap. III.) To face p. 40. ] MOTIVE POWER 41 til] we lose almost all that we have gained. But the bird has to deal with a material that seems far more shifty and ^independable than small pebbles or volcanic ashes, or than snow at its worst. He over- comes the difficulty by means of levers calculated to give the utmost rapidity of movement. The muscular effort required is great, but his muscles are strong, and it is long before they tire. Propulsion. — Phases of the Wing-stroke. But the bird has not only to lift himself, or to maintain the altitude he has already gained. He must also have onward momentum. Were this wanting, he could not even lift himself, for air has little or no supporting power when it has just been disturbed. He must, therefore, be perpetually advancing to fresh columns of air that have not yet been shattered by the beating of his wings. I have already pointed out that, when birds are flying in flocks, each takes care to keep clear of the backwash of the bird in front of him — takes care to avoid tracts of air that have already been disturbed ; that a Pigeon, when he has a string tied to his leg, cannot maintain himself in air, however wildly he may ply his wings, when he has reached the end of his tether ; that when a Kestrel hovers without advancing there is always a breeze, so that each wing-beat descends on fresh, unbattered air. For ordinary flight the wings must be so adjusted as to propel as well as lift. This the bird can effect only if the front part of the wing is lower than the hinder part. Thus the parallelogram of forces comes once more to his aid. Let f b (fig. 16) represent a section through the 42 THE FLIGHT OF BIRDS bird's wings, f being the front and b the back margin. The resistant air (w in the figure) will be equivalent to a wind blowing vertically upward. It will act at right angles to the plane of the wing, and the line representing its action will point not only upward but forward. During horizontal flight the front edge of the wing is slightly at a lower level than the back. But when the bird is rising and taking very Fig. 16. Diagram showing the effect of the lowering of the front margin of the wing. energetic strokes, then the required incline is not obtained by that method only. The wing moves forward as it descends, so that at the end of the down-stroke, instead of making a right angle with the body, it points more forward than outward. When the wing is in this position, there is the down- ward slope that is wanted, from its base at the shoulder to the tip. One method passes gradually into the other ; indeed, the wing is inevitably shoved forward when the air lifts its hinder margin. On PLATE V. B 1 £> Phases of the wing-stroke, continued from Plate IV. A : Last phase of the down-stroke. B : Beginning of the upstroke. C : Feathers bent upward during the up-stroke {see Marey, Vol des Oiseaux, p. 268, and Plate at end). D : Up-stroke continued. Continuation of series on Plate VI. {See Chap. III.) To face p. 42.] MOTIVE POWER 43 the forward-downward movement as giving the required incline I wish to lay stress, because treatises on flight do not, as far as I know, recognize this way of obtaining the downward slope, but speak only of the depressing of the front margin relatively to the back (see Pis. iv and v). Fig. 17. Gulls flying (after Marey). A. — 25 photographs per second. B. — 50 per second. I am now going to describe more in detail the different phases of the stroke. For this I must depend to some extent on Professor Marey's photo- graphs. Other photographers, myself among them, taking snapshots, have caught the various wing positions. He shows us the various phases in series : they follow one another at the rate of 25 or of 50 per second (see fig. 17 and Pis. iv, v, vi, vn). 44 THE FLIGHT OF BIRDS I will first give some account of a long and com- plete stroke. Preparatory to the down-stroke, the wing is raised till it points vertically upward, its front margin being turned in the direction of the bird's flight. There may then be a moment's pause, the wing, as it were, resting before it strikes its blow. In the case of gulls the next move seems to be a slight bend at the wrist- joint. After this begins the serious work. The wing descends with lightning speed, so fast indeed that this early phase of the down- stroke does not always appear in the series when photo- graphy has succeeded in depicting all the other phases. The great rapidity at this stage seems to indicate that it is not till the wing is approaching the horizontal that it begins to feel the resistance of the air and do its work of lifting and propelling. When this work is going on, the upward bending of the primary- feathers leaves us in no doubt about the fact. As the wing descends it points more and more forward. The way in which this is brought about is highly interesting. The big muscle which lowers the wing attaches to the front part of the lower face of the humerus (upper-arm bone) (see. Chap, vn, fig. 21). Its pull, therefore, tends to lower the front of the wing relatively to the hinder part by rotating the bone. But the air, acting on the feathers that spread out rearward, greatly aids the muscle, lifts the hinder part of the wing, and encourages the rota- tion. But the work of the air does not end here. As soon as the wing has an upward incline from front to back, it cannot but move forward ; the mere action of the air on a surface so inclined cannot but bring this about. Thus the upward incline from PLATE VI. Phases of the wing-stroke, con t inure I from Plate V. A : Nearly the same as Plate V., and from a dinVrenl point of view. B, C, D : Up- stroke continued. (See Chap. III.) [To face p. 45. MOTIVE POWER 45 front to back, without which the bird would not make headway, is obtained first by the raising of the back part of the wing relatively to the front, and later, as the stroke advances, by the forward movement of the descending wing, which brings it about that the extremity occupies a position lower than and in advance of the base. For a moment let us consider the working of the wings in combi- nation. With the body they form a kind of funnel — obviously one side of the funnel is missing, but this is unimportant. Caught in this funnel and deflected from the wing- surfaces, the air impinges upon the body and lifts it. When the wing has strained forward and downward till it can strain no further, the muscles at length relax. The wing is no longer rigidly extended, but slightly bent at the elbow-joint, and soon at the wrist also. If the bird is rising and has little onward momentum, the Elevator muscle does the work of lifting. The great flight- feathers, which during the down-stroke have been pressed close against one another and so have made the wing impervious to air, are now slightly rotated, so that interspaces are left which allow the air to pass, and thus the raising is effected without much opposition (see Chap, vn, fig. 24). If, on the other hand, the bird has much way on, the air itself effects the lifting and little work on the part of the muscles is required. After the strain of the down- stroke, the Depressor muscle ceases its contraction, and, perhaps, the Elevator gets to work. In any case the front margin of the wing is no longer de- pressed relatively to the hinder margin, but is lifted. The wing lets the wind have its way, and is carried 46 THE FLIGHT OF BIRDS back, almost unresisting, but occasionally, as photo- graphs show, it resists enough to cause a backward bending of the flight- feathers. It looks as if the bird, when his wings are being lifted by the rush of air into position for a fresh stroke, checked them for a fraction of a second in order to save himself from loss of altitude. It is possible this check in the course of the up -stroke may be the normal thing, or it may be only occasional (see PI. v). When the wing is thus raised by the rush of air — more strictly by the resistance of the air to its momentum — the flight- feathers are pressed against one another and there are no gaps to make the lifting easy. But no such help is required. The wing is blown back as far as it has freedom to go, and at the end of its rearward movement it is no longer facing as it was ; its under- surface is facing outward and its anterior margin is looking towards the bird's head. When the wing is in this position it is easy to raise it com- pletely and bring it forward ; in fact it moves edge- ways. Anatomy supplies very remarkable evidence that the raising of the wing requires little effort. Whereas the muscle which lowers the wing is red, ridged, and granulated, the Elevator muscle is paler and exposes a smooth surface when it is cut. The former is by far the better class of muscle, capable of long, unflagging effort. In ordinary horizontal flight most birds take a much shorter stroke than the one I have just described, nor is the wing pointed much forward (see PI. vn ). The upward incline from front to back that is needed is obtained by a slight lowering of the front margin relatively to the back, mainly PLATE VII. Gulls flying with short stroke. A : Lesser Black-backed Gulls. B : Black-headed Gulls ; the lower bird on the left with wings bent at the wrist, shows an early phase of the up stroke. (See Chap. III.) MOTIVE POWER 47 towards the extremity of the wing, and this gives propulsion enough. This short stroke is the one adopted by gulls in their ordinary leisurely flight. The wings give the air a sharp slap, and this with a bird so well built for flight and so skilled is very effective. It is to be noticed that, when gulls are taking short, leisurely strokes, the wings during the down- strokes are very distinctly curved from their base to their extremity. This curve prevents a too easy slipping away of the air and so increases the wing's lifting power ; but when a stronger stroke is taken the primary- feathers bend upward. The question how a bird lifts and propels himself I have now briefly answered. In other chapters I shall try to describe the build of this living flying- machine, to my thinking the noblest of all craft that sail the air (see Chaps, vn, vm, and xi). CHAPTER IV. STARTING. THE PRELIMINARY JUMP LOSS OF ALTITUDE BETWEEN WING- STROKES — THE WING'S FREEDOM TO ROTATE AEROPLANES MUSCLES BIG BIRDS AND SMALL. The Preliminary Jump. I have shown how a bird when flying maintains and propels himself, when we have given him in imagination a good start. We have now to study his method of starting. Let us suppose that he is standing on the ground ; if he is to fly he must somehow get clear of it. For this purpose his long, strong legs are of great service to him ; it is owing to them that he is so good a starter. Such is the bird that I think of as typical. Not only are his legs strong, but they are long, unless we compare them with those of a Wader or a Heron, birds that for their particular mode of life have developed an inordinate length of leg. The Puffin and the Swift, by their helplessness on level ground, call attention to the remarkable leg- power of the ordinary small bird, or bird of moderate size. The Swift and the Puffin cannot take the preliminary jump with which flight must begin. Bulky, short-legged birds all rise with difficulty ; the jump that should in a moment obtain for them freedom of wing action is beyond their power. PLATE VIII. A and B : Pigeons starting to fly. B : Avery hurried start. C : Pigeon rising with body inclined steeply upward. D : Herring-Gulls starting ; photographed in September, when they were moulting. (See Chap. TV.) STARTING 49 It is not that they are deficient in lifting power, for it is said that an Eagle can carry a weight as great as its own, and I have seen a Falcon flying off with a victim that was not far short of itself in point of size. Whether big or small, birds all fold their wings neatly upon their backs, where they cannot possibly interfere with freedom of leg action. In order to appreciate the excellence of this arrangement, we must compare the bird with the bat or with the pterodactyle, whose wings were remarkably bat-like. Extending, as they did, far back and attaching to his legs, they must have been as bad an encumbrance as long skirts. Though, no doubt, a fine flyer in a bat-like style when once launched on his way, the pterodactyle was a poor starter. The bird's legs, on the contrary, are not sacrificed to his wings (see PI. viii). Unless he is one of the specially bad starters, he jumps lightly into the air and is off. With regard to the heavy, lumbering way in which many big birds rise, a great deal is to be learnt by watching a Heron start to fly from level ground. Unlike the Condor, the Comorant, the Puffin, and the Swift, he has no difficulty in getting under way. True, he does not rise with a steep incline as an ordinary small bird or a Pigeon can, but a gradual ascent he carries out without difficulty. He drops from the top of his long legs on to the spacious fields of air and is at once clear of the earth, with ample room for the plying of his wings. On the other hand, the big short-legged birds and small birds like the Swift, with very short, feeble legs, have no room for a full sweep of their wings, 50 THE FLIGHT OF BIRDS unless they have, by some means other than a jump, attained some elevation. Hence the difficulty they have in starting from level ground. But even when this full sweep is possible big birds are still only capable of ascending with a gentle incline, and this is a fact that we must try to account for. Loss of Altitude between Wing-Strokes. To my thinking the explanation is this : since the big bird necessarily takes a longer stroke and requires more time to raise his wings for a fresh downward beat, he must inevitably lose more altitude between wing-strokes than the small bird. Professor Marey gives the rate of stroke for a number of birds as registered by means of scientific apparatus.* Strokes per second Sparrow .. 13 Duck 9 Pigeon 8 Marsh-Harrier .. 5| Screech-Owl . . 5 Buzzard . . 3 In the case of a Sparrow it is evident that there is no time for a drop between the strokes, though compared with those of most insects even a Sparrow's wings move slowly and heavily. The leisurely working of a Heron's wings is familiar to everyone who ever takes the trouble to observe birds ; usually he flaps along with only 130 strokes a minute or even slightly less. The Marsh-Harrier * See his Vol des Oiseaux, p. 100 k STARTING 51 has a much quicker beat, but even with such a rate as his a drop between the strokes is quite possible unless he has plenty of way on. And now we are getting to the explanation of the big bird's method of rising. In order to avoid losing altitude between the strokes, he must take care that he has momentum, and if he is to have momentum he must be content to ascend by a gentle incline. The Wing's Freedom to Rotate. Moreover, there is a want of freedom about his wing-movements which makes him incapable of anything but a very gradual ascent. If he were to incline his body steeply upward, after the manner of a small woodland bird that, making for a gap in the dense spreading boughs overhead, mounts almost vertically, he would have to rotate his wings in a way that is impossible for him ; he would have to lower the front margin relatively to the back, or else they would beat in such a way as to drive him backward instead of lifting him. In fact, he has too little freedom at the shoulder. He cannot set his wings as a steep ascent requires. The small bird's wings, on the other hand, rotate so freely that even when he sets his body with a steep upward slant he can still turn them over so that they have an up-and-down beat and raise him sky- ward. But though, speaking generally, the small bird is capable of a steeper ascent than the big bird, yet it would be a great mistake to imagine that if we were to arrange birds according to their weights, from the K 2 52 THE FLIGHT OF BIRDS lightest to the heaviest, we should at the same time be arranging them according to their angles of ascent — the angle which each makes with the horizon when the line of his ascent is as near to the vertical as his build and his powers allow. In the case of all very big, bulky birds, I believe the wing rotates reluctantly and with difficulty at the shoulder. The Gannet, Pelican, Cormorant, Eagle I have tested, and found that they have very little power of lowering the front edge of the wing relatively to the back ; one Eagle was a partial exception. I have never had a live or a freshly-killed Condor at my disposal, but there is reason to believe that the Condor is among the stiffest of the stiff. The way in which Condors are trapped in Chile — it is described by Darwin in his Journal of Researches (Chap, ix) — supplies indirect evidence of the great bird's limitations. The plan is to place a carcass "ona level piece of ground within an enclosure of sticks with an opening, and, when the Condors are gorged, to gallop up on horse- back to the entrance, and thus enclose them : for when this bird has not space to run it cannot give its body sufficient momentum to rise from the ground." In fact a Condor cannot rotate his wings and set them as they must be set if his line of ascent is to make a large angle with the horizon. As far as I know, all birds of great bulk have this defect, but when we come to birds of medium size we find great variations, and it soon becomes apparent that it is largely a question of habitat and environ- ment. The Pheasant and Duck — I have tested the Mallard, the Sheld-duck and the Teal as representa- tive of the Ducks — have great freedom of movement STARTING 53 at the shoulder, and can, moreover, point their wings forward so that at the finish of the down-stroke they have an upward incline from tip to base. On the other hand, the Partridge and the Herring-Gull have very little power of rotation. The striking contrast between the Partridge and the Pheasant throws a great deal of light upon the question. The Pheasant is a denizen of woods and has often to make for an opening in the branches that shows itself almost directly over his head. Heading straight for it, he points his body almost vertically upward, but, in spite of that, his wings have an up-and-down beat, and turn the concavity of their nether side towards the ground and their upper convex surface towards the sky. The Partridge, on the other hand, frequents plains and open fields where there are no entanglements to make a nearly vertical ascent even an occasional necessity. A Wild Duck sometimes finds it expe- dient to mount upward from her nest among the bushes in the same style as the Pheasant just described, and I have seen a Wigeon, without any apparent advantage to himself, shoot up thus from a large piece of open water. All birds, if we except those that frequent only open water, bare cliffs, bare hills or unwooded plains, may find, any moment, that they have to make a rapid ascent up a steep incline ; life itself may depend upon it. Hence great freedom at the shoulder is very common. I have observed it not only in the Pheasant and the Duck, but in the Jackdaw, Crow, Raven, Chough, Jay, Magpie, and Quail. No wing, I think, rotates more freely than that of the domestic Pigeon. Were it not for this, the bird in PL viii could not achieve, as he is 54 THE FLIGHT OF BIRDS evidently doing, the feat of an almost vertical ascent. The big and bulky birds then, such as the Gannet and the Condor, having a much slower wing-stroke than the light-weights, must make it their first object to attain momentum, otherwise they will lose altitude between the strokes. When they have got way on, they can rise, but the line of their ascent is a gentle slope. For a steeper ascent the set of their wings unfits them. But there are birds of medium size, such as the Duck and the Pheasant, which, striking with very great rapidity and rotating their wings as freely as any small bird, are capable of raising themselves almost vertically through the air. Aeroplanes. In a country like England, with trees and hedges almost everywhere, an aeroplane capable of a steep ascent is a great desideratum. Captain Brooke- Popham, writing on military aviation, says that " with no wind a fully-loaded machine, with observer, could get off a hard level field in a length of 120 yards and clear a fair hunting hedge at the end. Our Air Battalion " Farman " can do this in 90 to 100 yards without any difficulty."* Thus even the " Farman " requires a great deal more space than a Condor or a Comorant, birds noted as slow, lumbering starters, and though aeroplanes are constantly being improved it may well be doubted whether a steep ascent will ever be achieved. * The Army Review, January, 1912, p. 89. STARTING 55 Muscles. To rise in air before he has got up pace is hard work for any bird unless he can get the wind to help him, for a great strain is put upon the muscle that lifts the wing. And here I may call attention to the remarkable development of the Elevator muscle in the Pheasant. Its weight amounts to nearly one-third of that of the Depressor. It is very pale and has little lasting power, but for a brief effort it is very effective. Its development in the Duck is considerable, but not equal to what is found in the Pheasant. As soon as the bird begins to travel rapidly, there comes an easier time for the overtaxed Elevator, for the resistance of the air to the onward impetus is sufficient to lift the wings, and the impetus is due mainly to the work of the great Depressor muscle. Hence the Depressor not only lowers the wing, but indirectly lifts it. The Elevator is not fitted for long-sustained effort. The Depressor is a redder, rougher, more granulated muscle, and its different colour and texture are indicative of superior quality.* Big Birds and Small. As a rule, when a small bird flies, his line of flight is undulating. For him to rise is easy ; a few strong, rapid strokes lift him. He then partly flexes his wings and glides onward and slightly downward. This is very noticeable in the case of the Woodpecker. He flexes his wings more than most birds, and so the dipping character of his flight * For more on this subject, see p. 45 and Chap, vii : Muscles. 56 THE FLIGHT OF BIRDS is more conspicuous. The big bird, on the other hand, dare not lose altitude in so reckless a way, since when he has once lost it he with difficulty recovers it. He ploughs steadily on, whereas the small bird with a few rapid strokes gains altitude, then, like a bicyclist utilising his free wheel, glides restfully and rapidly onward, not minding the loss of some of the height he had gained. His stroke is more rapid, but he is able to take frequent easies.* I have now compared and contrasted big birds and small, but even the biggest birds that fly are not very big ; as compared with the larger mammals they are diminutive. Why, among all the birds that fly, are there none that weigh even half as much as, for example, a Zebra ? Helmholtz, dealing with this subject, produced a formula which, backed as it was by the authority of a great man, was too readily accepted, regardless of the fact that it was not founded on data obtained by experiment. There is nothing so misleading as mathematics when the premises are unsound. Helmholtz started with undeniable facts. If a bird's linear dimensions be multiplied by 4, then the area is multiplied by 16 (4 2 ), and the bulk, which must nearly correspond to the weight, by 64 (4 3 ). So far good. He showed that the weight increased more rapidly than the supporting area. But when he went on to maintain that the power required to lift the bird increased at a still more rapid rate — that, in the case I have taken, it would be 128 (4i) — then he was building a theory without a proper foundation * On big birds and small birds see Chap, i, pp. 18-22. STARTING 57 of proved fact. Even now, though the subject has been much studied, it would be very rash to venture on a formula. The big bird has great advantages. He can manage, as we have seen, with a relatively smaller expanse of wing, for the area, being greater absolutely, does not so readily allow the escape of air at the margins. Moreover, his wing, being longer absolutely, is a more powerful lever. The great weights which Eagles, for example, carry show that there is no deficiency of lifting power. But the big bird's wings have not the easiness of rotation at the shoulder- joint which makes it possible for a Green- finch, for instance, or a Pheasant, to rise with a steep incline ; he cannot put himself in the right attitude. It does not appear, however, that bulk in itself is any handicap. If this be so, it may well be asked why even big birds are quite small when compared with the larger mammals. I have already pointed out that the big bird, if his legs are short, has difficulty in beginning a flight, and so lacks a very important accomplish- ment. Then why are they not all mounted on stilts, like the Flamingo ? But legs of such length, or even the half of it, would for many birds be most incon- venient appendages, for a diving bird most of all. They would not help him to rise from the water, and they would be clumsy things beneath the surface. And thus among big birds there are many that for purposes of flight are handicapped by shortness of leg. Diminutive size brings with it another advantage. The small bird comes more rapidly to maturity. In his second spring, when he is not yet twelve months old, the Blackbird has already paired 58 THE FLIGHT OF BIRDS and is busy nesting. The Gannet is known, I believe, not to nest till his fifth year ; the Eagle comes to maturity later, possibly not till his tenth year. In rate of reproduction, therefore, a most important matter for the species, the small bird has a great advantage. And thus we can account for the small size of birds compared with that of mam- mals without recourse to any such theory as that of Helmholtz. The small bird is a better starter than the big, and he comes to maturity more quickly. CHAPTER V. STEERING. A VARIETY OF METHODS GOOD STEERERS AND BAD. A Variety of Methods. A bird can steer when his tail is gone. A Rook, when some accident has robbed him of this useful rudder and balancer, can still make shift. He is not like a ship left rudderless. It is evident, therefore, that the tail is not the bird's sole steering apparatus. If he wishes to steer to the left his usual method is to fling himself on his left side, the left wing pointing downward and the right wing upward, the two being in line with one another, while his head is pointed in the direction in which he wishes to travel (see PL ix). Then he can no longer progress along his former line of advance, for the expanse of his wings will check him. He travels, therefore, to the left, i.e. towards the point towards which his head is directed. But how does he effect the necessary change of balance ? There is no doubt that he occasionally gives a harder stroke with one wing than the other, a thing which the camera sometimes detects, though it is difficult for the eye to see it clearly. On Plate n are some photographs in which the wings have been caught 60 THE FLIGHT OF BIRDS at a moment when they were not held symmetrically. Another plan is to bend sideways at the waist, so as to move the centre of gravity towards one side. I was long doubtful whether this plan was actually adopted. At length I have obtained a photograph of a Gull, taken from below, where the waist, thus bent, is clearly shown, but unfortunately this photograph will not stand reproduction. Below I give some measurements which show that birds that are good steerers have greater suppleness of waist than clumsy steerers. The tail also is undoubtedly used for steering, though, I believe, it is more frequently of service in maintaining or restoring equilibrium. The rule is, as I have shown, that if the legs are long the tail is small, and as it is indisputable that long legs are very useful for balancing and of but little use in steering, we may infer that the tail is more a balancer than a rudder. Nevertheless, it is of much use in steering. Though its edges look to right and left, it can be made an effective rudder by the lowering of one side more than the other. Web-footed birds probably use their feet occa- sionally for steering purposes, but I doubt if their action counts for very much. When a Duck, for example, is hurtling through the air, if he lowers one foot, the resistance of the air will double it up, not expand it. It is very different when a Duck in swimming kicks backward ; the water acts upon the under-surface and spreads the webs to their fullest extent. Some birds when alighting use their feet in the same way. A Gannet, for in- stance, kicks hard in order to correct his balance PLATE IX. A and B : Gulls steering by throwing themselves on one side. A : Steering to the right and looking to the left. C and D : Pigeons showing the tail used as a rudder. (See Chap. V.) To fare p. 60. STEERING 61 and get into the right attitude for settling. It is a clumsy performance. The movement of the head to one side or the other has, no doubt, some slight influence on the balance. But the skull is very light, a great part of it as thin as paper, and one may occasionally see a Gull look to the left while he steers to the right (PI. ix ), just as a skater, who is quite at home on his skates, can make such minor movements without in any way upsetting his balance. This subject naturally brings to mind the wonderful way in which Mr. Cody showed the lateral stability of his biplane. He carried a passenger who stood at a distance of ten and a half feet from the centre where he himself was piloting. When a bird turns his head, it is far more probable that he is directing his eyes, to right or left, towards some object that has attracted his attention, than that he is regulating his balance. Mr. Bentley Beetham has a capital photograph* in which he has caught five Gulls in the act of turning their heads to the right. This may possibly be due, as he suggests, to a local current of air with which they all equally have to cope. But it seems far more likely that they have all suddenly caught sight of some object of interest, a fish or shoal of fish, at no great distance. What would one not give for a photograph of a large flock of birds scudding through the air high aloft — for a photograph taken at the moment when by some common impulse, as if following some leader, they each and all change their course ? Might we not see, if we could obtain such a photograph, * British Birds, Dec, 1910. 62 THE FLIGHT OF BIRDS all the heads turned one way, the object being to follow with the eye the movements of the leader or leaders of the flock ? Good Steerers and Bad. There are good steerers and bad steerers. Contrast the Swallow, the Lapwing, or the Sparrow- Hawk, those adepts at sudden swerving and doubling, with the Duck ! The Duck becomes the slave of his own ponderous momentum, and changes his course slowly and with effort. The Swallow, the Lapwing and the Sparrow-Hawk have all of them a fine expanse of tail, while the Duck's is small in area and, what there is of it, feeble. Efficiency or inefficiency of tail, no doubt, accounts for a great deal. But it is not the only factor to be considered. The Duck, though very strong, is lacking in agility ; the three other birds have far more suppleness, I believe, notably at one im- portant point. Though birds' backbones are, below the neck, very stiff, yet they allow of a good deal of bend, either up and down or sideways, at the waist. I have measured the amount of sideways bend in some few species and found that, as far as my far from complete evidence goes, good steerers Angle formed by backbone bending sideways at waist. Kestrel 141 5° Swallow 150° Swift 153-5° Common Tern . . . . 155° Kestrel (another specimen) . . 156° Domestic Duck . . . . 165° STEERING 63 have supple waists and poor steerers comparatively stiff ones, the Duck being a good deal the stiffest of those I have examined. It is difficult to account for the difference between the two Kestrels, but there is no doubt that the Duck is, of the birds in question, decidedly the stiffest. In fact these measurements make it a priori probable that suppleness of waist counts for a good deal. The result of the sideways bend must be that the bird is thrown on to its side — the attitude which is assumed when a sudden turn is to be made. For steering purposes, then, a bird has various methods at his disposal. He can take unequal wing-strokes or bend at the waist. Either of these means will put him in the attitude in which we see him steering to right or left, one wing pointing downward, the other upward. And he can use his tail as a rudder. Some birds may, no doubt, steer partly by means of their feet, but I doubt whether a foot is a very effective rudder. The Duck's webbed feet do not seem to make him an adept at turning. CHAPTER VI. STOPPING AND ALIGHTING. A bird alights without any jar. Let us imagine that when we first catch sight of him he is flying at some height above the ground. Wishing to descend, he will give his wings an upward slope and float down in the style usually preferred by Pigeons, or more probably he will slant his body, from tail to head, downwards and, partly flexing his wings, glide rapidly towards the earth. On nearing his landing place he will suddenly let his hind- quarters sink, and give his body an upward incline ; his wings, spread wide, present their whole under-surface to the air. This soon checks his momentum. But if he wants to make a very sudden stop he gives a stroke with his wings, and this, when the body is nearly upright, as a glance at one of the photographs {see PI. x) that illustrate the process will show, must bring him at once to a standstill. And just after landing, perhaps (notably if he is a Tern, a Lapwing or a Pigeon) he will raise his wings high above his head as if to stretch and refresh the muscles, in which attitude he looks very beautiful. A beautiful thing, too, is the folding of the wings ; it is all so quickly and so neatly done. A Tern or even a Pigeon when he alights has all the grace of the " Herald Mercury, new lighted on a heaven-kissing hill." PLATE X. A and B : Pigeons alighting. Though in B the feet are still clear of the window-ledge, I think the bird is giving his wings the final stretch that with many species is pre- liminary to the folding of them. C : Pigeon checking speed before alighting. (See Chap. VI.) To face p. 64. J STOPPING AND ALIGHTING 65 As birds are so careful in alighting to avoid all jar, it is remarkable that some species have habits which would seem likely to cause concussion of the brain. It is astonishing that the Nuthatch, for example, when he has been half the day hammering at nuts with tremendous vigour, yet suffers no bad conse- quences. But we must bear in mind that the bird's neck, with its peculiar saddle-and-rider vertebrae, is more supple than a snake, whereas the backbone, except just at the waist, is remarkable for its rigidity. Hence, probably, the care and dexterity in alighting that contrast so strikingly with the reckless use of the beak and head as a hammer by the Nuthatch, the Woodpecker, and other birds. It must be owned that there are some birds which, when they are alighting, are very unlike the "Herald Mercury." Among these we must count the Gannet. When he gets near his nest upon some cliff, he paddles hard with his legs, and at last settles clumsily down. Mr. Bentley Beetham (British Birds, May, 1911) has some very good illustrations of the Gannet 's style of alighting. One of them shows a bird that has so lost command of his movements, that he is flopping most ungracefully onto his nest. His breast is resting on the pile of dry seaweed, his outspread wings on the rock at either side. It is an astonishing attitude, but it must be borne in mind that the Gannet is in the habit of taking headers from a great height, and that his breast is shielded by air cavities beneath the skin, first-rate air-cushions that greatly reduce the shock when he dashes into the water. This may account for his descending breast foremost onto his nest. But the Cormorant, who has no 66 THE FLIGHT OF BIRDS air-cushions, though he is the Gannet's near relative, is also clumsy in alighting, and Mr. H. F. Witherby has described to me how Cormorants, wishing to alight upon a post, will sometimes make a bad shot, pass it, and have to turn and try again. A Gull will alight gracefully enough on the water, but when he is aiming at a particular spot on a rock, for example, he will often paddle as awkwardly as a Gannet. The Lapwing is in a very different category ; he is noted for his power of suddenly stopping, of making every possible turn. When you get near his nest he has a way of flying at you in threatening style, then suddenly checking himself and making off. He is a perfect master of bluff. A pair of Lapwings, by such menacing swoops and turns, will drive off a Crow whom they suspect of looking for their nest. In order to stop suddenly, what is wanted is a great expanse of feather, and I cannot help associating the Lapwing's remarkable wings — they are so strikingly broad at the extremity — with his well- known shock tactics. I have already remarked on the skill which small birds show in alighting on their perch. Were a Linnet not very clever at making all the required adjustments, she would seldom reach her nest, some- where deep in a gorse bush, without accident. CHAPTER VII. THE MACHINERY OF FLIGHT. THE BREASTBONE AND THE CONNECTED BONES MUSCLES AND QUALITY OF MUSCLE THE SCAFFOLDING OF THE WING PNEUMATIC BONES STDTFNESS OF WING EXPANSE OF BONE THE SPREADING OF THE WING STRUCTURE OF A FLIGHT- FEATHER — MOULTING — LEGS. When we carve a Partridge or a Grouse, since other questions seem at the time of more pressing interest, we seldom give a thought to the fact that we are slicing and disjointing what has been a most mar- vellous flying machine. Yet so it is. When we carve the breast, we first cut through the Great Pectoral, the powerful muscle that lowers the wing. Lying below it we find, easily distinguishable, a much smaller, lighter-coloured muscle — in the Grouse and its kin markedly lighter. This smaller muscle lies in the angle between the keel and the sternum (or breastbone) proper ; its work is the lifting of the wing. The Breastbone and the Connected Bones. The framework, the bony skeleton, is wonderfully adapted to the purpose to which it is put. It is most important, to begin with, that there should be a wide expanse of bone from which the great flight muscles may spring. And so the area of the sternum is increased by the large projecting keel. F 2 68 THE FLIGHT OF BIRDS. The Ostrich has no keel, and is incapable of flight. The Hoatzin, that curious South American bird, according to all observers a most feeble flyer, has the front part of the keel missing. In every case the shape of the keel has its significance and is well worth observing, since it indicates the style of flight. The Duck and the Guillemot, having very long breastbones and keels, have of course very long flight muscles and a long muscle is capable of more Fig. 18. Breastbones of (1) Guillemot — actual length of keel 5k inches. (2) Falcon. Drawn to scale. Compare that of the Adjutant, a Stork famed for his soaring. contraction than a short one. A glance at their breastbones, therefore, tells us that they are birds that fly with a long stroke. A Guillemot's wings are so small that a long, strong stroke is a necessity for him. A Duck, during horizontal flight, does not raise his wings very high, but the stroke is pro- longed till the wings can strain no further downward. THE MACHINERY OF FLIGHT 69 The Tern and the Falcon, on the other hand, have short, deep keels, and therefore short but big and strong flight muscles. Accordingly, their ordinary stroke is short but powerful. It is highly important also that there should be a firm pivot on which the wing may rest as it moves. During part of the wing's down-stroke there must be considerable Breastbone and connected bones of Adjutant — actual length of keel 4| inches. CI. : Clavicle ; Co. : Coracoid ; Sc. : Scapula. pressure inward, and were the skeleton to give, were it to supply only an unsteady wobbling pivot, vigorous flight would be out of the question. How is the pivot formed ? There is a strong bone, the coracoid, which springs from the front part of the breastbone and points forward and outward. It 70 THE FLIGHT OF BIRDS slopes more outward in strong than in weak flyers, and, as a rule, more in big birds than in small. At the shoulder -joint the coracoid is met by two other bones, the clavicle or merrythought (which varies much in make and strength), and the scapula or shoulder-blade, and the three bones together form a first-rate pivot, so that the strong wing-beats do not shatter the bird's framework. 2U Fig. 20. Clavicle of (1) Tern ; (2) Eagle. Drawn to scale. Muscles and Quality of Muscle. From the keel and from the outer part of the sternum (thus covering up the small Elevator muscle), and also from the clavicle, springs the Great Pectoral — the Depressor muscle ; it attaches by a short tendon to the front part of the under-surface of the humerus (upper-arm bone), and, attaching where it does, its pull tends to lower the front part of the wing relatively to the back. And springing as it does not only from the breastbone but also from the clavicle, it pulls the wing, not downward and backward, but simply downward. The underlying Elevator sends out a long tendon that passes THE MACHINERY OF FLIGHT 71 through a little tunnel formed by the three bones, which meet at the shoulder -joint, and after passing through the tunnel attaches to the upper side of the humerus, near to its preaxial or front edge. Its action is to erect the wing upon its pivot. Attaching Fig. 21. Humerus of right wing of Eagle. Actual length 6f inches. D : Flat area near the front margin of the under-surface, where the tendon from the Depressor muscle attaches. where it does, it must necessarily bring the wing into the proper position for beginning the next stroke with its front edge looking in the direction of flight. 72 THE FLIGHT OF BIRDS Sometimes, in order to carry out a movement, two muscles have to antagonise one another. When a bird wishes to check his speed suddenly, he lets his body hang downward, sometimes almost vertically, and holds his wings so that their under-surface faces in the direction of his flight. Now, if the Depressor were alone holding the wing, it would lower the front relatively to the back, and the resistance of the air would reinforce its action, making the wing turn its under-surface downward as in ordinary flight, whereas for checking speed it must face to the front. To orient it thus, by holding fast the front margin of the upper face of the humerus, is the work of the Elevator. The antagonising of one muscle by another is, of course, constantly going on. Were it not for this, with its steadying effect, no movement could be carried out with precision, no attitude could be maintained. There are, of course, other muscles which contribute to the working of the bird's wing, and, notably, there is a third pectoral, the office of which is to draw the wing back. But it is no part of this short treatise to describe the work of minor muscles. I have already alluded to the different qualities of muscle found in birds. Muscle of the highest quality — capable of an enormous number of very rapid contractions in rapid succession — consists of fibres in which the fine striation in the direction of the pull of the muscle is much obscured by irregular, highly granulated cross ridges. In colour it is a rich red. The fibres consist of small, contractible fibrillse, and are buried in a granular material known as sarcoplasm. Muscles which are capable of long- THE MACHINERY OF FLIGHT 73 continued exertion are rich in sarcoplasm, which would seem somehow to supply food to the muscle proper, to the contractile fibrillse ; at any rate this is a reasonable theory that explains the facts.* Now the muscle which in strong-flying birds is reddest and, in section, most ridged and granulated, is the Great Pectoral that lowers the wing. This is a muscle of great size, as I have already pointed out, and its output of unflagging energy is truly astonishing. The Elevator muscle is paler and smoother, and almost certainly of far inferior quality. Much less is demanded of it. As soon as the bird has got up pace, the resistance of the air lifts the wing, and the Elevator has little to do. In fact the great Depressor may claim to lift the wing, since to it is due the velocity which relieves the Elevator of its work. In the Blackcock, the Red Grouse and the Ptarmigan, the paleness of the Elevator forms a striking contrast against the rich red of the great muscle under which it lies. Among birds whose flight muscles I have examined, the Guillemot is the only one whose Elevator is as rich a red and almost as stringy as the Depressor. Is this explained by the fact that the bird uses his wings in swimming ? Has he to raise his wings in the water by muscular effort ? And is this due to the fact that when he swims he has less pace than when he flies ? In the Chicken, since it seldom uses its wings, all the Pectoral muscles are very pale and smooth. The wild Jungle Fowl, from which our domestic bird springs, uses its legs far more than its wings, and, I believe, domestication has not much altered the * See Starling's Physiology on quality of muscle (pp. 87, 88). 74 THE FLIGHT OF BIRDS character of the different muscles. Remarkable also is the large size of the Chicken's Elevator muscle. Its wild kinsman, haunting the jungle as it does, has to rise to its perch without any wind to help it, so that all the work of raising the wing must fall upon the muscle. But the effort required is not a pro- longed one, and so the muscle is pale. In the Sparrow-Hawk and, I believe, in other birds of prey, the Elevator is very small. Since they usually start to fly from a perch at some height above the ground, there is no difficulty in getting up speed enough to make the office of the Elevator largely a sinecure. The brown flesh on a Chicken's leg, though the human palate pronounces it inferior, is, nevertheless, when regarded as muscle, of distinctly better quality. The Fowl is primarily a runner, not a flyer ; hence, presumably, the darker colour and the greater strength and endurance of the leg muscles. More- over, these muscles are kept on the strain throughout the night; by the bending of the knee and ankle joints the machinery of muscles and tendons through- out the leg is set to work and the bird's toes strongly grip the perch. His weight keeps the legs bent, and the bending of the legs keeps muscles and tendons to their work. Were they to relax for a moment the bird would probably lose his life ; he would fall to the ground and some hungry carnivore, that happened to be prowling about, would seize him. But though in the Fowl the leg muscles are darker in colour than those of the breast and wings, they are not in section ridged and granulated like the great Pectoral of birds that are strong flyers. In the Moorhen, I have noted that the leg muscles are rather more ridged THE MACHINERY OF FLIGHT 75 than is the Elevator, but not so much as the De- pressor. In most birds that I have examined, I have found that the leg muscles show less ridging and granulation even than the Elevator. But the Moorhen makes great use of his legs. He is both a swimmer and a runner. When not swimming he is generally walking on the grass beside his pond or stream, busied with the search for worms. Before I leave the subject of quality of muscle I must point out that the inferiority of the pale to the red has been noticed also in thoroughbred horses. Mr. J. B. Robertson, in his very interesting paper on the Principles of Heredity applied to the Racehorse (p. 22), writes : "The pale fibres greatly predominate in the tissues of a pure sprinter, and the red fibres in those of a stayer." The big muscles, the work of which I have described, are all massed upon the sternum. Even the muscles which bend or straighten the wing at the elbow spring not from the humerus or upper- arm bone, but from the top of the coracoid and the anterior end of the shoulder-blade respectively. I once cut off the wing of a domestic Pigeon as close as possible to the body and found that it scaled just under £ oz. The bird weighed 13i oz. Thus the two wings together accounted for just under one- eighth of the whole. It is wonderful that such strength can be combined with such lightness. And not only is the wing, as a whole, light ; what weight it has belongs almost entirely to the bones and muscles of the near part. This Pigeon's wing balanced, when rested on a wire 2J inches from its base and 10 inches from its tip. The great primary 76 THE FLIGHT OF BIRDS feathers (i.e. those which arise from the hand) point mainly outward and only slightly backward. Hence the length of a bird's wing is due largely to feathers, and they are proverbially light things. Their strength is no less remarkable than their lightness. The Scaffolding of the Wing. Pneumatic Bones. The scaffolding of the wing is itself very light. The thickest of the bones, the humerus, is hollow in big birds that are strong on the wing. In some the bones are hollow right on to the finger-tips ; there is an opening in each bone at the near end (see fig. 22) ; a thin pulmonary membrane enters there, and thus they are filled with air that has passed through the lungs. The Gannet is a good example of this complete aeration. Many small birds, however, though first-rate flyers, have all their wing-bones solid ; the Swallow, for example. The Swift has only the humerus pneumatic. The differences be- tween nearly related birds are remarkable. The Gannet's remarkable pneumaticity I have already mentioned ; his near relative, the Cormorant, has only the humerus pneumatic. Such examples seem to make it clear that aeration is an adaptation to the life and habits of the particular species, not an unvarying character firmly established in certain orders of birds.* The fact that large birds have more aeration than small demands explanation. But the explanation * Besides the wing-bones, the Gannet has the following bones pneumatic : The vertebrae, the greater part of the sternum (not the keel), the ribs, the coracoid, the ischium and the femur. There are also ample air-cushions beneath the skin that covers the breast. THE MACHINERY OF FLIGHT 77 is not far to seek. The small bird would gain but little in lightness by the aeration of his bones, since each bone consists almost entirely of its exterior shell. The big bird, with his stout, bulky bones, will gain far more. Here is a case in which we may appropriately quote some geometrical facts. We have seen above that if we take two cubes, the side of one of which is twice that of the other, then a face of the larger one is four times the area of a face of the smaller one, and the cubic content of the larger is eight times that of the smaller. Thus an increase in the length and girth of a bone means a far greater increase of the space within the outer shell. Obvi- ously, then, a big bird stands to gain more in lightness by the hollowing of the bones. How he manages to do with so very little marrow — the bones having only a very thin lining — is another question to which I hope to return (see Chap. xi). How much the bigness of the bone has to do with pneumaticity is made clearer by the facts which follow. If we make measurements of the humerus of a Skua, or other Gull — the Gulls have very little aeration — and of an Eagle as the representative of the birds which have a great deal, we find that the girth of the Eagle's bone is disproportionate to the bird's superiority in length of wing. And the explanation, no doubt, is this ; the Eagle requires much greater strength in his wing -bones than does the Gull. Even a small increase in length of wing means a considerable increase in the pace at which the extremity will move. And, as we have seen, the resistance of the air increases as the square of the velocity. It is easy to see, then, that the Eagle and 78 THE FLIGHT OF BIRDS other big, long-winged birds have wing-bones of larger girth (of a girth, that is, disproportionate to their superiority in length of wing) in order that they may be able to bear the far greater strain put upon thern. A proportionate increase in weight would, no doubt, have caused difficulty, and this has been obviated by the aeration of the bones. Fig. 22. Humerus of (1) Skua; (2) Rhinoceros Hornbill ; (3) Sea Eagle — actual length 6f inches. Drawn to scale. F : Foramen, where the bronchial membrane enters, on the upper surface. To bring out this point more clearly I have taken the humerus of the Skua as the standard, and have calculated what would have been the length of the same bone in the Sea Eagle if it had been built on the same lines. After all this I have to admit that there remains a very puzzling case on which I cannot throw any light. The Hornbills are slow, heavy flyers, they are not very big, and they are the most pneumatic THE MACHINERY OF FLIGHT 79 Girth of humerus. Humerus. Aggregate length of wing-bones. Actual length. Length propor- tionate to girth. Actual length. Length propor- tionate to girth of humerus. Ins. Ins. Ins. Ins. Ins. Skua II H *i 13| 13f Sea Eagle 1A 6| 7 7 '20 20 T V 22^ of all birds. The huge hollow beak is characteristic of the whole skeleton (see fig. 22). Stiffness of Wing. Expanse of Bone. It is highly important that the bird's wing when extended should be rigid — its bony framework, that is. It has elasticity where it is wanted, the elas- ticity of the great feathers, which tends, as I have shown, to make the bird automatically stable during flight. The stiffness of the scaffolding of the bird's extended wing is most remarkable when we compare it with the human arm. If we extend one of our arms horizontally to its full length with the palm of the hand downward, we can, while still keeping the upper arm lifted, bend the elbow- joint and point the fore-arm vertically upward. As the bird's wing descends with great velocity through the air, it does not give at the elbow, but remains rigid. But the stiffness of the bird's wrist is still more striking ; when the wing is extended hardly any up-and-down movement at that point can 80 THE FLIGHT OF BIRDS take place. The rigidity there is more easily brought about owing to the fact that of the two small rows of wrist-bones the further one has been fused with the hand-bones. When the fore-arm is brought into line with the upper arm, the hand also falls almost into line and is held tight in that position by ligaments. As soon as the wing is flexed at the elbow, it bends also at the wrist — bends backward, that is, towards where digit 5 MC Skeleton of wing of Adjutant (from the elbow). D 1, 2, 3 : Digits 1, 2, 3. MC 1, 2, 3 : Metacarpals 1, 2, 3. R : Radius. RC : Radial carpal. Ul : Ulna. UC : Ulnar carpal. The marks left by the little pockets in which the secondary feathers are planted can be seen on the ulna. would be if it existed — and then at the latter joint much freer upward movement also is possible. And so the wing is rigid when rigidity is wanted, and pliant when pliancy is in demand {see fig. 23). THE MACHINERY OF FLIGHT 81 It is essential, too, that there should be a large expanse of bone, a firm foothold for the great flight- feathers. The lizard, or lizard-like reptile, from which birds are undoubtedly descended, had a very wobbly hand. Even the Archaeopteryx, the most ancient of birds, though it was beyond doubt a true bird and not a bird-like lizard, had three long fingers which show no sign of becoming fused together. With birds as we know them, things are very different. The three surviving metacarpals, or hand-bones, enlarged by the fusing with them of the further row of carpals or wrist -bones, are themselves fused together, the two of the three that are the most important fused both at the near and further ends. Of the three surviving digits two are fused ; the little thumb, that supports the bastard- wing, remains independent and insignificant ; and the result of all the fusing is that a broad, firm platform is provided for the feathers. Thus we see that the scaffolding of the wing when extended is remarkably rigid in spite of its lightness ; its wide expanse of feathers is remarkably elastic, and is planted on a broad, firm base. The Spreading of the Wing. The machinery by which the wing is spread is well worth study. The rapidity with which it does its work is wonderful, and the accomplished result is most beautiful. When the triceps muscle straightens the wing at the elbow- joint, it is straightened also at the wrist, not absolutely but nearly ; there are special muscles for giving the finishing touch. Of the two bones in the fore-arm the front one (the 82 THE FLIGHT OF BIRDS radius) is pulled towards the body when the elbow is straightened and this straightens the wrist. A further result inevitably accompanies the first straightening movement. There is an elastic liga- ment that stretches from the armpit to the extremity of the digits. Through this pass all the secondary flight-feathers and, in every bird in which I have traced its course, all the primaries (those that spring from the hand). Part of this ligament is shown in Fig. 24. Wing viewed from below (after Alix, with a slight alteration, see text). AM : Muscle and tendon supporting anterior mem- brane, Dl : Digit 1. El: Elastic ligament. F: Flexor carpi ulnaris, dividing into two. T : Tendons connecting muscle with quills. figure 24. But it is easy to see the whole of it in any bird's wing when the small covering feathers have been removed. The straightening of the wing stretches the ligament and the great flight-feathers spread like a fan.* {See PL xi.) * There are differences in different families. In some the liga- ment, instead of being pierced by the quills, fastens only to their under-side. THE MACHINERY OF FLIGHT 83 There is another ligament which fastens to the under-side of the quills of the secondaries nearer to their base. Moreover, the secondaries grow back- ward, and plant themselves each in a little pocket of very tough fibrous tissue. These pockets are firmly rooted in the hinder of the two bones of the fore-arm (the ulna) and leave, each of them, their mark on it (see fig. 23). Planted thus, the second- aries, though very firmly gripped, have freedom to rotate. The primaries, pointing outward as they do, run along above the hand and finger bones and are held with a still stronger grip. It is marvellous machinery which spreads the wings so rapidly, and which, having all its heaviest parts massed upon the body or very near to it, leaves them splendidly light, especially near their extremities. But we have not yet come to the end of the contrivances. During the down-stroke the secondaries press each against the one above it and so prevent the passage of air, whereas during the upstroke they let it pass. They are held fast by a number of stays ; there are the little pockets just described, the ligaments and a sheet of fibrous tissue extending from their bases to the great liga- ment. There are also little tendons that connect the secondaries with a muscle which, arising from the further end of the humerus, attaches its other extremity to one of the wrist-bones (carpals) and to one of the metacarpals, or hand-bones, beyond (see fig. 24). Working unopposed it bends the wrist. When the wing is straightened it helps by its opposi- tion to hold the wrist- joint tight, thus strengthening a point where the strain of flight is much felt. When 84 THE FLIGHT OF BIRDS the wing is bent the muscle lies in a slightly curved form — we notice it when we are getting the meat from between the two long bones of a chicken's wing. It is straightened out when the wing straightens, and this, combined with the outward slope assumed by the feathers as they spread, stretches the little tendons that arise from it and are fastened to the feathers. The tendons slope outward, away from the shoulder, and attach to the under-side of the quills, or rather to the fibrous tissue with which the quills are surrounded. M. Alix (in his Appareil locomoteur des Oiseaux) makes some of them at any rate attach to the further edge of the lower face of the quills, and gives them the credit for rotating the feathers during the down- stroke, so that they press against one another and make the wing impervious to air. But when I have scraped these little tendons away in the wing of a freshly-killed Pigeon I have found that, when I extended the wing, the feathers still took their proper position. The extending of the wing causes a stretching of all the important stays — the ligaments, the sheet of fibrous tissue, and these tendons — and the ligaments may claim the largest share of the credit for the spreading and marshalling of the feathers. During the up-stroke the secondaries are firmly tethered, but are no longer marshalled for action. There are interspaces as there are between the leaves of a folded fan that has seen much service. The feathers being now no longer pressed tight together, the air can pass between them. When, however, the bird is flying fast and the wings are lifted, not so much by muscles PLATE XI. A : Wing of Lark, showing the elastic ligament that holds the primary and secondary feathers. Another ligament nearer to the base is faintly shown. B : Primary wing -feathers of Black Vulture (on left) and Pink-footed Goose, upper surface ; of Heron, under surface ; and outer tail-feather of Heron. In all the inner web is the broader. (See Chap. VII.) [To face p. 85. THE MACHINERY OF FLIGHT 85 as by the resistance of the air, the feathers during the up -stroke are pressed against one another and the passage of the air is prevented. But this causes no trouble. The wing is very rapidly swept back- ward and upward ; it turns its front edge in the direction of flight, and when it is oriented thus it is easily straightened and moved forward into position for beginning the next down-stroke. During the down-stroke the air helps the living machinery in its work, making it still more effective. The outer webs of the feathers (see PI. xi) are very narrow compared with the inner ones, and the result is that the air acts much more strongly upon the latter during the down-stroke and also, when there is much momentum, during the up-stroke, with the result that each feather is rotated and has the inner side of its vane pressed very closely against the one that lies next to it and above it on the side nearer to the body. Though on the surfaces of the wing nothing is visible but feathers, yet for no small amount of the expanse two membranes can claim the credit. What is called the anterior membrane stretches from the head of the clavicle (or merrythought) to the hand (see fig. 24). In the Gannet it is of great breadth and is so hung that it not only increases the area of the wing, but sloping steeply downward, as it does, to form the front margin, it deepens the wing's concavity and makes it, near to the body, what a parachute should be. In most birds the membrane is slung with only a gentle slope from its front edge backward. In all it is at once stretched when the wing is spread. The other membrane lies farther 86 THE FLIGHT OF BIRDS back, in the armpit, and fastens the wing to the bird's side. It is worth while noticing that though muscles springing from the body in the main control the movements of the whole wing, yet there is a good deal of local independence. There are niceties of adjustment which depend on local muscles. Though the triceps extends the upper arm, the fore-arm and the hand, yet the two united fingers, and with them the great feathers they carry, are not under its sway but depend upon special muscles to spread them to the full and carry out minor movements. The little bastard-wing has also its own muscles — more muscles than one would expect to be at the service of so insignificant a piece of machinery. Structure of a Flight-Feather. For the bird flight without feathers is obviously an impossibility. If the scales of the bird's reptilian ancestors had remained mere scales, the ancestors would have remained reptiles still, condemned to crawl the earth. The scale has been glorified till it is hardly recognisable. First comes the quill with the dried remains of the pulp — the pulp that was there when it was alive and growing — still visible within it. Above is the rachis, or shaft, grooved down its front face (see Pis. xi and xn). From the shaft spring the barbs sloping towards the tip of the feather, from the barbs branch out the barbules or radii. From those of the barbules that are on the far side of the barb (the side farther from the base of the feather) spring the barbicels (diminutive of a diminutive !) that fasten barbule to barbule and PLATE XII. (From Life and Evolution.) The barbs, barbules and barbicels of a flight-feather. A and C x 90 diameters ; B x 250. A : A barb. C : A pair of barbs showing interlocking. B : A pair of barbules, showing the barbi- cels on the one nearer to the tip of the feather. To face p 86. THE MACHINERY OF FLIGHT 87 thus give the feather its elasticity, besides making it impervious to air. Compare for a moment a great flight-feather with one of those whose business is merely to clothe the bird and retain its animal heat — a poor, weak, fluffy thing. The barbicels have another name, hamuli or hooklets, from their shape. On the side of the barb which is nearer to the base of the feather, the barbules have no bar- bicels, but their endings resemble hairs, and these hair-like endings are neatly folded together so as to form a kind of hem. The opposite set of barbules lie over these smooth-ending ones, and among them they insert their hooklets. The hooklets slide along the barbules at moments of strain and stress, and to this, very largely, must be due the elasticity of the feather. The number of barbules is enormous ; according to Dr. Gadow, over a million in one large flight-feather. Here is another very interesting point: on the margin of the barbules opposite to that which is armed with barbicels are rough knobs. These are, I believe, the vestiges of barbicels which, being useless, are in process of disappearing. The feather is in fact thrice pinnate. The main shaft branches into barbs, the barbs into barbules, the barbules formerly branched, I believe, into barbicels on either side. But the barbules on the side nearer to the base of the feather have altogether lost their barbicels and have now only hair-like endings, while the other set have only one member of each pair properly developed, the other member being represented only by a mere vestige, so that there are only rough knobs to match the barbicels. 88 THE FLIGHT OF BIRDS From whatever point of view we look at it, a flight- feather is a wonderful thing. It provides a large expanse to support the bird's weight, it is elastic, it is light and at the same time strong. Moreover, it is renewed every year, and in such a way that the bird does not even for a day lose his power of flight. Moulting. A living machine differs from a man-made machine in many ways, and notably in this, that it has as an indispensable characteristic, the power of self -repair. When a bird's feathers are broken or worn out, they must somehow be replaced. Now nearly all birds that fly shed their flight-feathers gradually and in pairs, so that, though during the moult they are not at their best, yet they can always rise on the wing (see PI. vni, 4). The Goose has somehow earned a widespread reputation for stupidity, but the most stupid thing that he does is not, I believe, generally known. He moults so rapidly that for a time he is reduced to helplessness. In the island of Kolguev, the Samoyeds drive thousands of moulting geese, who can only swim or run, into great nets and thus provide themselves with a store of food for the Arctic winter. But, after aJl, this rapid method of moulting would involve but little danger for the Goose if man had not arrived on the scene, since his way of life became stereotyped. Many birds, not geese alone, have failed to find any means of escape from their enemy with his ever-changing method of attack. There are other birds which shed all the primary-feathers simultaneously without any disastrous consequences, THE MACHINERY OF FLIGHT 89 e.g. the Ducks. Mr. J. L. Bonhote has made a special study of the subject, and, quoting from him, I am able to give some very interesting and instruc- tive facts. The simultaneous moulting of the primaries occurs among the Divers, the Guillemots, the Razorbills, the Puffins, the Auks, and besides these among the Rails, e.g. the Moor-hens, the Land- rails, and the Crakes. Nearly all of them are water birds, and in the water they are fairly safe from their ordinary enemies. Some of them can dive, others can hide among reeds or under bushes ; the Geese by force of numbers can, probably, beat off a bird of prey. A sea bird, if he is to moult in this fashion, must be a true diver in habit and in plumage, if not in name, and thus we find that the Cormorant (who is always wanting to dry his wings on a rock) and the Gannet moult gradually. When we turn to land birds the case of the Corncrake does not surprise us, for though he is a migrant, yet in his day-to-day life he trusts more to his legs than to his wings. The case of the Coot is perplexing ; unlike the Moor- hen, he is capable of flight throughout his moult ; but, as Mr. Bonhote points out, he is the only member of the Rail family that frequents estuaries and open sheets of water, and he is, in addition, an indifferent diver.* Legs. I have already pointed out how some birds use their legs in balancing and, possibly, for steering. Indirectly the great leg-power possessed by most birds is a most important aid to flight. In order to * See " Eclipse Plumage and FHghtlessness " by J. L. Bonhote, in the Field, March 24, 1906. 90 THE FLIGHT OF BIRDS rise with ease from the ground, they must be able first to jump into the air. Most birds have an active springing gait, and are good jumpers, their legs being built much in the same way as those of a horse or an antelope ; the ankle-joint is raised high above the ground, and they walk or run upon their toes. The springiness is combined with remarkable light- ness. The bird's foot is made almost entirely of skin, bone, and tendon ; not like the human foot, fleshy and full of nerves. It is worked by long tendons attached to muscles that spring from the top of the leg- bone (the tibio -tarsus) or even from the base of the thigh-bone, and, running in grooves under the ankle-joint, give to the foot its springiness and to the toes their wonderful grip. The fusion of the two rows of ankle-bones with their bigger neighbours makes the ankle-joint an excellent pulley. Alto- gether the bird's wings have in the legs very able assistants. Comparatively few resemble the Swift in having very short and feeble legs ; but, as I have pointed out above, there are a considerable number, mostly big, bulky birds, which, owing to their shortness of leg and small power of jumping, have difficulty in starting to fly from level ground. CHAPTER VIII. VARIETIES OF WING AND OF FLIGHT. CURVE — NARROW AND BROAD WINGS STYLES OF FLIGHT FLIGHT IN FLOCKS THE WHIR OF WINGS. Curve. Narrow and Broad Wings. I shall first consider the question of curve, or, as the aviators call it, of camber — a question of the utmost importance. The wings of all birds are a good deal curved from front to back, so that as they descend they catch the air in a concavity and have plenty of lifting power. In some birds the front-to- back curve extends throughout the whole length of the wing, though in all there is some shallowing towards the tip. The Jay and the Red-legged Partridge (and their allies) supply good examples of a curve that shallows near the tip of the wing but does not disappear. In a Jay, of whose wings I made measurements, the depth of the curve — measured just on the near side of the starting-point of the bastard-wing — was 1 inch (see PI. xv and also xiii and xiv ). The breadth of the wing at this point was 5 1 inches. Thus the curve is 1 in 5|, a curve such as no aviator would ever think of using. But it must be remembered that during the down-stroke it is much reduced by the pliancy of the feathers, which yield to the pressure of the air. The span 92 THE FLIGHT OF BIRDS from wing-tip to wing-tip was only 18 inches. In fact it was a very short, broad, much-curved wing. The Pheasant's wing belongs to the same class ; it is very short (in a hen bird of which I took measure- ments, only 20 J inches) ; very broad (6 inches) ; and the curve in this particular bird was no less than 1J inches — If in 6 ! (see PI. xiii). The Hoopoe's wing is very much of the same build — unfortunately I cannot give exact measurements — and yet, when the season of migration comes round, many Hoopoes manage somehow to flap across the Mediterranean. As a rule migrating birds are characterized by wings of a very different make, and it sometimes happens that the wing is the feature in which they present the most striking contrast to their non-migratory kin. The clamorous Reed- Warbler, that is resident in Egypt, shows by his wings that he does not migrate — they are so very short. The migrant Reed- Warblers have decidedly longer and more pointed wings. As to the Hoopoe and its short, broad wings, it must be remembered that though it manages to cross the Mediterranean when the conditions are favourable, yet its migration flights are comparatively short. Some, I believe, do not migrate at all. If they are delayed by clouds while crossing the Mediterranean, their strength is apt to give out, whereas Herons and Swallows, to take two examples, have a reserve of vitality and will fly round a steamboat for hours, till, perhaps, the weather clears again, when with unflagging strength they will continue their journey. To pass on to the Red-legged Partridge, I found the depth of the curve to be 1 inch, as in the Jay, but St) . .3.9 * g o * 05 fi 2 88 1° -9 d £5 VARIETIES OF WING AND OF FLIGHT 93 the width of the wing was only 4| inches, whereas the Jay's was 5f inches wide. The wing-tip to wing- tip measurements were 18| inches, hardly an advance upon the Jay. It is a very short wing, and though narrower than the Jay's, it is a broad wing if we compare it with those of most birds. The Moor-hen, also, has a rounded and not very long wing, with a good deal of front-to-back curve, shallowing, however, very much near the extremity. The measurements in one specimen were : curve, T 7 ^- inch ; breadth, 4 £ inches ; wing-tip to wing-tip, 21 J inches. The breadth is very little greater than in the case of the Red-legged Partridge ; in length the Moor-hen has decidedly the advantage, and the gaps between the extremities of the feathers are much less ; in other words, the wing is more finished and more efficient. Here in England we look upon the Moor-hen as a stay-at-home bird, but it will cross the Alps in search of a warmer climate if need be. Frozen-out Moor -hens from northern or central Europe often accomplish this feat. In other birds we find the curve much more reduced towards the extremity of the wing ; in some it dis- appears almost entirely. In the Thrush the last inch and a half is nearly flat. In the Starling (PL xv ) there are quite 3 inches with very little curve. To take a bird of larger build, the Curlew's wing (PL xiii) shows a great reduction of curve as soon as it begins to taper to a point ; the last 4 inches are nearly flat. Moreover the curve has not the same character throughout. For a distance of some 9 inches from the body, a front margin, nearly an inch broad, is flat ; behind that the downward 94 THE FLIGHT OF BIRDS curve begins. Further out the whole breadth of wing is curved ; for some distance from the front margin there is an upward slope, then a downward slope sets in. In the nearer region, also, there is not a simple curve maintained throughout, but the feathers in one part are more bent down than in another, so that the back edge presents a curiously undulating line. The Sparrow-Hawk's wing has similar curves, but they are much less pronounced. What the exact significance of all this complication may be, presenting so marked a contrast to the uniform curves of an aeroplane, it is difficult to say. Probably there is some significance, since in birds that are strong on the wing the whole mechanism of flight down to minute details is so efficient. Sometimes we may see the wing-curves in their full beauty in a live bird, but it is only for a moment, and one wishes to look at them quietly and study them. The thing is, if one has a freshly-killed specimen, to cut off the wings close to the body and pin them out back downwards at their full stretch. Treated thus the wing retains its curves and a great deal of its beauty. If it is pinned face downwards and flattened, it conveys by no means so good an idea of what it was during the life of the bird. I have tried by photographs to show the outlines of various types of wing, the depth of their concavities, and the undulating curves seen in, for instance, the Curlew's wings. Some points come out distinctly, others the camera fails to see or does not see clearly. To show the gradual tapering and flattening of the farther half, perhaps no wings are better than those of the Gannet and the Tern (PI. xiv), and bb CO > .a -: fi o 6? ,j3 ts «5 © U © &Q E " — ,_, CO 2 — ^z^ -4-3 Z - r3 o o ^ 6 s £ bfl be o o o - 00 oa 5 -(j » X •^ £ .3 = CO c3 > © ( — M p —i CO P* »-* £ <<4 _ * k ... CO CD en *J 1 O t& <0 •* PO • ^ L2ii: Eeadley, F. W. The flight of birds. 1912/