324 THE AERONAUTICAL JOURNAL [October, 1914

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ON THE F L I G H T OF PTERODACTYLS.
BY
DR. E. H. HANKIN, M.A., F.R.MET. SOC., A.F.AE.S. (AGRA, INDIA),
AND
D. M. S. WATSON, M . S C , F.Z.S. (UNIVERSITY COLLEGE, LONDON).

I t is p e r h a p s u n f o r t u n a t e , f r o m t h e p o i n t of v i e w of a v i a t o r s , t h a t t h e e x t i n c t
flying r e p t i l e s k n o w n a s " p t e r o d a c t y l s " o r " p t e r o s a u r i a " n o l o n g e r e x i s t .
T h e r e c a n b e n o d o u b t , t h a t in t h e c a s e of t h e m o r e h i g h l y e v o l v e d m e m b e r s of
t h e g r o u p t h e o r g a n i s a t i o n w a s m o r e s p e c i a l i s e d for flight t h a n t h a t of a n y o t h e r
a n i m a l of w h i c h w e h a v e k n o w l e d g e . O t h e r flying a n i m a l s c a n w a l k , r u n , o r
s w i m , b e s i d e s fly. B u t in t h e c a s e of t h e h i g h e r p t e r o d a c t y l s t h e i r s t r u c t u r e is
s u c h t h a t it is difficult t o u n d e r s t a n d h o w t h e y c a n h a v e h a d a n y o t h e r m e a n s of
p r o g r e s s i o n t h a n flying. W i t h a b o d y little l a r g e r t h a n t h a t of a c a t t h e y h a d a
s p a n of w i n g a s s e r t e d in s o m e c a s e s t o h a v e r e a c h e d 21 feet o r m o r e . 1 These
h u g e w i n g s w e r e s o c o n s t r u c t e d t h a t it w a s i m p o s s i b l e for t h e m t o b e furled
a g a i n s t t h e b o d y a s h a p p e n s w i t h t h e w i n g s of b i r d s a n d b a t s . O n l y t h e o u t e r
half of e a c h w i n g could b e b e n t b a c k w a r d s in t h e d i r e c t i o n of t h e b o d y ( F i g . 1).
W i t h s u c h p a r t i a l l y furled w i n g s it m a y b e a s k e d h o w t h e y could p o s s i b l y s w i m
if t h e y e v e r a l i g h t e d o n t h e w a t e r o v e r w h i c h t h e y flew.

T h a t p t e r o d a c t y l s w e r e i n c a p a b l e of p r o g r e s s i n g a s q u a d r u p e d s is p r o v e d by
t h e fact t h a t , a s will b e f u r t h e r e x p l a i n e d b e l o w , t h e fore l i m b w a s i n c a p a b l e of
m o v e m e n t in a f o r e a n d aft d i r e c t i o n a t e i t h e r t h e s h o u l d e r o r e l b o w j o i n t s . 2

If t h e y e v e r w a l k e d o n t h e i r h i n d l e g s t h e y c o u l d o n l y h a v e d o n e so w i t h
t h e w i n g s extended, as otherwise t h e wing-tips would have trailed a l o n g the
ground.
P e r h a p s t h e m o s t f e a s i b l e m e t h o d of p r o g r e s s i o n for t h e m w h e n o n l a n d
is t h a t , h a v i n g a l i g h t e d o n t h e i r feet, t h e y fell o v e r o n t h e i r s t o m a c h s a n d p u s h e d
t h e m s e l v e s a l o n g , a f t e r t h e m a n n e r of p e n g u i n s , b y m e a n s of t h e h i n d l e g s ,
p e r h a p s w i t h a n o c c a s i o n a l s l i g h t lift f r o m t h e w i n g s for s u r m o u n t i n g a n o b s t a c l e .

T h a t t h e y c o u l d h a n g b y t h e i r h i n d l e g s f r o m t h e b o u g h s of t r e e s , a s is t h e
c u s t o m w i t h b a t s , is i m p r o b a b l e for v a r i o u s r e a s o n s . I t is n o t e v e r y b o u g h t h a t

1
A statement has recently appeared in a semi-popular scientific magazine to the effect that
the air of the present day could not support animals of such large span as the larger pterodactyls,
and that consequently the barometric, pressure in Cretaceous times must have been about twice
as great as it is now. But it may be doubted whether there is so great a difference between
birds of the present day and pterodactyls in respect of span as the author of this theory believes.
Pelicans reach a span of fifteen feet. Latham (" General History of Birds," Vol. X., p. 48, 1824)
gives records of the span of albatrosses of 10, n , 12, and 13 feet. He mentions the " Voyage
from England to India in the year 1754," by Ives (Edward), formerly Surgeon of Admiral Watson's
ship and of His Majesty's Hospital in the East Indies, on p. 5 of which appears :—•" An albatrose
(sic), a sea fowl, was shot off the Cape of Good Hope, which measured 17J feet from wing to wing.
A shark was also caught, and brought on board the Cumberland, with 72 young ones in her
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belly, each from 6 to 14 inches long." A gentleman in India gave to one of us a detailed account
of his shooting and measuring a bird, stated to be an adjutant, with a span of 18 feet. No
reason can be given for believing that the largest existing birds would become incapable of flight
were the earth to undergo a great reduction of atmospheric pressure. Large birds of heavy
loading, such as the black vulture, can soar as well at a height of two miles above the earth's
surface as at lower levels. At such a height the barometric pressure normally stands at about
20inches. There is reason for believing that pterodactyls were of much lighter loading than (he
black vulture.
2
A restricted range of fore and aft movement could occur at the wrist joint, but, as will be
shown later, this was so connected with movements of rotation as to be singularly ill adapted for
purposes of progression.
 October, 1914] THE AERONAUTICAL JOURNAL 325

could support their weight, which in the larger specimens is supposed to have
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reached 2olbs. A bat flies with head first and hind legs stretched out behind.
In order to g r a s p the bough with its feet it has t o check speed ahead, which it
does by " poise-flapping " at the same time rotating round the transverse axis.
As soon as its feet are thus brought forward they g r a s p the bough and the bat
rapidly furling its wings falls over in one direction or another to reach the head
downwards position. But in view of the large span of its wings the pterodactyl
could only attempt " poise-flapping " above the topmost twigs of the tree, which

FIG. I. PTERANODON. V I E W FROM BELOW, AND SIDE VIEW OF BODY W I T H WING

REMOVED.

twigs would be quite incapable of supporting its weight. If it chose a bough of

sufficient size and therefore nearer the trunk it is quite certain it could only h a v e
attempted poise-flapping with imminent risk of tearing or breaking its wings.
On the other hand, there can be no doubt that they were fully adapted for
https://doi.org/10.1017/S2398187300140290

life in the air. Throughout their organisation weight-saving has been carried
to an extreme. T h e hollow air-filled wing bones of vultures were heavy columns
compared with the delicate empty tubes that supported the wings of pterodactyls.
These tubes were made of hard bone material scarcely thicker than a visiting card.
At the two extremities the bones were strengthened internally by delicate struts-
and bands of bone of paper-like thinness. T h e bones of the hind limb, t h e
vertebrae, and even the phalanges of the wing-finger were hollow and filled with-
air. Striking illustrations of these facts may be seen in the specimens exhibited'
in the British Museum (Natural History), South Kensington.
 326 THE AERONAUTICAL JOURNAL [October, 1914

Remains of the larger pterodactyls are found in some cases in strata of

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marine origin under circumstances that make it probable that they habitually flew
a t some such distance as a hundred miles from the nearest land. 3 They seem to
have been mainly fish-eating in habit.

In view of the extreme specialisation of the pterodactyls for flight the construc-
tion of their wings is a matter of great interest. Owing to the above-described
delicacy of the bones the remains met with as fossils are usually so crushed that
it is impossible to discover the mode of action of the different joints. Some
exceptionally perfect examples of the bones of the pterodactyl Ornithodesmus
have recently come into the possession of the British Museum (Natural History).
From a study of these and other specimens we are now in a position to describe
the movements of which the different joints of the wing were capable. Our
account is founded entirely on the Cretaceous forms. The articular surfaces are
described from uncrushed specimens in perfect preservation from the English
Wealden and Cambridge Greensand. The different specimens agree in the shape,
number and arrangement of their articular facets, and further agree exactly with
those of similar bones of Pteranodon in the British Museum (Natural History).
Ornithodesmus is stated by R. W . Hooley 4 to have had a span of five metres
" allowing for the natural c u r v e . " If the wing bones were placed in contact, end
to end, in a straight line, a span of more than five metres would be obtained.
But as the bones of the wing were habitually held in a slightly flexed position at
the different joints Hooley reduces his estimate of the span to five metres.

As indicated by the name " p t e r o d a c t y l , " the wing is formed by the bones
of the arm together with the phalanges of one enormously elongated finger. The
structure of the wing was such as to permit movement at the shoulder, elbow,
wrist, and knuckle joints (Fig. i). The wing-finger is formed by four phalanges,
but these bones were fixed together by bony union in some cases and there is
no reason for thinking that movements could occur in any case at the different
joints of the finger.
In the case of birds and bats the shoulder joint is a ball and socket joint.
Thus the wing can not only be flapped up and down, but it can also be advanced
and retired. As has been shown by one of us such advancing and retiring is
used by the bird or bat for the purpose of changing the position of the " lift "
in relation to the centre of gravity and for thus causing movements round the
transverse axis. 5
But in the case of the later pterodactyls the shoulder joint was a hinge joint
that only permitted motion vertically up and down. A wide range of such move-
ment could occur as in the shoulder joint of birds. In that there is absolutely
no sign of curvature of the facet of the joint in the anterior posterior direction,
we regard it as certain that no advancing or retiring of the wing could occur at
the shoulder joint of the later pterodactyls. The flapping up and d o w n . a t the
shoulder joint was in a perfectly vertical direction. Thus, movement at this
joint was suitable for high speed flapping flight. Flapping downwards and
forwards, which is necessary for slow speed flight of birds, could not be accom-
plished by movements of the shoulder joint of pterodactyls without further adjust-
https://doi.org/10.1017/S2398187300140290

ments. It may be noted that there was no possibility of any movement of flexing
backwards of the wing against the body as occurs at this joint in birds. That

3
G. F. Eaton, "Osteology of Pteranodon." (Memoirs of the Connecticut Academy of Arts
and Sciences, New Haven, Vol. II., July, 1910.)
* " O n the Skeleton of Ornithodesmus latidens, an Ornithosaur from the Wealden Shales of
Atherfield (Isle of Wight)." (Quarterly Journal of the Geological Society, Vol. LXIX., June,
1913, No. 274, p. 372.)
5
E. H. Hankin, " Animal Flight," p. 156. (London : Iliffe & Sons, 1914.)
 October, 1914] THE AERONAUTICAL JOURNAL 327

is to say, pterodactyls could not furl their wings against the body by movements
a t the shoulder joint.
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W e will now consider the elbow. In the case of birds the elbow is a hinge
joint that permits of extending and flexing movements in the horizontal plane.
Flexing at this joint is used in furling the wing. In the case of pterodactyls the
elbow joint was also a hinge joint. The direction of permissible movement was
not horizontal as in birds. It was not vertically up and down as at the shoulder
joint of pterodactyls. It was between these two directions. Whereas flapping
downwards at the shoulder was vertically downwards, flapping downwards at the
elbow was not vertically downwards but in a direction that made an angle, so
far as we can judge, of nearly twenty degrees with the vertical. 6 Thus,
movement at the elbow joint would give flaps in a downwards and forward
direction. If there was nothing more to be said we might conclude that such
flapping downward and forward occurred and was required for slow speed flight,
as in the case with birds and bats. But before drawing this conclusion the
matter requires discussion.
In the first place birds advance their wings for slow speed flapping flight
both to compensate for the travelling backwards of the lift as speed decreases
and also in order that the flapping should produce more lift qua flapping propor-
tionate to the decrease of " pull " with its consequent decrease of life due to gliding.
T h a t a gliding action and consequent lift occurs during flapping flight is shown in
" Animal F l i g h t , " p . 222. Now the above suggestion as to the method of slow
speed flapping in pterodactyls is not excluded on the ground that the suggested
method would produce no " rotation up " round the transverse axis. It would be
an advantage to the pterodactyl if it could increase lift qua flapping while keeping
the long axis of its body horizontal and therefore in the most suitable position
possible for speed ahead. But the above suggestion is excluded on the ground
that it takes no account of the backward travel of the lift. This would require
to be compensated by an advance of the wings or of part of the wings. No
advancement is possible at either the shoulder or elbow joints. Neither, as will
be shown below, could there be advancement at the wrist joint lasting through
both the up and down strokes. But advancement could be produced, as will be
explained in a later paragraph, by increase of extension of the outer part of the
wing by movement at the knuckle joint. But if this occurred there would be
rotation upwards round the transverse axis and flapping from the shoulder joint
would then be downwards and forwards. Hence, the suggested movement at
the elbow joint would be superfluous.

In the second place the range of movement at the elbow joint is only about
30 degrees. This is quite insufficient for ordinary flapping flight, whether slow
or fast. It would only be sufficient for " half flaps," which in the case of birds
only occur in flight under certain infrequent atmospheric conditions.
Thus the first suggested explanation of the movement at the elbow joint
having failed we must look for another. Let us first examine more minutely the
structure of the joint.
. The bones that enter into this joint are the arm bone or humerus, and the
https://doi.org/10.1017/S2398187300140290

bones of the forearm, namely, the radius and ulna. In the human arm the outer
end of the radius is not fixed firmly to the outer end of the ulna. The attachment
permits of a certain amount of sliding so that the end of the radius can slide
round the end of the ulna. This movement is connected with rotation of the
hand by movement at the wrist joint. In pterodactyls the radius had no power

6
Imagine lines drawn along the greatest diameters of the facets at the two ends of the
humerus. When seen in an end-on view of the humerus these two imaginary lines make an
angle with each other, which we measured in two cases, and found to be i8° and 19 0 respectively.
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https://doi.org/10.1017/S2398187300140290

03
to
03

Trox-i yn- Q-I J)ist a I i

Cexr o (X-L Carpal
Wind vrtttaccLrp^X' *3
fed

ft)
©

o
ft)

L(5.ttT 0-1

F I G . 2. ORNITHODESMUS. V I E W FROM IN FRONT OF LEFT WRIST JOINT WITH BONES DISARTICULATED.

 October, i9i« THE AERONAUTICAL JOURNAL 329

of movement of this nature. But the radius could slide in the direction of its long
axis through a distance of nearly a quarter of an inch (in Ornithodesmus). T h e
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long axes of the two bones the radius and ulna are parallel to each other and
remain parallel during this movement. In other words the radius could take up
two extreme positions which we may describe as the " inward position " and the
" outward position." The elbow, as stated above, is a hinge joint. The surface
with which the radius articulated is not of quite regular curvature. T h e irregu-
larity is such that the radius assumes the " inward position " when the forearm
is raised up as far as possible. It assumes the " outward position " when the
forearm is directed downwards as far as possible.
T h a t this in-and-out displacement of the radius actually occurred is proved
by the fact that there are two articular surfaces on the radius and ulna where
they come into contact with each other near each extremity, which surfaces must
obviously have facilitated the movement.

P I G . 3. V I E W FROM BODY SIDE OF LEFT PROXIMAL CARPAL BONE. A BEFORE AND B

AFTER RECEIVING PUSH FROM RADIUS.

The fact that the radius had this singular power of movement obviously
throws no light on the function of the elbow joint. To discover the meaning of
the movement it is necessary to consider the effect of the movement of the radius
on the bones of the wrist joint.
A front view of the wrist joint with the bones disarticulated for the sake of
clearness is shown in F i g . 2. The distal 7 end of the radius is seen lying in front
of the distal end of the ulna. On the outer or wing-tip side of the joint is shown
the proximal end of the metacarpal of the wing finger. Between these bones are
https://doi.org/10.1017/S2398187300140290

shown the two chief bones of the wrist which are known as the " proximal carpal "
and the " distal c a r p a l . "
W e must first consider the movement that occurs on the proximal side that is
on the body side of the proximal carpal.
Fig. 3 shows the inner surface of the proximal carpal as seen in end on view

'The two terms, "distal," meaning the side furthest from the body or from the centre of
the object described, and " proximal," meaning nearest the body, are in common use in anatomy,
and might be found useful in other sciences.
 330 THE AERONAUTICAL JOURNAL [October, 1914

from the direction of the body of the animal. At the lower and posterior part of this
surface is seen a projection P . This fits into a corresponding depression on the
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end of the ulna. Thus, the lower side of the end of the ulna was pivoted at this
spot. The upper part of the outer end of the ulna rested against the curved
surface U F . The outer end of the radius rested against the facet R F .

As stated in previous paragraphs the radius can slide for a short distance
in the direction of its own length. W h e n it does this it pushes R F . This push
from the radius has the following consequences.

Firstly, there is a slight rotation of the wrist (relatively to the ulna) round the
pivot P . The direction of the rotation is indicated by the two diagrams in Fig. 3.
It results in a rotation upwards of the metacarpal bone round its long axis.

Secondly, there is rolling in the direction D C . Before the movement the

letter C and the letter D in the diagram may be regarded as having been equi-
distant from a pair of points placed one on each side of the end of the ulna.
After the rolling' one point on the ulna is further from D while the other point
is nearer to C. This movement is equivalent t o a retirement combined with
slight depression of the outer part of the wing. In the more perfect specimens
of the smaller pterodactyls in the South Kensington Museum the wing bones are

DfC

•OT.C

FIG. 4. V I E W FROM WING-TIP SIDE OF LEFT PROXIMAL CARPAL BONE. DFC, DFC,
FACETS FOR ARTICULATION OF DISTAL CARPAL BONE.

seen to be retired at the carpal joint through an angle of between 30 0 and 40 0

in the absence of any sign of displacement of the bones. It is probable that in life
the bones could only be retired through a slightly smaller angle.

In order to avoid confusion it is advisable to refer to the part of the wing

https://doi.org/10.1017/S2398187300140290

supported by the metacarpal (Fig. 1) as the " wing-front." The term may also
be applied to the part of the wing near the carpal joint in gulls. In each of these
cases the wing-front probably played a part in wing adjustments similar to that
of the wing-tip of vultures.

Now let us consider the movement at the outer surface of the bone we have
just been discussing. A view of this surface of the proximal carpal bone is shown
in F i g . 4. There are two articular surfaces D F C separated by a ridge. Move-
ment of the outer wing bones at this joint took the form of a rolling movement
 October, 1914] THE AERONAUTICAL JOURNAL 331

in the direction BA. The base of these wing bones may be regarded as equi-
distant from the letters A and B before the movement. After the movement part
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of the base had approached A and part had receded from B. This rolling move-
ment was of very slight extent. One of. the articular surfaces was larger than the
other. It is probable that slight sliding occurred on this larger surface. T h e
first of these movements must have slightly retired and markedly depressed the
wing-front. T h e second must have rotated it upwards.

The movements just described were those between the proximal carpal and
the distal carpal bones (Fig. 2), the so-called " intracarpal j o i n t . "
The movement between the distal carpal and the base of the wing metacarpal
was a simple rotation. As indicated in F i g . 5, on the outer surface of the distal

-MT-

I S C ~~
FIG. 5. END-ON VIEW FROM WING-TIP SIDE OF LEFT DISTAL CARPAL BONE,
P DEPRESSION FITTING PEG ON BASE OF WING METACARPAL (SHOWN IN
FIG. 2). MF, MF, ARTICULAR FACETS FOR ARTICULATION WITH BASE
OF WING METACARPAL. L S C ARTICULAR SURFACE OF LATERAL CARPAL.

carpal there is a depression P which fits a projecting peg on the inner end of the
metacarpal. The articular surface of the metacarpal sliding over the facets M F
caused a rotation of the wing metacarpal round the point P as a pivot. Thus, the
wing metacarpal could rotate round its long axis. T h e permissible movement
must have been of very small extent.

One more movement remains to be noticed, namely, that at the knuckle

joint K, Fig. 1. This was a hinge joint permitting extension and flexion in the
horizontal plane. This was equivalent to advancing and retiring not of the wing-
front but of the wing-tip. There is a projection at the base of the wing phalanx
https://doi.org/10.1017/S2398187300140290

which must have served for the attachment of an " extensor " tendon and which
may have limited the forward movement of the wing-tip. The fully advanced
position of the wing-tip is indicated by the dotted outline P ' in Fig. 1. This
full advancement must have been used for producing " rotation up " round the
transverse axis, since at no other part of the wing could movement occur suitable
for producing this effect. T h e probable position assumed in ordinary slow speed
gliding flight is indicated by the continuous line P . T h e wing-tip could be retired
till it pointed completely backwards or even could be turned in towards the body
of the animal.
 332 THE AERONAUTICAL JOURNAL [October, 1914

The movements of which the different parts of the wing are capable are
summarised in the following table:—
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JOINT. MOVEMENT. REMARKS.

SHOULDER (i) D o w n w a r d s Of wide range

\
ELBOW .. (2) Downwards and
slightly forwards, possi-
bly with slight rotation
downwards of wing-
front

Inseparably
' Fore-arm and (3) Rotation up of wing- connected
proximal carpal front

(4) Large retirement

(about 30 0 ) of wing-
front with slight de-
pression

W R I S T ...< Proximal carpal and (5) Slight retirement with

distal carpal large depression of
wing-front Inseparably
connected
(6) Rotation up of wing-
front

Distal carpal and (7) Rotation up of wing- Slight

i wing metacarpal front

KNUCKLE (8) Advancing and retir- Of wide range

ing of wing-tip

Of these different movements, that at the shoulder joint needs but little
discussion. In view of what is now known about the mechanics of flight, it is
permissible to suppose that movement at this joint was used principally for
flapping, that high-speed flapping occurred with wing-tips retired, and that the
wing-tips were advanced when flapping at slow speed and when starting or
https://doi.org/10.1017/S2398187300140290

ending a flight.
Movement of the elbow joint need not be separately discussed, as it could
only occur in unison with certain movements of the wrist.
As may be seen from the table, the different parts of the wrist joint were
capable of no less than five different movements, some of which were inseparably
connected. W e have to consider whether these movements occurred together or
in groups and what was their function.
It is a rule that if the same movement can occur at two positions in a joint
 October, 1914] THE AERONAUTICAL JOURNAL 333

it will usually occur at these two positions at the same time and in the same
direction. The movements occur in such a way that one reinforces the other. The
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reason for this fact is that " flexor " and " extensor " tendons are usually inserted
not on the bones of a wrist, for example, but pass over it and are inserted on the
bones of the hand beyond the wrist. But in the present case movements (3) and
(4) are inseparably connected with movement of the elbow (2). Hence if the
elbow is kept fixed, movements (5), (6), and (7) might occur independently of
(3) and (4)-
Movements (3) and (4) occurred with, and no doubt were at least to some
extent caused by, movement of the elbow. To understand the probable causation
of movements (5) and (6) it is necessary to refer to Fig. 2. On the underside of
the joint is seen a bone Pt, known as the " pteroid b o n e . " In the later pterodactyls
this is a small tapering splint of a bone that has the appearance of having
represented the insertion of a tendon. 8 As is indicated in Fig. 2, it was attached
t o the lateral carpal bone (by a movable joint), and this again was attached to the
distal carpal. The position of these bones was such that a pull on this supposed
tendon would have caused movements (5) and (6). Of these (6) was a rotation
up of the wing-front. It is not necessary to assume that there was a separate
tendon with similar function attached to the base of the wing metacarpal to
produce movement (7), for as soon as movement (6) had progressed far enough to
allow air pressure on the under side of the wing-front, this pressure would of
itself cause movement (7).
If we admit that the pteroid bone represents a pteroid tendon, we are obliged
to assume that there was another tendon whose pull would reverse the movement
caused by the pteroid. W e may provisionally refer to this supposed tendon as
the " antipteroid." W e have n o means of deciding whether this tendon was
inserted on to the distal carpal or whether running in a curved course over
that bone, it was inserted into the base of the wing metacarpal. In the latter case
its pull would reverse movement (7) besides (5) and (6). In either case, we are
probably safe in regarding movement (7) as merely a continuation of (6), and not
requiring separate consideration.
T h e different movements could have been combined in the following ways :—
A. If the elbow is depressed, then (apart from any effect from the pteroid)
there is—
(3) Rotation up of wing-front.
(4) Slight depression of wing-front (besides the depression due to fore-
arm movement).
(4) Large retirement of wing-front.
B. If elbow remains fixed (and also if it is depressed), the pull from the
pteroid causes—
(6) Rotation up of wing-front.
(5) Large depression of wing-front.
(5) Small retirement of wing-front.
8
The small size of this bone—9.6 centimetres in a pterodactyl of 15 metres span—renders it
unlikely that it supported a propatagium in the manner suggested by one of us. (" The
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Development of Animal Flight," Aeronautical Journal, Vol. XVI., p. 28, January, 1912.) In the
paper here quoted reasons are given (on aeronautical grounds) for believing that the earlier
pterodactyls had four wings, of which the hinder pair were supported by the hind legs. Stromer
has recently stated that one of his museum assistants recollects seeing a beautifully preserved
specimen of the pterodactyl Rhamphorhyncus in the shop of a dealer some ten years ago, and
that this specimen showed a wing membrane supported by the hind leg and passing from the
fifth toe of the hind foot to about half way along the tail. This statement appears to deserve
further investigation, and if the specimen is still in existence, it would be of great interest for it
to be examined and described. Stromer states that the tail fin of Rhamphorhyncus was horizontal,
and not vertical, as has hitherto been supposed to be the case. Thus the only instance that
has been adduced of the use of a vertical fir. by a flying animal turns out to be a myth. (Ernst
Stromer, " Bemeikungen zur Rekonstruktion eines Flugsaurier-Skelettes," Zeitschrift der
Deutschen Geologischen Gesellschatt, Vol. 62, 1910, Berlin, footnote on p. 88.)
 334 THE AERONAUTICAL JOURNAL lOctoier, ion

C. If elbow remains fixed, then the pull from the antipteroid tendon
causes :—
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(—6), (—7) Rotation down of wing-front.

(—5) Large elevation of wing-front.
(—5) Small advancement of wing-front.
Of these combinations, it may be noted that C could only occur in the absence
of A and B. On the other hand, A and B could occur either together or separately.
W e appear to be on fairly safe ground in concluding that the above is a
substantially accurate statement of the facts of the case as to the extraordinary
mechanism of the wrist joint of the pterodactyl. In the case of the human wrist
the " flexor " and " extensor " tendons work independently of rotation and in
whatever position of rotation assumed by the joint. The human wrist is a
joint adapted for many purposes. In the case of the pterodactyl the " flexor "
and " extensor " tendons could only work in connection with and in proportion
to rotation of the wrist. It is a joint adapted to one purpose. In view of the
high degree to which the movements are inter-related, it is impossible to imagine
that any of them are accidental. The movements have been adapted and
specialised for one purpose, and we are obliged to conclude that this purpose
was flight.
Besides the wrist joint, the whole organisation of the pterodactyl shows that
it was fully adapted and specialised for life in the air. But the most extraordinary
inference that we are obliged to draw from the structure of the pterodactyl is
that the muscles for flapping the wings were extremely weak in comparison with
those of birds. The evidence for this fact is the small size of the ridge on the
sternum to which these muscles are attached. If the small size of this ridge was
the only fact we knew about pterodactyls, we might well conclude that they were
land or sea animals that only occasionally rose into the air. But there is no
doubt, from other facts, that they were pre-eminently air-frequenting in their
habits. T h u s the small size of the flapping muscles leads us to suspect, not that
they flew worse than birds, but that they flew more scientifically.
The weakness of the flapping muscles makes it highly probable that their
habitual mode of flight was by soaring, rather than by flapping. The absence
of a wing-tip capable of lifting the wing-front to a large degree, as occurs in
the case of vultures and adjutants, indicates that their soaring could not have
resembled the slow speed circling of these birds of heavy loading. The retirement
of the wing-tip caused the wing of the pterodactyl to have an outline somewhat
like that of gulls and albatrosses. The implied suggestion that their flight was
like that of an albatross agrees well with the little we are able to infer about
their habits. 9
The weakness of the muscles for flapping the wings renders it highly
improbable that they could indulge in " poise-flapping," a form of flight that
requires rapid beats of large amplitude. Thus it is improbable that they poised
in the air and plunged down suddenly on their prey, as does the pied kingfisher.
The light construction of the wing bones and the weakness of the pectoral
https://doi.org/10.1017/S2398187300140290

muscles renders it improbable that they could indulge in any kind of flight that
9
The albatross and certain other of the larger soaring birds resemble pterodactyls more than
do other birds in the lightness of the structure of their wing bones. It is probable that the wing
membrane of the larger pterodactyls was attached to the hind legs. Hence the pull from this
membrane must have tended to draw the legs apart. The muscles that could have opposed this
tendency must have been weak, judging from the curiously small size of the pelvis. This weakness
of the muscles suggests that the wing membrane was extended across from one hind leg to the
other, an arrangement that would diminish the work required from the leg muscles. An analogy
for such an arrangement for the hinder part of tfie' volant membrane is given by the vampire
bats where there is an interfemoral membrane in the absence of a tail capable of aiding in its
support.
 Octoter, 1914] THE AERONAUTICAL JOURNAL 335

requires a sudden change of speed. It is not likely that they could dive through
the air by " shoulder d e s c e n t , " and check speed suddenly in the air in the manner
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employed by vultures. Neither is it likely that they could plunge into the water
as do gulls, in view of the fact that they were unable to furl the wings against
the body, and in view of the possible difficulty of rising again from the water in
any but the calmest weather.
If their flight was habitually like that of the albatross, and therefore at high
speed under certain atmospheric conditions, and if the frailness of their build
renders it improbable that they could check speed suddenly, it is probable that
they could check speed gradually while in the air and while gliding downwards.
Vultures perform this feat (in soarable air) by means of a wing adjustment
described by one of us elsewhere. 1 0 In view of the high speeds obtained in
soaring flight, as compared with the speed obtainable by flapping, it is probable
that this power of checking speed is possessed by all soaring animals. Perhaps
some of the wrist movements of pterodactyls were employed for this purpose.
The complication of the wrist movements of pterodactyls shows strikingly
that the study of animal flight is a complicated problem that we can only hope
to understand fully when we know more than we do at present of the nature of
bird flight and especially of soaring flight.
Our illustrations have been drawn by Mr. G. Howard Short, to whom we
are greatly indebted, from specimens preserved in the British Museum (Natural
History) at South Kensington.
10
" A n i m a l F l i g h t , " p . 121.
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