J. exp. Biol.

133, 33-58 (1987) 33

frinted in Great Britain © The Company of Biologists Limited 1987

GLIDING BIRDS: THE EFFECT OF VARIABLE WING

SPAN

BY VANCE A. TUCKER
Department of Zoology, Duke University, Durham, NC 27706, USA

Accepted 18 June 1987

SUMMARY
1. The equilibrium gliding performance of a bird is described by the relationship
between sinking speed (VB) and air speed (V). When V3 is plotted against V, the
points fall in a 'performance area' because the wing span is changed during gliding.
2. The lowest V, for each V in the performance area defines a 'maximum
performance curve'. This curve can be predicted by a mathematical model that
changes the wing span, area and profile drag coefficient (Crj,rr) of a hypothetical bird
to minimize drag. The model can be evaluated for a particular species given (a) a
linear function relating wing area to wing span, and (b) a 'polar curve' that relates
C D p r and the lift coefficient (C L ) of the wings.
3. For rigid wings, a single polar curve relates C D p r to CL values at a given
Reynolds number. The position and shape of the polar curve depend on the aerofoil
section of the wing and the Reynolds number. In contrast, the adjustable wings of a
laggar falcon (Falco jugger) and a black vulture (Coragyps atratus) gliding in a wind
tunnel have CL, and C D pr values that fall in a 'polar area' rather than on a curve. The
minimum values of C D pr at each C L bound the polar area and define a polar curve
that is suitable for evaluating the model.
4. Although the falcon and the vulture have wings that are markedly different in
appearance, the data for either bird are enclosed by the same polar area, and fitted by
the same polar curve for minimum Co,pr at each CL value. This curve is a composite
of the polar curves for rigid wings with aerofoils similar to those found in avian
wings. These observations suggest that the polar curves of other gliding birds may be
similar to that of the falcon and the vulture.
5. Other polar curves are defined by C L and C D pr values for the falcon and the
vulture gliding at a constant speed but at different glide angles. Each speed has a
different polar curve; but for a given speed, the same polar curve fits the data for
either bird.
6. The falcon and the vulture gliding in the wind tunnel at a given speed were
found to increase their drag by decreasing their wing span. This change increases
induced drag and probably increases C D pr for the inner parts of the wing because of
an unusual property of bird-like aerofoil sections: wings with such sections have
minimum values of C D p r at C L values near 1, while conventional wings have
minimum values of Co tPr at CL values near 0.

Key words: birds, glide polar, gliding performance, polar curve, soaring, wing span.
34 V. A. TUCKER

INTRODUCTION
During equilibrium gliding, all the forces on a glider are balanced; it neither
accelerates nor decelerates. The relationship between a glider's air speed and the
speed at which it sinks through the air completely describes the glider's aerodynamic
performance, and can be predicted from a relatively simple theory for rigid wings
(Welch, Welch & Irving, 1955; Pennycuick, 1975).
Gliding birds, however, change their wing span during flight. Anyone can observe
these birds gliding slowly on fully spread wings, then progressively flexing their
wings as they glide faster and faster. Hankin (1913) called this behaviour 'flex
gliding'. Flex gliding influences stability (Lighthill, 1975) and gliding performance
(Cone, 1964; Newman, 1958; Pennycuick, 1968; Raspet, 1960; Tucker & Parrott,
1970). In this paper, I shall analyse the relationship between lift and drag, for
different wing spans, in birds gliding at equilibrium and use that relationship to
predict maximum gliding performance.

THEORY
Consider a bird gliding at equilibrium at speed V along a flight path inclined at
angle 6 (the glide angle) to the horizontal (Fig. 1). Only two forces have significant
effects on the bird: gravitational force (the bird's weight, W) and aerodynamic force.
Since the sum of these two forces is zero, the aerodynamic force on the bird equals
the weight in magnitude but has the opposite direction.
The aerodynamic force has two components (Fig. 1): total drag (D), the
component Wsin0 that is parallel to the flight path, and lift (L), the component

Sinking speed

Fig. 1. Forces and velocities during equilibrium gliding. The glide path is inclined at
angle d to the horizontal. Lift and drag are perpendicular and parallel, respectively, to the
glide path. Air speed (V) is parallel to the glide path, and sinking speed (V,) is vertical
and downwards.
 Variable wing span in gliding birds 35
Wcos# that is perpendicular to the flight path. Sin0 and cosO can be replaced with
expressions involving the sinking speed through the air (V3) and the air speed (V):
D = WVs/V , (1)
L = W[l - (V s /V) 2 ]*. (2)
The total drag coefficient, C D , and the lift coefficient, C L , are computed by dividing
the force components by 0-5pSV2:
CD = 2D/(pSV 2 ) , (3)
CL = 2L/(pSV 2 ) . (4)
Air density (p) is 1 -23 kg m~3 in this study — the value at sea level in the US standard
atmosphere (von Mises, 1959), and S is the projected wing area, including the area of
the wings intercepted by the body.
The performance of a glider, described by the relationship between V and Vs,
depends on total drag, as can be seen by rearranging equation 1:
V3 = VD/W . (5)
I shall analyse performance by dividing total drag into additive components: induced
drag, profile drag and parasite drag. Many textbooks on aircraft aerodynamics
describe this method in detail (see, for example, Pope, 1951).

Induced drag
Induced drag, D,, arises whenever the wing produces lift. Lift results from a
higher pressure below the wing than above, and this pressure difference induces an
upward flow of air at the tips of the wing and a downward flow along the wing span.
The induced downward flow causes the induced drag on the wing (see Prandtl &
Tietjens, 1957, for a detailed discussion). Induced drag depends on wing span, b
(measured across the widest span of the wings), and is given by:
D| = 2kL2/(;rpb2V2) . (6)
The induced drag factor, k, has a value of 1 for a wing with a straight leading edge
when viewed from the front, constant downvvash and an elliptical distribution of lift
along its span. Actual wings typically have k values in the range of 1 — 1-1 (Reid,
1932), and some wings theoretically may have k values less than 1 (Cone, 1962; van
Dam, 1987). k also corrects the parasite drag of a complete aircraft at low speeds (see
section on Parasite drag below). I shall follow the common practice of using a value of
1 for k when dealing with isolated wings, and a value of 11 when dealing with a
complete aircraft. Spedding (1987) found a value of k of 1 -04 in a gliding sparrow
hawk, Falco tinnunculus; which is consistent with a value of 1-1 after correcting for
parasite drag at low speeds. (Some authors place the induced drag factor in the
denominator of equation 6, in which case it typically has values of 1 or less.)
The induced drag factor of bird wings may change during flight if the wings
Phange their shape and span so that the lift distribution is not elliptical. There are
insufficient data to account for these changes in the present theory, which assumes
36 V. A. TUCKER

that k remains constant during flight. However, the effect of increases in k will be
discussed qualitatively.
The induced drag is related to the induced drag coefficient, given by:
CD,i = 2D,/(pSV 2 ) . (7)
If the right side of equation 6 is substituted for Dj in the equation above, and L is
expressed in terms of CL, one obtains a simple expression for C& ,:

CD,, = kC L 2 /(^A) , (8)

where A is the aspect ratio of the wing, defined as the ratio of wing span to wing
chord. Wing chord is the mean width of the wing, defined as S/b. Therefore,
A = b 2 /S.

Profile drag
Profile drag, D pr , is the drag on the wing in addition to induced drag. It arises
from skin friction and pressure differences and is related to the profile drag
coefficient, given by:
CD,pr - 2D pr /(pSV 2 ) . (9)
The profile drag coefficient varies with the lift coefficient of the wing, and hence
with V. The relationship between Co.pr a n d CL depends on the shape of the wing's
cross-section (the aerofoil section, since the wing is an aerofoil) and the Reynolds
number (Re) of the wing, defined as
/ (10)
c' is the wing chord calculated from maximum wing area (Sma,<) and maximum wing
span (bmaA):
c' = Sniax/bma.x . (11)
H is the viscosity of the air. The ratio p/n has the value 68 436 sm" 2 for air at sea level
in the US standard atmosphere.
At a given Re, CL plotted against Co.pr for a rigid wing yields a polar curve. If the
wing is not twisted along its span and has the same aerofoil section everywhere, the
polar curve is nearly independent of aspect ratio (Prandtl & Tietjens, 1957) and is
characteristic of the aerofoil section.
Co.pr at a given C L decreases as Re increases, with a drop of several-fold over a
narrow range of Re values (Hoerner, 1965; Schmitz, 1960). The midpoint of this
range is the critical Re, which decreases in turbulent air or if the aerofoil surface is
rough. The critical Re for a smooth, bird-like aerofoil section in non-turbulent air is
75 000 (the Gottingen 801 section; Schmitz, 1960; similar to the USATS 4 section in
Fig. 4). Most soaring birds have Re values between 75 000 and 106. C D p r changes
gradually over this range according to the relationship:
F = 1-21 -0-226/tex 10~5 + 0-015\{Re X 10~5)2 , (12)
 Variable wing span in gliding birds 37
where F is the ratio CQ p r /(Co pr at ReX 10"5 = 1). I derived this relationship from
data in Hoerner (1965, fig. 17, pp. 6-13) and Schmitz (1960, fig. 6, p. 171) for a
variety of aerofoil sections at CL = 0-7.
As Re changes, the polar curve shifts right or left without markedly changing shape
(Schmitz, 1960). Thus, the CQ pr vs CL curve can be shifted to any Re between the
critical Re and 106 by calculating a correction term for Co,Pr at a CL of 0-7 and adding
that term algebraically to the CD,pr axis. Equation 12 was used to correct all polar
curves and C D p r values in this paper to Re X10~3 = 1. An example of the correction is
shown in the Appendix.

Parasite drag
Parasite drag, D par , is the drag on the aircraft exclusive of the drag on the wings
and is related to the parasite drag coefficient, given by:
CD,par = 2D par /(pS par V 2 ) . (13)
In flying birds, D par is the drag of the head, body, tail and feet. Spar is the cross-
sectional area of the drag-producing structures.
The parasite drag on an aircraft is commonly expressed in terms of the area of a flat
plate held perpendicular to the airflow. If the drag coefficient of the plate is assigned a
value of 1, a plate of area S par C Dpar has the same drag as the parasite drag of an
aircraft at speed V. The area of the plate is known as the equivalent flat plate area,
S,p:
Sfp = SparCD,par . (14)
If Sfp for an aircraft is known, parasite drag is given by:
D par = 0-5pSfpV2 . (15)
Sfp tends to increase at lower speeds as the tail of an aircraft drops, thereby
increasing the drag of the fuselage and tail. The change in Sfp is small, and it is
common practice to compensate for it by increasing the induced drag factor slightly
(Pope, 1951). Sfp also tends to increase at lower speeds as Re decreases (Goldstein,
1965).

Speed and sinking speed

The total drag on an aircraft is the sum of the three components:
D = D ; + D pr + D par , (16)
so, from equation 5,
V, = V(Di + D pr + D par )/W . (17)
After expanding the drag terms, this equation becomes:
V, = 2kL2/(jrpWb2V) + 0-5pV3(SCD,pr + Sfp)/W . (18)
38 V. A. TUCKER

Models
Equation 18 becomes a model of a glider if one allows b, L, CD pr , Sfpand V to vary
in the same way that they do in the real glider. The simplest model (I shall call it the
'constant span' model) describes a glider with rigid wings. Speed varies, but the wing
span, profile drag coefficient and the other variables on the right side of equation 18
remain constant. If sinking speed is plotted against speed for the constant span
model, a curve (1 shall call it the 'performance curve') results.
The performance curve is often called the 'glide polar', but I shall use the former
terminology to avoid confusion with the polar curves that relate lift and profile drag
coefficients for wings. Performance curves are usually drawn on a V vs V9 diagram
with V9.increasing downwards and V increasing to the right (Fig. 2).
A more realistic model for birds, which I shall call the 'maximum performance'
model, describes gliders with adjustable wings. Birds typically extend their wings to
maximum span when gliding slowly and flex glide at higher speeds. As wing span
changes, so do wing area, induced drag, and the lift and profile drag coefficients.
These changes make the product SCo.pr in equation 18 a function of wing span.
The relationship between sinking speed and speed calculated from equation 18 as
wing span varies need not be a performance curve as is the case with the constant
span model. If, at a given speed, b varies, then sinking speed varies as well. Birds can
also increase sinking speed by lowering their feet to increase Sfp (Pennycuick, 1960,
1968, 19716). As a result, sinking speed can have a range of values at each speed, and

0
A"*~\ * > ^Maximum performance
\ A ^ y ^ curve

20
nking :speed, \

\ \
40

\ . \
on Minimum performance \ *
60 curve ^^

" • ! • • • • • ia

20 40 60
Speed, V (ms" 1 )

Fig. 2. The performance area bounded by performance curves for a falcon, Falcojugger.
Speed for maximum performance is lowest at A, where C|. is maximum at 187. V, = V
along line BC - i.e. descent is vertical at each speed. Vertical descent is slowest at B,
where total drag = profile drag of the fully extended wings with Cn, pr = 1 - i.e. the wings
act as flat plates held perpendicular to the airflow. Vertical descent is fastest at C, where
total drag = parasite d r a g - i.e. the wings are fully folded. Solid lines calculated from the
theory in this paper, dashed lines interpolated.
 Variable wing span in gliding birds 39
the points relating sinking speed and speed fall in an area (Cone, 1964). I shall call
this the 'performance area' (Fig. 2).
The 'maximum performance' model describes the upper boundary of the
performance area. I shall call this boundary the 'maximum performance curve'.
Maximum performance in this sense means that a bird can glide at a particular speed
for the maximum time and distance through the air when it minimizes sinking speed.
If one considers the motion of the bird relative to the earth, the maximum
performance curve shows the minimum vertical wind velocity component required to
keep the bird from losing altitude and, for a given horizontal wind, indicates the
maximum distance over the ground that the bird can cover for a given loss in altitude.
The lower boundary' of the performance area is the 'minimum performance curve',
which describes the maximum sinking speed at each speed. At all but the lowest
speeds, maximum sinking speed occurs in vertical descent when Vs = V and
drag = weight. Vertical descent is slowest when the bird 'parachutes' on extended
wings (Fig. 2, point B), and fastest when the bird folds its wings and dives (Fig. 2,
point C).

The constant span model

In the constant span model, L is assumed to equal W (a constant); the only
variable on the right side of equation 18 is V. The equation for the performance curve
becomes:
V9 = k,/V + k 2 V 3 , (19)
where k] and k2 are constants, defined as follows:
k, = 2kW/(jrpb2) , (20)
k2 = 0-5p(SCD.pr + Sfp)/W . (21)
This model is useful to glider pilots (Welch et al. 1955) because it approximates the
characteristics of man-made gliders with simple wings. Such gliders can fly only at
the combinations of V and V, that fall on a single performance curve because their
rigid wings have lift and profile drag coefficients that fall on a single polar curve. The
glider cannot change its drag, and hence its sinking speed, without changing speed.
[For manoeuvres such as landing, gliders are usually equipped with panels on the
wings that may be raised to increase profile drag. Advanced gliders also have wing
flaps that can change the aerofoil section of the wing during flight — see, for example,
Jacobs (1986). The constant span model does not describe gliders that use these
devices.]
The constant span model has been used for gliding birds (Newman, 1958;
Pennycuick, \91\a,b, 1975, 1982; Blake, 1983). The simplicity of the model is
attractive, but a gliding bird need not hold its wing span and the other variables on
the right side of equation 18 constant at different speeds as the model requires. The
constant span model describes a performance curve that may be wholly within the
performance area, partly within it or wholly outside it, depending on the values
assigned to k) and ki in equation 19.
40 V. A. TUCKER

The maximum performance model

In the maximum performance model, wing span, wing area and the profile drag
coefficient may vary at each speed; their values are chosen to minimize sinking speed
at each speed. Under these conditions, equation 18 describes the maximum
performance curve.
Sinking speed may have a minimum value at a given speed when the wings are
flexed to less than their maximum span. Suppose, for example, that Co.pr were
constant. Then changes in wing span would have opposite effects on two of the drag
terms in equation 17: a reduction in span would increase induced drag but decrease
profile drag by reducing wing area (Newman, 1958; Pennycuick, 1968; Pennycuick &
Webbe, 1959; Tucker & Parrott, 1970). Indeed, a falcon gliding in a wind tunnel at
different speeds has been observed to adjust its wing span to near the theoretical
values for minimum sinking speeds (Tucker & Parrott, 1970).
Co.pr, however, varies with wing span because (1) CQ pr and CL are related by a
polar curve in bird wings as span varies, and (2) CL is related to S (equation 4),
which varies with wing span. To develop the maximum performance model, I shall
represent these two relationships by polynomials:
CD.pr = C 0 + Q C L + C 2 C L (22)
and
S = C 3 b + C4 , (23)
in which CQ, ..., C4 are constants.
The wing span for minimum sinking speed at each speed can be found by making
appropriate substitutions into equation 16, setting the derivative dD/db equal to 0
and solving for b. The minimum sinking speed can then be calculated.
First the terms in equation 16 are expanded:
D = 2kL2/(;rpb2V2) + 0-5pSCD,prV2 + 0-5pSfpV2 . (24)
For economy of notation, some of the constants in equation 24 may be combined into
new constants:
D = k 3 L 2 /(bV) 2 + k4SCD,prV2 + k5V2 . (25)
Before differentiation, CD,pr and S in equation 25 must be replaced with functions of
b- C o p r is a function of CL (equation 22), so
D - k 3 L 2 /(bV) 2 + k4SV2(C0 + C , C L + C2CL2) + k5V2 . (26)
CL is a function of S, so from equation 4
CL = ksL/^SV2) (27)
and

Replacing S with the right side of equation 23,

D = k 3 L 2 /(bV) 2 + Cok4(C3b + C4)V2 + Q k ^ L +
C 2 k 4 (k 6 L) 2 /[(C 3 b + C4)V2] + k5V2 . (29)
 Variable wing span in gliding birds 41
Differentiating,
dD/db = -2k 3 L 2 /(b 3 V 2 ) + C0k4C3V2 - C 2 k 4 C3(k 6 L) 2 /[(C 3 b + C4)V]2 (30)
and, replacing L with W[1-(V S /V) 2 ]* (equation 2),
dD/db = -2k3VV2[l - (V s /V) 2 ]/(b 3 V 2 ) + C0k4C3V2 -
C2k4C3(k6W)2[l - (V 3 /V) 2 ]/[(C 3 b + C4)V]2 . (31)
The value of b that makes the derivative equal to 0 is the wing span (b') for
minimum total drag. Replacing b with b', L with W[l — (V s /V) 2 ]* and D with
WVS/V in equation 25 yields
V9 - k3W[l - (V 3 /V) 2 ]/(b' 2 V) + V3(k4SCD,pr + k s )/W , (32)
where V, is the value of V3 used to calculate b'.
The first chosen value of V3 will probably not be a solution of equation 32 (i.e.
Vs will probably not equal V3), and I used Newton's method (Sokolnikoff &
Sokolnikoff, 1941) to estimate a new value of V3. This value is used to recalculate b',
and the whole process is repeated until values of Vs and b' are obtained that do satisfy
equation 32. The procedure is repeated with different values for V to obtain the
maximum performance curve.
The quantity Co in the above equations varies with Re, and hence with V. As Re
changes, the curve (equation 22) relating Co.pr to CL shifts to the right or left by an
amount related to F from equation 12. This shift is accomplished mathematically by
adding C(F—1) to Co, where C is the value of CD pr calculated from equation 22 for
C L = 0-7. (A computer program that calculates a maximum performance curve,
including Re effects, is available from the author.)

MEASUREMENTS
This section describes the data that will be used to evaluate the maximum
performance model and summarizes how they have been obtained. All the data have
been published, and details may be found in the original publications. Curvilinear
relationships between two variables are described with second-degree polynomial
equations (fitted by least squares) of the form:
y = ao + aix + a2x2 , (33)
where ao, aj and a2 are constants.

Lift and drag

Lift and drag can be determined by measuring W and the glide angle 6, or V, and
V, and then solving equations 1 and 2. In some cases, the measurements have been
made on birds gliding in natural conditions by using tracking devices, mounted
either on the ground or in a sailplane (Raspet, 1960; Pennycuick, 1960, 1971a), to
record the position of the bird relative to the observer.
It is difficult to make accurate measurements in nature for two reasons:
(1) tracking devices of sufficient accuracy are complicated and difficult to operate,
42 V. A. TUCKER

(2) not only the bird must be tracked, but the air as well. Equilibrium gliding
requires an inertial frame of reference relative to which the air does not move. The
frame may move relative to the observer (for example, if there is a wind), but its
movements must be measurable. Indeed, there may not be such a frame, for winds
often change in both speed and direction as a result of gusts and topographical
features (see McGahan, 1973, and Pennycuick, 1971a, for a discussion of these
problems).
The glide angle 9 can be measured more accurately in a wind tunnel than it can in
nature. If a bird is trained to glide freely in a wind tunnel tilted at the glide angle, the
bird remains stationary relative to the observer, and the motion of the air in the wind
tunnel is known. This technique has been used with a pigeon (Columba Iwia;
Pennycuick, 1968), a laggar falcon (Falco jugger, similar in size to the more familiar
peregrine falcon; Tucker & Parrott, 1970) and a black vulture (Coragyps atratus;
Parrott, 1970). These studies will be identified in this paper by references to 'the
pigeon', 'the falcon' and 'the vulture'.

Wing span and area

Wing span and area are changed in living birds during flight. Maximum wing span
(bmax) and area (S max ) can be measured on dead birds by stretching out the wings. At
lesser spans, the relationship between wing span and area was linear in the pigeon,
falcon and vulture gliding in a wind tunnel. This relationship can be estimated for
gliding birds from measurements of b max and Smax (see Appendix).

Equivalent flat plate area

Equivalent flat plate area of a bird can be measured by placing the body on a flight
balance in a wind tunnel (Pennycuick, 1968; Tucker, 1973). Sfp is a function of a
bird's linear dimensions and varies with body mass (Tucker, 1973):
S f p = O-OO334m°-660 . (34)
This equation was obtained from drag measurements made at nearly constant speeds
(11 — 12ms~') on birds with masses between 0-025 and 6-9kg. It therefore includes
Re effects due to body size but not those due to speed. Sfp did not change with speed
in the pigeon. Variations in Sfp with speed are ignored in the present study since the
drag due to Sfp is a small proportion of total drag at the speeds at which birds usually
glide.

P/vfile drag coefficient

Rigid zuings: polar curves for bird-like and conventional aemfoil sections
The profile drag coefficient, C D pr, may be determined by measuring the total drag
of a rigid wing and subtracting the induced drag from it. The total drag can be
measured by mounting the wing on a flight balance in a wind tunnel, and induced
drag can be calculated using equation 6. Alternatively, induced drag may be.
eliminated so that profile drag equals total drag. This is accomplished by mounting,
an untwisted, rectangular wing with a constant aerofoil section in a wind tunnel so
 Variable wing span in gliding birds 43
that it spans the tunnel from wall to wall (Pope & Harper, 1966). The profile drag
coefficient is obtained by dividing profile drag by 05pSV 2 .
The shapes of bird-like aerofoil sections are quite different from the shapes of
conventional ones. Bird wings (for example see Nachtigall, 1985) have highly
cambered upper and lower surfaces at the base of the wing and less cambered
surfaces towards the tip (Fig. 3). Wings of conventional aircraft have lower surfaces
that are nearly flat to convex, as in the Clark Y section (Fig. 4). This section was used
in many aircraft built between 1920 and 1940, and it often appears as an example in
textbooks on low-speed aerodynamics. Its aerodynamic characteristics are well-
known for a range of Re values, and it will be used in this paper as a reference for
comparison with the less conventional, bird-like sections.
There are also differences between the polar curves of bird-like aerofoil sections
and those of conventional aerofoils. Conventional sections have a minimum Crj,pr

Gottingen 462
RAF 19
Gottingen 400
1-8 Eiffel 35
1-6

1-4

1-2

10

0-8

0-6

0-4

0-2

0
002 004 006 008 010 012 014 016
Profile drag coefficient, C D

Fig. 3. Bird-like aerofoil sections and their polar curves. The sections progress (top to
bottom) from a highly cambered section typical of the base of a bird wing to a less-
cambered section typical of the tip of a bird wing. Shown for comparison are the polar
curve for minimum drag (equation 35) for the falcon and the vulture, and polar curves for
three different pigeon wing sections (Xachtigall, 1979). Polar curves have been corrected
to a Reynolds number of 105 and smoothed by fitting points to a second-degree
polynomial equation. Data for the bird-like sections and their polar curves may be found
in the National Advisory Committee on Aeronautics Technical Reports listed in
References. The number of the Technical Report is given in parentheses: Gottingen 462
(286), RAF 19 (93), Gottingen 400 (124), Eiffel 35 (93).
44 V. A. TUCKER

1-6

1-4

JM-2
5 10
'5
| 0-8
§ 0-6

0-4

0-2

0
:i
0 0-02 004 006 008 010 012 014
Profile drag coefficient, C D p r

Fig. 4. The top three aerofoil sections have the same thickness and progressively less-
cambered lower surfaces. The lift coefficient for the minimum profile drag coefficient
decreases as the camber of the lower surface decreases. The bottom aerofoil section
(Clark Y) is typical of those used in low-speed man-made aircraft flying at Reynolds
numbers between 1-5X105 and 4X106. Polar curves have been corrected to a Reynolds
number of 10s and smoothed by fitting points to second-degree polynomial equations
(except Clark Y data, which were fitted by free-hand curve). Data for these sections and
their polar curves may be found in the National Advisory Committee on Aeronautics
Technical Reports listed in References. The number of the Technical Report is given in
parentheses: RAF 19 (93), USATS 4 (93), Durand propeller 13 (93), Clark Y (244).

value at a C L value near 0, and C D p r is nearly constant for C L values between 0 and
1-0 (Clark Y, Fig. 4). Highly cambered bird-like sections such as the RAF 19
(Fig. 3) have a minimum value for C D p r near a C L value of 1, and C D p r increases
two- or three-fold as C L drops towards 0. In addition, the minimum C D p r values for
bird-like sections are higher than that for the Clark Y section.
These differences are related to the amount of camber of the lower surface of the
wing. The more the camber of the lower surface, the higher the C L at which the
minimum C D p r occurs (Fig. 4).
Not all studies of avian wing sections agree with the above description. Nachtigall
(1979) investigated model wings with the same aerofoil sections as those of the pigeon
wing. The polar curves for the model sections were similar to those for conventional
sections in that C D p r was at a minimum near a CL of 0 (Fig. 3). Also, the maximum
Co.pr values for the model sections were several times higher than those for the bird-
like or conventional sections in Figs 3, 4..

Adjustable wings: the polar area for bird wings

The profile drag of a wing attached to a living bird can be determined bj
measuring the total drag of the bird and subtracting induced drag and parasite drag.
 Variable wing span in gliding birds 45
In contrast to rigid wings, CD,pr and C L values for the adjustable wings of living birds
do not fall on a single polar curve. Birds may change wing span, aerofoil sections and
degree of wing twist from base to tip at a given speed. Consequently, their wings have
a range of CD,Pr values for each C L . The points representing this range fall within an
area (I shall call it the polar area) of the C D p r vs C L diagram rather than on a curve.
(Fig. 7 shows the polar area for the falcon and the vulture.)
Bird wings can have a polar curve if the bird glides under conditions that select one
curve from the infinite number that comprise the polar area. For example, the
pigeon, falcon and vulture wings had polar curves when the birds were gliding in
wind tunnels at their shallowest glide angles (i.e. with minimum sinking speed and
drag) for a given speed. These curves are the left-hand bounds of the polar areas, and
I shall call them 'polar curves for minimum drag' (e.g. Fig. 5).
The falcon and vulture also glided over a range of glide angles at a particular speed.
Measurements under these conditions yield polar curves that I shall call 'polar curves
for constant speed'. They will be described after the next section.

Polar curves for minimum drag

The C D pr and C L values for the falcon and the vulture fall on the same polar curve
for minimum drag (Fig. 5) when corrected to an Re value of 105. The equation for
the curve is:
CD,pr = 0-0349 - 0-0781CL + 0-0799CL2 . (35)
This curve is a composite of the polar curves for the bird-like aerofoil sections found
in different regions of the wing (Fig. 3). As the wing span varies, these sections are

002 004 006 008 0-10 012 014

Profile drag coefficient, C D

Fig. 5. Polar curve for minimum drag for the falcon (open circles) and the vulture (filled
circles), corrected to a Reynolds number of 105. C L increases as speed decreases. Data
from Tucker & Parrott (1970, table 2) and Parrott (1970, table 1).
46 V. A. TUCKER

1-8

1-6

1-4

V 1-2

10
Black vulture
0-8 (Raspet)

0-6

0-4 Red-shouldered hawk,

wing
0-2
White-backed
0 vulture Vulture (C D )
-005 0 005 010 015 0-20 0-25
Profile drag coefficient, C D
Total drag coefficient, C D

Fig. 6. Polar curves for minimum drag for various birds and the wing of a red-shouldered
hawk. Also shown are curves relating the lift coefficient to the total drag coefficient for the
falcon and the vulture. These total drag coefficients are less than the profile drag
coefficients alone of the hawk wing and the pigeon at some lift coefficients. All polar
curves have been corrected to a Reynolds number of 105 and smoothed by fitting points to
second-degree polynomial equations. See text for references.

exposed to the airflow to different degrees, so it is not surprising that the polar curve
for minimum drag shares points with the polar curves for the aerofoil sections.
The pigeon's polar curve for minimum drag has CQ pr values that are much greater
than those for the falcon and vulture at high CL values (Fig. 6), comparable at CL
values near 0 5 , and lower at C L values below 0-5. Pennycuick cautions that low CD,pr
values for the pigeon may be unreliable because of uncertainties in measuring
parasite drag.
Using data from gliding vultures, obtained by tracking from sailplanes (Raspet,
1960; Pennycuick, 1971a), presumably with the vultures at minimum sinking
speeds, I have constructed polar curves for minimum drag. The Co.pr values
calculated from Raspet's data for black vultures (see Appendix) are negative (Fig. 6),
probably because the vulture and the sailplane were gliding in different air masses
(Tucker & Parrott, 1970; Pennycuick, 1971a).
Pennycuick (1971a) assumed that C D , pr was independent of CL in African white-
backed vultures and estimated its value to be 00073 at an Re value of 2-9X103.
Correcting this value to Re — 103 yields 0011, somewhat lower than the minimum
Co,pr value of 0018 for the falcon and the vulture (Fig. 6).
The lift coefficients and total drag coefficients for dried bird wings mounted on a
flight balance in a wind tunnel have been measured for several species (Wither.,,
1981). A polar curve constructed from the data (see Appendix) for the one soaring
 Variable wing spat? in gliding birds 47

Table 1. Characteristics of different species used zcith the maximum perfonnance model
W tw, S
2
s,P2
Species (N) (m) (m ) s/s^ * (m )
Falcon 5-60 101 0- 112b + 0019 0-857b/bmM + 0-144 0-00214
Black vulture 17-5 1-37 0- 260b - 0-020 l-060b/b mM —0-060 0-00486J
African white-backed 52-8 2-18 0- 335b-0041+ l-060b/b mtx —0-060 00101
vulture
Fulmar 7-20 109 0- 073b + 0-034 0-700b/b mai + 0-299 0-00270|
Albatross 85-6 3-03 0- 062b + 0-425+ 0-307b/bnMX + 0-693 0-0140t

See text for references.

* Dimensionless relationship between wing span and wing area.
| See Appendix for derivation.
\ Calculated from equation 34.

bird he investigated, the red-shouldered hawk (Buleo lineatus), will be used here.
Although the measurements were made at one speed with a constant wing span, the
results are interesting when compared with those for the wings of living birds
(Fig. 6 ) .
T h e hawk wing had C Q p r values that are greater than the total drag coefficients of
the intact falcon and the vulture. Clearly, the dried wing is not aerodynamically
similar to living wings. Withers attributes the high drag of the dried wing to a low Re
value (0-5 X 10 5 , which is below the critical Re for smooth wings), surface roughness,
fluttering of feathers and wing twist. I n addition, the aerofoil sections of living bird
wings d u r i n g gliding can change markedly from those of anaesthetized or dead birds
(Biesel, Butz & Nachtigall, 1985).
It is remarkable that the same polar curve for m i n i m u m drag describes the data for
both the falcon and the vulture, as these birds are distinctly different in appearance
and size. At m a x i m u m span, the falcon's wings have pointed tips and an aspect ratio
of 7-7 (calculated from data in Table 1). T h e vulture's wings have square tips e n d i n g
in separated primary feathers, and an aspect ratio of 5-6. T h e vulture weighs three
times as m u c h as the falcon.
T h e polar curve for these birds is also similar to curves for rigid wings with bird-
like aerofoils, although it differs from the curve for the gliding pigeon. T h e pigeon is
a flapping bird rather than a gliding bird. Perhaps measurements on other flapping
birds will show that they too have different polar curves from gliding birds.
T h e s e observations suggest that polar curves for m i n i m u m drag may be similar to
equation 35 in all gliding birds with highly cambered aerofoils - a hypothesis that will
be interesting to test. In the absence of polar curves for m i n i m u m drag for gliding
birds other than the falcon and the vulture, I shall use equation 35 to evaluate the
m a x i m u m performance model for gliding birds in general.

Polar curves for constant speed

Both the falcon and the vulture glided in the wind tunnel over a range of glide
angles at a given speed. Data for both birds fell on the same polar curve for a given
48 V. A. TUCKER

1-8

1-6
66ms"1
1-4

(J1 1-2
fficient

10

0-8
§
Lift

0-6

0-4

0-2

002 004 006 008 010 0-12 014

Profile drag coefficient, C D

Fig. 7. Polar curves for constant speed for the falcon (open circles) and the vulture (filled
circles). Points for both birds at a particular speed are connected by lines. (To correct for
slight speed differences between the two birds, C L values for the falcon have been
interpolated from a curve relating V and CL-) At a given speed, the wing span of each bird
decreases as the profile drag coefficient increases. All polar curves are corrected to a
Reynolds number of 10s. Data from Tucker & Parrott (1970, fig. 5) and Parrott (1970,
fig. 2).

speed, but each speed had a different polar curve (Fig. 7). increased and wing
span decreased with increasing CQ pr at each speed.

Wing span and profile drag

How do birds change their wing spans to glide at various combinations of V and Vs
in the performance area? Either an increase or a decrease in wing span could increase
drag, depending on where the bird is flying in the performance area.
The falcon and the vulture when gliding at a given speed in the wind tunnel
invariably decreased their wing spans to increase drag. This change increased
induced drag, but paradoxically it also increased profile drag (as calculated in this
study) in both birds (Fig. 8). One might expect that profile drag would decrease with
wing span because of the reduction in wing area. The lift coefficient increased with
the reduction in wing area but not enough to account for the increase in C D p r
according to the polar curve for minimum drag (Fig. 5).
Pennycuick (19716) offered an explanation for the increased profile drag. African
vultures appear to increase drag when landing by twisting their wings so that the lift
increases at the outer parts of the wing and decreases at the inner parts of the wing.
Consequently, the lift distribution along the wings becomes less elliptical, and the
induced drag factor (k in equation 6) increases. The extra induced drag appears as,
profile drag, since k in this study has a constant value of 1-1.
 Variable wing span in gliding birds 49

10

-, °"9
8-4-9-1 m s " 1
If 0-8
•a
•3 0-7
I 0-6
11-2-14-

04
^
8-4-9-1 m s " 1
o
M 0-3
T3

ll-2-14-3ms
2
a. 01

0 •,
0-6 0-7 0-8 0-9 10
Wing span (m)

Fig. 8. Total drag and profile drag at different wing spans for the falcon gliding at
constant speeds in a wind tunnel. At a given speed, the tunnel was tipped to various glide
angles. The falcon responded by decreasing its wing span to increase drag and sinking
speed. Data for low speeds ( 8 4 and 9 1 ms" 1 ) are similar and are pooled. Data for high
speeds (11 -2, 12-3 and 141 m s " ' ) are also similar and are pooled. Data from Tucker &
Parrott (1970, fig. 5).

Twisting the wings as described above may also increase the actual profile drag of
the inner parts of the wings. These parts have aerofoil sections with highly cambered
lower surfaces. Polar curves for such sections have minimum C D p r values at C L
values greater than 1 (Fig. 3), and CD pr increases two-fold or more as C L drops
towards 0. If the wings are twisted so that the lift, and therefore C L of the inner wing,
drops towards 0, the profile drag of the inner wing will increase. In contrast, birds
flying on the maximum performance curve adjust their wing spans and keep their C L
values high, thereby avoiding the high-drag regions of the polar curves for their
wings.
Man-made gliders with simple wings have lower and lower C L values as they fly
faster and faster along their maximum performance curve. Consequently, designers
of these aircraft select aerofoils that, like the Clark Y, have minimum C D p r values
when CL is near 0 (Fig. 4).

PREDICTIONS
Maximum performance model
Given equation 35, very little additional information about a bird is needed to
predict a curve for maximum performance - only weight, maximum wing span and a
50 V. A. TUCKER

linear function relating wing area and wing span (Table 1). The predictions of the
maximum performance model can be compared with measured performance curves
of birds that presumably were gliding at the minimum sinking speed at each speed.
The falcon and the vulture in the wind tunnel fit this criterion when the wind tunnel
was tipped to the shallowest angles at which they would glide.
Sinking speeds were close to those predicted by the model, and wing spans showed
less agreement with the predictions (Figs 9, 10). The observed sinking speeds would
have been slightly lower if both birds had chosen different wing spans at some
speeds. The falcon usually kept its wings too flexed, and the vulture sometimes kept
its wings too flexed and at other times too spread out.
The predicted maximum performance curve for African white-backed vultures fits
the data of Pennycuick (1971a), presumably obtained at minimum sinking speeds.
Pennycuick used a constant span model to fit an empirical curve to the data (Fig. 11).
However, the data are too variable to discriminate between the two curves.
Pennycuick (1960) estimated a performance curve for fulmars (Fulmarus glaci-
alis) by tracking them while they soared near a cliff top. This curve is below the
predicted maximum performance curve (Fig. 12), suggesting that the fulmars were
not gliding at the minimum sinking speeds for their speeds.
The wandering albatross (Diomedea exulans) is one of the largest gliding birds;
its pointed, high aspect ratio wings are quite different from the wings of the falcon
and the vulture. The maximum performance model predicts poorer performance
(Fig. 12; Table 2) for this bird than has been estimated by others. For example,
Pennycuick (1982) estimated a maximum lift to drag ratio of 23-2 for the albatross,
compared to 14-2 estimated by the maximum performance model. Pennycuick used a

I • 1 1 i 1
c 10 -
C3
O y
Q. OO \
span/

0-75 o \
wing

Oo \
00 B 0-50 - o ON. -
c 3
E
S 0-25 m

E 1
0 |
1 1 | 1 1
1
>r 1 - -
•o"
u 2 *
8. 1
1 \
ink ing;

3 • -

4 \
on \
5 -, 1 1
\ , -
0 5 10 15 20 25 30
Speed, V(ms - ' )

Fig. 9. Predicted maximum performance curve and wing spans for maximum
performance in the falcon. Points represent measurements made at minimum glide angles
in the wind tunnel. Wing span is expressed as a fraction of maximum wing span.
 Variable wing span in gliding birds 51

10 15 20 25 30
Speed, V (ms" 1 )

Fig. 10. Predicted maximum performance curve and wing spans for maximum
performance in the vulture. Points represent measurements made a minimum glide angles
in the wind tunnel. Wing span is expressed as a fraction of maximum wing span.

_l i I 1 1 1 1
1 U

0-75 - -
O. J \
M E
0-50 -

o —
0-25
o
o- 1
1 I 1 0 1
0
T 1 1 o V 1
E -—~^>
o fo j? o
1 o "S J]£P o 9
o"
2 - °o 0 -
O o o

oo E 3 - 0
O °
-
0
4 - \
o

5 i i
10 15 20 25 30
Speed, V (ms" 1 )

Fig. 11. Bottom: predicted maximum performance curve (.1/), empirically fitted
performance curve (E) and measured data points for the African white-backed vulture.
The empirical curve assumes a constant wing span and a constant profile drag coefficient.
Measured data and empirical curve from Pennycuick (1971a). Top: predicted wing spans
for maximum performance. Wing span is expressed as a fraction of maximum wing span.
52 V. A. TUCKER

c 10
I
g1 0-75
Fulmar
| 0-50

| 0-25
0

> 1
Albatross
•T 2
oo E 3
2
4

5
0 5 10 15 20
Speed, V ( m s " ' )

Fig. 12. Predicted maximum performance curves and wing spans in the fulmar and the
wandering albatross. Points represent measurements on the fulmar (Pennycuick, 1960).
Wing spans are expressed as fractions of maximum wing span.

constant span model and a combined drag coefficient of 0-20 for profile and parasite
drag to make his estimate.
The combined drag coefficient is given by the sum Co.pr + Sf p /S, and a value for it
of 0-20 represents much lower profile and parasite drag values than those used in the
maximum performance model. Sfp in the maximum performance model is computed
from equation 34, so Sfp/S for the albatross is 0-023. This value alone is higher than
the combined drag coefficient of 0-20. If Sfp/S is halved to 0-012, then C D pr becomes
0-008 for the constant span model, which is still less than the lowest value of C D p r
(0-011) used in the maximum performance model. Albatrosses certainly look
streamlined in comparison with hawks and vultures and may indeed have better

Table 2. Predictions of the maximum performance model

Speed V Speed
Species (ms- 1 ) 1
* s, mm
(ms" ) (ms" 1 )
Falcon 8 0-83 10 11-9
Black vulture 10 0-97 12 11-6
African white-backed 12 102 14 117
vulture
Fulmar 10 0-95 12 11-5
Albatross 16 117 18 14-2

^8.mini minimum sinking speed for all speeds; occurs at speed shown in column immediately to
the left.
L/Dm^. maximum lift to drag ratio for all speeds; occurs at speed shown in column immediately
to the left.
 Variable wing span in gliding birds 53

0
1
O.
Q

D. 0-2
Q
V M
C—^> L r
,-. 0
1
(A 0
1
•o" 2

i.so 3
c 4
.s 5
10 15 20 25
Speed, V ( m s " ' )

Fig. 13. Comparison of predictions for the falcon by the maximum performance model
(M) and constant span model (C). The constant span model uses a wing span of 101 m
and a CR pr of 002. The bottom panel shows performance curves. Other panels show the
profile drag coefficient (Cn.pr), profile drag (D pr ) and induced drag (D,) at different
speeds.

performance than the maximum performance model predicts. Measurements on a

living albatross gliding in a wind tunnel would resolve the matter.
For all of the birds mentioned above, the model predicts that the wing span should
be maximum for maximum performance at all speeds up to a critical speed. Above
the critical speed, wing span should decrease as speed increases. All the birds
achieved their minimum sinking speed and maximum lift to drag ratio (Table 2) at
speeds below the critical speed, i.e. with their wings at maximum span.

Constant span model

A performance curve predicted by the constant span model has a different shape
from the maximum performance curve. For example, the maximum performance
model predicts that the falcon will have its maximum lift to drag ratio when gliding at
a speed of 10 ms" 1 (Table 2), with values for wing span and C D prof 101 m and 0-02,
respectively. Using these values in the constant span model, the predicted
performance curve is above the maximum performance curve at speeds less than
10ms~' (Fig. 13). Sinking speeds are too low because profile drag is too low
(Fig. 13). At speeds greater than 10ms" 1 , the sinking speeds are greater than those
for maximum performance because wing span, and hence profile drag, is too high.
54 V. A. TUCKER

Induced drag is low, but not low enough to compensate for the higher profile drag
(Fig. 13).
Although the constant span model is not accurate for the entire maximum
performance curve, it can predict a part of it. In the falcon, for example, the
predicted wing span and CD,pr values for maximum performance are nearly constant
at speeds near 10 ms" 1 (Figs 9 and 13, respectively). At these speeds, the constant
span model predicts nearly the same performance curve as the maximum perform-
ance model if one chooses the right wing span and CQ pr values (Fig. 13).

APPENDIX
Raspet's data for the black vulture
Raspet (1960) gives the following data for the black vulture: mass = 2-3 kg,
weight = 22-7 N, b = l-44m, S = 0-365 m2. He also gives sinking speeds (V8) for a
range of speeds (V) for a soaring vulture.
The profile drag coefficient (CQ pr ) before correction to Re X 10~5 = 1 is given by:
CD,pr. = 2 ( D - D , - D p a r ) / ( p S V 2 ) .
The drag terms on the right-hand side of this equation are given, respectively, by
equations 1, 6 and 15. Sfp comes from equation 34.
C D p r ' is corrected to C D p r at /texlCT 5 = 1 by the method described below for the
hawk wing. Reynolds number for Raspet's vulture is 68 426c'V, where c' is the wing
chord, given by S/b.

Withers's data for the red-shouldered hawk wing

Withers (1981) gives the following relationship between CL and Co at
ReXlO'5 = 0-5 and aspect ratio (A) = 3:
C D = 0-085 - 0-153CL + 0-555CL2 .
The profile drag coefficients (C Dpr ') at this Re are given by:
CD,pr' = C D - C L 7(*A) .
To correct C D p r ' to the equivalent profile drag coefficient (C D p r ) at/teXlO"^ = 1,
a term must be subtracted from all C D p r ' values. This term is 0-203(1 —l/F), where
0-203 is the value of C D p r ' at C L = 0-7, and F is computed from equation 12 for
5

Wing span and area for the zdiite-backed vulture

The relationship between wing span and wing area for the African white-backed
vulture was estimated from that for the black vulture, since both birds have similar
wing shapes. The dimensionless relationship between wing span and wing area for
the black vulture (Table 1) is:
x = l-060b/b m a x -0-060.
 Variable wing span in gliding birds 55

The maximum wing area (Smax) of the white-backed vulture is 0-69 m2, and b niax is
2-18 (Pennycuick, 1971a). Multiplying both sides of the equation by Smax and then
substituting values for Smax and b max yields the desired relationship:
S = 0-335b- 0-041 .

Wing span and area for gliding birds

The wings of some gliding birds are very different in appearance from those of the
falcon and the vulture. For example, the wandering albatross has wings with pointed
tips and an aspect ratio twice that of the falcon or the vulture. This section estimates
the relationship between span and area for a simplified, hypothetical wing of any
aspect ratio. The equations are then adjusted to estimate the relationship between
wing span and area for actual gliding birds. For the purposes of this paper, an
estimation will be made for the wandering albatross. First, consider wing span.
The hypothetical wing has three rigid, rectangular elements of lengths rj, r2 and r3
(Fig. 14), connected by joints at the shoulder, elbow and wrist. Two sets of elements
and the body width (be) make up the wing span. The maximum wing span is given
by:
r2 + r3) + b B . (36)
Expressing b^ as a proportion of b max ,
bB = k7bmax , (37)
equation 36 becomes, after substitution and rearrangement,
r 1 + r 2 + r3 = b m a x ( l - k 7 ) / 2 . (38)
If the angles at the wing joints remain equal as the wing flexes at angle a,
b = 2(ri + r2 + r3)sina- + b B (39)
or, after substitution,
= bmax[k7 + (l-k 7 )sinar] . (40)

Fig. 14. Hypothetical bird wing showing changes in wing area with flexing. See text for
explanation.
56 V. A. TUCKER

Now consider the area of the wing as it flexes. The wing loses area HIJK (Fig. 14)
at the wrist joint as the feathers overlap, but it gains area DEFG at the elbow joint as
overlapped feathers are exposed. The areas lost and gained at the wrist and elbow are
equal, so the net loss in area with flexing is due to overlap of the body by the base of
the wing (area ABC). The area lost (S') at the base of both wings is:
S' = c2tan(90 - a) , (41)
where c is the wing chord at the wing base, given by:
c = S max /b max . (42)
In a tapered wing, the chord at the base of the wing is larger than the value given
by the above equation, and the changes in area at the elbow and wrist are no longer
equal when the wing flexes. In addition, a may be constrained anatomically from
reaching 90° in an actual wing. For simplicity, I shall attribute the entire reduction in
area of a flexed, tapered wing to overlap of the body by a wing of chord e c at its base.
The 'taper factor', e, is 1 for a rectangular wing with equal flex angles at each joint.
Actual wings will have taper factors greater than 1, to be determined empirically.
Equation 41 for actual wings becomes:
S' = ec2tan(90 - o<) . (43)
The area of both wings when flexed at angle a is
S = Smax - e (S max /b max ) 2 tan(90 - a) . (44)
The relationship between b and S that results when a range of values for a is
substituted into equations 40 and 44 is approximately linear for b >£b m a x .
The above equations describe the measured relationships between wing span and
wing area for the falcon and the vulture when taper factors are 1*6 and 1-5,
respectively. Since the wing of the wandering albatross appears to be nearly
rectangular near the wing joints, I chose a taper factor of 1-2 for it. Using ky = 0-093,
Smax = 0-611 and b max = 3-03 for the albatross (data from Pennycuick, 1982) yields
the following relationship between wing span and area:
S = 0-062b + 0-425 . (45)

LIST OF SYMBOLS
A aspect ratio C C D , p r a t C L = 0-7, Re = K
a o , a i , ;*2 constants in polynomial Co,.. - , C 4 constants
equation cD total drag coefficient
a angle between humerus and CDi induced drag coefficient
body Co.ps parasite drag coefficient
b wing span CD,pr profile drag coefficient
b' temporary value of b used to
compute V5 cL lift coefficient
bB body width c wing chord at base
"max maximum wing span c' wing chord for maximum
wing span
 Variable wing span in gliding birds 57

D total drag Re Reynolds number

D, induced drag r r r
l i 2> 3 length of wing elements
Dpar parasite drag P density of air
profile drag projected wing area
Dpr
s
e taper factor S' wing area lost owing to flex
F correction factor for ing
Reynolds number Sfp equivalent flat plate area
k induced drag factor ^"max maximum wing area
k,,...,k ^ constants Spar parasite area
proportionality constant,
k7 e glide angle
b B = k 7 b max V air speed
L lift V3 sinking speed
m mass V3 temporary value of V, used
viscosity of air to compute Vs
nJZ ratio of circumference to di-
w weight
ameter of a circle

REFERENCES
BIESEL, W., BUTZ, H. & NACHTIGALL, W. (1985). Erste Messungen der Flugelgeometrie bei frei
Gleitfliegenden Haustauben {Columba livia var. domestica) unter Benutzung neu
ausgearbeiteter Verfahren der Windkanaltechnik und der Stereophotogrammetrie. In BIONA
Report 3, Bird Flight - Vogelflug (ed. W. Nachtigall), pp. 139-160. Stuttgart: Gustav Fischer.
BLAKE, R. W. (1983). Mechanics of gliding in birds with special reference to the influence of
ground effect. J. Biomech. 16, 649-654.
CONE, C. D. (1962). The theory of induced lift and minimum induced drag of non-planar lifting
systems. NASA Tech. Rept R-139.
CONE, C. D. (1964). A mathematical analysis of the dynamic soaring flight of the albatross with
ecological interpretations. Virginia Institute of Marine Science, Special Scientific Report 50,
1-104.
GOLDSTEIN, S. (1965). Modern Developments in Fluid Dynamics, vol. II. New York: Dover
Publications.
HANKIN, E. H. (1913). Animal Flight: A Record of Observation. London: Iliffe.
HOERNER, S. F. (1965). Fluid-dvnamic Drag. (Published by S. F. Hoerner).
JACOBS, D. (1986). The LS-6: one pilot's views. Soaring 50(1), 30-33.
LIGHTHILL, J. (1975). Aerodynamic aspects of animal flight. In Swimming and Flying in Nature,
vol. 2 (ed. T. Wu, C. J. Brokaw & C. Brennen), pp. 423-491. New York: Plenum Press.
MCGAHAN, J. (1973). Gliding flight of the Andean condor in nature, jf. exp. Biol. 58, 225-237.
NACHTIGALL, W. (1979). Der Taubenflugel in Gleitflugstellung: Geometrische Kenngrossen der
Flugelprofile und Luftkrafterzeugung.J. Orn. 120, 30-40.
NACHTIGALL, W. (ed.) (1985). B1ONA Report 3, Bird Flight -Yogelflug. Stuttgart: Gustav Fischer.
NATIONAL ADVISORY COMMITTEE ON AERONAUTICS (1920). Tech. Rept 93, 257-336. Washington:
US Government Printing Office.
NATIONAL ADVISORY COMMITTEE ON AERONAUTICS (1921). Tech. Rept 124, 421-474. Washington:
US Government Printing Office.
NATIONAL ADVISORY COMMITTEE ON AERONAUTICS (1926). Tech. Rept 244, 329-368. Washington:
US Government Printing Office.
NATIONAL ADVISORY COMMITTEE ON AERONAUTICS (1928). Tech. Rept 286, 139-183. Washington:
US Government Printing Office.
PCEWMAN, B. G. (1958). Soaring and gliding flight of the black vulture. J . exp. Biol. 35, 280-285.
PARROTT, G. C. (1970). Aerodynamics of gliding flight of a black vulture Coraqxps atratus.jf. exp.
Biol. 53, 363-374.
58 V. A. TUCKER

PENNVCUICK, C. J. (1960). Gliding flight of the fulmar petrel, jf. exp. Biol. 37, 330-338.
PENNYCUICK, C. J. (1968). A wind-tunnel study of gliding flight in the pigeon Columba livia.
J. exp. Biol. 49, 509-526.
PENNYCUICK, C. J. (1971a). Gliding flight of the white-backed vulture Gvps africanus.J. exp. Biol.
55, 13-38.
PENNYCUICK, C. J. (19716). Control of gliding angle in Ruppell's griffon vulture Gvps ruppellii.
J. exp. Biol. 55, 39-46.
PENNYCUICK, C. J. (1975). Mechanics of flight. In Avian Biology, vol. V (ed. D. S. Farner&J. R.
King), pp. 1—75. New York: Academic Press.
PENNYCUICK, C. J. (1982). The flight of petrels and albatrosses (Procellariiformes), observed in
South Georgia and its vicinity. Phil. Trans. R. Soc. Ser. B 300, 75-106.
PENNYCUICK, C. J. & WEBBE, D. (1959). Observations on the fulmar in Spitsbergen. Br. Birds 37,
321-332.
POPE, A. (1951). Basic Wing and Airfoil Theory. New York: McGraw-Hill.
POPE, A. & HARPER, J. J. (1966). Lowspeed Wind Tunnel Testing. New York: Wiley & Sons.
PRANDTL, L. & TIETJENS, O. G. (1957). Applied Hydro- and Aem-mechanics. New York: Dover
Publications.
RASPET, A. (1960). Biophysics of bird flight. Science 132, 191-200.
REID, G. R. (1932). Applied Wing Theory. New York: McGraw-Hill.
SCHMITZ, F. W. (1960). Aerodynamik des Fluginodells. Duisburg: Carl Lange Verlag.
SOKOLNIKOFF, I. S. & SOKOLNIKOFF, E. S. (1941). Higher Mathematics for Engineers and
Physicists. New York: McGraw-Hill.
SPEDDING, G. R. (1987). The wake of the kestrel (Falco tinnunculus) in gliding flight. J. exp. Biol.
127, 45-57.
TUCKER, V. A. (1973). Bird metabolism during flight: evaluation of a theory, jf. exp. Biol. 58,
689-709.
TUCKER, V. A. & PARROTT, G. C. (1970). Aerodynamics of gliding flight in a falcon and other
birds. J. exp. Biol. 52, 345-367.
VAN DAM, C. D. (1987). Efficiency characteristics of crescent-shaped wings and caudal fins.
iXature, Land. 325, 435-437.
VON MlSES, R. (1959). Theory of Flight. New York: Dover Publications.
WELCH, A., WELCH, L. & IRVING, F. G. (1955). The Soaring Pilot. London: Murray.
WITHERS, P. C. (1981). An aerodynamic analysis of bird wings as fixed airfoils. J. exp. Biol. 90,
143-162.