J. theor. Biol.

(1987) 129, 263-274

Flight of a Samara, AIsomitra macrocarpa

A K I R A A Z U M A AND YOSHINORI O K U N O

Institute of Interdisciplinary Research, Faculty of Engineering,

The University of Tokyo, Tokyo, Japan
(Received 2 December 1986)
The steady gliding flight of samples of Alsomitra macrocarpa samara was filmed
and analysed. By using the observed data, the flight performance of the samara was
made clear. The lift-to-drag ratio or the gliding ratio was about 3 ~ 4 and the rate
of descent was 0.3 - 0.7 m/sec, which was smaller than those of other rotary seeds.
The flight was so stable that samples were seen to take their optimal trimmed angle
of attack with a value between the maximum gliding ratio and the minimum rate
of descent. The aerodynamic function of the husk for the distribution of the seeds
was also revealed by making wind tunnel tests of the husk.
Nomenclature

AC
AR
b

Co
C~

Ca
CG
CL

G
cm
Cm
C

Ccenter

D
d
g
L
l
M
m
rl

Re
S
U

aerodynamic center
aspect ratio, = b 2 / S
wing-span
drag coefficient, = D/pV2S
minimum drag coefficient
two-dimensional drag coefficient, =d/pVgc
center of gravity
lift coefficient, =L/pV2S
two-dimensional lift coefficient, =l/pV2c
moment coefficient about the center o f gravity, =M/pV2c 2
two-dimensional moment coefficient, = m/~_pVZc2
wing chord
mean wing chord ? =fb/2
~-b/2 cdy/b
wing chord at the wing center
drag
maximum diameter of the husk, two-dimensional drag or spanwise drag
distribution
gravity acceleration
lift
two-dimensional lift or spanwise lift distribution
moment about the center of gravity
mass o f seed, two-dimensional moment
flight number
Reynolds number, = Vcce,ter/V
wing area
wind speed
263

0022-5193/87/230263+ 12 $03.00/0

1987 Academic Press Limited

264
V
W
XAC
O~o

y
A
P

A,

AZUMA

AND

Y.

OKUNO

flight speed
rate of descent
longitudinal position of the aerodynamic center from the leading edge
zero lift angle
gliding angle
swept angle of a line passing through one-quarter chords at wing center
and 3b/8 spanwise station
air density
Introduction

If a seed or a fruit has a film-like wing which is extremely lightweight with a large
surface, and if the center of gravity of the seed is located near but in front of the
aerodynamic center of the wing, then the seed can have a chance to fly by using
the lift generated by the wing in a glider-like translation during the fall, and to
make a long-distant journey in windy conditions. It is very rare to find such samara
which make a gliding flight without any auto-rotation. Many winged seeds such as
maple, black pine, santaloceous, linden, hornbeam, phoenix tree, ash and so on
autorotate during the fall in the wind (Norberg, 1973; McCutchen, 1977; Azuma &
Yasuda, 1985).
The glider-like flight of samara was introduced by Ahlborn (1897) and Hertel
(1963), and its flight mechanics were analysed by Kimura (1943). The present study
makes flight tests of samples of Alsomitra macrocarpa samara, in calm conditions
to specify their performance experimentally and theoretically. In addition, wind
tunnel tests of the husk were conducted to find its aerodynamic function in the
distribution of the seeds.
Construction of Seed and Wing
A pale species from Java, Alsomitra macrocarpa, has a sail wing as shown in
Fig. 1 and performs a stable gliding flight without any tail surface. The geometrical
characteristics of three typical seeds, and the mean and standard deviation of 10
seeds are given in Table 1. The dimensions are defined in Fig. 2.
The seed itself is very thin, about 1 mm in thickness, and is located nearly at the
center of gravity, which is a slightly forward position of the wing center. The wing
is also very thin (from a few ~m to some 10 p.m) and has a swept and tapered plan
form, twisted (washout) angle, reflected trailing edge, and adequately arranged
position of the center o f gravity (CG). An example of the mean camber line is
shown in Fig. 2(c).
By thin wing theory (Abbot & yon Doenhoff, 1959), the reflected aerofoil has
positive moment at positive angle of attack, as given in Table 2, and shifts the center
of pressure backward as the angle of attack increases, and thus has a tendency to
stabilize the pitching motion o f the seed.
The pitching stability is further strengthened by the sweep angle of the wing if
the center of gravity is located in front of the mean aerodynamic center. The washout

265

F L I G H T OF A S A M A R A

FIG. I. A seed of Atsomitra macroearpa.

TABLE 1

Geometrical configuration of seeds ( Alsomitra macrocarpa)

Number of samples
Items
Mass m (mg)
Wing-span, b (cm)
Wing area, S (cm z)
Aspect ratio, AR
Geometrical sweep angle, A ( d e g ) t
Wing loading, mg/S ( N / m 2)
Center of gravity, CG (cm)
(distance from leading edge)
Aerodynamic center, XAC (cm)

Typical examples
1

Mean value
of 10
3

samples

s.d.

264.0
14,1
60-3
3-3
11.3
0.438
1.6

314.1
13,8
66,9
2.8
11.5
0.461
1.8

289.1
14.6
55.3
3.9
21 '6
0.512
1.6

212.1
14.3
59.7
3"5
13.7
0.357
1.6

70.9
0.9
8.7
0,4
3.1
0.115
0.1

2"3

2" 1

2"4

2"2

0-1

+ Swept angle of a line passing through one-quarter chords at wing root and 3b/8 spanwise position.

of the wing and the dihedral, both of which result from the elastic deformation of
the wing during the flight, will prevent spiral instability (Perkins & Hage, 1949).
Many seeds are piled up one over another in a husk, which has a human head-like
form, as shown in Fig. 3, and is hung from a vine. Under windy conditions, the
husk is able to swing by the so-called Karman vortex (von Kfirmfin & Ruback,
1912), and to scatter the seeds in regular sequence. The configuration of the lip of
a hole, which is dug underneath the husk, is such as to introduce the wind inside
the husk, and to help the separation of the flying seeds from the remainder.
After separation, the respective seeds make a gliding flight, with an individual
performance which is determined by their own geometrical configuration and the
wind conditions at that time.

A. AZUMA AND Y. OKUNO

266

Wing span, b

c3./

f,~t....,~

~I~-Cenferof gFavity, CG

~angle,
mic center, AC
(a)

~" c/8 ~" c/8 ~ c/a "1" c/a ~' c/8

c/B "1 c/B "~ c/8 ~

(b)

~L 0'1
:~
_

0.05 /

o.o

\,

(e)

FiG. 2. Geometrical configuration of the seed. (a) Plan view. (b) Frontal view. (c) Profile (nondimensionalized by the semi-chord).

Flight Test of Plain Seeds

By adopting a method o f Rayner & Aldridge (1985), flight tests o f many seeds
of Alsomitra macrocarpa
were, as shown in Fig. 4, conducted in a calm room, and
filmed under a series o f flashed lights o f a stroboscope as shown in Fig. 5. A series
o f positions o f the respective gliding seeds in the space can be determined geometri-

267

F L I G H T OF A SAMARA
TABLE 2
Two-dimensional aerofoil characteristics
ltems

A
B
C
D
E
F
G

Zero lift
angle, a 0
(deg)

Moment coefficient
about the center
of gravity, cm

-0-53
0-28
0-09
-0-05
-0.49
0'08

2-36 x 10-2
3-04x 10-:
2-40 10-2
1-48 x 10-2
0-69 x 10-2
3'31 x 10-2

--

Mean

Aerodynamic
center, XAc/ C

1/4

--

-0.23

2.11 x 10-2

1/4

~~ ~.
I

ta)

(e)
FIG. 3. Husk of AIsomitra macrocarpa. (a) Plan view from top. (b) Plan view from bottom. (c) Side
view from left. (d) Frontal view.

cally f r o m t h e i r p i c t u r e s a n d i m a g e d scales in side a n d r e a r views, t a k e n b y c a m e r a s

(a) a n d (b) r e s p e c t i v e l y . I n the p r e s e n t test, t h e relative p o s i t i o n o f c a m e r a s with
r e s p e c t to the scales is fixed. T h e n the g l i d i n g s p e e d a n d rate o f d e s c e n t a r e c a l c u l a t e d
f o r t h e r e s p e c t i v e intervals. S e e d s t h a t m a d e c u r v e d flights, i n s t e a d o f s t r a i g h t g l i d i n g ,
w e r e e x c l u d e d f r o m t h e p r e s e n t a n a l y s i s . O n l y 10 flights o f t h r e e s e e d s listed in
T a b l e 1 were a v a i l a b l e f o r d a t a a n a l y s i s , b e c a u s e t h e y flew s t r a i g h t in f r o n t o f

268

A. A Z U M A

AND

Y. O K U N O

Camera(b)
Seed(~ S c o /

Flightpath /

Scale(b)

FIG. 4. Experimental apparatus.

the camera. The effect of measured error in position on the gliding p e r f o r m a n c e is

as follows: an error of 1 cm in distance will bring approximately 1% error in the
speed and 2% error in the aerodynamic coefficients.
The flight data and the resulting aerodynamic coefficients are given in Table 3.
In this table, mean values of the respective samples were not a simple ensemble
mean, but a weighted mean in which earlier flights were treated as more important.
That is to say, each d a t u m was multiplied by (1/2) "-~ as the flight n u m b e r (n)
increased, because, as can be recognized from Table 3, the p e r f o r m a n c e of the
respective seeds deteriorated successively with the flight number, because of small
deformations caused by the hard contact with either ground or side walls. The wing
is thin and fragile, but only one flight occurs in nature. The total mean was given
by the ensemble mean of the above weighted means of the respective seeds.
It is clear that the flight data are very scattered, even in the same seed. This
probably resulted from the nonlinear a e r o d y n a m i c characteristics of the wing,
because of the low Reynolds n u m b e r (such as ( 4 - 5 ) x 10 3 based on the mean wing
chord), the nonlinear deformation because of delicate construction of the thin wing,
and of the surrounding atmospheric conditions (such as temperature and humidity),
unnoticeable air currents in the room, and so on.
The mean lift-to-drag ratio of L/D=3.7, at the lift coefficient of CL =0"34, and
the mean rate of descent of w = 0.41 m / s e c show the high performance of the gliding
seed in comparison with the performance of rotary seeds, in which the rate of
descent is of the order of 1 m / s e c (Azuma & Yasuda, 1985).
When a typical seed is, for example, released at the height of 10 m on a calm day
without any wind, it flies with the speed of 1-5 m / s e c for a duration of 24 sec, and
lands at a horizontal distance of 37 m from the point of release.

F L I G H T OF A SAMARA

269

FtG. 5. An example of flight path flashed by a series of stroboscopic flashes. (a) Frontal view. (b)
Side view.

If a horizontal wind, the speed of which is 10 m/sec, exists, then the horizontal
distance o f flight is increased further by 10 m / s e c x 24 sec = 240 m.
It is very interesting to find that the flight is performed at a lift coefficient as small
as CL = 0,34, in the sense of aeronautical engineering. The following test was
performed to make clear why the lift coefficient is so small.

Flight Test of Modified Seeds

In the flight tests of the plain seeds it is impossible to determine the aerodynamic
characteristics o f the wing more precisely. In order to get a polar curve o f the wing,
the trimmed angle of attack must be changed forcedly. This was performed, as
shown in Fig. 6, by installing a thin column at the wing center, and by shifting the

270

A. A Z U M A

AND

Y. O K U N O

TABLE 3

Flight test results

Sample
No.

f
2

3
Total mean

Flight
No., 11

Flight
speed,
V (m/sec)

Rate of
descent,
w (m/sec)

Lift
coefficient, C L

Drag
coefficient,
Co

Lift-todrag rate,
L/D

1
2
3
Mean

1.33
1.34
1-38
1.34

0.38
0.39
0.34
0.38

0.398
0.387
0.372
0.391

0.120
0.119
0-095
0-116

3.33
3.26
3-93
3-40

1
2
3
4
Mean

1.37
1.47
1-51
1-20
1-40

0-28
0.34
0.47
0"35
0.33

0.392
0.337
0.314
0"499
0-374

0-082
0.081
0-102
0-151
0.089

4-79
4.16
3.06
3-30
4-29

1
2
3
Mean

t-69
1,94
2.13
1.82
1.52

0"46
0.58
0"70
0.53
0.41

0-281
0,211
0" t74
0.246
0.337

0-080
0.066
0-061
0-073
0,090

3-53
3-21
2.86
3.34
3.68

FIG. 6. Modified seed. An additional weight can be attached underneath the longitudinal column.

position o f an additional weight on the column. The column and the weight were
so thin and small that their aerodynamic characteristics, except for the trimmed
angle of attack and the resulted gliding speed, were not appreciably changed by
this modification from the original or plain seed.
In the present test, since the same seeds had to be tested several times, the tested
seeds were carefully caught by a soft basket made of a paper whenever they seemed
likely to collide with surrounding obstacles, and to suffer fatal damage.

FLIGHT

OF

271

A SAMARA

I'001

L~

"O~

~ 0.50

0-00

(Mo ,,ie, seeos

0.15

0-30

0.45

0.60

Alsornirnacrocarpa.
tra
Co

FIG. 7. P o l a r c u r v e o f

The data o f flights taking a weakly spiral path were utilized in this test, because
the flight at high lift coefficient showed a slight tendency to spiral instability. The
polar curve of a typical seed is shown in Fig. 7.

Aerodynamic Analysis of Wing

From the flight test data o f three-dimensional wings, the aerodynamic characteristics of the two-dimensional wings should be obtained. This was actually performed
as follows: having assumed a set of airfoil (or two-dimensional) data arbitrarily,
which is assumed invariant along the span, the three-dimensional aerodynamic
characteristics of the wing of known geometrical configuration could be obtained
by applying the local circulation method (LCM, Azuma et al., 1981, 1985). Then it
was easy to find an adequate set of airfoil data by which the three-dimensional wing
characteristics corresponded to the flight data of all tested wings. Shown in Fig.
8(a) and (b) are estimated results o f polar curves o f two- and three-dimensional
wings, and lift-to-drag characteristics of the ensemble mean of the test seeds at a
Reynolds number of Re ~ 4 103. The minimum drag coefficient Coo is about twice
the skin friction drag of a flat surface at the same Reynolds number. The wrinkles
on both surfaces of the wing seem to be unconnected to the drag rise at this small
Reynolds number.
It must be mentioned that the drag coefficient of the two-dimensional wing is
very large in the range of high angle-of-attack. This explains why the gliding flight
is performed at the low lift coefficient as stated before.
Shown in Fig. 9 are the lift-to-drag ratio and the rate o f descent of the threedimensional wing as a function of lift coefficient, which are calculated by the LCM
from the estimated two-dimensional aerodynamic characteristics. The maximum
lift-to-drag ratio and the minimum rate o f descent are respectively given at CL ----0"27
and CL = 0"53. However, the rate of descent is almost constant in the speed range

272

A. AZUMA

AND

Y. O K U N O

1-50

(a)
CLma, = 1.19

/ /

.+_-

2D polor / / /

CL mox " 0'87

/////
/

0-75

u
.J

057
r

0.00

0-15
'

0"3' 0
0"45
Orog coefficient Co, Cd

p-60

'i

1.50

(b)
Ct. max = I'19

. . . . .

0-75

c,~ = o.o38 ; 5.za 2

5"/00

0"00

le :14 (deg)

I 0I . 0

iI

15.0

20'0

Angle of attock, a (deg,)

FIG. 8. E s t i m a t e d c h a r a c t e r i s t i c s o f Alsomitra macrocarpa. (a) P o l a r c u r v e . (b) Ct-a , C a - a c u r v e o f
t h e airfoil.

5-00

LID

._6.=_
2"50
D

Alsomitro,

c, = o.34

0"00

- =

0"25

-- _

0"50

0"75

Lift coefficient, CL

FIG. 9. Performance of the estimated wing.

I'00

FLIGHT

OF

273

A SAMARA

of less than 1.7 m/sec. Therefore the optimal speed is considered to be the speed
of (L~ D)max. The ensemble mean of the lift coefficients in the flight test was CL = 0"34.
If wind is absent, the seed can get the highest distance by flying with a lift
coefficient of CL = 0"27, at which the lift-to-drag ratio and thus the gliding ratio (the
ratio of horizontal distance and the height loss) are maximum. However, if there is
a wind, then the flight is strongly dependent on the time until the seed reaches the
ground after the initiation of falling. This suggests that flight should occur at the
lift coefficient giving the minimum rate of descent. Thus, it is an interesting fact
that the actual lift coefficient of CL = 0.34 lies between two optimal lift coefficients,
the maximum lift-to-drag ratio (or the maximum gliding ratio) and the minimum
rate of descent, but is closer to the former because the minimum rate of descent is
almost constant beyond the selected value, CL > 0"34. Here, also, as seen in many
other locomotions, the living creature acts to get the optimal performance.

Contribution of the Husk

The geometrical configuration of the husk appears to contribute to the dispersal

of the winged seeds. Therefore, the husk was hung in a wind tunnel which had a
circular cross section of 3 m diameter in the test area. From the observation of
smoked flow of paraffin mist around the husk it was revealed that the flow approaching the mouth enters into the inside of the husk and then goes out from the rear
(downstream) part of the hole. Through this excursion of the flow, probably, the
respective seeds are removed from the husk successively.
As the wind speed increased, the husk hung in the wind tunnel was shaken by
the so-called "Karman vortex" series in the wake. Figure 10 shows the frequency
of the wake fluctuation, measured by a hot-wire anemometer, vs. the wind speed
U. The vertical line gives the standard deviation of the frequency increases with the
speed almost linearly, and the slope orthe "Strouhal number" based on the maximum
diameter of the husk, d, is approximately given by S = nd/U ~-0-17. The husk may
therefore be shaken with its own resonant frequency, which is determined by its

20"0

nd

..0- = o.17
p

I0-0
oD"

0-00

3-bo

6-bo

9.bo

12-00

Wind speed, U(m/s)

FIG. 10. Frequencyof the Karmanvortex series.

274

A, AZUMA AND Y. OKUNO

pendulum length. The shaken motion must be helpful for the dispersal of the
seed.

Conclusion
The geometrical characteristics of the wing of Alsornitra m a c r o c a r p a , such as the
slightly swept and twisted wing, the reflected trailing edge of the airfoil, the lightly
loaded wing and adequately arranged C G position, are well fitted to assure the
good performance and stability in gliding flight of the winged seed. The thin wing
with a sharp leading edge and adequate aspect ratio ( A R = 3 - - 4 ) produce the
appropriate lift-to-drag ratio ( L / D ~ 3 - 4) for the flight in small Reynolds number
( R e ~ 4 x 103). The low wing loading ( m g / S ~ 0.5 N / m 2) also guarantees a smaller
rate of descent ( w = 0 . 3 - 0 , 7 m/sec) than those of the rotary seeds. The flight is
performed at a lift coefficient of CL =0.34, which not only gives the maximum
gliding ratio but also guarantees approximately the minimum rate of descent. The
above small lift coefficient is adopted because o f the large drag coefficient at high
angles of attack.
By applying the local circulation method, the two-dimensional aerodynamic
characteristics of the wing section is revealed.
It is also made clear that the dispersal of the seeds is assisted by the wind
surrounding the husk, and the resonant pendulum motion of the hung husk.
The authors would like to express their sincere thanks to the Botanic Gardens of Indonesia
[Lembaga Biologi National L.I.P.I., Kebum Raya Indonesia (Hortus Botanicus Bogoriensis)],
Bogor-lndonesia, for supplying a husk and winged seeds of Alsornitra rnacrocarpa for this
study.
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