AIAA JOURNAL

Vol. 52, No. 5, May 2014

Technical Notes
Scaling of Aerodynamic Forces of ThreeDimensional Flapping Wings

II.

K. B. Lua, T. T. Lim, and K. S. Yeo

National University of Singapore, Singapore 117576,
Republic of Singapore
Downloaded by ORTA DOGU TEKNIK UNIVERSITESI on November 14, 2014 | http://arc.aiaa.org | DOI: 10.2514/1.J052730

DOI: 10.2514/1.J052730

I.

Experimental Apparatus and Procedure

The experiments were conducted in the same three-dimensional

flapping-wing mechanism as reported in [13]. Only the essential
features are discussed here.
The mechanism consisted of two wings, and each wing was
subjected to three axes of flapping motions, namely sweeping
motion, elevation motion, and rotational motion (see definition in
Fig. 1). The elevation angle refers to the angle between the
rotational axis and the horizontal plane; the angle of attack refers to
the angle between the wing chord and the horizontal plane; and the
sweeping angle is the angle between the projection of the rotational
axis on the horizontal plane and the line joining the centers of the two
coaxial shafts. The three axes of the flapping motions intersect at the
flapping center, which is located at the center of the gear box.
During the experiments, both the wings and the force transducer
were fully immersed in the center of the water, 1.5 1.5 1.2 m in
volume.
Fluid dynamic forces acting on the wing were measured by an inhouse-developed waterproof force transducer. Verification of the
force sensor was carried out by loading known weights at four corners
and at the center of the wing, and the maximum error was less than 3%
for the range of measured forces. During the experiments, the signal
was sampled at the sampling frequency of 100 Hz. The data were then
low-pass filtered through digital fourth-order Butterworth low-pass
filters with a cutoff frequency of 10 Hz (i.e., 30 times higher than the
maximum frequency of the flapping motions) and then digitally
smoothed with a 21-point mean smooth algorithm. The algorithm
calculated an arithmetic mean of the data points around a chosen
point: 10 points before and 10 points after the selected point. The
smoothing process clarifies but does not alter the trend of the forces.
Each set of the force measurements consisted of 10 flapping cycles
and was repeated 20 times for subsequent ensemble averaging. There
was a 4 min wait between successive runs to allow disturbances to
subside.
Three sizes of geometrically similar hawkmoth-like wings
(extracted from [4]) and fruit-fly-like wings were used (extracted
from [14]). The wings were all fabricated from a 1.5-mm-thick
aluminum sheet. For each wing profile,
to
p


the spans of the smallest
p



the largest wings are respectively 2 times shorter and 2 p
times




longer than that of the middle wing. The choice of the ratio of 2 is
dictated by our desire to test the wings of significant size differences.
Schematic drawings of the hawkmoth and fruit fly middle wings
mounted on the gear box are shown in Fig. 2. Table 1 lists the dimensions of the wing spans Rtip , relative distances from the flapping
center to the wing tip (R), and the location of the second moment of
area Rq
for the three
2

























wings. R2 is calculated using the equation

Introduction

The passion in developing flapping wing micro air vehicles has

attracted considerable attention in the study of unsteady aerodynamics of a flapping wing [13]. To enable sharing and comparing
of research results from different research groups, aerodynamic
forces are often scaled with the square of some reference velocity. In
the case of flapping-wing aerodynamics, complication arises because
velocity varies along the span of the wing since it is executing rotating
flapping motion. Some researchers use the average velocity at the
location of the second moment of area
V
2  (e.g., [46]) as the scaling
velocity. The use of V
2 is influenced by the blade element analysis of
[7,8], which shows that lift/drag under quasi-steady assumption is
proportional to V
22 . Many numerical analysts and experimentalists,
however, use average wing-tip velocity V
tip (e.g., [911]) as the scaling
velocity for unclear reason, probably for convenience.
In many computational studies, the flapping center is located at the
wing base. However, it is very difficult to do the same in experiments
due to the technical constraint of gear boxes and the force sensor
supporting the wing. Moreover, past study has shown that the insect
wing-flapping center may not coincide with the wing base as the
flapping motion is generated by body deformation [12]. Consequently, the virtual flapping center of the wings may be located either
outside or inside the insect body. The question arises as to whether
V
tip or V
2 is a more appropriate scaling velocity to normalize lift and
drag forces so that meaningful comparison can be made with results
from various research groups using different positions of flapping
center.
The desire to address this issue motivates the present investigation.
Force measurements are conducted on rigid wings with hawkmoth
and fruit fly wing profiles. Each wing profile has three geometrically
similar wings of various sizes. Because the absolute distance from the
wing base to the axis of rotation is fixed in the current flapping
facility, the dimensionless distance of the wing base from the flapping

varies with the wing size.
center (normalized by mean chord length c)
This leads to a scenario where the position of flapping center varies
from study to study.

R2  R0 r2 c drS, where S is the wing area, r is the distance from

the flapping center to the blade element, and c is the local chord length.
The experiments were conducted at Reynolds number Re  6012,
10,000 and 17,200; Re is based on the average velocity at the location


of the second moment of wing area (V
2 ) and mean chord length (c).
V
2 is calculated using the equation V
2  R2  I  n , where R2 is the
distance from the second moment of area to the flapping center, I is
the angular distance traveled by the wing for one complete cycle,
which is about twice the stroke amplitude, and n is the flapping
frequency. Re is varied by changing n . The reduced frequency kc is

V
2  cn

R2 In  cR

2 I.
defined as kc  cn

Received 11 April 2013; revision received 4 August 2013; accepted for

publication 21 August 2013; published online 31 January 2014. Copyright
2013 by the American Institute of Aeronautics and Astronautics, Inc. All
rights reserved. Copies of this paper may be made for personal or internal use,
on condition that the copier pay the $10.00 per-copy fee to the Copyright
Clearance Center, Inc., 222 Rosewood Drive, Danvers, MA 01923; include
the code 1533-385X/14 and $10.00 in correspondence with the CCC.
*Research Assistant Professor, Department of Mechanical Engineering, 9
Engineering Drive 1, Block EA, 07-08.

Professor, Department of Mechanical Engineering, 9 Engineering Drive 1,

Block EA, 07-08.

Associate Professor, Department of Mechanical Engineering, 9

Engineering Drive 1, Block EA, 07-08.

III.

Results and Discussion

In the discussion to follow, the results of the sixth flapping cycle,

which have reached a periodic state, are used. Re has insignificant
effect on the force coefficients for the range of Re (i.e., from 6012 to
1095

1096

AIAA JOURNAL, VOL. 52, NO. 5:

TECHNICAL NOTES

Coaxial shafts
Horizontal plane
Elevation motion
Rotational motion

Force transducer

Rotational axis

Downloaded by ORTA DOGU TEKNIK UNIVERSITESI on November 14, 2014 | http://arc.aiaa.org | DOI: 10.2514/1.J052730

Dummy
Gear boxes

Left wing

Right wing

Fig. 1

Schematic of the flapping-wing mechanism with fruit fly wings.

For ease of reference, if U
ref  V
2 , then the lift coefficient are denoted
as CL2 . Alternatively, if U
ref  V
tip , then the lift coefficient is
designated as CLt .
The hawkmoth wings were subjected to hawkmoth hovering
motion (HM, obtained from [4]) and simple harmonic motion
(SHM). The fruit fly wings were subjected to SHM, symmetric
rotation motion (SYR, obtained from [15]) and fruit fly motion (FF,
obtained from [16]). Here, only the results of hawkmoth wing
executing SHM motion and fruit fly wing executing SYR motion are
discussed because the conclusions for the other cases do not vary
significantly. In addition, only the results of the smallest hawkmoth
wing (i.e. HMsmall) executing HM motion are compared with
published results. The numerical values of R, R2 , R2 R, and c
for
each wing and I, n and kc for SHM, HM, and SYR motions for
Re  10; 000 are tabulated in Table 1. Note that the ratio of R2 R
reduces with wing size, which is a consequent of a fixed distance
between the wing base and flapping center.

R
R2

Rtip

a)

A. Hawkmoth Wings Executing Simple Harmonic Motion

R
R2

Rtip

b)
Fig. 2 Schematic of the a) hawkmoth, and b) fruit fly middle wings
mounted on the gear box.

17,200) considered here. Therefore, only the results for Re  10; 000
are shown here. Furthermore, only lift coefficients are shown as the
drag coefficient displays similar trend. The lift coefficient is defined by
CL 

2FL
U
2ref S

Sweeping motion

Location of second moment

of wing area

(1)

The motion profiles of SHM are shown in Fig. 3a, note that the
elevation angle was maintained at constant zero. Figure 3b shows the
CL2 for the three geometrically similar HM wings executing SHM
motion. The CL2 profiles show a slightly skewed sinusoidal pattern,
and the maximum lift is achieved during the second half of each
stroke, after the wing has reached a maximum velocity and started to
slow down, due to the pitching up motion of the wing that begins at
the middle of a stroke [13,17]. It can be seen from Fig. 3b that the
three wings exhibit similar CL2 profiles, although the largest wing
(HMlarge) displays slightly more fluctuations. These fluctuations
were caused by discrete microstepping action of the motors. With the
cutoff frequency of the low-pass filter set at 10 Hz in all cases, any
low-frequency fluctuation below the cutoff frequency would pass
through the filter, and the reverse is true for the higher-frequency
fluctuation. This explains why the other two wings flapping at higher
frequency have smoother force profiles.
In contrast, when V
tip is used as the scaling velocity, there are
distinct differences in the CLt profiles of the three wings (see Fig. 3c);
their magnitudes decrease with increasing wing size.
The above finding indicates that using V
tip to scale aerodynamic
forces causes significant discrepancy in the results. This is of utmost
important when comparing results from various research groups.
Based on the blade-element theory [7,8], the aerodynamic force
acting on a wing element is given by

AIAA JOURNAL, VOL. 52, NO. 5:

Table 1

1097

TECHNICAL NOTES

Wing and motion parameters

Wing type

Rtip , mm

R, mm

HMsmall
HMmiddle
HMlarge

176.78
252.00
340.00

291.38
366.60
456.00

HMsmall

176.78

291.38

FFsmall
FFmiddle
FFlarge

177.50
251.00
353.55

292.10
363.80
466.15

mm I, rad
R2 , mm R2 R c,
kc
n , Hz
SHM motion
195.03 0.67
55.91 4.1888 0.2150 0.2089
229.80 0.62
79.06 4.1888 0.2580 0.1256
273.05 0.60 102.58 4.1888 0.2818 0.0730
HM motion
195.03 0.67
55.91 4.0189 0.2241 0.2180
SYR motion
203.07 0.70
65.49 5.4105 0.1873 0.1803
239.38 0.66
92.23 5.4105 0.2237 0.0790
293.56 0.63 130.43 5.4105 0.2580 0.0620

150

60

Sweeping motion

40

120

(deg)

90

0
0

0.1

0.2

0.3

0.4

-20

0.5
t*

0.6

0.7

0.8

0.9

(deg)

Elevation motion

20

60

-40
-60
Rotational motion

30

-80

a)
Upstroke

Downstroke

HM
Large
HM-large
HM
Middle
HM-middle
HM
Small
HM-small

4.50

3.50

CL2

2.50

1.50

0.50
5.00
-0.50

5.10

5.20

5.30

5.40

5.50

5.60

5.70

5.80

5.90

6.00

5.90

6.00

t*

b)
Upstroke

Downstroke
HM-large
HM
Large
HM-middle
HM
Middle
HM
Small
HM-small

1.80

1.30
CLt

Downloaded by ORTA DOGU TEKNIK UNIVERSITESI on November 14, 2014 | http://arc.aiaa.org | DOI: 10.2514/1.J052730

80

0.80

0.30

5.00
-0.20

5.10

5.20

5.30

5.40

5.50

5.60

5.70

5.80

t*

c)
Fig. 3 a) Motion profiles of SHM motion. b) Transient CL2 and c) CLt profiles for the three sizes of HM wings executing SHM motion.

1098

AIAA JOURNAL, VOL. 52, NO. 5:

dFA  0.5CfA u2 dS

where is the instantaneous angular velocity, and r is the distance

from the flapping center to the blade element. In terms of the force
coefficient CFA for the whole wing, the aerodynamic force generated
is related to V 22 because

(2)

Therefore, the instantaneous aerodynamic force acting on the whole

wing is
ZR
ZR
2
FA  0.5 CfA u dS  0.5 CfA
r2 c dr
0

 0.52

TECHNICAL NOTES

ZR

FA  0.5CFA 2
CfA
r2 c dr

ZR

(3)

r2 c dr  0.5CFA 2 R22 S  0.5CFA V 22 S

(4)

150

80
Sweeping motion
Rotational motion

40

120

Elevation motion

90
0

0.1

0.2

0.3

0.4

0.5

0.6

0.7

0.8

0.9

(deg)

(deg)

20

t*

-20

60

-40
-60

30

-80

a)
Downstroke

Upstroke

3.00
FF
Large
FF-large
FF
Middle
FF-middle
FF
Small
FF-small

2.50
2.00

CL2

1.50
1.00
0.50
0.00
5.00

5.10

5.20

5.30

5.40

-0.50

5.50

5.60

5.70

5.80

5.90

6.00

5.90

6.00

t*

b)

1.40
FF
Large
FF-large
FF
Middle
FF-middle
FF
Small
FF-small

1.20
1.00
0.80
CLt

Downloaded by ORTA DOGU TEKNIK UNIVERSITESI on November 14, 2014 | http://arc.aiaa.org | DOI: 10.2514/1.J052730

60

0.60
0.40
0.20
0.00
5.00
-0.20

5.10

5.20

5.30

5.40

5.50

5.60

5.70

5.80

t*

c)
Fig. 4 a) Motion profiles of SYR motion. b) Transient CL2 and c) CLt profiles for the three sizes of FF wings executing SYR motion.

AIAA JOURNAL, VOL. 52, NO. 5:

Hence, the dynamic pressure defined as 0.5V 22 S is a proper

denominator to nondimensionalize the aerodynamic force of flapping
wings. On the other hand, the V 2tip does not carry these information.
This is the reason why V 2 provides adequate scaling when V tip
does not.

1099

TECHNICAL NOTES

stroke. This can be attributed to the large sweeping amplitude of

77.5 deg and the rotational motion (refer to Fig. 4a, note that the
elevation angle was maintained at constant zero). The minimum
angular distance between the trailing edges in this case is reduced
to less than 10 deg. Lehmann et al. [18] found that the effect of
wingwing interaction is significant when the angular separation
of the two wings is less than 1012 deg. Moreover, the largest wing
(FFlarge) has the least minimum angular distance between the
trailing edges, and its results are most affected by the wingwing
interaction.

B. Fruit Fly Wings Executing Symmetric Rotation Motion

The forces distribution of SYR is complicated by the wing

wing interaction, which occurs at the end and the beginning of each

180

80
Rotational motion
Sweeping motion

150

120

20
0

0.2

0.4

0.6

1 90

0.8

(deg)

, (deg)

40

t*

-20

Elevating motion 60
-40
-60

30

a)
Downstroke

Upstroke

6.00
HM-small
Aono and Liu (2008)

5.00
4.00

CV2

3.00
2.00
1.00
0.00
5.00

5.10

5.20

5.30

5.40

-1.00

5.50

5.60

5.70

5.80

5.90

6.00

t*

b)
Downstroke

Upstroke

2.50
HM-small
Aona and Liu (2008)

2.00

1.50
CVt

Downloaded by ORTA DOGU TEKNIK UNIVERSITESI on November 14, 2014 | http://arc.aiaa.org | DOI: 10.2514/1.J052730

60

1.00

0.50

0.00
5.00
-0.50

5.10

5.20

5.30

5.40

5.50

5.60

5.70

5.80

5.90

6.00

t*

c)
Fig. 5 a) Motion profiles of HM motion. Comparison of transient b) CV 2 and c) CV tip profiles for HM wings executing HM motion.

Downloaded by ORTA DOGU TEKNIK UNIVERSITESI on November 14, 2014 | http://arc.aiaa.org | DOI: 10.2514/1.J052730

1100

AIAA JOURNAL, VOL. 52, NO. 5:

Figure 4b shows the CL2 for the three sizes of FF wings executing
SYR motion. The three profiles show a rather complex and inconsistent trend as the wing size increases. In all cases, the force coefficient
profiles display local peak values at the beginning of a stroke and then
follow by a reduction to a local minimum before increasing to a
constant value for a short period and ending with the second peak at the
finish of the stroke. Although the first peaks of the three profiles
are similar, deviations in their results set in after the minimum values. It
can be seen from the figure that the larger wing produces smaller
magnitude. Also, at the region of relatively constant CL2 , although
both FFsmall and FFmiddle possess similar magnitude, the one
from FFlarge is noticeably smaller. It appears that the wingwing
interaction reduces the lift coefficient, probably by reducing the
strength of the leading-edge vortex, but this required further study. At
the end of each stroke, the last peak of the FFmiddle wing is the
largest, followed by FFlarge and FFsmall. The trends observed
during this period of the cycle are influenced by the combined effects
of wingwing interaction, acceleration, wake capture, and wing
rotation phenomena. It is anticipated that, if the kc and the same
minimum wingwing angle can be maintained for the three wings,
deviation in the results will be much smaller.
As for the CLt profiles, the trend is much clearer, as can be seen in
Fig. 4c. Here, a smaller wing generally results in a larger force coefficient,
except for the peak at the end of each stroke due to the wing rotation.
C. Comparison with Published Hovering Motion Results

Having established from our experiments that V
2 is the appropriate

scaling velocity to normalize aerodynamic forces, it is worthwhile to
compare with the published data in the literatures.
Here, we compare our results of the smallest hawkmoth wing (HM
small) executing HM motion with the computational study of Aono
and Liu [9], which used the same wing profile and flapping motion
obtained from [19]. However, unlike our experiment, their flapping
center was located at the wing base, and the hawkmoth insect body was
included in the computation. Based on the definition used here, their
Re  3214, R2 Rtip  0.52, and kc  0.5696. Aono and Liu [9]
presented their aerodynamic forces in terms of vertical force coefficient
CV and horizontal force coefficient CH . Under hovering condition, the
average CH is zero and the average CV equals the weight of the insect.
The V and H axes are tilted 15 deg from the L and D axes.
Due to space constraints, the comparison is made for the CV data
only. Accordingly, the CV data of [9] were first converted into CV2
and CVtip . This is accomplished by curve-fitting their CV profiles (see
Fig. 4 in [9]), noting that CV was defined as
CV 

FV
0.5URef 2 A

(5)

where Uref  2Rtip n , and is the angular distance traveled by the

wing for one single stroke (only sweeping angle is considered) and
2  4 rad. In our definition, includes the extra distance traveled
by the wing due to the elevating motion, and accordingly 2 is
slightly higher and equals 4.0189 rad. Therefore, CV is related to CVt
for one complete flapping cycle by the following expressions:
CVt  CV
4.018942

(6)

CV2  CVt
0.522

(7)

and

As for our results, they can also be converted into CV2 and CVt by
projecting the forces onto the CV axis that is tilted 15 deg from the
lift axis.
The motion profiles of HM are shown in Fig. 5a, and the comparison of the two sets of results is shown in Figs. 5b and 5c. In
Fig. 5b, the CV2 profiles from the two investigations match
reasonably well in both magnitude and pattern. The only obvious
discrepancy is the second peak in the CV2 profiles. In the case of CVt
(see Fig. 5c), the two studies show significantly different results.

TECHNICAL NOTES

IV.

Conclusions

Experimental studies have been conducted to assess whether V
tip

or V
2 is a more appropriate parameter to scale aerodynamic forces of
flapping wings. The results showed that V
2 is a more appropriate
scaling velocity to nondimensionalize aerodynamic forces because it
gives a better convergence of force coefficients of geometrically
similar wings, compared to using V
tip , which is very much apparatus/
configuration-dependent. This finding is supported by the comparison of our experimental results with the published computational
data. On the other hand, it is also observed that maintaining geometrical similarity without doing the same for kc and the minimum
wingwing angle can lead to a different CL2 due to the influence of
unsteady aerodynamic phenomena, including wingwing interaction, wake capture, and wing rotation.

References
[1] Ho, S., Nassef, H., Pornsinsirirak, N., Tai, Y. C., and Ho, C. M.,
Unsteady Aerodynamics and Flow Control for Flapping Wing Flyers,
Progress in Aerospace Sciences, Vol. 39, No. 8, 2003, pp. 635681.
doi:10.1016/j.paerosci.2003.04.001
[2] Platzer, M. F., Jones, K. D., Young, J., and Lai, J. C. S., Flapping-Wing
Aerodynamics: Progress and Challenges, AIAA Journal, Vol. 46, No. 9,
2008, pp. 21362149.
doi:10.2514/1.29263
[3] Shyy, W., Aono, H., Chimakurthi, S. K., Trizila, P., Kang, C. K., Cesnik,
C. E. S., and Liu, H., Recent Progress in Flapping Wing Aerodynamics
and Aeroelasticity, Progress in Aerospace Sciences, Vol. 46, No. 7,
2010, pp. 284327.
doi:10.1016/j.paerosci.2010.01.001
[4] Liu, H., Ellington, C. P., Kawachi, K., Van Den Berg, C., and Willmott,
A. P., A Computational Fluid Dynamic Study of Hawkmoth
Hovering, Journal of Experimental Biology, Vol. 201, No. 4, 1998,
pp. 461477.
[5] Sun, M., and Tang, J., Unsteady Aerodynamic Force Generation by a
Model Fruit Fly Wing in Flapping Motion, Journal of Experimental
Biology, Vol. 205, No. 1, 2002, pp. 5570.
[6] Birch, J. M., Dickson, W. B., and Dickinson, M. H., Force Production
and Flow Structure of the Leading Edge Vortex on Flapping Wings at
High and Low Reynolds Numbers, Journal of Experimental Biology,
Vol. 207, No. 7, 2004, pp. 10631072.
doi:10.1242/jeb.00848
[7] Weis-Fogh, T., Quick Estimates of Flight Fitness in Hovering Animals,
Including Novel Mechanisms for Lift Production, Journal of
Experimental Biology, Vol. 59, No. 1, 1973, pp. 169230.
[8] Ellington, C. P., The Aerodynamics of Hovering Insect Flight. 1. The
Quasi-Steady Analysis, Philosophical Transactions of the Royal
Society B, Vol. 305, No. 1122, 1984, pp. 115.
[9] Aono, H., and Liu, H., A Numerical Study of Hovering Aerodynamics
in Flapping Insect Flight, Bio-Mechanisms of Swimming and Flying,
Springer, Japan, 2008, pp. 179191.
[10] Dong, H. B., and Liang, Z. X., The Wing Kinematics Effects on
Performance and Wake Structure Produced by Finite-Span Hovering
Wings, 38th Fluid Dynamics Conference and Exhibit, AIAA Paper
2008-3819, June 2008.
[11] Park, H., and Choi, H., Kinematic Control of Aerodynamic Forces on
an Inclined Flapping Wing with Asymmetric Strokes, Bioinspiration
and Biomimetics, Vol. 7, No. 1, 2012, pp. 115.
doi:10.1088/1748-3182/7/1/016008
[12] Dudley, R., The Biomechanics of Insect Flight, Form, Function and
Evolution, Princeton Univ. Press, Princeton, NJ, 2000, pp. 3674.
[13] Lua, K. B., Lai, K. C., Lim, T. T., and Yeo, K. S., On the Aerodynamic
Characteristics of Hovering Rigid and Flexible Hawkmoth-Like
Wings, Experiments in Fluids, Vol. 49, No. 6, 2010, pp. 12631291.
doi:10.1007/s00348-010-0873-5
[14] Dickinson, M. H., Lehmann, F., and Sane, S. P., Wing Rotation and the
Aerodynamic Basis of Insect Flight, Science, Vol. 284, No. 5422, 1999,
pp. 19541960.
doi:10.1126/science.284.5422.1954
[15] Sun, M., and Tang, J., Lift and Power Requirements of Hovering Flight
in Drosophila Virilis, Journal of Experimental Biology, Vol. 205,
No. 16, 2002, pp. 24132427.
[16] Fry, S. N., Sayaman, R., and Dickinson, M., The Aerodynamics of
Hovering Flight in Drosophila, Journal of Experimental Biology,
Vol. 208, No. 12, 2005, pp. 23032318.
doi:10.1242/jeb.01612

AIAA JOURNAL, VOL. 52, NO. 5:

Downloaded by ORTA DOGU TEKNIK UNIVERSITESI on November 14, 2014 | http://arc.aiaa.org | DOI: 10.2514/1.J052730

[17] Lua, K. B., Lim, T. T., and Yeo, K. S., Aerodynamic Forces and Flow
Fields of a Two-Dimensional Hovering Wing, Experiments in Fluids,
Vol. 45, No. 6, Dec. 2008, pp. 10471065.
doi:10.1007/s00348-008-0527-z
[18] Lehmann, F. O., Sane, S. P., and Dickinson, M. H., The Aerodynamic
Effects of WingWing Interaction in Flapping Insect Wings, Journal of
Experimental Biology, Vol. 208, No. 16, 2006, pp. 30753092.
doi:10.1242/jeb.01744

TECHNICAL NOTES

1101

[19] Willmott, A. P., and Ellington, C. P., The Mechanics of Flight in the
Hawkmoth Manduca Sexta. 2. Aerodynamic Consequences of
Kinematic and Morphological Variation, Journal of Experimental
Biology, Vol. 200, No. 21, 1997, pp. 27232745.

R. Gordnier
Associate Editor