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|/. exp. Biol. 137, 221-252 (1988) 221 Printed in Great Britain © The Company of Biologists Limited 1988 FLIGHT PERFORMANCE OF A DRAGONFLY BY AKIRA AZUMA AND TADAAKI WATANABE Institute of Interdisciplinary Research, Faculty of Engineering, University of Tokyo, Tokyo, Japan Accepted 23 February 1988 Summary The dragonfly, Anaxparthenope Julius (Brauer) was observed in free flight, and a theoretical analysis of flight performance at various speeds was carried out. The variation with time of forces and moments acting on wings and body in steady trimmed flight was calculated by the local circulation method. Measures of flight performance, such as top speed, cruising speed and maximum endurance speed, were estimated from a necessary power curve required in steady flight and from the estimated available power. The results show that without using any novel unsteady aerodynamic force generated by a separated flow over the wings, the dragonfly can make steady trimmed flight at various flight speeds, from hovering to top speed. Introduction Application of the local circulation method (LCM) (Azuma et al. 1985) and a numerical computer simulation shows that a dragonfly, Sympetrum frequens, can make steady climbing flight without using an abnormally large lift coefficient (Norberg, 1975) or relying totally on unsteady aerodynamic forces (Savage et al. 1979). This result was obtained by considering the non-uniform and unsteady induced velocity distribution over a pair of stroke planes. The induced velocity can be precisely calculated from the Biot-Savart law which is known in electromagnetic theory and is used to determine the induced velocity distribution generated by wake vortices (see any book of hydrodynamics such as Newman, 1977). In the LCM, the time wise change of the induced velocity at any point in the stroke planes is given by multiplying the attenuation coefficient by the induced velocity generated by any preceding wing at the time that wing passed through that point. In Azuma et al. (1985), the effect of the trailing vortices on the derivation of the attenuation coefficient was introduced, but the effect of the shed vortices was neglected because the reduced frequency, which is a measure of the unsteadiness of the potential flow in the linear range of analysis and is given by k = Jtci/ U = I/AR1//1 in hovering flight, is very small relative to the large-aspect-ratio wing l»f dragonflies (see the next section for all definitions). Key words: dragonfly, flight dynamics, biomechanics, beating wing.
Transcript
Page 1: flight performance of a dragonfly - The Journal of Experimental Biology

|/. exp. Biol. 137, 221-252 (1988) 2 2 1Printed in Great Britain © The Company of Biologists Limited 1988

FLIGHT PERFORMANCE OF A DRAGONFLY

BY AKIRA AZUMA AND TADAAKI WATANABE

Institute of Interdisciplinary Research, Faculty of Engineering,University of Tokyo, Tokyo, Japan

Accepted 23 February 1988

Summary

The dragonfly, Anaxparthenope Julius (Brauer) was observed in free flight, anda theoretical analysis of flight performance at various speeds was carried out. Thevariation with time of forces and moments acting on wings and body in steadytrimmed flight was calculated by the local circulation method. Measures of flightperformance, such as top speed, cruising speed and maximum endurance speed,were estimated from a necessary power curve required in steady flight and fromthe estimated available power. The results show that without using any novelunsteady aerodynamic force generated by a separated flow over the wings, thedragonfly can make steady trimmed flight at various flight speeds, from hoveringto top speed.

Introduction

Application of the local circulation method (LCM) (Azuma et al. 1985) and anumerical computer simulation shows that a dragonfly, Sympetrum frequens, canmake steady climbing flight without using an abnormally large lift coefficient(Norberg, 1975) or relying totally on unsteady aerodynamic forces (Savage et al.1979). This result was obtained by considering the non-uniform and unsteadyinduced velocity distribution over a pair of stroke planes.

The induced velocity can be precisely calculated from the Biot-Savart law whichis known in electromagnetic theory and is used to determine the induced velocitydistribution generated by wake vortices (see any book of hydrodynamics such asNewman, 1977).

In the LCM, the time wise change of the induced velocity at any point in thestroke planes is given by multiplying the attenuation coefficient by the inducedvelocity generated by any preceding wing at the time that wing passed through thatpoint. In Azuma et al. (1985), the effect of the trailing vortices on the derivation ofthe attenuation coefficient was introduced, but the effect of the shed vortices wasneglected because the reduced frequency, which is a measure of the unsteadinessof the potential flow in the linear range of analysis and is given by k = Jtci/U = I/AR1//1 in hovering flight, is very small relative to the large-aspect-ratio wingl»f dragonflies (see the next section for all definitions).

Key words: dragonfly, flight dynamics, biomechanics, beating wing.

Page 2: flight performance of a dragonfly - The Journal of Experimental Biology

222 A. A Z U M A AND T. WATANABE

Two novel mechanisms for lift generation in insects were described by Weis-Fogh (1973,1975). The first one, called 'clap and fling', utilized the separated flowaround a pair of low-aspect-ratio wings (Lighthill, 1973, 1975; Maxworthy, 1979;Edward & Cheng, 1982). The second one, called 'flip', also utilized a pair ofvortices, a bound vortex and a shed vortex of opposite sense, generated by a rapidpronation of the anterior portion of the respective large-aspect-ratio wings.

Savage et al. (1979) studied the role of the vortices generated by a two-dimensional wing motion and unsteady effects on the lift generation and revealedthat the lift is developed during a 'pause' in the downstroke preceding thesupination and during the supination. Somps & Luttges (1985) made an exper-imental test to demonstrate the effect of unsteady separated flows and concludedthat large lift forces are actually produced by unsteady flow-wing interaction.However, we do not agree with this interpretation because of the existence ofphysically unexplained phenomena in the experimental data (see Appendix A).

In this paper, we analyse the free flight of a dragonfly, Anax parthenope Julius,from 16 mm cine films taken in a wind tunnel, to calculate the flight performance atvarious flight speeds by applying the LCM in a form extended to introduce theunsteady aerodynamic effects in a conventional sense, and to show that flight canbe performed without using the unsteady lift generated from the separated flow atleast in steady trimmed flight.

Definitions

AR aspect ratio of wing, AR = b 2 /Sa lift slopeb wing span (m)C D drag coefficient of three-dimensional wingCD f drag coefficient of body other than wingsCD. induced drag coefficientCD|1 profile drag coefficient, CD(1 = C D - CD.C L lift coefficient of three-dimensional wingC d drag coefficient to two-dimensional wing, Cd ( 1~ CD n

C G centre of gravityC/ lift coefficient of two-dimensional wing, Q = aa within unstalled

regionQ.max maximum lift coefficient Q = Q max beyond stallC m momen t coefficient of two-dimensional wing about the feathering or

elastic axisc chord length (m)D drag (N)FH horizontal force, positive forward (N)F v vertical force, positive upward (N)f beating frequency (Hz)/ drag area (m2)

Page 3: flight performance of a dragonfly - The Journal of Experimental Biology

Dragonfly flight 223

G acceleration measured in units of gravity accelerationg acceleration due to gravity, g = 9-80665 m s~2

I moment of inertia of a wing (kg • m2)k reduced frequency, k = jrcf/Uk aerodynamic parameterL lift (N)L body length/ spanwise airloading (N m~')M feathering moment, moment about feathering (elastic) axis (Nm)M C G pitching moment about the centre of gravity (positive for head-up)

(Nm)Mj pitching moment about an apparent joint on the root of the elastic

axis of the wing (Nm)m body mass (kg)mm musculature mass (kg)n nth order of higher harmonicsn load factorP necessary power (W)Pa available power (W)Pi induced power (W)Po profile power (W)P p parasite power (W)Q torque about an apparent joint (Nm)R distance from apparent joint to wing tip, R « b/s (m)Re Reynolds numberr spanwise station along the elastic axis (m)S wing area (m2)Sf maximum sectional area of body (m2)T period of one beating cycle (s)t time (s)u VuT

2 i Up2

UT tangential component of velocity relative to a blade element (ms~')Up perpendicular component of velocity relative to a blade element

(ms-1)V flight speed (m s ')v induced velocity (ms~')W weight of body = mg (N)x non-dimensional spanwise station, x = r /RxCG longitudinal distance of centre of gravity measured from front joint of

forewing (m)Xj distance between the first joints of forewing and hindwing

vertical distance of centre of gravity measured from hindwingangle of attack of wing section (degrees or radians)

/3 coning angle (degrees or radians)

Page 4: flight performance of a dragonfly - The Journal of Experimental Biology

224 A. AZUMA AND T. WATANABE

(3* beating angle projected in horizontal plane (degrees)

F flight angle (degrees or radians)y tilt angle of stroke plane (degrees or radians)d^,S2,A aerodynamic parameters related to the wing planformdy phase shift of flapping motion (degrees or radians)cL phase shift of feathering motion (degrees or radians)

0d

pAc/2

ViIJJ*

(X)

0

1n0-75RAcruisefhhovImaxminP,minroottip

body attitude (degrees)feathering angle (degrees or radians)air density (kgm~3)swept angle to half-chord line (degrees or radians)inflow angle (radians)azimuth angle or flapping angle (degrees or radians)amplitude of beating motion (degrees or radians)beating angle projected in vertical plane (degrees)angular velocity of beating motion, co = lizi (radians s~l)

Subscript or superscript

oth harmonic component (or steady state) or two-dimensional valuesthe first harmonic componentnth harmonic componentthree-quarter radius stationaerodynamic componentcruising flightforewinghindwinghovering stateinertial componentmaximum valueminimum valuepower minimumwing rootwing tip

Geometrical configuration

The general configuration of Anax parthenope Julius is shown in Fig. 1 andgeometrical characteristics of two dragonflies are given in Table 1. This dragonflyis considered to be one of the 'high performance' species: it has excellentmanoeuvrability and is a most active predator. It has a large wing load and highbeating frequency, enabling fast and skilful flight.

The beating can be represented both as the flapping (or heaving) motion of anelastic axis assumed to be a straight line roughly passing through a quarter chord ^the aerofoil section at any spanwise station and as a feathering (or pitchinJPmotion about the elastic wing axis. These motions are performed actively through

Page 5: flight performance of a dragonfly - The Journal of Experimental Biology

Dragonfly flight

Table 1. Geometric characteristics of two dragonflies

225

Body lengthMassWing span: forewing

hindwingWing area: forewing

hindwingAspect ratio: forewing

hindwingWing loadingCentre of gravity

Distance betweenthe first joints offorewing and hindwing

Estimated drag areaDrag coefficientMaximum section area

Abbreviation

Lmbf

bh

Sf

Sh

ARf=(bf)2/Sf

ARh = (bh)2/Sh

W/(Sf+Sh)XCGZCG

X.i

f=SfCDf

Sf

Units

m

kgmmm2

m2

Nm-2

mm

m

m2

m2

DragonflyA

7-5xlO~2

7-9X10-4

1-OxlO"1

9-7xlO~2

1-OxlO"3

1-2X10"3

107-83-53-OxMT3

6-OxlO"3

8-OxlO"3

9-4xlO~5

1-257-5X1Q-5

DragonflyB

8-OxlO"2

7-9X10'4

1-2x10"'1-lxlO"1

9-3 x 10"*l-3xKT3

15-79-33-53-OxlO"3

5-OxlO"3

8-OxlO"3

9-4xlO"5

1-257-5X10"5

two (front and rear) joints at the respective wing root (von Lendenfeld, 1881). Asshown in Fig. 1A,B, the flapping is assumed to be confined within a conical plane,the apex of which coincides with the front joint and the coning angle of which isdefined by fl, although the actual flapping motion deviates slightly from the conicalplane during the stroke (Fig. 2). The orbit of the three-quarter radius station of theelastic (or feathering) axis is called the 'stroke plane', which is considered to benormal to the cone axis.

In dragonflies, the cones related to the forewings are open towards the frontwhereas the cones related to the hindwings are open mainly towards the back.Therefore, the stroke planes of the forewings are located in front of the joints ofthe forewings and the stroke planes of the hindwings are located behind the jointsof the hindwings. The tilt angles of the two stroke planes are defined by yf and yh.

Mode of wing beat

Free dragonfly flight was observed in a wind tunnel and filmed with a 16 mmhigh-speed cine camera. By changing the wind speed in the tunnel the flight speedwas altered. To get a clear image of the feathering motion and the twistdistribution of the wings, three parallel stripes (lmm in width) were painted oneach left wing at three spanwise stations (Fig. 1A). The width and thus the mass ofthe stripes were so small that the beating mode and frequency were substantiallyunchanged by them.^ Observed modes of flight are given in Table 2. The orbits of the wing tips withPspect to the body, which roughly show the stroke planes of the respective wings,and the movements of the wings (measured at the three-quarter span position) in

Page 6: flight performance of a dragonfly - The Journal of Experimental Biology

226 A. AZUMA AND T. WATANABE

Inclination

Painted stripes ^ o y \ ^ stroke plane

U

Flight path angle

Body attitude

Stroke plane

T i p , p a t h Coning angle

Flapping amplitude

Wing position

Fig. 1. Schematic configuration of Anax parthenope julius and its stroke planes.(A) Side view of dragonfly in flight; (B) projection of stroke plane (view A).

inertial space, are shown in Fig. 2A-D for various flight speeds, V. The traces ofthe azimuth angles, 1/ and 1//1, which are defined by the flapping angles of theelastic axis projected to the stroke plane and measured from the horizontal line,are shown in Fig. 3A-D. They can be expressed by the first harmonic of a Fourierseries:

\p = xpo + ip\cos(cot + dy,). (1)

The values in the above and the following equations refer to either the forewing c*mthe hindwing.

However, as can be seen from Fig. 4A-D, the feathering angles at three span

Page 7: flight performance of a dragonfly - The Journal of Experimental Biology

Dragonfly flight

Table 2. Flight kinematics for dragonflies A and B; experiments 1-4

227

DragonflyExperiment

VelocityFlight angleBody attitudeBeating frequency

Stroke plane inclinationmeasured from horizontal line

forewinghindwing

measured from body axisforewinghindwing

Flapping amplitudeforewinghindwing

Phase difference of flappingmotion between forewing andhindwing

Phase difference between flappingand feathering

forewinghindwing

Calculated load factor

V(ms~')F (degrees)0 (degrees)f(Hz)

y1 (degrees)yh (degrees)

y' + 0 (degrees)yh + 0 (degrees)

i// (degrees)xjjh (degrees)

<5h (degrees)

01.O-75R

<5| ~~ 0l\o-75R

n

A1

0-7-12

2026-5

4038

6058

3626

51

89102

1-1

A2

1-5-1-11228-1

5548

6760

2526

61

9392

0-97

A3

2-34-84

29-0

5852

6256

2526

61

9195

1-25

B4

3-202-0

27-0

6368

6570

3834

93

8981

105

positions x = 0-25, 0-5 and 0-75, are expressed by the Fourier expansion seriesincluding higher harmonics as follows:

= 60+ 0ncos(n<wt + 8e) (2)

Table 3 shows the values of coefficients in the above Fourier expansion series ofthe experimental data. The first harmonic of the feathering angle of the respectivewings is, as can be seen from Fig. 5A-D, a function of the span position x,

01 = (0O-75R " 0root)(x/O-75) + 0root , (3)

where the coefficients, 0O-75R ~~ dTOO{ a nd 0root are also functions of the flight speed.

From these figures it can be seen that: (i) the beating frequency of the wings isalmost unaltered (f = 29-32Hz) with changes in flight speed; (ii) the tilt angle ofthe stroke planes with respect to the body axis gradually increases from about 40°to 70° as the flight speed increases; (iii) the phase difference between fore and hindpairs of wings is within 60°-90° and is not correlated with flight speed; (iv) theconing angle is about 8° in the forewing pair and about —2° in the hindwing pair

Page 8: flight performance of a dragonfly - The Journal of Experimental Biology

228 A. AZUMA AND T. WATANABE

Forewing . Root of wing

Body axis

Root of wing

Forewing

Vs" Root of wing

Body axis

Hindwing /

48 c

Root of wing

Fig. 2. Orbit of wingtip and the movements of the wings, measured at the three-quarters span positions. V, downstroke; A, upstroke; dashed line shows orbit of wing;left-hand plots show wing movements relative to the air; right-hand plots show wingmovements relative to the body. (A) Experiment 1 (V = 0-7ms~1, dragonfly A);(B) experiment 2 (V = l-5ms~J, dragonfly A); (C) experiment 3 (V = 2-3ms~\dragonfly A); (D) experiment 4 (V = 3-2ms~1, dragonfly B).

Page 9: flight performance of a dragonfly - The Journal of Experimental Biology

Dragonfly flight 229

Forewing

\S

A58"• Root of wing

Body axis

Hindwing

L_

52c

Root of wing

D

Forewing

• Root of wing

Body axis

Hindwing

65c

Root of wing

Fig. 2C,D

Page 10: flight performance of a dragonfly - The Journal of Experimental Biology

A. AZUMA AND T. WATANABE

Downstroke of forewmg Upstroke of forewingUpstroke of forewingDownstroke Upstroke of hindwing DownstrokeUpstroke of hindwing Downstroke

90

60

30

0

- 3 0

- 6 0

-90

Downstroke of forewing Upstroke of forewingDownstroke^ ^ Upstroke of hindwing JDownstroke

Forewing

THindwing

= 3-4xlO"2s

0 0-25T 0-5T 0-75T

90

60

30

0

-30

-601

Time

Downstroke of forewing Upstroke of forewingDownstroke, Upstroke orhindwing .Downstroke

= 3-6xl(T2s

0-25T 0-5T 0-75T

Fig. 3. Azimuth angle in the stroke plane for various flight speeds. O, forewing; <Xhindwing; lines are given by equation 1. (A) Experiment 1 (V = 0-7ms~', dragonflyA); (B) experiment 2 (V = 1-5ms~', dragonfly A); (C) experiment 3 (V = 2-3ms~',dragonfly A); (D) experiment 4 (V = 3-2m s"1, dragonfly B).

and is almost constant (within 10 % deviation) throughout the beating motion; (v)the flapping amplitude ranges from 25° to 40° and is not correlated with flightspeed; (vi) the feathering amplitude ranges from 40° to 60° in the forewings andfrom 30° to 40° in the hindwings and is also not correlated with flight speed; (vii)the phase difference between the flapping and feathering motion is 90°, which isconsidered to be optimal for efficiency at low beating frequency (Azuma, 1981);and (viii) since the beating motion is performed through two joints at therespective wing roots, in the feathering motion the wing is twisted linearly for agiven time - wash-out (twisted negatively towards the tip) in the downstroke andwash-in (twisted positively towards the tip) in the upstroke.

The wing beat modes were also observed in a smoke tunnel using a stroboscopicflash which made it possible to visualize the wake vortices of the beating wings andthe modes of the wing motion. The wake vortices were visualized by the paraffinmist method (Watanabe et al. 1986) (Fig. 6A,B). A series of trailing and shedvortices generated, respectively, by the span and time (or azimuthal) change of

Page 11: flight performance of a dragonfly - The Journal of Experimental Biology

180

150

120

90

60

30

0

Downstroke _j

0O-5R

0Q-75R

Upstroke , _0 '

O0-25R ^ ;

Forewing

T = 3-6xlO"2s

Dragonfly flight

180

150

120

231

Upstroke Downstroke

0 0-25T 0-5T 0-75T

Downstroke Upstroke 10U

150

120

90

G 601<u<u

£? 30

% n

Downstroke f Upstroke ,

8 ^ £R Hindwing

T = 3-6xlO~2s

0-25T 0-5T 0-75T

Downstroke Upstroke

#0 5R

T

, Downstroke

Hindwing= 3-4xlO-2s

0-25T 0-5T 0-75T

Downstroke Upstroke .Downstroke

Hindwing= 3-65xKT2s

0-25T 0-5T 0-75TTime

Fig. 4. Feathering angles with respect to the stroke plane. Span position, O, 0-75R;<0>, 0-50R; A, 0-25R; lines are given by equation 2. (A) Experiment 1 (V = 0-7ms"',dragonfly A); (B) experiment 2 (V = l-5ms~\ dragonfly A); (C) experiment 3(V = 2-3ms~\ dragonfly A); (D) experiment 4 (V = 3-2ms~', dragonfly B).

Page 12: flight performance of a dragonfly - The Journal of Experimental Biology

to to

Tab

le 3

. C

oeff

icie

nts

in F

ouri

er e

xpan

sion

ser

ies

of f

eath

erin

g m

otio

n ba

sed

on e

quat

ion

2

Dra

gonf

lyE

xper

imen

t

Fore

win

gA

mpl

itude

(de

gree

s)(0

-25R

/0-5

R/0

-75R

)

Phas

e (d

egre

es)

(0-2

5R/0

-5R

/0-7

5R)

Hin

dwin

gA

mpl

itude

(de

gree

s)(0

-25R

/0-5

R/0

-75R

)

Phas

e (d

egre

es)

(0-2

5R/0

-5R

/0-7

5R)

0o 0i 02 03 04 0i 02 03 04 00 01 02 03 04 01 02 03 04

91 24 5 3 2 95-1

24 -33 77 103 19 5 2 2

-163

-144

-107 -1

5

A 1 95 36 5 7 4 87-8

6 12 27 105 27 9 1 1

-169

-131 14

8-1

04

100 54 2 7 1 89

-88 52 -7 107 38 6 2 1

-178

-112 -3

7-2

16

86 25 4 4 1 77 144

-77 17 85 9 3 0 0

146

-140 10

-96

A 2 90 36 4 4 0 89 137

-65

-165 84 23 4 2 2

154

-100 -9

819

7

88 54 4 4 5 93 135

-68

113 81 30 6 2 2

154

-69

-52 83

77 14 3 1 3

104

-92

148

-68 83 10 3 0 4

147

-67

-106

-154

A 3 82 291 3 1 88 160 65 -3 82 22 9 4 2

163

-71

-86

-159

81 46 6 3 4 91 154 45 106 79 24 11 2 2

-164 -8

1-6

616

9

72 17 3 3 2 94-9

0 6 62 73 23 4 3 1

164

-158 10

9 58

B 4 79 30,

5 4 3 99 82 112

-101 74 30 3 2 3

167

-91

-25 50

83 48 6 4 4 99 98 142

-117 76 41 3 3 3

174

-88

-10 53

N C > D H H Z

Page 13: flight performance of a dragonfly - The Journal of Experimental Biology

Dragonfly flight 233

0 0-25 0-5 0-75 1

0 0-25 0-5 0-75 1 "0 0-25 0-5 0-75 1Span position

Fig. 5. Spanwise variation of feathering amplitude. First harmonic of observed data:O, forewing; A, hindwing; lines are given by equation 3. (A) Experiment 1(V = 0-7ms~', dragonfly A); (B) experiment 2 (V = l-5ms"1, dragonfly A); (C) ex-periment 3 (V = 2-3ms~1, dragonfly A); (D) experiment 4 (V = 3-2ms~\ dragonflyB).

bound vortices are clearly observed in wavy wake sheets of the respective wingpairs.

Aerofoil characteristicsTo determine the aerodynamic characteristics of the dragonfly wing aerofoil,

flight tests were conducted in a calm room using model gliders which had a mainwing composed of a pair of dragonfly hindwings, as shown in Fig. 7A,B. It wasnecessary to introduce a small swept angle for the main wing and to install a tailwing for steady gliding flight with a positive angle of attack.

Fig. 8A-C shows plots of data obtained for the main wings of the model glidersin which the trimmed angle of attack was changed by shifting the centre of gravitywith respect to the aerodynamic centre of the wing. The data for the gliders with a

wing, after removal of the contribution of the tail wing, are represented byircles and the data for the gliders without a tail wing are represented by triangles.

The data for which the angle of attack were measured are shown by solid symbolst

Page 14: flight performance of a dragonfly - The Journal of Experimental Biology

234 A. AZUMA AND T. WATANABE

Fig. 6. Wake pattern and embedded vortices. By changing the positions of smokefilaments two types of vortex images can be visualized; the rectilinear trace across thetrails is a supporting frame for the dragonfly. (A) Flow pattern induced mainly by thetrailing vortices; (B) flow pattern induced mainly by the shed vortices.

Page 15: flight performance of a dragonfly - The Journal of Experimental Biology

Dragonfly flight 235

Hindwing of dragonfly

S

Hindwing of dragonfly

10

Balance weight 120

Balance weight

Fig. 7. Model gliders using dragonfly wings. Numbers indicate lengths in mm.(A) Tail-less wing; (B) tailed wing.

in Fig. 8A-C. The solid curves in Fig. 8 show typical characteristics of these three-dimensional wings.

Utilizing the three-dimensional wing data, it is possible to determine theaerodynamic characteristics of the two-dimensional wing and the section aerofoilcharacteristics from simple wing theory (Diederich, 1952; Hoak & Ellison, 1968),as follows:

lift slope, a0 = (a/cosAc/2)/Vl - (2a/^AR) , (4)

induced drag coefficient, CD. = (CL2/^AR)(1 + (5l(52) + kA ,

profile drag coefficient, CD|) = CD — CD. ~ Cdo,

(5)

(6)

where AR, Ac/2 and a are the aspect ratio, the swept angle of the half chord line,and the lift slope of the three-dimensional wing, and <5ls <52, k and A areparameters related to the planform of the wing and are presented by charts ofDATCOM (Hoak & Ellison, 1968).

Fig. 9A shows the dragonfly aerofoil characteristics (or the aerodynamiccharacteristics of the two-dimensional wing, extrapolated from the above obtaineddata to hypothetical values in a large angle of attack range) as a function of angleof attack, in which the maximum lift coefficient is assumed to be Q max = 1-2. It is

Rot clear why the minimum drag coefficient is slightly lower than those expectedom the skin friction of a plate. Fig. 9B compares the data for polar curves for a

locust wing, fruit-fly wings and a dragonfly wing obtained by Jensen (1956), Vogel

Page 16: flight performance of a dragonfly - The Journal of Experimental Biology

236 A. AZUMA AND T. WATANABE

u

0-1 0-2 0-3Drag coefficient, CD

0-4

10

0-5

<C 0

- 0 - 5

-1-0

B

-20 -10 10 20 -20 -10Angle of attack, a- (degrees)

10 20

Fig. 8. Aerodynamic characteristics of three-dimensional wing. A, tailed wing;O, tail-less wing; filled symbols indicate that angles of attack were also measured.(A) Polar curve; (B) lift coefficient; (C) drag coefficient.

(1967) and Newman et al. (1977), respectively. The two-dimensional pitchingmoment around the aerodynamic centre, which is assumed to be a line connectedto the quarter-chord of the aerofoil along the span, is small and is thereforeneglected. Although the present curve shown by a solid line gives a higher profiledrag at large angles of attack than do the others, the above aerofoil data areutilized in the following calculations for both fore and hind pairs of wings.

Analysis by means of the local circulation method

Let us consider first the flight performance of a dragonfly at various speeds. The

Page 17: flight performance of a dragonfly - The Journal of Experimental Biology

Dragonfly flight 237

J1-5

U 1-0

0-5

S -0-5

- 1 0

-1-5,

A \\\

\\\

I, max ' i

-90 -60 -30 0 30 60 90Angle of attack, a (degrees)

U

1-4

1-2

10

c no

.a °'SssR 0-6

0-4

0-2

0

BLarge dragonfly Ujajst h i n d w i n g

wing model

\Re =

01 0-2 0-3 0-4Drag coefficient, Cd

Fig. 9. Aerofoil characteristics of a dragonfly and other wings. (A) Lift and drag:estimated values based on the experimental data of Anax parthenope julius. (B) Polarcurve: results from present observations and for large and small dragonfly wing model(from Newman et al. 1977), locust hindwing (Jensen, 1956) and cambered and flatfruitfly wings (Vogel, 1967).

spanwise variation of the airloading and the total aerodynamic forces andmoments acting on the wings are calculated by the local circulation method(LCM). A detailed description of the method is given by Azuma et al. (1985), andonly a brief explanation is given here.

^ A wing in beating motion is hypothesized to consist of a plurality of ellipticalRdngs, and the respective elliptical wings are supposed to operate in the inducedflow field generated by them. The timewise change of the induced velocity at any

Page 18: flight performance of a dragonfly - The Journal of Experimental Biology

238 A. AZUMA AND T. WATANABE

Vertical force

Horizontal forceWake sheet of hindwing

Flight directionWake sheet of forewing

Trailing vortices Shed vortices

Fig. 10. Mathematical model of wake vortices.

local (or spanwise and azimuthal) station is given by multiplying an attenuationcoefficient, which is decided by the trailing and shed vortices left in a simple (not afree) wake model as a function of the spanwise distance, by the azimuth angle.Thus, the calculation is time-consuming and yet considered to be sufficientlyaccurate, without suffering from the computational divergence which sometimesappears in free-wake analysis.

From the actual wake vortices shown in Fig. 6A,B> a mathematical model of thewake sheets can be derived from the tracings of beating wings (Fig. 10). Then theattenuation coefficients are determined by the Bio-Savart law from the trailingvortices, which are assumed to be combined with the tip vortices of the respectivewing tips, and from the shed vortices lying on the wake sheets.

Although the attenuation coefficient can be given as a function of span positionand azimuth angle, the spanwise distance is fixed at the three-quarter position tosimplify the calculations in the present analysis.

The method is based on the blade element analysis but is different from otherprevious studies as follows, (i) The aerodynamic coefficients are used in nonlinearforms as functions of angles of attack that include the stalled range, (ii) Theinduced velocity is not homogeneous on the stroke planes and is obtained by theLCM, which also includes the unsteady effect, (iii) The control inputs for freeflight in computer simulation are the feathering angles along the wing span, the^flapping angles, the tilt angles of the stroke planes, and the beating frequency fon

Page 19: flight performance of a dragonfly - The Journal of Experimental Biology

Dragonfly flight

Lift (dL)

Angle of attack

Induced velocity

ine

Projection of flight velocity

Beating speed n//cos/8

Inflow angle

239

Inflow angle

Pitch angle

Induced velocity

Projection of flight velocity

Beating speed n/>cos/3

Chord line

Fig. 11. Relative velocities and aerodynamic forces acting on a blade element.(A) Downstroke; (B) upstroke.

given flight speeds.When the flight conditions and the wing motion are known, the airloading, and

thus the total aerodynamic forces and moments as well as the inertial forces andmoments, can be calculated by the LCM.

By referring to Fig. 11, the lift, drag and feathering moment acting on a wingelement at spanwise station r with azimuth angle ip are given as follows:

dL = !pU2cQ((*)dr

dD = |pU2cCd(or)dr

dM = ipU2c2Cm(ar)dr

(7)

Page 20: flight performance of a dragonfly - The Journal of Experimental Biology

240

where

A. AZUMA AND T. WATANABE

a=6-<j>

UT = Vcosycosi/; + npcos/3

Up = V(sinycos/3 + cosysini//sin/3) + v

= tan-1(UP/UT).

(8)

Then, aerodynamic forces and moments generated by the wing element can begiven by:

dFv.A = dL[cos0(sinysini/;sin/3 + cosycos/3) + sin^sinycosi/;]

+ dD[cos0cosi/>siny- sin0(sinysini/;sin/3 + cosycos/3)]

dFH,A = dL[cos0(sinycos/? - cosysim//sin/3) - sin0cosycosi//]— dD[sin0(sinycos/3 — cosysin/3sim/>) + cos0cosycost//]

dQA = r(dLsin0

,A = dMj.A + dFv.AxCG -

where dMj A is the pitching moment acting on an apparent joint, which is ahypothetical joint at the root of the feathering (or elastic) axis, and is given by:

A = rdL(cos0cos7/; - sin0sini/;cos/3) -+ dMcosi/>cos/3.

cos0sini/;cos/

Similarly, by assuming that every angular motion is small and neglectingcoupling terms, the inertial forces and moments acting on a wing element can begiven by:

where

dFv.i = — dmrcos/3(i/»2sini/; — i/;cos7//)siny

dFH.i = -dmrcos/?(i/;2sini// - i/)cost//)cosy

dQj = dmr2cos2/fy

+ dFvjXcG - dF M J z C G

(11)

j = -dmr2cos/3sin^(^2sinip + T//COST//) . (12)

Then,.the total vertical force, horizontal force and pitching moment aboutcentre of gravity can be given by integrating the above elemental forces Qmoments along the wing span and by summing the individual components as

Page 21: flight performance of a dragonfly - The Journal of Experimental Biology

Dragonfly flight 241

follows:

F v = 2 | [(dFv.A + d F V J + dFhv,A + dFh

VJ)/dr]dr

F H = 2 j [(dF*H,A

M C G = 2 d M C G J + dMC G,A + dMCG,i)/dr]dr

(13)

where superscripts f and h show the values for forewing and hindwing, respect-ively. Their mean values (or time averages) for one period of beating motion aregiven by:

2n/co

F v = ((O/2JZ) j Fvdt0

2n/a>

H = {(O/2JT) J FHdto

Inlco

M C G = {(O/IJI) \ MC Gdt .0

(14)

Neglecting the feathering component, the power required to beat the wing is givenby:

2JI/W R

= (CO/2JI) j {2 j0 0

(15)

The vertical and horizontal forces are also normalized by the weight of thedragonfly and are expressed by the load factor, only the vertical component ofwhich is given by n in unit G, the ratio of the resulting acceleration and theacceleration due to gravity.

Performance

From the data given in Tables 2 and 3 for free flights, which were not exactlytrimmed flight (load factor n > 1), the necessary (mechanical) power is obtained asshown by the circles in Fig. 12. By referring to these data and by selecting modifiedfeathering angles as given in Table 4, the necessary power curve versus flight speedcan be calculated (Fig. 12). The curve passes through lower values than thoseobtained from the free-flight tests, because the calculated power is based onalmost completely trimmed flights (n ~ 1).

By assuming an available power to musculature mass ratio ofPa/mm = 260Wkg~1 (Weis-Fogh, 1975, 1977) and a musculature mass to totalmass ratio of mm/m = 0-25 (Greenwalt, 1962), the available power can be

Ptimated as Pa = 5-75xl0~2W. Then the top speed (Vmax) of this dragonfly is2ms" 1 (Fig. 12). This value can be increased either by reducing the estimated

drag area or by increasing the available power. The cruising flight speed (Vcruise,

Page 22: flight performance of a dragonfly - The Journal of Experimental Biology

242 A. AZUMA AND T. WATANABE

ox

ia 3

Available power Pa = 5-75 xlO"2W

\ Experiment 1\

Experiment'n-0-97

2 3 4 5 6 7Flight speed, V^s" 1 )

Fig. 12. Necessary power curve of the dragonfly, Anax parthenope (O, experimentaldata).

the maximum range speed), which is found at the point at which the necessarypower curve is at a tangent to a line drawn through the origin or (dP/dV)min, is3-5 m s"1 (Fig. 12). The minimum power speed (VP min, maximum endurance flightspeed) is 1-7 m s~'. The power required for hovering flight (PhOv) is estimated to be3-6xlO"2W.

Fig. 13A-D shows the time variations of the vertical and horizontal componentsof the aerodynamic force, torque about the flapping axis and moment about thecentre of gravity at V = 3-2 m s~'. They were calculated by the LCM and by bladeelement theory based on the constant induced velocity distribution. For thecalculation of the moment, the feathering moment around the elastic axis wasassumed to be zero (Cm = 0) because of ambiguous aerodynamic characteristicsdue to complex aerofoil configuration. It can be seen that by assuming a constantinduced velocity distribution some errors are introduced in these variations.However, as shown by the right-hand ordinate, their mean values are close to eachother in the vertical component of the force. Although the forces are in a well-balanced condition, Fy + Fy ~ W, and F'H + F v ^ 0, the mean moment about thecentre of gravity is not completely balanced but leaves a positive (pitch-up) valunder the assumption of zero feathering moment, Cm = 0. However, the valuenot too large and is within a range in which adjustment is possible by taking into

I

Page 23: flight performance of a dragonfly - The Journal of Experimental Biology

o x

X E o

_2

0-8

0-6

0-4

0-2

Dow

nstr

oke

of f

orew

ing

Ups

trok

e of

fore

win

g

Dow

nstr

oke

Ups

trok

e of

hin

dwin

gD

owns

trok

e

' = 3

-6xK

T2 s

-

0 -1 -2

3 s

or,

i o X 5

1

0-8

0-6

0-4

0-2 n

0-25

T0-

5T0-

75T

• C

0

-0-2

-0-4

-0-6

-0-8

-1

0

Dow

nstr

oke

of f

orew

ing

UD

Stro

ke o

f fo

rew

ing

Dow

nstr

oke

Ups

trok

e of

hin

dwin

gD

owns

trok

e

Hin

dwin

g

Fore

win

g

= 3

-6xl

(T2 s

00-

25T

0-5T

0-75

T

.9

- 0-2

-0-4

-0-6

-0-8

-10

i

10

0-8

0-6

0-4

0-2 0

-0-2

-0-4

-0-6

-0-8

-10

Hin

dwin

g

Dow

nstr

oke

of f

orew

ing

Mea

n va

lue

|L

CM

Ups

trok

e of

for

ewin

g

Dow

nstr

oke

Ups

trok

e of

hin

dwin

g ,

Dow

nstr

oke

O

0 I

-1

0-25

T0-

5T0-

75T

Dow

nstr

oke

of f

orew

ing

Ups

trok

e of

for

ewin

g

Dow

nstr

oke

Ups

trok

e of

hin

dwin

g D

owns

trok

e

T=

3-6x

lO~

2s

0-25

T0-

5T0-

75T

Tim

e

Fig

. 13

. A

erod

ynam

ic f

orce

s an

d to

rqu

e. E

xper

imen

t 4;

V =

3-2

ms

l , dr

agon

fly

B;

heav

y li

nes

show

the

val

ues

calc

ulat

ed b

y th

eL

CM

, th

in l

ines

the

val

ues

calc

ulat

ed b

y si

mpl

e m

omen

tum

the

ory

(v =

con

stan

t),

solid

lin

es f

orew

ing,

and

das

hed

line

s hi

ndw

ing.

(A)

Ver

tical

for

ce;

(B)

hori

zont

al f

orce

; (C

) to

rqu

e; (

D)

pitc

hing

mom

ent.

Grq

Page 24: flight performance of a dragonfly - The Journal of Experimental Biology

to

Tab

le 4

. M

odif

ied

coef

fici

ents

in

Fou

rier

exp

ansi

on s

erie

s of

feat

heri

ng m

otio

n fo

r tr

imm

ed

flig

ht

Dra

gonf

ly

A

A

A

BE

xper

imen

t 1

2 3

4

For

ewin

gA

mpl

itud

e (d

egre

es)

0O

89

90

90

78

80

82

70

77

81

(0-2

5R/0

-5R

/0-7

5R)

0 x

23

38

52

89 5890 61

90 6378 67 83 32

80 67 82 33

82 68 79 35

Hin

dwin

gA

mpl

itud

e (d

egre

es)

0O

83

82

79

71

72

74(0

-25R

/0-5

R/0

-75R

) 6 X

18

28

37

>B

lank

spa

ces

in t

he m

odif

ied

data

sho

w t

hat

the

valu

es a

re t

he s

ame

as i

n th

e ex

peri

men

tal

data

abo

ve a

nd h

ave

bee

n o

mit

ted

. ^ w w

Page 25: flight performance of a dragonfly - The Journal of Experimental Biology

Dragonfly flight 245

*>1

oXEZ,

o

que

10-8

0-6

0-4

0-20

-0-2

-0-4

-0-6

-0-8

-1-0

Downstroke of fore wing ̂ Upstroke of forewing

Inertial component Total torque

Aerodynamic component

T = 3-6xlO~2

0-25T 0-5T 0-75T

1

0-8

0-6

0-4

0-2

-0-6

-0-8

-1-0

Downstroke t Upstroke of hindwing t Downstroke

BTotal torque

. Aerodynamic component

0-25T 0-5T

= 3-6xlO"2s

0-75T

~ 2

- l

- 2

Downstroke of forewing , Upstroke of forewingDownstroke. Upstroke orhindwing ^)ownstroke

Inertial component= 3-6xlO~2s

2 £c

1 So

-l -3

- 2

0-25T 0-5TTime

0-75T

Fig. 14. Inertial and aerodynamic torque and force. Experiment 4; V = 3-2ms ',dragonfly B. (A) Forewing; (B) hindwing; (C) vertical force component.

account the feathering moment which is considered to be negative for a positivecamber (upward convex) in the aerofoil.

Fig. 14 shows the inertial torque of the respective wings and the verticalcomponent of the inertial force of the total wings in comparison with theaerodynamic component and the resultants. Although the respective inertial andaerodynamic components are of a comparable order of magnitude, the totaltorque and force are not large. The variation in the total vertical force is within 0Gand +3G in each beating cycle. As stated in Appendix A, it is important torecognize that the maximum value does not exceed n = 3G at the flight speed of3-2ms-J.

Fig. 15 shows the spanwise load distribution of vertical and horizontal com-ponents of the total force acting on the fore- and hindwings.

Fig. 16 shows the timewise variation of angle of attack on the stroke planes ofthe wings in cruising flight. It is interesting that a region of large positive angle ofattack is observed in the final stage of the downstroke near the wing tip of theforewing and a region of large negative angle of attack is observed in the earlystage of the upstroke near the midspan of the forewing.

Fig. 17 shows the share of the lift and drag components in the mean vertical

Page 26: flight performance of a dragonfly - The Journal of Experimental Biology

246 A. AZUMA AND T. WATANABE

force, F v . At very low speed, including hovering, a contribution of the dragcomponent on the vertical force cannot be neglected. This is not unconnected withthe fact that the dragonfly flies with its stroke plane tilted from the horizontal evenwhen hovering, although this increases the induced power required. This makesthe transition from hovering flight to other flight modes easier.

10r A

0-5

0

-0-51-0

0-5

0

0-5

-0-5 L

1 0 r

0-5

-0-5Ll-0r

0-5

j i i i

0 0-2 0-4 0-6 0-8 1-0 0 0-2 0-4 0-6 0-8 1 0Span position

Fig. 15A

Page 27: flight performance of a dragonfly - The Journal of Experimental Biology

Dragonfly flight 247

Appendix A

Fig. 18 is compiled from data obtained from an experiment on the dragonfly(Libellula luctuosd) (Somps & Luttges, 1985). The following characteristics of

10r B

0-5

0

-0-51 0

0-5

-0-51-0

0-5

-0-51 0

0-5

0

-0-510r

0-5

0

- 0 - 5 L

1 0 r

0-5

0

-0-510

0-5

-0-51-0

0-5

0

-0-5

1-0

0-5

0

- 0 - 5

j 1 1

0 0-2 0-4 0-6 0-8 1 0 0 0-2 0-4 0-6 0-8 1-0Span position

Fig. 15. Spanwise airload distribution. Experiment 4; V = 3-2ms~', dragonfly B;vertical load, horizontal load. (A) Forewing; (B) hindwing.

Page 28: flight performance of a dragonfly - The Journal of Experimental Biology

248 A. AZUMA AND T. WATANABE

Upstroke Downstroke

Upstroke

-10

Downstroke

Fig. 16. Timewise and spanwise variations of angle of attack. Experiment 4,V = 3-2ms"1, dragonfly B. (A) Forewing; (B) hindwing. Angles are given in degrees.

Drag component, FD/FV

00 10 20 30 4-0 5-0Flight speed, V (ms"1)

Fig. 17. The share of lift and drag components in the mean vertical force.

Page 29: flight performance of a dragonfly - The Journal of Experimental Biology

Dragonfly flight 249

\

2 4x.

- 4 L

9 18 27Time(sxlO~3)

36

Fig. 18. Beating motion and vertical force of the dragonfly, Libellula luctuosa (takenfrom Somps & Luttges, 1985). (A) Projected angle in vertical plane; (B) projectedangle in horizontal plane; (C) total lift.

wing motion and vertical forces (upward positive) can be obtained:

ipf = 0-223 + 0-852cos(cot) rad

iph = -0-048 + 0-763cos(otf + 1-274) rad

where

F v = [1-77 + 4-81cos(wt - 0-297)] x 10"2N

= 48° = 0-838 rad, yh = 55° = 0-960rad ,

(A.I)

(A.2)

(A.3)

(A.4)

(A.5)

Page 30: flight performance of a dragonfly - The Journal of Experimental Biology

250 A. AZUMA AND T. WATANABE

-100-25T 0-5T 0-75T

80

20

0

-2-0

-4-0

Downstroke

B

Upstroke

Total force of forewing

Aerodynamic component

7Inertial component = 3-6xl(T2s

0 0-25T 0-5T 0-75T

8-0

60

4-0

2-0

0

- 2 0

-4-0

Downstroke Upstroke Downstroke

Total force of hindwing

Aerodynamic component

' Inertial component

•y

= 3-6xlO~2s

0 0-25T 0-5TTime

0-75T

Fig. 19. Components of the vertical force calculated from the data of Libellulaluctuosa. (A) Total force; (B) forewing; (C) hindwing.

and the angular transformation between two definitions in Fig. 1 and Fig. 18 aregiven by

\p= - t a n ^ t a n i / ^ s i n y - tan/3*cosy) j

/3 = tan-'[cos7/>(tan/3*siny + tam//*cosy)] . J

Assuming that the above vertical force is evenly shared by the two pairs of wingand that the mass of the respective wings can be estimated by the similarity rule as

I

Page 31: flight performance of a dragonfly - The Journal of Experimental Biology

Dragonfly flight 251

derived from Anax parthenope, the vertical forces generated by the respectivepairs of wings are represented by:

F^ = {8-83 + [24-0/cos(0-637)]cos(o* - 0-934)} x 10~3 N 1

F v = {8-83 + [24-0/cos(0-637)]cos(o>t - 0-297)} x 10~3 N J

and are shown in Fig. 19 for the aerodynamic and inertial components. Because ofthe above assumption, the result is only a rough estimation of the vertical force,but the aerodynamic component of the vertical force in the respective wings can begiven over a wide range of the downstroke as described in the present paper.

However, in the experimental results obtained by Somps & Luttges (1985)(Fig. 18) there are two difficulties: the vertical force (over20G) and its mean value(5-3G) are both too large. These extremely large values are not realistic for theflight of living creatures, except when rapidly manoeuvring. The large valuesprobably result from the fact that the inertial force related to the body mass wasnot completely removed from the measurement of the vertical force.

The authors would like to thank Mr Masakatsu Takao, Mr Masayuki Kitamura,and other staff of the Japan Broadcasting Corporation (NHK) for stimulating us totry vortex-wake visualization by their scientific program on dragonflies. Theauthors are also indebted to Mr Shunji Oba, Iwata Agricultural High School, forcatching dragonflies, and Professor Keiji Kawachi and Mr Isao Watanabe, TheUniversity of Tokyo, who assisted in the computer programming and the wind-tunnel test, respectively.

ReferencesAZUMA, A. (1981). Unsteady fluiddynamics related to the motion of living creatures. / . Jap. Soc.

aeronaut. Space Sci. 29, 91-96 (in Japanese).AZUMA, A., AZUMA, S., WATANABE, I. & FURUTA, T. (1985). Flight mechanics of a dragonfly.

J. exp. Biol. 116, 79-107.DIEDERICH, F. W. (1952). A simple approximate method for calculating spanwise lift

distributions and aerodynamic influence coefficients. NACA TN 2751.EDWARD, R. H. & CHENG, H. K. (1982). The separation in the Weis-Fogh circulation-generation

mechanism. J. Fluid Mech. 120, 463-473.GREENWALT, C. H. (1962). Dimensional relationships for flying animals. Smithson. misc. Colls

144, 1-46.HOAK, D. E. & ELLISON, D. E. (1968). USAF Stability and Control DATCOM. Flight Control

Division, Air Flight Dynamics Laboratory, Wright-Patterson Air Force Base, Ohio, October1960. (revised August 1968).

JENSEN, M. (1956). Biology and physics of locust flight. III. The aerodynamics of locust flight.Phil. Trans. Soc. Ser. B 239, 511-552.

LIGHTHILL, M. J. (1973). On the Weis-Fogh mechanism of lift generation. J. Fluid Mech. 60,1-17.

LIGHTHILL, M. J. (1975). Mathematical Biofluid Dynamics. Philadelphia, PA: Society forIndustrial and Applied Mathematics.

MAXWORTHY, T. (1979). Experiments on the Weis-Fogh mechanism of lift generation by insectsin hovering flight. I. Dynamics of the "flying". J. Fluid Mech. 93, 47-63.

IEWMAN, B. G., SAVAGE, S. B. & SCHOUELLA, D. (1977). Model tests on a wing section of an™ Aeschna dragonfly. In Scale Effects in Animal Locomotion (ed. T. J. Pedley), pp. 445-477.

London: Academic Press.

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252 A. AZUMA AND T. WATANABE

NEWMAN, J. N. (1977). Marine Hydrodynamics. Cambridge, MA, London: MIT Press.NORBERG, R. A. (1975). Hovering flight of the dragonfly, Aeshna juncea L., kinematics and

aerodynamics. In Swimming and Flying in Nature, vol. 2 (ed. Y.-T. Wu, C. J. Brokaw &C. Brennen), pp. 763-778. New York: Plenum Press.

SAVAGE, S. B., NEWMAN, B. G. & WONG, D. T.-M. (1979). The role of vortices and unsteadyeffects during the hovering flight of dragonflies. J. exp. Biol. 83, 59-77.

SOMPS, C. & LUTTGES, M. (1985). Dragonfly flight: novel uses of unsteady separated flows.Science 228, 1326-1329.

VOGEL, S. (1967). Flight in Drosophila. III. Aerodynamic characteristics of fly wings and wingmodels. J. exp. Biol. 46, 413-443.

VON LENDENFELD, R. (1881). Der Flug der Libellen - Anatomie und Physiologie der Flugorganeder Insectan. Sitzungsberichte der Kaiserlichen Akademie der Wissenschagten LXXXIII,289-380.

WATANABE, I., AZUMA, A. & WATANABE, T. (1986). Wake vortex of flying dragonflies. FourthInternational Symposium on Flow Visualization, August 26-29, ENSTA, Paris, France.

WEIS-FOGH, T. (1973). Quick estimates of flight fitness in hovering animals, including novelmechanisms for lift production. J. exp. Biol. 59, 169-230.

WEIS-FOGH, T. (1975). Flapping flight and power in birds and insects, conventional and novelmechanisms. In Swimming and Flying in Nature, vol. 2 (ed. Y.-T. Wu, C. J. Brokaw &C. Brennen), pp. 729-762. New York: Plenum Press.

WEIS-FOGH, T. (1977). Dimensional analysis of hovering flight. In Scale Effects in AnimalLocomotion (ed. T. J. Pedley), pp. 405-420. London: Academic Press.


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